-# ■' - ■ '■ ■-'; '■ '^virr*'^.;'',.' *M'i '■# M^ — XXiT zf NATIONAL LIBRARY OF MEDICINE Bethesda, Maryland ^, Gift of Jack H. U. Brown, Ph.D, (pd. JfC i^^y^^r^. _ &C4s£i y^^c^u- 4*. ■M j/. f)( cl .#? CARPENTER'S ELEMENTS OF PHYSIOLOGY. ELEMENTS OF PHYSIOLOGY, INCLUDING PHYSIOLOGICAL ANATOMY. BY WILLIAM B. CARPENTER, M.D., F.R.S., F.G.S., EXAMINER IN PHYSIOLOGY AND COMPARATIVE ANATOMY IN THE UNIVERSITY OF LONDON: AND AUTHOR OF " THE PRINCIPLES OF HUMAN PHYSIOLOGY," AND '; THE PRINCIPLES OF GENERAL AND COMPARATIVE PHYSIOLOGY," ETC. SECOND AMERICAN, FRO 31 A NEW AND KEVISKD LONDON EDITION. WITH ONE HUNDRED AND NINETY ILLUSTRATIONS. PHILADELPHIA: BLANCHARD AND LEA. 1851. C. SHERMAN, PRINTER. PREFACE. The present volume owes its origin to a desire on the part of the Publisher, that an elementary treatise on Physiology should be added to the series of admirable Students' Manuals, on the various depart- ments of Medical Science, which he had previously issued. In carrying this desire into execution, the Author has endeavored to avoid inflicting upon the class for whose use the Treatise is especially intended, the injury of placing in their hands such a superficial and imperfect sketch of the science, as, whilst affording them but a limited amount of knowledge of its facts, should leave them very ill-informed as to its general doctrines. His object has rather been to convey to the Student as clear an idea as possible of those Principles of Physiology which are based on the broadest and most satisfactory foundation, and to point out the mode in which these principles are applied to the explanation of the phenomena presented by the living actions of the Human body. In this manner has the Author desired to prepare him for that more detailed study of the latter, which becomes necessary when Physiology is pursued (as it ought to be) in connexion with the changes produced in the living body by Morbific and Remedial Agents, and is thus taken as a guide in the study of the causes, prevention, and treat- ment of Disease—which should be the primary object of attention with every one who undertakes the practice of his Profession. Although this Manual combines in some degree the scope of the Author's "Principles of Physiology, General and Comparative," and of his "Principles of Human Physiology," yet it cannot be regarded as a mere abridgment of them, having been written for the most part with very little reference to them, and with every desire to make it complete in itself. As the matter of which these volumes are composed is itself condensed to the utmost practicable degree, it is manifestly impossible that the present Manual should contain more than a mere outline' of the subjects of which they treat. To them, therefore, he Vlii PREFACE. would refer such of its readers as may desire further information upon various topics which are here only slightly touched upon; and in them, also, will be found references to various original authorities, the intro- duction of which would be incompatible with the limited scope of a treatise like the present. The Author has only to add, that he feels most grateful for the kind appreciation which this Manual has experienced; and that, in the prepa- ration of the present Edition, he has used his best endeavors to render it still more worthy of a favorable reception. The whole treatise has been subjected to a most careful revision; many statements which the advance of science has shown to be doubtful or erroneous, have been omitted or corrected; and a considerable amount of new matter has been introduced. Of the First, Eleventh, and Twelfth Chapters, more especially, a considerable proportion has been entirely rewritten; and the Author ventures to believe that the doctrines which they contain will enable such as may master them to obtain a clearer comprehension of the facts of Physiological Science than they could previously have acquired. Regent's Park, London, September, 1851. TABLE OE CONTENTS. BOOK I. GENERAL PHYSIOLOGY. CHAPTER. PAGE. I. On the Nature and Objects of the Science of Physiology . . 17 1. General Characters of Organized Structures ... 18 25 44 53 56 2. Digestive Characters of Vital Actions 3. Of the Forces concerned in the production of Vital Phenomena 4. Of Degeneration and Death ..... 5. General Summary .... II. Of the External Conditions of Vital Activity . 1. Of Light, as a Condition of Vital Activity 2. Of Heat, as a Condition of Vital Activity . 3. Of Electricity, as a Condition of Vital Activity, 4. Of Moisture, as a Condition of Vital Activity 58 61 71 94 97 III. Of the Elementary Parts of Animal Structures . . . 105 1. Of the Primary Components of the Animal Fabric . . . 107 2. Of the Simple Fibrous Tissues . . . . . 119 3. Of the Basement or Primary Membrane .... 127 4. Of Simple Isolated Cells, employed in the Organic Functions . 130 5. Of Cells connected together, as permanent constituents of the Tissues 154 6. Of Cells coalesced into Tubes, with Secondary Deposit. . 196 BOOK II. SPECIAL PHYSIOLOGY. IV. Of Food, and the Digestive Process ..... 236 1. Sources of the Demand for Aliment ... • . 236 2. Of the Digestive Apparatus, and its Actions in general . . 252 3. Of the Movements of the Alimentary Canal . . . 257 4. Of the Secretions poured into the Alimentary Canal, and of the Changes -which they effect in its contents .... 264 5. Of Hunger, Satiety, and Thirst ..... 274 V. Of Absorption and Sanguification ..... 277 1. Of Absorption from the Digestive Cavity . . . 277 2. Of the Passage of Chyle along the Lacteals, and its admixture with the Lymph collected from the General System . .281 3. Of the Spleen, and other Glandular Appendages to the Lymphatic System ........ 286 4. Of the Composition and Properties of the Chyle and Lymph . 292 5. Of Absorption from the External and Pulmonary Surfaces . 296 6. Of the Composition and Properties of the Blood . . . 297 X TABLE OF CONTENTS. CHAPTER. VI. Or the Circulation of the Blood .... 1. Nature and Objects of the Circulation of Nutrient Fluid 2. Different forms of the Circulating Apparatus 3. Action of the Heart ..... 4. Movement of the Blood in the Arteries . 5. Movement of the Blood in the Capillaries 6. Movement of Blood in the Veins VII. Of Nutrition ...... 1. Selecting Power of Individual Parts 2. Varying Activity of the Nutritive Processes 3. Of Death, or Cessation of Nutrition 4. Disordered Conditions of the Nutritive Processes . VIII. Of Respiration ...... 1. Essential Nature and Conditions of the Respiratory Process 2. Different forms of the Respiratory Apparatus in the lower Animals 3. Mechanism of Respiration in Mammalia, and in Man . 4. Chemical Phenomena of Respiration 5. Effects of Insufficiency, or Suspension, of the Aerating Process IX. Of Secretion ........ 1. Of the Secreting Process in general, and of the Instruments by which it is effected ....... 2. Of the Liver, and the Bile ..... 3. Of the Kidneys, and the Urine ..... 4. Of the Cutaneous and Intestinal Glandulas 5. General Summary of the Excreting Processes X. Of the Development of Light, Heat, and Electricity, in the Animal Body ......... XI. Generation and Development 1. General View of the Nature of the Process • 2. Action of the Male . . . . 3. Action of the Female XII. Of the Nervous System 1. General view of the operations, of which the Nervous System is the Instrument ....... 2. Comparative Structure and Actions of the Nervous System 3. Functions of the Spinal Cord and its Nerves 4. Functions of the Medulla Oblongata 5. Functions of the Sensory Ganglia 6. Functions of the Cerebellum 7. Functions of the Cerebrum 8. Functions of the Sympathetic System XIII. Of Sensation, General and Special . 1. Of Sensation in general 2. Of the Sense of Touch . 3. Of the Sense of Taste 4. Of the Sense of Smell . 5. Of the Sense of Hearing 6. Of the Sense of Sight . PIOE. 304 304 305 320 327 331 343 343 345 350 352 357 357 363 375 382 389 393 393 400 40G 416 422 425 434 434 442 445 473 473 479 496 504 508 514 516 527 529 529 534 536 538 540 544 XIV. Of the Voice and Speech . 553 LIST OE WOOD ENGRAVINGS. FIO. 1. Simple isolated Cells, containing reproductive molecules 2. Fibrous structure of exudation-membrane ; after Gerber . . 3. Fibrous membrane lining egg-shell (original) .... 4. "White fibrous tissue of areolar tissue and tendon; after Gerber . 5. White fibrous tissue of ligament; after Gerber 6. Yellow fibrous tissue of ligamentum nuchse ; after Gerber 7. Development of fibres from cells; after Lebert 8. Ideal Section of a Joint ....... 9. Capillary vessels of Skin; after Berres .... 10. Capillary vessels of Intestinal villi; after Berres .... 11. Capillary vessels around orifices of Mucous follicles ; after Gerber . 12. Capillary vessels around follicles of Parotid Gland ; ditto 13. Distribution of Sensory nerves in Skin; after Gerber 14. Primary membrane, with germinal spots ; after Goodsir . 15. Primary membrane, showing component cells ; after Goodsir 16. Cells from Chorda Dorsalis of Lamprey; after Quekett . 17. Multiplication of Cartilage-cells by duplication; after Leidy 18. Parent-cells, with contained secondary cells, of cancerous structure; after Lebert . . . . . . f . 19. Cells from fluid of Herpes; after Addison .... 20. Oblique section of Epidermis ; after Henle .... 21. Epidermic cells from Conjunctiva; after Gerber 22. Portion of Choroid-coat, showing pigment cells ; ditto 23. Separate Pigment-cells; after Mandl ..... 24. Detached epithelium-cells from mucous membrane of mouth ; after Lebert 25. Pavement-epithelium from bronchial tubes ; after Lebert 26. Layer of cylindrical epithelium, with cilia; after Henle . 27. Follicles from liver of Crab, with contained secreting cells; after Goodsir 28. Follicles of Mammary gland, with contained secreting cells ; after Lebert 29. Secreting Cells of Human Liver ..... 30. Formation of Spermatozoa within cells; after Wagner 31. Diagram of Intestinal Mucous membrane, in intervals of digestion; after Goodsir ........ 32. Extremity of Placental villus; after Goodsir .... 33. Progressive stages of cell-growth, in Shell-membrane (original) 34. Progressive stages of coalescence of cells, in Shell-membrane (original) 35. Fusiform tissue of plastic exudations; after Lebert 36. Areolar and Adipose tissue; after Mandl .... 37. Capillary network around Fat-cells; after Berres 38. Cartilage of Mouse's ear; after Quekett .... 39. Section of Cartilage; after Schwann" ..... 40. Distribution of vessels on surface of Cartilage; after Toynbee 41. Nutrient vessels of Cornea; after Toynbee .... 42. Shell of Echinus (original) ...... 43. Sections of Shell of Pinna (original) ..... 44. Tubular shell-structure from Anomia (original) 45. Cancellated structure at extremity of Femur; after Toynbee Xll LIST OF WOOD ENGRAVINGS. FIG. 46. 47. 48. 49. 50. 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 74. 75. 76. 77. 78. 79. 80. 81. 82. 83. 84. 85. 86. 87. 90. 91. 92. 93. 94. 95. 96. 97. 98. 99. 100. 101. 102. 103. 104. 105. 106. 107. 108. Lacunae of Osseous substance ; after Mandl . Section of Bony Scale of Lepidosteus (original) . Network of Haversian canals, from vertical section of Tibia; after Mandl Transverse section of long bone; after Wilson Section of Cartilage, near seat of Ossification ; after Wilson Section of Cartilage, at the seat of Ossification; after Wilson Vessels of Dental Papilla; after Berres Oblique Section of Dentine ; after Owen . Vertical Section of human molar tooth ; after Nasmyth Development of Teeth ; after Goodsir Structure of Hair of Musk-deer and Sable (original) Structure of the Human Hair; after Wilson Fasciculus of striated Muscular fibre; after Mandl . Non-striated Muscular fibres; after Bowman Striated Muscular fibre separating into fibrillre Muscular fibre cleaving into disks; after Bowman Transverse section of muscular fibres ; after Bowman Structure of ultimate fibrillre of striated fibre (original) Nucleated fibres from non-striated muscle; after Wilson Fusiform contractile cells ; after Kolliker Nuclei in striated muscular fibres of foetus; after Bowman . Capillaries of Muscle ; after Berres Distribution of nerves in Muscle; after Burda,ch Components of gray substance of brain; after Purkinje Capillaries of Nervous centres ; after Berres Structure of Ganglion of Sympathetic ; after Valentin . Distribution of Sensory nerves in lip ; after Gerber . Capillaries at margin of lips; after Berres Section of Human Stomach ..... Mucous coat of small intestine, showing Villi, and orifices of follicles ; afte Boehm ........ Peyerian glandula; after Boehm .... Stomach of Sheep ....... Section of Stomach of Sheep, showing demi-canal; after Flourens Lobule of Parotid Gland ; after Wagner .... Gastric glanduloe ; after Wagner .... Orifices of Gastric tubuli; after Boyd .... Distribution of Capillaries in Intestinal Villus; after Berres Commencement of Lacteal in Intestinal Villus ; after Krause . Diagram of Lymphatic gland ; after Goodsir Epithelial cells of intra-glandular Lymphatic; after Goodsir Course of Thoracic duct ..... Appearance of inflamed Blood; after Addison . Vascular area of Fowl's egg; after Wagner . Diagram of the Circulation in Fish Diagram of the Circulation in Reptile Diagram of complete Double Circulation . Anatomy of Human Heart and Lungs Capillaries of Nervous centres; after Berres Capillaries of Glandular follicles; after Berres Capillaries of Conjunctival membrane; after Berres Capillaries of Choroid coat; after Berre3 Capillaries around orifices of mucous follicles; after Berres Capillaries in Skin of finger ; after Berres Capillaries in fungiform papilla of Tongue; after Berres Doris, showing branchial tufts ; after Alder and Hancock . One of the arborescent processes of gills of Doris; ditto Respiratory apparatus of Insects .... Diagram of different forms of Respiratory Apparatus (original) Capillaries of Gill of Eel (original) . . . . Section of Lung of Turtle ; after Bojanus Capillaries of Human Lung (original) Simple glandular follicles; after Miiller . Embryonic development of Liver; after Miiller LIST OF WOOD ENGRAVINGS. Xlll FIG. 109. 110. 111. 112. 113. 114. 115. 116. 117. 118. 119. 120. 121. 122. 123. 124. 125. 126. 127. 128. 129. 130. 131. 132. 133. 134. 135. 136. 137. 138. 139. 140. 141. 142. 143. 144. 145. 146. 147. 148. 149. 150. 151. 152. 153. 154. 155. 156. 157. 158. 159. 160. 161. 162. 163. 164. 165. Rudimentary Pancreas, from Cod; after Miiller . Mammary Gland of Ornithorhyncus ; after Miiller . Meibomian Glands; after Miiller . Portion of Cowper's Gland ; after Miiller Lobule of Lachrymal Gland; after Miiller Hepatic Follicles from Crab ; after Goodsir . Ultimate follicles from Mammary gland; after Lebert Surface of Lobule of Liver of Squilla; after Miiller Interior of Lobule of Liver of Squilla; after Miiller Liver of Tadpole; after Miiller ..... Distribution of Blood-vessels in Lobules of Liver; after Kiernan Connexion of Lobules of Liver with Hepatic vein ; ditto Distribution of Hepatic ducts around Lobules of Liver; after Kiernan Secreting Cells of Liver ...... Development of Kidney in embryo of Lizard; after Miiller Kidney of foetal Boa; after Miiller ..... Portion of Kidney of Coluber; after Miiller Fasciculus of tubuli uriniferi of Bird; after Miiller . Section of Kidney ....... Section of portions of Kidney, slightly magnified; after Wagner Distribution of vessels in Kidney ; after Bowman Vertical section of Skin ; after Wilson Hepatic Cells gorged with Fat; after Bowman Various stages of development of HjEmatococcus binalis; after Hassal Successive stages of development of simpler Algae; after Kutzing Diagram representing the three principal forms of the Generative process in Plants (original) ....... Successive stages of Segmentation of the vitellus of Ascaris; after Bagge Anatomy of the Testis ...... The Uterus and its appendages ...... Successive stages of Segmentation of Mammalian vitellus; after Coste Formation of the Mulberry mass ; after Coste .... Plan of Early Uterine Ovum; after Wagner . . Germ and surrounding parts; after Coste .... Vascular Area of Fowl's Egg ; after Wagner Diagram of Ovum, at commencement of separation of digestive cavity; afte Wagner ........ Diagram of Ovum, showing the formation of the Amnion ; after Wagner Diagram of Human Ovum, in second month, showing the Allantois; after Wagner ......... Diagram of the Circulation in the Ovum at the commencement of the forma tion of the placenta; after Coste Extremity of Placental Villus ; after Goodsir External membrane and cells of placental villus; after Goodsir Diagram illustrating the arrangement of the placental decidua ; after Goodsir Plan of the Foetal Circulation ...... Termination of portion of milk-duct in follicles; after Sir A. Cooper Portion of the Ganglionic tract of Polydesmus; after Newport Human embryo at sixth week; after Wagner Dissection of the Medulla Oblongata, showing the connexions of its tracts; after Solly (altered) .... Diagram of the relations of the Cerebrum to the Sensory Ganglia, as horizontal section (original) .... Diagram of the relations of the several parts of the Encephalon, as vertical section (original) ..... Capillary network at margin of Lips ; after Berres Distribution of tactile nerves in Skin; after Gerber . Capillaries of fungiform papilla of Tongue; after Berres Distribution of Olfactory nerve; after Wilson Diagram of the Auditory apparatus ; after Wilson Refraction of rays of light through convex lens Formation of images in eye .... Capillary network of Retina ; after Berres . Structure of the Larynx; after Willis several EXPLANATION OE PLATE I. The Figures in this Plate represent the Cells floating in the various animal fluids ; and they are all, with the exception of Figs. 4 and 5, copied from the representations given by M. Donne" in his "Atlas de l'Anatomie Microscopique." These representations are transcripts of Daguerreotype pictures, obtained from the objects, by a solar microscope, with a magnifying power of 400 diameters. Fig. 1. Red Corpuscles of Human Blood, viewed by their flattened surfaces ($ 215). Fig. 2. Red Corpuscles of Human Blood, adherent by their flattened surfaces, so as to form rolls;—at a, the entire surfaces are adherent; at b, their surfaces adhere only in part. Fig. 3. Red Corpuscles of Human Blood, exhibiting the granulated appearance which they frequently present, a short time after being withdrawn from the vessels. Fig. 4. Colorless Corpuscles of Human Blood (§ 214). Fig. 5. The same, enlarged by imbibition of water. Fig. 6. Red Corpuscles of Frog's Blood (§ 215). Fig. 7. The same, treated with dilute acetic acid;'the first effect of which is to render the nucleus more distinct, as at b; after which the outer vesicle becomes more transparent, and its solution commences, as at a. Tig. 8. The same, treated with water; at a is seen a corpuscle nearly unaltered, except in having the nucleus more sharply defined; at b, others which have become more spherical, under the more prolonged action of water; at c, the nucleus is quitting the centre, and approaching the circumference, of the disk; at d it is almost freeing itself from the envelope; and at e it has completely escaped. Fig. 9. Globules of Mucus, newly secreted (g 237). Fig. 10. The same, acted on by acetic acid. Fig. 11. Globules of Pus, from a phlegmonous abscess ($ 637). Fig. 12. The same, acted on by acetic acid. PJ.ATJE 1 tr:; mm m *..Jf: cV, !$&■ -Ms :NT *r*r- ^i $■ <© „Ay^ *r%A w m. 4 £** EXPLANATION OE PLATE II. The Figures in this Plate represent the principal forms of the Nervous centres in diffe- rent classes of animals. The 1st is copied from a Memoir by M. Blanchard; the 2d, 3d, and 4th, from Mr. Newport's delineations; the 5th to the 13th from the work of M. Guillot on the Comparative Anatomy of the Encephalon in the different classes of Verte- brata; and the last two from the work of M. Leuret on the same subject. Fig. 1. Nervous System of Solen; a, a, cephalic ganglia, connected together by a trans- verse band passing over the CEsophagus, and connected with the other ganglia by cords of communication; b, pedal ganglion, the branches of which are distributed to the powerful muscular foot; c, branchial ganglion, the branches of which proceed to the gills d, d, the siphons e, e, and other parts. On some of these branches, minute ganglia are seen; as also at/, /, on the trunks that pass forwards from the cephalic ganglia (§ 852). Fig. 2. Nervous System of the Larva of Sphinx ligustri; a, cephalic ganglia; 1-12, ganglia of the ventral cord (§ 856). Fig. 3. Thoracic portion of the Nervous System of the Pupa of Sphinx ligustri; a, b, c, three ganglia of the ventral cord; d, d, their connecting trunks; e, e, respi- ratory ganglia (g 862). Fig. 4. Anterior portion of the Nervous System of the Imago of Sphinx ligustri; a, cepha- lic ganglia; b, b, eyes ; c, anterior median ganglion, and d, d, posterior lateral ganglia of stomato-gastric system ; e, f, large ganglionic masses in the thorax, giving origin to the nerves of the legs and wings (§ 863). Fig. 5. Brain of the Perch, seen from above (§ 869). Fig. 6. The same, as seen from below. Fig. 7. Interior of the same, as displayed by a vertical section. The following references are common to the three preceding, and to the succeeding figures. a, a, Olfactory lobes or ganglia. b, b, Cerebral ganglia or Hemispheres. c, c, Optic lobes. d, Cerebellum. e, Spinal Cord. /, Pineal gland. g, Lobi inferiores (their precise character not determined). h, Pituitary body. i, Optic Nerves. xvi EXPLANATION OF PLATE II. Fig. 8. Brain of the Common Lizard, seen from above (\ 871). Fig. 9. The same, as seen from below. Fig. 10. The same, as displayed by a vertical section. Fig. 11. Brain of the Common Goose, as seen from above (g 872). Fig. 12. The same, as seen from below. Fig. 13. The same, as displayed by a vertical section. Fig. 14. Brain of the Sheep, viewed sideways (g 873). Fig. 15. The same, as displayed by a vertical section. In addition to the parts indicated by the preceding references, we have here to notice;—k, the corpus callosum ; I, the septum lucidum ; and m, the Pons Varolii. pl Art' 1/ Wrrf^ ■vi ? • ft! 1 i! ! '1 I / ■J11) > >-i; f If /if/ / i;X^t ^||g*r. .•<:T.:- ,t "l ^m\ j \ i 'VIST v r^' \ t? 9 t" :"T . '11 JO m iV .1; mL * '\ffr w$& of the intestinal canal. (Fig. 29.) On the other hand, Avhere theocjdinr- der-epithelium lies upon a concave surface, the free extremities, df tbie cells may be smaller than those which are attached.. Sometime* eaeh. 146 STRUCTURE AND ENDOWMENTS OF ANIMAL TISSUES. cylinder is formed from more than one cell, as is shown by the nuclei it contains; although its cavity'seems to be continuous from end to end. And occasionally the cylinders arise by stalk-like prolongations, from Vibratile or ciliated Epithelium;—a, nucleated cells, resting on their smaller extremities; 6, cilia. a tessellated epithelium beneath. The two forms of Epithelium pass into one another at various points; and various transitional forms are then seen,—the tessellated scales appearing to rise more and more from the surface, until they project as long-stalked cells, truncated cones, or cylinders. 234. Both these principal forms of Epithelial cells are frequently observed to be fringed at their free margins with delicate filaments, which are termed cilia; and these, although of extreme minuteness, are organs of great importance in the animal economy, through the extra- ordinary motor power with Avhich they are endowed. The form of the ciliary filaments is usually a little flattened, and tapering gradually from the base to the point. Their size is extremely variable ; the largest that have been observed being about l-500th of an inch in length, and the smallest about l-13000th. When in motion, each filament appears to bend from its root to its point, returning again to its original state, like the stalks of wheat when depressed by the wind; and when a number are affected in succession with this motion, the appearance of progressive waves following one another is produced, as when a wheatfield is agitated by frequent gusts. When the ciliary motion is taking place in full activity, however, nothing whatever can be distinguished, but the whirl of particles in the surrounding fluid; and it is only when the rate of movement slackens, that the shape and size of the cilia, and the manner in which their stroke is made, can be clearly seen. The motion of the cilia is not only quite independent (in all the higher animals at least) of the will of the animal, but is also independent even of the life of the rest of the body; being seen after the death of the animal, and pro- ceeding with perfect regularity in parts separated from the body. Thus isolated epithelium cells have been seen to swim about actively in Avater, by the agency of their cilia, for some hours after they have been detached, from the mucous surface of the nose; and the ciliary movement has been seen fifteen days after death in the body of a Tortoise, in which putrefaction was far advanced. In the gills of the River Mussel, which are among the best objects for the study of it, the movement endures with similar pertinacity. 235. The purpose of this ciliary movement is obviously to propel fluids over the surface on which it takes place; and it is consequently limited in the higher animals to the internal surfaces of the body, and always takes place in the direction of the outlets, towards which it aids SIMPLE ISOLATED CELLS.—EPITHELIUM. 147 in propelling the various products of secretion. The case is^ different, hoAvever, among animals of the loAver classes, especially those inhabiting the water. Thus the external surface of the gills of Fishes, Tadpoles, &c, is furnished with cilia; the continual movement of which reneAVS the water in contact with them, and thus promotes the aeration of the blood. In the loAver Mollusca, and in many Zoophytes, which pass their lives rooted to one spot, the motion of the cilia serves not merely to produce currents for respiration, but likeAvise to draAV into the mouth the minute particles that serve as food. And in the free-moving Animal- cules, of various kinds, the cilia are the sole instruments which they possess, not merely for producing those currents in the water which may bring them the requisite supply of air and food, but also for propelling their own bodies through the Avater. This is the case, too, with many larger animals of the class Acalephae (Jelly-fish), which move through the water sometimes with great activity, by the combined action of the vast numbers of cilia that clothe the margins of their external surfaces. In these latter cases it would seem as if the ciliary moA'ement were more under the control of the will of the animal, than it is where it is con- cerned only in the organic functions. In what way the will can influence it, hoAvever, it does not seem easy to say; since the ciliated epithelium- cells appear to be perfectly disconnected from the surface on Avhich they lie, and cannot, therefore, receive any direct influence from their nerves. Of the cause of the movement of the cilia themselves, no account can be given ; they are usually far too small to contain even the minutest fibrillae of muscle; and Ave must regard them as being, like those fibrillae, organs sui generis, having their own peculiar endowment,—which is, in the higher animals at least, that of continuing in ceaseless vibration, during the Avhole term of the life of the cells to which they are attached. The length of time during which the ciliary movement continues after the general death of the body, is much less in the warm-blooded than in the cold-blooded animals ; and in this respect it corresponds with the degree of persistence of muscular irritability, and of other vital endowments. 236. The Tessellated-Epithelium, as already mentioned, covers the Serous and Synovial membranes, the lining membranes of the blood- vessels and absorbents, and the Mucous membranes with their glandular prolongations, except where the cylinder-epithelium exists. It presents itself, with some modifications presently to be noticed, in the ultimate follicles of all glands, and also in the smaller bronchial tubes. In this latter situation it is furnished with cilia; and these are also found on the cells of the tessellated epithelium, which covers the delicate pia mater lining the cerebral cavities. The cylinder-Epithelium commences at the cardiac orifice of the stomach, and lines the whole intestinal tube; and, generally speaking, it lines the larger gland-ducts, giving place to the tessellated form in their smaller ramifications. A similar epithelium, furnished with cilia, is found lining the air passages and their various offsets,—the nasal cavities, frontal sinuses, maxillary antra, lachrymal ducts and sac, the posterior surface of the pendulous velum of the palate and fauces, the Eustachian tubes, the larynx, trachea, and bronchi,— becoming continuous, hoAvever, in the finer divisions of the latter, with 148 STRUCTURE AND ENDOAVMENTS OF ANIMAL TISSUES. the ciliated pavement-epithelium. The upper part of the vagina, the uterus, and the Fallopian tubes, are also furnished with a ciliated Cylin- der-Epithelium. The function of the cilia in all these cases appears to be the same; that of propelling the viscid secretions, Avhich would othenvise accumulate on these membranes, towards the exterior orifices, Avhence they may be carried off. 237. The simplest office which the Epithelium-cells of Mucous mem- branes perform, appear to be that of elaborating a peculiar secretion termed Mucus; Avhich is destined to protect them from the contact of air, or from that of the various irritating substances to which they are exposed, in consequence of their peculiar position and functions. This Mucus is a transparent semifluid substance, distinguished by its peculiar tenacity or viscidity. It is quite insoluble in water ; but is readily dis- solved by dilute alkaline solutions, from which it is precipitated again by the addition of an acid. A substance resembling Mucus may be pro- duced from any fibrinous exudation, or even from pus, by treating it Avith a small quantity of liquor potassae. The secretion of Mucus, like the formation of Epidermis, appears to take place with an activity pro- portioned to the degree of irritation of the subjacent membrane. On many parts of the mucous surface, a sufficient supply is afforded by the epithelium-cells which cover it; but in other situations, especially along the alimentary canal, the demand is much greater, and it is probably supplied not merely by the cells of the surface, but by those lining the crypts or follicles which are formed by involutions of it. 238. The Epithelium-cells which are thus being continually reneAved on the Mucous surfaces, commonly seem to have their origin in the granular germs diffused through the basement-membrane; but it is dif- ferent in regard to the cells of the follicles, which seem rather to occupy their cavity than merely to line their walls, and which appear to be in course of continual production from a germinal spot, or collection of re- productive granules, at the blind extremity of the follicle. This is the case in the ultimate follicles of the more complex glands; which may be regarded as so many repetitions of the simple crypts or follicles in the substance of the mucous membranes;—the only difference being, that Fig. 27. Fig. 28. Fig. 29. Two follicles from the liver of Carcinus Ultimate follicles of Mammary Secreting cells of manas (Common Crab), with their con- gland with their secreting cells, Human Liver; a, nu- tained secreting cells. a, a ;—b, b, the nuclei. cleus; 6, nucleolus; c, oil-particles. the former pour their secretion into a branch of a duct, which unites with the_ other ramifications to form a trunk; and this trunk conveys them to theirdestination in some cavity lined by a mucous membrane;—whilst the simple follicles or crypts at once pour forth their secretion upon the SIMPLE ISOLATED CELLS.—SECRETING CELLS. 149 surface of the membrane. The preceding figure (27) represents tAvo follicles of the liver of the Common Crab, which are seen to be filled with secreting cells; it seems evident, from the comparative sizes of these cells in different parts, that they originate at the blind extremity of the follicle, Avhere there is a germinal spot; and that, as they recede from that spot, they gradually increase in size, and become filled with their characteristic secretion, being at the same time pushed onwards towards the outlet by the continual new growth of cells at the germinal spot. In Fig. 28 are shown the corresponding ultimate follicles of the Mammary gland; filled, like the preceding, with secreting cells. 239. The whole of the acts, then, by which t-he separation of the dif- ferent Secretions from the Circulating fluid is accomplished, really con- sist in the growth and nutrition of a certain set of cells, usually covering the free surfaces of the body, both internal and external, or lining cavi- ties Avhich have a ready communication with these by means of ducts or canals.* These cells differ widely from one another, in regard to the kind of matter which they appropriate and assemble in their cavities; although the nature of their walls is probably the same throughout. Thus Ave find biliary matter and oil, easily recognisable by their color and refracting poAver, in the cells of the liver; milk in the cells of the Mammary gland; sebaceous or fatty matter in the cells of the sebaceous follicles of the skin ; and so on. All these substances are derived from the blood; being either contained in it previously, or being elaborated from its constituents by a simple process of transformation,—as, for example, that Avhich converts the albumen of the blood into the caseine of milk. Hence they may be considered as the peculiar aliments of the several groups of cells ; whose acts of nutrition are the means of drawing them off, or secreting them, from the general circulating fluid. When they have attained their full groAvth, and accomplished their term of life, their Avails either burst or dissolve aAvay, and thus the contents of the cells are delivered into the cavity, or upon the surface, at Avhich they are required. Now as all the canals of the glands open either directly outwards upon the surface, or into cavities which communicate with the exterior, it is evident that the various products of the action of these epithelial cells must be destined to be cast forth from the body. This Ave shall find to be the case ; some of them, as the bile and urine, being excretions, of Avhich it is necessary to get rid by the most direct channel; whilst others, like the tears, the saliva, the gastric fluid, the milk, &c, are separated from the blood, not so much for its purification, but be- cause they are required to answer certain purposes in the economy. 210. Now whilst thus actively concerned in the Nutritive functions of the economy, and exercising in the highest degree their poAvers of selection and transformation, these Secreting cells appear to have nothing to do with the operation of Reproduction. We have seen that they do not even regenerate themselves; all their energies being, as it were, con- centrated upon their OAvn growth ; and the successive production of new broods of them being provided for by other means. Throughout the organized creation, it appears to be necessary that the true act of Gene- * The Synovial secretion is perhaps the only one which is poured into a closed 150 STRUCTURE AND ENDOWMENTS OF ANIMAL TISSUES. ration should be performed by the reunion of the products of two dis- tinct orders of cells ; the coalescence of which produces a germ, that is the starting point of a new organism;—thus differing from the act of Reproduction by gemmation or budding, which essentially consists in an extension or out-groAVth from the original organism, by the subdivision of cells in the manner already described (§ 212). Among the lowest Cellular Plants, in which every cell is apparently similar to the rest, this operation is effected by the "conjugation" of any pair (as it would seem) of the cells which have been produced by multiplication from the original germ. But in all the higher organisms, both Vegetable and Animal, Ave find that certain cells are set apart for this purpose; and that there is an obvious distinction betAveen the "germ-cells," from within which the germ ultimately makes its appearance, and the " sperm- cells," which communicate to them a fertilizing influence. Still in the loAver tribes, both of Plants and Animals, we find that " sperm-cells" and "germ-cells" are deA'eloped in the midst of the ordinary tissues of the body; and it is only as Ave ascend the scale, and find the principle of division of labor carried out in other Avays, that we meet, as in Man, with particular organs set apart for their evolution, and find these organs appropriate respectively to distinct individuals. 241. The spermatic cells of Man are developed within the tubuli of the Testicle; where they appear to hold exactly the same relation to the membranous walls of those tubuli, as do the secreting cells to the tubes and follicles of the proper Glands, being, in fact, the representa- tives of their epithelial cells (Fig. 30, a). Each of these developes in its interior a variable number of secondary cells, or " vesicles of evolution ;" and within every one of these is produced a single thread-like body, dilated at one extremity, and possessed of a remarkable self-moving power, which.is termed a Spermatozoa. Sometimes the vesicles of evolu- tion remain enclosed within the parent-cell, until their spermatozoa have been completely developed, and have been set free by their rupture (b); Fig. 30. Formation of Spermatozoa within seminal cells; a, the original nucleated cell; 6, the same enlarged, with the formation of the Spermatozoa in progress; c, the Spermatozoa nearly complete, but still enclosed within the cell. and thus, when they have all performed their office, the parent-cell con- tains nothing but a bundle of spermatozoa (c), whose dispersion takes place as soon as its cell-wall gives way. From the very peculiar motion they possess, the Spermatozoa Avere long regarded as distinct and SIMPLE ISOLATED CELLS.—REPRODUCTIVE CELLS. 151 independent animalcules; it is now generally admitted, hoAvever, that they have no more claim to a distinct animal character, than have the ciliated epithelia of mucous membrane, which will likewise continue in movement Avhen separated from the body. Similar bodies are formed by all the higher Cryptogamic Plants; and it appears from late re- searches that their office, as in Animals, is to fertilize the contents of the "germ-cells," with which their self-moving poAver brings them into contact (chap, xi.) It is a curious fact that the seminal cells, in which the Spermatozoa are formed, are ejected from the gland in certain Crus- tacea, not only before they have burst and set free their Spermatozoa, but even long before the development of the Spermatozoa in their inte- rior is completed;—thus affording a complete demonstration of their independent vitality. 242. The " germ cells," in like manner, are very commonly developed among the lower Animals as the epithelia of the tubes or follicles which constitute the ovary; but in Man and the higher Animals, the ovary is a solid organ, and the germ-cells are developed in its substance, lying in the midst of the dense fibrous tissue which forms its parenchyma. These germ-cells, Avhich are known as "ovisacs," like the sperm-cells, develope secondary cells or ova in their interior; each ovisac, however, producing but a single ovum. The ovum, again, contains a tertiary cell, the germinal vesicle, whose contents appear to mingle with those of the sperm-cell in the act of fecundation, so that the fertilized germ is the result; the remaining contents of the ovum being the nutritive materials, at the expense of which this germ undergoes its first development (chap, xi.) 243. We now proceed to a class of cells, which are equally indepen-/ dent of each other, which begin and end their lives as cells, without' undergoing any transformation, but which form part of the substance of the fabric, instead of lying upon its free surfaces and being continually cast off from them. Still their individual history is much the same as that of the cells already noticed; and they differ chiefly in regard to the destination of their products. The first group of this class deserving a separate notice, is that which effects the introduction of aliment into the body ; of those kinds of aliment, at least, which are not received in solu- tion by any more direct means. Along the greater part of the intestinal tube, from the point at which the hepatic and pancreatic ducts enter it, to the rectum, we find the mucous membrane furnished with a vast num- ber of minute tufts or folds, by which its free surface is vastly extended; these are termed villi. They may be compared to the ultimate root- fibres of trees, both in structure and function; for each of them gives origin to a minute lacteal or chyle-absorbing vessel, which occupies its centre ; whilst it also contains a copious network of blood-vessels (Fig. 10, p. 127), which appears likewise to participate in the act of absorp- tion, by taking up substances that are in complete solution. Now at the end of every villus, there may be seen, whilst the process of digestion and absorption is going on, a cluster of minute opalescent globules, in the midst of which the origin of the lacteal is lost. These globules, whose size varies from l-1000th to l-2000th of an inch, are composed of a milky fluid, which is evidently the same with that which is found in 152 STRUCTURE AND ENDOWMENTS OF ANIMAL TISSUES. the lacteals ; and it is maintained by Prof. Goodsir, Avho first brought them into notice, that these globules are really cells, and that it is by Fig. 31. CO -f Diagram of mucous membrane of Jejunum, when absorption is not going on; a, epithelium of a villus; b} secreting epithelium of a follicle ; c, c, c, primary membrane, with its germinal spots of nuclei, d, d; e, genua of absorbent vesicles; /, vessels and lacteals of villus. their growth and nutrition that the milky fluid, or chyle, is selected from the contents of the digestive cavity. Their function, therefore, would be precisely the converse of that of the secreting cells already described ; whilst the history of their individual lives is the same. These absorbent cells draw their materials from the fluid in the digestive cavity, instead of from the blood; and Avhen they burst or liquefy, they set free their contents where they may be taken up by a lacteal and conveyed into the circulating current, instead of pouring them into a cavity through which they will be shortly expelled. In the intervals of the digestive process, however, the extremities of the villi are comparatively flaccid; and instead of cells, they shoAV merely a collection of granular particles (Fig. 31, e), which are considered by Prof. Goodsir to be cell-germs. There is considerable doubt, however, whether these supposed cells are anything else than oil-globules; and Avhether the real agents in the selection of chyle are not the epithelium-cells covering the villus (a), within which chylous-looking globules have been occasionally seen, when digestion was going on. 244. Although the Mucous membrane of the intestinal tube is the only channel through Avhich insoluble nutriment can be absorbed in the completely formed Mammal, and the only situation, therefore, in which we meet with these absorbent cells, there are other situations in Avhich similar cells perform analogous duties in the embryo. Thus the Chick derives its nutriment, whilst in the egg, from the substance of the yolk, by absorption through the blood-vessels spread out in the vascular layer of the germinal membrane surrounding the yolk ; Avhich vessels answer to the blood-vessels and lacteals of the permanent digestive cavity, and are raised into folds or villi as the contents of the yolk-bag are diminished. Now the ends of the vessels are separated from the fluid contents of the yolk-bag, by a layer of cells ; Avhich seems to have for its object to select and prepare the materials supplied by the yolk, for being received into the absorbent vessels. 245. In like manner, the embryo of the Mammal is nourished, up to ABSORBENT CELLS. 153 the time of its birth, through the medium of its umbilical vessels ; the ramifications of Avhich form tufts, that dip down, as it were, into the maternal blood, and receive from it the materials destined to the nutri- tion of the foetus, besides effecting the aeration of the blood of the latter, by exposing it to the more oxygenated blood of the mother. Now around the capillary loop of the foetal tuft, there is a layer of cells, closely re- sembling the absorbent cells of the villi; and these are enclosed in a cap of basement-membrane, Avhich completes the foetal portion of the tuft, and renders it comparable in all essential respects to the intestinal villus. It is again surrounded, however, by another layer of membrane and of cells, belonging to the maternal system;—the derivation and arrange- ment of which will be explained hereafter. The maternal cells (b, Fig. 32), may be regarded as the first selectors of nutriment from the circu- Fig. 32. Extremity of a placental villus:—a, external membrane of the villus, continuous with the lining mem- brane of the vascular system of the mother; 6, external cells of the villus, belonging to the placental decidua; c, c, germinal centres of the external cells; d, the space between the maternal and foetal portions of the villus; e, the internal membrane of the villus, continuous with the external membrane of the chorion; /, the internal cells of the villus, belonging to the chorion ; g, the loop of umbilical vessels. lating fluid of the parent: the materials, partially prepared by them, are poured into the cavity (d) surrounding the extremity of the tuft; and from this they are taken up by the foetal cells (/), Avhich further elabo- rate them, and impart them to the capillary loop (g) of the umbilical vessels. 24G. Thus we see that the several functions of Selection, Absorption, Assimilation, Respiration, Secretion, and Reproduction, are performed by the agency of cells in the Animal as in the Vegetable kingdom,—in the complex Human organism, as in the humblest Cryptogamic Plant: the only difference being, that in the latter there is a greater division of labor, different groups of cells being appropriated to different functions, in the general economy, whilst the history of their own processes of nutrition and decay is everywhere essentially the same. Thus we have seen that the Absorbent cells, at the extremities of the intestinal or pla- cental villi, select and draw into themselves, as the materials of their OAvn groAvth, certain substances in their neighborhood; which are still as much external to the tissues of the body, as are the fluids surrounding the roots of plants. Having come to their full term of life, they give up their contents to the absorbent vessels, which carry them into the general current of the circulation, where they are mingled with the fluid previously assimilated,—the blood. Whilst passing through the vessels, they are subjected to the action of the various cells (all of Avhich we have seen to be successive phases of the same type) which float in the circu- lating current; and by these they seem to be gradually assimilated, or 154 STRUCTURE AND ENDOAVMENTS OF ANIMAL TISSUES. converted into a substance of a more directly organizable character. The special function of the red corpuscles peculiar to Vertebrated animals, though not yet accurately knoAvn, seems intimately connected with the process of Respiration. Next we have various groups of cells, external to the vessels, on the free surfaces of the body ; whose office it is to draAV from the blood certain materials, which are destined for Secretion or separation from it; either for the sake of preserving that fluid in its re- quisite purity, or for answering some other purpose in the system. These cells grow at the expense of the substances which they draAY into themselves from the blood ; and on their dissolution, they cast forth their contents on the free surfaces communicating Avith the exterior of the body, to which they are in time conveyed. And, lastly, we have a special set of Generative cells, destined in the one sex to prepare the germs of neAV beings; and in the other to elaborate a product essential to their fertilization. 247. The cells which are thus the active instruments of the Organic functions, are usually produced and succeed one another with a rapidity proportional to the energy of those functions, though the causes Avhich influence their growth and decay are not always evident. Thus it is certain that, cceteris paribus, the rate of production of the Secreting cells depends upon the abundance of the materials supplied by the circu- lating current, which they are destined to eliminate from it. But this is by no means the sole condition of their development; for, as Ave shall see hereafter, these materials may accumulate unduly in the blood, through the insufficient activity of the cells Avhich are destined to sepa- rate them; whilst, on the other hand, the presence of certain substances in the blood appears to accelerate their production. Of these stimuli, Mercury is one of the most powerful; and Ave have continual opportu- nities of witnessing its effects, in giving an increased activity to the secreting actions. There is probably not a gland in the body, which is not in some degree influenced by its presence in the blood ; but the liver, the kidneys, the salivary glands, and the glandulse of the intestinal canal, appear to be those most affected by its stimulating poAvers. The action of the glands, in other words the development of the secreting cells, appears to be influenced by mental emotions ; being sometimes accele- rated, and sometimes retarded, through their agency. This is especially the case in regard to the secretion of Milk, Tears, Saliva, and Gastric juice. It seems probable that the influence thus manifested is partly exerted through the capillary circulation, Avhich is knoAvn to be powerfully affected by mental emotions, as in the acts of blushing and erection ; and that the increased production of the secretion is immediately due to the increased flow of blood to the gland. But there are other phenomena Avhich show that the development and actions of the secreting cells are more directly influenced by the nervous system ; these will be hereafter considered (chap. IX.) 5. Of Cells connected together as permanent constituents of the Tissues. 248. We now pass on to consider those Cells, which enter as compo- nent elements into the solid and permanent fabric of the body, and which CELLS CONNECTED TOGETHER IN SOLID TISSUES. 155 do not take so active a part in its vital operations. These we shall find to be usually more or less closely connected together, either by a general enveloping membrane, or by an intercellular substance, which is inter- posed between their walls, and holds them together by its adhesive properties. 219. The presence of a general enveloping membrane (where it is not a secondary formation) appears to depend upon the persistence of the original cell-walls ; which, instead of liquefying or thinning away, when distended by the multiplication of cells in their interior, are thickened or strengthened by additional nutrition. Such is perhaps the case with the sacculi in which the cells of Adipose tissue (§ 257) are often found clustered together; but this condition is usually much more obvious in many tumors, whose development depends upon an abnormal process of growth. 250. Where such enveloping membranes are wanting, we frequently find the component cells of the permanent tissues of Animals (like those of the higher plants) held together by an intercellular substance; which generally presents no distinct traces of organization; and which usually consists of Gelatine, or of a substance allied to it in composition. The proportion of this substance to the cells may vary in different cases; and very different characters may thus be presented by a tissue made up of the same elements. Thus the subjoined figure (33) represents a portion of one of the animal layers included betAveen the calcareous laminae of a bivalve shell: in Avhich we see on the one side a number of nuclei or incipient cells, scattered through a bed of homogeneous intercellular sub- stance, and bearing but a very small proportion to it; whilst the opposite end exhibits a set of polygonal cells, in close contact with each other, the intercellular substance being only represented by the thick dark lines, Avhich mark the boundaries of the cells, and which are rather thicker at the angles of the latter. BetAveen these tAvo extremes, we observe every stage of transition. 251. The presence of a very large amount of intercellular substance, through which minute cells are scattered at considerable interA'als (Fig. 33, a), is characteristic of various forms of Cartilage ; and more par- ticularly of that soft semi-cartilaginous structure, of which the Jelly-fish are for the most part composed. In other forms of cartilage, Ave find the cells more developed, and in closer proximity to each other, the proportion of the intercellular substance being at the same time dimi- nished (as seen at b and c, Fig. 33); but it is not often, save in the embryonic structures, that Ave find the cells in such close proximity, and the intercellular substance so nearly Avanting, as at d. Such examples do occasionally present themselves, hoAvever, even in the soft tissues. Thus the chorda dorsalis, Avhich replaces the vertebral column in the lowest Fishes, and of which the analogue is found in the embryos of the higher Vertebrata, is made up of a structure of this kind (Fig. 16). The true Skin in the Short Sun-fish, is replaced by a similar layer of cellular tissue, Avhich extends over the whole body, varying in thickness from one-fourth of an inch to six inches. And in the Lancelet (a little fish which is destitute of so many of the characters of a Vertebrated animal, that its right to a place in that division has been doubted), a considerable portion of the fabric is made up of a similar parenchyma. 156*- STRUCTURE AND ENDOAVMENTS OF ANIMAL TISSUES. 252. Now Ave shall find that one method, by which the requisite firm- ness and solidity are given to the animal fabric, consists in the depo- Fig. 33. Portion of shell-membrane, showing the origin of cells in the midst of horny intercellular substance; a, nuclei; b, incipient cells; c, the same further advanced, but separated by intercellular substance; d, the cells become polygonal by mutual pressure. sition of earthy substances in the interior of such cells, by a peculiar secreting action of their own. Thus in Shell, we find them completely filled up with carbonate of lime ; and in the enamel of Teeth with phos- phate of lime. When this is the case, there is a tendency to an apparent coalescence of the cells, by the obliteration of their partitions ; or rather, perhaps, by the removal of the whole intercellular substance from between them, the actual cell-walls being so very thin, that they are not distin- guishable. The incipient stages of this coalescence, as seen in another portion of the same membrane as that represented in the last figure, are shown in Fig. 34. At a, the nucleated cells are very distinct; and are separated by a large quantity of intercellular substance. At b, they approach each other more closely, the amount of intercellular substance being less ; the Avidest intervals are seen at the angles of the cells. At c, the approximation is much closer; and the cell-Avails are scarcely dis- tinguishable at the points Avhere they come into immediate contact. Proceeding further, we observe that the partitions are much less com- plete ; so that the originally distinct cellular character of the membrane Portion of shell-membrane, showing the gradual coalescence of distinct cells; at a, the cells separated by intercellular substance; at b, the partitions are thinner; and at c, they almost disappear. is chiefly indicated by the bright nuclei, which are regularly dispersed through it, and by the triangular dark spots, which show the remains of FUSIFORM CELLS. 157 the intercellular substance at the angles Avhere three cells join each other. The coalescence may be traced further than it is shown to do in the figure; so that if it were not for the evidence afforded by the transition-stages here represented, it would be difficult to prove that the membranous layer had its origin in cells. 253. These facts, respecting the gradual coalescence of cells, explain not merely certain appearances presented in Tooth, Shell, &c (here- after to be described); but also those which are exhibited by the Base- ment-membrane, as already detailed (§ 206). 254. There is no evidence, in the preceding case, that the cavities of . the cells coalesce ; and there is no reason Avhy they should do so. But we often find such a union, Avhere the production of a continuous tube is required. The long straight open ducts, through wThich the sap of Plants rises in the stem, are unquestionably formed by a coalescence of the cavities of cells of a cylindrical form, placed regularly end to end; and it seems probable that the netAvork of anastomosing vessels, through which the elaborated sap finds its way to the various parts of the vege- table fabric, is formed, in like manner, by the coalescence of cells, arranged obliquely and transversely in regard to one another. In like manner, the capillary Blood-vessels of Animals are usually believed to originate in toavs of cells, the cavities of which have run together by the obliteration of the transverse partitions ; as the persistent nuclei of such cells may be occasionally brought into vieAV in the walls of the ca- pillaries. And the same appears to be the origin of the tubular fibres of Muscular and Nervous tissue, which contain the elements characteristic of those tissues; these elements,—the fibrillse of muscle and the granular pith of the nerve-tube,—being evidently the secondary products of parent- cells, which seem to remain as their investing tubuli, in the Avails of which the original nuclei are often to be seen (§ 338 and 388). 255. Besides these changes, the original cells may often undergo marked alterations of form; and this quite independently of any pressure to which they may be subject. Thus the pigment-cells, as already men- tioned (§ 229), frequently exhibit a curious stellate form; arising from the development of radiating prolongations, which are put forth from the original spheroid. A form which is frequently assumed by the cells that are developed in fibrinous or plastic exudations, and which is also met Avith in the cells of tumors, both malignant (or Cancerous) and non-malignant, is that which has received the designation of fusiform or spindle-like, from its prolonged shape and pointed extremities. The various stages of transition, which may be observed betAveen the simple rounded cell and the fusiform cell, have been shoAvn in Fig. 7; and it is there seen that, Avhen the transformation has gone to its utmost extent, the nucleus of the cell is no longer visible, so that it bears a close re- semblance to a simple fibre. Such cells are found amongst the simple fibrous tissues, and, in the opinion of many, they give origin to them. The appearance of tissue composed of fusiform cells, is shown in Fig. 35; this is seldom met with as a permanent part of the normal fabric; but it is a frequent product of morbid action. 256. We now proceed with the description of the various tissues in the Human body, which are composed of cells united or transformed 158 STRUCTURE AND ENDOWMENTS OF ANIMAL TISSUES. in the foregoing manner; and we shall commence Avith Adipose or Fatty tissue, which may be considered as a sort of link, connecting the permanent tissues with those Avhich are more actively concerned in the processes of Nutrition, Secretion, &c. Fig. 35. Fusiform tissue of plastic exudations; a, fusiform bo- dies without nuclei; b, nu- cleated fusiform cells; c, granular intercellular sub- stance. 257. The Adipose tissue is composed of isolated cells, which have the power of appropriating fatty matter from the blood, precisely in the same manner as the secreting cells appropriate the elements of bile, milk, &c. These cells are sometimes dispersed in the interspaces of the Areolar tissue; whilst in other cases they are aggregated in distinct masses,— constituting the proper Adipose tissue. In the former case they are held in their places by fibres, that traverse the areolae in different direc- tions; whilst in the latter, each small cluster of fat-cells is included in a common envelope, on the exterior of which the blood-vessels ramify; and these sacculi are held together by areolar tissue. We are thus probably to regard each fatty mass in the light of a gland, or assemblage of secreting cells, penetrated by blood-vessels, and bound together by fibrous tissue; but having its follicles closed instead of open (which appears to be the early condition of the follicles of all glands, § 238): and consequently retaining its secretion within itself, instead of pouring it forth into a channel for excretion. Fig. 37. Capillary network around Fat-cells. 258. The individual fat-cells always present a nearly spherical or sphc< roidal form ; sometimes, however, when they are closely pressed together, Areolar and Adipose tissue ; a, a, fat-cells; 6, b, fibres of areolar tissues. ADIPOSE TISSUE. 159 they become somewhat polyhedral, from the flattening of their walls against each other. Their intervals are traversed by a minute network of blood-vessels (Fig. 37), from which they derive their secretion; and it is probably by the constant moistening of their walls with a watery fluid, that their contents are retained without the least transudation, although they are quite fluid at the temperature of the living body. If the watery fluid of the cell-walls of a mass of Fat be allowed to dry up, and it be kept at a temperature of 100°, the escape of the contained oily matter is soon perceptible.—By this provision, the fatty matter is alto- gether prevented from escaping from the cells of the living tissues, by gravitation or pressure; and as it is not itself liable to undergo change Avhen secluded from the air, it may remain stored up, apparently unal- tered, for almost an unlimited period. 259. The consistency, as well as the Chemical constitution, of the fatty matter contained in the Adipose cells, varies in different animals, according to the relative proportions of three component substances Avhich may be distinguished in it,—Stearine, Margarine, and Oleine. The two former are solid when isolated, and the latter is fluid ; but at the ordinary temperature of the warm-blooded animal, they are dissolved in it. Of these, Stearine is the most solid; and it is the most largely present, therefore, in the hardest fatty matter, such as mutton-suet. It is crystalline like spermaceti; it is not at all greasy betAveen the fingers, and it melts at 143°. It is insoluble in Avater, and in cold alcohol and ether: but it dissolves in boiling alcohol or ether, crystallizing as it cools. The substance termed Margarine exists along with stearine in most fats, but it is the principal solid constituent of Human fat, and also of Olive oil. It corresponds with Stearine in many of its proper- ties, and is nearly allied to it in Chemical composition ; but it is much more soluble in alcohol and ether, and it melts at 118°. On the other hand, Oleine, when pure, remains fluid at the zero of Fahrenheit's ther- mometer ; and it is soluble in cold ether, from which it can only be separated by the evaporation of the latter. It exists in small quantity in the various solid fats ; but it constitutes the great mass of the liquid fixed oils. The tendency of these to solidification by cold, depends upon the proportion of stearine or margarine they may contain. 260. All these substances are neutral compounds formed by the union of Stearic, Margaric, and Oleic acids, respectively, with a base termed Glycerine; this base may be obtained from any fatty matter, by treat- ing it with an alkali, which unites with the acid and forms a soap, setting free the Glycerine. They contain no Nitrogen ; and their pro- portion of Oxygen is extremely small in comparison with their amount of Carbon and Hydrogen: thus Stearine has 142 Carbon and 141 Hy- drogen to 17 Oxygen: and in the other substances the proportions are similar. The fatty bodies appear to be mutually convertible; thus mar- garic acid may be procured from stearic acid, by subjecting it to dry distillation ; and there is ample evidence that animals supplied with one of them may produce the others from it. 261. Since these Fatty matters are abundantly supplied by the Vege- table kingdom, and are found to exist largely in substances which were not previously supposed to contain them, it is not requisite to suppose, 160 STRUCTURE AND ENDOAVMENTS OF ANIMAL TISSUES. that Animals usually elaborate them by any transforming process from the elements of their ordinary food. The mode in Avhich they are taken into the blood, and the uses to Avhich they are subservient, will be here- after investigated: but it may be here remarked, that the portion sepa- rated from the circulating fluid to form the Adipose tissue, is only that Avhich can be spared from the other purposes, to which the fatty matters have to be applied. Hence the production of this tissue depends in part upon the amount of Fatty matter taken in as food; but this is not en- tirely the case, as some have maintained; for there is sufficient evidence that animals may produce fatty matter by a process of chemical trans- formation, from the starch or sugar of their food, Avhen there is an un- usual deficiency of it in their aliment. 262. The development of Adipose tissue in the body appears to an- swer several distinct purposes. It fills up interstices, and forms a kind of pad or cushion for the support of movable parts; and so necessary does it seem for this purpose, that even in cases of great emaciation, some fat is ahvays found to remain, especially at the base of the heart around the origin of the great vessels, and in the orbit of the eye. It also assists in the retention of the animal temperature by its non-con- ducting power; and Ave accordingly find a thick layer of it, in those warm-blooded mammals that inhabit the seas,—either immediately be- neath their skin, or incorporated with its substance. Its most important use, however, is to serve as a reservoir of combustible matter, at the expense of which the respiration may be maintained Avhen other mate- rials are deficient; thus Ave find that the respiration of hibernating animals is kept up, during the period when they cease taking food (§ 121), by the consumption of the store of fat which was laid up in their bodies, previously to their passing into that state; and it is also to be noticed that herbivorous animals, whose food is scanty during the winter, usually exhibit a strong tendency to such an accumulation, during the latter part of the summer, when their food is most rich and abundant, in order to supply the increased demand created by the low external temperature of the Avinter season. Other circumstances being the same, it appears that the length of time during which a warm-blooded animal can live without food, depends upon the quantity of fat in its body; for the rapid lowering of its temperature, which is the immediate cause of its death (§ 117), takes place as soon as the whole of this store has been exhausted. Of the means by which the fatty secretion is taken back again into the current of the circulation, Avhen it is required for use in the system, we know nothing whatever. 263. In order that it may be applied to the maintenance of the ani- mal heat, the fatty matter must be received back into the blood; and although Ave have no certain knowledge of the mode in which this is accomplished, yet it may be surmised to be as folloAvs. The Blood nor- mally contains a certain amount of fatty matter, held in solution by combination with its alkali; and should this be exhausted by the com- bustive process, the circulating current will draw into itself a fresh sup- ply from the interior of the fat-cells;—it having been shown by Mat- teucci that oleaginous particles will pass through animal membranes by CARTILAGE. 161 endosmose, to diffuse themselves through an aqueous liquid, provided the latter be alkaline. 264. In the simpler forms of Cartilage, we have an example of a tissue of remarkable permanence, composed entirely of cells scattered through an intercellular substance. This substance has received the distin- guishing appellation of Chondrine, which marks it as the solidifying ingredient of Cartilage (§177). All the Cartilages of the foetus,—those which are to be converted into bone, as well as those which are to remain unossified,—are composed of it; and yet, as soon as the process of Ossification commences, the chondrine is replaced by Gelatine, which is the sole organic constituent of the animal basis of bones. The per- manent cartilages, however, still contain only Chondrine; but if acci- dental bony deposits should take place in them (as frequently happens in old persons, especially in the cartilages of the ribs), the Chondrine gives place to Gelatine. This change of composition is coincident, as we shall hereafter see, with a complete change in texture; the basis of ■bony tissue not being Cartilage (as commonly imagined), but consisting of a substance much more nearly allied to the white fibrous tissue.—It is only in the pure cellular cartilages, in which the intercellular sub- stance presents no trace of organization, that Chondrine occurs. Those of the/fo-o-cartilages (§ 269), in which the intercellular substance has the characters of the White fibrous tissue, yield gelatine on boiling, in the manner of the ligaments and tendons; whilst those which contain much of the Yellow or elastic tissue, undergo very little change by boil- ing, and only yield, after several days, a small quantity of an extract which does not form a jelly, but which has the other chemical properties of Chondrine. 265. Besides the organic compounds already described, most Carti- lages contain a certain amount of mineral matter, which forms an ash when they are calcined. This ash contains a large proportion of car- bonate and sulphate of soda, together with carbonate of lime, and a small quantity of phosphate of lime; as age advances, the proportion of the soluble compounds diminishes, and the phosphate of lime predo- minates. This is especially the case in the costal cartilages, which almost invariably become converted into a semi-ossified substance, in old persons ; and it is remarkable that, even before they have themselves become thus condensed, they are united by ossific matter, when they have undergone fracture. Fig. 38. Cartilage of Mouse's ear. 266. When a pure Cellular Cartilage is examined microscopically its cells are seen to lie, sometimes singly, and sometimes in clusters of 162 STRUCTURE AND ENDOWMENTS OF ANIMAL TISSUES. two, three, or four, in cavities excavated in the intercellular substance. These occur at very variable distances; for in some instances they are packed together as closely as the cells of a vegetable parenchyma (Fig. 38); whilst in others the principal mass is composed of intercellular substance, through which the cells are interspersed at Avide intervals. From the various appearances Avhich may be observed in the same car- tilage at different stages of its groAvth, it would appear that the compo- nent cells multiply by the doubling process already described (§ 212); that they then separate from one another, each of them drawing toAvards itself (as it were) an envelope of intercellular substance; and that, by the repetition of the same process, the number of cells in the cartilage may be indefinitely multiplied. 267. Various stages of this history are shown in the accompanying figure (Fig. 39), which is taken from a section of the cartilaginous bran- chial ray of the larva or tadpole of the Rana esculenta, or Edible Frog. In the centre of the figure are shown three separate cellst which have evidently been at one time in closer proximity with each other. In one of these cells, the nucleus is seen to be developing two new cells in its interior; and a continuation of this process would give rise to the appear- ance shown at b, Avhere two cells are shown in close contact, being evi- dently the offspring of the same parent. Now if each of these cells in like manner developes tAvo others within itself, a cluster of four will be developed, as shown at a; and after a time, intercellular substance being Fig. 39. Section of the Branchial cartilage of Tadpole; a, group of four cells, separating from each other; 6, pair of cells in apposition; c, c, nuclei of cartilage cells; d, cavity containing three cells. accumulated around each, their walls will separate, and they will acquire the character of distinct cells. It would seem as if, in other cases, one of the first pair of cells developes another pair in its interior, whilst the other (from some unknown cause), does not at once proceed to do so; and thus only three cartilage-cells instead of four are clustered together in the cavity, as seen at d. 268. The primitive cellular organization now described is retained in some Cartilages through the whole duration of their existence. This is the case, for example, in most of the articular cartilages of joints; in the cartilaginous portion of the septum narium, in the cartilages of the alae and point of the nose, in the semilunar cartilages of the eyelids; in the cartilages of the larynx (with the exception of the epiglottis), the CARTILAGE. 163 cartilages of the trachea and bronchial tubes, the cartilages of the ribs, and the ensiform cartilage of the sternum. When partial ossific depo- sits take place, it is usually in the substance of cellular, rather than in that of fibrous cartilage. 269. When the intercellular substance, instead of being homogeneous, has a fibrous character, the tissue called Fibro-Cartilage is produced; and this may be either elastic or non-elastic, according as the yellow or the white form of fibrous structure prevails. In some instances, the fibrous structure is so predominant over the cellular, that the tissue has rather the character of a ligament than of a cartilage. The white fibrous structure is seen in all those cartilages, Avhich unite the bones by synchondrosis, and which are destined not merely to sustain pressure, but also to resist tension. This is the case especially in the substances which intervene betAveen the vertebrae, and which connect the bones of the pelvis ; these in adult Man are destitute of cartilage-corpuscles, ex- cept in and pear their centres; but in the lower Vertebrata, and in the early condition of the higher, the fibrous structure is confined to the exterior, and the Avhole interior is occupied by the ordinary cartilage- corpuscles. The yellow-fibrous or reticulated structure is best seen in the epiglottis, and in the concha of the ear; in the former of these, scarcely any trace of cartilage corpuscles remains ; and in the latter, the cellular structure is only to be met with towards the tip. 270. We have seen that the elements of the cellular tissues hitherto described, do not come into direct contact with the blood-vessels. The Epidermic and Epithelial cells are separated from them by the conti- nuous layer of basement-membrane, which forms the surface of the true skin, of the mucous membranes, of the glandular follicles produced from them, &c. In like manner, the cells of Adipose tissue are formed within membranous bags; around which the blood-vessels form a minute network. The cells of Cartilage are not nourished in any more direct manner; and are sometimes at a considerable distance from Fig. 40. Vessels situated between the attached synovial membrane, and the articular cartilage, at the point where the ligamentum teres is inserted in the head of the os femoris of the human subject, between the third and fourth months of foetal life;—a. the surface of the articular cartilage; b, the vessels between the articular cartilage and the synovial membrane ; c, the surface to which the ligamentum teres was attached ; d, the vein; e, the artery. the nearest vessels. It is certain that the substance of the permanent cellular Cartilage is not permeated, in a state of health, eAren by the minutest nutrient vessels; none such being brought into view under the 164 STRUCTURE AND ENDOAVMENTS OF ANIMAL TISSUES. highest magnifying power. They are, however, surrounded by vessels (Fig. 40), which form large ampullae or varicose dilatations at their edges, or spread over their surfaces; and it is by the fluid which is drawn from them by the Cartilage-cells, that the latter are nourished. The nutrition of a mass of Cartilage thus seems to bear a strong resem- blance to that of the thick fleshy Sea-weeds, which are in like manner composed entirely of cells, with intercellular substance disposed between them in greater or smaller amount. The cells in nearest proximity to the nutrient fluid, draw from it the requisite materials, and transmit these to the cells in the interior of the mass, receiving a fresh supply in their turn from the source in their OAvn neighborhood. When the Ar- ticular or other cellular Cartilages are inflamed, however, we find ves- sels passing into their substance; but these vessels are formed in an entirely neAV tissue, Avhich is the product of the inflammatory process, and cannot be said to belong to the Cartilage itself. 271. The temporary Cartilages, which have a like cellular structure, but which are destined to undergo metamorphosis into Bone, are equally destitute of vessels when their mass is small; but if their thickness ex- ceed an eighth of an inch, they are permeated by canals for the trans- mission of vessels. Still these vessels do not ramify with any minute- ness in the tissue; and they leave large islets, in which the nutritive process must take place on the plan just described. 272. The Fibro-Cartilages, formed as it were by the intermingling of two distinct elementary structures, have a degree of vascularity propor- tioned to the amount of the fibrous tissue which they contain ; but these vessels do not penetrate the cellular portions, where such are distinct from the mixed structure. 273. The Cartilaginous tissue appears to be more removed than almost any other in the body from the general tide of nutritive action. Its properties are simply of a physical character ; and they are not im- paired for a long time after the death of the tissue, its tendency to de- composition being very slight, so long as it is exposed to ordinary tem- peratures. It is protected by its toughness and elasticity from those mechanical injuries to Avhich softer or more brittle tissues are liable; and consequently it has little need of any active power of reparation. When loss of substance occurs as a result of disease or accident, this seems never to be repaired by real cartilaginous substance; but the space is filled up by a fibrous tissue developed from the reparative blastema (§ 213). It is in this tissue that the new vessels are found, which have been erro- neously- supposed to penetrate the cartilage when it becomes inflamed; the fact being, that the vessels are restricted to the "false membrane" formed in the inflammatory process, which takes the place of the carti- laginous tissue that has disappeared in consequence of imperfect nutri- tion or degeneration. 274. The Cornea of the Eye bears a superficial resemblance to Cartilage; but it corresponds rather with Fibrous Membranes in its elementary structure. Besides its anterior or conjunctival layer, which consists of epithelium, and its posterior layer of cells constituting the epithelium of the aqueous humor, the Cornea has been shown by Mr. Bowman to consist of three layers, of which the anterior and the poste- rior (which are very thin) have some of the characters of the yellow CELLS CONNECTED TOGETHER.—CORNEA. 165 elastic tissue, whilst the middle one, which forms its principal thick- ness, is composed of white fibres interlaced together in such a manner Fie. 41. Nutrient Vessels of the Cornea;—a, Superficial vessels belonging to the Conjunctival membrane and continued over the margin of the Cornea; b, Vessels of the Sclerotic, returning at the mIrgi™onhe Cornea as to form numerous lamellae, their interspaces or areolae having the form of tubes regularly arranged and constructed at intervals, so as not to be unlike rows of cartilage-cells, for Avhich in fact they have been mistaken.—Two sets of vessels, a superficial and a deep-seated, surround the margin of the cornea. The former (Fig. 41 a), belong rather to the Conjunctival membrane, which forms the outer layer of the cornea • and they are prolonged to the distance of l-8th or half a line from its margin, then returning as veins. The latter (b) do not pass into the true Cornea, but terminate in dilatations from which veins arise, just where it becomes continuous with the sclerotic. In diseased conditions of the Cornea, hoAvever, both sets of vessels extend themselves through it. Notwithstanding the absence of vessels in the healthy aondition of the corneal tissue, incised wounds of its substance commonly heal very readily, as is well seen after the operation for Cataract; but there is a danger in carrying the incision around a large proportion of its margin, lest the tissue should be too much cut off from the supply of nutriment afforded by the ampullae of the vessels that surround it. 275. The Crystalline Lens of the Eye approaches Cartilage, in its structure and mode of nutrition, more nearly than any other tissue. It may be separated into numerous laminae; which are composed of fibres that lock into one another, by their delicately-toothed margins. Each of these fibres appears to be made up of a series of cells, linearly arranged, which coalesce at an early period. The lens is not per- meated by blood-vessels ; at least after it has been completely formed • these being confined to the capsule. During the early part of foetal life, and in inflammatory conditions of the Capsular membrane, both its anterior and its posterior portions are distinctly vascular; but at a later period, only the posterior half of the Capsule has vessels distri- buted upon its surface. It has been shown by optical • experiments 166 STRUCTURE AND ENDOWMENTS OF ANIMAL TISSUES. devised for the purpose, that a moderate vascularity of the posterior capsule does not interfere with distinct vision; whilst if the anterior capsule were traversed by vessels, the picture on the retina Avould be no longer clear.—The substance of which the lens is composed, ap- pears to be soluble Albumen, or perhaps more closely resembles the Globulin of the blood. 276. The Vitreous body, which fills the greater part of the globe of the eye, also seems to possess a cellular structure; the cells containing a fluid, which is little else than water holding in solution a small quantity of albumen and saline matter; and the membrane which forms their walls being so pellucid as to be scarcely distinguishable. Indeed the cellular character of this substance is chiefly inferred from the fact, that when its capsule or enveloping membrane is punctured, even in several places, the contained fluid does not speedily drain away,—as it would do if it were merely contained in the interstices of an areolar tissue. The blood-vessels Avhich traverse the Vitreous body do not send branches into its substance ; and it must derive its nutriment from those which are distributed minutely upon its general envelope, and probably also from the large plexiform vessels of the ciliary processes of the Choroid coat. 277. Before proceeding to describe the structure of Bone, to which it seems natural to pass on from Cartilage, it Avill be useful to advert to the modes in which the tissues of Invertebrated animals are consolidated by deposits of solid matter, in order that they may afford the requisite support and protection, without that interstitial growth which is pecu- liar to the skeletons of the Vertebrated classes.—Commencing with the Polypifera, or Coral-forming animals, we observe that their strong axes or sheaths are destined only to give support to their softer structures, and that the parts once consolidated undergo no subsequent change. It was formerly imagined, that the stony Corals were " built up" by the animals which form them, somewhat in the same manner as a Bee con- structs its cell. But it is now fully demonstrated, that the calcareous matter (which here consists solely of the Carbonate of Lime) is combined with the living tissue ; and that the most solid mass of Coral thus has an organized basis. The proportion of earthy to animal matter, how- ever, is so great in these structures, that very little, if any, nutrient changes can take 'place in their tissues, when once it has become con- solidated. Such changes are not, however, required. The substance thus developed by the attractive power of the soft gelatinous tissues, which draw into themselves the small quantity of calcareous matter dis- solved in the surrounding water, is so little disposed to undergo change, that it will maintain its solidity for centuries^ and even when acted on by water or by heat, it does not undergo disintegration, for its calcareous particles arrange themselves in a neAV method, and become converted into a solid crystalline rock. Such rocks, the product of the metamor- phosis of ancient coral-formations, make up a large proportion of the external crust of the earth. The solid stem or sheath, once con- solidated, appears to undergo no further change in the living Coral- structure ; for its increase takes place, not by interstitial but by super- ficial deposit,—that is, not by the diffusion of new matter through its SHELLS OF MOLLUSCA. 167 whole substance, separating from each other the parts formerly de- posited but by the mere addition of particles to its surface and extre- mities. In this manner the growth of a solid Coral-structure may go to an enormous extent; the surface at which the consolidating action is going on, being the only part alive, that is exhibiting any vital change; and all the rest of the mass being henceforth perfectly inert. 278. In the class of Echinodermata, which includes the Star-fish, Sea-Urchin, &c, we find the calcareous structure presenting a very elaborate organization ; as an example of this, we shall select the shell of the Echinus, commonly known as the Sea-Egg. This shell is made up of a number of plates, more or less regularly hexagonal, and fitted together so as completely to enclose the animal, except at two points one of which is left open for the mouth, the other for the anus. On the surface of these plates are little tubercles, for the articulation of the spines, which serve as instruments of defence and of locomotion. The substance of the shell and of the spines is exactly alike ; being a sort of areolar tissue, consolidated by the deposition of calcareous matter, and having an innumerable number of interspaces or minute cancelli, freely communicating with each other. The arrangement of this calcareous netAvork in the spines is most varied and elaborate; and causes thin sections of them to be among the most beautiful of all microscopic ob- jects. The external and internal surface of each plate, in the shell of the living Echinus, is covered with a membrane, from Avhich its nutrition Fig. 42. Portion of the shell of the Echinus, showing at a the constituent plates, and at 6 the calcified areolar tissue, of which they are composed. is derived; this membrane dips down into the spaces between the ad- jacent plates ; but it does not penetrate the substance of the plates themselves, nor does it transmit vessels to their interior. A simi- lar membrane covers and encircles the spines; and it also connects these with the shell, being continuous with the membrane that envelopes the latter. Thus each plate and spine is itself completely extra-vascu- lar ; but it is enclosed in a soft membrane, which furnishes (whether by vessels or otherwise, has not yet been ascertained), the elements of its nutrition. 279. But we do not here find any evidence of interstitial growth; nor is there any reason why such should be required. For the tissue 168 STRUCTURE AND ENDOWMENTS OF ANIMAL TISSUES. of which it is composed, although of such extreme delicacy, is of great permanence, and does not exhibit the slightest tendency to decay, how- ever long it is preserved; so that, when once consolidated, it appears to undergo no further change in the living animal. The growth of the animal, however, requires a corresponding enlargement of its enveloping shell; and this is provided for by the simple process of superficial de- posit, through the subdivision of the whole shell into component plates. For by the addition of new matter at the edge of each plate, by the consolidation of a portion of the soft membrane that intervenes between the adjacent plates, the whole shell is enlarged, without losing its globular form. At the same time it is strengthened in a corresponding degree, by the consolidation of the soft tissue at the surface of each plate. And, in like manner, the spines are enlarged and lengthened by the progressive formation of new layers, each on the exterior of the preceding; so that a transverse section exhibits a number of concentric rings, like those of an Exogenous tree.—Thus even in the growth of this complex and elaborate structure, we recognise the principle of superficial deposit, which we shall find to be universal amongst the hard parts of the Invertebrata : notwithstanding that, at first sight it would have appeared impossible to provide on this plan for the gradual en- largement of a globular shell, completely enclosing the animal, and therefore required to keep pace with the latter in its rate of increase. 280. Among the Mollusca, we find the body sometimes altogether destitute of solid organs of support, protection, or locomotion,—as is the case, for example, in the Slug; and the movements are feeble and the habits inert, the muscles having no fixed points for their attachment, and acting Avithout any of the advantages of leverage. In other cases, we find the body more or less completely protected by the Shell; which is sufficiently large in some instances to cover the body completely, whilst in others it affords only a partial investment. The plan on which this shell is formed, however, is very different from that which has just been described; being much less complex. The Univalve shells, or those formed in one piece are always of a conical form ; the cone being sometimes simple, as in the Limpet; in other cases being spirally coiled, as in the Snail. Now the base of this cone is open; and through this, the animal can project its movable parts. When its increasing size requires additional accommodation, it is obvious that an addition to the large end of the cone will increase its diameter and its length at the same time; so as to afford the required space, Avithout any alteration in the form or dimensions of the older and smaller portions of the cone. This last, indeed, is frequently quitted by the animal, and remains empty ; being sometimes separated from the latter portions, by one or more partitions thrown across by the animal,—as is seen especially in the Nautilus and other chambered shells. Besides the new matter added to the mouth of the shell, a thin layer is usually formed over its whole interior surface; so that the lining of the neAV part is continuous with that of the old.—In the Bivalve shells, Ave trace this mode of in- crease Avithout any difficulty; especially in such shells as that of the Oyster, in which the successive laminae remain distinct. Each lamina is interior to the preceding, being formed on the living surface of the SHELLS OF MOLLUSCA. 169 animal; but it also projects beyond it, so as to enlarge the capacity of the shell; and as the separation of the valves affords free exit to those parts of the animal, which are capable of being projected beyond the she , there is obviously no need of any other provision to maintain the shell in its natural form.—Thus in the shells of the Mollusca, increase takes place at the surfaces and edges only. 281. The proportion of organic and calcareous matter in Shell differs considerably in the various tribes. The former is sometimes present in such small amount, that it can scarcely be detected; and the condition ol the calcareous matter then obviously approaches that of a crystalline deposit. But in other instances, the animal basis is very obvious- remaining as a thick consistent membrane, after all the calcareous mat- ter has been dissolved away by an acid. This membrane is formed of an aggregation of cells arranged with great regularity (Fig. 43, a); the cavities of which are filled with carbonate of lime in a crystalline state. Fig. 43. ra#ia elfHi ill» ISPlpf mm Prismatic cellular structure of shell of Pinna :-a, surface of lamina; b, vertical section. i?de;rr!rr ?f ^.cfs aPF0AaAches the hexagonal; their diameter varies m different shells from l-100th to l-2800th of an inch ; their thickness also is extremely variable, even in different parts of the same shell. irr^l^nnT^68 .meft.WltM lamina 0f such tenuitJ> ™ ^ot to mea- sure l-100th of an inch in thickness, whilst in other instances, a single WIT7- AVV thldTSS °f half an inch> or even (^ certain lar°ge hp1ni?vC1fl8)*0f T !rC,h 0r, m0re' In this case> the cells> instead of being thin flat scales like the tessellated-epithelium (§ 233), are long prisms somewhat like the cells of the cylinder-epithelium (Fig. 26), with their wals flattened against each other. The appearance Avhich is then presented by a vertical section of them, is represented in Fig. 43, b. The long prismatic cells are there seen to be marked by delicate trans- verse striae, and these, taken in connexion with other indications, appear to show, that every such prism is in reality formed by the coalescence of a pile of flat cells resembling those which are seen in the very thin lamina* just described; so that the thickness of the layer depends upon the number of the cellular laminae which have coalesced to form its component prisms. This character is of interest, as representing on a magnified scale a corresponding appearance in the Enamel of human 170 STRUCTUKE AND ENDOWMENTS OF ANIMAL TISSUES. Teeth, which we shall presently find to be formed upon the very same plan (§ 318). 282. We are to regard this kind of shell-substance, therefore, as formed by the secreting action of the epithelial cells covering the mantle of the animal,—which membrane, though it answers in position to the skin, has the soft, spongy, glandular character of a mucous membrane. These draw calcareous matter into their cavities, as a part of their own process of growth: this matter being supplied from the fluids of the vascular surface beneath. Now when these calcigerous cells are sepa- rated by intercellular substance, they remain distinct through the whole of their lives, and they form by their cohesion a tenacious membrane, that retains its consistency after the removal of the calcareous matter. -But this is only the case in certain groups of shells, chiefly belonging to the bivalve division. When the intercellular substance is wanting, and the cells come into close contact, their partitions become indistinct on account of their extreme tenuity ; and not unfrequently a fusion of the whole substance appears to take place, by the dissolution of the original cell-walls, so that it becomes more or less homogenous,—traces of the original cellular structure being here and there distinguishable (§ 252). 283. Sometimes where this fusion has taken place, so as to obliterate the original cell-structure, Ave find the almost homogeneous substance traversed by a series of tubuli, not arranged, hoAvever, in any very defi- nite direction, but forming an irregular netAvork (Fig. 44). These tubes vary in size from l-2000th to l-20,000th of an inch; but their general diameter, in the shells in which they most abound, is l-4500th of an inch. In the larger tubuli, something of a bed-like structure may occasionally be seen : as if their interior were occupied by rounded granules arranged in a linear direction. Although it might be supposed that this structure is destined to convey nutrient fluid into the substance of the shell, yet there is no evidence that such is the fact; and, on the contrary, there is ample evidence that, even in shells most copiously traversed by these tubuli, no processes of interstitial growth or renewal take place. The permanent character of the substance of all Shells, when once it is fully formed, is as remarkable as that of Coral; and as the adaptation of their Fig. 44. Tuhular shell-structure, from Anomia. size, to that of the animals to which they belong, is entirely effected by additions to their surfaces and edges, no interstitial deposit can have a share in producing it. SHELLS OF CRUSTACEA. 171 284. Among the Articulated classes, we still find that the skeleton is altogether external, and belongs therefore to the cutaneous system ; but it is formed upon a very different plan from the shells of the Mollusca, being closely fitted to the body, and enveloping every part of it; conse- quently it must increase in capacity, with the advancing growth of the contained structures. Moreover it is destined not merely to afford sup- port and protection to these, but to serve for the attachment of the mus- cles by which the body and limbs are moved; and the hard envelopes of the latter serve, like the bones of the Vertebrata, as levers by which the motor poAvers of the muscles are more advantageously employed. Again, the hard envelopes of the body and limbs are not formed of dis- tinct plates, like those of the Echinus-shell; but are only divided by sutures at the joints, for the purpose of permitting the requisite freedom of motion. It might have been thought that here, if anywhere, a process of interstitial growth would have existed, to adapt the capacity of the envelopes to the dimensions of the contained parts, as the latter increase with the growth of the animal; but, true to the general principle, that epidermic structures are not only extra-vascular, but that they undergo no change when they are once fully formed, we find that the hard enve- lopes of Articulated animals are thrown off, or exuviated, when the con- tained parts require an increase of room; and that a new covering is formed from their surface, adapted to their enlarged dimensions. 285. This is well knoAvn to occur at certain intervals in Crabs, Lob- sters, and other Crustacea ; which thus exuviate not merely the outer shell, with the continuation of the epidermis over the eyes, but also its internal reflection, Avhich forms the lining of the oesophagus and stomach, and the tendinous plates by which the muscles are attached to the lining of the shell. A similar moulting may be observed to occur in some of the minute Entomostracous Crustacea of our pools, every two or three jdays, even after the animals seem to be full grown. During the early groAvth of Insects, Spiders, Centipedes, &c, a similar moult is frequently repeated at short intervals ; but after these animals have attained their full groAvth, which is the case with Insects at their last change, no fur- ther moulting takes place, the necessity for it having ceased. This moulting is precisely analogous to the exfoliation and new formation of the Epidermis, in Man and most other Vertebrata; differing from it only in this, that the latter is constantly taking place to a small extent, whilst the former is completely effected at certain intervals, andvthen ceases. We have examples of a periodical complete moult in Vertebrata, howeA'er, among Serpents and Frogs. 286. The structure of the hard envelopes of Articulated animals cor- responds with that of the Epidermis and its appendages in Man. The firm casings of Beetles, for example, are formed of layers of epidermic cells, united together, and having their cavities filled by a horny secre- tion. The densest structure is found in the calcareous shells of the Crustacea; Avhich consists of a substance precisely analogous to the Dentine of Teeth (§ 311); covered on the exterior with a layer of pigment-cells. The calcareous matter consists chiefly of carbonate of lime ; but traces of the phosphate are also found. The animal basis has a firm consistent structure, resembling that of teeth. A thin ver- 172 STRUCTURE AND ENDOWMENTS OF ANIMAL TISSUES. tical section shows the tubuli running nearly parallel, but with occasional undulations, from the internal surface towards the external ; but no traces of the original calcigerous cells can be detected in the fully-formed shell, the process of fusion having gone so far as to obliterate them. The manner in which these tubuli are formed, will be presently con- sidered, under the head of Dental substance. 287. Noav the condition of the osseous skeleton of vertebrated animals is altogether different. Its purpose is still only mechanical; its sole use being, to afford support and protection to the softer textures, and to form inflexible levers by the action of the muscles, upon Avhich motion may be given to the different parts of the fabric. But it forms a part of the internal substance of their bodies; and as these grow in every part, and not merely by addition to this or that- portion, so must the Bones also, in order to keep pace with the rest of the structure. Hence we find them so formed, that the process of interstitial deposition may be continually going on in their fabric, as in that of the softer tissues; and the changes in their substance do not cease, even Avhen they have acquired their full size. The subsequent continuance of these changes appears destined, not so much to repair any waste occasioned by decom- position,—for this must be very trifling in a tissue of such solidity,—as to keep the fabric in a condition, in which it may repair the injuries in its substance occasioned by accident or disease. The degree of this reparative power is proportional, as we shall presently see, to the activity of the normal changes, which are continually taking place in the bone ; and is thus much greater in youth than in middle life, and greater in the vigor of manhood than in old age. 288. The structure of Bones is Avell adapted to demonstrate the dis- tinction between the tissues themselves, and those subsidiary parts, by which they are connected with the rest of the fabric. We have seen that Cartilage is essentially non-vascular ; that is, even when it exists, in a considerable mass, it is not traversed by vessels, but is nourished by absorption from the fluids contained in the vessels distributed on its exterior. Now every mass of Bone is penetrated by vessels ; nevertheless these do not penetrate its ultimate substance, and may be easily sepa- rated from it, leaving the bone itself as it was. In fact, as Prof. Goodsir observes, " a well-macerated bone is one of the most easily made, and, at the same time one of the most curious anatomical preparations. It is a perfect example of a texture completely isolated ; the vessels, nerves, membranes, and fat, are all separated, and nothing is left but the non- vascular osseous substance." Precisely the same may be said of the substance of a Tooth, from wrhich the vascular lining of the pulp-cavity has been removed; for it then possesses neither vessels, nerves, nor lymphatics ; and yet, as Ave shall presently see, it has a highly-organized structure, peculiar to itself. 289. The general characters of Osseous texture vary according to the shape of the Bone, and the part of it examined. Thus in the long bones we find the shaft pierced by a central canal, which runs continuously from one extremity to the other; and the hollow cylinder which sur- rounds this is very compact in its structure. On the other hand, the dilated ends of the bone are not penetrated by the large central canal; STRUCTURE OF BONE. 173 Fig. 45. Extremity of Os femo- nor are they composed of solid osseous substance. They are made up of cancellated structure, as it is termed ; that is, of osseous lamellae and fibres interwoven together (like those of areolar tissue, on a larger scale) so as to form a multitude of minute chambers or cancelli, freely communicating with each other, and with the cavity of the shaft; whilst the whole is capped Avith a thin layer of solid bone. Again, in the thin flat bones, as the scapula, we find the two surfaces composed of solid osseous texture, with more or less of cancellated structure interposed betAveen the layers. And in the thicker flat bones, as the parietal, frontal, &c, this cancel- lated structure becomes very distinct, and forms the diploe; this, however, is sometimes deficient, leaving a cavity analogous to the canal of the long bones: whilst the plates which form the surfaces of the bone (the external and internal tables of the skull), re- ^a^^l^^\^t semble in their thickness and solidity, as well as in of bone, in contact with the intimate structure presently to be described, the &, canceiii! shaft or hollow cylinder of those bones. Finally, we frequently meet (especially in the Ethmoid and Sphenoid bones) with thin lamellae of osseous substance, resembling those which elsewhere form the boundaries of the cancelli; these consist of but one layer of bony matter, and shoAV none of the varieties previously adverted to ; they are not penetrated by vessels, but are nourished only by their surfaces ; and they consequently exhibit to us the elements of the osseous struc- ture in their simplest form. It will be desirable, therefore, to commence with the description of these. 290. When a thin natural lamella of this kind is examined, it is found to be chiefly made up of a substance which is nearly homogeneous, sometimes exhibiting indistinct traces of a fibrous arrangement; this, however, may be generally resolved by prolonged boiling, into an assem- blage of minute granules, varying in size from l-6000th to l-14,000th Fig. 46. Lacunae of Osseous substance, magnified 500 diameters:—a, central cavity; 6, its ramifications. of an inch, Avhich are more or less angular in shape, and seem to cohere by the medium of some second substance, which is dissolved by the boiling. They are composed of Calcareous salts, apparently in chemical union Avith the Gelatine that forms the basis of the osseous substance. In the midst of this granular substance, a number of dark spots are to be observed, the form of Avhich is very peculiar. In their general out- line, they are usually somewhat oval; but they send forth numerous radiating prolongations of extreme minuteness, which may be frequently 174 STRUCTURE AND ENDOWMENTS OF ANIMAL TISSUES. traced to a considerable distance. These spots, known as the osseous corpuscles (sometimes termed the Purkinjean corpuscles, after the name of their discoverer), are highly characteristic of the true bony structure, being never deficient in the minutest parts of the bones of the higher animals, although those of Fishes are frequently destitute of them. These corpuscles were formerly supposed, from their dark appearance, to be opaque, and to consist of aggregations of calcareous matter which Avould not transmit the light: but it is now quite certain, that they are lacunce or open spaces ; and that the radiating prolongations from them, which are far smaller than the minutest capillary vessel, are canaliculi, or delicate tubes. Of these canaliculi, some may be seen to interlace freely with each other, whilst others proceed toAvards the surface of the bony lamella; and thus a system of passages, not by any means wide enough to admit the blood-corpuscles, but capable of transmitting the fluid elements of the blood, or matters selected from them, is established through the whole substance of the lamella. 291. The lacunae of Human bone have an average length of l-1800th of an inch; and they are usually about half as wide, and one-third as thick. The diameter of the canaliculi is from l-12,000th to l-20,000th of an inch. Their size and form differ greatly, however, in the different classes of Vertebrata; so that it is usually possible to refer a mere frag- ment of bone to its proper group, by an examination of its minute structure. The succeeding figure represents the arrangement of these lacunae and canaliculi in the bony scale of a fish (the Lepidosteus); which is almost the only existing representative of a large class of bony- scaled Fishes, that formerly tenanted the seas. Its lacunae will be seen to differ greatly in form from those of human bone ; and the canaliculi which proceed from them are much feAver in number.—The purpose of this penetration of the osseous texture by such a complicated apparatus of tubuli, can scarcely be anything else than the maintenance Fig. 47. Section of the bony scale of Lepidosteus;—a, showing the regular distribution of the lacunas and of the connecting canaliculi; 6, small portion more highly magnified. of its vitality by the continual percolation of nutrient fluid, drawn into the system of lacunae and canaliculi from the neighboring blood-vessels. Thus the nutrition of the ultimate osseous texture, though carried on upon the same general plan Avith that of Cartilage, differs in this :— that there is a provision in Bone for the ready transmission of nutrient matter through its texture, by means of minute channels, which does not exist in Cartilage; a difference obviously required by the greater solidity of the substance of the former, which does not allow of the diffused imbibition, that is permitted by the softer and moister nature of the latter. We shall presently find that these channels are only STRUCTURE OF BONE. 175 formed at a late stage of the development of bone, where the remaining tissue has acquired its completest consolidation. m 292. Now, as already remarked, the simple structure just described is found, not merely in the delicate plates which form the thinnest part of certain bones in Man, but also in those lamella?, which form the walls of the cancelli of the larger and thicker bones. Every one of these lamellae repeats, in fact, the same history. The cancelli are lined by a membrane derived from that of the cavity of the shaft, over which blood-vessels are minutely distributed; between these blood-vessels and the osseous texture is a layer of cells; and from the materials selected and communicated by these, each lamella is nourished, through its system of radiating canaliculi and nutritive centres. The cancelli, at the time of their formation in the foetal bone, are entirely filled with such cells; which appear (as will be presently explained) to be the descendants of the cells of the original cartilage; but in the adult bone, a large proportion of them are filled with fatty matter, which they secrete into their cavities.—The Vessels of the cancellated struc- ture at the extremities of the long bones are derived from those of the medullary cavity, which is penetrated by large trunks from the exterior; and ,n the flat bones, they form a system of their own, connected with the vessels of the exterior by several smaller trunks. 293. The solid osseous texture which forms the cylindrical shafts of the long bones, and the thick external plates of the denser flat bones, is not cut off from nutritive action in the degree in which it might seem to be; tor it is penetrated by a series of large canals, termed the Haversian (after Clopton Havers, their discoverer), which form a net- work in its interior, and which serve for the transmission of blood- vessels through its substance (Fig. 48). These canals, in the long bones, run for the most part in a direction parallel to the central cavity ; and they communicate with this, with the external surface, with the cancelli, and Avith each other, by frequent transverse branches; so that the whole system forms an irregular network, pervading every part of the solid texture, and adapted for the establishment of vascular communications throughout. The diameter of the Haversian canals varies from l-2500th to l-200th of an inch, or more; their ^ average diameter may be stated at about l-500th of an inch. They are lined by a membrane which is continuous with that of the external surface, and which carries this inwards, so to speak, to form the lining membrane of the cen- tral cavity, and of the cancelli;—and the cavity of the tube encloses a single twig of an artery or vein. Thus we may consider the whole Osseous texture as enclosed in a membranous bag; on which blood-vessels are minutely dis- tributed, and which is so carried into the bone by involutions and prolongations, that no part of the latter is ever far removed from a vascular surface. Haversian Canal, seen on a lon- gitudinal section of the compact tissue of the shaft of one of the long bones; 1, arterial canal; 2, venous canal; 3, dilatation of another venous canal. 176 STRUCTURE AND ENDOWMENTS OF ANIMAL TISSUES. 294. In the adult bone, the cells which fill the remaining cavity of these canals secrete fatty matter. This is particularly evident in the case of the central cavity, which may be considered as an immensely dilated Haversian canal, where they constitute the medulla or marrow. It does not appear that these take any active part in the nutrition of the bone; indeed, in the bones of Birds the shaft is entirely hollow, and air is admitted into it from the lungs, so that its lining membrane is rendered subservient to the aeration of the blood. 295. The arrangement of the elementary parts of the osseous sub- stance around the Haversian canals is very interesting and beautiful. When a transverse section of a long bone is made, the open orifices of the longitudinal canals present themselves at intervals, sometimes con- nected by a transverse canal where the section happens to traverse this. Around these orifices, we see the osseous matter arranged in*the form of cylinders, which appear to be marked by concentric circles (Fig. 49, 1, 2). Now when one of these circles is minutely examined, it is found to be made up of a series of lacunae, analogous to those already de- scribed; these, however, are seldom or never so continuous as to form a complete circle. The long sides of the lacunae are directed, the one Fig. 49. Minute structure of bone, drawn with the microscope from nature. Magnified 300 diameters. 1. One of the Haversian canals surrounded by its concentric lamellae. The lacunae are seen between the lamellse; but the radiating tubuli are omitted. 2. An Haversian canal with its concentric laminse, lacunas, and. radiating tubuli. 3. The area of one of the canals. 4, 4. Direction of the lamellae of the great medullary canal. Between the lamellae at the upper part of the figure, several very long lacuna; with their tubuli are seen. In the lower part of the figure the outlines of three other canals are given, in order to show their form and mode of arrangement in the entire bone. towards the Haversian canal (3) in the centre, the other toAvards the circular row next beyond it. .And when the course of the canaliculi is traced, it is found that these converge on the inner side towards the central canal, inosculating with those of the series next within, whilst those of the outer side pass outwards in a radiating or diverging direc- tion, to inosculate with those of the series next external. Thus a com- plete communication is formed, by means of this system of radiating canaliculi and intervening lacunae, between the central canal and the COMPOSITION OF BONE. 177 outermost cylindrical lamella of bony matter ; and each of these lamellae derives its nourishment from the vessels of the central canal, through the lamellae which intervene between itself and the vascular membrane lining that tube. 296. Thus every one of the Haversian canals is the centre of a cylindrical ossicle, which is complete in itself, as far as its elementary structure is concerned, and Avhich has no dependence on, or connexion with, other similar ossicles. These are arranged, hoAvever, side by side, like sticks in a fagot; they are bound together by a thin cylinder of bone, on the exterior of all, Avhich derives its nourishment from the periosteum, or enveloping membrane ; in like manner, the hollow bundle is lined by a similar cylinder, which surrounds the great medullary cavity, and is nourished by its vascular membranes; and the spaces that here and there intervene between the ossicles are filled up with laminae, which are parallel to those of the external and internal cylinders, and which seem to derive their nutriment from them (Fig. 49, 4). In this manner, the whole structure acquires great density and solidity.— The structure of the outer and inner tables of the skull, and of other thick solid layers of bone, is precisely similar; except that the Haver- sian canals have no such definite directions, and form an irregular net- work. 297. Thus we see that each of the lamellae of bone surrounding an Haversian canal, or bounding the cancelli, may be regarded as a repe- tition of the simple bony plate, which draAvs its nourishment direct from the vascular membrane covering its surface, by means of its system of lacunae and canaliculi. The membrane lining the Haversian canals, cancelli, and central medullary cavity, is an internal prolongation of that which clothes the exterior ;—just as the mucous membranes, with their extensions into glandular structures, are internal prolongations of the true skin. Every Haversian canal and every cancellus are repe- titions of each other in all essential particulars, their form alone being different. The central medullary canal is but an enlarged Haversian canal or cancellus. And the Avhole cylindrical shaft is a collection of ossicles, each of which is a miniature representation of itself, being a hollow cylinder, Avith a central vascular cavity. 298. The principal features of the Chemical constitution of Bone are easily made evident. After all the accessory parts have been removed, and nothing remains but the real osseous texture, this may be separated, by simple processes, into its tAvo grand constituents,—the animal basis, and the calcareous matter. The latter may be entirely removed by maceration of the bone in dilute Muriatic or Nitric acid ; and a substance of cartilaginous appearance is then left, which, Avhen submitted to the action of boiling water for a short time, is almost entirely dissolved aAvay, and the solution forms a dense jelly on cooling. The same sub- stance, Gelatine, may be obtained by long boiling under pressure, from previously unaltered bone; and the calcareous matter is then left in a friable condition. By submitting a bone to a heat sufficient to decom- pose the animal matter, Avithout dissipating any of the earthy particles, we may obtain the whole calcareous matter in situ; but the slightest violence is sufficient to disintegrate it. The bones of persons long 178 STRUCTURE AND ENDOWMENTS OF ANIMAL TISSUES. buried are often found in this condition; their form and position being retained until they are exposed to the air, or are a little shaken, when they crumble to dust. The proportion of the earthy matter of Bones to the animal basis may be differently stated, according as we include, in our estimate of the latter, the contents of the medullary cavity, the Haversian canals, and the cancelli, or confine ourselves to that portion only of the animal matter Avhich is united with the calcareous element in the proper osseous tissue. According to the recent experiments of Dr. Stark,* the relative amount of the two elements, in the latter estimate, is subject to very little variation, either in the different classes of animals, or in the same species at different ages, the animal matter composing about one-third, or 33^ per cent., and the mineral matter two-thirds, or 66^ per cent. The degree of hardness of bones does not altogether depend, therefore, on the amount of earthy matter they may contain; for the flexible, semi-transparent, easily-divided bones of Fish contain as large an amount of animal matter, as the ivory-like leg-bones of the Deer or Sheep. The usual analyses of Bone, however, have been made upon the former kind of estimate : and they show that the proportion of the earthy matter to the whole of the animal substance contained in bone varies much in different animals, in the same animal in different ages, and even in different parts of the same skeleton. The reason of this will be apparent, Avhen the history of the growth of Bone has been explained; since there is a gradual filling up of all the cavities at first occupied by fat-cells, vessels, &c, which does not cease with adult age, but which continues during the whole of life. In this manner the bones of old persons acquire a high degree of solidity, but they become brittle in proportion to their hardness. From the same cause, the more solid bones contain a larger proportion of bone-earth than those of a spongy or cancellated texture ; the temporal bone, for exam- ple, containing 63J per cent., whilst the scapula possesses only 54 per cent. In the former of these bones, the proportion is nearly th©\ same as that which exists in pure osseous tissue, the amount of the remaining tissues which it includes being very small, on account of the solidity of the bone; but the latter contains in its cancelli a large quantity of blood-vessels, fat-cells, &c, which swell the proportion of the animal matter from 33J to 46 per cent. 299. The Lime of bones is for the most part in a state of Phosphate, especially among the higher animals ; the remainder is a carbonate. In Human bones, the proportion of the latter seems to be about one- sixth or one-seventh of the whole amount of bone-earth. In the bones of the loAver animals, however, the proportion of Carbonate is greater; and it is curious that in callus, exostosis, and other irregular osseous formations in the higher animals, the proportion of the Carbonate should be much greater than in the sound bone. In caries, however, the proportion of the Carbonate is less than usual. The composition of the Phosphate of Lime in Bones, is somewhat peculiar; eight pro- portions of the base being united with three of the acid. According to Professor Graham, it is to be regarded as a compound of two tribasic * Edinburgh Medical and Surgical Journal, April, 184o. COMPOSITION OF BONE. 179 phosphates; one atom of the neutral phosphate (in which one propor- tional of the acid is united Avith tAvo of lime and one of water), being united Avith two proportionals of the alkaline phosphate (in which one part of acid is united with three of the base), together with an atom of water, Avhich is driven off by calcination. Besides these components, some Chemists assert that a small quantity of Fluoride of calcium is present in Bone; but this is rather doubtful, since it has been shoAvn by Dr. G. 0. Rees that the solvent action upon glass, which has been supposed to be characteristic of fluoric acid, may be imitated by phos- phoric acid in combination with water, Avhich, if heated upon glass of inferior quality until it volatilizes, will act upon it with considerable energy.—Other saline matters, such as phosphate of magnesia, oxides of iron and manganese, and chloride of sodium, are found in bones in small amount. 300. The first development of Bone is usually preceded by the for- mation of a Cartilaginous structure, which occupies the place after- Avards to be taken by the bone; and it is commonly considered that the bone is formed by the calcification of the cartilage-substance. This, however, does not appear to be the case, as will be presently shoAvn; and it Avould probably be more correct to say that the cartilage is super- seded by bone. Moreover, Bone is frequently developed in the sub- stance of Fibrous membranes ; and the structure produced by this intra- membranous ossification cannot be distinguished from that which is generated by the intra-cartilaginous.—We shall commence the history of the development of Bone, with the period in Avhich its condition resembles that of the permanent Cartilages. As already mentioned, there is no essential difference between the temporary and permanent Cartilages, in regard to their ultimate structure; the former, however, are more commonly traversed by vessels, especially when their mass is considerable. These vessels, however, do not pass at once from the exterior of the cartilage into its substance; but they are conveyed in- wards along canals, which are lined by an extension of the perichondrium or investing membrane, and which may thus be regarded'as so many involutions of the outer surface of the cartilage. These canals are espe- cially developed at certain points, which are to be the centres of the ossifying process ; of these puncta ossificationis, we usually find one in the centre of the shaft of a long bone, and one in each of its epiphyses; in the flat bones there is one in the middle of the surface, and one in each of the principal processes. Up to a late stage of the ossifying process, the parts which contain distinct centres are not connected by bony union, so thatthey fall apart by maceration ; and even when they should normally unite, they sometimes remain separate,—as in the case of the Frontal, bone, in which we frequently meet Avith a continuation of the sagittal-suture down the middle, dividing it into two equal halves, which have originated in two distinct centres of ossification. It is inte- resting to remark that, in the two lowest classes of Vertebrata,—Fishes and Reptiles,—we find the several parts of the osseous system present- ing in a permanent form, many of the conditions which are transitory in the higher; thus the different portions of each vertebra, the body, lateral arches, spinous and transverse processes, &c, which have their 180 STRUCTURE AND ENDOWMENTS OF ANIMAL TISSUES. distinct centres of ossification, but Avhich early unite in Man, remain permanently distinct in the loAver Fishes ; the division of the frontal bone, just adverted to, is constant amongst Fishes and Reptiles ; and in these classes we meet with a permanent separation of the parts of the occipital and temporal bones, which, being formed from distinct centres of ossification, are at first distinct in the higher animals. 301. During the formation of the punctum ossificationis, and the spread of the vessels into the cartilaginous matrix, certain changes are taking place in the substance of the latter, preparatory to its conversion into bone. Instead of single isolated cells, or groups of tAvo, three, or four, such as we have .seen to be characteristic of ordinary Cartilage (§ 267), we find, as Ave approach the ossifying centre, clusters made up of a larger number, Avhich appear to be formed by a continuance of the same multiplying process as that already described (Fig. 50). And Fig. 50. Section of Cartilage near the seat of ossification; each single cell having given birth to four, five, or six cells, which form clusters. These clusters become larger towards the right of.the figure, and their cells more numerous and larger; their long diameter being l-1500th of an inch. when we pass still nearer, we see that these clusters are composed of a yet greater number of cells, which are arranged in long rows, Avhose direction corresponds with the longitudinal axis of the bone ; these clus- ters are still separated by intercellular substance, and it is in this, that the ossific .matter is first deposited (Fig. 51). Thus if we separate the Fig. 51. The same cartilage at the seat of ossification: tbe clusters of cells are arranged in columns ; tho intercel- lular spaces between the columns being l-3250th of an inch in breadth. To the right of the figure, osseous fibres are seen occupying the intercellular spaces, at first bounding the clusters laterally, then splitting them longitudinally and encircling each separate cell. Tbe greater opacity of the right hand border is due to a threefold cause, the increase of osseous fibres, the opacity of the contents of the cells, and the multipli- cation of oil-globules. cartilaginous and the osseous substance at this period, we find that the ends of the rows of cartilage-cells are received into deep narrow cups of CONVERSION OF CARTILAGE INTO BONE. 181 bone, formed by the transformation of the intercellular substance be- tween them. Immediately upon the ossifying surface, the nuclei, which were before closely compressed, separate considerably from one another, by the increase of material within the cells ; and the nuclei themselves become larger and more transparent. These changes constitute the first stage of the process of ossification, Avhich extends only to the calcifica- tion of the intercellular substance ; in this stage there are no blood- vessels directly concerned. The bony lamellae thus formed, mark out the boundaries of the cancelli and Haversian canals, which are after- wards to occupy a part of the space that is hitherto filled by the rows of cartilage corpuscles. 302. Up to this point, there is no essential difference in the accounts of those Avho have most carefully studied the process of ossification ; but in regard to the history of its subsequent stages, there is much discre- pancy; and this especially with respect to the origin of the bone-lacunae, which some regard as metamorphosed cartilage-cells, others as the spaces originally occupied by their nuclei, whilst others do not regard them as in any way derived from the cartilage-cells, but consider them as a new formation. Much may doubtless be urged in favor of each view; the author's OAvn observations incline him to the latter, and lead him to regard the lacuna? as cells, Avhich like the pigment-cells of Batrachia, ifec, have sent out the stellate prolongations that constitute the canali- culi. All stages of gradation may be traced, between simple rounded cavities,—whose correspondence in size Avith the cells that are scattered in the midst of the consolidating blastema leaves scarcely any doubt of their identity with these,—and the lenticular lacuna with numbers of canaliculi proceeding from it. These gradations are particularly well seen during the process of ossification ; so that it seems probable that the radiating extension of the cells takes place during the consolidation of the surrounding tissue.—It is an additional argument against the idea that the bone-lacunae in any Avay originate from the cartilage-cells, that they are found to present exactly the same characters in bone Avhich is developed in the substance of fibrous membrane (after the manner to be presently described), and in the formation of which, there- fore, cartilage has had no participation. 303. Although, in a large proportion of the skeleton, the formation of Bone is thus preceded by that of cartilage, yet such is by no means invariably or necessarily the case ; for the flat bones, such as the scapula, and those forming the roof of the skull, haA'e usually only a centre of cartilage, beyond Avhich the ossifying process extends in membrane only. This membrane is chiefly composed of fibrous fasciculi, corre- sponding with those of the Avhite fibrous tissues; but amongst these are seen numerous cells, some about the size of blood-discs, but others two or three times larger, containing granular matter; and a soft amorphous or faintly granular matter is also found interposed amidst the fibres and cells. The process of ossification here seems essentially to consist in the consolidation of the fibres by earthy matter; for the first bony deposit is seen as an irregular reticulation, very loose and open towards its edges, and there frequently presenting itself in the form of distinct spicula, Avhich are continuous with fasciculi of fibres in the surrounding 182 STRUCTURE AND ENDOWMENTS OF ANIMAL TISSUES. membrane. The limits of the calcifying deposit may be traced by the opaque and granular character of the parts affected by it; and it gradu- ally extends itself, involving more and more of the surrounding mem- brane, until the foundation is laid for the entire bone. Everywhere the part most recently formed consists of a very open reticulation of fibro- calcareous spicula, whilst the older part is rendered harder and more compact by the increase in the number of these spicula. As the process advances, and the plate of bone thickens, a series of grooves or furrows, radiating from the ossifying centre, are found upon its surface; and these, by a further increase in thickness, occasioned by a deposit of ossific matter all around them, are gradually converted into close canals (the Haversian), which contain blood-vessels, and are lined by processes of the investing membrane. The lacunae and canaliculi seem to take their origin in the cells which are interspersed among the fibres, their prolongations extending themselves, and insinuating themselves through the spaces left between the interlacing fibres, whilst the process of cal- cification is going on. 304. The first osseous tissue which is formed by either of these pro- cesses, has an irregular cancellated structure, analogous to that which is found at the extremities of the long bones in adults. This is gradu- ally modified by changes Avhich essentially consist in absorption and new deposition; for the absorptive process first unites minute areolae into larger ones, by removing their partitions ; and it is upon their in- terior walls that neAV osseous lamellae are noAV deposited, from materials supplied by the blastema they contain. It is by a process of this kind, that the central medullary cavity is first formed in the bones of young animals. At an early period, no such cavity exists, and its place is occupied by small cancelli; this is the permanent condition of the bones in most Reptiles. The cancelli gradually enlarge, hoAvever ; and those within the shaft coalesce Avith one another until a continuous tube is formed, around which the cancelli are large, open, and irregular. At the same time, the diameter of the surrounding shaft is increasing by the process of interstitial groAvth just described ; so that the size of the medullary cavity at last becomes greater than that of the whole shaft Avhen its formation commenced. The aggregation of the osseous matter in a holloAV cylinder, instead of a solid one, is the form most favorable to strength, as may be easily proved upon mechanical principles. The same arrangement is adopted in the arts, Avherever it is desired to obtain the greatest strength with a limited amount of material. 305. The growth of Bones takes place by the addition of new tissue to the part already formed; but this addition may take place in three modes,—namely, by the development of neAV bone in the cartilage yet remaining between the different centres of ossification; by the develop- ment of new bone in the membrane covering the surface; and by the interstitial formation of neAV layers Avithin the Haversian canals and cancelli of the part already formed, by which the requisite solidity is given to it. Of the first process Ave have the most characteristic ex- ample in the increase in length of a long bone, by the ossification of the cartilage which intervenes betAveen the shaft and the epiphyses, and Avhich continues to groAV, up to the time of the final union of these REGENERATION OF BONE. 183 parts. Thus it Avas long since proved by the experiments of Hales and Hunter, that the growth of a long bone takes place chiefly towards the extremities ; for they found that, when metallic substances were inserted in the shaft of a growing bone of a young animal, the distance between them was but little altered after a long interval, whilst the space be- tAveen the extremities of the bone had greatly increased. And it seems that, at a later period, when the epiphyses have become completely united to the shaft, an elongation continues to take place, by the slow ossification of the articular cartilage.—Again, the bone is progressively increased in thickness, by the gradual production of neAV osseous matter upon its surface; this production being effected by the conversion of the inner layer of the periosteum, the fibres of which are found to be continuous Avith those of the animal matrix of the surface of the bone.— And it is by the successive formation of neAV layers of osseous tissue, one Avithin another, giving the appearance of concentric rings Avhen the Haversian canals are cut across (Fig. 49), that the proportion of hard to soft parts in bone is gradually increased; the calibre of the Haver- sian canals being correspondingly diminished. Of this we have a curious exemplification in the antlers of the deer, in Avhich the cavity of the canals is gradually choked up by the formation of osseous tissue, until the vascular supply is cut off, and the death of the bone is the result. 306. The difference in the relations of the Osseous substance to the Arascular netAvork, at different ages,—accounting for the variations in the rapidity of its nutrition and reparation,—is well displayed in the effects of Madder. This substance has a peculiar affinity for Phosphate of Lime; so that Avhen the latter is formed by precipitation in a fluid tinged with madder, it attracts color to it in its descent, and falls to the bottom richly tinted. Now Avhen animals are fed with this sub- stance, it is found that their bones become tinged with it, the period required being in the inverse proportion to their age. Thus in very young'animals a single day suffices to color the entire skeleton, for in them there is no osseous matter far from the vascular surfaces; Avhen sections are made, hoAvever, of the bones thus tinged, it is found that the color is confined to the immediate neighborhood of the Haversian canals, each of Avhich is encircled by a crimson ring. In full-groAvn animals, the bones are very slowly tinged; because the osseous texture is much more consolidated and less permeablje to fluid than in earlier life; and because the vascular membrane lining the Haversian canals is removed further from the outer and older layers of osseous tissue which surround them, by the interposition of newer concentric layers, which diminish the diameter of the canals. In the bones of half-groAvn animals, a part of the bone is nearly in the perfect condition, Avhile a part is new and easily colored; so that the action of this substance enables us to distinguish the new from the old. 307. The Regeneration of Bone, after loss of its substance by disease or injury, is extremely complete; in fact there is no other structure of so complex a nature, Avhich is capable of being so thoroughly repaired. Although the regenerative poAver appears to be so much less in Verte- brated animals, than it is in the loAver Invertebrata, yet it is probably 184 STRUCTURE AND ENDOAVMENTS OF ANIMAL TISSUES. not at all lower in reality,—the neAV structures actually formed being as complex in the one case as in the other. It is noAvhere, perhaps, more remarkably manifested, than in the re-formation of nearly an entire bone, Avhen the original one had been lost by disease ; all the attach- ments of muscles and ligaments, as Avell as the external form and inter- nal structure, being ultimately found as complete in the neAV bone, as they originally Avere in that which it has replaced. Much discussion has taken place in regard to the degree, in which the different membra- nous structures, that surround bone and penetrate its substance, par- ticipate in its regeneration; some having supposed the periosteum to have the power of itself forming neAV bone, others attributing the same power to the membrane lining the medullary cavities. It appears cer- tain, hoAvever, that new osseous tissue may be formed in a great variety of modes. It has been ascertained by Mr. Paget* that it may be pro- duced through the intermediation of perfect fibrous tissue, either when this previously existed as such (as the periosteum or interosseous mem- brane), or Avhen it has been neAvly formed by the fibrillation of the plastic fluid effused as the material for reparation. The agency of the perios- teum is seen in many cases of necrosis, in Avhich that membrane has been completely detached from the dead shaft, and ne>v bone has been generated from its interior. The ossification of a neAvly-produced fibrous membrane is believed by Mr. Paget to be the ordinary mode of repara- tion of fractures of the skull; and it takes place in a manner essentially the same as that of the original intra-membranous development of bone. But new bone may also be formed, according to that most excellent ob- server, by ossification of the fibrous tissue in its rudimental state (§ 193). In abnormal bone-growths, it sometimes appears as if the tissue had been formed by the ossification of cells; but more commonly the calci- fication takes place in an earlier stage of tissue-production, that of the "nucleated blastema," in which a granular osseous deposit is seen, Avhich gradually increases so as to form the lamellae of a fine cancellous texture, at the same time inclosing the nuclei, which seem to occupy the places afterwards to remain as the lacuna). It is seldom that the reparation of bone takes place through the intermediation of cartilage ; though this is occasionally formed, rather, perhaps, in the loAver animals than in the human subject. 308. The reparation of Bone, after disease or injury, takes place ex- actly upon the same plan as its first formation. A plastic or organiza- ble exudation is first poured out from the neighboring blood-vessels, and this forms a sort of bed or matrix, in Avhich the subsequent processes take place. At first all new bone possesses a minutely cancellous struc- ture, much like that of the foetal bones in their first construction; but this gradually assimilates itself to the structure of the bones which it repairs, its outer portions acquiring a more compact laminated struc- ture, while its interior substance acquires Avider cancellous spaces, and a perfect medulla. When the shaft of a long bone of an animal has been fractured through, and the extremities have been brought evenly to- gether, it is found that the new matter first ossified is that which oc- cupies the central portion of the deposit, and which thus connects the * Lectures on Reproduction and Repair, Medical Gazette, 1849. FORMATION OF TEETH. 185 medullary cavities of the broken ends, forming a kind of plug that enters each. This was termed by Dupuytren, by whom it Avas first dis- tinctly described, the provisional callus. This is usually formed in the course of five or six weeks, or less in young subjects; but at that period the contiguous surfaces of the bone itself are not cemented by bony union ; and the formation of the permanent callus occupies some months, during which the provisional callus is gradually absorbed, and the con- tinuity of the medullary canal restored, in the same manner as it was at first established. The permanent callus has all the characters of true bone. It seems to have been established by the observations of Mr. Paget, however, that these statements do not usually apply to the case of Man ; in Avhom, Avhen the limb is kept at rest, the union between the fractured ends is accomplished by ossification of the substance connect- ing them, without the intermediation of a provisional callus; this being only formed Avhen the portions of the bone are kept in continual move- ment. 309. The most extensive reparation is seen, when the shaft of a long bone is destroyed by disease. If violent inflammation occur in its tissue, the death of the fabric is frequently the consequence,—apparently through the blocking-up of the canals with the products of the inflam- matory action, and the consequent cessation of the supply of nutriment. It is not often that the Avhole thickness of the bone becomes necrosed at once; more commonly this result is confined to its outer or its inner layers. When this is the case, the neAV formation takes place from the part that remains sound; the external layers, Avhich receive their vas- cular supply from the periosteum and from the Haversian canals con- tinued inAvards from it, throAving out neAV matter on their interior, Avhich is gradually converted into bone; whilst the internal layers, if they should be the parts remaining uninjured, do the same on their exterior, deriving their materials from the medullary membrane and its prolonga- tions into their Haversian canals. But it sometimes happens that the Avhole shaft suffers necrosis; and as the medullary membrane and the entire system of Haversian canals have lost their vitality, reparation can only take place from the periosteum, and from the living bone at the tAvo extremities. This is consequently a very sIoav process; more especially as the epiphyses, having been originally formed as distinct parts from the shaft, do not seem able to contribute much to the regene- ration of the latter. 310. We next proceed to the Teeth, Avhich are organs of mechanical attrition, developed in the first part of the alimentary canal, for the purpose of comminuting the food conveyed into it. Their place of ori- gin is altogether different from that of bone, as they commence in little papillary elevations of the mucous membrane covering the jaw ; but the substance from which they are formed is the same primitive cellular tissue, as that in which Cartilage itself originates. We may best understand the structure and development of the Teeth in Man, by first inquiring into the characters presented by those of some of the lower animals, and the history of their eArolution. In the foetal Shark, the first appear- ance of the tooth is in the form of a minute papilla on the mucous mem- brane covering the jaAvs; the substance of this papilla is composed of ispherical cells, which are imbedded in a kind of gelatinous substance 186 STRUCTURE AND ENDOAVMENTS OF ANIMAL TISSUES. resembling that of incipient cartilage; whilst its exterior is composed of a dense, structureless, pellucid membrane. The cellular masses not at first permeated by vessels ; but a small arterial branch is distributed to each papilla, and? spreads out into a tuft of capillaries at its base (Fig. 52). The papilla gradually enlarges, by the formation of neAV cells at the part immediately adjacent to the blood-vessels, which supply the material requisite for their development; and Avhen it has acquired its full size, the process of calcification takes place, by which it is converted into Dentine, the substance most characteristic of teeth. 311. This Dentine, Avhich in the Elephant's tusk is knoAvn as Ivory, is a firm substance, in which mineral matter predominates to a greater extent than in bone; but wdiich still has a definite animal basis, that retains its form Avhen the calcareous matter has been removed by mace- ration in acid. In every 100 parts, the animal matter forms about 28; and of the mineral portion, phosphate of lime constitutes about M\ parts, carbonate of lime 5-£ parts, and phosphate of magnesia and soda, Avith chloride of sodium, about 21 parts. When it is fractured, it seems to possess a fibrous appearance; the fibres radiating from the centre of the tooth toAvards its circumference. But Avhen a thin section of it is submitted to the microscope, it is seen that this fibrous appearance is due to a peculiar structure in the dentine, which the unaided eye cannot discover. The dentinal substance is itself very transparent; but it is traversed by minute tubuli, which appear as dark lines, generally in very close approximation, running from the internal portion of the tooth towards the surface, and exhibiting numerous minute undulations, and sometimes more decided curvatures, in their course (Fig. 53). They STRUCTURE AND DEVELOPMENT OF DENTINE. 187 Fig. 53. occasionally divide into two branches, Avhich continue to run at a little distance from one another in the same direction ; and they also fre- quently give off small lateral branches, Avhich again send off smaller ones. In some animals the tubuli may be traced at their extremities into minute cells, or cavities, analogous to the lacunae of bone; and the lateral branchlets occasionally terminate in such cavities, which are called the intertubular cells. The diameter of the tubuli at their largest part, averages l-10,000th of an inch ; their smallest branches are im- measurably fine. It is impossible that CAren the largest of them can receive blood, as their diameter is far less than that of the blood-discs; but it is probable that, like the canaliculi of bone, they may absorb nu- trient matter from the vascular surface, Avith which their internal ex- tremities are in relation. 312. In the Teeth of Man and of most Mammalia, we find the central portion IioIIoav; and lined, in the living tooth, by a vascular membrane. This cavity, with its vascular wall, is analogous to a large cancellus or Haversian canal of Bone; and, as Ave shall presently see, it is formed in a similar manner. Upon the walls of the cavity, all the tubuli open; and they radiate from this towards the surface of the upper part of the tooth, as shown in the accom- panying figure. The central cavity is con- tinued as a canal through each fang or root; and the dental tubes in like manner radiate from this, toAvards the surface of the fang.—In the teeth of many of the lower animals, how- ever, Ave find a netAvork of canals extending through the substance of the tooth, instead of a single cavity; and these canals are fre- quently continuous Avith the Haversian canals of the subjacent bone, so that the analogy betAveen the tAvo is complete. From each canal the dentinal tubuli radiate, just in the manner of the canaliculi of bone (§ 295); and thus we may regard a tooth of this kind as repeating, in each of the parts surrounding one of these canals, the structure of the human tooth. 313. The process by which the cellular mass, or pulp, of the dental papilla becomes converted into the Dentine of the perfect tooth, has not been so clearly made out, as to be beyond all question. The folloAving, hoAvever, is the account given of it by Mr. Tomes,* Avho has very care- fully examined it. The dentinal pulp is at first composed of a mass of nucleated cells, held together by a meshwork of delicate fibres and bands constituting an imperfect form of areolar tissue, the interspaces betAveen them being occupied by a homogeneous plasma. In the part which is nearest to the coronal surface of the tooth Avhen calcification is about to commence, the areolar tissue has usually disappeared, and its place is occupied by a finely-granular gelatinous substance. The cells are at first disposed Avithout any regularity in the midst of this; but Oblique section of Dentine of hu- man tooth, highly magnified, show- ing the parallel tubuli. * Lectures on Dental Physiology and Surgery. 188 STRUCTURE AND ENDOAVMENTS OF ANIMAL TISSUES. immediately preceding the conversion of the pulp into dentine, the cells are seen to enlarge, and to become arranged in lines, nearly parallel to each other, and perpendicular to the coronal surface of the tooth ; they then elongate in such a manner, that their extremities come into appo- sition ; and they finally coalesce, so that each toav of cells forms a continuous tube. Whilst this change is in progress, the gelatinous intercellular substance is becoming consolidated by calcareous deposit; Avhich also hardens tbe thick Avails of the tubes, so that only their inte- rior, formed by the coalescence of the original cell cavities, remains uncondensed, thus constituting the dentinal tubuli. This process takes place first on the surface of the pulp, and gradually extends inwards. As the more external and larger cells become hardened, the inner ones increase in size, assume the linear arrangement, and in their turn be- come converted, writh the intercellular substance, into tubular dentine; until at last the great bulk of the pulp is thus transformed, leaving only a comparatively small portion, Avhich, with its nerves and blood-vessels, occupies the central cavity of the tooth. 314. Thus the substance of the outer portion of the pulp is actually converted into dentine, and does not form it by a process of excretion, as Avas formerly supposed. Sometimes it happens that the normal changes are interrupted, and that some of the original meshAvork remains persistent; and it is probably to this that the appearance of large cells, not unfrequently seen in Human teeth (Fig. 53) is due.—Although in the most characteristic form of Dentine, no blood-vessels exist, yet there are certain species, both among Mammals, Reptiles, and Fishes, in which the Dentine is traversed by cylindrical prolongations of the central cavity, conveying blood-vessels into its substance ; and the presence of these medullary canals, giving to the Dentine a vascular character, thus increases its resemblance to bone.—The central portion of the pulp is sometimes converted into a substance still more nearly resembling bone, having its stellate lacunae as well as its vascular canals. This change is normal or regular in certain animals, as in the extinct Iguanodon and Icthyosaurus, and in the Cachalot or Sperm-whale ; and the ossified pulp bears a close resemblance to the bones of the respective animals, although it is not formed in continuity with them. A similar change occurs in the Human tooth ;—sometimes, it would appear, rapidly, as the result of disease; but in general more slowly, increasing gradually with the advance of age. 315. It is not easy to ascertain the amount of nutritive change that takes place in the substance of Dentine, Avhen it is once fully formed. When young animals are fed with coloring-matter, it is found to tinge their teeth, as well as their bones; and if the tooth be in process of rapid formation at the time of the experiment, the progressive calcifica- tion of the pulp from Avithout imvards, is marked by a series of concen- tric lines. Even in the adult, some tinge Avill result from a prolonged use of this substance; and it has been noticed that the teeth of persons who have long suffered from Jaundice sometimes acquire a tinge of bile. These facts sIioav that, even after the complete consolidation of the dentine, it is still pervious to fluids : and that in this manner it may draAv into itself, from the vascular lining of the pulp-caA'ity, a substance SUCCESSION OF TEETH.—STRUCTURE OF ENAMEL. 189 capable of repairing its structure, is proved by the circumstance that a neAV layer of hard matter is occasionally throAvn out upon a surface Avhich has been laid bare by caries. 316. In those simple teeth Avhich consist solely of Dentine, the mode of production already described,—that of the consolidation of a papilla upon the mucous membrane of the mouth,—is all which is requisite. When the formation of the tooth itself is complete, it may remain attached only to the mucous membrane, which is the case in the Shark, or it may groAV downwards, by the addition of neAV dental structure at its base, until it comes in contact Avith the bone of the jaw. Where it is only attached to the mucous membrane, as in the Shark, it is very liable to be torn away; but a new tooth, formed from a distinct papilla, is ready to replace it; and this process is continually repeated, the development of neAV papillae being apparently unlimited. On the other hand, Avhere the root of the tooth comes in contact with the jaAV, it may completely coalesce with it, which is the case in many Fishes, the Ha- versian canals of the bones being continued as medullary canals into the dentine; or it may send long spreading roots into the bone, Avhich are united to it at their extremities. In the classes of Fishes and Rep- tiles (with scarcely any exceptions) the teeth are by no means perma- nent, as among Mammalia; but new teeth are continually succeeding the old ones. The mode in Avhich these teeth originate, by small buds from the capsules of the preceding, will be understood when the capsular development of all the higher forms of the dental apparatus has been described. 317. It is obvious that there is no provision, in the simple calcifica- tion of the dental papilla, for any variations of density, other than those which may result from the different degrees of hardness in the substance of the dentine itself. Now in the teeth of Man and most other Mam- mals, and in those of many Reptiles and some Fishes, Ave find two other substances, one of them harder, and the other softer, than Dentine; the former is termed Enamel; and the latter Cementum or Crusta petrosa. For the development of these, a peculiar modification of the apparatus is requisite. 318. The Enamel is composed of long prismatic cells, exactly resem- bling those of the prismatic shell-substance formerly described (§ 281), but on a far more minute scale ; the diameter of the cells not being more, in Man, than l-5600th of an inch. The length of the prisms corresponds with the thickness of the layer of enamel; and the tAvo sur- faces of this layer present the ends of the prisms, which are usually more or less regularly hexagonal. The quantity of animal matter in the enamel of the adult is extremely minute,—not above 2 parts in 100; and it is only at a very early age that the true character of the animal structure can be distinctly seen. Of the 98 parts of mineral matter in the enamel, 885- consist (according to Berzelius) of phosphate of lime, 8 of carbonate of lime, and 1| of phosphate of magnesia. The course of the prismatic cells is more or less Avavy; and they are marked by nu- merous transverse striae, resembling those of the prismatic shell-sub- stance, and probably originating in the same cause,—the coalescence of a line of shorter cells, to form the lengthened prism. The Enamel is 190 STRUCTURE AND ENDOAVMENTS OF ANIMAL TISSUES. Vertical section of human molar tooth :—1, enamel; 2, ce- mentum or crusta petrosa; 3, dentine or ivory; 4, osseous ex- crescence, arising from hypertro usually destitute of tubuli; but Mr. Tomes has shoAvn that it is occa- sionally penetrated by prolongations of the tubuli of the dentine, and that this peculiarity, Avhich is occasional and abnormal in Man, is cha- racteristic of the teeth of many Marsupials. In density and resisting power, the Enamel far surpasses any other organized tissue, and ap- proaches some of the hardest of mineral sub- stances. In Man, and in Carnivorous animals, it covers the crown of the tooth only, with a simple cap or superficial layer of tolerably uni- form thickness (Fig. 54, 1), which follows the surface of the dentine in all its inequalities; and its component prisms are directed at right angles to that surface, their inner extremities resting in slight but regular depressions on the exterior of the dentine. In the teeth of many Herbivorous animals, however, the Enamel forms (with the cementum) a series of vertical plates, which dip doAvn (as it Avere) into the sub- stance of the dentine, and present their edges, alternately with it, at the grinding surface of the tooth; and there is in such teeth no con- tinuous layer of dentine over the crown. The purpose of this arrangement is evidently to pro- vide, by the unequal wear of these three sub- „ stances,—of Avhich the Enamel is the hardest phy of cementum; 5, cavity 5 6, ' pi osseous ceiis at outer part of and the Cementuui the softest,—for the constant maintenance of a rough surface, adapted to tri- turate the tough vegetable substances on which these animals feed.— The Enamel is the least constant of the Dental tissues. It is more fre- quently absent than present in the teeth of the class of Fishes; it is Avanting in the entire order of Ophidia (Serpents) among existing Rep- tiles ; and it forms no part of the teeth of the Edentata (Sloths, &c.) and Cetacea (Whales) amongst Mammals. 319. The Cementum, or Crusta Petrosa, has the characters of true Bone; possessing its distinctive stellate lacunae and radiating canaliculi. Where it exists in small amount, we do not find it traversed by medullary canals; but, like Dentine, it is occasionally furnished with them, and thus resembles Bone in every particular. These medullary canals enter its substance from the exterior of the tooth; and consequently pass to- wards those, which radiate from the central cavity towards the surface of the dentine, where it possesses a similar vascularity, as was remarka- bly the case in the teeth of the extinct Megatherium. In the Human tooth, however, the Cementum has no such vascularity. It forms a thin layer, which envelopes the root of the tooth, commencing near the ter- mination of the capping of Enamel (Fig. 54, 2). This layer is very sub- ject to have its thickness increased, especially at the extremity of the fangs, by hypertrophy, resulting from inflammation; and sometimes large exostoses are thus formed (Fig. 54, 4), which very much increases the difficulty of extracting the tooth. When the tooth is first developed, the Cementum envelopes its crown, as Avell as its body and root; but FORMATION OF DENTAL CAPSULE. 191 the layer is very thin where it covers the Enamel, and being soft, it is soon Avorn aAvay by use. In the teeth of many Herbivorous Mammals, it dips down with the Enamel to form the vertical plates of the interior of the tooth ; and in the teeth of the Edentata as Avell .as of many Rep- tiles and Fishes, it forms a thick continuous envelope over the whole of the surface, until worn away at the crown. 320. The.development of these additional structures is provided for by the enclosure of the primitive papilla, from Avhich the Dentine is formed, within a Capsule, which, at one period, completely covers it in: betAveen the inner surface of the capsule, and the outer surface of the dentinal papilla, a sort of epithelium is developed, by the calcification of which, the Enamel is formed; and the Cementum is generated by the conversion of the capsule itself into a bony substance. The pro- cesses by Avhich this capsular investment is produced, and the tooth completed and evolved, -will now be briefly described, as they occur in the Human foetus. 321. The dental papillae begin to make their appearance, at about the seventh week of embryonic life, upon the mucous membrane cover- ing the bottom of a deep narrow groove (Fig. 55, a,) that runs along the edge of the jaw (Fig. 55, b); and during the tenth week, processes from the sides of this "primitive dental groove," particularly the ex- ternal one, begin to approach one another,#so as to divide it, by their meeting, into a series of open follicles, at the bottom of which the pa- pillae may still be seen. At the thirteenth week all the follicles being completed, the papillae, which were at first round blunt masses of cells, begin to assume forms more characteristic of the teeth which are to be developed from them; and by their rapid growth, they protrude from Fig. 55. Successive stages of the development of the deciduous or temporary teeth, and of the origin of the sacs of the permanent set. the mouths of the follicles (Fig. 55, c). At the same time, the edges of the follicles are lengthened into little valve-like processes, or oper- cula, which are destined to meet and form covers to the follicles (Fig. 55, d). There are tAvo of these opercula in the Incisive follicles, three for the Canines, and four or five for the Molars. And by the fourteenth Aveek, the tAvo lips of the dental groove meet over the mouths of the follicles, so as completely to enclose each papilla in a distinct capsule (Fig. 55, e). At this period, before the calcification of the primitiA'e pulps commences, a provision is made for the production of the second or permanent molars; Avhose capsules originate in buds or offsets from the upper part of the capsules of the temporary or milk-teeth (Fig. 55,/). 192 STRUCTURE AND ENDOAVMENTS OF ANIMAL TISSUES. These offsets are in the condition of open follicles, communicating Avith the cavity of the primitive tooth; but they are gradually closed in, and detached altogether from the capsules of the milk-teeth (Fig. 55, WSM ii^Ss^; iP ==\i ^^^^^i •,! WM ~J-\i_7\(.j^-— e. Dorsal ganglion of Sympathetic nerve of Mouse;—a, b, cords of connexion with adjacent sympathetic ganglia; c, c, c, c, branches to the viscera and spinal nerve; d, ganglionic globules or ceUs; e, nervous fibres crossing the ganglion. course into relation with the vascular matter, which occupies the inte- rior of the ganglion, in one or more of the modes already specified (§ 378). Some of the fibres may terminate in the cells of the ganglion, 222 STRUCTURE AND ENDOWMENTS OF ANIMAL TISSUES. and new ones may originate in them; so that there is no constant cor- respondence between the number of fibres which enter a ganglion, and of those who pass out of it.—The only exception to the general fact, that the vesicular matter occupies the centre of the ganglia, occurs in the brain of Vertebrata, in which it is chiefly disposed on the exterior, forming the cortical envelope. The reason for this variation is probably to be found in the very large amount of this substance, which the brain of the higher Vertebrata contains; and in the necessity of the free access of blood-vessels to it, which is provided for by a great extension of its surface beneath the investing vascular membrane (pia mater), more readily than it could be in any other mode. 381. But the vesicular matter is not found in the central masses only of the Nervous system ; for it presents itself also at those parts of the surface or periphery which are peculiarly destined to receive the im- pressions that are to be conveyed to the central organs. Thus on the expansion of the optic-nerve which forms the retina, there is a distinct layer of ganglionic corpuscles, or nerve-cells, with a minute plexus of vessels, possessing all the essential characters of the vesicular substance of the brain ; and something of the same kind has been seen in con- nexion with the corresponding expansion of the olfactive and auditory nerves. Moreover, the study of the history of the development of these organs has shown that the vesicular matter of the retina is an offshoot (so to speak) from that of the optic ganglion, that of the labyrinth of the ear being in like manner an offshoot from that of the auditory gan- glion. Thus it is obvious that the fibres of the connecting nerve are interposed betAveen the cells of the peripheral and those of the central organs, for the sake of preserving that connexion between them which would otherwise have been interrupted; and that the vesicular matter is the active agent in the origination of those changes which take place as a consequence of sensory impressions, whilst for the conduction of such changes, the fibrous structure is alone required. 382. The ultimate distribution of the nerve-fibres in the skin and tongue, however, has not been so clearly made out, nor can their rela- tion to cells be distinctly traced. These fibres are distributed, for the most part, to the papillse, in which they can be frequently seen to form Fig. 72. Fig. 73. Distribution of the tactile nerves at the surface of the lip; as seen in a thin perpendicular section of the skin. loops (Fig. 72), accompanied by similar loops of blood-vessels (Fig. 73): but no such loops can be seen in the fungiform papilke of the tonguej CHEMICAL COMPOSITION OF NERVOUS MATTER. 223 the continuity of whose nerve-fibres cannot be distinguished near their apices. Here, however, the history of the development of the nervous plexuses in the skin seems to show, that the fibres may be considered as commencing in a peripheral ganglionic expansion ; for it has been shown by the observations of Prof. Kolliker upon the tail of the tadpole, that the nervous plexuses are formed in the same manner as the capillary network; namely, by the inosculation of the prolongations of radiating cells, whose centres are at a considerable distance from each other. Hence it is probable that cell-nuclei, or some equivalent centres of force, remain in connexion with the peripheral nerve-fibres of the skin and tongue, as with those of other sensory surfaces. Sometimes the ultimate plexuses seem to be formed in the Skin, as in Muscle, by the subdivision of the primitive fibres (§ 341). 383. We have now to speak briefly of the Chemical Composition of the Nervous matter;—a consideration which will be presently shoAvn to be of much importance. As formerly remarked (§ 7), the vital activity of a tissue is usually greater, as the proportion of its solid to its fluid contents is less; and this rule holds good most strikingly in regard to the Nervous substance, the vital activity of which is far greater than that of any other tissue, and the solid matter of which usually consti- tutes no more than a fourth, and occasionally does not exceed an eighth of its entire weight. The proportion of water is greatest in infancy and least in middle life; and it has been observed to be under the average in idiots. Of the solid matter of the brain, about a third consists of fibrine or albumen; Avhich is probably the material of the membrane of the tubuli, as Avell as of the tissue that connects them.—It is chiefly Avith the Fatty matter, Avhich constitutes about a third of the solid substance, that the attention of Chemists has been occupied. This is stated by M. Fremy (one of the most recent analysts) to contain, besides the ordinary fatty matters, and Cholesterine or biliary fat, tAvo peculiar fatty acids, termed the Cerebric and the Oleo phosphoric. Cerebric acid, when puri- fied, is white, and presents itself in crystalline grains. It contains a small proportion of Phosphorus; and differs from the ordinary fatty matters in containing Nitrogen, as also in containing twice their propor- tion of oxygen. Oleo phosphoric acid is separated from the former by its solubility in ether; it is of a viscid consistence; but when boiled for a long time in Avater or alcohol, it gradually loses its viscidity, and resolves itself into a pure oil, Avhich is elaine, while phosphoric acid remains in the liquor. The proportion of phosphorus in the brain is considerable ; being from 8 to 18 parts in 1000 of the whole mass, or from l-20th to l-30th of the whole solid matter. It seems to be unusually deficient in the brain of idiots. The remaining; third and sometimes more, is composed of a substance termed Osmazome (Avhich seems to be a proteine-compound in a state of decomposition), together Avith saline matter. No satisfactory examination has yet been made into the com- parative composition of the vesicular and fibrous substances ; but accord- ing to Lassaigne, the former contains much more Avater than the latter, and little colorless fat, but nearly 4 per cent, of red fat, which does not exist in the other. 384. Various circumstances lead to the belief, that the Nervous tissue, 224 STRUCTURE AND ENDOAVMENTS OF ANIMAL TISSUES. during the whole period of active life, is continually undergoing changes in its substance, by decay and renewal. We know that, after death, it is one of the first of all the animal tissues to exhibit signs of decompo- sition ; and there is no reason to suppose, that this tendency is absent during life. Hence for the simple maintenance of its normal character, a considerable amount of nutritive change must be required. But many circumstances further lead to the conclusion, that, like all other tissues actively concerned in the vital operations, Nervous matter is subject to a waste or disintegration, which bears an exact proportion to the activity of its operations;—or, in other words, that every act of the Nervous system involves the death and decay of a certain amount of Nervous matter, the replacement of which will be requisite in order to maintain the system in a state fit for action. We shall hereafter see, that there are certain parts of the Nervous system, particularly such as put in action the respiratory muscles, which are in a state of unceasing, though moderate, activity; and in these, the constant nutrition is suffi- cient to repair the effects of the constant decay. But those parts, which operate in a more powerful and energetic manner, and which therefore waste more rapidly when in action, need a season of rest for their repa- ration. Thus a sense of fatigue is experienced, when the mind has been long acting through its instrument—the brain ; indicating the necessity for rest and reparation. But when sleep, or cessation of the cerebral functions, comes on, the process of nutrition takes place with unchecked energy, counterbalances the results of the previous waste, and prepares the organ for a reneAval of its activity. In the healthy state of the body, when the exertion of the nervous system by day does not exceed that, which the repose of the night may compensate, it is maintained in a condition which fits it for constant moderate exercise; but unusual demands upon its poAvers,—whether by the long-continued and severe exercise of the intellect, by excitement of the emotions, or by the com- bination of both in that state of anxiety which the circumstances of man's condition too frequently induce,—occasion an unusual waste, which requires, for the complete restoration of its poAvers, a prolonged repose. 385. There can be no doubt that (from causes which are not known), the amount of sleep required by different persons, for the maintenance of a healthy condition of the nervous system, varies considerably; some being able to dispense Avith it, to a degree which would be exceedingly injurious to others of no greater mental activity. Where a prolonged exertion of the mind has been made, and the natural tendency to sleep has been habitually resisted, by a strong effort of the Avill, injurious results are sure to follow. The bodily health breaks down, and too frequently the mind itself is permanently enfeebled. It is obvious that the nutrition of the Nervous system becomes completely deranged; and that the tissue is no longer formed, in the manner requisite for the dis- charge of its healthy functions. 386. As the amount of Muscular tissue that has undergone disinte- gration, is represented (other things being equal), by the quantity of urea in the urine, so do we find that an unusual waste of the nervous matter is indicated by an increase in the amount of phosphatic deposits. AVASTE AND RENEAVAL OF NERVOUS SUBSTANCE. 225 No others of the soft tissues contain any large proportion of phospho- rus ; and the marked increase in these deposits, Avhich has been con- tinually observed to accompany long-continued wear of mind, whether by intellectual exertion, or by anxiety, can scarcely be set down to any other cause. The most satisfactory proof is to be found in cases, in Avhich there is a periodical demand upon the mental powers; as, for example, among clergymen, in the preparation for, and discharge of, their Sunday duties. This is found to be almost invariably followed by the appearance of a large quantity of the phosphates in the urine. And in cases, in Avhich constant and severe intellectual exertion has impaired the nutrition of the brain, and has consequently weakened the mental poAver, it is found that any premature attempt to renew the activity of its exercise, causes the reappearance of the excessive phos- phatic discharge, which indicates an undue waste of nervous matter. 387. As the disintegration of the Nervous* System is thus propor- tional to its exercise, so must its reparation make a corresponding de- mand upon the nutritive processes. And accordingly we find, that it is very copiously supplied with blood-vessels; and that the amount of food, appropriated to its maintenance in an active condition, is very considerable. This Ave know from the fact, that persons of active minds, but sedentary bodily habits, commonly require nearly as much food as those, in whom the waste of the Muscular system is greater, and that of the Nervous system less, in virtue of their bodily activity and the less energetic operation of their minds. 388. The nerve-fibres appear to originate, according to the observa- tions of Prof. Kolliker, in cells which become fusiform by elongation (§ 193), and Avhich then coalesce at their extremities; and these seem to increase, after the first formation of the trunks, by the longitu- dinal subdivision of fusiform cells Avhich had not previously undergone complete metamorphosis into fibres, as well as by the development of cells de novo. The nuclei of the original cells may be frequently seen in the nerve-tubes at a later period, lying between their membranous walls and the substance deposited in their interior. The earliest condi- tion of both forms of nerve-fibre appears to be precisely the same; but the gelatinous remains in a state nearly resembling this; whilst the tubular is developed into a higher form. Various gradations, indeed, may be traced between the two. The vesicular matter appears to be in a state of continual change, as is the case with all cells whose functions are active. The appearances observed by Henle in the cortical sub- stance of the brain lead to the belief, that there is as continual a suc- cession of nerve-cells, as there is of epidermic cells; their development commencing at the surface, where they are most copiously supplied with blood-vessels from the investing membrane, and proceeding as they are carried toAvards the inner layers, Avhere they come into more immediate relation with the fibrous portion of the nerve-structure. This change of place is probably due to the continual death and decay of the mature cells, where they are connected Avith the fibres; and the constant pro- duction of neAV generations at the external surface,—thus carrying the previously-formed cells inwards, in precisely the same manner that the epidermic cells are progressh'ely carried outwards. r 15 226 STRUCTURE AND ENDOWxMENTS OF ANIMAL TISSUES. 389. The regeneration of Nervous tissue that has been destroyed, takes place very readily in continuity Avith that Avhich is left sound. This may be more easily proved by the return of the sensory and motor endowments of the part, whose nerves have been separated, than by microscopic examination of the reunited trunks themselvef, which is not ahvays satisfactory. All our knoAvledge of the functions of the nervous system leads to the belief, that perfect continuity of the nerve- tubes is requisite for the conduction of an impression of any kind, whether this be destined to produce motion or sensation; and various facts, well knoAvn to Surgeons, prove that such restoration may be com- plete. In the various operations which are practised for the restoration of lost parts, a portion of tissue removed from one spot is grafted, as it were, upon another; its original attachments are more or less completely severed,—frequently altogether destroyed,—and new ones are formed. Now in such a part, so,long as its original connexions exist, and the new ones are not completely formed, the sensation is referred to the spot from which it was taken ; thus when a new nose is made, by partly detaching and bringing down a piece of skin from the forehead, the patient at first feels, when anything touches the tip of his nose, as if the contact Avere really Avith his forehead. After time has been given, however, for the establishment of new connexions Avith the parts into whose neighborhood it has been brought, the old connexions of the grafted portion are completely severed; and an interval then ensues, during which it frequently loses all sensibility; but after a time its power of feeling is restored, and the sensations received through it are referred to the right spot. A more familiar case is the regeneration of Skin, containing sensory nerves, which takes place in the well-managed healing of wounds involving loss of substance. Here there must ob- viously be, not merely a prolongation of the nerve-tubes from the sub- jacent and surrounding trunks, but also a formation of new sensory papillae. A still more striking example of the regeneration of Nervous tissue, however, is to be found in those cases (of which there are now several on record), in which portions of the extremities, that have been completely severed by accident, have been made to adhere to f.he stump; and have, in time, completely recovered their connexion with the Ner- vous as with the other systems, as is indicated by the restoration of their sensory and motor endowments.—Of the degree in which the vesi- cular substance of the Nervous system may be regenerated, Ave have no certain knowledge; but there can be little doubt, from the activity of its usual nutritive changes, that a complete reproduction may be effected in cases of loss of substance, where it can commence from a neighboring mass of the same tissue. 390. We have now to inquire into the conditions, under which the peculiar properties of the Nervous System are manifested in an active form; and it will first be desirable to explain, somewhat more in detail, the nature of the different operations to which it is subservient. These operations present themselves in their most complex form, in Man and the higher animals; but they may often be most satisfactorily studied in the lower. In the first place, when an impression is made upon any part of the surface of the body by mechanical contact, by heat, elec- PROPERTIES OF NERVOUS SYSTEM. 227 tricity, or any other similar agent,—or upon the organs of special sense (the eye and ear, the nose and tongue), by light or sound, by odorous or sapid bodies,—these impressions, in the healthy and wakeful state of the Nervous system, are felt as sensations ; that is, the mind is rendered conscious of thern. Now there can be no doubt that the mind is imme- diately influenced, not by the impression in the remote organ, but by a certain change in the condition of the brain, excited or aroused by that Avhich has originated elsewhere. For if the communication with the brain be cut off, no impression on the distant parts of the nervous system is felt, notwithstanding that the mind remains perfectly capable of re- ceiving it. The mind, then, is only rendered conscious of external objects, by the influence which they exert upon the brain, or upon a certain part of it, Avhich, being the peculiar seat of sensation, is called the sensorium. Hence we recognise, in the process by which the mind is rendered con- scious of external objects, three distinct stages; first, the reception of the impression at the extremities of the sensory nerve ; second, the conduction of the impression, along the trunk of the nerve, to the sensorium; third, the change excited by it in the sensorium itself, through Avhich sensation is produced. Here, then, the change in the condition of the nervous system commences at the circumference, and is transmitted to the centre; and the fibres which are concerned in this transmission are termed sensory. 391. On the other hand, when an emotion, an instinctive impulse, or an act of the will, operates through the central organs to produce a muscular contraction, the first change is in the condition of the vesicular substance of those organs. The influence of this change is transmitted by the motor nerves to the muscles, among which they are distributed ; and the desired movement is the result. Here, too, we have at least three stages; first, the origination of the change by an impression act- ing on the central organs ; second, the conduction of that change along the motor nerves; and third, the stimulation of the muscles to contrac- tion. But the operation here commences at the centre ; and the effects of the change in the brain are transmitted to the circumference, by a set of nervous fibres which are termed motor. The complete distinct- ness of these tAvo classes of fibres Avas first established by Sir C. Bell. It is best seen in the nerves of the head, of which some are purely sen- sory, and others purely motor; but it may also be clearly proved to exist at the roots of the spinal nerves (although their trunks possess mixed endowments), the posterior being sensory, whilst the anterior are motor. 392. But although sensations can only be felt through the brain, and voluntary motions can only be produced by an action of the mind through the same organ, yet there are many changes in the animal body, in which the nervous system is concerned, Avhich yet do not involve the operation of the brain, being produced without our consciousness being necessarily excited, and Avithout any act of the will, or even in opposi- tion to its efforts. Of these actions, the spinal cord of Vertebrata, and its prolongation within the cranium, are the chief instruments; in the Invertebrate animals, they are performed by various ganglia, which are usually disposed in the neighborhood of the organs to which they 228 STRUCTURE AND ENDOWMENTS OF ANIMAL TISSUES. minister. If the spinal cord of a Frog be divided in its back, above the crural plexus, so as entirely to cut off the nerves of the lower ex- tremities from connexion Avith the brain, the animal loses all voluntary control over these limbs, and no sign of pain is produced by any injury done to them. But they are not thereby rendered motionless; for various stimuli applied to the limbs themselves will cause movements in them. Thus if the skin of the foot be pinched, or if a flame be applied to it, the leg will be violently retracted. Or, if the cloaca be irritated by a probe, the feet will endeavor to push aAvay the instrument. We have no reason hence to believe, that the animal feels the irritation, or intends to execute these movements in order to escape from it; for motions of a similar kind are exhibited by men, who have suffered injury of the lower part of the spinal cord, and who are utterly unconscious, either of the irritation which their limbs receive, or of the actions which they perform. 393. We are not to suppose, however, that the stimulus acts at once upon the muscles, without the nervous system being concerned at all; throwing them into contraction by its direct influence. For it is quite certain, that unless the nervous trunks remain continuous with the spinal cord, and unless the part of the spinal cord with which they are connected remains sound (although cut off from connexion with the parts above, and with the brain), no action will result. If the trunks be divided, or either of the roots by which they are connected with the spinal cord be severed, or the lower portion of the spinal cord itself be injured, no stimulation will cause the muscular move- ments just described. A very good example of this necessity for the completeness of the nervous trunks, which convey impressions to and from the central organ, is found in the movements of the iris, for the contraction and dilatation of the pupil. Here the stimulus of light upon the retina gives rise to a change in the condition of the optic nerve; which, being transmitted to a certain portion of the encephalon with which that nerve is connected, excites there a motor impulse; and this impulse is conveyed through a distinct nerve (a branch of the third pair) to the iris, occasioning contraction of the pupil. Every one knows that this adjustment of the size of the pupil to the amount of light, is effected without any exertion of the will on his OAvn part, and even without any consciousness that it is taking place. It is performed, too, during profound sleep; when the influence of light upon the retina excites no consciousness of its presence,—when no sensation, therefore, is produced by it. 394. The class of actions thus performed, is termed reflex ; and we see that every such action involves the folloAving series of changes. In the first place, an impression is made upon the extremity of a nerve, by some external agent; just as when sensation is to be pro- duced. Secondly, this impression is transmitted by a nervous trunk to the spinal cord in Vertebrata, or to some ganglionic mass which answers to it in the Invertebrata. But instead of being communicated by its means to the mind, and becoming a sensation, it immediately and necessarily executes a motor impulse; Avhich is reflected back as it were to certain muscles, and, by their contraction, gives rise to a CONDITIONS OF NERVOUS ACTIONS. 229 movement. We shall hereafter see, that nearly all those movements in the animal body, which are immediately connected with the mainte- nance of the organic functions,—such as those of respiration, degluti- tion (or swallowing), the expulsion of the faeces, urine, and foetus, &c, —are performed in this manner. 395. Noav there is strong reason to believe that the changes which take place in the nervous trunks are of the same nature, whatever may be the source from which they proceed,—whether, for example, the movement is simply reflex, whether it proceed from a mental emotion, or whether it be executed in obedience to an act of the will. It was formely supposed that all the afferent or centripetal fibres pass up to the Brain, and that all the efferent or centrifugal fibres pass down from the same organ; the Spinal Cord being looked upon as little else than a bundle of nerves. It is now known, however, that by far the greater part of the fibres of any trunk terminates in the central organ, to Avhich that trunk at first proceeds; and that the Spinal Cord may be consi- dered as a series of such ganglionic centres, each receiving the afferent fibres, and giving origin to the efferent of its own segment. So, again, the special sensory nerves, the olfactive, optic, auditory, and gustative, terminate in their own ganglionic centres, which lie at the base of the brain, in immediate connexion with the summit of the spinal cord, and Avhich are quite independent of the cerebrum. The apparatus for receiving impressions, and for originating motions, is thus complete in itself; and the addition of the cerebrum does not make any essential difference in its operations, save that this sensori-motor apparatus (as it may be termed) is made to act through its means as the agent of the mind, in addition to its functions as the instrument of the automatic movements. We shall hereafter see (chap. XII.), that the difference betAveem Instinct and Intelligence is closely connected with the development of the cerebrum ; but that this organ, even in that highest grade of development which it possesses in Man, has no other connexion with the sensory organs than that which it acquires through its relation with the sensory ganglia, and has no more power of exciting muscular movement, than by playing (so to speak), upon the spinal cord, whose efferent fibres respond to its mandates, just as they Avould do to the stimulus, of an impression primarily acting through that organ. 396. Of the mode by which the effects of changes in one part of the Nervous system, are thus instantaneously transmitted to another, nothing Avhatever is known. There is evidently a strong analogy betAveen this phenomenon, and the instantaneous transmission of the Electric poAver along good conductors ; but the relation is much more intimate than this, for Electricity is capable of exciting Nerve-force whilst, conA'ersely, Nerve-force can excite Electricity. Thus a very feeble galvanic current transmitted along a motor nerve, serves to excite con- tractions in the muscles supplied by it; and in like manner, a galvanic current transmitted along any of the sensory nerves, give rise to a sensation of the kind to Avhich the nerve ministers. Moreover we shall hereafter see, that certain animals are capable of generating Electric power in a very remarkable manner (chap, x.) ; and that the 230 STRUCTURE AND ENDOAVMENTS OF ANIMAL TISSUES. nervous force is essentially concerned in this operation. But, on the other hand, it is quite certain that the influence transmitted along the nerves of the living body is not ordinary electricity ; for all attempts to procure manifestations of electric changes in the state of nerves, that are acting most energetically on muscles, have completely failed; and a nerve remains capable of conveying the influence of electricity, when it has been rendered unable to transmit the influence of the brain, as by tying a ligature round it, or by tightly compressing it between the forceps, Avhich gives no interruption to the one agency, while it completely checks the other.—Notwithstanding, then, the strong analogy which exists between these two powers, we are not Avarranted in regarding them as identical; but they have towards each other that relation of reciprocity, which exists between Electricity and Heat, or between Electricity and Magnetism, each being convertible into the other in a certain definite ratio (§ 53). 397. It is more desirable, however, that we should understand the conditions under which the phenomena of the Nervous System take place, than that we should spend much time in discussing the identity of its peculiar powers with any others in Nature. The conducting power of the nervous fibres appears to remain with little decrease for some time after death, especially in cold-blooded animals; for we can, by pinching, pricking, or otherwise stimulating the motor-trunk, give rise to contractions in the muscles supplied by them, exactly as during life. This poAver is much lessened by the influence of narcotics; so that if a nen^ous trunk be soaked in a solution of opium, belladonna, or other powerful narcotic, it ceases to be able to convey the effects of stimuli to the muscles, some time before the muscles themselves lose their con- tractile power. On the other hand, it seems to be exalted by various irritating influences; so that, when the nervous trunk has been treated with strychnia, or when it has been subjected to undue excitement in other ways, a very slight change is magnified (as it were) during its transmis- sion, and produces effects of unusual intensity. 398. Now although the conducting power of the fibrous structure will continue for a time, after the circulation through it has ceased, the peculiar endowments of the vesicular substance, by which it originates the changes which the former transmits, are only manifested, when blood is moving through its capillaries. Thus if the circulation through the brain cease but for a moment, total insensibility, and loss of the power of voluntary motion, immediately surpervene. The brain is supplied with blood through four arteries,—the two internal carotids, and the two vertebrals ; and by the communication of these with each other through the circle of Willis, the circulation will still be kept up, if only one of them should convey blood into the cavity of the cranium. Hence it is necessary that the flow of blood should be checked through all of them, in order that the functions of the brain should be suspended; and the suspension is then complete and instantaneous. The best method of effecting this was devised by Sir Astley Cooper. He tied both the carotid arteries in a dog : which, for the reasons just mentioned, did not produce any decided influence on the functions of the brain, the circulation being kept up through the vertebrals. But upon compress- EFFECTS OF STIMULANTS UPON NERVOUS POWER. 231 ing the latter, so as to suspend the flow of blood through them, imme- diate insensibility, and loss of voluntary power, were the result. When the compression Avas taken off, the animal immediately returned to its usual state ; and again become suddenly insensible, Avhen the pressure was reneAved. Although the functions of the brain were thus suspended, those of the spinal cord were not; as was shown by the occurrence of convulsive movements. But in the state called Syncope, or fainting, the suspension of the circulation, by a failure in the heart's action, causes an entire loss of poAver in both these centres ; and a complete cessation of muscular movement is the result. This condition may come on instantaneously, under the influence of powerful mental emo- tion, or of some other cause, which acts primarily in suspending the heart's action, and consequently in checking the circulation ; the insen- sibility, and loss of muscular power, are secondary results, depending upon the suspension of the powers of the nervous centres, consequent upon the cessation of the Aoav of blood through them. 399. The due activity of the vesicular nervous matter is not only dependent upon a sufficient supply of blood, but it requires that this blood should be in a state of extreme purity; for there is no tissue in the body, whose functions are so readily deranged, by any departure from the regular standard in the "circulating fluid,—Avhether this con- sist in the alteration of the proportions of its normal ingredients, or in the introduction of other substances which have no proper place in it. One of the most fertile sources of disturbance in the action of the brain, consists in the retention of substances within the blood, which ought to be excreted from it. We shall hereafter see, that three of the largest and most important organs in the body,—the lungs, the liver, and the kidneys,—have it for their special office, to separate from the circulating fluid the products of the decomposition, which is continually taking place in the body; and thereby to maintain its purity and its fitnesss for its important functions. Now if these, from any cause, even partially fail in their office, speedy disturbance of the functions of the nervous centres is the result. Thi^s if the lungs do not purify the venous blood of its impregnation of carbonic acid, or restore to it the proper proportion of oxygen, the functions of the brain^ are seriously affected. The sensations become indistinct, the Avill loses its control over the muscles, giddiness and faintness come on, and at last complete insensibility supervenes. Corresponding symptoms occur, though to a less serious degree, when the excretion of carbonic acid is but slightly impeded. Thus Avhen a number of persons are shut up in an ill-ventilated apartment, for a sufficient length of time to raise the proportion of carbonic acid in the air to 1 or 2 per cent., the continued purification of their blood by respiration is but insufficiently performed, for reasons which will be stated hereafter (chap, viii.) ; and the car- bonic acid accumulates in their blood in a sufficient degree, to produce headache and obtuseness of the mental powers.—Similar results take place, as will be shown hereafter, from the retention of the substances which ought to be drawn off by the liver and kidneys: tl ese, when they accumulate in even a trifling degree, produce torpor of the functions of the brain ; and when their proportion increases, complete cessation of 232 STRUCTURE AND ENDOWMENTS OF ANIMAL TISSUES. its powers is the result, their action being precisely that of narcotic poisons. Various substances introduced into the blood may exert similar influences; depressing the activity of the vesicular substance of the nervous centres, and consequently producing torpidity, not merely in regard to the reception of impressions, and the performance of volun- tary motions, but also in the mental operations generally. 400. On the other hand, various conditions of the blood, especially those depending on the presence of certain external agents produce an undue energy in the functions of the nervous centres; which energy, however, is almost invariably accompanied by irregularity, or want of balance among the different actions. Of this we have a familiar example in the operation of alcohol. Its first effect, when taken in moderate quantity, is usually to produce a simple increase in the activity of the cerebral functions. A further dose, however, occasions not merely an increase, but an irregularity ; destroying that power of self-control, which is so important a means of balancing the different tendencies in the healthy condition of the mind. And a still larger dose has the effect of a narcotic poison; producing diminution or suspension of activity in all the functions of the brain. In some persons, this is the mode in which the alcohol acts from the first, its stimulating effects being altogether wanting.—A similar activity is usually produced by the respiration of the Nitrous Oxide, which seems to increase all the powers of the mind, save that of self-control, which it diminishes ; the individual, while under its influence, being the slave of his impulses, which act on his muscular system with astonishing energy. Very analogous to this, is the incipient stage of mania; which is simply an undue energy of the cerebral func- tions at first in some degree under the control of the will, but after- wards, increasing to an extent that renders the individual completely powerless over himself; and showing itself in the intensity of the sensa- tions produced by external objects, in the vividness of the trains of thought (which, being entirely uncontrolled, succeed each other with apparent irregularity, though probably according to the laAvs of associa- tion and suggestion), and in the violence of the muscular actions. Such a state may continue for some time, without the intervention of sleep; but the subsequent exhaustion of nervous power is proportioned to the duration of the excitement; and frequent attacks of mania almost invariably subside at last into imbecility. 401. In these cases of undue excitement, there is obviously an increase in the supply of blood to the head; as indicated by the suffusion of the face, the injection of the conjunctiva, the throbbing at the temples, the pulsation of the carotids; and we find that measures which diminish the activity of the circulation through the brain, are those most effec- tual in subduing the excitement. But it does not at all follow, that this undue action of the brain should be connected with an excess in the whole amount of nutritive material, and should require general deple- tion for its treatment. In fact, a very similar class of symptoms may present itself under two conditions of an entirely opposite kind,— inflammation, accompanied with an increase in the proportion of fibrine in the blood, and requiring treatment of a lowering kind,—and irritation, depending on a state of blood in which there is a deficiency of solid DEPENDENCE OF NERVOUS POWER ON SUPPLY OF BLOOD. 233 materials, and requiring a strengthening and even a stimulating regimen. The skill of the practitioner is often put to the test, in the due discrimination between these states. 402. The preceding examples mark the influence of various causes upon the actions of the vesicular matter of the brain; others might be adduced to show, that the vesicular substance of the spinal cord is also liable to have its poAvers depressed or excited; but these will be best adverted to hereafter, when the distinct functions of that organ are under consideration (chap, xii.) We may simply notice, that the stimu- lating effect of Strychnia is peculiarly and most remarkably exerted upon the vesicular substance of the spinal cord; and that a correspond- ing state, in which violent convulsive actions are excited by the most trifling causes, sometimes presents itself as a peculiar form of disease, named Tetanus, which may be either idiopathic, depending probably upon a disordered condition of the blood, or traumatic, consequent upon the irritation of a wound. 403. But, as formerly remarked, it is not in the Nervous centres only, that changes originate. Whenever an impression is made upon the surface of the body, or upon the organs of special sense, Avhich, being conducted to the nervous centres, either excites a sensation in the brain (§ 390), or a reflex action through the spinal cord (§ 392), the reception and propagation of such impression at the extremities of the sensory nerves requires a set of conditions of the same kind with those, Avhich we have seen to exist in the nervous centres. In fact, if Ave re- gard the course of the motor nerves as commencing in the nervous centres and terminating in the muscles, we may with equal justice con- sider that of the sensory nerves as originating in their peripheral extremi- ties, and terminating in the sensorium. And, as already stated (§ 381), precisely the same kind of vesicular structure exists in some (probably in all) of the peripheral expansions of the sensory nerves, as makes up the gray substance of the brain and spinal cord. Noav it is easily shown, that the circulation of the blood through these parts is just as necessary for the original reception of the impressions, as is the circulation through the brain to their reception as sensations, and to the origination of motor impulses by an act of the will. We find that anything Avhich retards the circulation through a part supplied by sensory nerves, diminishes its sensibility; and that if the flow of blood be completely stagnated, entire insensibility is the result. A familiar example of this is seen in the effects of prolonged cold; which, by diminishing, and then entirely checking, the Aoav of blood through the skin, produces first numbness, and then complete insensibility of the part. This result, however, may be partly due to the direct influence of the cold upon the nerve-vesicles themselves; depressing their peculiar vital powers (§ 97). The same effect is produced, however, when the supply of blood is checked in any other way; as, for example, by pressure on the artery, or by obstruction in its interior. Thus when the main artery of a limb is tied, numbness of the extremities is immediately perceived; and this continues, until the circulation is re-established by the collateral branches, when the usual amount of sensibility is restored. Again, in the gan- grene which depends upon obstruction of the arterial trunks by a 234 . STRUCTURE AND ENDOAVMENTS OF ANIMAL TISSUES. fibrinous clot in their interior, diminution of sensibility, consequent upon the insufficient circulation, is one of the first symptoms. 404. On the other hand, increased circulation of blood through a part produces exaltation of its sensibility; that is, the ordinary im- pressions produce changes of unusual energy in its sensory nerves, This is particularly evident in the increased sensibility of the genital organs of animals during the period of heat; and in those of Man, Avhen in a state of venereal excitement. Moderate warmth, friction, exercise, and other causes which increase the circulation through a part, also augment its sensibility; and this augmentation is one of the most constant indications of that state of determination of blood, or active congestion, which usually precedes inflammation, and which exists in the parts surrounding the centre of inflammatory action. But it must be borne in mind, here as elseAvhere (§ 401), that such exaltation of func- tion in a limited part, is quite consistent with general debility; and in fact we may often observe, that the tendency of such local affections is particularly great, when the blood is in a very poor condition. (See chap, v.) 405. To sum up, then, we may compare the vesicular substance, wherever it exists, to a galvanic combination: the former being capable of generating nervous influence, and transmitting it along the fibrous structure, to the part on which it is to operate; in the same manner as the latter generates electric power, and transmits it along the con- ducting wires, to the point at which it is to effect a decomposition or any other change. In one of the most perfect forms of the galvanic battery (that invented by Mr. Smee), although the metals remain in- serted in the acid solution, and are consequently always ready for action, no electricity is generated until the circuit is complete; and the ivaste of the zinc, produced by its solution in the acid, is therefore exactly proportional to the electric effects to which it gives rise. The condition of the nervous system, in the healthy and waking state, bears a close analogy to this; for it is in a state constantly ready for action, but waits to be excited; and its waste is proportional to the activity of its function.—The vesicular matter, diffused over the surface of the body, is inactive, until an impression is made upon it by some external agent; but a change then takes place in its condition (of which we know no more, than that the presence of arterial blood and a certain amount of warmth are necessary for it), which is transmitted to the central organs by the sensory trunks. It Avould appear that the excite- ment of this change has a tendency to increase the afflux of blood to the part; thus when a lozenge or some similar substance is allowed to lie for a time in contact with the tongue, or with the side of the mouth, a roughness is produced, which is due to the erection of the sensory papillae, by the distension of their blood-vessels.—On the other hand, the change in the vesicular matter of the central organs, by which motion is produced in the distant muscles, may be excited either by the stimulus conA'eyed by the afferent nerves (as in reflex action, § 392), or by an act of Mind. This act may be voluntary, originating in the will; or it may be instinctive or emotional, resulting from certain states of CONNEXION BETAVEEN THE MIND AND BRAIN. 235 mind excited by sensations, and altogether independent of the will. Of the mode m Avhich the mind thus acts upon the nervous system, we know nothing whatever, and probably never shall be informed in our present state of being But it is sufficient for us to be aware of the physiological fact of the peculiar connexion between the mind and the brain ; a connexion so intimate, as to enable the mind to receive through tlm body a knowledge of the condition of the Universe around it, and to impress on the body the results of its own determinations; and of such a nature, that the regularity of the working of the mind itself is dependent upon the complete organization of the brain in the first in- stance, upon the constant supply of pure and well elaborated blood, and upon all those^ influences which favor the due performance of the nutritive operations in general. BOOK II. SPECIAL PHYSIOLOGY. CHAPTER IV. OF FOOD, AND THE DIGESTIVE PROCESS. 1. Sources of the Demand for Aliment. 406. The dependence of all Organized beings upon food or aliment, must be evident from the facts stated in the preceding portion of this Treatise. In the first place, the germ requires a large and constant supply of materials, with which it may develope itself into the perfect being, by the properties with which it is endowed. In all but the lowest tribes of Plants, we find the materials required for the earliest stages of the process prepared and set apart by the parent. Thus in the seed, the germ itself forms but a small proportion of the Avhole substance, the principal mass being composed of starchy matter which is laid up there for its nutrition ; and the act of germination consists in the appropriation of that nutriment by the germ, and the consequent development of the latter, up to the point at Avhich it becomes inde- pendent of such assistance, and is able for itself to procure, from the soil and atmosphere that surround it, the materials for its continued growth. So in the egg of the Animal, the principal mass is composed of Albumen and oily matter ; the germ itself being, at the time the egg is first deposited, a mere point invisible to the naked eye ; these mate- rials serve as the food or aliment of the germ, which gradually draws them to itself, and converts them into the materials of its own struc- ture ; and at the end of a certain period, the young animal comes forth from the egg, ready to obtain for itself the food which is necessary for its continued increase in size. 407. In many instances among the lower animals, the form in which the young animal emerges from the egg is very different from that which it is subsequently to assume ; and the latter "is only attained by a process of metamorphosis. This change has been longest known, and most fully studied, in the case of Insects and Frogs; which were for- merly thought to constitute an exception to all general rules in this respect,—the Insect coming forth from the egg in the state of a Worm, SOURCES OF DEMAND FOR FOOD. 237 and the Frog in the condition of a Fish. But it is now known that changes of form, as complete as these, occur in a large proportion of the lower tribes of Animals; so that the absence of them is the excep- tion. The fact seems to be, that the supply of nutriment laid up within the egg, among the lower classes, is by no means sufficient to carry on the embryo to the form it is subsequently to attain ; and its development is so arranged, that it may come into the world in a condition which adapts it to obtain its own nutriment, and thus to acquire for itself the materials for its further development. Thus the Insect, in its larva or Caterpillar state, is essentially a foetus in regard to its grade of develop- ment ; but it is a foetus capable of acquiring its own food. In this con- dition it attains its full growth as regards size, though its form remains the same ; but it then, in passing into the Chrysalis state, reassumes (as it were) the condition of an embryo within the egg,—the development of various new parts takes place, at the expense of the nutriment stored up in its tissues,—and it comes forth as the perfect Insect. In many of the loAver tribes, the animal quits the egg at a still earlier period in comparison ; thus it has been lately shoAvn by M. Milne Edwards, that some of the long marine worms consist only of a single segment, form- ing a kind of head, when they leave the egg; and that the other seg- ments, to the number (it may be) of several hundred, are gradually developed from this, by a process that resembles the budding of Plants. 408." Up to the period, then, when the full dimensions of the body have been attained, and the complete evolution of all its organs has taken place, a due supply of food is necessary for these purposes. In the Plant, nearly the whole of the alimentary materials taken into the system, are thus appropriated; the extension of its structure going on almost indefinitely, and the waste occasioned by decay being compara- tively small. Thus the carbon, which is given out by the respiratory process in the form of carbonic acid, bears but a small proportion to that Avhich is introduced by the decomposition of that same gas, under the influence of light (§ 81). And the fall of the leaves, which takes place once a year or more frequently, and Avhich gives back a large quantity of the matter that has undergone the organizing process, does not occur, until by their means a considerable addition has been made. to the solid and permanent substance of the tree. 409. This is not the case, however, with the animal. Its period of increase is limited. The full size of the body is usually attained, and all the organs acquire their complete evolution, at a comparatively early period. The continued supply of food is not then requisite for the ex- tension of the structure, but simply for its maintenance ; and the source of the demand lies in the constant waste, to which, during its period of activity, it is subjected. We have seen that every action of the Nervous and Muscular systems involves the death and decay of a cer- tain amount of the living tissue,—as is indicated by the appearance of the products of that decay in the Excretions, and a large part of the demand for food will be consequently occasioned by the necessity for making good the loss thus sustained. Hence we find that the demand for food bears a close relation to the activity of the animal functions; so that a diet, Avhich would be superfluous and injurious to an individual 238 OF FOOD AND THE DIGESTIVE PROCESS. of inert habits, is suitable and beneficial to one who is leading a life of continual exertion ; and this difference manifests itself in the require- ments of the same individual who makes a change in his habits—the indolent man acquiring an appetite by vigorous exertion, and the active man losing his disposition to hearty feeding by any cause that keeps him from his accustomed exercise. We see precisely the same contrast between Animals of different tribes, whose natural instincts lead them to different modes of life. The Birds of .most active flight, and the Mammals which are required to put forth the greatest efforts to obtain their food, need the largest and most constant supplies of nutriment; but even the least active of these classes stand in remarkable contrast with the inert Reptiles, whose slow and feeble movements are attended with so little waste, that they can sustain life for weeks and even months, with little or no diminution of their usual activity, without a fresh supply of food. 410. The waste and decay just adverted to, however, do not affect the muscular and nervous tissues alone ; for all the operations of nutri- tion involve it to a certain extent. It has been already shown that the acts of absorption, assimilation, respiration, secretion, and reproduc- tion ;—all those, in. fact, by which the material for the nutrition of the nervous and muscular tissues is first prepared, and subsequently main- tained in the requisite purity,—are effected in the Animal, as in the Plant, by the agency of cells, which are continually dying and requiring renewal. In most Vegetables, the death of the parts concerned in these functions takes place simultaneously, as soon as they have per- formed them; the whole crop of leaves ceasing at once to perform its proper actions, and dropping off;—to be replaced by another, at an in- terval that solely depends upon the temperature under which the tree is living (§ 99). In the evergreen, however, the process bears a close resemblance to that which we observe in the Animal; for the leaves die one by one, and not simultaneously ; and are constantly undergoing replacement, so that the vigor of the system and the actiA'ity of its nutritive processes never suffer a complete suspension. 411. In the Animal body, the different classes of cells, to which allusion has been made, are in like manner constantly undergoing death and renewal; and this with a rapidity proportioned to the energy of their functions. Hence a supply of food is as requisite, to furnish the materials of their growth, as it is in Plants to furnish the materials of the growth of the leaves. A large part of these materials are subse- quently used for other purposes in the economy; thus, as the leaves prepare the sap Avhich is to nourish the woody stem, and to form new shoots, so do the absorbing and assimilating cells prepare and furnish the fluid elements of the blood, which are to repair the waste of nerve and muscle, bone and cartilage, &c. But still a considerable amount is expended in the simple nutrition of these organs themselves, whose duration is transient, and whose solid parts are cast off as of no further use. Thus the skin and all the mucous surfaces are continually form- ing and throwing off epidermic and epithelial cells, whose formation re- quires a regular supply of nutriment; and only a part of this nutriment (that which occupies the cavity of the cells) consists of matter, that is SOURCES OF DEMAND FOR FOOD. 239 destined to serve some other purpose in'the system, or that has already ansAvered it; the remainder (that of which the solid walls are composed) being furnished by the nutritive materials of the blood, and being henceforth altogether lost to it.—Thus every act of Nutrition involves a Avaste or decay of Organized tissue. 412. We may observe a marked difference, however, between the amount of aliment required, and the amount of waste occasioned by the simple exercise of the nutritive or vegetative functions in the building- up and maintenance of the animal body, and that which results from the exercise of the animal functions. The former are carried on, with scarcely any intermixture of the latter, during foetal life. The aliment, in a state of preparation, is introduced into the foetal vessels; and is conveyed by them into the various parts of the structure, which are developed at its expense. The amount of waste is then very trifling, as we may judge by the small amount of excretory matter, the product of the action of the liver and kidneys, which has accumulated at the time of birth ; although these organs have attained a sufficient development to act with energy when called upon to do so. But as soon as the movements of the body begin to take place with activity, the waste in- creases greatly; and Ave even observe this immediately after birth, when a large part of the time is still passed in sleep, but when the actions of respiration involve a constant employment of muscular power.—In the state of profound sleep, at subsequent periods of life, the vegetative functions are performed, with no other exercise of the animal powers, than is requisite to sustain them ; and we observe that the waste, and the demand for food, are then diminished to a very low point. This is well seen in many animals, which lead a life of great activity during the warmer parts of the year, but which pass the winter in a state of profound sleep, Avithout, hoAvever, any considerable reduction of tempe- rature ; the demand for food, instead of being frequent, is only felt at long intervals, and the excretions are much reduced in amount. And those animals which become completely inert, either by the influence of cold, or by the drying-up of their tissues, do not suffer from the pro- longed deprivation of food; because not only are their animal functions suspended, but their nutritive operations also are in complete abey- ance ; and the continual decomposition of their tissues, Avhich Avould otherwise be taking place, is checked by the cold or desiccation; so that the Avhole series of changes which goes on in their active condition, is completely at a stand. 413. But there is another most important cause of demand for food, amongst the higher Animals, which does not exist either amongst the loAver Animals, or in the Vegetable kingdom. We have seen (chap, ii.) that Mammals and Birds, and to a certain extent Insects also, are able to sustain the heat of their bodies at a fixed standard, and thus to become independent of variations in external temperature. This they are enabled to do, as will be explained hereafter, by a process strictly analogous to ordinary combustion; the carbon and hydrogen which are directly supplied by their food, or which have been employed for a time in the composition of the living tissues and are then set free, being made to unite with oxygen introduced by the respiratory process, 240 OF FOOD AND THE DIGESTIVE PROCESS. and thus giving off as much heat as if the same materials were burned in a furnace. And it has been further shown, that the immediate cause of death in a warm-blooded animal, from Avhich food has been entirely withheld, is the inability any longer to sustain the temperature, which is requisite for the performance of its vital operations (§ 117). Hence Ave see the necessity for a constant supply of aliment, in the case of warm-blooded animals, for this purpose alone : and the demand will be chiefly regulated by the external temperature. When the heat is rapidly carried off from the surface, by the chilling influence of the surrounding air, a much greater amount of carbon and hydrogen must be consumed within the body, to maintain its proper heat, than when the air is nearly as warm as the body itself; so that a diet which is appropriate to the former circumstances, is superfluous and injurious in the latter; and the food which is amply sufficient in a warm climate, is utterly destitute of power to enable the body to resist the influence of severe cold. This is a fact continually experienced; both in the ordinary recurrence of changes of temperature in our own climate; and, still more remarkably, when the same individual is subjected to the extremes of heat and cold, in successively visiting the tropical and frigid zones. 414. Thus we find that in the Animal body, aliment is ordinarily ,' required for four different purposes. First, for the original construc- ■ tion or building-up of the organism. Second, to supply the loss occa- ! sioned by its continual decay, even when in a state of repose. Third, i to compensate for the waste occasioned by the active exercise of the I nervous and muscular systems. And Fourth, to supply the materials for the heat-producing process, by which the temperature of the body is kept up.—The amount required for these several purposes will vary according to the conditions of the body, as regards exercise or repose, I and external heat or cold. It is also subject to great variation with ,' difference of age. During the period of growth, it might be anticipated J that a larger supply of food would be required, than when the full stature has been attained; but a very small daily addition Avould suffice in the case of a child or youth, to produce the entire increase of aAvhole \ year. Yet every one knoAvs that the child requires much more food than the adult, in proportion to his comparative bulk. This results from the much more rapid change in the constituents of his fabric; Avhich is evident from the large proportional amount of his excretions, from the \ quickness with which the effects of illness or of deficiency of food mani- 1 fest themselves in the diminution of the bulk and firmness of the body, from the short duration of life when food is altogether Avithheld, and \ from the readiness with which losses of substance by disease or injury !are repaired, when the nutritive processes are restored to their full activity. The converse of all this holds good in the aged person. The excretions diminish in amount, the want of food may be sustained for a longer period, losses of substance are but slowly repaired, and everything indicates that the interstitial changes are performed with comparative \ sloAvness; and, accordingly, the demand for food is far less in propor- | tion to the bulk of the body than it is in the adult, and may be even V^ absolutely less than in the child of a fourth of its weight. INFLUENCE OF VARIATIONS IN SUPPLY OF FOOD. 241 415. The demand for food is increased by any cause, which creates an unusual drain or waste in the system. Thus an extensive suppu- rating action can be sustained only by a large supply of highly-nutri- tious food. The mother, who has to furnish the daily supply of milk which constitutes the sole support of her offspring, needs an unusual sustenance for this purpose. And there are states of the system, in which the solid tissues seem to possess an unusual tendency to decom- position, and in Avhich an increased supply of aliment is therefore re- quired. This is the case, for example, in diabetes; one of the first symptoms of which disease is the craving appetite, that seems as if it would never be satisfied. And there can be no doubt that, putting aside all the other circumstances that have been alluded to, there is much difference amongst individuals, in regard to the rapidity of the changes which their organism undergoes, and the amount of food re- quired for its maintenance. 416. The influence of the supply of food upon the size of the indivi- dual, is very evident in the Vegetable kingdom ; and it is most strikingly manifested, Avhen a plant naturally growing in a poor dry soil is trans- ferred to a damp rich one, or Avhen we contrast two or more individuals of the same species, groAving in localities of opposite characters. Thus, says Mr. Ward, " I have gathered, on the chalky borders of a wood in Kent, perfect specimens in full flower of Erythrcea Centaurium (Com- mon Centaury), not more than half an inch in height; consisting of one or tAvo pairs of most minute leaves, with one solitary flower: these were growing on the bare chalk. By tracing the plant toAvards and in the wood, I found it gradually increasing in size, until its full development was attained in the open parts of the wood, where it became a glorious plant, four or five feet in elevation, and covered with hundreds of flow- ers." On the other hand, by starvation, naturally or artificially induced, Plants may be dwarfed, or reduced in stature : thus the Dahlia has been diminished from six feet to two; the Spruce Fur from a lofty tree to a pigmy bush; and many of the trees of plains become more and more dwarfish as they ascend mountains, till at length they exist as mere underwood. Part of this effect, however, is doubtless to be attributed to diminished temperature; which, as already remarked, concurs with deficiency of food in producing inferiority of size. 417. Variations in the supply of food would not appear to be effec- tual in producing a corresponding variety of size in the Animal king- dom : this is not, however, because animals are in any degree less dependent than Plants upon a due supply of food; but because such a limitation of the supply, as would dwarf a Plant to any considerable extent, would be fatal to the life of an Animal. On the other hand, an excess of food, which (under favorable circumstances), would pro- duce great increase in the size of the Plant, would have no correspond- ing influence on the Animal; for its size appears to be restrained within much narrower limits,—its period of groAvth being restricted to the early part of its life, and the dimensions proper to the species being rarely exceeded in any great degree. Even in the case of giant individuals, it does not appear that the excess of size is produced by an over-supply of food ; but that the larger supply of food taken in is called for by the 242 OF FOOD AND THE DIGESTIVE PROCESS. unusual wants of the system,—those wants being the result of an extra- ordinary activity in the processes of growth, and being traceable rather to the properties inherent in the system, than to any external agencies. Thus we not unfrequently hear of children, who have attained an extra- ordinary size at the age of a few years ; and this excess of size is usually accompanied with other marks of precocious development. We shall hereafter see, that a provision exists in the Digestive apparatus, which absolutely prevents the reduction and preparation of the food, in any amount greatly surpassing that which the wants of the system demand (§ 474); and it is probably to this cause, in part, that we are to attri- bute the small degree of influence exerted by an excess of food, in pro- ducing an increased development of the Animal frame. 418. The influence of a diminished supply of food, in producing a marked inferiority in the size of Animals, is most effectually exerted during those early periods of growth, in which the condition of the system is most purely Vegetative. Thus it is well known to Entomo- logists, that, whilst it is rare to find Insects departing widely from the average size on the side of excess, dwarf-individuals, possessing only half the usual dimensions, or even less, are not uncommon ; and there can be little doubt that these have suffered from a diminished supply of nutriment during their larva state. This variation is most apt to present itself in the very large species of Beetles, which pass several years in the larva state; and such dwarf specimens have even been ranked as sub-species. Abstinence has been observed to produce the effect, upon some Caterpillars, of diminishing the number of moults and accelerating the transformation ; in such cases, the Chrysalis is more delicate, and the size of the perfect Insect much below the average. 419. One of the most remarkable examples known, of the effect of food in modifying the development of Animals, is to be found in the economy of the Hive-Bee. In every community, the majority of indi- viduals consists of neuters; which may be regarded as females, having the organs of the female sex undeveloped; and Avhich, whilst incapable of reproduction, perform all the labors of the hive. The office of con- tinuing the race is restricted to the queen; who is the only perfect female in the community. If by any accident the queen be destroyed, or if she be purposely removed for the sake of experiment, the bees choose tAvo or three from amongst the neuter larvce, wdiich are being nurtured in their appropriate cells; and these they cause to be developed into perfect queens. The first operation is to change the cells in wThich they lie into royal cells; these differ considerably from the ordinary ones in form, and are of much larger dimensions. This is accomplished by breaking down the walls of the surrounding cells, removing the eggs or grubs they may contain, and rebuilding the central cell upon an en- larged scale, and upon the same plan as the royal cells in which the queens are ordinarily reared. Whilst this operation is going on, the maggot is supplied with food of a very different nature from the farina or bee-bread (composed of a mixture of pollen and honey), which has been stored up for the nourishment of the workers; this food being of a jelly-like consistence and pungent stimulating character. After the usual transformations, the grub becomes a perfect queen ; differing from EFFECTS OF EXCESS OF FOOD. 243 the neuter bee, into which it would have otherwise been changed, no? only in the development of the reproductive system, but in the general form of the body, the proportionate shortness of the wings, the shape of the tongue, jaws, and sting, the absence of the hollows on the thighs in Avhich the pollen is carried, and the loss of the power of secreting Avax. 420. That insufficiency of wholesome food, continued through succes- sive generations, may produce a marked effect, not merely upon the stature, but upon the form and condition of the body, even in the Human race, appears from many cases, in which such influence has operated on an extensive scale. Thus there are parts of Ireland inha- bited by a population descended from those who were treated by the English as rebels two centuries since, and who were driven into moun- tainous tracts, bordering on the sea, where they have been since exposed to the two great brutalizers of the human race, hunger and ignorance. The present race is distinguished physically from the kindred race of Meath and other neighboring districts, Avhere the same causes have not been in operation, by their Ioav stature (not exceeding five feet two inches), their pot-bellies and bow-legs; whilst their open projecting mouths, with prominent teeth and exposed gums, their advancing cheek- bones and depressed noses, bear barbarism in their very front. " These spectres of a people that once were well-grown, able-bodied, and comely, stalk abroad into the daylight of civilization, the annual apparitions of Irish ugliness and Irish want."—The aboriginal population of New Holland, as a whole, presents a similar aspect; and apparently fromj the operation of the same causes. 421. When a larger quantity of azotized food (§429) is habitually consumed than the Avants of the system require, it is not converted into solid flesh; but it is got rid of by the various processes of excretion. The increased production of Muscular fibre depends, as we have already seen (§ 362), upon nothing so much as the exercise of the muscle. It cannot take place unless the blood supply it with the materials; but no degree of the richness of the blood can alone produce it. Consequently, the accumulation of nutritive matter in the blood is so far from being a con- dition of health, that it powerfully tends to produce disease,—either of an inflammatory character, if the fibrine predominate,—or of the he- morrhagic character, if the red corpuscles predominate. This state is most apt to present itself in those who live well and take little exercise; and the remedy for it is either to diminish the diet, or to increase the amount of exercise, so as to bring the tAvo into harmony. 422. The continued over-supply of food has several injurious effects: it disorders the digestive processes, by stimulating them to undue activity, and lays the foundation for a complete derangement of them; it giAres a predisposition to the various diseases of repletion, as already noticed; and it throAvs upon the excreting organs much more than their proper amount of labor, besides tending to produce a depraved condi- tion of the matters to be drawn off by them, which renders the proper act of excretion still more difficult. When this is the case various disor- ders arise, caused by the retention, within the circulating current, of substances Avhich are very noxious to the general system, and which become most fertile sources of disease. What are commonly regarded 244 OF FOOD AND THE DIGESTIVE PROCESS. as diseases of the biliary and urinary organs, are really, in a large pro- portion of cases, nothing else than disordered actions of those organs, occasioned by the irregular mode in which the products of decomposition are formed within the blood, and dependent upon some error in diet, either as regards quantity or quality. Thus the "lithic acid diathesis," in which there is an undue proportion of that substance in the urine, and of which Gout is a particular manifestation, is due, not to disorder of the kidney, but to an undue production of lithic acid in the blood; so long as the excreting action of the kidney is sufficient to prevent its accumulation in the blood, so long the general health is but little affect- ed; but whenever that action receives a check, various constitutional symptoms indicate that the system is disturbed by the presence of this product of decomposition. And though our remedies may be rightly directed, in part, to facilitating its escape through the kidneys, yet the radical cure is to be sought only in the regulation of the diet, and in the prevention of the first production of the substance in question.—Similar remarks might probably be applied to the disorders of the Liver; but we are, from various causes, far less perfectly acquainted with their character than we are with those of the Kidney. 423. There is only one tissue, the increase of which is directly pro- duced by an over-supply of food. This is the Adipose or fatty. It is formed almost entirely at the expense of the non-azotized constituents of the food (§ 430); the walls of the cells, into which the fatty matter is secreted, being the only part of this tissue that is derived from the proteine-compounds of the blood. The production of the adipose tissue is most directly favored by the presence of a large amount of fatty matter in the food ; but it may also be effected, as will be presently shown, by the conversion of starchy and saccharine substances into fatty compounds. It cannot occur, unless there be in the food a larger pro- portion of substances that can be thus appropriated, than is sufficient to maintain the heat of the system by the respiratory process. Conse- quently, whatever increases the demand for heat is unfavorable to the deposition of fat; and vice versd. The fattening of animals is now brought to a regular system; and experience has showrn that rest and a warm temperature, with food containing a large amount of oily mat- ter, are most conducive to the accumulation. Rest acts by keeping the respiration at a low standard; for it will hereafter be shown (chap. viii.), that a much larger proportion of carbonic acid is throAvn off when the body is in active movement than when it is in repose. External warmth has the same effect; the demand upon the calorifying power being diminished, and more of the combustible material being left, to be stored up as fat. « 424. The deposition of fat affords a supply of combustible matter, against the time when it may be needed; and it is consequently found, that the duration of life in warm-blooded animals, when they are com- pletely deprived of food, is in a great degree proportional to the amount of fat they have previously accumulated. There is no sufficient reason to believe that fatty matter can be converted, within the animal body, into a proteine-compound, which can serve for the nutrition of the mus- cular and other tissues. But the greatest and most constant waste, VALUE OF DIFFERENT MATERIALS OF FOOD. 245 when an animal is undergoing starvation, is that which is occasioned by the heat-producing process; this, so long as the supply lasts, is kept up by the store of fat, which is gradually consumed; and when it is com- pletely exhausted, the temperature falls, hour by hour, until life can no longer be sustained (§ 117). The use of this store of fat, in supplying any temporary deficiency in the food, becomes evident from such expe- riments ; for when it has been completely exhausted, the Avithholding of a single meal proves fatal, from the want of power to sustain the calori- fying process. We find that animals, Avhich are likely to suffer from deficiency of food in the Avinter, or which spend that period in a state of quiescence, have a tendency to accumulate a store of fat in the autumn; which tendency seems principally to depend upon the nature of their food. We observe it chiefly in those Birds and Mammals which live upon seeds and grains; and these, when ripe, contain a large quan- tity of oily matter, which thus becomes a valuable store against the time of need. There are many birds such as the beccafico, so much esteemed in Italy, which are described, if killed at this season, as being " lumps of fat." 425. It is well known to breeders of cattle that some varieties or breeds have a much greater tendency to the production of Adipose tis- sue than others placed under the same circumstances ; and the former are therefore selected to undergo the fattening process. Corresponding differences may be met with among different individuals of the Human race; some persons having a remarkable tendency to the production of fat, under circumstances Avhich do not seem by any means favorable to it, whilst others appear as much indisposed to this deposit. The latter condition we notice particularly in that temperament which is commonly termed the "bilious;" and it is important to bear in mind that, Avhere such an indisposition exists, any superfluity of fatty matter in the food taken into the system must be excreted again through the liver, instead of being retained and stored up in the body. It is very desirable, therefore, that such persons should abstain from any excess of this kind ; since an habitual call upon the liver, to relieve the system of a super- fluity of fatty matter, is certain to produce a disordered state of that organ; and in order to prevent it, the diet should be altered, so as to include less of fatty matter, or the amount of exercise should be in- creased so that it may be burned off by the additional respiration which then takes place. ^X~ 426. We see, then, that the amount of food which can be properly „ appropriated by the system varies considerably in different individuals, and in the same individual under different circumstances. Consequently it is impossible to give any general rule, which shall apply to every one alike. The average quantity required by adult men, leading an active life, and exposed to the ordinary vicissitudes of temperature in their own climate, seems to be from 30 to 36 ounces of dry aliment. But a healthy condition may be kept up on scarcely more than half this allow- ance, if the muscular poAA*ers are but little exerted, and the surrounding temperature be high ; provided that it consists of substances of a nutri- tious kind, united in proper proportions. 427. The value of different substances as aliment depends in the first 246 OF FOOD AND THE DIGESTIVE PROCESS. place upon the quantity of solid matter they contain ; being of course the greater as the solids form the larger proportion of the entire weight. : Many esculent vegetables contain so large a quantity of water, that the ; nutriment they afford is very slight in proportion to their bulk.—Next it depends upon the proportion of digestible matter which the solid parts include; for it is not every substance containing the requisite ingre- dients that is capable of being reduced to a state which enables it to be absorbed. Thus woody fibre is composed of the same elements as starch-gum; but it passes out of the intestinal canal unchanged and therefore affords no nutriment. In the same manner, the horny tissues of animals though nearly allied to proteine in their composition, are completely destitute of nutritive properties to man and the higher ani- mals, because not capable of being reduced by their digestive process ; though certain insects appear capable of living exclusively upon them. 428. But when the watery and indigestible parts of the food are put out of consideration, and our attention is directed only to the soluble solids, we find a most important difference in the chemical composition of different substances, which renders them more or less appropriate to v the different purposes Avhich have to be answered in the nutrition of the body. It has been already pointed out, that Vegetables possess the -, power of combining the elements furnished by the inorganic world into two classes of compounds,—the ternary, consisting of oxygen, hydrogen, and carbon,—and the quarternary, which consist of these elements, with <- the addition of azote or nitrogen. These two classes are hence termed the non-azotized, and the azotized. 429. Now the azotized compounds are required for the reparation of the waste of the muscular tissue, and for the general nutrition of the body; consequently, unless the food contain a sufficient proportion of these substances, the body must be insufficiently nourished, and the strength must diminish, even though other elements of the food be in superabundance. The azotized substances formed by Plants are essen- tially the same, as already shown (§ 174), with those Avhich are fur- nished by the Albuminous solids and fluids of animals; but the quantity of them is usually small in proportion to the non-azotized, being consi- i derable only in the Corn-grains, in the seeds of Leguminous plants, and in some other products, which the universal experience of ages has demonstrated to be the most nutritious of Vegetable substances. The other azotized compounds existing in the animal body may be elaborated by the transformation of these proteine-compounds ; so that when they are duly supplied, the system cannot become enfeebled for want of sup- port.—But there is another azotized compound, Gelatine, that is fur- nished by Animals, to which nothing analogous exists in Plants; and this, although it cannot sustain life by itself, is a valuable adjunct to the proteine-compounds. For as the gelatinous tissues suffer waste in com- mon with the others, it is evident that if the gelatine be supplied already prepared, it may be at once applied to their nutrition ; and thus the pro- portion of proteine, which they would otherwise require, is not demanded, and the labor of transformation is also saved. Further, there is this great advantage in combining a proportion of gelatine Avith the food,— especially when the digestive powers are feeble,—that being already in a VALUE OF DIFFERENT MATERIALS OF FOOD. 247 state of perfect solution, it is taken up at once by the simple act of physi- cal absorption or endosmose, instead of requiring any preliminary pre- paration .or exertion of vital activity for its absorption. But there is no evidence that Gelatine can ever be transformed into a proteine-compound, and can thus be applied to the nutrition of the muscular and other fibrous tissues ; and the presumption, derived from the result of various experi- ments, is very strong the other wray. 430. The Non-azotized compounds, which are presented to us in great abundance in the Vegetable kingdom, exist under various forms; of 0->-T^ which the principal are starch, sugar, and oil. The two former may be •' < regarded as belonging to one class, the Saccharine ; because we know that starch and the substances allied to it may be converted into sugar* by simple chemical processes, and that this transformation takes place T readily both in the Vegetable and Animal economy. On the other hand, the Oily matters contained in vegetable and animal food, are usually " ranked as a distinct group of alimentary substances; and it has been _ maintained that, under no circumstances, has the Animal the power of elaborating fatty matter from starchy or saccharine compounds. But this is now knoAvn to be an unfounded limitation; since the transforma- tion of a saccharine into a fatty compound takes place in the case of „ bees, which form wax when fed upon pure sugar, and it has been recently shown that it may take place in the laboratory of the Chemist, butyric acid (the fatty acid of butter) being one of the products of the fermenta- tion of sugar, taking place under peculiar circumstances. It appears, / indeed, to be one office of the Liver to effect this transformation, as will be explained hereafter. 431. The great use of these substances in the Animal economy, is to support the respiratory process, and thus maintain the temperature of the body. We have seen that, in the compounds of the Saccharine group (in Avhich Starch is included), the amount of oxygen is no more than sufficient to form water with the hydrogen of the substance (§ 12), so that the carbon is free to combine with the oxygen taken in by the lungs, and thus becomes a source of calorifying power. Again, in the oily matters taken in as food, the proportion of oxygen is far smaller, so that they contain a large quantity of surplus hydrogen, as well as of carbon, ready to be burned off in the system, and thus to supply the heat re- quired. This is obviously the ordinary destination of the alimentary matters belonging to these classes; and the greatest economy in the choice of diet is therefore exercised, when it is composed of azotized sub- stances in sufficient amount to repair the waste of the system, and of non-azotized compounds which include free carbon and hydrogen in suf- ficient quantity to develope (with the aid of other processes) the requisite amount of heat by combination with oxygen. But if there be a deficiency in either of these kinds of aliment, the body must suffer. Should the supply of duly-prepared azotized matter be less than is required to re- pair the waste of the albuminous and gelatinous tissues, then these di- minish in bulk and in vital poAver, though the heat of the body may be kept up to its proper standard. But if the non-azotized matter should be supplied in sufficient amount, or in a form in which it cannot be appropriated, the heat of the body cannot be sustained in any other • way, than by drawing upon the store of fat previously laid up. 248 OF FOOD AND THE DIGESTIVE PROCESS. 432. Various circumstances lead to the belief, that the saccharine compounds are thus carried off by the respiratory process, within a short time after they have been introduced into the system. They have not been detected in the chyle drawn from the lacteal absorbents; but there seems reason to believe that, in consequence of their ready solubility, they are directly taken up by the blood (§ 493), and that they are so rapidly burned off there, as to escape notice in that fluid. But it has been lately shown by Dr. Buchanan, that, if the blood be examined within a short time after a meal consisting in part of farinaceous and saccharine substances, a very appreciable quantity of saccharine matter is found in it. This soon disappears, however, being eliminated or sepa- rated from the blood by the action of the lungs. In fact it is very pro- bable, that a large proportion of the matter thus taken in never enters the general circulation at all; as the blood of the mesenteric veins pro- ceeds to the lungs, after passing through the liver, before it is trans- mitted to the systemic arteries, and may there lose its saccharine matter, as fast as this is taken in from the stomach. After a meal containing the ordinary admixture of saccharine, oily, and albuminous compounds, it is probable that the saccharine are first received into the blood, and are the first to be eliminated from it; and that, by the time they have been all consumed, the oily matter, introduced through the more circuitous channel of the lacteal system, is ready to answer the same purpose. If these are exhausted before a fresh supply of food is taken in, cold as well as hunger is experienced ; and the body is in this condition pecu- liarly liable to suffer from any depressing causes, such as a low external temperature, poisonous miasmata, &c.; hence the prudence of avoiding exposure to such influences upon an empty stomach. 433. We can thus in part account for the fact, Avhich universal expe- rience has established, that in Avarm-blooded animals, a mixture of azo- tized and non-azotized substances is the diet most conducive to the welfare of the body ; and that, in all but the purely carnivorous tribes, the diet provided by Nature consists not only of albuminous, gelatinous, and oily substances, such as are furnished by the flesh and fat of animals, but also of saccharine or farinaceous matter. This is the diet to which Man is evidently best adapted ; and it is remarkable hoAV completely accordant is his use of the ordinary materials of food, with the principles now es- tablished by chemical and physiological research, in regard to the wants of his bodily system, and the best mode of supplying them. Thus, good AA'heaten bread contains, more nearly than any other substance in ordi- nary use, that proportion of azotized and non-azotized matter, Avhich is adapted to repair the Avaste of the system, and to supply the neces- sary amount of combustible material, under the ordinary conditions of civilized life in temperate climates; and we find that the health and strength can be more perfectly sustained upon that substance, than upon any other taken alone. The addition of a moderate quantity of butter increases its heat-producing powers: and this is especially useful Avhen the temperature is Ioav, under which condition there is usually an in- creased disposition to the employment of fatty matters as articles of food. On the other hand, if the body be subject to violent exertion, advantage is gained by increasing the proportion of the proteine-com- pounds, by the addition of animal flesh; and, under any circumstances, VALUE OF DIFFERENT MATERIALS OF FOOD. 249 there is an economy in the use of gelatine, in the form of soup, which diminishes the demand for other azotized matter. The use of animal flesh, hoAvever, as a principal article of diet, except when the individual is lead- ing the incessantly-active life of a carnivorous animal, is very far from being economical, and is positively injurious to the welfare of the body. 434. On the other hand, in rice, potatoes, cassava-meal, and similar substances, the farinaceous or saccharine components form so very large a proportion of the Avhole mass, and the proteine-compounds are present in so very small an amount, that they are insufficient to support the bodily vigor when taken alone, unless a larger quantity be ingested, so as to supply the requisite proportion of azotized matter. But Avhen these substances form part of a mixed diet, the other ingredients of Avhich consists of animal flesh, a much smaller quantity of them suffices; and the same kind of combination is then formed, as exists in the single article of bread. Those in Avhose diet the farinaceous elements predo- minate largely, and the azotized compounds exist in the smallest amount compatible with the maintenance of the bodily vigor, are exempt from many diseases incident to those Avho live more highly; thus among the potato-eating Irish, and the oatmeal-feeding Scotch, gout is a disease never heard of; whilst among the richer classes of the same countries, there is no peculiar exemption from it. 435. The oily constituents of food are most abundant in the diet of the inhabitants of frigid zones, who feed upon Avhales, seals, and other animals loaded with fat, and Avho devour this fat with avidity, as if in- stinctively guided to its use. It is by the enormous quantity of this substance taken in by them, that they are enabled to pass a large part of the year in a temperature beloAV that of our coldest winter, spending a great portion of their time in the open air ; as well as to sustain the extreme of cold, to Avhich they are occasionally subjected. And in con- sequence of its being more sloAvly introduced into the system than most other substances, a larger quantity may be taken in at one time, Avith- out palling the appetite ; Avhilst its bland and non-irritating character favors its being retained until it is all absorbed. In this manner, the Esquimaux and Greenlanders are enabled to take in 20 or 30 pounds of blubber at a meal; and, Avhen thus supplied, to pass several days without food. On the other hand, among the inhabitants of warm climates there is comparatively little disposition to the use of oily matter as food ; and the quantity of it contained in most articles of their diet is comparatively small. 436. In the Milk, Avhich is the sole nutriment of the young Mammalia, during the period immediately succeeding their birth, Ave find an admix- ture of albuminous, saccharine, and oleaginous substances; AYhich in- dicates the intention of the Creator, that all these should be employed as components of the ordinary diet. The Caseine or cheesy matter is a proteine-compound ; the Butyrine of butter is but a slight modification of its ordinary fats ; and its Sugar differs from that in common use, only by its larger proportion of Avater. The relative amount of these ingre- dients in the milk of different animals is subject, as Ave shall hereafter see, to considerable variation; but they constantly exist, at least in the milk of the Herbiv ^rous Mammalia, and of those Avhich, like Man, sub- sist upon a mixed diet. But it has been recently asserted, that the milk 250 OF FOOD AND THE DIGESTIVE PROCESS. of the purely Carnivorous animals is destitute of Sugar, consisting, like their food, of proteine-compounds and fatty matter only. 437. No fact in Dietetics is better established, than the impossibility of long sustaining health, or even life, upon any single alimentary prin- ciple. Neither pure albumen or fibrine, gelatine or gum, sugar or starch, oil or fat, taken alone for any length of time, can serve for the due nu- trition of the body. This is partly due, so far as the non-azotized com- pounds are concerned, to their incapability of supplying the waste of the albuminous tissues. This reason does not apply, hoAvever, to the pro- teine-compounds ; which can not only serve for the reparation of the body, but can also afford the carbon and hydrogen requisite for the sus- tenance of its temperature. The real cause is to be found in the fact, that the continued use of single alimentary substances excites such a feeling of disgust, that the animals experimented on seem at last to prefer starvation, rather than the ingestion of them. Consequently it is quite impossible to ascertain, by such experiments, the nutritive poAver of the different alimentary principles; no animal being capable of sus- taining life upon less than tAvo of them at least. The same disgust is experienced by Man, when too long confined to any article of diet, which is very simple in its composition ; and a craving for change is then ex- perienced, which the strongest will is scarcely able to resist. Thus, in the treatment of Diabetes, a disease in which there is an undue tendency to the production of sugar in the system, it is very important to abstain completely from the introduction of saccharine or farinaceous matters in the food; but the craving for vegetable food, which is experienced when the diet has long consisted of meat alone, is such as to make per- severance in the latter very difficult; and a means has been latterly devised of supplying this want without injury, by the use of bread from which the starchy portion has been removed, the gluten or azotized matter alone being eaten.* 438. The organic compounds, which have been enumerated as sup- plying the various wants of the system, would be totally useless without the admixture of certain inorganic substances, which also form a con- stituent part of the bodily frame, and which are constantly being voided by the excretions, especially in the Urine. These substances have various uses in the system. Thus common Salt, or the Chloride of Sodium, appears to afford, by its decomposition, the muriatic acid which is concerned in the digestive process, and the soda which is an important constituent of the bile. Its presence in the serum of the blood, also, and in the various animal fluids which are derived from this, probably * As an illustration of the advantage of this treatment, even in unpromising cases, the author may cite an instance which has come under his own observation. The patient was a man of 72 years of age; the disease had lasted at least a year, and was decidedly on the increase; considerable loss of flesh and of muscular vigor had taken place ; and the quantity of sugar in the urine was such as to make it quite sweet to the taste. By the careful restriction of his diet to animal flesh and gluten-bread, this in- dividual kept the disease in complete check for more than 'five years; he gained flesh, and improved in strength ; and his urine lost its sweetness. Having two or three times ventured upon a return to his ordinary diet, his old symptoms immediately manifested themselves, warning him of the necessity of perseverance in the strict regimen pre- scribed for him. He died at last, at the age of 77 years, of old age, rather than of any specific disease. NECESSARY MATERIALS OF ANIMAL FOOD. 251 aids in preventing the decomposition of the organic constituents of these fluids.— 1 hosphorus has been supposed, until recently, to be chiefly re- quisite as one of tne materials of the nervous tissue (§ 383); and also, when acidified by oxygen, to unite with lime in forming the bone-earth by which bone is consolidated. But there is reason to believe, from the results of late inquiries, that the acid and alkaline phosphate of lime and soda are very important constituents of the various fluid secretions, and have a large share in their respective actions.—Sulphur exists in small quantities in several animal tissues ; but its part appears to be by no means so important as that performed by phosphorus.—Lime is required for the consolidation of the bones, and for the production of the shells and other hard parts that form the skeletons of the Invertebrata ; and also as the base of the acid phosphate, which has been just referred to as an important constituent of the animal fluids.—Lastly, Lron is an essential constituent of Haematosine; and is consequently required for the production of the red corpuscles of the blood in Vertebrated animals. 439. These substances are contained, more or less abundantly, in most of these articles generally used as food; and where they are defi- cient, the animal suffers in consequence, if they be not supplied in any other Avay. Thus common Salt exists, in no inconsiderable amount, in the flesh and fluids of animals, in the milk, and in the substance of the egg; it is not so abundant, however, in Plants; and the deficiency is usually supplied to herbivorous animals in some other way. Thus/salt is pur- posely mingled with the food of domesticated animals; and in most parts of the world inhabited by wild cattle, there are spots where it exists in the soil, and to which they resort to obtain it. Such are the "buffalo-licks" of North America.—Phosphorus exists also, in combi- nation with proteine-compounds, in all animal substances composed of these ; and in the state of phosphate, combined with lime, magnesia, and soda, it exists largely in many vegetable substances ordinarily used as food. The phosphate of lime is particularly abundant in the seeds of the grasses; and it also exists largely, in combination with caseine, in Milk.—Sulphur is derived alike from vegetable and animal substances. It exists, in union with proteine-compounds, in flesh, eggs, and milk; also in several vegetable substances; and, in the form of sulphate of lime, in most of the river and spring water that we drink. 440. Lime is one of the most universally diffused of all mineral bodies; for there are very few Animal or Vegetable substances in which it does not exist. The principal forms in which it is an element of Ani- mal nutrition, are the carbonate and phosphate. Both these are found in the ashes of the grasses, and of other plants used as food; the phos- phate of lime being particularly abundant (as already mentioned) in the corn-grains. The production of these cannot take place, to their fullest extent, unless the soil previously contain phosphate of lime in a state in which the plant can receive it; and it is now understood, that the diminished fertility of many lands is due, in great part, to the exhaus- tion of the soil as regards this ingredient. The restoration of the alka- line and earthy phosphates to the soil, in the form of manure, is the obvious means of preserving its fertility; but so long as a very large proportion of the excrements of animals (the materials of which are originally derived from the earth, through the vegetables it supplies) is 252 OF FOOD AND THE DIGESTIVE PROCESS. allowed to run to waste, so long will it be necessary that the requisite amount of phosphate of lime should be draAvn from foreign sources. 441. The phosphate of lime, as already mentioned, seems to perform important offices of a chemical nature in the animal economy, besides being the chief solidifying ingredient of bones and teeth; but the car- bonate Avould seem principally destined to mechanical uses only ; and Ave find it predominating, or existing as the sole mineral ingredient, in those non-vascular tissues of the Invertebrated animals, Avhich give sup- port and protection to their soft parts (§ 277). The degree of develop- ment of these tissues depends in great part upon the supply of carbonate of lime which the animals receive. Thus the Mollusca which inhabit the sea, find in its waters the proportion of that substance which they require; but those dAvelling in streams and fresh-water lakes, which contain but a small quantity of lime, form very thin shells; Avhilst the very same species inhabiting lakes, which, from peculiar local causes, contain a large impregnation of calcareous matter, form shells of re- markable thickness. The Crustacea which periodically throw off their calcareous envelope (§ 285), are enabled to renew it Avith rapidity by a very curious provision. There is laid up in the Avails of their stomachs a considerable supply of calcareous matter in the form of little concre- tions, which are commonly known as "crabs' eyes." When the shell is throAvn off, this matter is taken up by the circulating current, and is thrown out from the surface, mingled with the animal matter of which the shell is composed. This hardens in a day or tAvo, and the neAV covering is complete. The concretions in the stomach are then found to haA'e disappeared; but they are gradually replaced before the supply of lime they contain is again drawn upon. The large amount of carbo- nate of lime which is required by the laying Hen, is derived from chalk, mortar, or other substances containing it, which she is compelled by her instinct to eat; and if the supply of these be Avithheld, the eggs which she deposits are soft on the exterior,—not being destitute of shell, as commonly supposed,—but having the fibrous element of the shell (§ 181) unconsolidated by the intervening deposit of chalky particles. 2. Of the Digestive Apparatus, and its Actions is general. 442. It has been already pointed out, that the nature of the food of Animals is so far different from that of Plants, as to require the prepa- ratory process of Digestion, before its nutritious part can be taken up by the absorbent vessels and received into the system. This process may be said to have three different purposes in vieAv: the reduction of the alimentary matter to a fluid form, so that it may become capable of absorption ; the separation of that portion of it which is fit to be assi- milated or converted into organized texture, from that Avhich cannot serve this purpose, and which is at once rejected; and the alteration, to a certain extent when required, of the chemical constitution of the former, Avhich prepares it for the important changes it is subsequently to undergo. The simplest conditions requisite for the accomplishment of these purposes are the folloAving: a fluid capable of performing the solution, and of effecting the required chemical changes; a fluid capable of separating the excrementitious matter, by a process analogous to SIMPLEST FORMS OF DIGESTIVE APPARATUS. 253 chemical precipitation; and a cavity or sac in which these operations may be performed. 443. In the loAvest Animals, Ave find this cavity formed upon a very simple plan ; the digestive sac being a mere excavation in the solid tissue of the body, lined with a membrane which is an inverted continuation of the external integument, and communicating with the exterior by one orifice only, through Avhich food is drawn in, and excrementitious matter rejected. In the little Hydra, or fresh-Avater Polype, the external covering of the body and the lining of the stomach correspond so closely in their structure, their actions differing only with their situation, as to be mutually convertible; for the animal maybe turned completely in- side-out, without its functions being deranged. The fluid necessary to dissolve the food, knoAvn by the name of "gastric fluid" or "gastric juice" is secreted in the Avails of the stomach; and, from the trans- parency of the tissues, the Avhole process may be watched. The prey is frequently, and indeed generally, introduced alive, by the contractile poAver of the arms, which coil round it, and gradually draw it into the mouth or entrance to the stomach; and its movements may often be observed to continue for some time after it has been swallowed. In a little time, however, its outline appears less distinct, and a turbid film partly conceals it; the soft parts are soon dissolved and reduced to a fluid state; and any firm indigestible portions which the body may con- tain, are rejected through the aperture by which it entered. The nutri- tive matter is absorbed by the Avails of the stomach, every part of which appears to be endoAved Avith equal power in this respect: and it is con- veyed to the remoter parts of the arms by the simple imbibition of one part from another, without any proper circulation through vessels. 444. In Polypes of a higher conformation, however, the digestive cavity is provided with a second orifice: from the dilated cavity or stomach, an intestinal tube proceeds; and this has a termination dis- tinct from the mouth, though often in its neighborhood. The food, be- fore entering the stomach, is submitted to a poAverful triturating appa- ratus, resembling the gizzard of birds, by Avhich it is broken doAvn; and in the digestive cavity it is submitted, not merely to the action of the gastric fluid, but also to that of the bile, which is secreted in little follicles in the Avails of the stomach, and Avhich is poured into its cavity during the process of digestion,—being easily recognised by its bright yelloAv color. The excrementitious matter is rejected in the form of little pellets, through the intestinal tube. 445. As Ave ascend the Animal scale, we find the digestive apparatus gradually increased in complexity; but its essential characters remain the same. Near the entrance to the stomach, we usually find an appa- ratus for effecting the mechanical reduction of the food, by Avhich its subsequent solution may be rendered more easy. This may consist of a set of teeth ; either fixed in the mouth, as in Mammalia and Reptiles ; or more particularly besetting the pharynx, as in Fishes; or attached to the Avails Of the stomach, as in Crustacea. Or it may be formed by the tongue, conArerted into a sort of rasp ; as in the common Limpet, which thus reduces the sea-Aveeds that constitute its chief food. Or the same purpose may be answered by a gizzard, or first stomach, with dense 254 OF FOOD AND THE DIGESTIVE PROCESS. muscular and tendinous walls ; such as we find in the grain-eating Birds, and many Insects, and in certain Molluscs and Polypes. But where the food is already composed of very minute particles, or is received in a liquid state (as in the case of those animals which live upon the juices of others), or is easily acted on by the gastric juice, no such preparation is requisite. 446. Before the food reaches the true digestive stomach, it is some- times delayed in a previous cavity, in order that it may be macerated in fluid, and may be thoroughly saturated with it. This is the purpose of the crop of Birds, and of the first stomach of Ruminant animals. When this incorporation with fluid is not effected before the food is sub- jected to the triturating process, it usually takes place concurrently with it; and in those animals which reduce their food in the mouth by the process of mastication, there is a special secretion of fluid into that cavity, for this purpose; this fluid is termed Saliva, and the act by Avhich it is incorporated with the food is termed insalivation. ]The mechanical reduction of the aliment, and its incorporation with fluid, constitute, as we shall hereafter see, a very important preparation for the true digestive process./ 447. This process, among the higher animals, takes place exclusively, I or nearly so, in the stomach ; the form of which varies Avith the charac- ter of the food. When this is of a nature to be easily acted on by the gastric fluid, the stomach is a simple enlargement of the alimentary canal, almost in the direct line betAveen the oesophagus and the intestinal Fig. 74. A vertical and longitudinal section of the Human stomach and duodenum, made in such a direction as to include the two orifices of the stomach. 1. The oesophagus ; upon its internal surface the plicated arrange- ment of the cuticular epithelium is shown. 2. The cardiac orifice of the stomach, around which the fringed border of the cuticular epithelium is seen. 3. The great end of the stomach. 4. Its lesser or pyloric end. 5. The lesser curve. 6. The greater curve. 7. The dilatation at the lesser end of the stomach, which has received from AVillis the name of antrum of the pylorus. This may be regarded as the rudiment of a second stomach. 8. The rugae of the stomach formed by the mucous membrane : their longitudinal direction is shown. 9. The pylorus. 10. The oblique portion of the duodenum. 11. The descending portion. 12. The pancreatic duct and the ductus communis eholedochus, close to their termination. 13. The papilla upon which the ducts open. 14. The transverse portion of the duodenum. 15. The commencement of the jeju- num. In the interior of the duodenum and jejunum, the valvulse conniventes are seen. tube ; so that there is little provision for the delay of the food in its cavity. But when the aliment is such as to be less easily reduced, and DIGESTIVE APPARATUS OF MAN. 255 requires to be submitted to the action of the gastric fluid for a longer period, the stomach forms a more considerable enlargement, and is placed more out of the direct line betAveen the oesophagus and the commence- ment of the intestine. The former condition obtains in the Carnivora, and particularly in those which live more upon blood than upon flesh,— such as Weasels, Stoats, &c, in which this part of the alimentary tube is almost straight; the latter condition is found among the Herbivora, and the provision for the delay of the aliment attains its greatest com- plexity in the Ruminant animals. The form of the human stomach (Fig. 74) is intermediate between that of purely carnivorous and purely herbivorous animals. As in the former, there is a direct passage from the cardiac orifice, or entrance of the oesophagus, to the pyloric orifice, or commencement of the intestine ; but there is also a considerable dilatation or cul de sac, which is out of that line; and it appears that, during the digestive process, there is a constriction across the stomach, Avhich separates the cardiac portion from the pyloric, and causes the retention of the food in the dilated part or large extremity. The gas- tric fluid is still secreted in the walls of this organ, by scattered follicles Avhich pour their products into its cavity through separate orifices; but the bile is elaborated by a distinct organ, altogether removed from it, Avhich transmits its secretion by a single duct, that opens into the intes- tinal tube at a short distance from its commencement; and at the same point is delivered the pancreatic secretion, which, as we shall hereafter see (§ 480), takes an important share in the preparation of the alimentary products. 448. The action of the Stomach is restricted, in the higher animals, to the reduction of the food by the solvent poAvers of the gastric juice], and to the absorption (by the vessels in its walls) of those parts of it* which are in a state of the most perfect solution. The changes whichi are produced by the admixture of the biliary and pancreatic fluids take) place in the intestine; and the principal part of the nutritive elements! of the food are taken up by the absorbent vessels of the walls of the \ intestine, after that process has been accomplished. It would seem as! if the preparation of the food for absorption Avere not by any means I completed, in this first portion of the alimentary canal; for it is still( destined to pass through a long and convoluted tube, which is sometimes j extended to an extraordinary degree; and in this passage it is gradually exhausted of its nutritious matter. The length of the intestinal canal/ bears a close relation to the character of the food. In the Carnivorous! animals, Avhose aliment is easily dissolved and prepared for conversion t into blood, the intestine is comparatively short; thus in the Lion and 1 other Felines it is no more than three times the length of the body; / and in some of the bloodsucking Bats, it is almost straight and simple, j On the other hand, in Herbivorous animals it is of enormous length;] thus in the Sheep it is about twenty-eight times as long as the body. In animals Avhose diet is mixed, its length is intermediate between these extremes ; thus in Man, the whole length of the intestinal tube is \ about thirty feet, or between five and six times that of the body. The / intestine is of much smaller diameter along its first portion, than it is : nearer its termination; and it is consequently distinguished into the / 256 OF FOOD AND THE DIGESTIVE PROCESS. small and the large. In the small intestine, which constitutes in Man about five-sixths of the Avhole, the surface of the mucous membrane is greatly extended by the valvuloe conniventes, which are folds or dupli- catures, often several lines in breadth, not entirely surrounding the intestine, but extending for about one-half, or three-fourths of its cir- cumference. These are wanting at the lower part of the ileum. The whole surface of the mucous membrane of the small intestine, below the entrance of the biliary ducts, is thickly covered with villi, or little root-like projections, in Avhich the proper absorbent vessels originate. No proper valvulse conniventes exist in the large intestine; the only extensions of the mucous membrane being crescentic folds at the edges of the sacculi or pouch-like dilatations in its walls; and the villi are comparatively few in number, gradually disappearing towards the termi- nation of the intestine. 449. The mucous membrane of the alimentary canal, through its whole course, is studded with the orifices of numerous scattered glands which lie in its thickness, or immediately beneath it. The simplest of these are the follicles of Lieberkuhn, which are small pouches, formed by an inflexion of the mucous surface, analogous to the follicles of other mucous membranes and apparently destined for the elaboration of the p. _r protective secretion (§ 237, see Fig. 31, b). These follicles in the small intestine, are very simple in their character, and not very deep; and their apertures, which are small, are situated for the most part around the bases of the villi. In the large intestine they are more prolonged, especially towards the extremity of the rectum, where they vnious coat of smaii in- form a distinct layer, the component tubes of the"foiircies'oTueberkFhn wmcn are visible to the naked eye; they probably sued with tenacious white form the peculiarly thick and tenacious mucus of that part. These mucous follicles become parti- cularly evident when the membrane is inflamed; for they then secrete an opaque whitish matter, which is absent in the healthy state, and which distinguishes their orifices (Fig. 75). A modified kind of these follicles, rather more complex in structure, is found abundantly in the stomach; where it is concerned in the secretion of the gastric fluid (§ 469). 450. The coats of the intestine contain other glandulae, of which some appear destined, like the preceding, to elaborate fluids of use in the system; Avhilst others serve rather to draw off from the blood certain products of decomposition, which are to be excreted from it. The former are known as the glands of Brunner, and the latter as those of Peyer, after the names of their respective discoverers. The glands of Brunner are situated in the duodenum, and lie, not in the mucous but in the sub-mucous tissue. Though their size is only about that of a hemp-seed, they are of very complex structure, consisting of several hundred follicles, clustered round the ramifications of an excretory duct, so as to resemble the salivary glands (Fig. 79); and each pours its secretion through a single orifice into the intestinal tube. Athough nothing is certainly knoAvn of the properties of the fluid secreted by these ALIMENTARY CANAL AND ITS MOVEMENTS. 257 glandulae, yet there is strong reason to belieA'e, from their position and character, that it assists the pancreatic and biliary secretions in pre- paring the alimentary materials for absorption (§ 480).—The glands of Peyer are either solitary or agminated; the latter form large patches, which are made of aggregations FiS- 76. of the former. Each solitary gland in its closed r.-^7p\ state consists of a spheroidal vesicle (Fig. 76, a), a -^y''^;'M Avhich is half imbedded in the mucous membrane, i':-}^} but which also forms an elevated projection above ■^m^r jt; and this projection is surrounded by a ring or zone of openings which lead into an annular cluster of Lieberkuhnian follicles. On rupturing one of these vesicles, its cavity is found to contain a gray- ish white matter, interspersed with cells in various Pe^n^^u^fShe stages of development; and these products appear to large intestine:—a, as seen b. o l.i , • n i • t from above, when closed; b, e set tree by the spontaneous opening oi the vesicle, as seen in section, after ha- which takes place when it has become mature, by desSofSer^uhnoneeit0her the thinning away of its wall at its most projecting side- part (Fig. 76, b). In any one of the agminated glands, some of the vesicles are usually found to be open, and others closed. The closed condition is not, as was once supposed, peculiar to the Peyerian glan- dulae ; since, as will be shown hereafter, it is the general rule for other glandular follicles in an early stage of their development to be equally closed (§ 718). Of the nature of the secretions of the Pey- erian glandulae, nothing has been positively ascertained; but some probable inferences from well-known facts will be stated hereafter (§ 749). 3. Movements of the Alimentary Caned. 451. The food which is conveyed to the mouth, is grasped with the lips, by a muscular effort, which is voluntary in the adult under ordinary circumstances, but Avhich may be performed automatically when the influence of the will is withdrawn; in the infant, as among the lower animals, the action seems purely automatic,- the nipple of the mother being firmly grasped by the lips when introduced between them, even after the brain has been removed.—By the act of mastication, Avhich then succeeds, the food is triturated and mingled with the salivary secretion ; and is thus prepared for the further process of solution, to which it is to be subjected in the stomach. The degree of this prepa- ration, and the form of the instruments by which it is effected, vary in different animals, according to the nature of the food. In those Carni- vora whose aliment consists exclusively of flesh, very little mastication is nece'ssary, because this substance is very readily acted on by the gastric fluid; and Ave accordingly find the molar teeth raised into sharp cutting edges, and Avorking against each other with a scissors-like action (the only one permitted by. the articulation of the jaw), so as simply to divide the food. On the other hand, in those Herbivora Avhose food consists of tough vegetable substances, such as the leaves of grasses, or the stems and roots of other plants, we find the molar or 17 w 258 OF FOOD AND THE DIGESTIVE PROCESS. grinding teeth peculiarly adapted to its reduction; their surface being extended horizontally, and being kept continually rough, by the alter- nation of vertical plates of different degrees of hardness; and the lower jaw being so connected with the skull, that great freedom of motion is permitted. In Man we find an intermediate conformation, as regards both the teeth and the articulation of the jaw; for the molar teeth possess broad surfaces which are covered with a continuous coat of enamel, but which are raised into rounded tubercles ; and the articula- tion of the jaw allows it a degree of freedom, which is much greater than that possessed by the Carnivora; although inferior to that which, exists in many Herbivora. The whole apparatus of Mastication is so formed in Man, as to lead to the conclusion that he is destined to live on a mixed diet, composed in part of animal flesh, and in part of vege- table substances that are sufficiently soft to be reduced by the simple act of crushing, or by the slight trituration for which the molar teeth are adapted. 452. The mechanical reduction of the food by Mastication, and the incorporation of the Salivary secretion with its substance, constitute a very important step in the Digestive process. We shall hereafter see that the operations, to which the alimentary matter is subjected in the stomach, are of a purely Chemical nature ; and this preparation is exactly of the same character as that, which the Chemist finds it advantageous to make, Avhen he is operating on a substance of difficult solution. For nothing is so favorable to the action of the solvent, as the previous reduction of the matter to be dissolved, and its thorough incorporation with (he fluid that is to act upon it. We shall hereafter see, that the relative properties of the Saliva and of the gastric fluid are such, that, by the minute admixture of the food with the former, the latter finds access to every particle of it. Hence the practice of eating so rapidly, that Mastication and Insalivation are insufficiently performed, is extremely injurious ; since it throws more work upon the Stomach than it ought to perform, by rendering its solvent action more difficult. There can be no doubt that, by the prolonged continuance of , it, a foundation is laid for the distressing complaint termed Dyspepsia, or difficulty of digestion; »and where any form of this complaint exists, too much attention cannot be paid to the efficient reduction of the food in the mouth. 453. When the aliment has been sufficiently triturated, it is con- veyed into the Pharynx by the act of Deglutition or swalloAving. , , •'i This act involves a great many distinct movements, into a minute '" y description of which we shall not here enter; but it is desirable that its general nature should be well understood. It is one of those most .purely reflex in its character (§ 394), and is not capable of being per- ^y formed or even controlled by a voluntary effort. This stateme'nt may seem inconsistent with the fact, that we SAvallow w-hen we will; but it is not so in reality. The muscular movements which are concerned in »W deglutition, are called forth by nerves that proceed from the spinal cord, not from the brain ; these motor nerves are excited to action, by the contact of solid or fluid matters Avith the mucous surface of the fauces, and in no other* way. The impression produced by the contact is MOVEMENTS OF C3SOPHAGUS. 259 conveyed to the Medulla Oblongata, or that portion of the spinal cord which lies Avithin the cranium, by afferent nerves that terminate in it; and, in immediate respondence to that impression, a motor impulse is transmitted from it, which calls the muscle into the combined action necessary to produce the movement. Now this contact also produces a sensation, provided the brain be sound and awake, because nervous fibres proceed from the mucous surface to the brain as well as to the spinal cord ; but this sensation is not a necessary link in the chain of actions, by Avhich the movement is produced; for the act of Deglutition takes place during profound sleep, when all sensation is suspended, and it may be excited even after the brain has been removed. It seems to be voluntary, under ordinary circumstances, simply because it is by an act of the will, that the matter to be swallowed is carried backAvards into contact with the fauces; but that it is not so in reality, is shoAvn by the fact, that when this impression has once been made with sufficient force, we cannot by any effort of the will, prevent the action. We have a good example of. this in the following circumstance, of no very unfrequent occurrence. The tickling of the upper part of the fauces with a feather is often practised to induce vomiting ; but if the end of the feather be carried too low down, it excites the act of degluti- tion instead; the feather is grasped by the pharnyx and draAvn down- Avards; and if it be not held tenaciously between the fingers, it is drawn from them and carried doAvmvards into the stomach. 454. The carrying-back of the alimentary matter, so that it reaches the fauces or upper part of the pharynx, is principally accomplished by the tongue ; when it has passed the anterior palatine arcnes, these con- tract and close over the tongue, so as to prevent the return of the food / into the mouth; and at the same time the posterior palatine arches and the uvula are so drawn together, as to prevent its passage into the > posterior nares. The larnyx is drawn forward beneath the root of the tongue, and the epiglottis is pressed down over the rima glottidis, so that nothing can enter the latter, unless draAvn towards it by an act of inspiration. When fairly Avithin the pharynx, the alimentary matter is seized by the constrictors which enclose that part of the alimentary tube, and is drawn downwards by them into the oesopha- gus, which is the cylindrical continuation of it. The continued action of the constrictors serve to propel the food along the oesophagus ; their movement being of a reflex nature, excited by the contact of the substance contained in the tube, with its lining membrane,—which produces an impression that is transmitted to the medulla oblongata, and is reflected back as a motor impulse to the muscles. We have sv here a distinct case of reflex action without sensation; for we have . \. no consciousness of the ordinary passage of food down the oesophagus, unless it occasion pressure on the surrounding parts through its bulk, or unduly irritate the lining membrane by its high or low temperature or its acrid qualities; and yet it may be shoAvn by experiment, that the >■ completeness of the nervous circle is requisite for the excitement of the movement which Avill not take place when it is interrupted either by division of the nerves, or by destruction or paralysis of the medulla oblongata. 260 OF FOOD AND THE DIGESTIVE PROCESS. 455. The progress of the food along the (Esophagus is aided by I* j r , the action of the muscular coat peculiar to it. This is composed of fi'f'3'''*^ ,} nfHhe non-striated fibre ; and, like that of the intestinal canal further on, ji!" J.*^* it is usually stimulated to contraction by the direct contact of the l'^^*"^1' "stimulus, and not either by the will, or by the reflex action of the spinal '-' ' ' cord. The movement produced by it is of the peristaltic or wave-like >••* kind; the contractions being limited to one portion of the tube, and r ': being propagated along it from above downwards. This action con- ", ;//* tinues after the division of all the nerves supplying the oesophagus; T',--'"'" and it cannot, therefore, be dependent upon the brain or spinal cord. It may be observed to take place in a rhythmical manner (that is, at short and tolerably regular intervals), whilst a meal is being SAvallowed; but as the stomach becomes full, the intervals are longer and the ■ • . wave-like contractions less frequent. The degree in which the action of the oesophagus alone, without that of the surrounding muscles, is capable of propelling the food into the stomach, seems to vary in diffe- rent animals. When the latter are paralysed in the Dog, by section of the nerves that supply^them, the food that has entered the oesophagus is still propelled into the stomach; but this is not the case in the Rab- bit, the action of its oesophageal fibres not being sufficient to carry the food onwards to the stomach, though it will expel it from the divided extremity of the tube when it is cut across. The usual peristaltic movements of the oesophagus are reversed in Vomiting : and this rever- ; sion has been observed even after the separation of the stomach from the oesophagus, as a consequence of the injection of tartar emetic into the veins. k 456. At the point where the oesophagus enters the Stomach,—the ; cardiac orifice of the latter,—there is a sort of sphincter, or circular muscle, which is usually closed. This opens when there is a sufficient pressure on it, made by the accumulated food propelled by the move- ments of the oesophagus above; and it then closes again, so as to*retain the food in the stomach. The closure is due to reflex action ; for when the nerves supplying it are divided, the sphincter no longer contracts, and the food regurgitates into the oesophagus. The opening of the car- diac orifice is one of the first acts which takes place in vomiting. 457. In ruminating animals, there is a very remarkable conforma- Fig. 77. Compound Stomach of Sheep :—a, oesophagus ; 6, paunch; c, second, or honeycomb stomach; d, third stomach, or many-plies; e, fourth stomach or reed;/, pylorus. tion at the lower end of the oesophagus, which is destined to regulate the passage of food into the different compartments of the stomach, DIGESTIVE APPARATUS OF RUMINANTS. 261 according as it has been submitted to the second mastication, or not. The oesophagus (Fig. 77, a) does not terminate at its opening into the first stomach or paunch (b), but it is continued onwards as a deep groove with two lips (Fig. 78): by the closure of these lips it is made to_ form a tube, which serves to convey the food onwards into the third stomach ; but Avhen they separate, the food is allowed to pass either into the first or the second stomach. When the food is first swallowed, it undergoes but very little mastication; it is consequently firm in its consistence, and is brought down to the termination of the oesophagus in dry bulky masses. These separate the lips of the groove or demi-canal (Fig. 78, e, e), and pass into the first and second stomachs. After they have been macerated in the fluids of these cavities, they are returned to the mouth by a reverse peristaltic action of the oesophagus; this return takes place in a very regular manner, the food being shaped into globular pellets by compression within a sort of mould formed by Fig. 78. Section of part of the Stomach of the Sheep, to show the demi-canal of the oesophagus; the mucous mem- brane is for the most part removed, to show the arrangement of the muscular fibres. At a is seen the termination of the oesophageal tube, the cut edge of whose mucous membrane is shown at 6. The lining of the first stomach is shown at c, c; and the mucous membrane of the second stomach is seen to be raised from the subjacent fibres at d. At e, e, the lips of the demi-canal are seen bounding the groove, at the lower end of which is the entrance to the third stomach or many-plies. the demi-canal, and these pellets being conveyed to the mouth at regular intervals, apparently by a rhythmical movement of the oesophagus. It is then subjected to a prolonged mastication within the mouth (the "cheAving of the cud"), by which it is thoroughly triturated and im- pregnated Avith saliva ; after Avhich it is again swallowed in a pulpy semi-fluid state. It noAV passes along the groove which forms the con- tinuation of the oesophagus, Avithout opening its lips ; and is thus con- veyed into the third stomach (Fig. 77, d), whence it passes to the fourth 262 OF FOOD AND THE DIGESTIVE PROCESS. (e), in which alone the true digestive process takes place. Noav, that the condition of the food as to bulk and solidity is the circumstance which determines the opening or closure of the lips of the groove, and which consequently occasions its passage into the first and second stomachs, or into the third and fourth, appears from the experiment of Flourens; who found that when the food, the first time of being SAvallowed, was artificially reduced to a soft and pulpy condition, it passed for the most part along the demi-canal into the third stomach, as if it had been ruminated, —only a small portion finding its way into the first and second stomachs. How far the actions of this curious apparatus are dependent upon ner- vous influence,—or how far they are due to the exercise of the contrac- tility of the muscular fibre, directly excited by the contact of the sub- stances with the lining membrane of the tubes and cavities,—has not yet been clearly ascertained. 458. The food, when introduced into the stomach, and submitted to the solvent action of its secretions, is also subjected to a peculiar move- ment, which is effected by the muscular walls of that organ. The pur- pose of this motion is obviously to keep the contents of the stomach in that state of constant agitation, which is most favorable to their chemical solution ; and particularly to bring every portion of the ali- mentary matter into contact Avith the walls of the stomach, so as to be subjected to the action of the fluid, which is poured forth from them during the digestive process. The movement is produced by the alter- nate shortening and relaxation of the various fasciculi, which are dis- posed in almost every direction throughout the muscular wall of the stomach; and it seems to produce a kind of revolution of the contents of the stomach, sometimes in the direction of its length, and sometimes transversely. Its result is well shown in the hair-balls, which are occa- sionally found in the stomachs of animals, that have swallowed hair from time to time through licking their skins ; the component hairs not being pressed together confusedly, but being worked together in regular directions, and so interwoven that they cannot be readily separated. As digestion proceeds, the dissolved fluid escapes, little by little, through the pyloric orifice, which closes itself firmly against the passage of solid bodies; and this motion continues until the stomach is completely emptied; when it ceases, until food is again introduced. The bulk of the alimentary mass diminishes rapidly, as the solvent process is near its completion ; and the separation of the fluid product or chyme is aided by a peculiar action of the transverse fasciculi, which surround the stomach at about four inches from its pyloric extremity. These shorten in such a manner, as to produce a sort of hour-glass separation between the portions of the stomach on either side of it; and the fluid solution, being received by the pyloric or smaller portion, is pumped away through the pylorus; whilst the solid matter yet undissolved is retained in the larger division. 459. The degree in which these movements are dependent upon the nervous system, or are under its control or direction, has not yet been clearly ascertained. Distinct movements may be excited in the stomach of a Rabbit, if it be distended with food, by irritating the par vagum soon after the death of the animal; these movements seem to commence MOVEMENTS OF STOMACH AND INTESTINE. 263 from the cardiac orifice, and then to spread themselves peristaltically along the Avails of the stomach; but no such movements can be excited if the stomach be empty. On the other hand, there is distinct proof that all the movements necessary to digestion may take place after the section of that nerve; although the first effect of the operation appears to be to suspend them completely. It is probable that the movements of the stomach are more regular and energetic in Herbivorous animals, Avhose food is difficult of digestion, than they are in the Carnivora, whose aliment is dissolved with comparative facility. 460. From the time that the ingested matter enters the Intestinal tube, it is propelled onAvards by the peristaltic contractions of its mus- cular coat; which are excited, independently of all nervous influence, by the contact of the aliment, or by that of the secretions mingled Avith it in its passage along the canal. These last appear to have an impor- tant effect; for we find that, when the bile-duct is tied, so as to prevent the bile from entering the intestine, constipation always occurs; whilst an increase of the biliary and otheV secretions, consequent upon the action of mercury or upon any other cause, produces an increased peristaltic movement, and a more rapid discharge of the excrementitious matter. During the passage of the alimentary matter along the small intestine, as we shall see hereafter, a large proportion of its fluid is removed, by the absorbent poAver of the villi; and the residue is again brought, therefore, to a more solid consistence. This residue consists in part of those portions of the aliment, which are not capable of being dissolved or finely divided, so as to be received by the absorbents; and in part of the matters poured into the alimentary canal, by the various glands that discharge their contents into it, for the purpose of being carried out of the body. The faeces Avhich are thus formed, are pro- pelled through the large intestine, by the continued peristaltic action of its walls, until they arrive at the rectum. 461. That the ordinary peristaltic action of the intestinal canal is independent of nervous influence, is sufficiently indicated by the fact, that it will continue when the tube is completely separated from all connexion with the nervous centres; as well as by the difficulty, already adverted to (§ 353), of exciting contractions in the muscular coat by any stimulation of its nerves. All the nerves of the intestine, from its commencement at the pyloric orifice of the stomach to its termination at the anus, are derived from the ganglia of the sympathetic system; but there is evidence that those Avhich influence its movements are really derived from the spinal cord (see chap, xii.) Although the will has no influence whatever on the peristaltic movement, yet the emotions seem to affect it; and it is probably to convey their influence, that the intestinal canal is supplied Avith motor nerves. It is also furnished with sensory nerves, which form part of the trunks of the sympathetic system, but which really pass onwards to the brain ; these do not, how- ever, make us conscious of the passage of the alimentary matter along the canal, so long as it is in a state of health; but in various diseased conditions, they give rise to sensations of the most painful nature. 462. For the occasional discharge of the faeces from the rectum, and for the retention of them at other times, we find the outlet or anal 264 OF FOOD AND THE DIGESTIVE PROCESS. orifice provided with an additional muscular apparatus, Avhich is con- nected with the spinal system of nerves. The act of Defecation is due to the pressure upon the contents of the rectum, which is occasioned by the combined contraction of the diaphragm and the abdominal muscles ; whilst, on the other hand, the retention of the faeces is due to the con- tractile power of the sphincter muscle which surrounds the anus. The action of the sphincter ani, like that of the sphincter of the cardia, is a reflex one; dependent upon the connexion of the muscle, by excitor and motor nerves, with the spinal cord. If the lower portion of the cord be destroyed, or if the nerves be divided, the sphincter loses its con- tractile power, and becomes flaccid. When in proper action, however, its power is sufficient to prevent the escape of the contents of the rec- tum ; until the expulsive force becomes very strong, in consequence either of the quantity of faeces which have accumulated, or the acridity of their character. In either case, the impression made upon the mucous membrane of the rectum is conveyed to the spinal cord; and, by a reflex motor impulse, the mMscles of defecation are thrown into combined action, the resistance of the sphincter is overcome, and the faeces are expelled. An unduly irritable state of the mucous membrane, or a disordered state of the excrementitious matter (resulting from the irritating character of the substances swalloAved, from the acrid charac- ter of the secretions poured into the canal, or from an unusual change in the aliment during the digestive process), may occasion unduly frequent calls upon the muscles of defecation, which the sphincter is unable to resist. On the other hand, if the progress of the faeces be delayed in the large intestine, by deficient peristaltic movement, they accumulate higher up, and the act of defecation is not excited. 463. Although the sphincter ani on the one hand, and the muscles of defecation on the other, are called into action by the reflex poAver of the spinal cord, and are so far involuntary in their operation, yet they are also in some degree subject (in Man at least) to the influence of the will. The resistance of the sphincter may be increased by a voluntary effort, when it is desired to retain the faeces in opposition to the power of the expulsors; and it is only when the latter operate with excessive force, that they can overcome it. On the other hand, the expulsors may be called into action, or may be aided, by the will, when the stimu- lus to their movement received through the spinal cord Avould not other- wise be strong enough; and the fasces may thus be evacuated hy a voluntary effort, at a time when they Avould not othenvise be discharged. 4. Of the Secretions poured into the Alimentary Canal, and of Changes which they effect in its contents. 464. The whole Mucous Membrane of the Alimentary canal, from the mouth to the anus, is covered during health with that peculiar viscid secretion, termed mucus, of which the characters have been already described (§ 237). This is formed, partly, on the free surface of the membrane itself, but chiefly in the numerous follicles or depressions by which that surface is increased; and it appears destined for the protec- tion of the delicate highly vascular membrane from undue irritation, by i SALIVARY GLANDS AND THEIR SECRETIONS. 265 the contact of the substances which are passing through the alimentary ,A ■ ; tube. When these are unusually acrid, the secretion of mucus is aug-- mented in quantity, and is increased in viscidit}% so as to form an effec- tive sheath to the membrane, which Avould otherwise suffer severely.' When this secretion is deficient, the membrane is irritated by the con- tact of any but the blandest substances; and the class of remedies termed demulcents are useful in coating and protecting it. 465. JDuring the mastication of the food in the mouth, the Salivary f secretion is poured in, for the purpose of being mingled with it, and of v^ri rendering the act of mastication more easy. This secretion is formed gJtt-w.-^W by three pairs of glands,—the Parotid, the Sub-lingual, and the Sub-.* maxillary ; these arc composed of minute follicles, "^ Avhose diameter is about l-1000th of an inch, con- FiS- 79- nected together by branches of their duct, upon Avhich they are set like grapes upon their stalk, JjjjM jj|k -vX'^' *"* surrounded by a plexus of blood-vessels, and bound iHp^ffifBKfoi1 -« ^JP^1, together by areolar tissue. Within the follicles |||JH^^^HS^^^ are the true secreting cells (§ 238); by whose groAvth and development, the material of the se- cretion is separated from the blood. These sali- vary cells are often to be recognised in the saliva; they must not, however, be confounded with the epithelium-cells of the mucous membrane of the mouth, Avhich are much larger. The fluid ob- ""'""' ' iL^ir1 tained from the mouth is not pure saliva; for Lobule of Parotid Gland of ;,?, ,, ,, ,. ,i •, ir» • • i i • i new-born infant, filled with the mUCUS OI the mOUth itself IS mingled With mercury; magnified 50 dia- the secretion from the salivary glands. If the meters- ^ proportion of the former be considerable, it gives to the fluid of the ; {, u-,o mouth an acid reaction; whilst, if the latter be predominant (Avhich it is directly before, and during, the act of eating), the fluid of the mouth has an alkaline reaction. It may be sometimes observed, that the saliva of the mouth will strike a blue color with reddened litmus- paper, whilst it turns blue litmus-paper red; thus shoAving the presence both of an acid and an alkali in a state of imperfect neutralization. ^ 466. The solid matter of the Salivary secretion is about 1 per cent.^C, of the whole; and this consists in part of animal principles, and in „, part* of saline substances. The animal matter consists of ozmazome,,/p^ Atvvvi mucus, and a peculiar substance termed ptyaline or salivary matter ;j ,/ Avhich is soluble in water and insoluble in alcohol, and which is yet dif- ferent from both albumen and gelatine. This substance appears to have a decided effect in producing the metamorphosis of certain alimen- tary substances, on Avhich it acts like a ferment. Starch may be^ comrerted into sugar, and sugar into lactic acid, by its agency ; and, if\ acidified, it has a certain solvent poA\Ter for caseine, animal flesh, and/ ' . other proteine-compounds. Its chemical nature has not yet been pre- * t • cisely determined. The saline constituents of the Saliva are nearly \ *vv-v/l identical with those of the blood; the chlorides of sodium and potassium' *'- form considerably more than half; and the remainder consists chieflyv> ftp of the tribasic phosphate of soda, to Avhich the alkaline reaction of- the } fluid is due, Avith the phosphates of lime, magnesia, and iron. It is of' 266 OF FOOD AND THE DIGESTIVE PROCESS. the earthy phosphates, that the tartar which collects about the teeth is ;T~r "\. chiefly composed, the particles of these being held together by about cj''4^ 20 per cent, of animal matter; and the composition of the concretions, Avhich occasionally obstruct the salivary ducts, is nearly the same. 467. The quantity of Saliva formed during the twenty-four hours, has been estimated at from 15 to 20 ounces; but on this* point it is im- possible to speak with certainty. The secretion is by no means con- stantly flowing; indeed, it is almost entirely suspended, when the masticator muscles and tongue are at perfect rest, unless it be excited by any mental cause; and hence it is, that the mouth becomes dry during sleep, if it be not kept closed. The Aoav of Saliva takes place s just Avhen it is most wanted; that is, \rhen food has been taken into the mouth, and when the operation of mastication is going on. But it will also take place, especially in a hungry person, at the sight, or even at the idea, of savory food; as is implied by the common expression of the "mouth watering" for such an object. The influence thus exercised over it by the emotional state of the mind, is probably conveyed to the salivary glands by the Fifth pair; which contains many of the gray or organic filaments, and which seems to take the place, in the Head, of a distinct sympathetic system. 468. Having been conveyed into the stomach, the food is submitted «.', • to the action of the gastric fluid, Avhich is secreted in the walls of that organ. This fluid is not present in the empty stomach; its secretion being excited by the presence of food, or by the irritation of the walls V^V. of the organ by some solid body. In the intervals betAveen the diges- tive process, the mucous membrane is of a light pink hue; but it - becomes more turgid with blood, when the presence of food calls for ;., the activity of its secreting processes. It is of a soft and velvet-like appearance; and it is constantly covered with a very thin transparent vis- cid mucus, which has neither acid nor alkaline reaction. By applying V aliment or other stimulants to the internal coat of the stomach, and by observing the effect through a magnifying glass, numerous minute pa- £' pillae can be seen to erect themselves upon the mucous membrane, so as f to rise through the coating of mucus ; and from these is poured forth a %A^ . pure, limpid, colorless, slightly viscid fluid, having a distinctly acid fy "P\" • reaction, which is the Gastric juice. This fluid is secreted by follicles, which are lodged in the walls of the stomach, and which closely resemble ^'; those that elsewhere secrete mucus; but they are usually of more complex structure, and are more numerous. 469. If the Mucous membrane of the stomach be divided by a section perpendicular to its walls, it is seen to be made up, as it were, of tubu- lar follicles closely applied to each other ; their blind extremities resting upon the submucous tissue, and their open ends being directed towards the cavity of the stomach. In some situations these tubuli are short and straight; in other parts they are longer, and present an appearance of irregular dilatation or partial convolution (Fig. 80, 1). This is their usual character, especially near the cardiac orifice of the stomach; but :; . " ryear the pyloric orifice they have a much more complex structure (Fig. '80, 2). These tubular follicles are arranged in bundles or groups, and . are surrounded and bound together by a fine areolar membrane; and GASTRIC FLUID. 267 this also serves to convey vessels from the submucous tissue, which ramify among the follicles, and supply the materials for their secretion. Ffc. 80. Fig. 81. i:;y place in the stomach ; but a longer time is required to make it. This i. • is easily accounted for by the difference of the conditions ; for no ordi- nary agitation can produce the same effect Avith the curious movements of the stomach (§ 458); fresh gastric fluid is poured out, as it is wanted, during the natural process of digestion; and the continual removal of PROPERTIES OF PEPSINE AND OF GASTRIC FLUID. 269 the matter Avhich has been already dissolved, by its exit through the pylorus, is of course favorable to the action of the solvent upon the remainder. The quantity of food, which a given amount of gastric fluid can dissolve, is limited; precisely as in the case of the acidulous solu- ^X-JSth tion of pepsine. The marked influence of temperature upon its action/ is shoAvn by the fact, that fresh gastric fluid has scarcely any influence^ on the matter submitted to it, when the bottle is exposed to cold air,) instead of being kept at a temperature of 100°. Hence the use of a large quantity of cold water at meal-times, or of ice afterwards, must retard the digestive process. 473. The pulpy substance, which is the product of the reducing action of the gastric juice, is termed Chyme. Its consistence will of 7 'SJIawvj course vary, in some degree, with the relative quantity of solids and ( "" /' liquids ingested. In general it is grayish, semifluid, and homogeneous; and possesses a slightly acid taste, but is otherwise insipid. When the food has been of a rich oily character, the Chyme possesses a creamy aspect; but when it has contained a large proportion of farinaceous matter, it has rather the appearance of gruel. The state in which the various alimentary principles exist in it, has not yet been accurately determined ; the following, however, may be near the truth.—The Pro-1 , teine-compounds, Avhether derived from Animal or Vegetable food, are 'A^ all reduced to a state of solution, if the gastric digestion have been properly performed; and in this state, they have all the properties of Albumen.—Gelatine will be dissolved or not, according to its previous^, condition; if it exist in a tissue from which it cannot readily be ex- , tracted, it Avill pass forth almost unchanged; but Avhen ingested in a ^ t / , state of solution, it remains so ; and if it have been preAaously prepared '^^^w for solution by boiling, its solution is completed in the stomach. Its condition, hoAvever, is altered in the process; for it loses the power of gelatinizing, and cannot be precipitated by chlorine; so that it cannot be detected as such either in the blood or the chyle into which it is ' received.—The Gummy matters of Vegetables are dissolved, Avhen they's n exist in a soluble form; as in the case of pure gum, pectine, and dex- r 'V\*J*V trine or starch-gum. It does not appear, however, that any further \^^JJ:'%\^\j conversion of Starch is effected by the gastric fluid; for if no saliva be '/1^9'^ admitted into the stomach, no sugar is generated there by the meta- morphosis of the starch Avhich it may contain. But the continued intro- duction of the saliva ordinarily occasions the continuance of the process, /,, J - although the presence of the free acid of the gastric fluid in some degree W a,u^ ■'■ interferes Avith it; and it is not until this has been neutralized by thel^^U**-1- admixture of the biliary and pancreatic secretions, that the metamor->'<■ -vVk<•■'•' phosis of the starch is actively renewed. Any sugar that may have'*? *vfu been taken in as such, or that may have been produced from starch by the ! converting power of the saliva, is reduced to the state of complete solu- tion.—Oily matters do not appear to be in any way acted upon, other-. wise than by being set free by the solution of the envelopes which may*)^t.;-. ; •- ^tf have contained them (e. g. fat cells), and by being dispersed through J 7!^tX the mass ; their state of division, however, does not seem to be yet fine^ '°~ ,,/^j enough to uIIoav of their absorption.—Most other substances, as resins, Avoody fibre, horny matter, yellow fibrous tissue, &c, pass unchanged^ 'w- 270 OF FOOD AND THE DIGESTIVE PROCESS. from the stomach, and undergo no subsequent alteration in the intes- tinal canal; so that they are discharged among the faeces as completely useless. ~l- 474.'We have now to notice the conditions, under which the Gas- / trie fluid is secreted ; the knowledge of which is of great practical im- portance. We have seen that it is not poured forth, except Avhen food is introduced into the stomach, or when its Avails are irritated in some other mode; and there is reason to believe, that it is not preAdously secreted and stored up in the follicles, but that the act of secretion itself is due to the stimulus applied to the mucous membrane. The quantity of the fluid then poured into the stomach, hoAvever, is not regulated by the amount of food ingested, so much as by the Avants of the system ; and as only a definite quantity of food can be acted on by a given amount of gastric juice, any superfluity remains undissolved for some time,—either continuing in the stomach until a fresh supply of the solvent is secreted, or passing into the intestinal canal in a crude state, and becoming a source of irritation, pain, and disease. The use of a small quantity of salt, pepper, mustard, or other stimulating substances, appears to produce a gently stimulating effect upon the mucous mem- brane, and by causing an increased afflux of blood, to augment the •'■•'■' quantity of the gastric fluid poured forth. Any excess of these or other irritants, however, produces a disordered condition of the mucous mem- brane, Avhich is very unfavorable to the digestive process. It becomes red and dry, with an insufficient secretion of mucus; the epithelial lining is abraded, so that the mucous coat is left entirely bare ; and '■"'irregular circumscribed patches of a deeper hue, sometimes with small aphthous crusts, present themselves here and there on the walls of the stomach. Similar results follow excess in eating. When these changes are inconsiderable, the appetite is not much impaired, the tongue does not indicate disorder, and the digestive process may be performed; but , , ., >;" if they proceed further, dryness of the mouth, thirst, accelerated pulse, • ,; foulness of the tongue, and other symptoms of febrile irritation, mani- fest themselves ; and no gastric secretion can then be excited by the stimulus of food. Similar results may follow the excitement of the ,. emotions ; and those of a depressing nature seem especially to produce a pale flaccid condition of the mucous membrane, which is equally unfavorable to the due secretion of gastric fluid. 475. That the amount of the secretion is ordinarily proportioned to r,.•■■ ■ ithe Avants of the system,—that the introduction of any superfluous ?. ' aliment into the stomach is not only useless but injurious, as giving rise / to irritation,—that incipient disorder of the stomach may occur, render- ing it less fit than usual for the discharge of its important duties, without manifesting itself by the condition of the tongue,—that when the tongue does indicate disorder of the stomach, such disorder is usually consider- able,—and that every particle of food ingested, in such states as prevent the secretion of gastric fluid, is a source of fresh irritation,—are truths which cannot be too constantly kept in mind. There can be no doubt that the habit of taking more food than the system requires, is a very prevalent one ; and that it is persevered in, because no evil result seems to folloAV. But when it is borne in mind that this habit must keep the SECRETION OF GASTRIC FLUID. 271 stomach in a state of continual irritation, however slight, it can scarcely be doubted that the foundation is thus laid for future disorder of a more serious kind. Two circumstances especially tend to maintain this prac- tice in adults, independently of the mere disposition to gratify the palate. One is the habit of eating the same amount of food, as during the period of growth, when more Avas required by the system. The other is the custom of eating too fast; and this is injurious,—both by preventing sufficient mastication, and thus throwing on the stomach more than its proper duty,—and also by causing an over-supply of food to be ingested, before there is time for the feeling of satisfaction to replace that of hunger (§ 486). 476. Of the Albuminous, Saccharine, and other matters dissolved in >,. . the Chyme, there is reason to believe that part are absorbed through the blood-vessels so copiously distributed on the walls of the stomach (§ 492). The remainder, with the undissolved matters, pass into the duodenum, Avhere the chyme is mingled with the biliary and pancreatic secretions.—The secretion of Bile is evidently a process of the highest / importance in the economy; as we may judge alike from the size of the' -• Liver and the supply of blood it receives, and from the rapidly fatal effects of its suspension. That a part of it is purely excrementitious, and is poured into the intestinal tube for the purpose of being carried, out of the body, cannot be questioned; but there is strong evidence, that a part of it is destined to be absorbed again, after performing some action of importance upon the contents of the alimentary canal.—Iri :> all but the very lowest animals, we find traces of a bile-secreting appa^ f- ; ratus ; and this is almost constantly situated in the immediate neigh-; borhood of the stomach. In many cases, the secretion is poured directly, into the cavity of that organ; but in most, it is conveyed (as in Man) into the intestinal tube near its commencement. Hence it seems clear from the disposition of the biliary apparatus, that it has a purpose to serve in connexion with the digestive function, and is not destined solely for the elaboration of a product which is to be cast out of the body; since, if the latter were the case, that product would be carried out immediately, like the urinary excretion, and would not be discharged into the alimentary canal high up.—This conclusion is confirmed by experiment; for it has been shown that, if the bile-duct be divided, and be made to discharge its contents externally through a fistulous orifice in the walls of the abdomen, instead of into the intestinal canal, those '\ animals Avhich survive the immediate effects of the operation, exhibit indications of the imperfect performance of the digestive operation. At first they eat much, but their food does not seem to impart to them an adequate amount of nutrition; afterwards they lose their appetite, be- come thin, and usually die after an interval of some months passed in this state. If, however, they be allowed to lick the orifice, so as to receive the fluid discharged from it into their stomachs, these injurious results do not folloAV.—The observation of disease in the human subject leads to similar conclusions ; for, when the biliary secretion is deficient, or its floAV into the intestine is obstructed, the digestive processes are evidently disordered; the peristaltic action of the bowels is not duly performed; the faeces are white and clayey; and there is an obvious 272 OF FOOD AND THE DIGESTIVE PROCESS. insufficiency in the supply of nutriment prepared for the absorbent vessels. jjt 477. On the other hand, that one great object of the secretion is to withdraAV from the Blood certain products of the decomposition of the tissues, which would otherAvise accumulate in it, and Avould be deleterious to its character, is shown by evidence yet more decisive. We find that the action of the liver is constant, and not occasional, like that of the ,.,»' Salivary and Gastric glands; and that, if anything interfere with the secreting process, and thereby cause the accumulation of the elements of the bile in the blood, the effects of their presence are immediately manifested in the disorder of other functions, especially those of the nervous system (§ 399); and the continued suspension of the function leads to a fatal result, unless the elements of the bile are drawn off (as sometimes happens) by the urinary organs. When the secreting action of the liver has once been performed, an obstruction to the discharge of the bile into the intestine does not seem to be so immediately injurious. The fluid accumulates, and distends the bile-ducts and the gall-bladder; and Avhen they^ are completely filled, part of it is reabsorbed into the blood, apparently in a changed condition, since it does not then produce the same injurious effects, as result from the accumulation of the same materials, previously to the action of the Liver upon them. The color- ing matter seems to be very readily taken back into the circulating system; and is deposited by it in almost every tissue of the body. 478. Although the secreting action of the Liver is constant, yet the discharge of bile into the intestine is certainly favored by the presence of chyme in the latter. The purpose of the gall-bladder is obviously to permit the accumulation of bile, when it is not wanted in the intestine; and we find it most constantly present in those tribes of animals, which live upon animal food, and Avhich therefore take their aliment at inter- vals ; Avhilst it is more frequently absent in those herbivorous animals, in which the digestive process is almost constantly going on. The middle coat of the bile-ducts is clearly muscular, and has a peristaltic action like that of the intestinal canal; this action may be excited by galvanism, or by irritation of the branches of the Sympathetic nerve, by which it is supplied. The mucous coat of the ductus choledochus is disposed in valvular folds, in such a manner as to prevent the reflux of the bile or of the contents of the intestine; and a still further security is afforded by the valvular covering to the orifice of the duct, Avhich is furnished by the mucous covering of the intestine itself. The flow of bile into the intestine, when its presence is needed there, is commonly imputed to the pressure of the distended Duodenum against the gall- bladder ; but it is probable that the contractility of the muscular coat of the duct itself, which may be excited either through the sympathetic nerve, or by irritation at the orifice of the duct (as in the case of the Salivary glands), is the real cause of the discharge of the fluid. It is an interesting fact, Avhich proves how much the passage of the Bile into the Intestine is dependent upon the presence of aliment in the latter, that the gall-bladder is almost invariably found turgid in persons Avho have died of starvation ; the secretion haA'ing accumulated, through the want of demand for it, although there was no obstacle to its exit. SECRETION OF BILE. 273 479. The composition of the Bile, and the structure of the organ which elaborates it, will be more appropriately considered hereafter, Avhen the Secreting apparatus generally is being described (chap, ix.) At present we have to inquire what is the precise effect of its admixture with the products of digestion, and what is the purpose which this ad- mixture serves. In the first place, it may be stated that biliary matter is essentially a soap, formed by the union of a fatty acid with a soda- base ; and that it serves the purpose of neutralizing the acidity of the chyme, which is derived from the gastric juice; the biliary acids falling, v • doAvn as an insoluble precipitate, Avhen thus deprived of their soda. Further, the bile shares with the pancreatic fluid in that emulsifying poAver, by which the fatty matters of the food are reduced to a state of such fine division, as to be rendered capable of being absorbed; and thus it happens that the introduction of these matters into the system, through the medium of the lacteal absorbents (§ 494), does not take place until after the chyme has been mingled Avith the biliary and pan- creatic secretions. Again, it has been asserted (but the fact has not been fully substantiated), that the admixture of biliary matter produces a conversion of saccharine in fatty compounds. When fresh bile is mingled with neAvly-formed chyme, in a glass vessel, the mixture sepa- ja^ rates into three distinct parts; a reddish-brown sediment at the bottom, ^" a wdiey-colored fluid in the centre, and a creamy pellicle at the top.*, The central stratum probably contains the albuminous, gelatinous, sac- "■ charine, and other matters in a state of solution; the superficial pellicle may be looked upon as consisting chiefly of oleaginous matter destined for absorption ; whilst the sediment, partly consisting of the unreducible portion of the food, and partly of the biliary matter itself, is evidently excrementitious. 480. Tho Pancreatic secretion has a chemical constitution very ana- \ logous to that of Saliva; but the peculiar organic compound which it contains, has been found by M. Bernard to possess a power of emulsify- ing fatty matter, when mingled with it; and there is strong reason to believe that the chief purpose of this secretion is to effect such a change in the condition of the oleaginous constituents of the chyme, as may prepare them for absorption. But further, the neutralization of the acid of the gastric fluid now allows the metamorphosis of starch to be recom- menced ; and as there is eAddence that the production of sugar continues to take place during the passage of the chymous mass along the small intestines, in animals Avhose food is partly or completely vegetable, the pancreatic fluid, which has been experimentally ascertained to possess - this power, is probably the chief agent by Avhich the conversion is effected. It may be surmised, further, that the glandulae of Brunner (§ 450) par- ticipate in the functions of the Pancreas; being, perhaps, the chief agents in the elimination of that "succus entericus," which has been experimentally found to concur with the biliary and pancreatic fluids in the emulsification of fatty matters. 481. During the passage of the contents of the Intestine, now aug- mented by the biliary secretion, along the canal, the nutritious portion '" ' is gradually withdrawn by the absorbent vessels on its Avails; and the excrementitious matter alone remains, increased in amount by the pro- 18 274 OF FOOD AND THE DIGESTIVE PROCESS. ducts of the secretion of the Peyerian and other glandulae, with Avhich the mucous lining of the lower intestines is studded. Many of the loAver animals are furnished, at the part where the small intestine enters the large, with a caecum, resembling that which in Man is termed the vermi- form appendage of the caecum, but greatly exceeding it in size. Some- times Ave find two caeca instead of one ; and these are much prolonged, so as to form tubes of considerable length. It has been ascertained that, in herbivorous animals, a distinctly acid secretion is formed by the cascum, during the digestive process; and there is reason to believe, that the food there undergoes a second process, analogous to that to which it has been submitted in the stomach, and fitted to extract from it any undissolved alimentary matter which it may still contain. There is no reason to believe, however, that any such process takes place in Man, whose real caecum is rudimentary,—the part of the intestine which has received the name, being merely the dilated commencement of the colon. The act of Defecation, by Avhich the excrementitious matter is discharged, has been already noticed (§ 462); the Absorption of nutritive matter will be treated of in the succeeding Chapter. 5. Of Hunger, Satiety, and Thirst. 482. The want of solid aliment is indicated by the sensation of Hunger; and the deficiency of fluid by that of Thirst. On the other hand, the presence of a sufficiency of food or liquid in the stomach is indicated by the sense of Satiety. These sensations are intended as our guides, in regard to the amount of aliment we take in. What is the real seat of these sensations, and on what conditions do they de- pend? 483. The sense of Hunger is referred to the stomach, and seems im- mediately to depend upon a certain condition of that organ; but what that condition is, has not yet been precisely ascertained. It is not pro- educed by mere emptiness of the stomach, as some have supposed; for, if the previous meal have been sufficient, the food passes entirely from the cavity of the stomach, before a renewal of the sensation is felt. It cannot be due to the action of the gastric fluid upon the coats of the stomach themselves ; because this fluid is not poured into the stomach, except when the production of it is stimulated by the irritation of the secreting follicles. It has been attributed to distension of the gastric follicles by the secreted fluid ; but there is no evidence that the fluid is secreted before it is wanted; and moreover, it is well known that mental emotion can dissipate in a moment the keenest appetite, and it is diffi- cult to imagine how this can occasion the emptying of the follicles. Perhaps the most satisfactory view is that, which attributes the sense of hunger to a determination of blood to the stomach, preparing it for the secretion of gastric fluid; since this is quite adequate to account for the impression made upon the nerves; and it accords with what has just been stated of the influence of mental emotions, since we know that these have a powerful effect upon the circulation of blood in the minute vessels (§ 603). 484. Although the sense of Hunger is immediately dependent, in SENSE OF nUNGER AND SATIETY. 275 great part at least, upon the condition of the stomach, yet it is also in- dicative of the condition of the general system ; being extremely strong, when the body has undergone an unusual waste Avithout a due supply of food, even though the stomach be in a state of distension ; whilst it is not experienced, if, through the general inactivity of the system, the last supply has not been exhausted, even though the stomach has been long empty. It is well known that, when food is deficient, the attempt to allay the pangs of hunger by filling the stomach with non-nutritious substances, is only temporarily successful; the feeling soon returning with increased violence, though it has received a temporary check. The reason for this is obviously, that the general system has received no satisfaction, although the stomach has been caused to secrete gastric fluid by the contact of solid matter with its Avails: so that although the state on which hunger immediately depends, has been for a time relieved, this state is soon renewed, unless the solid matter introduced into the stomach be of an alimentary character, and be dissolved and carried into the system. 485. When the food is nutritious in its character, but of small bulk experience has shoAvn the advantage of mixing it with non-nutritious substances, in order to give it bulk and solidity ; for if this be not done, it does not exert its due stimulating influence upon the stomach; the gastric juice is not poured forth in proper quantity ; and the result is, that neither is the sense of hunger relieved, nor are the wants of the body satisfied. Thus the Kamschatdales are in the habit of mixing earth or sawdust with the train-oil, on which alone they are frequently reduced to live. The Veddahs or wild hunters of Ceylon, on the same principle, mingle the pounded fibres of soft and decayed wood with the honey on which they feed when meat is not to be had; and on one of them being asked the reason of the practice, he replied, " I cannot tell you, but I know that the belly must be filled." It has been found that soups and fluid diet are not more readily converted into chyme than solid aliment, and are not alone fit for the support of the body in health; and it is often to be observed, in disordered states of the stomach, that it can retain a small quantity of easily digested solid food, when a thin broth would be rejected. 486. The sense of Satiety is the opposite of Hunger; and like it, depends on tAvo sets of conditions,—the state of the stomach, and that of the general system. It is produced in the first instance by the ingestion of solid matter into the stomach, which gives rise to the feel- ing of fulness ; but this is only a part of the sensation which ought to be experienced ; and it is only Avhen the act of digestion is being duly performed, and nutritive matter is being absorbed into the vessels, that the peculiar feeling of satisfaction is excited, which indicates that the wants of the system at large are being supplied.—It has been very justly remarked by Dr. Beaumont, that the cessation of the demand set up by the system, rather than the positive feeling of satiety, should be the guide in regulating the quantity of food taken into the stomach. The sense of satiety is beyond the point of healthful indulgence ; and is Nature's earliest indication of an abuse and overburden of her powers 276 OF FOOD AND THE DIGESTIVE PROCESS. to replenish the system. The proper intimation is the pleasurable sen- sation which is experienced, when the cravings of the appetite are first allayed ; since, if the stomach be sufficiently distended with wholesome food for this to be the case, it is next to certain that the digestion of that food will supply Avhat is required for the nutrition of the body. It is only when the substance with which the stomach is distended, is not of a digestible character, that the feelings excited by the state of that organ are anything but a correct index of the wants of the system. 487. The Par Vagum is evidently the nerve, which conveys to the sensorium the impression of the state of the stomach, and which is therefore the immediate excitor of the sensation of hunger or of the feeling of satiety. But it is evident from experiments upon animals, that it is not the only source, through which they are incited to take food, and are informed Avhen they have ingested enough ; and it is probable that the Sympathetic nerve is the channel, through Avhich the wants of the system are made known, and through which, in particular, the feeling of general exhaustion is excited, that is experienced when there has been an unusual waste, or Avhen the proper supply has been too long withheld. 488. The conditions of the sense of Thirst are very analogous to those of hunger ; that is, it indicates the deficiency of fluid in the body at large; but the immediate seat of the feeling is a part of the ali- mentary canal,—not the stomach, however, but the fauces. It is re- lieved by the introduction of fluid into the circulating system, through any channel; whilst the mere contact of fluids with the surface to which the sensation is referred, produces only a temporary effect, un- less absorption takes place. If liquids be introduced into the stomach by an cesophagus-tube, they are just as effectual in allaying thirst, as if they Avere swallowed in the ordinary manner; and the same result follows the injection of fluid into the veins (as was most remarkably the case when this method of treatment Avas practised in the Asiatic Cholera), or the absorption of fluid through the skin or the lower part of the alimentary canal. The deficiency of fluid in the body may arise,—and Thirst may consequently be induced,—either by an un- usually small supply of fluid, or by excessive loss of the fluids of the body, as by perspiration, diarrhoea, &c. But it may also be occasioned by the impression made by particular kinds of food or drink upon the alimentary canal; thus salted or highly-spiced meat, fermented liquors when too little diluted, and other similar irritating agents, excite thirst; the purpose of Avhich sensation is evidently to cause the ingestion of fluid, by which these substances may be diluted, and their irritating action prevented. ABSORPTION AND SANGUIFICATION. 277 CHAPTER V. ABSORPTION AND SANGUIFICATION. 1. Absorption from the Digestive Cavity. 489. So long as the Alimentary matter is contained in the digestive cavity, it is as far from being conducive to the nutrition of the system, as if it Avere in contact with the external surface. It is only when absorbed into the vessels, and carried by the circulating current into the remote portions of the body, that it really becomes useful in main- taining the vigor of the system, by replacing that which has decayed, and by affording the materials for the various organic processes Avhich are continually going on. Among the Invertebrated animals, Ave find the reception of alimentary matter into the circulating system to be entirely accomplished through the medium of the veins, Avhich are dis- tributed upon the walls of the digestive cavity. We not unfrequently observe, that the intestinal tube is completely enclosed within a large venous sinus, so that its whole external surface is bathed with blood ; and into this sinus, the alimentary materials Avould appear to transude, through the walls of the intestinal canal, to become mingled with the blood, and to be conveyed with its current into the remote portions of the body. Among the Vertebrata, we find an additional set of vessels, interposed betAveen the walls of the intestine and the sanguiferous sys- tem, for the purpose, as it would seem, of taking up that portion of the nutritive matter Avhich is not in a state of perfect solution, and of pre- paring it for being introduced into the current of the blood. These vessels are the lacteals or absorbents. They are very copiously distri- buted upon the walls of the smaller intestine, commencing near the entrance of the biliary and pancreatic ducts; the walls of the large intestine are less abundantly supplied with them, and they do not show themselves in the villi Avhich are found on some parts of the lining mem- brane of the stomach, although the walls of that viscus are supplied with lymphatic absorbents. 490. Nevertheless it is quite certain, that substances may pass into the current of the circulation, which have been prevented from passing further than the stomach; thus, if a solution of Epsom-salts be intro- duced into the stomach of an animal, and its passage into the intestine be prevented by a ligature around the pylorus, its purgative action will be exerted nearly as soon, as if the communication between the stomach and intestines had been left quite free; or if a solution of prussiate of potash be introduced into the stomach under similar circumstances, the presence of that salt in the blood may be speedily demonstrated by chemical tests. It appears from the experiments of MM. Tiedemann and Gmelin, that when various substances were min- gled with the food, Avhich, by their color, odor, or chemical properties might be easily detected,—such as gamboge, madder, rhubarb, camphor, 278 ABSORPTION AND SANGUIFICATION. musk, asafoetida, and saline compounds,—they Avere seldom found in the chyle, though many of them were detected in the blood and in the urine. The coloring matter appeared to be seldom absorbed at all; the odorous substances were generally detected in the venous blood and in the urine, but not in the chyle ; whilst, of the saline substances, many Avere found in the blood and in the urine, and only a very few in the chyle. 491. This passage of substances in a state of perfect solution, from the stomach into the blood-vessels, is probably due to the operation of that peculiar modification of Capillary Attraction, which is called En- dosmose. When tAvo fluids differing in density are separated by a thin animal or vegetable membrane, there is a tendency to mutual admixture through the pores of the membrane ; but the less dense fluid will transude Avith much greater facility than the more dense; and consequently there will be a considerable increase on the side of the denser' fluid; Avhilst very little of this, in comparison, will have passed towards the less dense. When one of the fluids is contained in a sac or cavity, the flow of the other toAvards it is termed Endosmose, or floAV-inAvards; whilst the contrary current is termed Exosmose or floAv-outwards. Thus if the caecum of a fowl, filled Avith syrup or gum-Avater, be tied to the end of a tube, and be immersed in pure water, the latter Avill penetrate the caecum by Endosmose, and will so increase the volume of its contents, as to cause the fluid to rise to a considerable height in the attached tube. On the other hand, a small proportion of the gum or syrup will find its way into the surrounding fluid by Exosmose. But if the caecum were filled with water, and Avere immersed in a solution of gum or sugar, it would soon be nearly emptied, the Exosmose being much stronger than the Endosmose. It is in this manner that Ave may cause the flattened corpuscles of the blood to be distended into spheres, by treating them with water; or may empty them almost completely, by immersing them in syrup (§ 215); since their contents are more dense than the surrounding fluid in the first case, so that they will be augmented by Endosmose; whilst they are less dense in the second, so as to be diminished by Exosmose. 492. Now it seems to be in this manner, that substances contained in the cavity of the stomach, and perfectly dissolved by its fluids, are received into the blood-vessels; for as the blood is the fluid of greater density, it will have a tendency to draw towards it, by Endosmose, the saline and other matters, which are in a state of perfect solution in the stomach. The Mucous membrane, which forms the inner wall of that organ, is most copiously supplied with blood-vessels; partly, indeed, that they may afford the materials of the gastric secretion; but partly, also, that they may take up the substances, which are capable of entering the circulating current by this direct channel. The move- ment of blood through the vessels, tends to accelerate the permeation of liquids through their walls, in a very remarkable degree: as may be shown by the following simple experiment. If a membranous tube, such as a piece of the -small intestine or of a large vein of an animal, be fixed by one extremity to an opening at the bottom of a vessel ABSORPTION OF SOLUBLE MATTERS INTO THE VEINS. 279 filled with Avater, and have a stopcock attached at the other extre- mity, and be then immersed in Avater acidulated with sulphuric or hydrochloric acid, it will be some time before the acid will penetrate to the interior of the tube, Avhich is distended with water; but if the stopcock be opened, and the water be allowed to discharge itself, the presence of the acid will be immediately discovered (by tincture of litmus) in the liquid which flows out. Thus the continuance of the Cir- culation is obviously one of the most important of the conditions of Absorption. It is whilst passing through the system of capillaries, which forms a minute plexus immediately beneath the free surface of the mucous membrane, that the blood thus receives an admixture of the soluble matters contained within the digestive cavity; and hence it is that these substances are detectible in the blood of the gastric and mesenteric veins, sooner than in any part of the arterial system. They are very rapidly diffused, however, through the general circulation ; and may even show themselves in the excretions within so short a period, that it is obvious that they must have been absorbed immediately on their introduction into the stomach. Thus Mr. Erichsen found that he Avas able to detect the presence of ferrocyanide of potassium in the urine, within one minute after it had been swallowed in solution. This, however, was only Avhen it Avas taken after a long fast; more commonly the absorption is less rapid; and if the substance be introduced within an hour or two after a full meal, it may be as much as half an hour before its presence in the urine gives evidence of its having been received into the circulating current. Although it is difficult to speak with certainty on the point, yet there appears a strong probability, that, both in the stomach and intestinal tube, the absorption of nutritive matters in a state of perfect solution (such as gum, sugar, pectine, gelatine, and soluble albumen) is thus accomplished through the medium of the blood- vessels ; Avhich also take up the chief supply of water that is required by the system. It is difficult else to see the purpose of the extraordinary vascularity of the mucous membrane, and in particular of those filaments Distribution of Capillaiies in the Villi of the Intestine. or narrow folds, termed villi, which so thickly cover its surface. Each of these villi is furnished Avith a plexus of minute blood-vessels, of which the larger branches may even be seen with the naked eye, when they are distended with blood or with a colored injection. By these villi, the vascular surface of the mucous membrane is enormously extended. In Man, they are commonly cylindrical or nearly so, and are from about a quarter of a line, to a line and a half in length; but 280 ABSORPTION AND SANGUIFICATION. in many of the loAver animals, they are spread out into broader laminae at the base, and are connected together so as to form ridges or folds. 493. The nutritive materials taken up by the blood-vessels of the ali- mentary canal, are not conveyed directly into the general circulation; for they are first submitted to the agency of the Liver. All the veins which return the blood from the gastro-intestinal capillaries, converge into the portal trunk, which distributes it to the various portions of that secreting apparatus; and there is strong reason to believe, that not merely is the fluid there depurated of some matters Avhose presence Avould be injurious, but that the Liver exercises a poAverful assimilating action upon the proper nutritive substances, rendering them fitter to become components of the Blood. For the blood of the portal vein, when examined during digestion, is found to contain a large proportion of albumen and comparatively little fibrine ; whilst in that Avhich has passed through the liver, the amount of fibrine has undergone a large increase. Again, it appears that fatty matters are elaborated in the liver, either from saccharine substances, or from albuminous compounds; for even when no fat can be detected in the blood of the vena portae, that of the hepatic vein contains it in considerable amount. So, again, it appears that the liver elaborates from some other constituents of the blood a saccharine compound (diabetic sugar), which is destined for im- mediate elimination by the lungs, and which, being much more readily carried off by the respiratory process than either grape-sugar or cane- sugar, may be regarded as its most appropriate pabulum. Further, if white of egg mixed Avith Avater be injected into any of the systemic veins, distinct evidence of the presence of albumen is speedily traceable in the urine ; showing that this substance has not been properly assimilated. But if the same fluid be injected into the portal system, no trace of its presence in the urine is found. So, again, when a solution of sugar is injected into the general venous system, this substance soon sIioavs itself in the urinary excretion; but if the same injection be made into the vena portas, so that the sugar is obliged to pass through the liver, no such elimination takes place, it being then assimilated with the blood. The liver, however, is not required to effect a corresponding change in the fatty matters taken up from the food; for these are received into the blood through the absorbents, rather than through the sanguiferous vessels ; and it is found that if fatty matters be injected into the general circulation, no effect is produced on the urine.* 494. Every one of the intestinal Villi, however, also contains the commencement of a proper lacteal vessel; the portion of the absorbent system specially adapted for the reception of alimentary matters from without, being thus distinguished, on account of the milky aspect of the fluid Avhich is found Avithin it. The folloAving figure (83) represents the appearance offered by the incipient lacteals, in a villus of the jejunum of a young man, who had been hung soon after taking a full meal of farinaceous food. The trunk that issues from the villus is formed by the confluence of several smaller branches, whose origin it is difficult to trace ; but it is probable that they form loops by anastomosis with each * See the recent Lectures and Memoirs of M. CI. Bernard, in "L'Union Me"dicale," and the " Gazette Me"dicale," for 1850. ABSORPTION OF ALCOHOL, ETC.—MESENTERIC GLANDS. 281 other, so that there is no proper free extremity in any case. It is quite certain that the lacteals never open by free orifices upon the surface of the intestine, as Avas formerly imagined. And there seems good reason to believe, that either by the cells asserted by Prof. Goodsir to be developed Avithin the free extremities of the villi, or by the epi- thelial cells which cover their extremities (§ 243), a selection is made of those substances Avhich are proper FiS- 83- to be received into the special Absorbent system. 495. It is particularly important to keep in view the difference betAveen the tAvo modes, by which ali- mentary substances are introduced into the system, when we are treating those disordered states, in which the digestive process is imperfectly performed, or is altogether suspended. There can be little doubt that the immediate cause of death, in many diseases of exhaustion, is the want of power to main- tain the heat of the body; the stomach not being r i . v . p i i.i -ii i Uneoltae Intestinal Villi, able to digest iood, and the special absorbent poAver with the commencement of of the lacteals being altogether suspended, so that alacteal- the inanition is as complete, as if food were altogether withheld. Now under such circumstances, it becomes a matter of the greatest im- portance to present a supply of combustible matter, in such a form that it may be introduced into the circulating system by simple Endosmose ; and the value which experience has assigned to broths and to thin fari- naceous solutions, and still more, to diluted alcoholic drinks, frequently repeated, under such circumstances, seems to depend in great part upon the facility with which they may be thus absorbed. The good effects of alcohol, cautiously administered, are no doubt owing in part to its specific influence upon the nervous system; but that they are also due to its heat-producing power, appears from the results of the administra- tion of frequently-repeated doses, in states of utter exhaustion,—the temperature of the body being kept up so long as they are continued, and falling when they are intermitted (§ 118). As the alcohol is thus burned off nearly as fast as it is introduced, it never accumulates in sufficient quantity, to produce its usual violently-stimulating effects upon the nervous system. 2. Passage of the Chyle along the Lacteals, and its admixture with the Lymph collected from the general System. 496. The Lacteal vessels, which commence on the surface of the in- testines, run together on their walls, and form larger trunks, which converge and unite with each other in the mesentery; and the main trunks thus formed then enter certain bodies, which are commonly knoAvn as the " mesenteric glands." Their structure, hoAvever, does not seem to correspond with that of the proper glands ; as they are simply composed of lacteal trunks, convoluted into knots, and dilated into larger cavities, amongst which bloodvessels are minutely distributed. These blood-vessels have no direct communication with the interior of the lacteals; but are separated from them by the membranous walls of 282 ABSORPTION AND SANGUIFICATION. both sets of tubes. The epithelium, which lines the absorbent vessel, undergoes a marked change Avhere the vessel enters the gland, and be- comes more like that of the proper glandular follicles in its character. Instead of being flat and scale-like, and forming a single layer in close apposition with the basement-membrane, as it does in the lacteal tubes before they enter the gland and after they have emerged from it, we find it composed, within the gland, of numerous layers of spherical nu- cleated cells (Figs. 84 and 85); of which the superficial ones are easily detached, and appear to be identical with the cells that are found float- ing in the chyle. The purpose of the cells will be presently inquired into. Fig. 84. Fig. 85. Diagram of an Absorbent Gland, showing the Portion of intra-glandular Absorbent, showing intra-glandular network, and the transition from along the lower edge the thickness of the germi- the scale-like epithelia of the exlra-glandular ab- nal membrane, and upon it, the thick layer of sorbents, to the nucleated cells of the intra-glan- glandular epithelial cells. dular. 497. After emerging from the mesenteric glands, the lacteal trunks converge, with occasional union, until they discharge their contents into the receptaculum chyli, which is situated at the front of the body of the second lumbar vertebra. Into the same cavity are poured the contents of a part of the other divisions of the Absorbent system; which is dis- tributed through the body in general, and which, from the transparency of the fluid or lymph it contains, is termed the lymphatic system. From the receptaculum chyli, arises the thoracic duct; which passes upwards in front of the spine, receiving other lymphatic trunks in its course, to terminate at the junction of the left subclavian and jugular veins ; where it delivers its contents into the sanguiferous system (Fig. 86). A smaller duct receives some of the lymphatics of the right side, and there termi- nates at a corresponding part of the venous system; but it does not receive any of the contents of the lacteals. 498. The Lymphatic system is evidently allied very closely to the lacteal, in its general purposes; and makes its first appearance in the same class of animals, namely, in fishes. The vessels of which it is composed are distributed through most of the softer tissues of the body, and are particularly abundant in the skin. They have never been'found to commence by closed or open extremities ; but seem to form a net- work, from Avhich the trunks arise. In their course they pass through glandulae, disposed in different parts of the body, which exactly re- semble in structure those which are found upon the lacteals in the mesentery. And they at last terminate, as already shown, in the same general receptacle with the lacteals. Hence it cannot be reasonably doubted that the fluid which they absorb from the various tissues of the body, is destined to become again subservient to nutrition ; being poured back into the current of the blood, along with the new materials, which LACTEAL AND LYMPHATIC SYSTEMS. 283 Fig. 86. are now for the first time being introduced into it. That the special Absorbent apparatus of Vertebrated animals has for part of its functions, to effect a change in the materials absorbed, and thus to aid in fitting them for introduction into the blood, seems apparent from the facts of Comparative Anatomy ; which show that, the more distinct the blood is from the chyle and lymph, the more marked is the provision for delaying the latter in the absorbent system, and for subjecting it to preliminary change. 499. The course of the Absorbent vessels in Fishes is short and simple; they are not furnished with glands ; and they pour their contents into the blood-vessels at several different parts of the body. In this class the blood contains fewer red corpuscles, and its coagulating power is feebler, than in any other Vertebrata. And in the lowest tribes, in which the Vertebrated character is almost en- tirely wanting, and in Avhich the blood is almost pale, no special ap- sorbent system has yet been dis- covered.—In Reptiles, the length of the Absorbent vessels is remarkably increased by their doublings and convolutions; so that the system ap- pears to be more highly developed, than in either of the warm-blooded classes. But this superiority is not real; for there is yet no trace of the glands, Avhich concentrate, as it were, the assimilation power, of a long se- ries of vessels. Moreover, we often find the lymphatics of this class fur- nished with pulsating dilatations, or lymphatic hearts ; which have for their office to propel the lymph into the venous system. In the Frog there ar e t wo pair s of these; one situ- ated just beneath the skin (through Avhich its pulsations are readily seen in the living animal), immediately behind the hip-joint; the other pair being more deeply seated at the upper part of the chest. The former receive the lymph of the posterior part of the body, and pour it into the veins proceeding from the same The course and termination of the Thoracic Duct. 1. The arch*of the aorta. 2. The thoracic aorta. 3. The abdominal aorta; showing its principal branches divided near their origin. 4. The arteria innominata, dividing into the right carotid and right subclavian arteries. 5. The left carotid. 6. The left subclavian. 7. The su- perior cava formed by the union, of, 8, the two venae innominatae; and these by the junction, 9, of the internal jugular and subclavian vein at each side. 10. The greater vena azygos. 11. The termination of the lesser in the greater vena azygos. 12. The receptaculum chyli; several lymphatic trunks are seen opening into it. 13. The thoracic duct dividing opposite the middle of the dorsal vertebras into two branches, which soon reunite ; the course of the duct behind the arch of the aorta and left subclavian artery is shown by a dotted line. 14. The duct making its turn at the root of the neck, and receiving several lymphatic trunks previously to terminating in the posterior aspect of the junction of the internal jugular and subclavian vein. 15. The termination of the trunk of the ductus lymphaticus dexter. 284 ABSORPTION AND SANGUIFICATION. part; the latter collect that which is transmitted from the anterior part of the body and head, and empty their contents into the jugular vein. Their pulsations are totally independent of the action of the heart, and of the respiratory movements ; since they continue after the removal of the former, and for an hour of two subsequently to the death and com- plete dismemberment of the animal. They usually take place at the rate of about sixty in a minute; but they are by no means regular, and are not synchronous on the two sides. 500. In Birds, we find the Absorbent system existing in a more per- fect form; its diffused plexuses and convolutions being replaced by glands; in which the contained fluid is brought into closer proximity with the blood, and in which it is subjected to the influence of assimilating cells. These, however, are not very numerous; being principally found on the lymphatics of the upper extremities. The absorbents, in this class, terminate principally by two thoracic ducts, one on each side, which enter the jugular veins by several orifices. There are, however, two other entrances, as in Reptiles, into the veins of the lower extremity; and these are connected by two large dilatations of the lymphatics, which are evidently analogous to the lymphatic hearts of Reptiles, but which have little or no power of spontaneous contraction.—In Mam- malia, the Absorbent system presents itself in its most developed and concentrated state. The vessels possess firmer walls, and are more copiously provided with valves, than in the classes beneath; and the glands are much more numerous, particularly upon the vessels that receive or imbibe substances from without,—as those of the digestive cavity, the skin, and the lungs. The terminations of the absorbents in the veins are usually restricted, as in Man, to the single point of en- trance of the thoracic duct on either side ; but they are sometimes more numerous ; and certain variations in the arrangement of the thoracic ducts, which occasionally present themselves as irregularities in Man, are the ordinary condition of these parts in some of the lower Mam- malia. 501. With regard to the source of the matters absorbed by the Lymphatics, it is difficult to speak with certainty. We shall presently see that their contents bear a close resemblance to the fluid element of the blood, or " liquor sanguinis," in a state of dilution; it is very probable that they partly consist of the residual fluid, Avhich, having escaped from the blood-vessels into the tissues, has furnished the latter with the materials of their nutrition, and is now to be returned to the former. But they may include, also, those particles of the solid frame- work which have lost their vital powers, and which are, therefore, not fit to be retained as components of the living system, but Avhich have not undergone a degree of decay which prevents them from serving, like matter derived from the dead bodies of other animals, as a material for re- construction, when it has been again subjected to the organizing process. 502. It was formerly supposed (and the doctrine Avas particularly in- culcated by the celebrated John Hunter) that the office of the Lymphatic system is to take up and remove all the effete matter, that is to be cast out of the body, being no longer fit for its nutrition. But for such a supposition there is no adequate foundation. It seems absurd to ima- ABSORPTION BY THE LYMPHATICS. 285 gine, that this effete matter Avould be mingled with the newly-ingested aliment, and Avould be poured back with it into the general current of the circulation, instead of being at once carried out of the system. And the idea is directly negatived, as we shall presently see, by the actual composition of the lymph draAvn from these vessels; the solid matter of which consists, in great part at least, of substances of a nutritive cha- racter. It is true that other substances are occasionally found in the lymphatics; thus, when the gall-bladder and bile-ducts are over-dis- tended with bile, in consequence of some obstruction to its exit, the lymphatics of the liver are found to contain a biliary fluid. In like manner, the lymphatics in the neighborhood of a large abscess have been found to contain pus. When the limb of an animal, round the upper part of Avhich a bandage is tied, is kept for some hours in tepid milk, the lymphatics of the skin are found distended with that fluid. And when saline solutions are applied to the skin, they are usually detected more readily in the lymphatics, than in the veins. But these facts only prove, that the lymphatics very readily imbibe soluble sub- stances with which they are in proximity; and this imbibition seems to take place on the same physical principles, as the imbibition of soluble substances by the veins of the intestinal canal. 503. The more ready absorption of such substances by the lympha- tics, than by the veins, of the cutaneous surfaces,—contrary to Avhat obtains in the alimentary canal,—is easily accounted for, by the very abundant distribution of the lymphatics in the skin, and the ready access which fluids can obtain to their walls. In other tissues it is different: thus it appears that saline matters injected into the lungs are detected much sooner in the serum of the blood, than they are in the lymph ; and make their appearance earlier in the left cavities of the heart, to which they Avould be conveyed by the pulmonary vein, than in the right, which they would reach through the thoracic duct and descending cava. This is obviously due to the minute distribution of the blood-vessels upon the walls of the air-cells; which makes them far more ready channels for the imbibition of fluid, than the lym- phatics could be.—In regard to the occasional absorption of pus from the cavity of an abscess or of an open ulcer, by the lymphatics, it is to be remarked that the absorbent vessels must themselves probably be laid open by ulceration ; since in no other way can we understand the entrance of the globules, so large as those of pus, into their interior. 504. In regard to the cause of the movement of the. chyle and lymph along the absorbent vessels, from their commencement to their termina- tion in the central receptacle, no very definite account can be given. The middle coat of these vessels has a fibrous texture; and the fibres bear some resemblance to that of the non-striated muscle. In the thoracic duct, this fibrous structure is more evident; and distinct con- tractions have been excited in it, by irritating the sympathetic trunks from which it receives its nenres, and the roots of the spinal nerves with which those trunks are connected. Hence it seems probable, that there is a sort of peristaltic contraction of the walls of the absorbents, analogous to that which takes place in the intestinal tube, serving to driA'e their contents slowly onwards; their reflux being prevented by 286 ABSORPTION AND SANGUIFICATION. the valves, with which they are copiously furnished. Moreover, it is probable that the general movements of the body may concur with the contractile power of the absorbent vessels themselves, to urge their con- tents onwards; for almost every change in position must occasion increased pressure on some portion of them, which will propel the fluid contents in the sole direction permitted by the valves, and thus give them an additional impulse towards the trunks, in which they are col- lected for delivery into the blood-vessels. 3. Of the Spleen, and other Glandular Appendages to the Lymphatic System. 505. The structure and functions of the Spleen, and of certain other organs allied to it in character, have been among the most obscure subjects in Anatomy and Physiology; and they are far from having been yet fully elucidated. There seems sufficient evidence, however, for regarding them in the light of appendages to the Absorbent system, and as concerned, like it, in the process of Sanguification, or the preparation of Blood. Hence this appears to be the most appro- priate place for such a brief notice of them as the present state of our knowledge admits. 506. The Spleen is certainly to be regarded as an organ of compound structure, having at least two sets of functions to fulfil. It is essen- tially composed of a fibrous membrane, which constitutes its exterior envelope, and which sends prolongations in all directions across its in- terior, so as to divide it into a number of minute cavities of irregular form, freely communicating with each other. In many animals, this fibrous envelope, and the prolongations or trabecular which it sends through the substance of the organ, are distinctly muscular ; containing a large proportion of the peculiar fusiform contractile cells formerly described (§ 337). These, however, do not present themselves in the Human spleen; and its trabeculae do not appear to have any contractile property. The areolae formed by the trabecular tissue, commonly known as the splenic follicles, are differently occupied in different ani- mals. In the Ruminants they are lined by a continuation of the splenic vein, which dilates into a cavernous structure, capable of re- ceiving a very large quantity of blood. In Man, however, they have no communication with the splenic vein, and are chiefly occupied by the Malpighian corpuscles and the parenchymatous tissue, which, in the Ruminants, are limited to the partitions between the venous cells. The Malpighian corpuscles of the spleen are whitish spherical bodies, which are always connected with the smaller arteries, like currants with their stalks; being sometimes in immediate contact with them, but more commonly being connected by a peduncle. Their size, when fully formed, varies from l-3d to l-6th of a line. Each of them contains, as its constant and essential elements, nucleated cells from l-4000th to l-2500th of an inch in diameter, pale and faintly granular, together with free nuclei, as well as larger cells of l-2000th of an inch in diameter, Avhich sometimes contain AA'hat appear to be red blood-cor- t puscles. These are enclosed in a capsule, which has no orifice, and STRUCTURE AND FUNCTIONS OF THE SPLEEN. 287 Avhich appears to be comparable to the elementary vesicles of other glands, before they have acquired an outlet by the rupture of their Avails (§ 718). Whatever may be the function of these bodies, it is pro- bable that it must be completed by the rupture of the capsules and the discharge of their contents; and that new corpuscles are continually in course of development.—The true splenic parenchyma consists in great part of cells which correspond in appearance with those of the Malpi- ghian corpuscles; but in addition to these, there are cells which bear a strong likeness to the colorless ' granule-cells' of the blood (§ 217), and others which resemble young red-corpuscles. These elements, also, present indications of being in a state of continual development and degeneration; and form small irregular groups of various sizes, which are clustered especially on the sheaths of the vessels, the trabecular partitions, and the exterior of the Malpighian capsules.—A considerable part of the contents of the splenic areolae has been found by Prof. Kolliker to consist of blood-corpuscles in various stages of degenerative metamorphosis. These are aggregated in small masses, each of which acquires an investing membrane, that may contain from one to twenty corpuscles; and the blood-cells gradually diminish in size, and undergo a transition into pigment-granules ; so that the containing cells are converted into pigmentary granule-cells, which at last, by a gradual loss of color of the granules, become quite pale. Such cells are found in the blood drawn from the splenic vein, the vena portae, and the in- ferior cava; and occasionally in that of other veins. These curious collections of degenerating blood-corpuscles, however, are not peculiar to the spleen, for they have been found by Prof. Kolliker in other parts, such as the substance of the muscles. 507. In regard to the functions of the Spleen, great uncertainty still exists. It appears from the foregoing account of its structure, that it may be regarded as an organ of duplex character, and probably of double function. In the Ruminants, the cavernous dilatations of the veins enable it to hold, upon occasion, a large quantity of blood; and their walls are so elastic, that their cavities may be greatly distended Avith a very moderate force; the Spleen of the sheep, which weighs about 4 oz., being easily made to contain about 30 oz. of water. This peculiar distensibility evidently points to the Spleen, as a kind of reservoir, connected Avith the Portal circulation, for the purpose of relieving the portal vessels from undue pressure or distension, under a great variety of circumstances. The portal system is well known to be destitute of valves, so that the splenic vein communicates freely with the Avhole of it; and thus, if any obstruction exists to the flow of blood through the liver, or any peculiar pressure elsewhere prevents the mesenteric veins from dilating to their full extent, the general circula- tion is not disturbed, the Spleen affording a kind of safety-valve. That any cause of congestion of the Portal system peculiarly affects the Spleen, has been proved by experiment; for, after the portal vein has been tied, the spleen of an animal that previously Aveighed only 2 oz., has been found to increase to 20 oz.—Again, the Spleen appears to serve as a reservoir, into Avhich superfluous blood may be carried, during the digestive process. When the alimentary canal is distended with 288 ABSORPTION AND SANGUIFICATION. food, and a great afflux of arterial blood takes place to the mucous membrane, the veins of the portal system will be liable to increased pressure from Avithout, whilst their contents will be augmented by the quantity of fluid neAvly absorbed from the alimentary canal. In this, as in the preceding cases, the distensibility of the spleen makes it a kind of safety-valve, by which undue distension of the portal system is relieved. It has been ascertained that its maximum volume is attained about five hours after a meal, Avhen the process of chymification is at an end, and that of absorption is taking place with activity; and the in- crease is proportional rather to the amount of the fluids ingested, than to that of the solids.—Although the Human Spleen has no true cavern- ous structure, yet its veins are obviously very distensible, so that a great accumulation of blood may take place in it. Thus, in Asphyxia, Avhen the circulation of blood is checked in the Lungs, and Avhen the stagna- tion extends itself backAvards to the right side of the heart, the vena cava, and thence to the portal system, the Spleen is often found after death to be enormously distended with blood. And in the cold stage of intermittent fever, in which a great quantity of blood is driven from the surface towards the internal organs, the Spleen receives a large portion of it, so that its increased size becomes quite perceptible; and in cases of confirmed Ague, the Spleen becomes permanently enlarged, forming what is popularly known as the "ague-cake." 508. But besides this safety-valve function, there can be little ques- tion that the Spleen performs some other, which is related more closely to the nutritive operations, and which in some degree correspond with that performed by the Absorbent glandulae. The multitude of glandu- lar cells in immediate relation with blood-vessels, and the appearances of rapid development and degeneration which these present, taken in connexion with the fact that there is no other outlet for the products of their action than that which is afforded by the veins, clearly indicate that whatever this product may be, it is destined to form part of the blood; and that the Spleen is, therefore, an organ of sanguification. This view is confirmed by the remarkable fact, ascertained by recent experiments, that after the spleen has been extirpated, the lymphatic glands of the neighborhood increase in size, and cluster together as they enlarge, so as to form an organ which at least equals the original spleen in volume. This circumstance explains the reason for the almost invariable negative result of the extirpation of the spleen; for although the operation has been frequently practised, with the view of deter- mining the functions of the organ by the symptoms presented by the animals after its removal, no decided change in the ordinary course of their vital phenomena has ever been observed, and the health, if at all disturbed for a time, is afterwards completely regained. Now if the principal function of the Spleen be the same with that of the lymphatic glands in general, it is easy to understand hoAV its loss may be at once compensated by an increased action on their part, and how it may be permanently replaced by an increased development of certain of those bodies.—It is worthy of remark, that a Spleen is found in all Verte- brated animals which have a distinct Absorbent system; but that no orgaji exactly corresponding with it exists in the Invertebrata, which STRUCTURE AND FUNCTIONS OF THE SPLEEN. 289 are destitute of that system,—although the distensible cellated cavities, apparently destined to perform its safety-valve function, exist in some of the higher among them. This is an additional reason for regarding its parenchymatous portion as essentially a part of the assimilating ap- paratus of the Absorbent system. 509. It would further seem as if the Spleen were specially concerned in the development of the red corpusdes of the blood; since its paren- chyma contains cells which resemble these in various stages of develop- ment. But this organ also appears to promote the disintegration of those red corpuscles Avhich have become effete; and this so powerfully, that the blood of the splenic vein contains a far less proportion of red corpuscles, and a far greater proportion of albumen, than that of any other vessels in the body. It is supposed by Prof. Kolliker, that the dissolved blood-corpuscles are subservient to the formation of bile; the coloring matter of which is nearly allied to that of the blood. 510. The Supra-Renal Capsules seem to correspond with the Spleen in their essential structure, whilst in the arrangement of their compo- nent parts, they bear more resemblance to the Kidney. Their exterior or cortical portion, in Man and the Mammalia generally, is formed of straight arteries, which divide into a minute capillary network; and from this arise venous branches, which form a minute plexus through the internal or medullary substance, pouring its contents into a large central cavity, Avhich is the dilated commencement of the supra-renal vein. In the lower Vertebrata, hoAvever, there is no distinction of cor- tical and medullary substance; the distribution of vessels being nearly the same throughout. As in the Spleen, we find in the interspaces of the vascular plexus a parenchymatous structure; partly composed of free cells and nuclei in various stages of development; and partly con- sisting of closed granular vesicles, within which similar cells and nuclei are included. There is ground for asserting that these vesicles are themselves at first in the condition of simple cells, and that the cells Avhich they include are a secondary product. In Man and the Mam- malia, these vessels are confined to the cortical substance, and have the form of elongated tubes, lying betAveen its parallel blood-vessels. Thus, then, the supra-renal capsules possess the essential structure of a gland, in every respect save the Avant of an excretory duct; and Avhatever may bo the product of their cell action, this must be received back into the blood again. The fluid contents of these bodies are rich in proteine- compounds and in fat; and it can be scarcely doubted that these mate- rials here undergo an elaboration, Avhich renders them better fitted for the nutrition of the system. It does not seem unlikely that these bodies-, like the Spleen, have a double function; and that, besides participating in the general actions of the Absorbent glandulae, they^may serveas-a divftrticnjnm forthe Renal circulation, when from any cause the se- curing funHTorTbf imeKidneys is retarded or checked, and the move- ment of blood through them is stagnated.—The Supra-Renal capsules of Man attain a very large size early in foetal life, surpassing the true Kidneys in dimension up to the tenth or twelfth week; but they affcer- Avards diminish relatively to the latter, and are evidently subordinate organs during the Avhole remainder of life. In most of the lower ani- 290 ABSORPTION AND SANGUIFICATION. mals, hoAvever, these bodies retain through life the same relative deve- lopment. 511. The Thymus Gland is another body, which seems referable to the same group; having all the essential characters of a true gland (§ 714), save an excretory duct; and its function being evidently con- nected, during the early period of life at least, Avith the elaboration of nutritive matter, which is to be .reintroduced into the circulating cur- rent. Its elementary structure may be best understood from the simple form it presents Avhen it is first capable of being distinguished in the embryo. It then consists of a single tube, closed at both ends, and filled with granular matter; and its subsequent development con- sists in the lateral growth of branching off-shoots from this central tubular axis. In its mature state, therefore, it consists of an assem- blage of glandular follicles, Avhich are surrounded by a plexus of blood- vessels ; and these follicles all communicate with the central reservoir, from which, hoAvever, there is no outlet. The cavities of the follicles contain a fluid, in which a number of corpuscles are found, giving it a granular appearance. These corpuscles are for the most part in the condition of nuclei; but fully developed cells are found among them, at the period when the function of this body seems most active. The chemical nature of the contents at this period, closely resembles that of the ordinary proteine-compounds.—It has been commonly stated, that the Thymus attains its greatest development, in relation to the rest of the body, during the latter part of foetal life; and it has been considered as an organ peculiarly connected with the embryonic condition. But this is a mistake; for the greatest activity in the growth of this organ manifests itself, in the Human infant, soon after birth ; and it is then, too, that its functional energy seems the greatest. This rapid state of growth, however, soon subsides into one of less activity, which merely serves to keep up its proportion to the rest of the body; and its increase usually ceases altogether at the age of about two years. From that time, during a variable number of years, it remains stationary in point of size; but, if the individual be adequately nourished, it gradually assumes the character of a mass of fat, by the development of the cor- puscles of its interior into fat-cells, which secrete adipose matter from the blood. This change in its function is most remarkable in hiber- nating Mammals; in which the development of the organ continues, even in an increasing ratio, until the animal reaches adult age, when it includes a large quantity of fatty matter. The same is the case, gene- rally speaking, among Reptiles. It is an important fact in the history of this organ, that it is not to be detected in Fishes; and does not appear to exist, either in the tadpole state of the Batrachian reptiles, or in the Perennibranchiate group; so that we may regard it as essen- tially connected with pulmonic respiration.* j 512. Various facts lead to the conclusion, that the function of the | Thymus, at the period of its highest development, is that of elaborating ? and storing-up nutritive materials, to supply the demand Avhich is pecu- $ liarly active during the early period of extra-uterine life. The elabo- * See Mr. Simon's admirable Prize Essay on the Thymus Gland. THYMUS GLAND. 291 rating action probably corresponds with that, which is exerted by the glands of the Absorbent system ; and the product, as in the preceding cases, seems to be taken back into the circulation. The provision of a store of nutritive matter seems a most valuable one, under the circum- stances in which it is met Avith; the waste being more rapid and variable than in adults, and the supply not constant. Thus it has been noticed that, in over-driven lambs, the thymus soon shrinks remarkably ; but that it becomes as quickly distended again during rest and plentiful nourishment. As the demand becomes less energetic, and as the sup- plies furnished by other organs become more adequate to meet it, the Thymus diminishes in size, and no longer performs the same function. It then obviously serves to provide a store of material, not for the nutri- tion of the body, but for the respiratory process, when this has to be carried on for long periods (as in hibernating Mammals, and in Rep- tiles), Avithout a fresh supply of food.—It is possible that the Thymus gland may further stand in the same relation to the Lungs, as the Spleen to the Liver, and the Supra-Renal capsules to the Kidneys ; that is, as a diverticulum for the blood transmitted through the bronchial arteries (which are the nutritive vessels of the Lungs), before the Lungs acquire their full development in comparison with other organs, or when any cause subsequently obstructs the circulation through their capillaries. 513. The Thyroid Gland bears a general analogy to the Thymus; but its vesicles are distinct from each other, and do not communicate with any common reservoir. They are surrounded, like the vesicles of the true glands, with a minute capillary plexus ; and in the fluid they contain, numerous corpuscles are found suspended, which appear to be cell-nuclei, in a state of more or less advanced development. This body is supplied with arteries of considerable size; and with peculiarly large lymphatics. Though proportionably larger in the foetus than in the adult, it remains of considerable size during the whole of life.—It ap- pears, from the recent inquiries of Mr. Simon,* that a Thyroid gland, or some organ representing it in place and office, exists in all Verte- brated animals. It presents its simplest form in the class of Fishes ; in some of Avhich it appears to consist merely of a plexus of capillary vessels, connected with the origin of the cerebral vessels, and capable, by its distensibility, of relieving the latter, in case of any obstruction to the proper movement of blood through them. In the higher forms of this organ, the glandular structure,—consisting of the closed vesicles over Avhich the capillary plexus is distributed, and of their cellular con^ tents,—is superadded : and the organ then appears, likejjii£ Spifiga, to be destined for two different uses ; namely, to^sgjxe as a diverticulum to>-tke_Cerebral circulation; and to aid in theidaboratiofPof nutritive matter/which Is "probably taken up by the Absorbent system, to be again poured by it into the general current of the circulation. *—— 514. Thus the Spleen, the Supra-Renal Capsules, the Thymus Gland, and the Thyroid Gland, all seem to share in the preparation of the nu- tritive materials of the blood; in fact, Ave may regard them all as toge- ther constituting an elaborating apparatus, which is precisely analogous Philosophical Transactions, 1844. 292 ABSORPTION AND SANGUIFICATION. to that of the ordinary glands, but of Avhich the elementary parts are scattered through the body instead of being collected into one compact structure, and of Avhich the product is received back into the blood, in- stead of being discharged through an efferent duct, upon the surface, or into an open cavity, of the body. And it is a remarkable confirma- tion of this vieAv, that Prof. Goodsir should have ascertained that the last three of these organs are in continuity Avith each other, during the early part of foetal life; and that they are in reality portions of the blastoderma or germinal membrane (chap, xi.), which retain their ori- ginal simplicity of structure, whilst other parts undergo changes of form and texture, and which continue to perform their original function, save that the materials of their elaborating action are now supplied by the blood instead of by the yolk. The probable uses of these bodies, as diverticula to the circulation through other organs, render them liable to occasional distension with blood; and it seems determined that this blood shall not lie useless, but shall be subservient to the action in ques- tion ; the gland-cells that line the cavities of the organ withdraAving cer- tain constituents of the blood, to restore them to the circulating current, in a state of more complete preparation for the operations of Nutrition. Their function is very probably vicarious ; that is, the determination of blood is greatest (through the state of the other organs), at one time to one of these bodies, and at another time to another. Hence the effects of the loss of any one of them are not serious; as the others are ena- bled in a great degree to discharge its duty. 4. Composition and Properties of the Chyle and Lymph. 515. The chief chemical difference between the Chyle and the Lymph, consists in a much smaller proportion of solid matter in the latter, and in the almost entire absence of fat, which is an important constituent of the former. This is well shown in the folloAving comparative analyses, performed by Dr. G. 0. Rees, of the fluids obtained from the lacteal and lymphatic vessels of a donkey, previously to their entrance into the thoracic duct; the animal having had a full meal seven hours before its death. Water, - - - Albuminous matter (coagulable by heat), Fibrinous matter (spontaneously coagulable), Animal extractive matter, soluble in water and alcohol, Animal extractive matter, soluble in water only, Fatty matter, ------ Salts;—Alkaline chloride, sulphate and carbonate, with traces of alkaline phosphate, oxide of iron, 100000 100000 The Lymph obtained from the neck of a horse has been recently analyzed by Nasse, with nearly the same result. He found it to contain 95 per cent, of Avater; and the 5 per cent, of solid matter was chiefly composed of albumen and fibrine, with Avatery extractive,—scarcely a trace of fat being discoverable. The proportions of saline matter were found to be remarkably coincident Avith those Avhich exist in the serum Chyle. Lymph. 90-237 95-536 3-516 1-200 0-370 0-120 0-332 0-240 1-233 1-319 3-601 a trace 0-711 0-585 CHARACTERS OF THE CHYLE. 293 of the blood ; as might be expected, from the fact, that the fluid portion of the lymph must have its origin in that which has transuded through the blood-vessels: the absolute quantity, however, is rather less. A similar analysis of the Chyle of a cat by Nasse, has given results very closely correspondent Avith that of Dr. Rees; for the proportion of water Avas 90-5 per cent.; and of the 9-5 parts of solid matter, the albumen, fibrine, and extractive amounted to more than 5, and the fat to more than 3 parts. Dr. Rees has also analyzed the fluid of the Thoracic duct of Man, Avhich consists of chyle with an admixture of lymph ; and he found this to contain about 90-5 per cent, of water, 7 parts of albumen and fibrine, 1 part of aqueous alcoholic extractive, and not quite one part of fatty matter, with about J per cent, of salines. The composition of this fluid more resembles that of the lymph than that of the chyle; the proportion of the fatty to the albuminous matter being small. This was probably due to the circumstance, that the sub- ject from which it Avas obtained (an executed criminal) had eaten but little for some hours before his death. 516. The characters of the Chyle are not the same in every part of the Lacteal system ; for the fluid undergoes a very important series of changes in its characters, in its transit from the Avails of the intestines to the receptaculum chyli. The fluid draAvn from the lacteals that tra- verse the intestinal walls, has no power of spontaneous coagulation; whence we may infer that it contains little or no Fibrine. It contains Albumen in a state of complete solution, as we may ascertain by the influence of heat or acid in producing coagulation. And it includes a quantity of fatty matter, which is not dissolved, but suspended in the form of globules of variable size. The quantity of this evidently varies with the character of the food ; it is more abundant, for instance, in the chyle of Man and the Carnivora, than in that of the Herbivora. It is generally supposed that the milky color of the chyle is owing to the oil-globules ; but Mr. Gulliver has pointed out that it is really due to an immense multitude of far more minute particles, AA'hich he has described under the name of the molecular base of the chyle. These molecules are most abundant in rich, milky, opaque chyle; whilst in poorer chyle, Avhich is semi-transparent, the particles float separately, and often exhibit the vivid motions common to the most minute mole- cules of various substances. Such is their minuteness, that, even with the best instruments, it is impossible to determine either their form or their dimensions with exactness; they seem, hoAvever, to be generally spherical; and their diameter may be estimated at between l-36,000th and l-24,000th of an inch. Their chemical nature is as yet uncertain; they are remarkable for their unchangeableness, when submitted to the action of numerous reagents, Avhich quickly affect the proper Chyle- corpuscles ; whilst their ready solubility in Ether would seem to indi- cate that they are of an oily or fatty nature. ^ 517. The milky aspect which the serum of blood sometimes exhibits, is due to an admixture of this molecular base. It may be particularly noticed, Avhen blood is drawn a feAV hours after a full meal, that has been preceded by a long fast. By recent experiments it has been found, that the serum begins to shoAV this turbidity about half an hour after the 294 ABSORPTION AND SANGUIFICATION. meal has been taken; and that the turbidity increases for some hours subsequently, after Avhich it disappears. The period at which the dis- coloration is greatest, and the length of time during Avhich it continues, vary according to the digestibility of the food. When the serum is allowed to remain at rest, the opaque matter rises to the surface, pre- senting very much the appearance of cream ; and when separately examined, it has been found to contain a proteine-compound, mingled with oily matter,—the relative amount of the tAvo appearing to depend in part upon the characters of the food ingested. Hence it would seem probable that the molecular base of the chyle is partly derived from albuminous matter of the food; more especially as it is known that oily particles, when introduced into an albuminous fluid, become surrounded with a pseudo-membranous pellicle. The gradual disappearance of the turbidity of the serum, indicates that the substance which occasioned it no longer exists as such in the circulating current; being either drawn off by the nutritive or secretory operations, or being converted by the assimilating process into the ordinary constituents of the blood. 518. During the passage of the Chyle along the lacteals, towards the Mesenteric glands, it undergoes two important changes; the presence of Fibrine begins to manifest itself by the spontaneous coagulability of the fluid ; and the oil-globules diminish in proportional amount. The fibrine appears to be formed at the expense of the albumen ; as this latter ingredient undergoes a slight diminution. It is in the chyle drawn from the neighborhood of the mesenteric glands, that we first meet with the peculiar floating cells, or chyle-corpuscles, formerly adverted to (§ 214), in any number. The average diameter of these is about l-4600th of an inch ; but they vary from about l-7000th to l-2600th,—that is, from a diameter about half that of the human blood- corpuscles, to a size about one-third larger. This variation probably depends in great part upon the period of their growth. They are usually minutely granulated on the surface, seldom exhibiting any regular nuclei, even when treated Avith acetic acid ; but three or four central parti- cles may sometimes be distinguished in the larger ones. These corpus- cles are particularly abundant in the chyle obtained by puncturing the mesenteric glands themselves; and there can be little doubt, that they are identical with the altered epithelium-cells, which line the lacteal tubes in their course through those bodies (§ 496). 519. The glandular character of these cells, and their continued pre- sence in the circulating fluid, seem to indicate that they have an important concern in the process of assimilation,—that is, in the con- version of the crude elements derived from the food, into an organizable matter adapted to the nutrition of the body; in other words, in the conversion of Albumen into Fibrine; which change would seem to take place to a considerable extent in the Mesenteric glands. For it is only in the Chyle which is drawn from the lacteals intervening between the mesenteric glands and the receptaculum chyli, that the spontaneous coagulability of the fluid is so complete, as to produce a perfect separa- tion into clot and serum. The former is a consistent mass, which, when examined with the microscope, is found to include many of the chyle-corpuscles, each of them being surrounded Avith a delicate film of PROGRESSIVE ELABORATION OF BLOOD. 295 oil; the latter bears a close resemblance to the serum of the blood, but has some of the chyle-corpuscles suspended in it. Considerable diffe- rences present themselves, however, both in the perfection of the coagu- lation, and in its duration. Sometimes the chyle sets into a jelly-like mass ; which, without any separation into coagulum and serum, liquefies again at the end of half an hour, and remains in this state. The coagu- lation is usually most complete in the fluid drawn from the receptaculum chyli and thoracic duct; and here the resemblance between the floating cells, and the white or colorless corpuscles of the blood, becomes very striking. 520. The Lymph, or fluid of the Lymphatics, differs from the Chyle, as already remarked, in its comparative transparency: its want of the opacity or opalescence, which is characteristic of the latter, being due to the absence, not merely of oil-globules, but also of the "molecular base." It contains floating cells, which bear a close resemblance to those of the Chyle on the one hand, and to the colorless corpuscles of the Blood on the other; and these, as in the preceding case, are most numerous in the fluid which is drawn from the lymphatics that have passed through the glands, and in that obtained from the glands them- selves. Lymph coagulates like chyle ; a colorless clot being formed, which encloses the greater part of the corpuscles. The lacteals may be regarded as the Lymphatics of the intestinal Avails and mesentery ; performing the function of interstitial absorption, as well as effecting the introduction of alimentary substances from without. During the inter- vals of digestion, they contain a fluid, Avhich is in all respects conform- able to the lymph of the lymphatic trunks. 521. Thus by the admixture of the aliment newly introduced from without, with the matter Avhich has been taken up in the various parts of the system, and by the preparation which these undergo in their course toAvards the thoracic duct, a fluid is prepared, which bears a strong resemblance to blood in every particular, save the presence of red corpuscles. Even these may sometimes be found in the contents of the thoracic duct, in sufficient amount to communicate to them a perceptible red tinge; but it is doubtful whether they have not found their way thither accidentally,—some of the lymphatic or lacteal trunks, which have been divided in the dissection necessary to expose the duct, having taken up blood by their open mouths, and rapidly transmitted it into the general receptacle. The fluid of the thoracic duct may be compared to the blood of Invertebrated animals; from which the red corpuscles are almost or altogether absent; but which contains white or colorless corpuscles, and which possesses but a slight coagulating power, in consequence of its small proportion of fibrine. And we hence see, why these animals should require no special absorbent sys- tem ; since the blood-vessels convey a fluid, which is itself so analogous to the chyle and lymph to be absorbed, that the latter may be at once introduced into it, Avithout injuring its qualities. 296 ABSORPTION AND SANGUIFICATION. 5. Absorption from the External and Pulmonary Surface. 522. Although the Mucous Membrane of the Alimentary Canal is the special channel for the introduction of nutritive or other substances into the system, it is by no means the only one. The Skin coA'ering the body, and the Mucous Membrane prolonged into the Lungs, are also capable of absorbing liquids and vapors, and of introducing them into the Circulation ; although they serve this purpose less in Man and the higher animals, than in some of the loAver. Their utility in this respect is best shown, when, from peculiar circumstances, the function of the digestive cavity cannot be properly performed; and when, therefore, the system has been more than usually drained of its fluids, and stands in need of a fresh supply. Thus shipwrecked sailors, and others, who are suffering from thirst, owing to the Avant of fresh water, find it greatly alleviated, or altogether relieved, by dipping their clothes into the sea, and putting them on Avhilst still wet, or by frequently immersing their own bodies. In a case of dysphagia, in which neither solid nor fluid nutriment could be introduced into the stomach, the patient was kept alive for a considerable time, and his sufferings greatly alleviated, by the administration of nutritive clysters, and by the immersion of his body in a bath of tepid milk and Avater, night and morning. Under this system, the weight of the body, which had previously been rapidly diminishing, remained stationary, although the amount of the excretions was increased : and the use of the bath had a special influence in assuaging the thirst, which was previously distressing. It appeared that the water of the urinary excretion, amounting to from 24 oz. to 36 oz. per day, must have been entirely supplied from this latter source. Again, a man who had lost nearly 31bs. by perspiration, during an hour and a quarter's labor in a very hot atmosphere, regained 8 oz. by immersion in a warm bath at 95° for half an hour. In these cases it appears probable, from the experiments already noticed (§ 502), that the Lymphatics, rather than the blood- vessels, are the chief agents in the absorbing process; not, however, from any powers peculiar to them, but merely on account of the thin- ness of their walls, and their very copious distribution in the skin. 523. Absorption may also take place from an atmosphere saturated with watery vapor. Of this we have a very curious proof in the Frog; whose urinary bladder (which serves as a sort of reservoir for water) has been observed to be refilled, after having been emptied, by placing the animal in an atmosphere loaded with watery vapor. Numerous instances are on record which prove that such absorption may take place in Man, to a very considerable extent; though the proportion introduced through the Skin, and through the Lungs, can- not be exactly ascertained. The ready introduction of volatile matter into the system, through the latter channel, is a matter of familiar experience; thus if we breathe an atmosphere through which the vapor of turpentine is diffused, it soon produces the characteristic odor of violets in the urinary secretion. And it is probably in this manner, that a large number of those putrescent miasmata and other zymotic poisons are introduced, which are such fertile causes of disease. COMPOSITION OF THE BLOOD. 297 6. Of the Composition and Properties of the Blood. 524. Having traced the steps by which the Blood is elaborated, and prepared for circulation through the body, and having (in the former part of the volume) inquired into the characters of its chief consti- tuents, Ave have now to consider the fluid as a whole, to study the usual proportions of these constituents, and the properties which they impart to it. 525. The Blood, whilst circulating in the living vessels, may be seen to consist^of a, tlranRpairpnt».,nfiady.Jin1orl£a&JLiud, termed Liquor Sanguinis ; in which the Corpuscles, to which the blood owes its red hue, as well as the white or colorless corpuscles, are freely suspended and carried along by the current. On the other hand, when the blood has been drawn from the body, and is allowed to remain at rest, a spontaneous coagulation takes place, separating it into Clot and Serum. The Clot is composed of a network of Fibrine, in the meshes of which the Corpuscles, both red and colorless, are involved ; and the Serum is the same with the liquor sanguinis deprived of its Fibrine. When the Serum is heated, it coagulates, showing the presence of Albumen. And if it be exposed to a high temperature, sufficient to decompose the animal matter, a considerable amount of earthy and alkaline Salts remains. Thus we have four principal components in the Blood:— namely, Fibrine, Albumen, Corpuscles, and Saline matter. In the circulating Blood they are thus combined:— Fibrine A Albumen v In solution, forming Liquor Sanguinis. Salts J lied Corpuscles,—Suspended in Liquor Sanguinis. But in coagulated blood they are thus combined:— D , n , > Crassamentum or Clot. Red Corpuscles $ Albumen ) t> ... , ,. „ ^ a ,. > Remaining in solution, forming Serum. A certain amount of Serum, hoAvever, is involved in the Crassamentum ; and can only be separated by cutting the clot into thin slices, and carefully washing it. 526. The components of the Blood may be separated, and their amount estimated, in various Avays. Thus, if fresh-drawn blood be continually stirred with a stick, or be " Avhipped" with a bunch of twigs, the fibrine coagulates in the form of strings, which adhere to the Avood, and may thus be AvithdraAvn ; whilst the red corpuscles then remain suspended in the serum, gradually sinking to tbe bottom in virtue of their greater specific graATity. On the other hand, the Red Corpuscles may be separated, in those animals in Avhich they are large enough, by passing the blood through a filter ; haA'ing previously mingled with it some substance which retards, but does not prevent its coagula- tion* (§ 185). The liquor sanguinis is thus separated from the blood- * This experiment cannot be performed -with Human blood, because the corpuscles are small enough to pass through the pores of any filter that allows the liquor sanguinis to permeate it; but it answers very well with Frog's blood. 298 ABSORPTION AND SANGUIFICATION. discs; and the former coagulates, whilst the blood-discs are retained upon the filter. The experiment convincingly proves, that the act of coagulation is not due to the red corpuscles, as Avas at one time ima- gined.—The ordinary act of coagulation, by withdrawing the Fibrine and Corpuscles, makes it easy to estimate the proportion of Albumen and of Saline matter in the Blood, Avhen due alloAvance is made for the quantity of Serum retained in the Clot; and the relative propor- tions of these may be determined, by evaporating the fluid, so as to obtain the Avhole amount of solid matter it contains, and by then cal- cining the residuum, so as to ascertain how much of this is a mineral ash,—the remainder being chiefly Albumen.—The solid matter of the blood also contains various Fatty substances, which may be removed from it by ether. Some of these appear to correspond Avith the con- stituents of ordinary Fat (§ 261); whilst another contains phosphorus, and seems allied to the peculiar fatty acids of Neurine (§ 383); and another has some of the properties of Cholesterine, the fatty matter of the Bile (§ 724).—Besides these, there are certain substances known under the name of Extractive ; one group of which is soluble in water, and another in Alcohol. Of the precise nature of these, little is knoAvn. They have been aptly termed " ill-defined" animal principles; and it is probable that they may include various substances in a state of change or disintegration, which are being eliminated from the Blood by the processes of Excretion. 527. The general result of numerous recent analyses of the Blood may be thus stated. The whole amount of solid matter is rather greater in the Male than in the Female; being, on the average, about 210 parts in 1000 in the former, and 200 in the latter. This diffe- rence, hoAvever, chiefly depends on the larger proportion of red corpus- cles contained in the blood of the male. The proportion of Albumen seems more constant than that of the other constitutents of Blood; seldom varying beyond 5 or 6 parts, in either sex, above or below 70 in 1000. The quantity of Corpuscles appears liable to considerably greater variation ; the superiority on the side of the Male, however, being very strongly marked in the maximum and minimum, as well as in the average. We may regard its average in the Male as about 132in 1000 parts of blood; but it may fall to 110-5 parts, without the health being seriously affected; whilst, on the other hand, it may arise to 186 without any manifestation of disease. In the Female, its average may be about 120 parts in 1000 ; but it may fall to as little as 71'4, and may rise to 167, consistently with ordinary health. The range of variation is thus much greater in the Female than in the Male ; the minimum being considerably less, in the former, than half the maximum: whilst in the latter it is much more. This is probably due in part to the fact, that the loss by the Catamenial discharge may produce a great tempo- rary depression in the proportion of the Corpuscles. The aArerage proportion of Fibrine seems to be no more than 2*5 in the Male ; and though it may rise to as much as 3-5 or even 4, without disordering the system, it does not seem to fall below 2, in the state of ordinary health. The average in the Female is probably about 2-3; the proportion may rise to 3, or fall to 1*8; but the variation seems less considerable in the COMPOSITION OF THE BLOOD. 299 Female than in the Male.—Much is probably yet to be learned, regard- ing the influence of different kinds of food recently taken, on the pro- portion of these constituents of the blood ; and it does not seem un- likely, from what has been already stated (§ 517), that the quantity of fatty matter is especially liable to variation, in accordance Avith the amount contained in the food, and the time which has elapsed since the last meal. 528. The Saline constituents of the blood, obtained by drying and incinerating the whole mass, usually amount to betAveen 6 and 7 parts in 1000. More than half of their total quantity is composed of the Chlorides of Sodium and Potassium ; and the remainder is made up of the tribasic Phosphate of Soda, the Phosphates of Lime and Magnesia, Sulphate of Soda, and a little Phosphate and Oxide of Iron. Of these the chief part are dissolved in the Serum ; but the Earthy Phos- phates, which are insoluble by themselves, are probably combined Avith the proteine-compound (§ 175); and the Iron is contained, chiefly or entirely, in the red corpuscles. It is difficult to speak with certainty, from the examination of the ashes of the blood, as to the state of the saline constituents of the circulating fluid. Thus the Serum has an alkaline reaction ; and this has been supposed to be due to the pre- sence of alkaline Carbonates. Moreover, the presence of the Lactates of potash and soda seems probable; for it is certain that lactic acid is normally introduced into the blood, and is also eliminated from it; and the rapidity Avith which the lactates are removed as such, or are converted into carbonates, seems to afford a sufficient explanation of the difficulty in demonstrating the presence of this acid in the circula- ting fluid. Although the ashes of the entire mass of blood do not effer- vesce on the addition of an acid, effervescence takes place, Avhen acid is added to the ashes of the Serum; showing the existence in it, either of alkaline Carbonates, or of Lactates which have been reduced to the state of Carbonates by incineration. When the entire mass of blood is incinerated, hoAvever, enough phosphoric acid is produced from the phos- phorized fats, to neutralize the alkaline carbonates, and thus to prevent their presence from being recognised. The alkaline reaction of the blood, however, is certainly dependent in part upon the presence of the tribasic phosphate of soda, which appears to confer upon the serum a special poAver of absorbing carbonic acid. 529. The following appear, from the considerations stated in the pre- ceding part of the Volume, to be the chief uses of the principal consti- tuents of the Blood, in the general economy. The Fibrine is the material, Avhich is most completely prepared for organization, and Avhich supplies what is requisite for the nutrition of the larger proportion of the solid tissues of the body. It is, therefore, being continually AvithdraAvn from the blood by the nutritive operations ; and the demand appears to be supplied, in part by the influx of Fibrine that has been prepared in the Absorbent system, and in part by the continued transformation of Albu- men, which takes place during the circulation of the Blood. If a proper amount of Fibrine be not present in the Blood, its physical properties are so far altered, by the diminution of its viscidity, that it Avill not circulate through the capillaries as readily as before ; a certain degree 300 ABSORPTION AND SANGUIFICATION. of viscidity having been experimentally found to be favorable to the movement of fluid through glass or metallic tubes of small bore.—The Albumen of the blood is the raw material, at the expense of A\Thich not only the Fibrine, but many other substances are generated during the nutritive process. All the Albuminous compounds of the Secretions, the Horny matter of the Epidermic tissues, the Gelatine of the simple fibrous tissues, the solid materials of the Red Corpuscles, and other sub- stances, may be regarded as almost certainly produced by the transfor- mation of the Albumen of the Blood : and a continual supply of this from the food is therefore requisite, to preserve the due proportion in the circulating fluid.—The Red Corpuscles, Avhich (it will be remembered) are almost exclusively confined to Vertebrated animals, appear to be more connected Avith the function of Respiration, than Avith that of Nutrition ; and the stimulating action of Arterial blood, especially upon the Muscular and Nervous tissues, appear chiefly to depend upon their presence. It has been observed in particular, that their presence is more effectual in stimulating the heart's action, than is that of either of the other constituents of the blood. In addition to what has been already stated (§ 219), in reference to their continual disintegration and reneAval, it may be mentioned, that when the blood of one animal Avas injected by Magendie into the veins of another having discs of very different size and form, the original Red Corpuscles soon disappeared, and were replaced by those characteristic of the species, in whose veins the fluid was circulating. 530. The use of the Saline matter is evidently in part to prevent de- composition in the circulating Blood; but also to supply the mineral materials, requisite for the generation of the tissues, and entering into the composition of the secretions. It is by the saline and albuminous matters in conjunction, that the specific gravity of the Liquor Sanguinis is kept up to the point, at which it is equivalent to that of the contents of the Red Corpuscles; and it is only in this condition, that the forma- tion of the latter can duly take place. The Fatty matters of the blood are evidently derived from the food, either directly, or by a transforma- tion of its farinaceous ingredients (§ 430); and they are chiefly appro- priated to the maintenance of the combustive process. But there is reason to believe, that Oleaginous matter performs a most important part in the incipient stages of Animal nutrition ; and that its presence is not less essential to the formation of "cells, than is that of the albumi- nous matter which forms their chief component, all nuclei being observed to include fatty particles. That which may be superfluous is either de- posited in the cells of Adipose Tissue, or it is eliminated by the Liver, the Sebaceous follicles of the Skin, and, in the female Avhen nursing, by theJVlammary glands. 531. The proportion of these components of the Blood is liable to , undergo changes in disease, which extend far beyond the widest limits \ which have been mentioned as consistent with health. Thus, the ; quantity of Fibrine exhibits a remarkable increase in Inflammation; the \ amount then found in the blood being from 5 or 6 parts in 1000 to 9, • 10, or even 10£, according to the extent and intensity of the disease. ;On the other hand, it presents a remarkable diminution in Typhoid COMPOSITION OF BLOOD IN DISEASE. 301 fevers; the quantity being sometimes as little as 0-9. If any decided Inflammation should develope itself, however, in the course of the Fever, the proportion of Fibrine rises accordingly. A deficiency of Fibrine in the blood predisposes to Hemorrhages, Congestions, &c, either into the substance of the tissues, or on the surface of membranes ; and these conditions are well knoAvn to be of frequent occurrence as complications of febrile disorders. An excess of Fibrine is not much affected by copious bleeding, even if this be frequently repeated; but there is reason to think, that the administration of Mercury has a tendency to restrain its production. 532. It is difficult to say what amount of Red Corpuscles should be regarded as excessive; since, as we have seen, they may augment to a great degree, without disturbing the health. When they are present in an amount much above the average, they seem concerned in pro- ducing the condition termed Plethora; which marks a "high condi- tion" of the system, and which borders upon various diseases, espe- cially those of Congestion, and Hemorrhages. To these a peculiar liability then exists; because, although the proportion of Fibrine in the blood is not absolutely low, it is low in reference to that of the Red Corpuscles. Plethoric persons do not seem more liable to Inflamma- tion, than are those of Aveakly constitution. The quantity of the Red Corpuscles is rapidly diminished by frequent bleeding ; and hence it is loAvered .by repeated Hemorrhages. On the other hand, it is speedily restored to its usual standard under the influence of nutritious diet, if the digestive powers have not been too much weakened to make use of this.—The proportion of Red Corpuscles undergoes a marked diminu- tion in various forms of Aneemia; and particularly in Chlorosis. In severe cases of this latter disease, it has been found as low as 27 in 1000; and it not unfrequently sinks to 40 or 50. The marked influ- ence of the administration of Iron, in favoring the reproduction of Red Corpuscles, has been already noticed (§ 219). 533. The proportion of Albumen in the blood seems less liable to change, except in the condition termed Albuminuria, in which a large quantity of Albumen appears in the Urine. When this condition is permanently established, it is indicative of the existence of serious organic disease of the kidney; but it may occur for a short time under the influence of simple congestion of that organ, which causes an escape of the Albuminous part of the blood, together with the water which is filtered off (as it were) in this gland (§ 728). Now Avhen Albuminuria is fully established, there is a marked diminution in the quantity of Albumen in the serum of the blood; and this diminution is constantly proportional to the amount of Albumen present in the Urine.—The proportion of Saline matter appears to undergo less alte- ration in disease, than that of the other constituents of the blood; and has not been found to have a regular correspondence, either in the way of excess or diminution, with any particular morbid state. 534. The condition of the Blood may be affected, not merely by alteration in the proportions of its normal ingredients, but by the pre- sence of other substances;—either such as are generated in it, and are constantly being eliminated from it in health, but have accumulated to 302 ABSORPTION AND SANGUIFICATION. an abnormal degree;—or such as have found their way into it from without. Thus, Carbonic Acid, Urea and Lithic Acid, Cholesterine and other elements of Bile, and other matters which it is the office of the Excreting organs to remove, may accumulate in the blood, and may become fertile sources of disease, by their injurious influences. The introduction of various Mineral substances, by absorption from Avithout, changes the composition of the normal elements of the Blood, and thus affects their vital properties; thus strong saline solutions diminish or destroy the coagulating power of the Fibrine. But the most remarkable cases of depravation of the Blood, by the introduc- tion of matters from without, are those which result from the action of ferments,—exciting such Chemical changes in the constitution of the fluid, that its Avhole character is speedily changed, and its vital pro- perties are altogether destroyed. Of such an occurrence we have a marked example in the various forms of malignant fevers; in which the introduction of a very minute quantity of noxious matter into the blood, either through the lungs or through the skin, produces a speedy alteration in the characters of the whole mass of the blood, the func- tion of every organ in the body is disordered, and decomposition of the solids and fluids takes place to a considerable extent, even before the circulation ceases, and whilst consciousness yet remains. The train of symptoms produced by the bite of venomous Serpents, and of rabid animals, appears referable to the same cause,—the alteration in the condition of the whole current of blood, by the introduction of a minute quantity of a substance that acts as a ferment. 535. The Coagulation of the blood, as already explained, depends upon the passage of its Fibrine from the fluid state to the solid (§ 184); consequently, if the Fibrine be separated from the other elements, no coagulation takes place. On the other hand, if the amount of Fibrine be larger than ordinary, the coagulum possesses an unusual degree of firmness. The length of time which elapses before coagulation, and the degree in which the Clot solidifies, vary considerably ; in general they are in the inverse proportion to each other. Thus, if a large quantity of blood be withdrawn from the vessels of an animal at the same time, or within short intervals, the portions that last flow coagu- late much more rapidly, but much less firmly, than those first obtained, in consequence of the diminished proportion of fibrine. On the other hand, when the fibrine is in excess, its coagulation is unusually de- layed. From this delay an important change results, in the mode in which the coagulation takes place; for the red corpuscles, instead of being uniformly diffused through the coagulum, have time to sink to the bottom, in virtue of their greater specific gravity; and the upper part of the clot is consequently made up of Fibrine, almost exclusively, whilst the lower is chiefly formed by the aggregation of the red cor- puscles. Hence the upper layer is almost destitute of color (whence it has received the name of buffy coat), and is remarkably tenacious in its character; whilst the lower is very deep in hue, and very friable in consistence. When the fibrillated network forming the buffy coat undergoes the slow contraction, which is characteristic of highly-elabo- COAGULATION OF BLOOD—BUFFY COAT. 303 rated fibrine subsequently to its coagulation, it draAvs in the edges of the upper surface of the clot, giving it a cupped appearance. 536. The Buffy Coat may present itself under a great variety of conditions; and it can no longer, therefore, be regarded, as it formerly was, a sign of the Inflammatory state. It is most fully developed when acute Inflammation exists; because in that condition all the circumstances which favor it are present. That it may be produced by any cause, which occasions delay in the coagulation of the blood, is evident from the fact, that healthy blood may be made to exhibit it, by adding a solution of a neutral salt, which retards, but does not prevent its coagulation. But the blood may coagulate with its ordinary rapidity, or even more speedily than usual; and may yet exhibit the Buffy Coat. And, moreover, the separation of the Fibrine and the Red Corpuscles may take place in films of blood so thin, as not to admit of a stratum of one being laid over the other ; the tAvo elements separating from each other laterally, and the films acquiring a speckled or mottled appearance, equally characteristic of the Inflammatory con- dition with the Buffy Coat itself. Hence the separation must be due in such cases, to other causes than gravity: and recent observations have accounted for it, by shoAving that the Red Corpuscles have an unusual attraction for one another in the inflammatory state, causing their coalscence in piles and masses; whilst the particles of Fibrine have also a peculiar strong attraction for each other. Thus there is a poAverful tendency, that draws together the components of each kind, and consequently tends to separate them from the others; and when this separation takes place, the difference in the specific gravity of the Fig. 87. The microscopic appearance of a drop of blood in the inflammatory condition. The red corpuscles lose their circular form, and adhere together ; the white corpuscles remain apart, and are more abundant than usual. two elements decides their respective situations.—The peculiar ten- dency of the Red corpuscles to unite, in the Inflammatory state, serves to distinguish this condition eA7en in a single drop of blood; and it is then that the White corpuscles may be most easily distinguished, as they are seen apart from the rest of the mass, having no tendency to 304 CIRCULATION OF NUTRITIVE FLUID. unite with it. In fact, the white corpuscles are not found in company with the red, in the ordinary coagulum, but rather with the fibrinous portion; and when they are peculiarly abundant, as they usually are in Inflammatory blood, they may form a considerable proportion of the buffy coat. 537. The Buffy Coat may present itself, without the least increase in the normal quantity of Fibrine, and without any approach to the Inflammatory state; simply because the Fibrine is present in exces- sive amount, in relation to the amount of Red corpuscles, the latter being much below their usual proportion. Thus in severe Chlorosis, the buffy coat is almost as strongly marked, as in the severest Inflam- mation ; but the two conditions are at once distinguished by the rela- tive proportions of solid matter in the blood, as indicated by the size of the Coagulum. For in Chlorosis, the coagulum is very small, [in consequence of the reduced proportion of Corpuscles, and is almost invariably found floating in the serum ; whilst in the ordinary Inflam- matory condition, it is of full size, frequently adhering to the side of the vessel. CHAPTER VI. OF THE CIRCULATION OF THE BLOOD. 1. Nature and Objects of the Circulation of Nutrient Fluid. 538. The nutritive fluid,—the elements of which are thus partly taken up and prepared by the Absorbent system, but are in great part also imbibed through the Blood-vessels distributed upon the walls of the digestive cavity, and assimilated by the liver (§ 493),—is carried into the various parts of the system, by the act of Circulation. This move- ment answers various purposes. It furnishes all the tissues, which are to deriAre nutriment from the Blood, with a constantly-reneAved supply of the materials which they severally require; and in this manner it is subservient to the growth, not only of those tissues which form part of the solid structure of the body, but also of those various cells, covering its free surfaces, which are being continually cast off and renewed, and which, in the course of their development, separate from the blood the products that are to perform ulterior purposes in the economy, or are to be removed as altogether effete. Thus the Circulation is subsement to the functions of Nutrition and Secretion. In the exercise of these functions, different materials are draAvn from the blood by the several tissues it supplies. Thus the nutrition of the muscle requires fibrine; that of the nerve requires fatty-matter; that of the bone draws off gela- tine and earthy salts; that of the hepatic cells removes the fatty matter and other elements of bile; that of the milk-cells (during lactation) separates albuminous, fatty, and saccharine substances;—and so on. Thus various portions of the blood, when returning from the several CIRCULATION OF NUTRITIVE FLUID. 305 organs through which they have been transmitted, have undergone very different changes by the nutritive and secreting processes, according to the function of the organs which they have supplied ; and if the same portion of the circulating fluid were constantly being transmitted to each organ, and returning from it, its composition would speedily undergo a change that Avould render it no longer fit for its purposes. By the union of the different local circulations, however, into one general circu- lation, this change is prevented, and the whole mass of the blood is maintained in its normal or regular condition ; for as its composition is such, as to supply all parts of the body, in the state of health, with the proportions of nutritive material which they respectively need; and as the returning currents are all mingled together in the vessels, before being again distributed to the system, each part supplies what the other has been deprived of, and thus the normal proportion of ingredients in the whole mass of the blood is constantly kept up, whilst in each of its separate streams it is undergoing an alteration of a different kind. 539. But these processes alone might be carried on by the aid of a much less rapid Circulation, than that which exists in Man and the higher animals. We do, in fact, occasionally meet with examples in Avhich they continue for some time, under an almost total stagnation of the current. There are others, however, which require a much more rapid and uninterrupted movement of the circulating fluid. We have already seen that, for the action of the Nervous and Muscular tissues, oxygen is necessary; and the amount of that gas contained in the blood circulating through these tissues Avould be very speedily exhausted, if it were not continually renewed ; whilst the carbonic acid, which is formed at the expense of that oxygen, would speedily accumulate to an injurious degree, if it were not carried off as fast as it is produced. Hence we find that, in all Animals, the maintenance of the Respiration, by carry- ing Oxygen from the respiratory surface into the different parts of the system, and by conveying back Carbonic acid to be thrown off at the Respiratory surface, is one of the great purposes of the Circulation of the blood; and its extreme importance is shoAvn by the very speedy check, which the interruption of this function produces in the movement of the blood, in warm-blooded animals. Thus in a Bird or Mammal, completely cut off from Oxygen, the circulation in the lungs will come to a stop, which stoppage will necessarily extend itself over the whole body, in little more than three minutes. We find, then, that the rate of the circulation in different animals bears a relation to the energy of their Respiration; and this energy is closely connected with the general activity of their functions, but particularly with that of the Nervous and Muscular systems, which are most dependent for the exercise of their powers upon a continually fresh supply of oxygen, and upon the un- ceasing removal of the carbonic acid which is generated in their sub- stance. 2. Different forms of the Circulating Apparatus. 540. It is desirable that the Circulating apparatus should be first studied in its very simplest form,—that which it possesses in Plants and 20 306 CIRCULATION OF NUTRITIVE FLUID. in the lowest tribes of Animals; as in this way alone can the forces which are concerned in the movement of the fluid, be rightly appre- ciated. In all the higher Plants, the ascending or crude sap is to be distinguished from the elaborated or descending sap. The former of these fluids should be compared rather with the chyle than with [the blood of Animals ; for it is not yet fully prepared to take part in the nutrition and extension of the structure. But there are some circum- stances attending its movement, Avhich throw light upon other more complicated phenomena. The ascending sap consists principally of water; which is imbibed, together Avith various substances which it holds in solution, by the delicate tissue at the soft extremities of the root-fibres, or spongioles. The poAver of forcing upwards. a column of sap, which exists in these bodies, and Avhich seems due to Endosmose (§ 491), is shoAvn by very simple experiments. If the stem of a Vine, or of any tree in which the sap rises rapidly, be cut across when in full leaf, the sap continues to flow from the lower extremity ; and this with such force as to distend wTith violence, or even to burst, a bladder tied firmly over the cut surface. If, instead of a bladder, a bent tube be attached to this, and mercury be poured into it so as to indicate the pressure exerted, it is found that the rise of the sap takes place with a force equal to the pressure of from one to three atmospheres (from 15 to 45 lbs. upon the square inch)—or even more. Thus the ascent of the sap is partly due to a powerful vis a tergo, or impelling influence, derived from the point where the absorption takes place. 541. But, on the other hand, if the upper extremity be placed with the cut surface of the stem in water, a continued absorption of that fluid will take place, as is evidenced by the withdrawal of the water from the vessel; the fluid Avhich is thus taken up, however, is not retained within the stem and branches, but is carried into the leaves, and is thence dissipated by exhalation. It is obvious, then, that the vis a tergo is not the sole cause of the ascent of the sap; but that a vis a froute also exists, by which the fluid is drawn toAvards the parts in which it is to be employed. This is further made apparent by a few simple experiments. If a branch, when thus actively absorbing fluid, be carried into a dark room, the absorption and ascent of fluid imme- diately cease almost completely ; and are renewed again, so soon as the leaves are again exposed to light. Noav we know, from other experi- ments, that light stimulates the exhaling process (§ 87), whilst darkness checks it; and the cessation of the demand in the leaves thus produces a cessation in the absorption at the lower extremity of the stem. And this is the case, also, in the natural condition of the plant; as is easily shoAvn by immersing the roots in water, and observing the respective quantities which are removed by absorption during sunshine, shade, and darkness. On the other hand the movement of the sap may be excited, when it would not otherwise take place, by the production of a demand at the extremities of the branches ; thus if a branch of a vine growing in the open air, be introduced into a hot-house, and be subjected to artificial heat during the winter, its buds will be developed, its leaves will expand, and these will draAv fluid to themselves through the roots and stems, which are still inactive as regards the remainder of the tree. CIRCULATION OF ELABORATED SAP. 307 And the natural commencement of the movement of the ascending sap, which takes place with the returning warmth of spring, has been ex- perimentally shown to occur, in the first instance, not in the neighbor- hood of the roots, but nearest the extremities of the branches; the exhalation of fluid from the expanding buds being the first process, and a demand for fluid being thus created, which is supplied by the flow that is thus excited in the lower part of the stem,—this, again, being supplied from the roots, which are thus caused to recommence their absorbent function. 542. Thus we see that, in the ascending sap, the movement is en- tirely regulated by the demand for fluid occasioned by the actions of the leaves ; even though it is in great part dependent on the via a tergo which has its seat in the spongioles. Not even this force, however,— poAvcrful as it has been shoAvn to be,—can produce the continuance of the upward flow, when the exhalation from the leaves is checked by darkness, and when the demand occasioned by the action of these organs is consequently suspended. 543 The movement of the descending sap offers numerous points which deserve to be carefully considered. This fluid is strictly compa- rable to the blood of animals; having undergone a preparation or elaboration in the leaves, which adapts it to the nutrition and extension ot the structure, and to the formation of the various secretions of the plant. A great part of the fluid of the ascending sap has been lost by exhalation: and the remainder, thus concentrated, receives a large additional supply of solid matter through the agency of the green cells of the leafy parts, which take in carbon from the atmosphere (§ 83) • so that it now includes a considerable amount of gummy matter in the state prepared for being converted into solid tissue, as well as numerous other compounds. Now this elaborated sap seems to be conveyed into the various parts of the system, partly by transmission from one cell to another, but partly through the agency of a network of vessels, which takes its origin in the leaves, and extends along the branches to the stem and roots, chiefly in the bark of those parts. These vessels are strictly analogous to the capillaries or small blood-vessels of Animals • but they differ from them in this,—that the capillary network of Ani- mals communicates on either side with larger trunks, being formed, in fact, by the interlacement or anastomosis of their minutest branches' — whilst the network of nutritive vessels in Plants is everywhere con- tinuous with itself, not having any communication Avith large vessels, so that the fluid prepared in the leaves commences a circulation there which is continued on the same plan, until it has found its way to its' remote destination in the roots. 544. The natural movement of the elaborated sap through these vessels may be studied, under favorable circumstances, with the assis- tance of the microscope ; the requisite conditions being, that the part should be sufficiently transparent for the vessels to be distinctly seen that the sap shall contain globules in sufficient number to allow its' movement to be distinguished by their means, and that the circulation should be observed without the separation of the organ examined from the rest of the Plant, Avhich would produce irregular movements by the 308 CIRCULATION OF NUTRITIVE FLUID. escape of the sap from the wounded part. These conditions may be attained in many Plants;—most conveniently, perhaps, in the stipules of the Ficus elastica, one of the trees which affords the largest supplies of Caoutchouc; and it is then found that the movement takes place in the following manner. Distinct currents are seen, passing along the straightest and most continuous vessels, and crossing by the lateral connecting branches of the network. These currents follow no deter- minate direction ; some proceeding up, and others down; some to the left, and others to the right; not unfrequently a complete stoppage is seen in one or more of the channels, without any obvious obstruction; and the movement then recommences, perhaps in the opposite direction. The influence of a force, developed by the act of circulation, which determines the direction of the movement, appears from this : that if a tube be cut off, so as to give its contents an equally free exit at both ends, the sap only flows out at one extremity. The movement is retarded by lowering the temperature of the surrounding air, and it is completely checked by extreme cold; it is capable of being renewed by moderate warmth; and a further addition of heat increases its rapidity. By a strong electric shock, the force by which the liquid is propelled seems to be altogether destroyed; for the movement then ceases entirely. 545. Noav it is quite certain that this circulation cannot be due to any vis a tergo ; both because it is not constant in its direction in par- ticular vessels ; and because there is no organ in Avhich any propelling force, that could extend itself through such a complex system of vessels, may be developed. Nor can it be in any way due to the force of gravity; for although this may assist the descent of the fluids through the stem, it is totally opposed to its ascent from the ends of its branches toAvards their origin, when, as often happens, the latter are at the higher level. Moreover, it may be noticed that this circulation takes place most readily, in parts that are undergoing a rapid development; and that its energy corresponds with the vitality of the part. Further, it may be observed to continue for some time in parts that have been completely detached from the rest; and on which neither vis a tergo, nor vis a fronte, can have any influence. It is evident, then, that the force,— whatever be its nature,—by which this continued moA'ement is kept up, must be developed by the processes to which that movement is subser- vient ; in other words, that the changes involved in the acts of nutrition and secretion are the real source of the motor power. The manner in which they become so, is the next object of our inquiry; and on this subject, some new views have recently been put forth by Prof. Draper, which seem to account well for the phenomena. 546. It is capable of being shown, by experiments on organic bodies, that, if two liquids communicate with each other through a capillary tube, for the walls of which they both have an affinity, but this affinity is stronger in the one liquid than in the other, a movement will ensue; the liquid which has the greatest affinity being absorbed most energeti- cally into the tube, and driving the other before it. The same result occurs when the fluid is drawn, not into a single tube, but into a net- work of tubes, permeating a solid structure; for if this porous struc- CIRCULATION IN PLANTS AND LOWER ANIMALS. 309 ture be previously saturated with the fluid, for which it has the less degree of attraction, this will be driven out and replaced by that for which it has the greater affinity, when it is permitted to absorb this. Now if, in its passage through the porous solid, the liquid undergo such a change, that its affinity is diminished, it is obvious that, according to the principle just explained, it must be driven out by a fresh supply of the original liquid, and that thus a continual movement in the same direction would be produced. 547. Now this is precisely that which seems to take place in the organized tissue, permeated by nutritious fluid. The particles of this fluid, and the solid matter through which it is distributed, have a certain affinity for each other; which is exercised in the nutritive changes, to which the fluid becomes subservient during the course of its circulation. Certain matters are drawn from it, in one part, for the support and increase of the woody tissue ; in another part, the secreting cells demand the materials Avhich are requisite for their growth,—as starch, oil, resin, &c.; and thus in every part that is traversed by the vessels, there are certain affinities between the solids and the fluids, Avhich are continually being developed afresh by acts of growth, as fast as those which pre- viously existed are satisfied or neutralized by the changes that have already occurred. Thus in the circulation of the elaborated sap, there is a constant attraction of its particles towards the walls of the vessels, and a continual series of changes produced in the fluid, as the result of that attraction. The fluid, which has given up to a certain tissue some of its materials, no longer has the same attraction for that tissue; and it is consequently driven from it by the superior attraction then pos- sessed by the tissue for another portion of the fluid, which is ready to undergo the same changes, to be in its turn rejected for a fresh supply. Thus in a groAving part, there is a constantly-reneAved attraction for the nutritive fluid, Avhich has not yet traversed it; whilst, on the other hand, there is a diminished attraction for the fluid, which has yielded up the nutritive materials required by the particular tissues of the part; and thus the former is continually driving the latter before it. 548. But the fluid which is thus repelled from one part, may still be attracted toAvards another; because that portion of its contents which the latter requires, may not yet have been remoAred from it. And in this manner, it would seem, the flow of sap is maintained, through the whole capillary netAvork, until it is altogether exhausted of its nutritive matter. The source of the movement is thus entirely to be looked for in the changes Avhich take place in the act of growth; and the influence of heat, cold, and other agents, upon the movement is exercised through their power of accelerating or retarding those changes.—The fluid which thus descends through the stem and roots, seems to be at last almost en tirely exhausted ; a portion of it appears to find its way into the interior of the stem, and to be mingled with the ascending current; but all the rest seems to have been entirely appropriated by the different tissues, through which it has circulated. Thus there is no need of any general receptacle, into which it may be collected, and from which it may take a fresh departure;—such as is afforded by the heart of Animals. And as the purpose of this circulation is only to supply the nutritive mate- 310 CIRCULATION OF NUTRITIVE FLUID. rials, and not to convey oxygen,—this element being but little required in the vegetative processes, and being supplied by other means,—the same energy and rapidity are not required in it, as need to be provided for in the higher Animals. 549. A condition of the Circulating system very similar to this, exists in several of the lower animals, as well as in the embryo-state of the higher. In the very lowest no blood-vessels are required, for the same reason that no sap-vessels exist in the loAvest Plants;—namely, because every part absorbs and assimilates nutritious fluid for itself, so that it does not require a supply from vessels. As, in the sea-Aveeds, the Avhole substance is nourished by direct absorption from the fluid in contact with the external surface, every part of which seems endowed with the same absorbent poAver, so in Zoophytes do we find; that the whole sub- stance is nourished by direct absorption from the internal surface, which forms the lining of the digestive cavity. In the same manner, the Aeration of the animal fluids,—or the exposure of them to the air contained in water, by Avhich they may part with carbonic acid and imbibe oxygen,—is provided for, not by any special respiratory organs, but by the contact of Avater with every part of the soft external and internal surfaces. Further, as the substance of their body is nearly of the same kind in every part, they do not require the continual inter- change of the fluid distributed to its several portions. Thus no circu- lation is necessary, in these simple animals, either for the nutrition of their tissues, or for the aeration of the fluids. The same is the case with others of the loAver tribes ; as well as with the embryo of the higher Animals, at the earliest period of their development. Thus the lowest Entozoa, or parasitic worms, have a digestive cavity channelled out, as it Avere, in their soft gelatinous tissues; and from the walls of this, the nourishment is drawn by the several component parts of those tissues, without the mediation of vessels. And the embryo even of Man, in its early condition, consists of an aggregation of cells, each of which absorbs for itself from the nutritious fluid with Avhich it is surrounded, and goes through all its functions independently of the rest. 550. Proceeding a little higher, we find the first appearance of proper vessels in the higher Entozoa, and in the Echinodermata. These vessels take up the nutritive fluid from the Avails of the digestive cavity, on which they are spread out, just as the roots of Plants do from the soil. They then unite into trunks, by which the fluid is conveyed to the more distant parts of the structure, in the same manner as the ascending sap is conveyed to the leaves by the vessels of the stem and branches; and these trunks again subdivide, and form a netAvork of capillary vessels, which are dispersed through the several parts of the fabric ; some of them being very abundantly distributed upon a portion of the surface, which is particularly destined to perform the respiratory function. Through these capillary vessels, the fluid seems to move in very much the same manner, as through the system of anastomosing vessels in Plants ;—that is, its motion is due, rather to forces which are developed during its circulation, than to any vis a tergo derived from the contractile power of a propelling organ. But there is this difference: that, after having traversed the minute vessels, and yielded up to the tissues a part CIRCULATION IN ARTICULATED ANIMALS. 311 of the solid matter which it contains, the fluid is collected^ again by other trunks, Avhich convey it back to the point from which it started; there it is mingled with the fluid that has been newly absorbed, and with that which has undergone aeration; and it is then distributed, as before, through the general capillary network of the body. 551. Now this is very much the condition of the Human embryo, at the time Avhen vessels are first developed in its substance. These ves- sels are formed by the coalescence of cells ; and from the contents of these cells, Avhich have been imbibed from the yolk, the first blood seems to be derived. The first formation of blood-vessels takes place, not in that part of the embryonic structure which is to be developed into the perfect animal, but in a membranous expansion from it, which surrounds the yolk, and which answers the purpose of a temporary stomach. A capillary network is formed in a limited portion of this membrane, termed the vascular area (Fig. 88); and this is not by the branching of Fig. 88. Vascular area of Fowl's eggs, at the beginning of the third day of incubation;—a, a, yolk; 6, b, b, b, venous sinus bounding the area; c, aorta; d, punctum saliens, or incipient heart; e,e,area pellucida; /,/, arteries of the vascular area; g, g, veins; h, eye. larger trunks, these trunks being subsequently formed by the reunion of the capillaries. The first movement of the blood is towards the cen- tral spot, in Avhich the organs of the permanent structure are being evolved; and it takes place before the incipient heart has acquired any muscularity, so that it must be quite independent of any contractile force exerted by that organ. Here too, then, we perceive that the circulation is essentially capillary ; and that it is sustained by forces very different from those, of Avhich the action is most evident to us in the higher animals. 552. As Ave ascend the animal scale, however, we find that provision is made for a more regular and vigorous Circulation of the Blood, than that which exists in the lowest classes. Even in the class of Echino- dermata (including the Star-fish and Sea-urchin), a portion of the prin- cipal vessel is peculiarly endowed with contractile power; and this may be seen in constant pulsation, like the heart of the higher animals, alternately contracting, to propel the fluid it contains, through the ves- 312 CIRCULATION OF NUTRITIVE FLUID. sels that issue from it, and then dilating, to receive a fresh supply from the vessels that pour their contents into it. It seems quite certain, however, from the extent of the vascular system of these animals, that the influence of such a pulsatile cavity must be quite insufficient to keep up the movement of blood through it. A similar provision is observable in the lower tribes of Worms, in which this contractile vessel lies along the back; propelling the blood forwards, by a sort of peristaltic move- ment, through trunks which pass out at its anterior termination; and receiving it again after it has circulated through the system, by vessels which enter at its posterior extremity. In the higher orders of Worms, in the Myriapoda or Centipede tribe, and in Lnsects, we find this dorsal vessel divided by transverse partitions containing valves, into separate cavities which answer to the different segments of the body. Each of these is, to a certain extent, the heart of its own segment, receiving and propelling blood by trunks which open into it; but they all participate in the more general circulation just described, a large portion of the blood being poured into the hindermost segment, transmitted forwards from cavity to cavity through the valves which separate them, and at last propelled through trunks that issue from the most anterior segment. In some instances we find that two or three of these trunks, on either side, pass round the oesophagus, and reunite beloAv it, so as to enclose it in a sort of collar; and they form a main trunk by this union, which runs backwards along the under surface of the body, and which distri- butes the blood to its different organs by lateral branches. These subdivide into a capillary network, and the returning vessels, Avhich originate in this network, pour the blood which has circulated through it into the posterior cavity of the dorsal vessel.—Still it is very evident from the observation of the circulation in those transparent species in Avhich the Avhole process can be distinctly watched under the Microscope, that the contractile power of the dorsal vessel is far from sufficient of itself to sustain the Circulation; and that the movement of the blood through the capillary network is in part due to forces deve- loped during its progress, being often retarded or accelerated in parti- cular spots, without any visible change in the propelling force of the eentral organ. Moreover, the blood, during some part of its course, almost always escapes from the proper vessels into lacunce channelled among the tissues; and over its flow through these, no central impelling organ can have much influence. 553. In most of these animals, there are distinct organs of Respira- tion, confined to some one part of the body ; and we often find that the vessels which convey blood to them, are furnished with distinct con- tractile portions, like so many supplementary hearts, for the purpose of propelling the blood through them more energetically. In proportion as we ascend the series of Articulated animals, do we find for the most part, a more vigorous and regular circulation, both for the nutrition of the system, and for the transmission of the blood through the respira- tory organs; but there is an exception in the case of insects, which deserves special notice. In this class, the circulation is much less vigo- rous than it is in other Articulated animals of similar complexity of structure; though it might have been anticipated, that the extraordinary CIRCULATION IN MOLLUSCS. 313 activity of their movements would necessitate a corresponding rapidity in the circulating current, especially for the purpose of conveying an extraordinary supply of oxygen to the nervous and muscular systems. But this is provided for in another way; the air being conveyed to these tissues not through the blood, but by direct transmission through the minute ramifications of the air-tubes or tracheae, which penetrate the very smallest organs of the body (§ 659). 554. The condition of the Circulating apparatus in the Embryo of higher animals, at a period a little advanced beyond that just alluded to, presents a striking analogy with that last described ; for the heart, at the time of its first formation, seems like a mere dilatation of the princi- pal vascular trunk, having thickened walls, in which, after a time, mus- cular fibre begins to be developed, and the contractile power manifests itself. The pulsation of this heart, hoAvever, does not seem to extend its influence immediately through the vascular area ; the capillary circu- lation in which, remains for some time in great degree independent of it. There is no resemblance inform, hoAvever, between the dorsal vessel of Insects, and the incipient heart of the higher animals; since the latter is never much prolonged, and speedily becomes doubled (as it were) upon itself; and its first division into distinct cavities is merely for the purpose of separating its receiving portion, or auricle, from its propelling portion, or ventricle. But the general condition of the Circulating system is much the same in the two cases; and it is further alike in this,—that it is not always easy to show that the vessels haA^e distinct walls, as they frequently seem like mere channels excavated in the tissues. 555. We may next turn our attention briefly to the condition of the Circulating apparatus in the Molluscous classes, Avhich has lately been found to present some very peculiar characters. In these it would seem as if the moving power were more concentrated in the heart, than in the preceding; for this organ seems no longer like a mere dilatation of the vascular trunk, but is a distinct sac with muscular walls, usually having at least tAvo cavities, an auricle and a ventricle. The usual course of the Circulation is the following. The blood, expelled from the ventricle of the heart, passes along the main systemic artery, or aorta; which distributes it to the body at large. It is then collected again, and transmitted to the respiratory organs ; in Avhich it is exposed, either to the air contained in the surrounding Avater, or (in the terres- trial Molluscs) more directly to the atmosphere; and from these it is returned to the heart, to be again transmitted to the system.—Thus we see that the heart of these animals receives and impels aerated blood; and that its office is, to send that blood to the capillaries of the general system. Hence it may be called a systemic heart. 556. The blood, in the first part of its course, passes through dis- tinct vessels : it has been lately shoAvn, however, that in the Molluscs in general, the blood which has passed through the systemic capillaries, and is on its Avay to the respiratory organs, is no longer thus confined, but that itjneanders through passages or lacunar, which are channelled out in the tissues, and which even communicate freely with the abdomi- nal cavity in Avhich the viscera lie ; so that their whole exterior is 314 CIRCULATION OF NUTRITIVE FLUID. bathed by the circulating fluid. It is perhaps in this part of its course, that it most readily takes up the fresh nutrient materials, which have been prepared by the digestive process, and which Avould, under such circumstances, find their way with comparative facility from the inner surface of their walls, to the outer.—After being thus diffused, in its venous or carbonized state, through the substance of the tissues and through the visceral cavity, it is again collected into distinct trunks; and these convey it to the respiratory organs.—Now although it cannot be doubted, that the impelling power of the heart is the chief cause of the movement of the blood through the systemic vessels, yet it would seem impossible to suppose, that this power can be exerted over the unrestrained currents, in which it is diffused through the body, after passing through the systemic capillaries ; and it can scarcely be doubted, that its passage through the capillaries of the respiratory organs is due to the power which is developed in themselves, under the conditions already alluded to. 557. There is a very curious phenomenon to be observed in the cir- culation of some of the loAvest Molluscs ; namely, the continual reversal of the course of the current. The heart, in these animals, is much less perfectly formed, than in the higher tribes; and seems more like the mere contractile dilatation of the principal trunk, which is the sole representative of that organ in the Echinodermata. The circulating fluid is sometimes transmitted first to the system;. and, after being dis- tributed to its different parts by jhe ramifications of the main artery, it meanders through the channels excavated in its tissues; and then flows toAvards the respiratory surface, after passing over which, it re- turns to the heart. But after a certain duration of its flow in this direction, the current stops, and then recommences in the contrary direction,—proceeding first to the respiratory organs, and then to the system in general. It would seem as if in this, one of the lowest forms of animals possessing a distinct Circulation, the central poAver were not yet sufficiently strong, to determine the course which the fluid is to take : so that it undergoes continual vacillations. In a group of Com- pound Polypes, to which this class of Molluscs has many points of affinity, there is a movement of fluid through the stem and branches, which in like manner continually changes its direction. This move- ment, hoAvever, can scarcely be regarded in the light of a proper Cir- culation ; since the tubes in which it occurs are in direct communication with the digestive cavities of the Polypes. But the flow seems altogether independent of any mechanical propulsions ; and takes place most ener- getically and regularly towards parts in Avhich new growth is going on. 558. We have noAv to consider the chief forms in which the Circu- lating apparatus presents itself in the Vertebrated classes ; and first in that of Fishes. We have here, as in Molluscs, a heart with two cavi- ties, an auricle and a ventricle ; this heart, however, is not placed at the commencement of the systemic circulation, but at the origin of the respiratory vessels. The blood which it receives and propels, is venous or carbonized; this is transmitted along a main trunk, which speedily subdivides into lateral branches or arches ; and these distribute it to the fringes of gills, that hang on the sides of the neck. By the action CIRCULATION IN FISHES. 315 of the water on the gills, the blood is aerated in its passage through them ; and it is then collected by a series of converging vessels, which re- unite to form the great systemic artery, or aorta. By the ramifications of this artery, the blood, now FiS- 89- aerated, is distributed through the system, and affords the requisite nourishment and stimulation to its tissues. Returning from the systemic capil- laries in a venous state, the blood of the head and anterior portion of the body finds its Avay at once into the great systemic vein, or vena cava, by Avhich it is conveyed back to the auricle of the heart; but that which has traversed the capillaries of the posterior part of the body, and of the ab- dominal viscera, is conveyed by a distinct system of veins to the liver and the kidneys. In these organs, the veins again subdivide into a network of capillaries, which is distributed through the secreting structure, and which serves to afford to the secreting cells the materials of their develop- ment. This is termed the portal system of vessels. ~. -171 ,1 -n • /• .1 v -ii-i .1 Diagram of the Circulating .rJrom the capillaries of the liver and kidneys, the Apparatus of Fishes:— a, the blood is finally collected by the hepatic and renal trunkVuppiylngniebranrfiia'i veins, which convey it into the vena cava; where ^ll^X^^^n- it is mingled with the blood that has not passed veycd *y «> «> the branchial r i , i-i 11 veins, to /, the aorta, which through those organs, and is thus conveyed to the distributes it to the system; Vionrl- thence it is collected, and re- neart. turned to the auricle, by the 559. The heart of Fishes, then, belongs to the ^^ich unite in the vena respiratory circulation. It propels venous blood to the capillaries of the gills, in Avhich it is aerated; returning from these, the aerated blood is transmitted through a second set of capillaries, those of the system, in which it again becomes venous : whilst a portion of this blood is made to traverse a third set of capil- laries, those of the liver and kidneys, before it is again subjected to the propelling poAver of the heart. Noav as the heart, instead of being stronger than it is in animals with the complete double circu- lation presently to be described,—in which the greater part of the blood propelled by it only traverses one set of capillaries, and never more than tAvo,—is much Aveaker in proportion, it is evident that here, too, a supplementary poAver must exist, by which the flow of blood through the capillaries is aided, and on Avhich, indeed, the portal circu- lation must greatly depend. 560. An extremely interesting aspect of the circulating apparatus is presented by the Amphioxus or Lancelot; an animal which presents the general form of a Fish, and which can scarcely be referred to any other group ; but in Avhich the characters of the Vertebrated series are de- graded (as it were) to the level of the lower Molluscous and Vermiform classes. The blood, which is white, moves through distinct vessels, but there is no proper heart; and the vascular trunks present several dila- tations, in different parts, which have muscular walls, and show con- tractile power. Thus the circulation is carried on, not through the 316 CIRCULATION OF NUTRITIVE FLUID. agency of a central impelling organ, as in other Fishes ; but by a power Avhich is scattered or diffused through various parts of the system of blood-vessels, as in the lower Invertebrata.—The respiratory apparatus, also, is formed upon a type much lower than that of Fishes ; for it con- sists simply of a dilatation of the first part of the alimentary canal, or pharynx, upon the walls of which the blood is distributed in divided streams, its cavity being filled with water, which serves to aerate the blood. This is precisely the type, on which the respiration is effected, in those lowest Molluscs, of which mention has just been made, as ex- hibiting alternations in the direction of the circulating current (§ 557). In other respects, however, the arrangement of the vascular system in this extraordinary animal corresponds with that which obtains in Fishes. 561. It is requisite that, in the class of Fishes, the whole of the A'enous blood returned from the system should pass through the respi- ratory organs before being again transmitted to the body ; since the aerating action of the small quantity of air diffused through the Avater, would otherAvise be insufficient for its renovation. But in Reptiles, all of which breathe air during their adult condition, the case is very different; for if the whole current of their blood were exposed to the atmosphere, before being again sent to the body, the quantity of oxygen conveyed into the tissues would be too great, and would have an over-stimulating effect. The plan of the Circulation is, therefore, differently arranged in Reptiles. We find the heart to consist of three cavities ; tAvo auricles and one ventricle. From the ventricle issues a single trunk, which speedily sub- divides ; some of its branches proceeding to the lungs, and others to the body. The blood which is trans- mitted through this trunk, is of a mixed character, as we shall presently see ; being neither fully aerated, nor yet highly carbonized. It contains sufficient oxygen, to stimulate the nervous and muscular sys- tems of these comparatively inert animals ; whilst it tio^ln'RTptnes^-a.'sin^e a^so contains enough of carbonic acid, to require ventricle, receiving the being exposed to the atmosphere through the medium aerated blood from 6, the « ,° . * m, , , i i • 1 1 ii 1 pulmonary auricle, and ve-ot the lungs. Ihe blood which has passed through temic^u^cfe^nd^o^l- tne systemic capillaries, and which has been thereby tonthPeartu0imon\?iXceadfira^ rendered completely venous, is returned to one of ries a, and part to the sys- the auricles—the systemic—by the vena cava. On the other hand, the blood which has passed through the capillaries of the lungs, and which has been thereby rendered com- pletely arterial, is returned through the pulmonary vein to the other auricle,—the pulmonary. Thus one of the auricles exclusively receives aerated, and the other carbonated blood ; and as both pour their con- tents into the common ventricle, the blood which that cavity contains and propels is of a mixed character. 562. Various modifications of this form of Circulating apparatus exist in the different groups of reptiles. In the lowest among them, CIRCULATION IN REPTILES. 317 which breathe permanently by gills like Fishes, besides possessing imperfectly-developed lungs, the apparatus exhibits a blending of both plans; for a small portion of the blood, which is propelled by each contraction of the ventricle, passes directly to the lungs; the principal part of it being at once distributed to the gills, as in Fishes. After passing through these, it is transmitted to the general system; and on returning thence, in a completely venous state, it is mingled with the blood which has been arterialized in the lungs. This latter, hoAvever, bears so small a proportion to the rest, that, if the aeration were not partly effected by the gills, it would be insufficient for the wants of the animal. The tadpoles of the common Frog and Water Newt, as well as of other species which, like them, begin life in the general condition of Fish, present a similar condition at one period of their change. At first, the whole aeration is effected by means of gills, the lungs being in a rudimentary or undeveloped state; and the entire circulation is carried on as in Fishes, the pulmonary vessels being scarcely traceable. As the lungs begin to be developed, however, a portion of the blood is sent to them ; and at the same time, communicating passages which previously existed, betAveen the vessels that convey blood to the gills, and those that return it from them, are increased in size; so that a certain proportion of the blood is transmitted to the system, Avithout having passed through the gills at all. By a further increase in the diameter of these, the whole current of blood takes this direction, the gills being no longer serviceable; and as, at the same time, the lungs are attaining their full development, the aeration which they effect in the blood transmitted to them becomes sufficient, and the whole circula- tion is thus permanently established on the Reptilian type. 563. On the other hand, among the higher Reptiles, we find the circulating apparatus presenting approaches to the form it possesses in Birds and Mammals. For the ventricle is divided, more or less completely, into tAvo cavities, one of which propels aerated blood to the system, whilst the other transmits venous blood to the lungs. A certain amount, of mixture of arterial and venous Wood always takes place, however, either in the heart itself or in the vessels; so that the blood which the body receives, is never purely arterial. But this mixture is sometimes effected in such a manner, that pure arterial blood is sent to the head and anterior extremities; though the re- mainder of the body receives a half-aerated fluid. This is accomplished in the Crocodile, by a provision very similar to that which exists in the foetus of warm-blooded animals, (chap, xi.) The portal circulation in Reptiles is carried on nearly upon the same plan as in Fishes. It receives the blood from the posterior extremities and from the tail, as well as from the abdominal viscera; and this blood is distributed by the portal capillaries, not only through the liver, but also through the kidneys, although the latter also receive arterial branches from the aorta. The fact that the kidneys are supplied from the general portal circulation in Fishes and Reptiles, has an important bearing on the difference in the arrangement of their OAvn vessels, which will be here- after shown to exist, betAveen the kidneys of these animals and those of Birds and Mammals (§ 728). 318 CIRCULATION OF NUTRITIVE FLUID. 564. In the warm-blooded division of the Vertebrated series, which includes the classes of Birds and Mammals, we find the whole circu- lation possessed of a greatly-increased energy; but the distinguishing peculiarity of the apparatus in these animals, is that conformation of the heart and vessels which secures a complete double circulation of Fig. 91. the blood; that is, which provides for the aeration of every particle of the venous blood which has re- turned from the system, before it is again sent into the tissues. The heart may be regarded as con- sisting of tAvo distinct parts,—a systemic heart, like that of the Molluscs, forming its left side,—and a respiratory heart, like that of Fishes, constituting its right. Each of these parts has a receiving cavity or auricle, and an impelling cavity or ven- tricle. The cavities of the two sides are completely separated from one another, in the adult state at least; though their Avails are united, for economy of material. It is obvious that much is saved in this manner ; since, as the contractions of the auricles and of the ventricles on the two sides occur simul- taneously, the pressure of blood in the one is partly Diagram of the Circulating , • i i ,i xi ^i l -. ? Apparatus in Mammals and antagonized by that on the other, wherever it acts KiT^tes-MeMcwai on the wall that is common to both. This antago- th^IffiMridetfeerighi ™sm *s not complete, however ; since the systemic ventricle, propelling venous ventricle contracts with far greater force than the blood through e, the pulmo- . i,i ni, , i ii nary artery, to /, the capii- pulmonary ; and the wall between them must be inriciefrl«ilto| tof ^d capable of resisting the difference of pressure on wood fro^ tte pulmonary jts tw0 gjjgg thus occasioned. The blood which is vein, and, delivering it to the ' left ventricle, h, which pro- returned from the system, in a venous state, through pels it, through the aorta, i, , , ,i • i , • i i i • i • to the systemic capillaries, the vena cava to the right -auricle, and Avhich is 4ew vdns, and carried %& poured by it into the right ventricle, is impelled by ctavtahe6heartthr°ughthe7ena tne latter through the capillaries of the lungs, v where it undergoes aeration. Returning thence in an arterialized state, it is conveyed into the left auricle, and thence flows into the left ventricle; by which it is propelled through the great systemic artery or aorta, and through its ramifications to the general system. ♦* 565. The greater part of the blood Avhich has been rendered venous by passing through the systemic capillaries, is collected by the systemic veins, and is returned directly to the heart through the vena cava. But a portion is still employed for the distinct circulation, which is destined to supply the materials for the secreting action of the liver. The blood that has traversed the capillaries of the walls of the alimentary canal, and of the other viscera concerned in digestion, is collected again by the converging veins into a large venous trunk, the vena portse, by which it is distributed through the liver. This vessel, although formed by the convergence of veins, and conveying venous blood, has really the character of an artery in an equal degree; for it subdivides and ramifies after its entrance into the liver, so as to form a netAVork of capillaries, from which the blood is again collected, and thence trans- CIRCULATION IN MAMMALS. 319 mitted by the hepatic vein to the vena cava. Thus that portion of blood Avhich supplies the liver with the materials of its secreting action, passes through two sets of capillaries, between the time of its leaving Fig. 92. Anatomy of the Human Heart and Lungs. 1. The right ventricle; the vessels to the right of the figure are the middle coronary artery and veins; and those to its left, the anterior coronary artery and veins. 2. The left ventricle. 3. The right auricle. 4. The left auricle. 5. The pulmonary artery. 6. The right pulmonary artery. 7. The left pulmonary artery. 8. The remains of the ductus arteriosus. 9. The arch of the aorta. 10. The superior vena cava. 11. The right arteria innominata, and in front of it the vena innominata. 12. The right subclavian vein, and behind it its corresponding artery. 13. The right common carotid artery and vein. 14. The left vena innominata. 15. The left carotid artery and vein. 16. The left subclavian vein and artery. 17. The trachea. 18. The right bronchus. 19. The left bronchus. 20, 20. The pulmonary veins; 18, 20, form the root of the right lung; and 7, 19, 20, the root of the left. 21. The superior lobe of the right lung, 22. Its middle lobe. 23. Its inferior lobe. 24. The superior lobe of the left lung. 25. Its inferior lobe.' the heart and its return to it. The portal circulation in Birds, as in Reptiles and Fishes, receives the blood from the posterior part of the body, and from the extremities; but the portal blood is only conveyed to the liver; the kidneys being supplied by the renal artery. 566. This perfect form of the Circulating apparatus is only attained, in the warm-blooded animal, after a series of transformations, which strongly remind us of the permanent forms presented by the vascular system in Fishes and Reptiles. Thus in the embryo of the Chick at about the 60th hour, and in that of the Dog at about the 21st day, the curved and dilated tube, of which the heart previously consisted (§ 554), is found to be distinctly divided into an auricle and a ven- tricle. From the latter originates the main arterial trunk, which divides into four pairs of lateral branches; and these pass round the pharynx, precisely in the position and direction of the arterie's of the gills of Fishes. They do not, however, distribute the blood to gill- tufts; for none such are developed in the embryo of the warm-blooded animal: but they meet again below the pharynx, to form a trunk, Avhich supplies the general circulation. Within a short period, how- ever, the Avhole plan of the circulation undergoes a change. The auricle 320 CIRCULATION OF NUTRITIVE FLUID. and the ventricle are each divided by a partition, that is developed in the middle of the heart; and thus the two auricles and the tAvo ven- tricles are formed. Whilst this is going on, a change takes place also in the vessels that arise from the heart; for the arterial trunk, that was previously single, undergoes a division into two distinct tubes ; one of which is connected with the left ventricle, and becomes the aorta, whilst the other originates in the right ventricle, and becomes the pul- monary artery. Of the four pairs of branchial arches, some are sub- sequently obliterated; whilst others undergo changes that end in their becoming the arch of the aorta, the right and left pulmonary arteries, and the right and left subclavians. 567. The muscular power of the heart is much greater in the warm- blooded than in the cold-blooded Vertebrata, in proportion to the ex- tent of the circulation which it is concerned in maintaining; and it is r_evidently destined to take a much larger share in the propulsion of the fluid, than it is in the lower tribes. Marry Bh-ysiologisLs^indeecLAfe of opinion that the jnojjenient gXJ^hsJolpo^^_entirely'due to the actum, ofLtdiehearj; and~tnis view appears toBe sup"ported by the results of [numerous experiments upon the circulation. But it is very difficult, if """"hot""impossible, to make experiments that shall be really satisfactory upon this point; and it appears safer to trust to the " experiments ready prepared for us by Nature," as Cuvier termed them,—namely, those lower forms of animated being, in which various diversities of structure present themsehres, and in which we can study the regular and undisturbed effects of these. Thus we have seen that, in Plants and the lowest Animals, Avhich have no central impelling cavity, the movement of the nutritive fluid is entirely dependent upon the power that is diffused through the network of vessels in which it circulates. As Ave ascend the series, we find an organ of impulsion developed upon a certain part of the vascular system, whose object it is to give increased energy and regularity to the movement. And ascending still higher, we find the moving power gradually concentrated, as it were, in this organ; yet it is not altogether withdrawn from the capillary net- work, as we shall see from several facts to be presently adduced. The particular actions of the Heart, the Arteries, the Capillaries, and the Veins, will now be considered in more detail. • 3. Action of the Heart. 568. The Heart is a hollow muscle, endowed in an eminent degree with the property of irritability; by which is meant, the capability of being easily excited to movements of contraction alternating with relaxa- tion (§ 347). At first sight, its actions seem different from that of the muscles which are called into action by the impulse of the will; for in these there is apparently no such alternation, the state of contraction being kept up as long as the will operates. But it has been already explained that, even in these, the individual fibres are probably in a state of continual alternation of contraction and relaxation, during their active condition,—one set taking up the action, whilst another is return- ing to the state of relaxation. Hence the chief peculiarity in the Heart's MOVING POAVERS OF THE CIRCULATION. 321 action consists in this,—the whole mass of fibres of each division of the organ contract and relax together. The contraction of the tAvo ventricles is perfectly synchronous, as is that of the two auricles; but the contraction of the auricles is synchronous Avith the dilatation of the ventricles, and vice versa. The regularity of this alternation, however, is somewhat disturbed, when the irritability of the heart is becoming exhausted ; and both sets of movements will continue, when the auricle and ventricle have been separated from one another. The regular suc- cession, in the natural state, is doubtless in part due to the fact, that the transmission of blood from the auricle into the ventricle, by the contraction of the former, is the stimulus Avhich most effectually excites the latter to contraction; Avhilst the ventricle is contracting, the auricle, now free to dilate, is distended by the flow of blood from the veins that open into it; and this flow stimulates it to renewed contraction, just at the time when the contraction of the ventricle has been completed, and its state of relaxation enables it to receive the blood poured in through the orifice leading from the auricles. 569. In the living animal, the auricular and ventricular movements succeed one another with great regularity; and, when the circulation is proceeding with vigor, scarcely any appreciable pause can be dis- covered betAveen the different acts. The contraction or systole of the Auricles takes place precisely at the same moment with the dilatation or diastole of the Ventricles; and, as soon as the latter are full, and the former are empty, the diastole of the Auricles and the systole of the Ventricles, immediately succeed. The systole of the Ventricles occa- sion the propulsion of blood into the arterial system; and this action produces the pulse, as will be explained hereafter. And it also corre- sponds Avith the impulse or stroke of the heart against the parietes of the chest. This impulse is not produced, as some have supposed, by the swinging of the entire heart forwards; but by the peculiar mode in Avhich the Ventricular systole takes place. In the contraction of its walls, every dimension is lessened; but shortening is the most percep- tible change, the vertical diameter of the Ventricle being the greatest. Owing to the peculiar spiral disposition of the fibres of the heart, its apex is not simply draAvn upwards by their contraction, but it is made to describe a spiral movement, from right to left, and from behind for- wards ; and it is in this manner, that it is caused to strike against the side of the chest. 570. The systole of the Ventricles is immediately followed by their diastole; but the commencement of this has been observed to occur at a small interval previous to the contraction of the Auricles; and some- times a brief interval of repose may be noticed, separating the first stage of the Ventricular diastole, which may be partly due to the simple elas- ticity of the walls of the Ventricles, from the second, which is accompa- nied by the systole of the Auricles, and in which the blood of the latter is forcibly propelled into them. When the circulation is being carried on regularly, the blood is propelled into the Ventricles with sufficient force to dilate them strongly; so that the hand closed upon the heart is opened with violence. Even the auricles dilate with more force than it seems easy to account for by the vis a tergo of the blood in the venous 21 322 CIRCULATION OF NUTRITIVE FLUID. system; which is small compared with that which the fluid possesses in the arteries. 571. The natural movements of the Heart are accompanied by cer- tain sounds, which are heard when the ear is applied over the cardiac region; and an acquaintance with these sounds and with their causes is of much importance, since the alterations which they undergo in dis- ease, afford us some of our most accurate information in regard to the nature of the morbid affection. Concurrently with the impulse of the heart against the chest, a dull and prolonged sound is heard; this, which is termed the first sound, marks the ventricular systole, and is synchronous Avith the pulsation in the arteries. The second sound, which is short and sharp, folloAVS immediately upon the conclusion of the first; and it must therefore be produced during the first stage of the Ventricular diastole, before the systole of the Auricles has com- menced. It is followed by a brief interval of repose, which occurs during the remainder of the Ventricular diastole and the Auricular systole; and this is succeeded by a recurrence of the first sound. If the whole period between two successive pulsations be divided into four parts, it is estimated that the first sound usually occupies two of these; and the second sound, and the interval, one part each. 572. Now in order to understand the causes of these sounds, it is necessary to sfudy the course of the blood through the heart a little more in detail. When the Ventricles, distended with blood, are con- tracting upon their contents, they eject them forcibly through the narrow orifice of the aorta and pulmonary artery ; and the semilunar valves, which guard these orifices, are thrown back against the walls of the arteries. The regurgitation of the blood into the auricles is prevented by the action of the mitral and tricuspid valves ; but the flaps of these do not suddenly fall against each other, when the blood first begins to press them together; being restrained by the chordae tendinece. The connexion of these Avith the carnear columnar, which form part of the ventricular walls, and contract simultaneously with them, appears to have this use,—that the flaps of the valves, which are completely thrown back during the preceding rush of blood from the auricles to the ven- tricles, may be drawn into a favorable position for the blood to get behind them and bring them together, so as completely to close the orifice. As soon as the ventricular diastole begins to take place (even before the contraction of the auricles has commenced), there will be a tendency of the blood, that has just been propelled into the aorta and pulmonary artery, to flow back to the heart; but this regurgitation is completely prevented by the semilunar valves of these orifices, which are immediately filled-out by the backAvard tendency of the blood, and which meet in such a manner as completely to close the orifices. This closure is much more sudden than that of the mitral and tricuspid valves, being altogether unrestrained. 573. The first sound is certainly in part due to the impulse of the heart against the thoracic parietes; as is prOved by the fact, that when the impulse is prevented, the sound is much diminished in intensity: also by the circumstance, that, when the ventricles contract with vigor, the greatest intensity of the sound is over the point of percussion. But SOUNDS OF THE HEART. 323 that it is not entirely due to this cause, is also sufficiently evident from tAvo circumstances;—its prolonged character, which could scarcely be given by a momentary impulse;—and its continuance, though with diminished intensity, Avhen the parietes of the chest are wanting, and even after the complete removal of the heart from the body. Moreover, the duration of the first sound is much increased by any morbid state of the orifices of the ventricles, which obstructs the exit of the blood. Much discussion has taken place as to the cause of that part of it, Avhich is not due to the impulse; some having attributed it to the muscular contraction of the walls of the ventricles, others to the flow of blood over the irregular surfaces of their interior, and others to the rush of the fluid through the narrow orifices leading to the aorta and pulmonary artery. There can be little doubt, that the first and last of these causes are both concerned in producing the sound. For as a sound may be distinctly heard by means of the stethoscope, when the heart is con- tracting vigorously out of the body, and Avhen no blood is propelled by it, nothing else than muscular contraction can be then regarded as its source; and there is other evidence, that sound may be produced by this cause, since the vigorous contraction of any other large muscle gives rise to a continued tingling, which may be heard through the stethoscope. But when the heart is contracting in its natural position, and is propelling the blood with its ordinary vigor, the sound is heard in its greatest in- tensity at the base of the heart, i. e., at the origin of the great arteries; and since any obstruction to the exit of the blood through them increases the intensity as well as the length of the sound, it can scarcely be doubted that it is partly due to the rush of the blood through the con- tracted entrances of these vessels. A very similar sound, known as the "bruit de soufllet" or bellows-sound, may be heard through the stetho- scope, over any large artery, when it is compressed, so as to permit the passage of blood less readily than usual. Thus the ordinary first sound maybe regarded as composite in its nature; being made up of the sound produced by the impulse of the heart against the parietes of the chest, of the muscular sound occasioned by the forcible contraction of the thick walls of the ventricles, and of the sound generated by the friction of the particles of blood against each other, and against the boundaries of the narrowing orifices which lead into the vessels. 574. The cause of the second sound is simpler, and more easily un- derstood. It is due to the sudden filling-out of the semilunar valves Avith blood, at the moment when the ventricular systole has ceased, and Avhen the commencing diastole produces a tendency to the regurgitation of blood from the aorta and pulmonary artery. The sudden passage of the valves, from a state of complete relaxation to one of complete ten- sion, occasions a sort of click ; A\Thich is the second sound of the heart. That this is the real cause, has noAv been fully demonstrated. If one of the valves be hooked back against the side of the artery, by the in- troduction of a curved needle, so that a reflux of blood is permitted, the sound is entirely suppressed. And if the complete closure of the valves be preArented by disease, so that their tension is diminished, and a cer- tain amount of regurgitation takes place, the second sound is no longer heard in its proper intensity: Avhilst, on the other hand, a sound analo- 324 CIRCULATION OF NUTRITIVE FLUID. gous to the first, and sometimes prolonged over the whole interval of repose, indicates the reflux of the blood into the ventricles. When the semilunar valves are thickened by a morbid deposit, their surface rough- ened, and their opening narrowed, the first sound becomes harsher and sharper; and the second sound acquires the same character,—the back- ward as well as the forward flow of the blood being affected by this cause. 575. The natural movements of the mitral and tricuspid valves, appear to be accomplished with perfect freedom from sound; for the size of the orifices which they guard prevents any considerable friction of the blood, in its flow from one cavity to the other ; and their closure when the ventricular systole begins, does not take place Avith the rapidity and suddenness of that of the semilunar valves. But when their structure is changed by disease, their action is not so noiseless; and they give rise to various morbid sounds, Avhich are heard in addi- tion to the ordinary sounds, and wThich may even obscure them alto- gether. In the same manner, the ordinary movements of the heart do not produce any audible friction-sound, between the two surfaces of the pericardium, that which covers the heart, and that Avhich lines the pericardial sac. These surfaces are kept moist, in health, by the serous fluid constantly exhaling from them; and they are extremely smooth, so that they glide over one another noiselessly. But if they become dry, as in the first stage of inflammation, a slight creaking is heard, accompanying both the ordinary sounds of the heart, and somewhat resembling the rustling of paper. And if they are roughened by the deposit of inflammatory exudations, this "to and fro" sound becomes of a harsher character. 576. The walls of the left ventricle are considerably thicker than those of the right; and the contractile power is greater. This difference is obviously required, by the difference in length between the systemic and the pulmonary vessels ; the amount of force* necessary to drive the blood through the latter, being far inferior to that which is requisite to propel it through the former. The average thickness of the walls of the left Ventricle is about 4J lines ; being somewhat greater than this at the middle of the heart, and less at its apex. The average thickness of the walls of the right ventricle is not more than 1J line ; being a little greater than this at the base, and less at the apex of the heart. The left auricle is somewhat thicker than the right. The capacities of all the four cavities are nearly equal; each of them, in the full-sized heart, holding about tAvo ounces, of fluid. The Ventricles are, perhaps, a little larger than their respective Auricles ; but there is no very positive difference in capacity, between the Ventricles and Auricles of the two sides. 577. The quantity of blood which is propelled at each Ventricular systole, cannot, therefore, exceed two ounces ; and it is probably some- Avhat less, as the ventricles do not seem to empty themselves completely at each contraction. Now the whole quantity of the blood seems to be about one-fifth of the entire weight of the body; so that it will amount to about 28 lbs. in an individual of 140 lbs. weight. Allowing 75 pul- sations to a minute, 150 oz. (or 9 lbs. 6 oz.) of blood would pass through CIRCUMSTANCES AFFECTING RATE OF PULSE. 325 each ventricle of the heart in that time; consequently nearly three minutes would be required for the passage of the entire mass of the blood through the whole circle of its movement, if its rate be entirely determined by the impulses it receives from this central organ. But it appears, from various experiments, that the rate of circulation is much more rapid than this. For if a solution of any salt, easily detectible in the blood be injected into one of the large veins near the heart, it may be traced in the arterial circulation in from 15 to 20 seconds after- wards ; during which interval it must have traversed the whole pul- monary system of vessels, and passed through both sides of the heart. And if the salt be one, which acts poAverfully on the heart itself,—as is the case with Nitrate of Baryta or Nitrate of Potass,—this action is manifested almost at the same moment with the appearance of the salt in the arteries of other parts; thus showing that it has been conveyed by the coronary arteries into the capillaries of the heart itself. The period required for the transmission of a saline substance from the veins of the upper part of the body to those of the lower,—which can scarcely be accomplished through any more direct channel than the current that returns to the heart, then passes through the lungs back to the heart again, and then flows through the systemic arteries and capillaries to the veins,—is accomplished in little more than 20 seconds, even in an animal so large as a Horse. It appears, then, that even the vigorous and constant action of the Heart is not alone sufficient to maintain the circulation at its ordinary rate; and we are not justified, therefore, in excluding those sources of movement in the higher animals, which evidently exert so important an influence in the loAver. 578. The force with which the heart propels the blood is such, that if a vertical pipe be inserted into the Carotid artery of a horse the blood> will sometimes rise in it to a height of 10 feet. From com- parative experiments upon other animals, it has been estimated that the vigorous action of the heart in Man would sustain a column of blood in his aorta about 7£ feet high; or, in other words, that the force Avith which the heart ordinarily propels the blood through the aorta, is equal to that which would be generated by the weight of a column of blood of the same size, and 7* feet high; which weight would be about 4J lbs. But the force which must be exerted by the heart to sustain such a column, may be shown, upon physical principles, to be as much greater than this, as the area of a plane passing through the base and apex of the left ventricle is greater than that of the transverse section of the aorta ; and as the proportion of these areae is about 3 : 1, the real force of the heart may be stated at about 13 lbs. _ 579. The number of contractions of the heart, in a given time, is liable to great variations within the limits of health, from several causes ; the chief of which are diversities of Age and Sex, amount of Muscular exertion, the condition of the Mind, the state of the Digestive system, and the period of the Day. The following are the points of greatest importance, in regard to the action of these several influences. 326 CIRCULATION OF NUTRITIVE FLUID. Age.—The pulse of the newly-born infant averages from 130 to 140 per minute ; and this rate gradually diminishes, until, in adult age, the pulse averages from 70 to 80; and in the decline of life from 50 to 65. Sex.—The pulse of the adult female exceeds that of the adult male in frequency, by about 10 or 12 beats in a minute; and it is also more liable to disturbance from other causes. Muscular Exertion.—The effect of this in accelerating the pulse is well known; but as the amount of change depends upon the degree of exertion, no general statement can be made on the subject. The con- tinued influence of a moderate degree of muscular exertion, is shown by the effect of posture upon the pulse. Thus the pulse is on the average from 7 to 10 beats faster (per minute) in the standing than in the sitting posture; and 4 or 5 beats faster in the sitting than in the recumbent posture. This amount of variation is temporarily increased by the muscular effort required for the change of posture; but this soon sub- sides into the continued rate, which the permanent maintenance of the new posture involves. There are certain states of the system, in which the heart's action is increased to a most violent degree, by a simple change of posture; and in which, therefore, it is necessary that even this slight movement should be made with gentleness and caution. Mental Condition.—The action of the heart is peculiarly influenced, as every one is aware, by the excitement of the emotions. This is a fact to Avhich, however familiar, the medical practitioner should con- stantly direct his attention. The trifling agitation occasioned by the entrance of the medical man will produce, in many patients, such an acceleration of the pulse, as would be very alarming, if its true cause were not known. And the real rate of the pulse cannot be ascertained, until time has been permitted for the agitation to subside; which is favored, also, by the influence of a gentle manner and tranquillizing conversation. The operation of the intellectual powers does not seem to affect the rate of the heart's movement in any other way, than by inducing a general state of feverishness, if it be too long or too ener- getically kept up. State of the Digestive System.—The pulse is quickened during the digestion of a meal; but no exact numeral statement can be made on this subject. Period of the Day.—The frequency of the pulse appears to be some- what greater in the morning than it is in the evening; and the tem- porary action of any of the preceding causes, more quickly subsides in the evening than in the morning. 580. The movements of the heart have rjeen supposed to depend upon a constant supply of nervous influence, generated by the cerebro- spinal system, and transmitted through the sympathetic nerve, the branches of which are copiously distributed to it. And this idea seemed to derive support from the fact, that, when the brain and spinal cord are removed, or when large portions of them are suddenly destroyed, by crushing or by the breaking-up of their substance in any other mode, the movements of the heart are arrested. But it has been shown that the brain and spinal cord may be gradually removed, without any such consequence; and the occasional production of foetuses EQUALIZATION OF FLOAV IN THE ARTERIES. 327 destitute of those centres, but possessing a regularly-pulsating heart, is another proof that .the movements of this organ do not depend upon a supply of nervous influence derived from them. Still they are capable of being influenced by impressions transmitted through the nerves. It has been ascertained by Valentin, that, after the heart has ceased to beat, its contractions may be re-excited by stimulating the roots of the Spinal Accessory nerve, or of the first four Cervical nerves; the influence of that stimulation being conveyed to the heart by the Sympathetic system, the cardiac portion of which communicates with these nerves. Irritation of the Par Vagum, also, has a tendency to accelerate the heart's action, or to re-excite it when it has ceased; but the complete severance of both its trunks produces little disturb- ance in the regularity of the movement. The action of the heart may be also affected more directly through the Sympathetic system; thus it is excited by irritation of the cervical ganglia, especially the first; whilst continued pressure upon the cardiac nerve, by an enlarged bronchial gland, has appeared to be the cause of its occasional suspen- sion. It is without doubt through its nervous connexions, and probably through the sympathetic system, that the heart receives the influence of mental emotions. 581. The movements of the heart may be suspended, or altogether checked, by sudden and violent impressions on the nervous centres, even though these do not occasion any perceptible breach of substance. Thus in concussion of the brain, there is not merely insensibility, but also a complete suspension of the circulation, occasioned by a failure of the heart's power. This suspension may be permanent, so that ani- mation cannot be restored; or it may be temporary, as in ordinary fainting. The well-knoAvn influence of blows upon the epigastrium, in producing sudden death, is probably to be attributed to a similar cause, —namely, the shock thus communicated to the extensive plexus of gan- glionic nerves, radiating from the semilunar ganglia, and proceeding to the abdominal vriscera. Violent impressions upon other nervous expan- sions may produce a dangerous weakening of the heart's contractile power; this is the case, for example, with extensive burns, which may produce faintness, and even death, especially in children, by the depres- sion Avhich they induce. Many other causes of sudden suspension of the heart's action might be enumerated; but they may be generally traced to a strong impression upon the nervous system; though of the mode in which this operates Ave know nothing. 4. Movement of the Blood in the Arteries. 582. The Blood, thus propelled from the Heart into the Arteries by a series of interrupted jets, Avould continue to flow in the same manner, if it were not for the equalization of its movement, effected by the pro- perties of the arterial walls. This influence is exerted by the middle or fibrous coat, which consists in part of yellow elastic tissue (§ 189), and in part of non-striated muscular fibre (§ 337). The proportion of these two components varies in arteries of different calibre; the muscular 328 CIRCULATION OF NUTRITIVE FLUID. tissue being thicker in the smaller branches, and the elastic tissue being found in larger amount in the main trunks. 583. It is chiefly to the simple physical property of Elasticity, thus possessed by the Arterial tubes, that Ave owe the equalization of the flow of blood; and we may hence understand the reason Avhy the trunks that; are in nearest connexion with the heart, should be those most endowed with it. If a forcing-pump were to inject water, by successive strokes, into a system of tubes with perfectly unyielding walls, the flow of fluid at the further extremities of these tubes would be as much interrupted as its entrance into them. But if the pump be connected with an air-vessel (as in the common fire engine), so that a part of the force of each stroke is expended in compressing the air, the expansion of this, during the interval betAveen the successive strokes, produces a continuous flow of water along the tubes. Or if the tubes themselves were endowed with a certain degree of elasticity, which should allow them to dilate near their commencement, so as to receive the new charge of fluid, and Avhich should occasion a continued pressure upon the fluid during the interval of the stroke, the same equalizing effect Avould be produced. This is precisely the case with the arterial system; the intermittent jets, by which the blood is propelled from the heart, are speedily converted into a continued stream ; so that, at even a mode- rate distance from the heart, the only indication of its interrupted action is presented by the greater or less rapidity of the flow; and this gives rise, when an artery is divided, to an alternate rise and fall of the jet of blood, and, in the ordinary circulation, to the phenomenon called the pulse. This is due to an increase in the dimension of the arterial tube, both in length and breadth, with each additional ingress of blood; the increase in length is the more considerable of the two effects, and causes the artery to be somewhat lifted from its seat. During the intervals, a quantity of blood corresponding to that which had entered, escapes by the further extremity of the tube; and thus the artery is enabled to contract to its previous dimensions, and to return to its bed. We may compare the pulse, therefore, to a wave, which commences in the heart, and travels onwards through the arterial system. 584. In the large arteries near the heart, the pulsation is always precisely synchronous with the ventricular systole; but it takes place somewhat later in the arteries at a distance from the heart; the time required for the transmission of the wave being proportioned to the degree in which the walls of the arteries yield to it. If they were quite rigid, the egress at one extremity must take place at the precise moment, that the fluid is forced into the other. On the other hand, if the walls be too easily distensible, they yield to the propelling force in such a degree that it is entirely expended upon them; and the fluid is not moved onward at all or but very sloAvly. In the healthy state of the arterial walls, they should contract upon their contents Avith sufficient force to equalize the flow of blood, and to prevent the pulse-Avave from occupying more than one-sixth or one-seventh of a second, in its propa- gation to the remotest arteries of the system; and the pulse should be full, producing a prolonged but gentle elevation beneath the finger, and capable of resisting moderate pressure. This condition is dependent EFFECTS OF MUSCULARITY OF ARTERIES. 329 in great part upon the due tonicity of the muscular coat of the arteries (§ 365). When this tonicity is in excess, the walls of the arteries are too rigid ; the pulse at the wrist is felt to occur exactly at the same time Avith the ventricular systole; and its character is that of strength, incom possibility, and sustained poAver, though it may be even sloAver than usual. This is the case in Avhat is commonly termed " high corfdi- tion" of the system ; which predisposes to inflammatory disorders, but which renders it less susceptible than usual to the influence of mala- ria, contagious miasmata, or other causes of a depressing character. On the other hand, when the tonicity of the arteries is less than it should be, their walls yield too much to the pulse-wave ; so that the pulse at the wrist is often felt even after the second sound is heard ; and the pulse itself is jerking, unsteady, and too easily compressible. This loose relaxed state of the vessels is the most unfavorable that can be to regularity and vigor of the circulation ; and it manifests its ill effects in the general condition of the system, which is then peculiarly prone to suffer from the agency of malaria, infectious miasmata, or any other depressing causes. 585. Although many Physiologists have denied that the Arteries possess real Muscular contractility in any degree, yet there can be no longer any doubt on the subject; since numerous experimenters have succeeded in producing distinct contraction in their walls, by the appli- cation of those stimuli, Avhich act upon muscular fibre in general. More- over it has been ascertained, that when an artery is dilated by the blood injected into it from the heart, it reacts with a force superior to the impulse to Avhich it yielded ; and that, if a portion of an artery from an animal recently dead, in which the vital properties are still preserved, and a similar portion from an animal that has been dead some days, in Avhich nothing but the elasticity remains, be distended Avith equal force, the former contracts to a much greater degree than the latter after the distending force is withdraAvn.—One use of this contractile poAver may very probably be, to assist the Heart in maintaining the flow of blood; for if the arterial walls yield rapidly to the ingress of blood, and then, contract upon their contents with a force greater than that which dis- tended them, the current must necessarily be propelled onwards with greater force. This supplementary propelling force, on the part of the arteries, may serve as a compensation to that diminution of the heart's power which must result from the increased friction of the blood against the Avails of the vessels occasioned by their subdivision ; and we thus observe even in the highest animals, some traces of that diffused agency on Avhich the Circulation is so much more dependent in the loAver tribes.. 586. It seems probable, however, that one chief use of the Muscu-_ larity oT The Arterial walls, consists in its regulation of lhe_diameter ofTnVfubes, In liceoToTance with the quantity ofMoj>pMo_l^ conducted through^henTTcT any party the proper ainouTiFT^emg determined by c^uTnltancesattlTeTrrire:—Such local changes may form a part of the regular series of actions of the human body, as Avhen the Uterine and Mammary arteries undergo enlargement, at the periods of pregnancy and parturition ; and they occur still more frequently in diseases, which are attended by increased action of particular organs. In such cases, 330 CIRCULATION OF NUTRITIVE FLUID. it cannot be vis a tergo of the Heart, that occasions the enlargement of certain arterial trunks, and of no others; since any increase in its pro- pulsive power would affect all alike. It must be, therefore, through a power inherent in themselves, that the dilatation takes place ; and there seems much reason for attributing to the Sympathetic system of nerves a control 0Arer this poAver, and consequently the office of regu- lating the local distribution of blood, in accordance Avith the Avants of the different parts. It is well known that the nerves of this system are copiously distributed upon the arterial Avails ; and it has been experi- mentally shown, that they have the power of producing contractions in the larger arteries. Moreover, there is every reason to believe that the diameter of the Capillary blood-vessels, and the rate of the move- ment of the blood through them is much influenced by these nerves (§ 603); and it seems highly probable, therefore, that they should have a corresponding influence upon the size of the trunks, from Avhich these capillaries are derived. 587. The arterial system possesses nearly the same relative capacity in every part; that is, if a section could be made through all the sys- temic arteries at a certain distance from the heart, the united areas would be found equal to that of the aorta ; and those of the branches of the pulmonary arteries Avould equal those of their trunk. This re- sults from the fact, that, at every subdivision, the united areas of the branches are almost precisely equal to that of the trunks from which they proceed ; although the united diameters of the former far exceed that of the latter. According to a well-known mathematical laAv, the areas of circles are as the squares of the diameters ; consequently, in making such comparisons, it is necessary to square the diameters of the trunk and those of the branches, and to contrast the former with the sum of the latter. Thus a trunk Avhose diameter is 7, may subdivide into two branches, each having a diameter of nearly 5 ; for the square of 7 is 49, and twice the square of 5 is 50. Or a trunk whose diameter is 17 may subdivide into three branches whose diameters are 10, 10, and 9 J (making 29J as the sum of the diameters); for the square of the diameter of the trunk is 289, wdiile the sum of the squares ef those of the branches is 290J. It appears, hoAvever, from Mr. Paget's recent admeasurements, that there is seldom an exact equality betAveen the area of the trunk and that of its branches; the area sometimes increas- ing, and sometimes diminishing. The former seems the general rule in the upper extremities; the latter in the loAver. Thus the area of the trunk of the external carotid is to that of its branches, as 100 to 119; Avhilst the area of the abdominal aorta, just before its final division, is to that of its branches as 100 to 89. 588. In almost every part of their course, the ramifications of the arteries communicate freely with each other, by anastomosis ; and this communication is most important, as affording the means by which the circulation is sustained, Avhen the current through the main trunk is ob- structed. There is scarcely an artery in the body except the aorta, which may not be tied, with the certainty that the blood will still be conducted to its destination, by the collateral circulation. At first, the quantity which thus passes is very insignificant, and is by no means ARRANGEMENT OF CAPILLARY VESSELS. 331 sufficient to supply what is needed ; thus, when the femoral artery has been tied for popliteal aneurism, the limb becomes cold, and the sensi- bility of the surface and its muscular power are alike diminished. In a few hours, hoAvever, its warmth returns, and its sensibility and muscular poAver are restored ; indicating that its circulation has been already re- established through the collateral branches. And Avhere an opportunity presents itself at a subsequent period for examining the state of the vessels in such a limb, it is found that an extraordinary enlargement has taken place in arteries that were previously of insignificant size, which form a communication betAveen the branches that issue above and below the interruption. Moreover, it is commonly found that the main trunk has become completely impervious above the part where it Avas obliterated by the ligature, up to the point at which the nearest lateral branch is given off.—Even the abdominal aorta has been tied in dogs, without fatal results ; the circulation in the posterior part of the body, and in the hinder extremities, being then maintained chiefly by the inosculation of the external mammary artery with the epigastric, upon the parietes of the abdomen. 5. Movement of Blood in the Capillaries. 589. The ultimate ramifications of the arteries pass so insensibly into those of the Veins, that no definite line of demarcation between them can be draAvn; and although we are in the habit of speaking of the " Capillaries" as a distinct systeni of vessels, yet it ought to be strictly borne in mind, that they differ only in size from the vessels from which they receive their blood on the one side, and into which they pour it on the other. It was at one time supposed, that they Avere merely channels or passages, excavated in the tissues, having no definite walls of their OAvn. This is probably true of them in the lower tribes of Animals; and it may also be the case at an early stage of their development in the higher. But Avhen their formation is complete, they undoubtedly possess walls of a fibrous texture, as distinct as those of the arteries and veins, though of extreme thinness. From the occa- sional appearance of bodies resembling cell-nuclei, in the substance of the Avails of the capillaries, it has been thought that their tubes are formed, in the first instance, by the coalescence of cells arranged in a linear direction ; and this idea receives confirmation from the fact, that the ducts of Plants are undoubtedly formed in this manner, and not by the mere retirement of the tissues on either side, leaving an intervening channel. The closely-reticulated structure usually formed by the capil- laries, has been commonly regarded as distinguishing them both from the arteries and the veins; and it is not unusual to speak of the arteries as delivering the blood into the " capillary netAvork," and of the veins as receiving the fluid that has traversed this. Such expressions are not incorrect as implying the simple fact, that betAveen the arteries and the veins is a netAvork of minute vessels, through Avhich the blood has to travel Avhen proceeding from one to the other; but these vessels must not be regarded as belonging to a distinct class, being nothing 332 CIRCULATION OF NUTRITIVE FLUID. else than the minutest subdivisions of the veins and arteries, which commonly inosculate freely with each other. 590. The degree of this inosculation, and the consequent form of the capillary netAvork, are subject, however, to very great variations ; and these may be generally shown to have a relation to the form of the ultimate elements of the tissues, Avhich are traversed by the capillaries. Thus we have seen in the capillaries of Muscle, that the major part run parallel to the course of the fibres, lying in the minute interspaces be- tAveen them (Fig. 67); a few transverse branches serving to connect them with each other. A similar distribution prevails in the capillaries of the Nervous trunks ; but those of the Nervous centres are arranged in the form of a minute network, so as completely to traverse every part of the structure (Fig. 93). Again, we observe that the capillaries of Glands form a minute network around the secreting follicle (Fig. 94); and a similar arrangement prevails in the capillaries of the air- cells of the lungs, which are set so closely together, that it would seem Fig. 93. Fig. 94. as if the purpose were to cover the surface with blood as completely as possible, consistently with its being retained Avithin vessels, and not spread out into a continuous film (Fig. 106). A network of very much the same character is found in the villi of the mucous membrane (Fig. Fig. 95. Fig. 96. , Capillary Network in simple mucous mem- brane of palpebral conjunctiva. 82), on the ordinary surface of simple mucous membrane (Fig. 95), and on that of the choroid coat of the eye (Fig. 96). Where the surface of the mucous membrane is depressed into follicles, the arrangement of the DISTRIBUTION OF THE CAPILLARIES. 333 capillaries has an evident reference to these (Fig. 97) ; whilst, on the other hand, Avhere the surface of the skin is raised up into sensory papillse, the capillary network sends looped prolongations into them which are found accompanying their nerves (Figs. 98 and 99). 591. It cannot be supposed that the arrangement of the vessels has any further influence upon the function of the part they traverse, than Fig. 97. Fig. 98. that which it derives from the regulation of the supply of blood afforded to each indiAridual portion of the structure. The form of the capillary Fig. 99. Capillary Network of fungiform papilla of the tongue. network is evidently determined by that of the elements of the tissues permeated by it; these are the real operative instruments in every part; and the distribution of the blood-vessels is so arranged, as to afford them precisely the amount of nourishment they respectively require. Thus Ave have seen, that there are many living parts, possessing most important functions, in the human body, which are not in any direct relation Avith blood-vessels, and which yet derive their whole nutriment, and the materials of their functional operations, from the blood. This is the case, for example, Avith the whole of the epithelial and epidermic cells; and also with the articular cartilages, and the substance of the teeth. Even in bone, the islets between the Haversian canals, Avhich are completely unpenetrated by vessels, are of considerable size. Such islets must everywhere exist, between the meshes of the capillary net- Avork ; so that the question of the vascularity or non-vascularity of a tissue is one of degree only;—the ultimate fibre of muscle or nerve, and the cells and fibres of other tissues, being as completely non-vascular, as the entire substance of a tooth or of an articular cartilage ; the latter being nourished, like the former, by imbibition from the surrounding vessels. 592. The term " Capillary" may be employed in an extended or a 334 CIRCULATION OF NUTRITIVE FLUID. restricted sense; in the former it includes all the minute vessels which pass betAveen the arteries and the veins; in the latter it is applied only to those which admit no more than a single file of blood-discs at once, and excludes those Avhich admit two, three, or even four rows, even although they establish a direct communication from one side of the network to the other. The former application of the term is the most convenient, although perhaps not the most strictly accurate; and it will be therefore here employed in its extended sense. And this is rendered more correct by the fact, that the size of the individual capillaries is by no means permanent; an enlargement often taking place in one, and a contraction in another, at the same time: so that vessels, Avhich were previously true capillaries, no longer remain such; and passages, which were previously of far greater calibre, are reduced to the average diameter. 593. The opinion was long entertained, that there are vessels adapted to the supply of the Avhite or colorless tissues ; carrying from the arteries only the fluid portion of the blood, or liquor sanguinis, and leaving the rest behind. No other such vessels have been really ob- served, however, than the capillaries in a state of unusual contraction, as just now mentioned. And it may be safely affirmed, that the suppo- sition of their existence is not required. For any one who observes the smallest capillary vessels under the microscope, may perceive, that the current of blood Avhich passes through them is almost free from color,—as the red corpuscles themselves appear to be, when spread out in a single layer. Tissues which are rather scantily permeated by such vessels, therefore, may still be white; and it is only where the netAvork is very close, and the quantity of blood Avhich passes through it is consequently great, that a perceptible color will be communicated by the red corpuscles. And we have seen, that the idea that Nutrition can only be carried on by direct communication with vessels, is entirely unfounded; the tissues into Avhich no bloodvessels can be traced, being adapted to nourish themselves, like cellular Plants, by the imbibition of fluid at their surfaces, on Avhich vessels are (for the most part) copiously distributed. 594. That the blood can only minister to the operations of Nutrition, Secretion, &c, whilst it is circulating through the Capillaries, is eAddent from several considerations. The thickness of the walls of the larger vessels interposes an effectual barrier to its transudation ; and so com- pletely is the blood cut off even from penetrating these, that they do not derive their own nourishment from the blood which Aoavs through their tubes, but from a capillary netAvork in their own substance, which is supplied by vessels from collateral branches,—these being termed the vasa vasorum. MoreoA'er it is by the inosculation of the capillaries alone, that the minute network is formed, Avhich serves to bring the blood into proximity with the minute parts of the tissues to be nourished; thus let it be supposed that the minute arteries of Muscle were to ter- minate in veins, without undergoing further subdivision, the islets left betAveen their anastomosing branches would be far too large, and the nutritive materials Avould consequently not be supplied with sufficient readiness, eA'en supposing that it could freely permeate the Avails of VARYING SIZE OF THE CAPILLARIES. 335 these vessels.—The capillaries, then, must not be regarded as altogether distinct in their endowments, from the vessels with which they are con- nected on either side; but merely as intended, by their minute sub- division and inosculation, to bring the blood into sufficiently close rela- tion with the tissues they are to nourish, and to allow a greater degree of transudation of its elements by the comparative thinness of their Avails. 595. When the Aoav of blood through the Capillaries of a trans- parent part, such as the web of a Frog's foot, is observed with the microscope, it appears at first to take place with great evenness and regularity. The influence of the contractions of the heart may be seen to extend itself into the smaller arteries ; the blood moving onwards in them with a somewhat jerking motion. But this influence alto- gether disappears in the capillary network; the flow of blood through this being even and continuous, except when the action of the heart is becoming Aveak and irregular, or when its influence is impeded by obstruction in the vessels leading to the part,—the blood being then impelled by a succession of jerks, with intervals of complete repose. But on watching the movement for some time, various changes may be observed, which cannot be attributed to the heart's influence, and Avhich shoAV that a certain regulating or distributive power exists in the Avails of the capillaries, or in the tissues which they traverse. Not only do Ave occasionally perceive some of the tubes enlarging, so as to admit several files of blood-discs instead of one, whilst others that previously received several now only admit one;—but we also see vessels coming into vieAV, which were not previously noticed, whilst other vessels seem to become obliterated. This apparently new forma- tion and obliteration of vessels, hoAvever, does not really take place; for a "more close examination shows, that the former of these appear- ances is due to the entrance of red corpuscles into passages which existed before, but which were in such a state of contraction as enabled them only to admit the fluid portion of the blood; whilst, by a con- verse change in certain capillaries, from the dilated to the contracted state, the appearance of obliteration is produced, the red corpuscles being excluded, and the transparent fluid of the blood being alone transmitted by them. 596. But these are by no means all the irregularities which may be detected by a close scrutiny of the Capillary circulation. The velocity of the current is liable to great and sudden variations, which cannot be accounted for by any change in the heart's action, or in the supply of blood afforded by the arteries; and this change may manifest itself, either in the Avhole capillary network of a part, or in a portion of it; the circulation taking place with diminished rapidity in one part, and with increased energy in another, though both are supplied by the same trunk. These variations are sometimes manifested by the complete change in the direction of the movement, in certain of the transverse or communicating branches ; this movement taking place, of course, from the stronger toAvards the weaker current. Not unfrequently an entire stagnation, of longer or shorter duration, precedes the reversal of the direction. Irregularities of this kind are most frequent 336 CIRCULATION OF NUTRITIVE FLUID. when the heart's action is enfeebled or partially interrupted ; and it Avould thus appear, that the local influences by Avhich they are produced are overcome by the propelling power of the central organ, when this is acting Avith its full vigor. When the Avhole current has nearly stag- nated, and a fresh impulse from the heart renews it, the movement is seldom uniform through the entire plexus supplied by one trunk ; but is much greater in some of the tubes than in others,—the variation being in no degree connected with their size, and being very different in its amount at short intervals. 597. All these circumstances indicate that the movement of blood through the capillaries is very much influenced by local forces ; although these forces are not sufficiently poAverful, in the higher animals, to maintain it alone. And from other facts it appears, that the condi- tions necessary for the energetic Aoav of blood through these vessels, are nothing else than the active performance of the nutritive and other operations, to Avhich they are subservient. The examination of a single one of these processes, Avill afford us the requisite proof. The blood when circulating through the systemic capillaries, yields a portion of its oxygen to the tissues it permeates, and receives from them carbonic acid. On the other hand, when passing through the pulmonary capil- laries, it gives up its carbonic acid to the atmosphere, and imbibes a fresh supply of oxygen. Noav if either of these changes be prevented from taking place, a retardation and even a complete stagnation of the blood will take place,—the flow through the capillaries being noAV resisted, instead of accelerated, by the relation which the blood bears to the tissues. Thus it has been shoAvn, that if an animal be partially deprived of oxygen, so that the arterial blood is not duly aerated (rather resembling the ordinary venous blood), and cannot exert its proper action on the tissues, the pressure upon the walls of the systemic arteries is increased, although the supply of blood propelled by the heart, and the propulsive poAver of the heart itself are diminished; and this plainly indicates a retardation in the systemic capillaries, producing an undue accumulation in the arteries. On the other hand, the sus- pension of the supply of oxygen to the lungs, either by an obstruction in the air passages, or by the substitution of some other gas, brings the pulmonary circulation to a stand in a very short time, the blood not being able to undergo its usual changes in the capillaries of those organs; and by this stagnation, the whole movement of blood is speedily checked. The readmission of oxygen, if the suspension of the circulation have not been too long continued, occasions the renewal of the movement in the capillaries, and thence in the Avhole circle of A'essels; and this even after the heart has ceased to propel blood towards the lungs. 598. The principles already noticed (§ 547), as put forth by Prof. Draper, seem fully adequate to explain these phenomena. The arterial blood,—containing oxygen Avith which it is ready to part, and being prepared to receive in exchange the carbonic acid which the tissues set free,—must obviously have a greater affinity for the tissues, than venous blood, in Avhich both these changes have been already effected. Conse- quently, upon mere physical principles, the arterial blood, which enters the systemic capillaries on one side, must drive before it, and expel on MOVEMENTS OF BLOOD IN CAPILLARIES. 337 the other Bide of the network, the blood which has become venous whilst traversing it. But if the blood which enters the capillaries have no such affinity, no such motor power can be developed. On the other hand, in the capillaries of the lungs, the opposite affinities prevail. The venous blood and the air in the pulmonary cells have a mutual attrac- tion, which is satisfied by the exchange of oxygen and carbonic acid that takes place through the walls of the capillaries; and when the blood has become arterialized, it no longer has any attraction for the air. Upon the very same principle, therefore, the venous blood will drive the arterial before it in the pulmonary capillaries, whilst respira- tion is properly going on; but if the supply of oxygen be interrupted, so that the blood is no longer aerated, no change in the affinities takes place whilst it traverses the capillary network; the blood, continuing venous, still retains its need of a change, and its attraction for the walls of the capillaries; and its egress into the pulmonary veins is thus resisted, rather than aided, by the force generated in the lungs. 599. The change in the condition of the blood, in regard to the relative proportions of its oxygen and carbonic acid, is the only one to which the Pulmonary circulation is subservient; but in the Systemic circulation, the changes are of a much more complex nature,—every distinct organ attracting to itself the peculiar substances which it requires as the materials of its own nutrition, and the nature of the affinities thus generated being consequently different in each case. But the same law holds good in all instances. Thus the blood conveyed to the liver by the portal vein, contains the materials at the expense of which the bile-secreting cells are developed; consequently the tissue of the liver, which is principally made up of these cells, possesses a certain degree of affinity or attraction for blood containing these materials ; and this is diminished, so soon as they have been drawn from it into the cells around. Consequently the blood of the portal vein will drive before it, into the hepatic vein, the blood which has traversed the capillaries of the portal system, and which has given up, in doing so, the elements of bile to the solid tissues of the liver. The same principle holds good in every other case. 600. We are noAV prepared, therefore, to understand the general principle, that the rapidity of the circulation of a part will depend in great measure upon the activity of the functional changes taking place in it,—the heart's action, and the state of the general circulation, remaining the same. When, by the heightened vitality, or the unusual exercise, of a part, the changes which the blood naturally undergoes in it are increased in amount, the affinities which draw the arterial blood into the capillaries are stronger, and more speedily satisfied, and the venous blood is therefore driven out with increased energy. Thus a larger quantity of blood will pass through the capillaries of the part in a given time, without any enlargement of their calibre, and even though it be somewhat diminished; but the size of the arteries by which it is supplied soon undergoes an increase, which adapts it to supply the increased demand. Any circumstance, then, which increases the func- tional energy of a part, or stimulates it to increased nutrition, will occasion an increase in the supply of blood, altogether irrespectively 22 338 CIRCULATION OF NUTRITIVE FLUID. of any change in the heart's action. This principle has long been known, and has been expressed in the concise adage, " Ubi stimulus, ibi fluxus;" which those Physiologists Avho maintain that the Circu- lation is maintained and governed by the heart alone, cast into un- merited neglect. 601. An undue acceleration of the local circulation, arising from an excess of functional activity in the part, and unaccompanied by any other change, constitutes the state knoAvn as active congestion, or deter- mination of blood. This may be artificially produced by the applica- tion of gentle stimulants; and it is usually the first change that occurs, when their action proves sufficiently violent to produce Inflammation. From that state, however, it is distinguished by this important charac- ter,—that there is merely an exaltation of the natural function, but no change. Moreover we shall presently see that, in Inflammation, there is a stagnation of blood, not an acceleration. We frequently meet with cases, in which this active congestion becomes very manifest: especially in persons of active minds, who exert their mental poAvers too violently, and who thereby induce an habitually-increased flow of blood towards the head, manifested in the increased pulsation of the carotids, the suffusion of the face and eyes, and the heat of the surface. The balance of the circulation being thus disturbed, there is almost invariably a diminished energy of the movement of blood in other organs, especially the extremities; as indicated by their habitual cold- ness and lividity. In the treatment of such a state (which is often the precursor of serious disease), it should be our object to restore the circulation in the extremities, by friction, exercise, &c.; and to abate the flow of blood toAvards the head, by restraining the functional activity of the brain, by the application of cold to the surface, by keeping the head high during sleep, and other means of similar tendency. 602. There is another condition of the capillary circulation, also known under the name of Congestion, which is precisely the opposite of the preceding. In this state, there is deficient functional energy in the part, and the circulation through it is consequently retarded,—as in the lungs where there is a partial obstruction to the aeration of the blood. The same cause produces a deficient tonicity of the Arteries, and allows their walls to be unduly distended by the vis a tergo of the blood; and consequently there is a great accumulation of blood in the part, with a retarded movement. This condition, like the preceding, predisposes to Inflammation, although in a different mode, as will be explained hereafter (§ 631). It is relieved by causes which promote the action of the part; thus congestion of the lungs, occasioned by the effusion of fluid into the air-cells, Avhich creates an obstacle to the aera- tion of the blood, disappears when that effusion is absorbed. And congestion of the liver, the result of deficient secreting poAver in the organ, is relieved by mercurial and other medicines, which promote the flow of bile by stimulating the growth of the hepatic cells. 603. The Capillaries, like the Arteries, possess a power of contrac- tion and dilatation, which seems to be very much under the influence of the Nervous System, and particularly of that part of it which conveys the influence of the Emotions. We have a visible example of this in- MOVEMENT OF BLOOD IN THE VEINS. 339 fluence in the act of blushing, Avhich consists in a sudden enlargement of the capillaries and small vessels of the surface; whilst the converse state of pallor, Avhich often alternates with it under the influence of strong emotion, is evidently due to an unusual contraction of the same vessels. But the effects of this influence are no less sensible in other cases ; and particularly in the regulation of the quantity of certain secretions, in accordance with the mental state, or the condition of the system generally. To the mode in which this regulation is effected, the act of blushing seems to afford us the key; for it indicates that the supply of blood afforded to the glands, may be entirely governed by the influence of the nervous system upon the calibre of the arteries. Thus the nursing mother, at the sight, or even at the thought, of her child, Avhen the usual time for suckling approaches, feels a rush of blood to the breast, exactly resembling that which takes place to the cheeks in blushing, and popularly termed "the draught;" this rush occasions an almost immediate increase in the secretion. In like manner we may explain the influence of the mental state upon the amount of the secre- tions of the lachrymal, the salivary, and many other glands ; its influence upon their quality, must probably be effected through changes in the condition of the blood itself. 604. The supply of Nervous agency from the Cerebro-spinal system, has been clearly proved to exert no direct influence in maintaining the capillary circulation ; since the latter continues as usual, after all the nerves of a part have been divided. This is obviously due to the fact, that the operations of nutrition, secretion, &c, are essentially indepen- dent of this agency. But as they are in some degree influenced by it, so will the capillary circulation be affected, through its connexion with them. In this manner Ave are to explain the effect of violent impres- sions upon the nervous centres in bringing to a stand, not merely the action of the Heart (§ 581), but the Capillary circulation all over the body. The general vitality of the system appears to be at once de- stroyed ; so that the capillary circulation, which may usually be seen to continue in the web of a frog's foot for some time after the interruption of the heart's action, is immediately suspended by crushing the brain Avith a hammer. 6. Of the Movement of Blood in the Veins. 605. The Arenous system is formed by the reunion of the small trunks, Avhich originate in the capillary network ; and it carries back to the heart the blood Avhich has been transmitted through this. This blood is dark or carbonated in the systemic veins; whilst it is bright or oxy- genated in the pulmonary veins. The structure of the veins is essen- tially the same vvith that of the arteries; but the fibrous tissue of their middle coat less decidedly exhibits the characters, either of the yellow elastic tissue, or of non-striated muscle. Still it possesses no inconside- rable amount of Elasticity; and a certain degree of Muscular contrac- tility also. The Avhole capacity of the Venous system is at least twice and perhaps more nearly three times, that of the Arterial; and the rate of motion of the blood in it must be proportionably sloAver. 340 CIRCULATION OF NUTRITIVE FLUID. 606. The movement of the Blood through the Veins is, without doubt, chiefly effected by the vis a tergo, or propulsive force, which results from the contractile power of the heart and arteries, aided by the power generated in the capillary vessels. The intermittent flow, which is caused by the interrupted action of the former, is usually so far equalized during the passage of the blood through the capillary network, that no pulsation can be shoAvn to exist in the veins; but instances occasionally present themselves, in which a venous pulse may be clearly perceived. The Venous Circulation is affected, however, by certain other causes, which exert little influence on the movement of blood in the Arteries. One of these is the frequently recurring action of Muscles, to which the Veins are peculiarly subject, on account of their position. In every instance in which Muscular movement takes place, a portion of the Veins of the part will undergo compression; and as the blood is pre- vented by the valves in the veins, from being driven back into the small vessels, it is necessarily forced onwards towards the heart. As each set of muscles is relaxed, the veins that were compressed by it fill out again, to be again compressed on the renewal of the force. Thus we see how the general Muscular movements of the body have an important influence in maintaining the Venous Circulation,—how continued exer- cise, involving the alternate contraction and relaxation of several groups of muscles, must send the blood more rapidly towards the heart, and thus increase the rapidity of its pulsations,—and how the sudden and simultaneous action of a large number of muscles after repose (as when we rise up from the sitting or lying, to the standing posture), may drive the blood to the heart with so violent an impetus, as to produce even fatal results, if, by any diseased condition of that organ, it should be rendered unable to dispose Avith sufficient rapidity of the quantity of blood thus transmitted to it. 607. The Respiratory movements exert a slight influence upon the flow of blood through the large veins near the heart; for as the chest is a closed cavity, in Avhich a partial vacuum is produced by the act of In- spiration, whilst its contents are compressed by the act of Expiration, the former state will favor the movement of blood from the large veins on the exterior of the cavity, towards the heart, whilst the latter condi- tion will retard it. This produces the phenomenon termed the respira- tory pulse ; which may be seen in the veins of the neck and shoulders in thin persons, and especially in those who are suffering from pulmonary diseases. The influence of the Respiratory movements is made evident by introducing a tube into the Jugular vein, the lower end of which dips into water; for an alternate elevation and depression of the water into the tube is then witnessed, showing the suction poAver of the Inspi- ratory movement, and the expellent force of the Expiratory act. On the other hand,_ the Expiratory movement, while it directly tends to cause accumulation in the veins, will assist the heart in propelling the blood in the Arteries; and by the combined action of these two causes is produced, among other effects, the rising and sinking of the Brain, synchronously with expiration and inspiration, which are observed when a portion of the cranium is removed. 608. A pulsatory movement may be occasioned in the great veins RESPIRATORY PULSE—VENOUS PULSE. 341 near the heart, by another cause entirely distinct from the preceding; namely, the regurgitation of blood from the ventricle into the auricle, and thence into the venae cavae, during the ventricular systole ; and the pul- sation thus occasioned is synchronous, therefore, with that in the arteries (proceeding backwards, however, from the heart), instead of correspond- ing with the respiratory movement. This regurgitation may take place, not from any disease in the valves on the right side of the heart, but simply from over-distension of its cavities, resulting fr.om any obstruc- tion to the circulation of blood through the lungs; for Avhen this occurs, the tricuspid valve does not completely close, and allows a portion of the blood to escape from the ventricle backwards into the auricle and venae cavae. This want of complete closure, constituting what has been termed the "safety-valve function" of the tricuspid valve, has been par- ticularly noticed in diving animals, in which the circulation through the lungs is liable to be temporarily suspended. The venous pulsation which is thus produced, may be noticed in almost every case of long- standing dyspnoea; especially Avhen this is accompanied (as it usually is) by hypertrophy and dilatation of the right ventricle of the heart. 609. The Venous circulation is much more liable than the Arterial, to be influenced by the force of Gravity; and this influence is particu- larly noticeable, when the tonicity of the vessels is deficient. The fol- lowing experiments performed by Dr. Williams, to elucidate the influence of deficient firmness in the walls of the vessels, and of gravitation, over the movement of fluids through tubes, throw great light on the causes of Venous Congestion. A tube with two equal arms having been fitted to a syringe, a brass tube two feet long, having several right angles in its course, Avas adapted to one of them, whilst to the other was tied a portion of a rabbit's intestine four feet long, and of calibre double that of the brass tube, this being arranged in curves and coils, but without angles and crossings. When the two ends were raised to the same height, the small metal tube discharged from two to five times the quantity of water discharged in a given time by the larger but mem- branous tube; the difference being greatest, Avhen the strokes of the piston were most forcible and sudden, by which the intestine was much dilated at its syringe end, but conveyed very little more water. When the discharging ends were raised a feAV inches higher, the difference in- creased considerably, the amount of fluid discharged by the gut being much diminished ; and when the ends were raised to the height of eight or ten inches, the gut ceased to discharge, each stroke only moving the column of Avater in it, and this subsiding again, without rising high enough to overflow. When the force of the stroke was increased, the part of the intestine nearest the syringe burst. 610. From these experiments it is easy to understand, how any defi- ciency of tone in the Venous system will tend to prevent the ascent of the blood from the depending parts of the body, and will consequently occasion an increased pressure on the walls of the vessels, and an aug- mentation in the quantity of blood they contain. All these conditions are peculiarly favorable to the escape of the watery part of the blood from the small vessels ; and this may either infiltrate into the areolar tissue, or it may be poured into some neighboring serous cavity, pro- 342 CIRCULATION OF NUTRITIVE FLUID. ducing dropsy. Thus it happens, that such effusions may often be traced to that state of deficient vigor of the system, which peculiarly manifests itself in want of tone of the blood-vessels ; and that it is re- lieved by remedies which tend to restore this. In many young females of leuco-phlegmatic temperament, for example, there is a tendency to swelling of the feet, by oedematous effusion into the areolar tissue, in consequence of the depending position of the limbs; the oedema disap- pears during the night, but returns during the day, and is at its maxi- mum in the evening. And the congestion which frequently manifests itself in the posterior parts of the body, towards the close of exhausting diseases, in Avhich the patient has lain much upon his back, is attributable to a similar cause ; of such congestion, effusions into the various serous cavities are frequent results ; and such effusions, taking place during the last hours of life, are often erroneously regarded as the cause of death. To the same cause we are to attribute the varicose state of the veins of the leg, which is so common amongst persons of relaxed fibre, and especially in those whose habits require them to be much in the erect posture; and thi3 distension occasionally proceeds to complete rupture, the causes of which are fully elucidated by the experiments just cited. 611. It has been thought that the circulation Avithin the Cranium takes place under different conditions from that of other parts of the body. For as the cranium is a closed cavity,—a certain part of which is occupied by the cerebral substance and its membranes, the remainder being filled up with blood,—it has been argued that the amount of blood in the vessels of the brain must be ahvays the same ; and that any disturbance of its circulation must be due to a difference in the relative quantity of blood in the arteries and the veins. This idea appeared to derive support from the results of experiments, which showed that the blood is retained in the vessels within the cranium of animals bled to death, unless an opening be made in the skull, so as to allow the air to exert the same pressure upon these vessels, as upon those of other parts. But such experiments do not at all sanction the assertion, that the quantity of blood within the cranium is constant; on the contrary, we have reason to believe that it undergoes as much change as in other parts. For although the cerebral substance be incompressible, yet its bulk is subject to constant variation, according to the quantity of fluid it contains; and the presence of the cerebro- spinal fluid in the sub-arachnoid cavity of the brain and spinal cord, appears to be peculiarly destined to favor this continual change,—the proportions of it contained in the spinal and the cerebral cavities, re- spectively, being governed by the bulk of the other contents of the cranium. Thus if the vessels of the cerebrum be in their ordinary state of fulness, a certain amount of fluid is present in the sub-arachnoid cavity of the brain ; this will be pressed out into the spinal portion of the cavity, if the cerebral vessels be unusually distended with blood; Avhilst it will be increased from the latter source, so as to fill up the vacant space within the cranium, if the cerebral vessels be unusually empty. OF NUTRITION. 343 CHAPTER VII. OF NUTRITION. 1. Selecting Power of the Individual Parts. 612. The Blood which is carried into the different parts of the system, by the Circulating apparatus, is the source from Avhich all the organs and tissues of the body derive the materials of their growth and development; and, as we have seen, it is distributed by the Capillaries of the several tissues, with a degree of minuteness, which varies accord- ing to the activity of the nutrient operations taking place in the indi- vidual parts. Thus, in Nerve and Muscle, Mucous Membrane and Skin, a constant decay of the old, and a development of new tissue, are taking place, when these organs are in a state of functional activity; and a copious supply of blood is carried through every part of their substance; whilst in Cartilage and Bone, Tendon and Ligament, the amount of interchange is very small, and is effected by a much less minute reticulation of capillary blood-vessels. 613. The materials of the nutritive process being prepared in the blood, the process of nutrition is the act of each individual part; which grows and developes itself, in virtue of its own inherent poAvers, as long as the requisite conditions are supplied. The mode in which this takes place, in each individual tissue, has been already explained in the for- mer part of this Treatise. We have seen that, in the great majority of cases, the act of Nutrition is, in fact, a process of cell-growth; and that it takes place under the same conditions as the production of the simple isolated cell, which constitutes the whole of the humblest forms of Cryptogamic Vegetation,—namely, that it grows from a germ, Avhich draws to itself the materials of its nutrition, and gives to some of them a neAV arrangement, whereby they form the cell-Avail, whilst others are introduced into the cell-cavity,—and then when it has passed through its regular series of changes, it dies, and sets free its contents. We have seen that, in some cases, the germs are prepared by previously- existing cells of the same kind; whilst in others they are furnished by certain "nutritive centres," which seem to be constantly engaged in the preparation of them, deriving their materials from the blood. Fre- quently it would seem as if the original or parent-cell is able to con- tinue the production of secondary cells to an unlimited extent, even though it may have itself undergone a considerable change of form. Thus the ultimate follicles of Glands seem to be at first closed cells, which subsequently open at the part nearest to the duct, and establish a connexion with it; and having thus changed their condition, they go on developing new generations of secreting cells in their interior, from their own nuclei or germinal centres, to an unlimited extent. In like manner, the parent-cells of Muscular Fibre, which have coalesced to form the tubular Myolemma, seem to continue to develope new fibrillae from their nuclei, notwithstanding their change of form. 344 OF NUTRITION. 614. The selecting power, which is possessed by the germs of each kind of tissue, and which enables them to draAv from the blood the materials which they severally require for their development, manifests itself also in the mode in which substances, that are abnormally present in the blood, affect the condition and development of the solid tissues. Thus we find that the presence of a certain quantity of Arsenic in the blood, will produce a state of irritation in all the Mucous membranes of the body. The continued introduction of Lead into the circulating system, occasions a modification in the nutrition of the extensor muscles of the fore-arm, producing the form of partial paralysis commonly termed "wrist-drop" and the existence of this modification is shown by the presence of lead in the palsied muscles. Here we have to remark the symmetrical nature of the affection, consequent upon the occurrence of the same disorder in the corresponding parts of the two sides of the body; for these muscles appear to have the same kind of tendency to attract lead from the circulating current, in a degree that is equal on the two sides, as they have to draw from the blood the materials of their regular growth, and to develope themselves in an exactly similar manner. In like manner, the cutaneous eruptions, which are occasionally produced by the internal exhibition of iodide of po- tassium, are found to be almost precisely symmetrical; the presence of the medicine in the blood being the occasion of a disordered nutrition of certain parts of the skin; and the selecting power of particular spots being evinced, by the exact correspondence of the parts affected on the tAvo sides. 615. The same appears to be the case with regard to substances, whose presence in the blood is rather the result of a disordered condi- tion of the digestive and assimilating processes, than of their direct introduction from without. Thus in Lepra and Psoriasis,—chronic diseases of the Skin, which seem to have their origin in a disordered state of the blood, rather than in the solid tissues affected,—we find a remarkable tendency to the repetition of the patches on the two sides of the body, or on the corresponding parts of the limbs; and this we must attribute to the peculiar attraction, existing between the solid tissues of those parts, and the morbid matter circulating through them. So in those chronic forms of Gout and Rheumatism, which modify the nutrition of the joints, producing a deposit of "chalk-stones," or per- manent distortion and stiffening, we almost invariably find the corre- sponding joints of the two sides affected. The chief exceptions to the general principle, that the presence of morbid or extraneous matters in the blood affects corresponding parts alike, are found to exist where there is much febrile disturbance, or where local causes produce a peculiar tendency to disorder of a single part. The nearer the character of the morbid process is to that of the ordinary nutritive operations, the more nearly does it approach these, in the symmetry with which it developes itself.* * See Dr. W. Budd's valuable Paper on the "Symmetry of Disease," in vol. xxv. of the Medico-Chirurgical Transactions. VARYING ACTIVITY OF THE NUTRITIVE PROCESSES. 345 2. Varying Activity of the Nutritive Processes. 616. The nutritive operations go on with very great variations in their_ relative activity, under different circumstances. As a general rule it may be stated that, the greater the demand for the functional activity of the organ or tissue, the more energetic is its nutrition; and vice versd. Now this is readily understood, when it is considered, that the active state of many structures essentially consists in an act of nutrition ; thus the energy of the secreting processes is really de- pendent, as we have seen, upon the growth of the secreting cells, which make up the essential part of the gland, and the energy of the absorb- ing and assimilating process is dependent upon the development of the cells, which select and elaborate the nutrient matter. This growth is regulated mainly by the supply of blood; being increased by the afflux of blood towards the part, in consequence of the influence of the nerves upon the vessels, or through any other change in the current of the circulation. Thus the secretions are increased in amount, by emo- tions of the mind, that act (probably through the sympathetic nerve) in regulating the calibre of the arteries supplying their respective glands: or the interruption of the function of one gland shall occasion an increased nutrition, and consequently an augumented secretion, in its fellow,—as when one of the kidneys is hypertrophied, through a dis- ease in the other, that renders it incapable of performing its office. Still it would appear, that there may be variations in the activity of these organs, resulting from causes inherent in themselves (of the nature of which we know little or nothing); and that here, as elsewhere, active nutritive operations will promote the circulation of blood through the parts, whilst a languid state of the function will retard it.- 617. In certain other tissues, however, the functional activity would seem to consist rather in a waste or decay of structures previously developed ; this is the case especially in Nerve and Muscle, which are found to undergo disintegration, in exact proportion to the degree in which they are exercised; whilst the degree in which this waste is repaired, depends upon the supply of nutritive material, the quiescent state of the part, and other circumstances. But even here we find that functional activity occasions increased nutrition; in the same manner as burning a lamp with a high flame increases the amount of fluid draAvn up by the wick. For neither the nerves nor the muscles can act with energy, without a large supply of arterial blood; and this is drawn to them on the principles already mentioned (§ 60) as in- creasing the energy of the local circulation. The determination of blood to the parts, thus established, favors their increased nutrition; and thus we find that, under favorable circumstances, any set of Mus- cles, which is habitually exercised, undergoes a great increase of de- velopment ; whilst, in like manner the Nervous centres, if too great a demand be made upon their activity, are liable to become hypertrophied (especially in young persons), and may thus become subject to disor- ders, which temporarily or permanently destroy their powers. In these cases, then, the functional activity determines the increased supply of blood, and occasions the augmented growth; and increased nutrition will 346 OF NUTRITION. rarely take place in these tissues without an especial stimulus of this kind. Thus we find that, when a larger supply of nutritive matter is introduced into the circulation, than is required to repair the Avaste of these tissues, they do not undergo an increased development in conse- quence ; but an augmented nutrition is produced, either in the adipose tissue, or in the glandular structures by Avhich the superfluous matter is eliminated from the system. 618. Augmented nutrition, or Hypertrophy, then, may result in cer- tain organs, from an excessive supply of their nutrient materials; as in the case of the kidney, just mentioned; or as in the enlargement which Ave not unfrequently meet with in the livers of those, who have resided long in warm climates, and Avho have not sufficiently restricted their supply of non-azotized food to the small amount required for respiration at an elevated temperature, thereby sending an over-supply of that par- ticular class of bodies, to be separated from the blood by the liver. Or, in other cases, the increase of functional activity may be the imme- diate cause of the increased nutrition ; and this we see, not only in the nervous centres and voluntary muscles, which are put in action by the will, but in parts over Avhich the mind has no control. Thus the heart becomes hypertrophied, when an obstruction exists in the pulmonary or systemic circulation, to overcome which, increased energy of contraction is required; and in the same manner the muscular coats of the urinary and gall-bladder acquire an extraordinary increase of thickness, when long-continued obstruction, by calculi or stricture in the canals issuing from them, impedes the free exit of their contents. Sometimes, how- ever, a local hypertrophy takes place, which cannot be accounted for in either of these modes; as when a single finger is enlarged out of all proportion to. the rest, or the whole of one hand increases to a much greater size than the other, by the existence (as it would seem) in the individual part of that tendency to unusual development, which, when it affects the whole body uniformly, produces a gigantic stature. 619. Now a precisely reversed series of conditions diminishes the activity of the nutrient processes, and induces a state of Atrophy. If there be a deficiency in the general amount of nutriment introduced into the system by absorption, a general atrophy results ; and the waste being more rapid than the supply, there is a diminution in the volume of all the tissues excepting the nervous, Avhich seems to have its nutri- tion kept up even to the last, at the expense of all the rest. Such a condition results not merely from the Avant of food, but also from the Avant of power to assimilate it; and thus emaciation may take place to an excessive degree, when food of the most nutritive character is copiously supplied, and when the appetite for it is vehement; in conse- quence of disorder in the mesenteric glands, or in some other part of the apparatus particularly concerned in the elaboration of fibrine. A partial atrophy may result, in like manner, from a deficiency of the materials required for the formation of an individual tissue or organ; thus the adipose tissue throughout the body, may be atrophied, in con- sequence of an absence of those materials in the food, which are capable of being converted into fatty matter. Or a particular organ may be atrophied, by a diminution of the circulating current that should flow VARYING ACTIVITY OF THE NUTRITIVE PROCESSES. 347 to it, either in consequence of obstruction in the trunk or by the par- tial diversion of thestream of blood in another direction; and thus the liver, which is much more developed in the foetus, relatively to the rest of the body, than it is in the adult, undergoes a partial atrophy immediately after birth, in consequence of the change in the whole course of the cir- culation Avhich takes place as soon as the lungs are expanded. 620. But partial atrophy may also take piace from causes inherent in a particular organ. Thus we occasionally meet Avith limbs, which are " blighted," never attaining their due size relatively to the remainder of the body, yet not exhibiting any defect of organization. Here there would seem to be, from some unknown cause, a deficient power of growth; analogous to that which, when acting on the body in general, confines it Avithin dwarfish dimensions.—One of the most frequent causes of partial atrophy, hoAvever, is Avant of functional activity in the organ ; and this is particularly the case in regard to the Muscular and Nervous systems. Thus, as already remarked (§ 348), any set of Muscles that is long disused, becomes partially atrophied; which is probably due to the feebleness and languor of the circulation, consequent upon the absence of the demand for arterial blood. As soon as the parts are called into use again, their nutrition improves. So, also, in regard to the Nerves ; the nutrition of both the fibrous and vesicular structures appears to be entirely dependent upon the activity of their function ; and as the former are inert without the latter, we may say that the due nutrition of the nervous system entirely depends upon the functional activity of the vesicular matter. Of this we have a well-marked illus- tration in the fact, that when the cornea has been rendered so opaque by disease or accident, as to prevent the penetration of any light to the interior of the eye, the retina and the optic nerve lose after a time their characteristic structure ; so that scarcely a trace of the peculiar globules of the former or of the nerve-tubes of the latter can be found in them. 621. In the healthy condition of the organism, however, the nutri- tion in every part of the body goes on in a degree sufficient to keep it constantly ready for the performance of its appropriate function; a regular supply of the requisite materials being furnished in the aliment, and being prepared by the assimilating processes; and the products of the Avaste or decay of the tissues, together with such alimentary mate- rials as may be superfluous, being carried off by the excreting opera- tions. When the nutrition and the waste are equal, the weight of the body remains the same ; and this is commonly the case in adult age. But during the earlier periods of life, the powers of growth are greater; the demand for food is very large in proportion to the bulk of the body; and though the Avaste is rapid, and the excreting process very active (as evinced by the large amount of urea and of carbonic acid set free), the growth predominates over the decay, and the development of the whole structure proceeds at a gradually-decreasing rate, until the full stature and bulk are attained. The energy of the nutritive process is particularly manifested, in the rapidity and completeness with which severe injuries, occasioned by disease or accident, are repaired. In advanced life, on the contrary, although the Avaste is comparatively small, the renewing processes are enfeebled in a still greater degree; 348 OF NUTRITION. and there is a gradual diminution in the stature and bulk of the body, and in its physical powers. All the functions are performed with de- creased energy ; and the comparative inertness of the nutritive processes is seen in the difficulty with Avhich the effects of severe injuries are repaired, in the length of time requisite for the purpose, and frequently in the imperfection of the result. 622. During the successive periods of life, there are many remarkable changes in the relative nutrition of different organs ; Avhich we can attri- bute to nothing else, than to inherent differences in their own powers of development. Thus, during the early stages of foetal existence, the greatest energy of growth is seen in certain parts which are to answer but a temporary purpose, and which are afterwards completely atrophied. This is the case, for example, with the Corpora Wolffiana, which seems to ansAver the purpose of temporary kidneys, and in connexion with which the permanent kidneys and the genital organs are developed ; and of these bodies, though of large size in the early embryo, and evidently of great importance, no trace whatever is afterwards to be discovered. So in regard to the Supra-Renal capsules, the Thymus and Thyroid glands, and other organs, we find their proportional size the greatest, and their function evidently the most active, during foetal existence and in early infancy; after which their bulk diminishes in proportion to the rest of the body, and their functional activity seems almost at an end. 623. Even in the relative development of the organs which form essential parts of the permanent structure, Ave find considerable varia- tions at different periods of life. Thus the eArolution of the generate system does not usually take place, until the rest of the body is approaching its maturity; but cases sometimes occur, in which this apparatus attains its full development, both in the male and the female, at a very early period of childhood, and seems capable of performing its functions. In the Human species, these organs, when once evolved, remain always in a state of preparation for the performance of their function, unless thay are atrophied through complete disuse, or have lost their vigor by age, or through excessive demands upon their acti- vity ; but in most of the lower animals, the development of these organs is periodical through the whole of life, taking place at a certain season of the year, and being greatly influenced, it would appear, by the external temperature, and by the supply of food. Thus in the Sparrow the testes are no larger than mustard-seeds, during the greater part of the year ; but in the spring, they acquire the size of large peas, and it is then only that they possess any procreative power. 624. We are not always to judge of the degree of development of organs, hoAvever, by their size alone ; for the completeness of their struc- ture, and their aptitude for the performance of their functions, must also be taken into the account. Thus in the new-born infant, the organs of Digestion and Assimilation, though of small size, are so completely formed as to be able at once to take on the duty of receiving and pre- paring the nutritive materials, provided these are supplied in a form adapted to their powers; the Circulating apparatus is fully adequate to transmit the products of the action of those organs to the body in general, and to bring back the results of its continual decay; and the VARIATIONS OF NUTRITION WITH AGE. 349 Respiratory organs, together with other parts of the Excretory appa- ratus, are so completely evolved, as to be able to ^separate the effete matter, and to cast it out of the system with an energy equivalent to that of the organs by which new matter is introduced and appropriated. On the other hand, the Brain, although of larger comparative size at birth, than at any subsequent period of life, is but very imperfectly developed; for its structure is not yet so far completed, as to prepare it for a state of high functional activity. In fact it would seem as if the use of the organ, as called forth by the new circumstances in which the infant is placed as soon as it comes into the world, is essential to its complete development; and the same may be said of the Muscular system. 625. During the whole period of infancy and childhood, the current of nutrition seems peculiarly directed towards the brain ; for though its size does not continue to increase, in proportion with that of the remainder of the body, its structure is evidently being rendered more perfect, and its functional activity is excited with remarkable facility. Hence it is peculiarly liable to be acted on by various causes which may produce disease ; and the operation of remedies, which specially affect that organ, is far more poAverful than at any other period of life. Thus, whilst, a child will bear a fourth, or even a third of the dose of a purgative adequate for an adult, it is strongly affected by an eighth, or even a twelfth of the dose of a narcotic or a stimulant that would be required to produce a corresponding effect in middle life. This peculiar impressibility of the nervous system, resulting from the activity of the nutrient processes which are taking place in it, manifests itself also in other ways ; thus children are peculiarly liable to have its poAvers de- pressed by any sudden shock, such as a dIoav, or an extensive burn or laceration; whilst, on the other hand, if the depression be not fatal, they recover from its effects much more speedily than an adult would do from a similar condition. 626. During the periods of youth and adolescence, the chief energy of development (except in regard to the generative system, already noticed), appears to be directed towards the Muscular apparatus ; Avhich then increases in vigor, in a degree which surpasses its increase of size ; and the circulating and respiratory organs, upon whose energetic action there is then a corresponding demand, are peculiarly liable to disturbance of function, inducing disease in themselves or in other parts. The maladies of this period are for the most part of a sthenic or inflam- matory character; resulting, as we shall presently see, from the exces- sive activity of the assimilating processes, which are disposed to produce more fibrine than the wants of the body require. Or if, on the other hand, there be an imperfect elaboration of the nutrient materials, as happens in the tubercular diathesis, its effects are peculiarly liable to manifest themselves at this period, when the demand for nutritive matter is greatly augmented by the activity of the muscular system. G27. In adult age, there should be such a balance of all the functions, arising from the due de\7elopment and proper use of each organ, as may preserve the body in the state of health and vigor, without any marked change in the relative dimensions of its different parts, through a long 350 OF NUTRITION. series of years. The digestive, assimilating, and excreting organs, as they were the first to come to maturity, are commonly the first to fail in their activity ; but this is very generally the result of over-exertion of their powers, the amount of food introduced into the stomach being rarely (among the higher and middle classes of society at least) kept down to the real wants of the system. The muscular apparatus usually experiences the effects of this diminished nutrition, sooner than the nervous system ; the vigor of the latter being often sustained in a remarkable degree (as shown by the energy of the mental operations) through a protracted life, when it has not been overtasked at an earlier period. The very slight impairment of the nutrition of the nervous system, during the general emaciation which results from a wasting dis- ease, or during that more gradual decline of the bodily vigor which is consequent upon advancing age, is a phenomenon which strongly marks it out as the part of the body, to the maintenance of whose integrity everything else is subservient; and this is still more remarkably shown in the phenomena of starvation, in which state, notAvithstanding the dis- appearance of the Avhole of the fat, and the reduction of the weight of the body in general by about 40 per cent., the nervous system appears to lose little or none of its substance (§ 117). 3. Of Death, or Cessation of Nutrition. 628. The general cessation of the NutritiAre operations, in Death, usually depends, as formerly explained (§ 65), upon the cessation of the supply of Nutriment, in consequence of the stagnation of the Circulating current; and this stagnation may result from the direct operation of three causes ; namely,—failure in the propulsive power of the Heart, or Syncope,—obstruction to the flow of blood through the pulmonary capillaries, consequent upon a deficient supply of air, or Asphyxia,— and a disordered state of the blood itself (§ 534), which at the same time weakens the power of the heart, and prevents the performance of those changes in the systemic capillaries, Avhich afford a poAverful auxiliary to the circulation; a mode of death, for which the term Ne- crarmia has been proposed. Each of these conditions may be dependent upon a variety of remote causes, which cannot be here particularized. But it is evident that, when either one of them has been established, the nutritive processes must speedily cease, although they may continue for a short time at the expense of the blood in the capillaries of the part. The cooling of the body is another cause of their cessation ; and this is one reason Avhy molecular death (or the death of the individual organs and tissues) follows so much more closely on somatic death (or the ces- sation of the circulating and respiratory functions), in warm-blooded than in cold-blooded animals. In either case, however, the solid tissues may preserve for a time their independent vitality; and changes may take place in them, which indicate the continuance of their nutritive actions to a certain extent, even when they have been disconnected from the body. There are undoubtedly cases, however, in which the loss of vital power is as complete and immediate in the solids as in the fluids; the want of ability to avail themselves of nutriment being as decided in VARIOUS CAUSES OF DEATH. 351 the former, as the deficiency of supply is in the latter. This is seen, for example, Avhen death results from a sudden and violent shock, which destroys the vitality of the Avhole system alike (§ 604); molecular death being here consentaneous with somatic. 629. But as each component part of the Animal fabric has an indi- vidual life of its own, so must it have a limited duration of its own ; the period of termination of its vital activity, or its death, being quite inde- pendent of that of the body at large, excepting in so far as the opera- tions of the latter are requisite to afford it a constant supply of appro- priate nutriment, and to maintain its temperature at the proper eleva- tion. It is perfectly compatible, on the other hand, with the Life of the entire organism, that certain parts of it should be continually in course of decay and reneAval; and, in fact, we find that the most im- portant parts in the vital functions are performed by tissues whose indi- vidual duration is comparatively brief, but which are renewed as fast as they degenerate. We have a Avell-marked example of this in the case of the leaves of trees, which are the chief agents in the preparation of the nutritious fluid, at whose expense the permanent tissues of the trunk and branches are generated ; and although there is nothing in the Ani- mal body at all comparable to the complete exuviation which commonly takes place in the Plant at the close of the season of vegetative activity, yet there is a continual death and separation of parts that have per- formed their function, which in the end makes up a much larger aggregate. Thus there is scarcely a less complete renewal of the epidermis in Man, in the course of twelve months, than there is in Serpents, Frogs, &c, which throw it off periodically ; the only difference being, that in the one case the whole is exuviated and renewed at once, Avhilst in the other there is a continual interchange. In the exuviation of the antlers of the Deer, and of the milk-teeth of all Mammalia, we have very marked examples of this limitation of the life of individual parts, even in the highest Animals ; and as a general proposition it may be stated, that every part must degenerate, when it has gone through the whole series of changes in which its Life consists, and that it must then either die and decay, or must be so altered in its constitution, as to be able to remain inactive without further change. 630. Hence we see that the duration of vital activity must be carteris paribus, in the inverse ratio of its energy ; that is, the life of any part, or of the entire organism, must be shortened by any excess of func- tional activity ; whilst it may be prolonged by such a degree of repose, as does not involve an impairment of its nutrition. We see this most remarkably exemplified in the case of cold-blooded animals; the dura- tion of whose lives, after they have sustained some fatal injury (such as the removal of the heart or of the lungs), or are placed in any other circumstances incompatible with its continuance, is in the inverse pro- portion to the elevation of the temperature to which they are exposed, and therefore to the degree of their vital activity (§ 128). Now although this variation is comparatively little observable in the rate of life of that portion of the fabric of Avarm-blooded animals which is concerned in their organic functions (the temperature to which it is subjected being nearly constant), it is clearly seen in those organs Avhose functional 352 OF NUTRITION. activity is more under the control of the individual, and is therefore less constant. Thus, in Man, we continually notice that the duration of the powers of the Brain and the Generative system is the longest, when these organs have been moderately exercised; and that it is much cur- tailed by the excessive use of either. The duration of their activity, however, is not increased by partial or entire disuse of the organs ; for this induces a state of atrophy, on the principles already mentioned. Now we have every reason to believe, that what is true of individual parts and organs, is, true also of the whole structure; and that the existence of the entire bodily fabric may thus come to an end, without any special disease, in consequence of the limit originally set to its powers of self-renovation. It is but rarely, however, that this occurs; the various accidents of life, the neglect of ordinary precautions for the preservation of health, and hereditary tendencies to various kinds of , morbid action, being too frequently the means of cutting off the term of Human existence, long before its natural expiration. 4. Disordered Conditions of the Nutritive Processes. 631. Having thus passed in review the general conditions, under which the ordinary Nutritive processes take place, it may be well to add a few words in relation to two of their abnormal states; one or other of which is concerned in a very large proportion of the diseases that afflict the human race. In one of these, there is a tendency to the excessive production of fibrine in the blood; whilst in the other, there is a want of the proper nutritive power in the tissues, which is appa- rently due to an imperfect elaboration of that important material. The one of these conditions is termed Inflammation ; whilst the other, Avhich is less active, but more insidious, is known as the Tubercular Diathesis. 632. The extraordinary tendency to the production of Fibrine in the blood, which has been already noticed (§ 531) as one of the most im- portant characters of Inflammation, seems to be always conjoined with a depressed vitality of the tissues of some part of the body, which indis- poses them to the performance of their regular nutritive operations ; and this part may undergo a variety of changes, according to the degree in which it is affected. The depressed condition of its nutritive operations involves, on the principles explained in the preceding chapter, a languor in the movement of blood through it, together with a distensible state of the capillaries, which causes them to contain a far greater amount of that fluid than under ordinary circumstances. On the other hand, there is a tendency to the production of an increased amount of plastic mate- rial in the blood, as if for the reparation of the part whose vitality is lowered. What is the immediate cause of this production is still doubt- ful ; but we see the consequences of the deficiency of it in those asthenic or unhealthy Inflammations, which so frequently involve the destruction of a large amount of tissue ; the degeneration of the part first affected soon extending itself to others, if there be no limit set up by the repara- tive powers of the blood.—In ordinary or sthenic Inflammation, of which the increase of fibrine in the blood, and a diminution in the power of appropriating it on the part of the tissues, are the most characteristic suppuration; ulceration; gangrene. 353 phenomena, the simplest result is the effusion of fibrinous matter, or organizable lymph, into the substance of the part inflamed, or upon the nearest free surface; and thus is produced a condensation of the tissue, or a new growth upon the membrane. But when the depression of vitality is more complete, the tissue at that spot gradually dies and dis- integrates ; and Avhilst itself undergoing such changes, it gives origin to similar changes in the effused fibrine, which it converts from a plastic or organizable deposit, into an aplastic or unorganizable one, namely, pus, the cells of Avhich degenerate without passing into any higher or more permanent form of tissue, whilst the liquid through which they are dispersed has lost its coagulating power. Thus is produced the Suppurating process; which may either take place in a cavity thus ex- cavated in the substance of a tissue or organ; or on a free surface. In either case, the surrounding tissues, which are less inflamed, and in which the vitality is impaired but not destroyed, become consolidated by a deposition of organizable fibrine, which prevents the infiltration of pus through their substance. If this should not occur, through a want of power to generate well-elaborated fibrine, the suppurating process extends itself rapidly, with the most calamitous results; the properties of pus being such, as to produce a tendency to decomposition, both in the blood, and in the solid tissues into the substance of which it may be carried. 633. Another consequence of Inflammation is Ulceration, which is a breach of surface caused by the same process as that Avhich forms the cavity of an abscess,—namely, the degeneration of the inflamed tissue, and the removal of its particles, either by absorption, or by solution and ejection in pus. Many ulcers commence as abscesses near the surface, which at last come to open upon it; and others are pre- ceded by inflammation of the superficial tissues, which die and are thrown off, leaving a vacuity, which may be subsequently increased by the extension of the degeneration of the deeper parts. These may either die and be thrown off en masse, constituting what is known as the "sloughing ulcer," or they may disintegrate more slowly, and may be dissolved in the discharge from the ulcerated surface. This dis- charge, when proceeding from a spreading ulcer, is usually of a thin ichorous quality, and has the power of exciting unhealthy action even in healthy parts to Avhich it may be applied; and it is its change to what has been designated as "laudable pus," that indicates the cessa- tion of the destructive, and the commencement of the reparative pro- cess (§ 636). 634. The state of Gangrene, which consists in the entire loss of vitality of the part, with a complete cessation of the circulation through it, is commonly regarded as a result of Inflammation, when this process occurs in its most intense form; but it may be more rightly considered as the ultimate consequence of the causes which produce Inflammation. For it is an essential part, as Ave have seen, of the condition of Inflam- mation, that the vitality of the affected tissues should be lowered; and thus there is in them always a tendency to death, which is most com- pletely developed in Gangrene. We have a Avell-marked example of this complete destruction of the life of a part, by the intense operation 354 OF NUTRITION. of causes, which, when less potent, occasion Inflammation, in the case of frost-bites produced by Cold; for this agent at the same time pro- duces contraction of the blood-vessels, and depression of the vital powers of the solid tissues, proceeding to the complete destruction of them; whilst in the parts adjoining those which are actually killed, the inflammatory state is developed, an effusion of fibrine being produced, which serves to plug up the mouths of the vessels, and thus to prevent hemorrhage, when the mortified part drops off. Here we see, that the violent action of cold completely destroys the vitality of the part most exposed to it; and this by its direct influence on the properties of the organized structure. No inflammation can take place in the part thus killed, because the vital processes are altogether brought to an end. But inflammation takes place in the adjoining parts, which are less seriously affected; for the depression of their vital powers occasions the result already adverted to,—namely, the production of an increased amount of fibrine in the blood, and an infiltration of this substance into their tissues. The same is the case, with regard to the operation of other powerful agents; such as those which (like Caustic Potass, or Sulphuric Acid) destroy the vitality of the parts to which they are ap- plied, by the chemical decomposition of their tissues. The Inflammatory process is set up, not in the parts which are killed by the application, but in the surrounding tissues, whose vitality has been simply depressed; and thus, when the slough, or dead part, is cast off, there is a prepara- tion for the development of new tissue to supply its place, from the superabundant plastic materials of the surrounding parts. 635, If, then, we limit the term Inflammation, as there seems reason to do, to that state, in which there is a tendency to stagnated circula- tion, with increased production of Fibrine, in the vessels of the part, we see that the Gangrene cannot be a result of that process, which is one rather of reparation than of destruction. But Gangrene proceeds, where we can distinctly trace its causes, from the violent operation of the same agents, as those which, in a less degree, produce Inflammation. And where this last process is not set up at the line of demarcation between the living and the dead parts, Gangrene, like Suppuration, has a tendency to spread; the influence of the decay, which is taking place in one part, having a tendency to propagate itself, to the adjoining tissue ; and a constantly-extending destruction being thus produced. 636. We have now to speak of those reparative processes, by which the effects of disease or injuries are more or less perfectly recovered from.—The healing of a simple wound may take place by the direct adhesion of its walls, when they can be drawn closely together; but more frequently it is accomplished by the intermediation of a thin layer of "coagulable lymph," which may be thrown out for the purpose of reparation, without the existence of inflammatory action. But the reparation of wounds, in which there has been so great a loss of sub- stance that neither direct nor indirect adhesion can take place, is accomplished by the gradual development of new tissue from the "nucleated blastema" with which the cavity is first filled. This, however, may occur in two very different modes; and from the inqui- ries of Mr. Paget it appears that the determination of one or the other # REPARATIVE PROCESS—GRANULATION. 355 of them is chiefly dependent on the condition of the wound, as to seclu- sion from air, or exposure to it. When the reparative effusion is poured out into a subcutaneous wound, the "nucleated blastema" appears to be gradually developed into fibrous tissue without any loss, ^and usually with freedom from local inflammation (beyond what may be the immediate result of the injury), as well as from constitutional irritation. This process seems to take place naturally in cold-blooded animals, even in superficial wounds; the contact of air not producing that disturbance in it, which it occasions in warm-blooded animals. And Nature frequently endeavors (so to speak) to bring it about in the superficial wounds of warm-blooded animals, by the formation of a large scab, which protects the exposed surface; but this happens much less frequently in the Human subject, than it does among the lower animals; the unnatural conditions in which a large proportion of the more civilized races habitually live (especially deficient purity of the air, continual excess in diet, and the frequent abuse of stimulants), being obviously unfavorable to it. The application of steam to wounded surfaces has been found to favor the reparation by the most healthy process; and the formation of an artificial scab by means of resinous unguents has also been practised Avith advantage. It is the duty of the Surgeon to endeavor to promote it by every means in his power; since it is the method of healing, which is not merely the most desirable as regards its economy of nutritive material and freedom from constitutional irritation, but which most completely supplies the loss of substance, so that the cicatrix does not contract. The newly- formed fibrous tissue becomes vascular, by the extension of loops or arches from the adjacent capillaries, and of other loops from these; and subsequently other structures—such as bone, lymphatics, and nerves,—may be developed in it. True Cartilaginous tissue, and the higher form of Muscular fibre, however, seem never to be thus generated de novo in the new tissues of a repaired part; so that wounds of Carti- lages and Muscles are united by simple fibrous texture. 637. In an open wound, on the other hand, which is healing by the process termed Granulation, the "nucleated blastema" is rapidly deve- loped into cells, amongst which vessels speedily extend themselves; but the vitality of this tissue is very low, and that part of it which is ex- posed to the air passes into the condition of pus, its cells being either imperfectly developed from the first, or speedily undergoing degenera- tion. Thus there is a constant waste of plastic material, the amount of which, in the case of an extensive suppurating sore, must be a serious drain upon the system; whilst at the same time, the local inflammation is greater, and gives rise to more or less of constitutional disturbance; and the formation of neAV tissue is so much less complete, that by its subsequent degeneration, and removal by absorption, a contracted cica- trix is produced, Avhich is different from the original texture. The new tissue is here produced by a metamorphosis of cells into fibres; and this change is taking place in the deeper part of the granulation-structure, whilst the more superficial is degenerating into pus.—The difference between the two modes of reparation now described, is often one of life and death, especially in the case of large burns of the trunk in children; 356 OF NUTRITION. for it frequently happens that the patient sinks under the great consti- tutional disturbance occasioned by a large suppurating surface, although he may have survived the immediate shock of the injury. 638. If the Fibrine of the Blood, however, be not well elaborated, it does not possess its due organizability; and thus instead of being con- verted, either when effused as an Inflammatory product, or in the ordi- nary Nutritive process into solid tissue, proper to the part in Avhich it is deposited, it is liberated from the vessels in a state, which prevents any but a very imperfect structure from being developed by it, and which tends to very speedy degeneration. This is the condition of the Tubercular substance, which is so often found to replace the proper tissue, especially in the lungs; being slowly deposited there, by a sort of degradation of the regular nutritive operations; and being effused in larger quantity, when the inflammatory process is set up. There is every degree of gradation between the plastic or organizable deposit of well-elaborated Fibrine, the caco-plastic or imperfectly-organizable mat- ter of Tubercle, and the aplastic or non-organizable matter of Pus. The microscopic examination of tubercular deposits shows, that they some- times contain fully-developed cells and fibres, analogous to those of fibrinous exudations; but that more frequently, the cells and fibres are imperfectly formed, and are accompanied by a large quantity of a gra- nular substance, which strongly resembles coagulated Albumen; and that in many cases, there is scarcely any trace of organization in the mass. The greatest degree of organization is found in the semi-trans- parent, miliary, gray, and tough yellow forms of Tubercle ; the least in the opaque, crude, or yellow Tubercle.—It is the opinion of some emi- nent Pathologists, that Tubercular matter is always deposited in the first instance in the cellular form; but that it tends to undergo a rapid and complete degeneration. 639. The constitutional state, which predisposes to this perversion of the ordinary nutritive operations, and which is knoAvn as the Tuber- cular Diathesis, may be the result of the continued operation of any causes, that tend to depress the vital powers; such as insufficient nutri- tion, habitual exposure to cold and damp, protracted mental depression, &c.; or it may be derived from the operation of the same or other causes on the ancestors of the individual, being one of those disorders which has a peculiar tendency to become hereditary. The treatment must be directed to the invigoration of the system by good food, active exercise, pure air, warm clothing, and cheerful occupations; and by the due employment of those means, at a sufficiently early period, many valuable lives may be saved, which would otherwise fall a sacrifice to Tubercular disease in the lungs, or other important organs.—Much reason has lately presented itself for the belief, that a deficiency of appropriate oleaginous constituents in the food exerts a marked influ- ence in the production of the Tubercular diathesis. This would appear to be indicated by the very marked benefit which has been derived in the treatment of Pulmonary Consumption and other tubercular diseases, from the use of Cod-liver oil, or of other easily assimilated fish-oils. And the same view is confirmed by the remarkable exemption of the Icelanders (whose diet is extremely oleaginous) from Tubercular dis- NATURE AND CONDITIONS OF THE RESPIRATORY PROCESS. 357 eases, notwithstanding that the general habits of the people would seem peculiarly favorable to their production. _ 640. There is another remarkable class of diseases, resulting from a disordered condition of the nutritive processes ;—those, namely of a malignant nature. We not unfrequently meet with abnormal growths of a fatty, cartilaginous, fibrous, or bony structure; which appear to originate in some perverted action of the part itself, and which have little tendency to reappear in the same part, when they have been removed, still less, to reappear in distant parts. But the various forms of Malignant or Cancerous disease are peculiar in this,—that they are composed of cells, sometimes of a globular form (see Fig. 18), sometimes elongated or spindle-shaped, having a power of rapid°multi- plication, and not capable of changing into any kind of normal tissue. When a truly cancerous growth has once established itself in any part of the body, it may increase to an unlimited extent, obtaining its nourishment from the blood-vessels in its neighborhood, and destroy- ing the surrounding parts by its pressure, as well as by drawing-off their supply of aliment. When it has developed itself to a consi- derable degree in one part, it is very liable to make its appearance in others, especially when the original growth has been removed ; and hence the judicious surgeon is disinclined to remove a Cancerous growth of any but the most limited kind; knowing that the disease is almost certain to reappear. There is a strong analogy between such Cancerous growths, and the low forms of Fungoid Vegetation, which develope themselves in the interior of the higher Plants, and even in Animal bodies; and in both cases, the disease may be propagated by inoculation from one individual to another. But still it appears pro- bable that Cancerous disease, like tubercular, is of constitutional origin ; and the peculiar tissue which characterizes it, is perhaps to be regarded simply as_tlm manifestation of the presence of a morbid matter in the blood, which is thus removed from the circulating current; just as fatty matter is removed by an increased formation of Adipose tissue, or as the elements of the excretions are eliminated by an increased growth of the gland-cells of which they are the appropriate pabulum. CHAPTER VIII. OF RESPIRATION. 1. Essential nature and Conditions of the Respiratory Process. 641. The function of Respiration essentially consists in an inter- change of oxygen and carbonic acid, between the blood of the Animal and the surrounding medium ; carbonic acid being given out by the blood, and oxygen entering in its stead. It has been already noticed (§ 84) that this function is performed likewise by Plants; although, in consequence of their deriving a large part of their food from the 358 OF RESPIRATION. atmosphere by a converse process—the absorption of carbon and the liberation of oxygen,—their true respiration is commonly overlooked. It may, therefore, be regarded as common to all Organized beings. Every one is conscious, in his own person, of the imperative demand for the due performance of this operation. If the breath be purposely held for a few seconds, a feeling of distress is experienced, Avhich increases every moment, and at last prompts irresistibly to the respi- ratory movement. And if the admission of air to the lungs be in any way prevented, the respiratory movements are at first increased in energy, violent efforts are made to obtain the needed supply; these are succeeded by irregular convulsive actions, and at the same time insensibility comes on; and within a short time all movement ceases, the circulation of the blood is suspended, and a stop is put to all the vital operations of the body. This state, which is termed Asphyxia, usually comes on, in a warm-blooded animal, within ten minutes of the time when the respiration is completely checked ; thus affording the most convincing proof of the importance of that function in the Animal economy. In many cold-blooded tribes, hoAvever, a much longer sus- pension may be borne with impunity ; as also by warm-blooded animals, when the general activity of their functions is lowered in the state of hibernation (§ 121). We shall now inquire into the sources of the necessity for this interchange of oxygen and carbonic acid; and the mode in which the suspension of it acts upon the system at large. 642. All Organized bodies, as already explained, are liable to con- tinual decay, even whilst they are most actively engaged in performing the actions of Life; and one of the chief products of that decay is car- bonic acid. A large quantity of this gas is set free, during the decom- position of almost every kind of organized matter; the carbon of the substance being united with oxygen supplied by the air. Hence we find, that the formation and liberation of carbonic acid goes on with great rapidity after death, both in the Plant and in the Animal; and that it takes place also, to a very great extent, in the period that often precedes the death of the body, during which a general decomposition of the tissues is going on. Thus in Plants, as soon as they become unhealthy, the extrication of carbon in the form of carbonic acid takes place in greater amount, than its fixation from the carbonic acid of the atmosphere ; and the same change normally takes place during the period that immediately precedes the annual fall of the leaves, their tissue being no longer able to perform its proper functions, and giving rise by its incipient decay, to a large increase in the quantity of car- bonic acid set free. The same thing probably happens in the Animal body, during the progress of many diseases which are attended with an extraordinary tendency to decomposition in the solids and fluids; for in such cases the blood usually exhibits an unusually dark hue, indicating that it has not been properly freed from the usual amount of carbonic acid which it has received from the tissues. It has not yet been accu- rately determined, however, Avhether there is an increase in the amount of carbonic acid actually thrown off in such cases. 643. Hence, the first object of the Respiratory process, which is common to all forms of Organized being, is to extricate from the body SOURCES OF EXCRETION OF CARBONIC ACID. 359 the carbonic acid, which is one of the products of the continual decom- position of its tissues. The softness of many of the tissues of Animals, and the large quantity of fluid contained in their bodies, render them more prone than Plants to this kind of decomposition ; and, in warm- blooded animals, the high temperature at which the fabric is usually maintained, adds considerably to the degree of this tendency ; so that the waste of their tissues, from this cause alone, is as much greater than that of cold-blooded animals, as the latter is than that of Plants. But when the temperature of the Reptile is raised by external heat to the level of that of the Mammal, its need for respiration increases, owing to the augmented Avaste of the tissues. When, on the other hand, the Avarm-blooded Mammal is reduced, in the state of hibernation, to the level of the cold-blooded Reptile, the waste of its tissues diminishes to such an extent, as to require but a very small exertion of the respiratory process to get rid of the carbonic acid, which is one of its chief products. And in those animals Avhich are capable of retaining their vitality when frozen (§ 136), or Avhen their tissues are completely dried up (§ 159), the decomposition for the time is entirely suspended, and consequently there is no carbonic acid to be set free. 644. But another source of Carbonic acid to be set free by the Respiratory process, and one wdrich is peculiar to Animals, consists in the rapid changes which take place in the Muscular and Nervous tissues, during the period of their activity. It has been already shown (§ 361), that there is strong reason to believe the Avaste or decomposi- tion of the Muscular tissue to be in exact proportion to the degree in which it is exerted; every development of muscular force being accom- panied by a change in the condition of a certain amount of tissue. In order that this change may take place, the presence of Oxygen is essential; and one of the products of the union of oxygen with the elements of muscular fibre, is carbonic acid. The same may probably be said of the Nervous tissue (§ 384). Hence it may be stated as a general principle, that the peculiar waste of the Muscular and Nervous substances, which is a condition of their functional activity, and which is altogether distinct from the general slow decay that is common to these tissues with others, is another source of the carbonic acid Avhich is set free from the animal body: and that the amount thus generated will consequently depend upon the degree, in which these tissues are exercised. In animals which are chiefly made up of the organs of vegetative life, in whose bodies the nervous and muscular tissues form but a very small part, and in Avhose tranquil plant-like existence there is but very little demand upon the exercise of these structures, the quantity of carbonic acid thus liberated will be extremely small. On the other hand, in animals, whose bodies are chiefly composed of muscle, and whose life is an almost ceaseless round of exertion, the quantity of carbonic acid thus liberated is very considerable. 645. We are enabled to trace the connexion betAveen the amount of muscular exertion, and the quantity of carbonic acid set free in the act of respiration, in the class of Insects, better than in any other. They have no fixed temperature to maintain ; and they are consequently not in the condition of Avarm-blooded animals, in which the quantity of 360 OF RESPIRATION. carbonic acid set free is kept up to a more regular standard by the provision to be presently noticed. On the other hand, they are pre- eminent among all Animals, in regard to the energy of their muscular power in relation to the bulk of their bodies ; and the waste of muscu- lar tissue during their state of activity must therefore be very great. Thus a Humble Bee has been found to produce one-third of a cubic inch of carbonic acid in the course of a single hour, during Avhich its whole body Avas in a state of constant movement, from the excitement consequent upon its capture ; and yet during the whole twenty-four hours of the succeeding day, which it passed in a state of comparative rest, the quantity of carbonic acid generated by it was absolutely less. 646. Besides these sources of Carbonic acid, which are common to all animals, there is another, Avhich appears to be peculiar to the two highest classes, Birds and Mammals. These are capable of maintain- ing a constantly-elevated temperature, so long as they are supplied with a proper amount of appropriate food ; and their power of doing so appears to depend upon the direct combination of certain elements of the food, with the oxygen of the air, by a process analogous to com- bustion ; these elements having been introduced into the blood for that purpose, but not having formed a part of any of the solid tissues of the body, unless they have been deposited in the form of fat. The nature of these substances has been already noticed (§ 430). It is quite clear that they cannot be applied in their original form, to the nutri- tion of the tissues that originate in proteine-compounds ; and it is tole- rably certain that, in the ordinary condition of the body, they undergo no such conversion, as Avould adapt them to that purpose. The Liver seems to afford a channel, by which some of the fatty matters are drawn off from the blood; but even these seem, in part at least, to be reab- sorbed (§ 725), and to be thrown off by the respiratory process. 647. The quantity of Carbonic acid, that is generated directly from the elements of the food, seems to vary considerably in different animals, and in different states of the same individual. In the Carni- vorous tribes, which spend the greater part of their time in a state of activity, it is probable that the quantity Avhich is generated by the waste or metamorphosis of the tissues is sufficient for the maintenance of the required temperature,—and that little or none of the carbonic acid set free in respiration is derived from the direct combustion of the materials of the food. But in Herbivorous animals of comparatively inert habits, the amount of metamorphosis of the tissues' is far from being sufficient; and a large part of the food, consisting as it does of substances that cannot be applied to the nutrition of the tissues, is made to enter into direct combination with the oxygen of the air, and thus to compensate for the deficiency. In Man and other animals, which can sustain considerable variations of climate, and can adapt themselves to a great diversity of habits, the quantity of carbonic acid formed by the direct combination of the elements of the food Avith the oxygen of the air, will differ extremely under different circumstances. It Avill serve as the complement of that Avhich is formed in other ways; so that it Avill diminish with the increase, and will increase with the diminution, of muscular activity. On the other hand, it will vary in SOURCES OF CARBONIC ACID IN THE ANIMAL BODY. 361 accordance with the external temperature; increasing with its diminu- tion, as more heat must then be generated; and diminishing with its increase.—In all cases, if a sufficient supply of food be not furnished, the store of fat is drawn upon ; and if this be exhausted, the animal dies of cold (§ 117). 648. To recapitulate, then; the sources of Carbonic acid in the Ani- mal body are threefold.—1. The continual decay of the tissues ; which is common to all organized bodies ; which is diminished by cold and dryness, and increased by warmth and moisture; which takes place with increased rapidity at the approach of death, whether this affect the body at large, or only an individual part; and which goes on un- checked, when the actions of nutrition have ceased altogether.—2. The metamorphosis, which is peculiar to the Nervous and Muscular tissues; which is the very condition of their activity; and which, therefore, bears a, direct relation to the degree in which they are exerted.—3. The direct conversion of the carbon of the food into carbonic acid; * which is peculiar to warm-blooded animals; and which seems to vary in quantity, in accordance with the amount of heat to be generated. 649. Now the function of Respiration has for its object, not merely to extricate ^the Carbonic acid which is generated in the system, but likewise to introduce the Oxygen Avhich is required to form that car- bonic acid; the proportion of oxygen in the tissues, and in the combus- tible materials of the blood, not being sufficient for this purpose. Hence it is not enough, that the carbonic acid should be removed; for this may be accomplished by causing an animal to breathe an atmo- sphere which contains no oxygen. Any cold-blooded animal, such as a Frog or a Snail, may be kept in hydrogen or nitrogen for several hours or even days; and will give out, during that time, an amount of car- bonic acid nearly as great, as if it had been respiring atmospheric air. But the continued production of carbonic acid must have a limit, occa- sioned by the want of oxygen, and death will then supervene.—On the other hand, a supply of oxygen may be freely afforded; and yet the presence of even a small amount of carbonic acid in the surround- ing atmosphere (in addition to that which is normally present in it, § 81) will impede the extrication of that substance from the blood; and if the excess be considerable, the carbonic acid will not be set free at all; so that the same injurious results follow, as if respiration Avere al- together prevented from taking place. 650. These two actions are accomplished by the very same act; ad- vantage being taken of the property of "mutual diffusion," Avhich is common to all gaseous substances that do not unite chemically with one another. In virtue of this property; Hydrogen, the lightest of gases, and Carbonic acid, one of the heaviest, when introduced into the same vessel, will be found in a short time to have uniformly mixed, notwith- standing the difference of their specific gravities, which are as 1 to 22. Now this intermixture will take place, when the two gases are sepa- rated by a porous septum ; each gas passing towards the other, by an action resembling the Endosmose and Exosmose of liquids (§ 491). And it may also take place, when one of the gases is diffused through a liquid; provided that the other gas is likewise capable of being 362 OF RESPIRATION. absorbed by the liquid. In this manner, as already mentioned, the surface of venous blood, enclosed in a bladder, may be made to exhibit the arterial hue, by suspending the bladder in an atmosphere of oxygen; for the carbonic acid of the blood, and the surrounding oxygen, will overcome by their mutual attraction the obstacle inter- posed by the bladder ; and the former will be lifted out, so to speak, and will be replaced by the latter. It has been found by experiment, that the free carbonic acid diffused through blood, may be more completely extricated from the liquid, by exposing it to hydrogen, than by placing it under the vacuum of an air-pump ; for in the latter case there is nothing to replace it, and the attraction between the gas and the liquid tends to resist the exhausting influence of the vacuum; whilst in the former, the blood receives one gas in exchange for the other, so that the whole force of the tendency to mutual diffusion is exercised in lift- ing out the carbonic acid. 651. The immediate purpose of the organs of Respiration, then,— whatever may be the variety in their form,—is this: to expose the blood to the air, in a state of such minute division as to present a very extended surface, a thin membrane only being interposed between them. For this purpose we find a certain organ, or set of organs, spe- cially set apart in all the higher animals; and this is formed by a prolongation of the general surface, either externally or internally, according to the mode in which the respiration is accomplished. Thus in Fishes and aquatic Molluscs, the blood is aerated by exposure, not directly to the atmosphere, but to the air which is dissolved in the water they inhabit; and the respiratory apparatus is formed in them of an extension of the external surface, at a particular part, into innu- merable delicate fringe-like processes, the gills (Fig. 100); every divi- sion of which contains a network of blood-vessels (Fig. 104); so that Fig. 100. Doris Johnstoni, a Nudibranchiate Gasteropod, showing the luft of external gills. the amount of blood which is exposed to the surrounding medium at any one time, is collectively very great, although the quantity contained in each gill-filament is very minute. On the other hand, in all the air- breathing Vertebrata, the blood is exposed to the atmosphere, through the medium of an internal membranous prolongation, which is conti- nuous Avith the mucous membrane lining the mouth and nostrils; this forms a pair of sacs, termed lungs, communicating with the back of the mouth by means of a tube called the trachea or windpipe, through which air is freely admitted to the cavities thus formed (Fig. 105). RESPIRATION IN MOLLUSCS. 363 The blood is minutely distributed on the walls of these sacs by a close network of Capillary vessels (Fig. 106); and not only on the external walls, but also on numerous partitions by which the cavities are sub- divided with more or less minuteness, so as greatly to extend the vas- cular surface. 652. Such is the essential nature of the Respiratory apparatus ; but in order that it may be carried into that vigorous operation, which is required in the higher classes of animals, various supplementary arrange- ments are made, for the purpose of promoting the due influence of the air upon the blood. In the first place, the capillary vessels of the respi- ratory surface are connected with arterial trunks, which issue immedi- ately from the heart, and which thus convey a constant stream of blood from that organ; whilst they give origin to venous trunks, which ter- minate directly in the heart, and which are ready to convey back to it the blood that has undergone aeration. Thus by the energetic action of the heart, and by the force generated in the capillaries of the lungs (§ 598), a constant renewal is effected in the blood, which is exposed to the air through the medium of these organs. On the other hand, the renewal of the blood Avould be useless, unless a fresh supply of air were continually being introduced, and that Avhich had been vitiated, by the loss of its oxygen and the admixture of carbonic acid, were removed; and this is effected by a series of muscular movements, which are adapted for the alternate expulsion of the vitiated air from the lungs, and for the introduction of a fresh supply of pure air from the atmosphere. These movements are kept up by a certain part of the nervous system; but they are not dependent upon any exertion of the will, for they con- tinue during profound sleep, and in other states in which even conscious- ness is altogether suspended. 2. Different forms of the Respiratory Apparatus in the lower Animals. 653. Before proceeding to consider, in more detail, the structure and actions of the respiratory apparatus in Man, Ave may advantageously glance at the mode, in which this function is effected in the lower ani- mals.—In the lowest and simplest, Avhich are inhabitants of the water, Ave do not find any special apparatus for the aeration of the fluids of the body ; this being accomplished by the exposure of them to the surround- ing medium, through the thin integument; and the interchange of the layer of water (holding air in solution) in contact with the aerating sur- face, is effected either by the general movements of the body, or by the action of the cilia (§ 234) which produce the currents necessary for this purpose. Not unfrequently, the internal surfaces—such as the walls of the stomach and of other cavities,—seem as much concerned in this function as the external, or even more so ; these cavities being distended with Avater taken in through the mouth, and this water being frequently renewed by the ejection of that which has been vitiated, and by the in- troduction of a fresh supply. This is the case in the Sea Anemone, for example, and in many other polypes ; and there are certain higher forms of the same class, in Avhich there is a great dilatation of the pharynx, Avhich seems peculiarly destined for the aeration of the fluids, 364 OF RESPIRATION. —being filled with water, and then suddenly emptied at tolerably regu- lar intervals. 654. In the various classes of the Molluscous sub-kingdom, we find the respiration provided for, by the adaptation of distinct organs for the purpose. As most of the animals of this group are inhabitants of the water, the respiration is usually carried on by means of gills, rather than by any organ resembling a lung. The latter is found, however, in a feAV species; such as the Snail, Slug, and other terrestrial air-breathing Molluscs, and usually consists of a simple cavity, situated in the back, communicating directly with the air through an aperture in the skin, and having its Avails covered Avith a minute network of blood-vessels. The form and position of the gills differ extremely in the several classes of Molluscous animals. In the lowest the respiratory surface is formed, as in the higher Polypes, by a dilatation of the Pharynx; but sometimes, instead of surrounding a large cavity, it forms a special riband-like fold of membrane, passing from one end of it to the other, on which the blood is minutely distributed. In this group of animals, there is a regular system of canals for the conveyance of the blood; but these, in many parts of the system, and especially on the respiratory membrane, do not seem to be furnished with distinct walls, and are rather mere channels excavated in the tissues. And the circulation is liable to a continual change in its direction, the blood being sometimes transmitted to the respiratory surface before it proceeds to the body, and sometimes after it has traversed the other tissues (§ 557). The Avater in contact with the respiratory surface is continually renewed by the action of the cilia, with which it is thickly covered. 655. In certain of the Molluscs, inhabiting bivalve shells, we find that the internal surface of the mantle or skin that lines the valves, is the special organ of respiration ; the external water having free access to these by the separation of the skin along the edges of the valve, so that it enters the cavity in which the vriscera are lodged, and bathes their exterior. But in most bivalve molluscs, the internal surface of the mantle is doubled (as it were) into four riband-like folds, which are delicately fringed at their edges, and which have, in fact the same essential structure as the gills of higher animals (§ 663). To these the blood is transmitted, Avhen it has been rendered venous by traversing the vessels of the body generally : and in these it is exposed, through a surface Avhich is greatly extended by the minute division of the fringes, to the action of water introduced from without, and constantly renewed by ciliary action. In many of these animals, as in the common Oyster, the two lobes of the mantle are so completely separated, that the Avater can still enter freely between the valves, but in general, they are more or less united, so that the cavity in which the gills lie is partially closed. There is always a provision, however, for the free access of water from without by means of two apertures, one Tor its entrance and the other for its ejection, and in certain species which burrow deeply in sand or mud, these apertures are furnished with long tubes, or siphons, which convey the water from near the entrance of the burroAV, and carry it thither again. In these also, a continual flow of water over the respi- RESPIRATION IN WORMS AND CRUSTACEA. 365 ratory surface is maintained by the vibration of the cilia, with which they are clothed. 656. The position of the gills, in the Mollusca of higher organization, is extremely variable. Sometimes they are disposed upon the external surface of the body, and form delicate leaf-like or arborescent appendages (Fig. 101); whilst in other FiS- 101- cases they are enclosed in a special cavity or gill- chamber, to which water is freely admitted from Avithout; a continual interchange being provided for, either by ciliary action, or by muscular move- ments specially adapted for the purpose. The blood is conveyed to them, after having become venous in traversing the capillaries of the general system, by means of large channels and sinuses excavated in the several parts of the body (§ 556); and after being aerated in the gills, it returns to , the heart to be again conveyed to the system. In One of the art^'went pro- the Cuttle-fish tribe, there are supplementary S JllTtL^ltS hearts at the origin of the branchial arteries, or and enlarsed- vessels that distribute blood to the gills; and these have evidently for their purpose, to render the respiratory circulation more energetic, and thus to increase the aeration of the blood, in the degree required for the vigorous habits of these animals, Avhich present a remarkable contrast to the sluggish, inert character of the Mollusca in general.—In these classes, taken as a whole, the respiration is low in its amount. The blood contains no red corpuscles, excepting perhaps in the highest class; and the change in its composition, which is effected by the air, is con- fined, therefore, to the fluid plasma, or liquor sanguinis. And as it is not exposed directly to the air, except in a few species, but to the air contained in the water inhabited by the animals, this change cannot be very energetically performed. But as the life of these animals is chiefly vegetative,—as their movements, except in the highest classes, are few and feeble,—and as they maintain no independent heat,—there is but little need of that interchange, which it is the object of the respiratory process to effect; and these animals can sustain the complete suspension of it for a long time. _ 657. Among many of the Articulated tribes, the respiration is car- ried on upon a similar plan. In some of the lowest, such as the Tape- worm of the intestinal canal, there is no special provision for the aera- tion of the fluids; the soft integument permitting the extrication of car- bonic acid, and imbibition of oxygen, in the required degree. This is but very small, however; the life of these animals being almost purely vegetative. In the Marine Worms, which constitute a numerous and interesting group, endowed with considerable locomotive powers, and leading a life of almost constant activity, there is, on the other hand, a special provision for this function ; the blood being transmitted, in the course of its circulation, to a series of gill-tufts, which are composed of a delicate membrane prolonged from the external surface of the body, and which sometimes have the form of branching trees, and sometimes of delicate brushes made up of a bundle of distinct filaments. In either 366 OF RESPIRATION. case, the filaments are traversed by blood-vessels, and are adapted to bring the blood into close relation with the surrounding water; and the continual interchange of the latter is provided for by the restless move- ments of the body. The tufts are sometimes arranged along every seg- ment of the body ; and their multiplication prevents them from indivi- dually attaining any considerable size. In other cases, they are disposed at intervals; and they are then larger, being less numerous. Their most beautiful development is where they are present on the head only, the rest of the body being enclosed in a shelly or sandy tube, as in the Ser- pulse and Terebellee. The gill-tufts then frequently present the appear- ance of a flower, endowed, when alive, Avith the most brilliant and deli- cate hues. In many animals of this group, there is a small supplemen- tary heart at the base of every one of the vessels that distribute the blood to the gills ; and this is obviously designed to aid in the respiratory circulation, for which the feeble action of the dorsal vessel would not furnish sufficient power (§ 552). 658. The higher Articulated classes are, for the most part, adapted to atmospheric respiration, on the plan to be presently explained; but there is one class, that of Crustacea, whose respiration is still carried on through the medium of water. In the lowest forms of this group, there is no special respiratory apparatus; the general surface being soft enough to admit of the required aeration of the fluids through its own substance, and the animal functions being performed with so little activity, that a very small amount of interchange is required. In the higher orders, however, Avhose bodies are encased Avithin a hard envelope, we find external gills, like those of many Molluscs; and these are attached to the most movable parts of the body,—one or more pairs of legs being in some instances kept in constant agitation, for the purpose of producing currents in the surrounding fluid, that may serve for the aeration of the blood. In the Crab-tribe, Avhich constitutes the highest family of this class, the gills are themselves enclosed within a cavity, formed by a sort of doubling of the hard integument of the under side of the body ; and a constant stream of Avater is maintained through this, by means of a peculiar valve, situated in the exit pipe; the continual movement of the valve causing a regular stream of water to issue from the gill-chamber, and thus occasioning the entrance of a constantly-fresh supply. In these, also, we find a dilatation, the walls of which seem to have contractile powers, at the commencement of each artery that dis- tributes the blood to the gills; and this connects the venous blood from the various channels, in Avhich it has meandered through the body. It is by the enclosure of the gills within a cavity, and by the consequent protection of them from the drying influence of the air (which would prevent their function from being duly performed), that Crabs and other allied species are enabled to live for a considerable time out of water; and the Land-Crabs, as they are termed, are adapted to spend the greater part of their lives at a distance from the sea, by means of a special glandular apparatus within the gill-cavity, which secretes a fluid that preserves the surface of the gills in the moist condition requisite for the aeration of the blood through its membrane. Thus the Land-Crabs are RESPIRATION IN INSECTS AND SPIDERS. 367 air-breathing animals (except at certain seasons, when they frequent the sea-shores), although they breathe by gills. 659. In Insects and other proper air-breathing Articulata, however, the character of the respiratory apparatus is very different. The tran- sition from one form to the other is effected through such animals as the Leech and the Earthworm, which seem able to live almost equally well in air or water, and whose respiration appears to be carried on chiefly, if not entirely, through the medium of the external surface alone. These animals are furnished with a series of small sacs, disposed at regular intervals along each side of the body, and opening by a row of pores, which are termed spiracles or stigmata; but these sacculi do not seem to participate in the respiratory function, their office being rather to secrete a protective mucus. But in the Myriapods, these sacculi are respiratory organs, and communicate more or less freely with each other. And in Insects, the spiracles, instead of forming the entrances to so many distinct sacs, open into a pair of large tubes, one of which tra- verses the body on either side, along its whole length. These tubes, termed tracheoe, have many communications with each other across the body ; and they branch out into innumerable prolongations, the ultimate ramifications of which are distributed to every portion of the system. They occasionally present dilatations of considerable size (Fig. 102, a); especially in the thoracic region of the body, in those insects which are endoAved with great powers of flight. These dilatations or air-sacs appear destined to serve as reservoirs of air, during the time that the insect is upon the wing, its spiracles being then partially closed ; and they may also be useful in diminishing the specific gravity of the body. The air- tubes are prevented from having their cavity obliterated through the pressure of the surrounding parts, by means of an elastic spiral fibre; Avhich winds round them, between their outer and inner membrane, from one extremity to the other (Fig. 102, b) ; and which answers the purpose of the cartilaginous rings and plates, in the trachea and bronchi of air-breathing Vertebrata. Fig. 102. A Respiratory apparatus of Insects : a, air vesicles and part of tracheal system of Scolia horlorum. b, por- tion of one of the great longitudinal trachese of Carabus auratus, with one of its spiracles. 660. In this manner, the air that is introduced through the spiracles is carried into every part of the body, and is brought into immediate relation with the tissues to be aerated; so that the carbonic acid which 368 OF RESPIRATION. they set free is communicated at once to the atmosphere, instead of being taken up by the blood; and the oxygen they require is imbibed in the same manner. And thus we see how the respiration of this inte- resting class, which is unequalled for its energy when thebody is in a state of activity, is provided for without an active circulation of blood, and Avithout the presence of red corpuscles,—Avhich elsewhere seem to be essential conditions of the interchange of oxygen and carbonic acid between the air and the tissues, wherever this takes place to any great extent. 661. In the Spider tribe, we return to a more concentrated form of the respiratory apparatus; but, notAvithstanding that it is limited within much narrower dimensions externally, it exposes a very large amount of surface on its interior. It consists of a series of sacs, much less numerous than in the lower Articulata, and not communicating with each other. Their lining membrane, however, is doubled into a series of folds, which lie in proximity with each other, like the leaves of a book, and which thus present a very extensive surface within a very small space. Over this surface the blood is distributed in a minute capillary network; and thus it comes into immediate relation with the air, which is received into the cavity through its aperture or spiracle. The alternate admission and expulsion of air seem to be provided for, as in Insects, by movements of the body, which first empty the cavities or air-tubes by compression, and then allow them to be refilled by their own elasticity, the pressure being relaxed. The respiratory cavities in the Spider-tribe have received the name of pulmonary branchiar, from their analogy, on the one hand, with the lungs of higher animals, and, on the other, with the branchial sac or gill-cavity of the higher Crustacea, the gills in which are formed by prolongations of the lining membrane, like the leaf-like folds in the air-cavities of the Spider-tribe. 662. The accompanying diagram will giAre an idea of the relations of these different forms of the respiratory apparatus, amongst themselves, and to that of Vertebrata. Let the line A B represent the general Fig. 103. Diagram illustrating different forms of the Respiratory apparatus: a, simple leaf-like gill; b, simple respiratory sac; c, divided gill; d, divided sac; e, pulmonary hranchia. surface of the animal; the continuations of that line on its upper side being its external prolongations; and those below, its internal prolon- gations or reflexions. Now at a is seen the character of the simple foliaceous or leaf-like gill, such as is found in the lower aquatic animals; presenting merely a flat expanded surface in contact with the water, over which the blood may be distributed. At b is shown a correspond- RESPIRATION IN FISHES. 369 ingly simple inversion; such as that which forms the respiratory sac of the Leech, having the blood-vessels distributed upon its walls. A higher form of the gill, such as is found in Fishes and in the higher aquatic Invertebrata, is seen at c, the surface being greatly extended, by subdivision into minute filaments. A more complex form of the pulmonary apparatus, such as is found in the higher Vertebrata, is shown at d, the blood being distributed, not merely to its outer Avails, but to the minute partitions which subdivide its cavity into cells. And at e is represented the respiratory organ of the Spider-tribe, which bears an obvious resemblance to the lung of the Vertebrated animal, shown at d; whilst it is evidently as nearly allied to the gill shown at c, provided this be imagined to be sunk within a cavity formed by a depression of the external surface, instead of projecting beyond it.— Thus we see how very close is the real resemblance between all the forms of the respiratory apparatus, however unlike each other they may at first sight appear to be. 663. Thegills of Fishes correspond with those of the higher Mollusca in all essential particulars; but they are more largely developed in pro- portion to the size of the body; and they are placed in a situation, that enables them to receive a more regular and constantly-changed supply, Fig. 104. Capillary network of a pair of [leaflets of the gills of the Eel:—a, a, branches of the branchial artery con- veying venous blood; b, b, branches of the branchial vein, returning aerated blood. The disappearance of the dark shading of the network, as it traverses the gill, is designed to indicate the change in the character of the blood, as it passes from one side to the other. both of blood and water. The gills are suspended to bony ,or cartilagi- nous arches, of which three, four, or more, are fixed on either side of the neck ; and the fringes hang loosely within the cavity, which commu- nicates on the one hand with the mouth, and on the other with the ex- terior of the body. The mechanism of respiration is very complex in these animals; and is evidently adapted to produce the most effectual aeration possible. The mouth is first distended with water; and its muscles are then thrown into contraction, in such a manner as to expel the water, through the aperture on either side of the pharynx, into the gill-cavity. At the same time, the bony arches are lifted and separated from each other, by the action of muscles especially adapted to this purpose; so that the gill-fringes may hang freely, and may present no obstacle to the flow of the Avater betAveen them. When they have been thus bathed with the aerating liquid, and their blood has undergone the necessary change, the water is expelled through the outward aperture on each side of the back of the neck ; which is furnished with a large flap or valvular cover, termed the operculum. In some of the cartila- 370 OF RESPIRATION. ginous Fishes, each branchial arch is enclosed in a separate cavity; which communicates on the inner side with the pharynx by an orifice peculiar to itself, and by another orifice with the external surface. Thus there is a series of external openings, instead of a single one, on each side of the neck ; and these sometimes amount to six or seven, as in the Lamprey, reminding us of the spiracles of Articulated animals; whilst there is a corresponding series of internal openings in the pha- rynx on either side, or into a tube that communicates with it. 664. It is well known, that most Fishes speedily die when removed from the water; and it can be easily shown, that the deficient aeration of the blood is the immediate cause of their death. But as it might have been expected, that the atmosphere Avould exert a much more energetic influence upon the blood contained in the gills, than that which is exercised by the air contained in the Avater, the question naturally arises, how this deficient aeration comes to pass. It is chiefly due to the two following causes:—the drying-up of the membrane of the gills themselves, where it is exposed to the air, so that the aeration of the blood is impeded ;—and the flapping together of the filaments of the gills, which no longer hang loosely and apart, but adhere in such a manner as to prevent the exposure of the greater portion of their surface to the air. Those fishes can live longest out of water, in Avhich the external gill-openings are very small so that the gill-cavity may be kept full of fluid; and there are certain species which are provided, like the Land-crab, with a particular apparatus for keeping the gills moist, and which perform long migrations over land in search of food even (it is said) ascending trees. These are exceptions to the general rule. 665. The respiration of Fishes is much more energetic than that of any of the lower aquatic animals ; and this is partly due to the great extension of the surface of the gills, partly to the provision just ex- plained for maintaining a constant Aoav of fresh water over their sur- face, and partly to the position of the heart at the base of the main trunk that conveys the blood to the gills (§ 588), by which the regular propulsion of that fluid through these organs is secured. Their blood too, is furnished with red corpuscles ; which give important aid in con- veying oxygen from the gills to the remote tissues of the body, and in returning the carbonic acid to be excreted. The proportion of these varies^ considerably, in the different species of the class, being very small in those that approach most nearly to the Invertebrata ; and there is even an entire absence of them in one remarkable fish, the Amphi- oxus or Lancelot; whilst they are present in large numbers in the blood of certain Fishes, which have great vascular activity, and can maintain a high independent temperature. Q6Q. It would seem, however, that not even this high amount of respiration is ahvays sufficient for Fishes which live in small collections of water, where their temperature is liable to be greatly augmented by the heat of summer ; under which condition, there is an increased prone- ness to disintegration in their tissues, and a corresponding necessity for the extrication of carbonic acid and for the absorption of oxygen. Many fresh-water fishes, under such circumstances, may be seen to come to the RESPIRATION IN REPTILES. 371 surface and to swallow air; and it would seem as if the interior of the intestinal canal then served the purpose of a respiratory surface, the air being expelled from the anus, deprived of a large part of its oxygen, and highly charged with carbonic acid. 667. In addition to their apparatus for aquatic respiration, many Fishes are provided, in their air-bladder, with the rudiments of the air- breathing apparatus of higher animals; although it is only in certain species, which approach Reptiles in their general organization, that this really affords any aid in the aeration of the blood. The air- bladder in its simplest condition is entirely closed; and it is then obviously incapable of taking any share in the respiratory function, although it seems to be an organ of some importance to the animal, in regulating its specific gravity, and altering its position in the water. In other cases, it communicates with the intestinal tube by a short, Avide canal, termed the ductus pneumaticus; and this may serve to admit air, Avhich is taken into the alimentary tube by the process of swalloAving just mentioned. In the Reptilian Fishes, just adverted to, the air-bladder forms a double sac, which is evidently the representative of the double lung of the air-breathing Vertebrata; and it communi- cates with the back of the mouth by a regular trachea or windpipe, which has a muscular valve at its commencement, serving to open or to close its orifice. Some of these fishes are able to live for a consider- able time out of wrater, their respiration being maintained by these rudimentary lungs; and they can also make a hissing sound, by the expulsion of the contents of the air-sacs through the narrow glottis, or entrance to the trachea. 668. The condition of the Respiratory apparatus, and the mode in which the function is performed in the class of Reptiles, are peculiarly interesting; as it is in this class, that we first meet with the complete adaptation of the Vertebrated structure to the aeration of the blood by the direct influence of the atmosphere. Their general habits of life require but a very feeble amount of aeration, especially at moderate temperatures ; their muscular and nervous systems being usually exer- cised in a very low degree; their movements being sluggish, and their perceptions obtuse. In fact, they may be considered, on the whole, as the most vegetative of all Vertebrated animals. In accordance with this character, the lungs are so constructed as not to expose any .very large amount of blood to the air at any one time; and, as we have already seen (§ 563), only a portion of the stream of the circulation is diverted to the lungs ; the main current being sent to the system, with only that amount of aeration, which it has derived from the admixture of the portion of blood that has been aerated in the lungs, with the A'enous current that has last been returned from the system. 669. The lungs of Reptiles are, for the most part, capacious sacs, occupying a considerable part of the cavity of the trunk; but they are very slightly subdivided, so that the amount of surface they can expose is really small. Where any subdivision exists, it is usually at the upper extremity of the lung, near the point of entrance of the bronchial tube; and where there is no actual subdivision of the cavity, we usually find that its surface is extended in this situation, by the formation of a number of little depressions or pouches in its walls, upon which the 372 OF RESPIRATION. Fig. 105. Section of the Lung of the Turtle. blood-vessels are minutely distributed. The greatest amount of subdi- vision is seen in the lungs of the Turtle tribe; but even in these, the partitions scarcely form a complete division at any part of the lungs; and the ultimate air-cells are of very large size (Fig. 105). The air- sacs of Reptiles are not filled, like those of Mammalia, by an act of inspiration, but hy a process of swallowing, which is comparatively tedious ; and, from the small amount of aerating surface, in proportion to the amount of air thus received into the cavity, one inflation of the air- sacs lasts for a considerable time. When the replacement of oxygen by carbonic acid has proceeded to an extent that renders the air no longer fit to remain in the lungs, these cavities are emptied by pressure exercised upon them by the muscles of the trunk ; and the slow exit of the air through the narrow glottis is accom- panied by a prolonged hissing sound, which is the only sort of voice that is possessed by the greater part of the Reptile class. The lungs are again filled by the swallowing-process; and all goes on as before. 670. Now in the Frog tribe, which forms the lowest order of Reptiles (and which is sometimes ranked as a distinct class, under the title of Amphibia), the respiration during the early or Tadpole state, is aquatic; being carried on by means of gills, and conducted exactly upon the plan of that of Fishes. The lungs are not developed, until a period long subsequent to the ani- mal's emersion from the egg; and as soon as they are ready to come into play, an alteration begins to take place in the circulating system, by which the current of blood is diverted towards them, and away from the gills (§ 562). This change takes place to its full extent in the Frog, Toad, Newt, and their allies; which henceforth have a respiration and a circulation exactly analogous to that of Reptiles in general; but it is checked in the Proteus, Siren, and other species, Avhich form the peren- nibranchiate group,—so called from the persistent character of their gills, which still remain in action, the lungs never being sufficiently de- veloped to maintain the respiration by themselves. The curious influ- ence which Light possesses on this metamorphosis, has been already referred to (§ 95). 671. This order Batrachia is further distinguished from other Rep- tiles, even when the metamorphosis is complete, by the softness and nakedness of the skin, which is destitute of the scales and horny plates that cover it in the Lizards, Serpents, and Tortoises. The skin of the Frog tribe is a very important organ of respiration, being richly sup- plied with blood-vessels, and exposing their contents to the influence of the air, under circumstances nearly as favorable as those afforded by the imperfectly-developed lungs of these animals. Thus a Frog, from which the lungs have been removed, will live a considerable time at a moderate temperature, if its skin be freely exposed to a moist air; for, in consequence of the peculiar mode in Avhich the circulation is carried RESPIRATION IN REPTILES AND BIRDS. 373 on in these animals (§ 561), the interruption to the flow of blood through the lungs does not, as in the higher classes, produce a stagna- tion of the general current through the body; and the blood receives, in its course through the skin, a sufficient amount of aeration for the support of life. Indeed, at a low temperature, the influence of water on the skin is sufficient (by means of the air included in the liquid) to remove the small amount of carbonic acid then ready for excretion, and to supply the requisite amount of oxygen; and Frogs may thus live beneath the water for any length of time, without coming to the surface to breathe.^ But with the rise of the temperature of their bodies, their blood requires a higher degree of aeration; and they then come to the surface to take in air by the mouth. Avhich aerates the blood through the lungs. It appears that, during the heat of summer, the pulmonary respiration, and the influence of the water on the skin, are not sufficient; as it is found that Frogs die, if they are confined to the water under such circumstances,—their natural habit being to quit the water at such times, so that the air may exert its full influence on their skin as Avell as on their lungs. They do not, however, quit the neighborhood of water, and soon die if exposed to a dry atmosphere, for, if the skin become dry, its aerating function can be no longer performed. The same result happens if the passage of gases through the skin be impeded by smearing it over with any unctuous substance. We shall presently find reason to believe that this cutaneous respiration is a very important part of the function, even in Man and Mammalia. 672. The class of Birds presents a most striking contrast to that of Reptiles, in regard to the energy of the respiratory function, and the extent of the apparatus destined to its performance. The air-cells are considerably diminished in size, so that the extent of surface over which they expose the blood to the air is greatly increased; and there even seems reason to believe that the air comes into direct contact Avith the vessels of the very close capillary plexus which intervenes between the air-cells. But the subdivision of the lungs is not carried to the same degree of minuteness as it is in Mammalia; and the required extent of surface would not be afforded by the lungs alone. In addition to these organs, we find large air-sacs, communicating with them, disposed in different parts of the body,—such as the abdominal cavity, the inter- spaces among the muscles, the spaces betAveen the muscles and the skin, &c. These very greatly increase the respiratory surface, their lining membrane being extremely vascular, and adapted to expose the blood to the influence of the air. In most Birds, the bones themselves are hollow, and the lining membrane of their cavities serves as an addi- tional aerating surface, the air being introduced into the interior of the bones, by canals that communicate directly with the lungs. So free is this communication, that the respiration has been known to be main- tained through the fractured humerus of an Albatross, when an attempt was made to destroy the bird by compressing its trachea. Thus the respiratory surface is extended into the remoter parts of the system, very much as in Insects; and the hollowness of the bones, together with the presence of numerous air-sacs in different parts of the body, contribute to diminish its specific gravity. The large quantity of air 374 OF RESPIRATION. thus included in different portions of the frame, also serves, like that contained in the air-sacs of Insects, as a reservoir for the supply of the principal aerating organs during active flight, when the respiratory movements are less free. 673. The mechanism of Respiration in Birds is very different from that which produces the respiratory movements in Mammalia. The cavities of the chest and thorax are not yet separated by a diaphragm ; except in a very small number of species, that approach most nearly to the next class. But, on the other hand, the whole cavity of the trunk is more completely enclosed in a bony casing, the ribs being connected with the sternum by osseous prolongations from the latter, instead of by cartilages, and the sternum itself being so largely developed, as to cover almost the entire front of the body. Now the natural condition of this bony framework is such, that, when no pressure is made upon it, the cavity it encloses is in a state of distension; and the state of emptiness can only be produced by a forcible compression of the frame- work, through an exertion of muscular power. The lungs, instead of being freely suspended in the cavity of the chest, as in Mammalia, are attached to the ribs; and their OAvn tissue is endoAved with a degree of elasticity, which causes them to dilate when they are permitted to do so. In the state of distension, therefore which is natural to the cavity of the trunk, the lungs are expanded, and fill themselves with air, which they draw in through the trachea; and this condition they retain, until, by the action of the external muscles upon the bony framework, the cavity of the trunk is diminished, and the air is expelled from the lungs and air-sacs, which are again filled as soon as the pres- sure is taken off. As the air-sacs chiefly communicate with the part of the lungs that is most distant from the trachea, the air has to traverse the whole extent of those last organs, both when it is being drawn into the air-sacs, and when it is being expelled from them; so that it is made to serve for the aeration of the blood in the most effectual manner. 674. Thus, although the respiratory apparatus of Birds does not possess the highly concentrated development which we shall find it to present in Mammals, it serves, by the extension of the aerating surface through the body, to bring the air and the blood into most intimate relation; and the energy of the function is further provided for, by the mode in which the pulmonary circulation is carried on (a distinct heart, as it Avere, being provided for it, § 564), as well as by the arrangement of the blood-vessels, which transmit to the respiratory organs the whole of the blood that has been returned in a carbonated state by the great veins of the system. The very large proportion of red corpuscles con- tained in the blood gives additional effect to these provisions. The very high amount of respiration which is natural to Birds, and which cannot be suspended even for a short time without fatal consequences, has a direct relation (as already explained) with their extraordinary muscular activity, as well as with the high bodily temperature which they are fitted to maintain, and which cannot be lowered in any great degree without the suspension of their other functions. Birds are peculiarly susceptible of impurities in the atmosphere; and it has been shown by experiment, that if a Bird, a Mammal, and a Reptile, be RESPIRATION IN BIRDS AND MAMMALS. 375 placed together in a limited quantity of air, which gradually becomes vitiated by their respiration, the Bird will die first, the Mammal next, and the Reptile last. Or if the Bird be placed alone in a limited quantity of air, and be left until the atmosphere is so vitiated as to be no longer capable of supporting its life, a Mammal will still live for a time in that atmosphere; and when it is no longer fit to sustain the life of the Mammal, the Reptile may still breathe it without injury for a considerable period. There is strong reason to believe, indeed, that, in former epochs of the Earth's history, when the Reptile class was predominant, supplying the place of Mammals on land, and of Birds in the air, the atmosphere was so highly charged writh carbonic acid, as not to be capable of sustaining the life of the higher air-breathing Ver- 3. Mechanism of Respiration in Mammalia and in Man. 675. It is in the class of Mammalia that we find the Respiratory apparatus presenting its highest degree of concentration; and the arrangements for its action the most complete. The Lungs are divided into cavities of extreme minuteness, so that the extent of surface Avhich they expose is enormously increased. These cavities, or air-cells, are all connected with the trachea by means of the bronchial tubes and their minute subdivisions; but, on account of the minuteness of these passages, a considerable force would be required to inflate the air-cells with air, if their distension Avere to be accomplished by the propulsion of air through the trachea, as we have seen to be the normal mode of inspiration in Reptiles. Moreover, if the air were introduced in this manner, the air-cells would be the last portions of the pulmonary struc- ture that would be distended by it, as well as the first to be emptied Avhen the air is forced out again by external pressure. The mechanism of Respiration in Mammalia, however, is so arranged, that the air is most effectually drawn into the lungs, instead of being forced into them; and the distension of the air-cells is far more complete than it could be rendered in the latter method, besides being accomplished in a much shorter time. 676. The general principle of the operation is this. The lungs are suspended in a cavity that is completely closed, being bounded above and around by the bony framework of the thorax, the interspaces of which are filled up by muscles and membranes, and being entirely cut off from the abdomen below by the diaphragm. Under ordinary cir- cumstances, the lungs completely fill the cavity; their external surface, covered by the pleura, being everywhere in contact with the pleural lining of the thorax. But the capacity of the thoracic cavity is sus- ceptible of being greatly altered by the movements of the ribs, and by the actions of the diaphragm and abdominal muscles, as will presently be explained in more detail. When it is diminished, the lungs are compressed, and a portion of the air contained in them is expelled through the trachea. On the other hand, when it is increased, the elasticity of the air Avithin the lungs causes them immediately to dilate, so as to fill the vacuum that would otherwise exist in the thoracic 376 OF RESPIRATION. cavity; and a rush of air takes place down the air-tubes, and into the remotest air-cells, to equalize the density of the air they include (which has been rarefied by the dilatation of the containing cavities) Avith that, of the surrounding atmosphere. 677. The diameter of the ultimate air-cells of the Human lung varies from about the l-200th to the l-70th of an inch. Their shape is irregular, and their walls are, for the most part, flattened against each other. Each of the ultimate ramifications of the bronchial tubes communicates with a cluster of these air-cells grouped around it; those which are in immediate proximity with the tube open into it oy well- defined circular apertures, and the others communicate with it by open- ing into these and into each other. Each air-cell is lined by an exten- sion of the mucous membrane from the bronchial tubes; but this does not seem to be furnished with an epithelial covering. Between the adjacent air-cells, is a network of fibrous tissue, that forms the connect- ing medium by which they are held together; this tissue appears to be of the elastic kind. The pulmonary arteries subdivide into branches, whose ultimate ramifications form an extremely minute capillary plexus; and this is disposed betAveen the walls of the adjacent air-cells, so that each portion of this plexus comes into relation with the air (through the lining membrane of the contiguous air-cells) on both sides,—an arrange- ment which is obviously the most favorable that can be to the aeration of the contained blood. It has been calculated by M. Rochoux, that the number of air-cells grouped around each terminal bronchus is little less than 18,000; and that the total number in the lungs amounts to six hundred millions. If this estimate be even a remote approximation to the truth, it is evident that the amount of surface exposed by the walls of these minute cavities, must be very many times greater than that of the whole exterior of the body. Fig. 106. Arrangement of the Capillaries of the air-cells of the Human Lung. 678. The larger bronchial tubes are more or less cartilaginous; but the smaller branches do not possess any such deposit in their walls, though still retaining their circular form. We find in the latter a fibrous structure, which seems to possess the properties of non-striated muscle; and by this, the diameter of these tubes appears to be governed. The STRUCTURE OF THE LUNGS IN MAN. 377 contractility of the walls of the smaller bronchi may be excited by chemical, electrical, or mechanical stimuli applied to themselves ; though it is not so readily caused to manifest itself by stimulating the nerves. By the continued influence of galvanism, bronchial tubes of a line in diameter have been made to contract, until their cavity Avas nearly oblite- rated ; and it has been found by Volkmann that a similar effect may be produced by galvanising the Par Vagum. Supposing the muscular fibres of the bronchial tubes to contract during expiration, the effect of such contraction would be to diminish both the length and the diameter of the tubes, and thus to force out their contained air. Whether such contraction, alternating with relaxation, takes place automatically, as a part of the ordinary rhythmical movements of respiration, has not yet been clearly made out; but in its tonic form, it manifests itself strongly in certain diseased conditions, especially in spasmodic Asthma, which appears essentially to consist in a contracted state of the smaller bronchial passages, occasioning an interruption to the passage of air through them. It is interesting to observe, that the contractility of the muscular walls of these tubes has been experimentally found to be greatly diminished by the application of vegetable narcotics, especially stramonium and belladonna,—substances which are well-known to have a powerful remedial influence in spasmodic Asthma. 679. The Lungs themselves appear to be, almost entirely, passive instruments of the Respiratory function. Their contraction when over- distended, and their dilatation after extreme pressure, may be partly due to the elasticity of their structure ; which seems to produce, Avhen acting by itself, a moderately-distended state of the air-cavities. This, too, is the condition that seems most natural to the cavity of the chest; the fullest dilatation, or the most complete contraction, of which it is capa- ble, being only accomplished by a forcible effort. 680. The dilatation of the cavity of the chest, which constitutes Inspiration, is accomplished by two sets of movements;—the elevation of the ribs, and the depression of the diaphragm. From the peculiar mode in which the ribs are articulated with the spinal column at one extremity, and from the angle which they make with the cartilages that connect them to the sternum at the other, the act of elevation tends to bring the ribs and the cartilages more into a straight line, and to carry the former to a greater distance from the median plane of the body, whilst the sternum is also throAvn fonvards. Consequently the eleva- tion of the ribs increases the capacity of the thorax, upwards, fonvards, and laterally. The moArement is chiefly accomplished by the Scaleni muscles, Avhich draw up the first rib; and by the Intercostals, which draAv the other ribs into nearer proximity with each other, so that the total amount of movement in each rib increases as we pass from above downwards,—every one being drawn up by its connexion with the one above it, and being drawn nearer to it by the action of its own intercostals. The elevation of the ribs is further assisted by the Ser- ratus magnus, and by other muscles connected Avith the spine and the scapula; and when the respiratory movement is very forcibly performed, the scapula is itself drawn upwards, by the muscles that descend to it from the neck, thus producing an increased elevation of the ribs, and 378 OF RESPIRATION. an unusual enlargement of the upper part of the thoracic cavity.—When the Expiratory action is to be performed, the descent of the rib is occasioned by the muscles of the spine and abdomen, which proceed upwards from the loAver part of the trunk ; and this action is aided by the elasticity of the costal cartilages. 681. In the ordinary act of inspiration, however, the Diaphragm performs the most important part. The contraction of this muscle changes its upper surface, from the high arch that it forms Avhen relaxed and pushed upwards by the viscera below, to a much more level state; though it never approaches very closely to a plane; being some- what convex, even when the fullest inspiration has been taken. When thus drawn down, it presses upon the abdominal viscera, and causes them to project forwards, which they are allowed to do, by the relaxa- tion of the abdominal muscles. In tranquil breathing, this action is alone nearly sufficient to produce the requisite enlargement of the tho- racic cavity ; the position of the ribs being very little altered. In the expiratory movement, the diaphragm is altogether passive; for, being in a state of relaxation, it is forced upwards by the abdominal viscera, which are pressed inwards by the contraction of the abdominal muscles. These last, therefore, are the main instruments of the expiratory move- ment; diminishing the cavity of the chest by elevating its floor, at the same time that they draw its bony framework into a narrower com- pass. 682. In this manner, by the regularly-alternating dilatation and con- traction of the thoracic cavity, the air within the lungs is alternately in- creased and diminished in amount; and thus a regular exchange is secured. This exchange, however, can only affect at any one time a certain proportion of the air in the lungs ; thus it is probable, that the quantity remaining in these organs after ordinary expiration is above 100 cubic inches, whilst the amount usually expired is not above 20 cubic inches. Indeed if it were not for the tendency of gases to mutual diffusion, the air in the remote air-cells might never be renewed.—If any aperture exist, by which air could obtain direct access to the pleural cavity, the lungs would not be dilated by its enlargement; for the vacuum would be supplied much more readily, by the direct ingress of the air (provided the aperture be large enough), than by the distension of the lung. Thus a large penetrating wound of the thoracic cavity may completely throw out of use the lung of that side ; and the same result will follow, when an aperture forms by ulceration in the sub- stance of the lung itself, establishing a free communication betAveen the pleural cavity and one of the bronchial tubes; so that, of the air which rushes in by the trachea, to compensate for the enlargement of the tho- racic cavity, a great part goes at once into that cavity, without con- tributing to the distension of the lungs, and therefore without semng for the aeration of the blood. 683. The number of the respiratory movements (that is, of the acts of inspiration and expiration taken together) may be probably estimated at from 14 to 18 per minute, in a state of health, and of repose of body and mind. Of these, the greater part are moderate in amount, involv- ing little movement except in the diaphragm; but a greater exertion, , MECnANISM OF RESPIRATION IN MAN. 379 attended with a decided elevation of the ribs, is usually made at every fifth recurrence. The frequency of the respiratory movements, hoAv- ever, is liable to be greatly increased by various causes, such as violent muscular exertion, mental emotion, or quickened circulation ; whilst it may be diminished by the torpidity of the nervous centres, on whose agency the movement depends,—as we see in apoplexy, narcotic poison- ing, &c. An acceleration seems very constantly to take place in dis- eases, which unfit a part of the lung for the performance of its func- tion ; and the rate bears a proportion to the amount thus thrown out of use. Thus, the usual proportion between the respiratory movements and the pulse being as 1 to 4-£ or 5, it may become in Pneumonia as 1 to 3, or even in severe cases 1 to 2 ; the increase in the number of respiratory movements being much greater in proportion, than the aug- mentation of the rate of the pulse. But it must be remembered by the practitioner, that a simply hysterical state may produce, in young females, an extraordinary acceleration of the respiration; the number of movements being sometimes no less than 100 per minute. There will be a great increase, also, in the number of inspirations, when the regular movements are prevented from being fully performed, by any cause that affects their mechanism, even whilst the lungs themselves are quite sound. Thus in inflammation of the pleura, or pericardium, or in rheumatic affections of the intercostal muscles, the full action of the ribs is prevented by the pain which the movements produce ; and the same is the case in regard to the diaphragm, when the peritoneum or the abdominal viscera are affected with inflammation. Under such circumstances, there is an involuntary tendency to make up for the de- ficiency in the amount of the respiratory movements, by an increase in their number. 684. The combined actions of the respiratory muscles, which have been noAV explained, belong to the group termed reflex ; being the result of the operation of a certain part of the nervous centres, which does not im'olve the will, or even sensation, and which may continue when all the other parts of the nervous centres have been removed. In the Invertebrated Animals, we commonly find a distinct ganglionic centre set apart for the performance of the respiratory movements ; and the division of the nervous centres in Vertebrated animals, which is the seat of the same function, may be clearly marked out, although it is not so isolated from the rest. It is, in fact, that segment of the Medulla Oblongata and upper part of the Spinal Cord, wrhich is connected with the 5th, 7th, and 8th pairs of cephalic nerves, and with the phrenic. The entire brain may be removed from aboAre (by successive slicing), and the whole spinal cord may be destroyed beloAV; and yet the respi- ratory movements of the diaphragm will still continue,—those of the intercostal and other muscles being of course suspended, by the destruc- tion of that portion of the cord from Avhich their nerves arise. But if the spinal cord be divided, betAveen the point at which it receives the 5th and 8th pairs of nerves, and that at which it gives origin to the phrenic, the movements of the diaphragm immediately cease; and this is the reason Avhy death is so instantaneous, in cases of luxation or frac- ture of the higher cervical vertebrae, causing pressure upon the spinal 380 OF RESPIRATION. cord just beloAV its exit from the cranium; whilst if the injury take place below the origin of the phrenic nerve, life may be prolonged for some time. 685. The Respiratory movements, like other reflex actions (§ 394), depend upon a stimulus of some kind, originating at the extremities of the nerves, propagated towards the centre by the afferent trunks, and giving rise to a motor impulse, which is transmitted along the efferent or motor nerves to the muscles, and which occasions their contraction. Now the importance of the respiratory function to the maintenance of life which has already been sufficiently pointed out, necessitates an ample provision for its due performance ; and thus we find that the stimulus for the excitement of the movements may be transmitted through several channels. Its chief source, no doubt, is in the lungs; and arises from the presence of venous blood in the capillaries, and of carbonic acid in the air-cells. Under ordinary circumstances,—that is, when the blood is being duly aerated, and the air being properly renewed, —the impression thus made upon the nerves of the lungs is so faint, that we cannot perceive it, even when we specially direct our attention to it. But if we suspend the movements for a moment or two, we immediately experience a sensible uneasiness. The Par Vagum is obviously the channel, through which this impression is conveyed to the nervous cen- tres ; and it is found that, if the trunk of this nerve be divided on both sides, the respiratory movements are greatly diminished in frequency. Hence it is undoubtedly one of the principal excitors of the respiratory movements. 686. But the sensory nerves of the general surface, and more parti- cularly the sensory portion of the Fifth pair, which supplies the face, are most important auxiliaries, as excitor nerves ; the inspiratory move- ment being peculiarly and forcibly excited by impressions made upon them, especially by the contact of cold air or water with the face. The poAver of the impression made by the air upon the general surface, and particularly upon the face, in exciting the inspiratory movement, is well seen in the case of the first inspiration of the new-born infant, which appears to be excited solely in this manner. An inspiratory effort is often made, as soon as the face has emerged from the Vagina of the mother; whilst, on the other hand, if the face be prevented from coming into contact Avith cool air, the inspiratory effort may be wanting. When it does not duly take place, it may often be excited by a slap with the flap of the hand upon the nates or abdomen; a fact which shows the special influence of impressions upon the general surface, in rousing the motor impulse in the Medulla Oblongata, and in causing its transmission to the muscles. The deep inspirations which follow a dash of cold water upon the face, or the descent of the cold douche or of the divided streams of the shower-bath upon the body, or the shock of immersion in the cold plunge-bath, all testify to the powerful influence of such impressions in the adult; and the efficacy of other kinds of irritation of the skin, such as beating with holly- twigs, in maintaining the respiratory movements in cases of narcotic poisoning, shows that the required impressions are not restricted to the contact of cold air or water. It seems probable, from various facts, REFLEX CHARACTER OF THE RESPIRATORY MOVEMENT. 381 that the presence of venous blood in the arterial capillaries of the system, and the consequent stagnation in the current through them (§ 597), may exert an influence through the Sympathetic nerves: which may be transmitted, by the copious inosculations of that system with the Par Vagum, to the Medulla Oblongata; and which may there serve as a valuable auxiliary in exciting the respiratory movements. 687. Of the mode in which the impressions, thus transmitted to the Medulla Oblongata, act in exciting the motor impulses which issue from it, nothing is known; but these impulses, directed along the phrenic, intercostal, and other nerves, produce the requisite movements. When the stimulus is unusually strong, various nerves and muscles are put in action, which do not co-operate in the ordinary movements of inspiration ; and it; may sometimes be observed, that movements are thus excited in parts, which will not act in obedience to the will, being to all appearance completely paralysed. This fact shows how com- pletely the class of actions in question is independent of the influence of the mind; but we must not lose sight of the control which the mind, especially in the higher classes of animals, possesses over them. Va- rious actions of the respiratory muscles, particularly those of weeping and laughing, are the most direct means of expressing the passions and emotions of the mind; and are involuntarily excited by these. And, again, the respiratory actions are placed in a certain degree under the control of the Will; in order that they may be subservient to the pro- duction of vocal sounds, and to the actions of speech, singing, &c. The Avill cannot long suspend the respiratory movements; for the stimulus to their involuntary performance soon becomes too powerful to be any longer resisted. And it is well that it should be so ; for if the perfor- mance of this most important function were left to our own choice, a few moments of forgetfulness would be productive of fatal results. But it is to the power which the will possesses, of directing and controlling the respiratory movements, that Ave owe the faculty of producing arti- culate sounds, and thus of holding the most direct and intimate con- verse with each other. 688. It is essential for the due performance of the respiratory move- ments, that the portion of the nervous centres, on Avhich they depend, should be in a state of activity. This is the case, under ordinary cir- cumstances, throughout life. The state of perfect quiescence, to which the Brain is liable, never affects the Medulla Oblongata; and the respi- ratory movements are consequently kept up with as much regularity and energy (in proportion to the requirements of the system), during our sleeping, as during our Avaking hours. But if any cause induce torpidity of the medulla oblongata, the respiratory movements are then retarded, or even suspended altogether; and all the consequences of the cessation of the aeration of the blood speedily develope themselves (§ 706). This is seen in apoplexy ; when the pressure, or other cause of suspended activity, which at first affected the brain alone, gradually propagates its influence downwards. The same is the case in narcotic poisoning ; in which also the brain is the first to be affected, and may suffer alone; but if the noxious influence be propagated to the medulla oblongata, it manifests itself in the retardation of the respiratory move- 382 OF RESPIRATION. ments, and, when sufficiently poAverful, in their complete suspension. Under such circumstances, it is requisite to resort to all possible means of keeping up the respiratory movements; and when these fail, arti- ficial respiration may be successfully employed. For if, by such means, the circulation can be prevented from failing for a sufficient length of time, the ordinary processes of nutrition go on, the poisonous matter is gradually decomposed, or eliminated by the secreting organs; and the nervous centres resume their usual functions. A torpid condi- tion of the medulla oblongata, inducing a retardation of the respiratory movements, seems to be one of the morbid conditions attendant upon typhoid fever; and probably depends in the first instance upon a disor- dered state of the blood, which does not exert its usual vivifying influ- ence. In such cases, the proportion of the respiratory movements to the pulse sinks as low as 1 to 6, or even 1 to 8; and thus the due aera- tion of the blood is not performed, and its stimulating properties are still further diminished. 4. Chemical Phenomena of Respiration. 689. Having now fully considered the means, by Avhich the Atmo- sphere and the Blood are brought into relation in the lungs, we have to examine into the results of their mutual action. It will be remem- bered that the Atmosphere contains about 21 per cent, of Oxygen to 79 of Nitrogen, by measure; or 23 parts of Oxygen to 77 of Nitrogen, by weight. The changes which it undergoes in Respiration may be considered under four heads:—1. The disappearance of Oxygen, which is absorbed. 2. The presence of Carbonic Acid, which has been exhaled. 3. The absorption of Nitrogen. 4. The exhalation of Nitrogen. Of these, the first two are by far the most important.—It Avas formerly supposed that the Oxygen which disappears, is the precise equivalent of the Carbonic Acid Avhich is set free (the latter gas containing its own bulk of the former); and that the union of the absorbed oxygen with the carbon to be eliminated, takes place in the lungs. It is now known, hoAvever, that the carbonic acid is given out ready formed, its production having taken place at the expense of oxygen preA'iously contained in the blood; and that a much larger proportion of oxygen is usually absorbed, than is contained in the carbonic acid exhaled, the difference sometimes exceeding the third part of the carbonic acid formed, [whilst it is sometimes so small that it may be disregarded. This diversity seems to depend, partly upon the constitution of the species experimented on, and partly upon the degree of development of the individual, but in great part upon the nature of the food; it having been established by the recent experiments of MM. Regnault and Reiset, that the quantity of oxygen absorbed into the system is much greater on an animal diet, than on a farinaceous. It is certain that, of this absorbed oxygen, a part must enter into combination with the sul- phur and phosphorus of the original components of the body, converting these into sulphuric and phosphoric acids ; and the remainder must enter into other chemical combinations, very probably uniting with the hydro- AMOUNT OF CARBONIC ACID EXHALED. 383 gen of the fatty matter, to form part of the Avater which is exhaled from the lungs. 690. This interchange would seem to depend upon the tendency which all gases have to mutual admixture, when they are separated by a porous septum. According to the law discovered by Prof. Gra- ham, the relative volumes of the gases which will thus replace each other, are inversely as the square-roots of their specific gravities; thus, the specific gravity of oxygen being to that of hydrogen as 16 to 1, the replacing volume of oxygen is to that of hydrogen as 1 to 4. The same holds good, when one of the gases is absorbed by a liquid; pro- vided the replacing gas be also capable of being absorbed to the same extent. On this principle, the replacing volume of oxygen is to that of carbonic acid as 1174 to 1000; but as the actual amounts inter- changed do not constantly follow this ratio, it is obvious that they are liable to be modified by other conditions, these being chiefly (it seems probable) the relative quantities of the two gases already present in the blood, and the relative facility with which they are absorbed into it or extricated from it. 691. It is difficult to form an exact estimate of the actual quantity of Carbon, thrown off from the lungs in the form of Carbonic Acid during any lengthened period: since the amount disengaged during experiments carried on for a limited time, cannot, for many reasons, be taken as affording a fair average. Thus the quantity will vary with the external temperature, with the state of previous rest or activity, Avith the length of time that has elapsed since a meal, and particularly with the general development of the body. The amount of carbonic acid exhaled is greatly increased by external cold; as is shown in the results of such experiments as the following.—Small Birds and Mam- mals having been enclosed in a limited quantity of air, for the space of an hour, at ordinary temperatures, the quantity of carbonic acid they produced was noted. The experiment Avas then repeated at a tempera- ture nearly approaching that of the body; and was performed a third time at a temperature of about 32°. The following are the comparative amounts. Temp. 59°—6S° Temp. 86°—106° Temp, ahout Grammes. Grammes. Grammes, A Canary, 0-250 0129 0-325 A Turtle-dove, 0-684 0-366 0-974 Two Mice, 0-498 0-268 0-531 A Guinea-pig, 2-080 1-453 3-006 Thus it Avould appear that the quantity of carbonic acid exhaled between 86° and 106° is not much more than half of that which is exhaled be- tAveen 59° and 68°; and is only about tioo-fifths of that which is given off at 32°. 692. The quantity of Carbonic Acid exhaled during exercise, and for a certain time after it, and also after a full meal, is considerably in- creased ; Avhilst on the other hand, it is greatly diminished during sleep. Thus a person who was excreting 145 grains of carbon per hour, whilst fasting and at rest, excreted 165 after dinner, and 190 after breakfast and a Avalk: whilst he only excreted 100 during sleep. The variation 384 OF RESPIRATION. with the general development of the body, and also with the sex and age, is considerable. Thus, the exhalation is almost always greater in males, than in females of the same age, at every period of life except childhood. In males, the quantity increases regularly from eight to thirty years of age, remaining nearly stationary until forty ;—thus it averages 77*5 grains of carbon per hour at eight years; 135 grains at fifteen; 176*7 grains at twenty; and 189 grains from thirty to forty. Between forty and fifty, there is a well-marked diminution, the average being then 156 grains; and the diminution continues up to extreme old age, when the amount exhaled scarcely exceeds that which is extricated at ten years of age; thus, .between sixty and eighty, it was 142*5 grains; and in a man of a hundred and two, it was only 91*5 grains. These average results, however, are widely departed from in individual cases; an extraordinary development of the muscular system being al- ways accompanied by a high rate of extrication of carbon; and vice versd. Thus a man of remarkable muscular vigor, whose age was twenty-six years, exhaled 217 grains of carbon in an hour; a robust man of sixty exhaled 209*4 grains; and an old man of ninety-tAvo, who in his younger days had possessed uncommon muscular power, and who preserved a remarkable degree of energy, still gave forth at the rate of 151 grains per hour. On the other hand, a man of slight muscular development, at the age of forty-five, only exhaled 132 grains; and an- other at the age of seventy-six, only 92-4 grains. 693. In females, nearly the same proportional increase goes on, up to the time of puberty; when the quantity abruptly ceases to increase, and remains stationary so long as menstruation continues regular. The average quantity of carbonic acid exhaled by girls nearly approaching puberty, is about 100 grains per hour ; and it remains at this standard until nearly the close of menstrual life. At the period of the cessation of the catamienia, it undergoes a perceptible increase; the average be- tAA'een forty and fifty years of age, being about 130 grains per hour; and the quantity exhaled in a woman of great muscular development, and of forty-four years of age, rising to 152-4 grains in an hour. After the age of fifty, or thereabouts, the quantity decreases, as in men. It is remarkable that, during pregnancy, there is the same increase in the exhalation of carbon, as there is after the final cessation of the cata- menia; and the same takes place, if the menstrual discharge be tem- porarily suspended, through any other cause. 694. It is obviously difficult, then, to obtain exact estimates, from any experiments conducted for a short time only, of the total amount of Carbon thrown off during a lengthened period; since the condition of the individual varies so greatly at different times; and the variation amongst different individuals is so great. Moreover, of the total amount of carbon excreted in a gaseous form, a certain part is undoubtedly set free from the skin; but the proportion of this does not seem to be con- siderable. As a means of measuring the whole quantity of carbonic acid set free, without causing the respiratory movements to be per- formed in any unnatural manner, Prof. Scharling constructed an air- tight chamber, of dimensions sufficient to allow an individual to remain in it for some time without inconvenience; and so arranged, that he AMOUNT OF CARBONIC ACID EXHALED. 385 could eat and drink, read, or sleep Avithin it. This was connected with an apparatus, by Avhich the air was continually renewed; and the air drawn off was carefully analysed, in order to determine the quantity of carbonic acid contained in it. The average per hour, in different states having been ascertained, it was calculated that, allowing seven hours for sleep in adults, and nine hours for children, the total amount of carbon consumed in the tAventy-four was as follows: No. Weighing. Grains. Oz. Troy. 1. A male, aged thirty-five,.....'131 lbs. 3387 or 7-0 2. A male, aged sixteen,......115^ lbs. 3453 or 7-2 3. A soldier, aged twenty-eight, . . . 164 lbs. ' 3699 or 7-7 4. A girl, aged nineteen,......Ill lbs. 2540 or 5-3 5. A boy, aged nearly ten,.....44 lbs. 2054 or 4-3 6. A girl, aged ten,.......46 lbs. 1930 or 40 695. This estimate is perhaps rather too low, as it does not take sufficient account of the great increase which is produced by exercise. Another method has been adopted by Prof. Liebig, who endeavored to ascertain the total amount of carbon excreted from the body in the form of carbonic acid, by comparing the amount of carbon taken in as food, with that contained in the faeces and urine ; the difference being set down to the account of respiration. His estimate amounts to the very large sum of 13*9 oz. of solid carbon per day, which he considers to be thus set free by the lungs and skin; but this is almost certainly above the truth. The observations were made upon a body of soldiers, who Avere subjected to severe daily exertion ; and they were far from being exactly conducted, many of the items being set doAvn by guess only, whilst of others no account whatever was taken. We may per- haps consider 10 or 11 oz. as more nearly representing the amount of carbon consumed by adult men exposed to severe exertion ; whilst from Prof. Scharling's experiments it may be inferred, that from 7 to 8 oz. of carbon are thrown off during the twenty-four hours, by the lungs and skin of adult men not using much active exertion ; to Avhich another ounce or tAvo may be added, as the increased quantity excreted during moderate exercise. In a subsequent series of experiments, Prof. Schar- ling ascertained the proportion of carbonic acid given off from the general surface of the human body, to be from about l-30th to l-50th of the whole amount; so that, adopting l-40th as the average, out of eight ounces of carbon exhaled, only one-fifth of an ounce is set free in the form of carbonic acid from the Skin. 696. If we assume 10 oz. or 4800 grains of solid Carbon, as the total amount excreted from the lungs and skin of a male adult, using active exercise, in the course of twenty-four hours, Ave find that the •^volume of carbonic acid thus generated must be nearly 37,000 cubic inches, or more than 21 cubic feet. Of this, about 16 cubic feet are probably extricated from the lungs. But it is probable, that about 10 cubic feet per day is nearer the ordinary average. Now it has been ascertained, that the whole quantity of air Avhich passes through the chest during that time under similar circumstances, is about 266 cubic feet; so that the proportion of carbonic acid contained in the expired air seems to average about 4 per cent. It is certain, however, that 25 386 OF RESPIRATION. this proportion may rise much higher; particularly when the respira- tory movements are slowly and laboriously performed. Now in order that the blood should be properly aerated, it is requisite that the air should contain no previous impregnation of carbonic acid; since the diffusion of even a moderate percentage of that gas through the inspired air, seriously impedes the exhalation of more. Thus it was found by Messrs. Allen and Pepys, that when 300 cubic inches of air were respired for three minutes, only 28J inches of carbonic acid (or somewhat more than 9 per cent.) were present in it; though the rate of its production in a parallel experiment, in which fresh air was taken in at each inspiration, Avas 32 cubic inches per minute, or 96 cubic inches in three minutes. It appears from the experiments of Dr. Snow, that the presence of carbonic acid in the atmosphere acts more delete- riously on the system, in proportion as the normal quantity of oxygen has been reduced; and hence, that the substitution of carbonic acid for oxygen by the respiratory process, vitiates the air far more effectually than the introduction of a surplus of carbonic acid, the normal quantity of oxygen being still present. He concludes from his experiments upon the lower animals, that 5 or 6 per cent, of carbonic acid cannot exist in an atmosphere respired by Man, without danger to life ; and that less than half this amount would soon be fatal, when it is formed at the expense of the oxygen of the air. A still smaller proportion is capable of producing very injurious results. Thus the discomforts occasioned by the presence of a crowded audience in a church, lecture- room, or theatre, which is not provided with sufficient ventilation, are due in great part to the continued respiration of air, which becomes loaded in the course of an hour or two with carbonic acid gas, to the amount of from one-half or two per cent.,—as has been ascertained both by direct experiment, and by calculation. And there can be no reasonable doubt, that the habitual respiration of such air, in the nar- row and noisome dwellings of the poor, or in croAvded factories and workshops, has a tendency to produce, both directly and indirectly, much loss of physical and mental vigor, and also to blunt the acute- ness of the moral feelings ; its influence being specially noticed in in- creasing the predisposition to Epidemic diseases, and in augmenting the fatality of their attacks. 697. The effects of a simple deficiency of Oxygen in the respired air, are experienced by those who breathe a rarefied atmosphere, such as that which exists on the summits of high mountains. All persons who have made such ascents, have experienced the insufficiency of rarefied air to sustain the degree of respiration required for active exertion. As long as the body remains at rest, no inconvenience is perceived; but as soon as the muscular system is put into action, the insufficiency of the supply of oxygen is manifested by the feeling of distress and languor; which becomes so severe, that the individual, if unused to sueh ascents, is obliged to stop and take breath at every few steps. The necessity for doing so will be easily understood, when it is remem- bered that Avhen the pressure of the atmosphere is reduced to half its usual amount, the bulk of a given weight of air will be twice as great as at the surface of the earth, or the same measure will Aveigh only CHANGES EFFECTED IN THE BLOOD. 387 half as much. Consequently, when the chest is completely filled with air, the real quantity of oxygen included*-in it, is only half of that which is drawn in by a corresponding inspiration at the earth's surface. 698. With regard to the absorption and exhalation of Nitrogen, it seems probable that both these processes are constantly going on; but that their relative activity varies under different conditions. Thus it has been ascertained by MM. Regnault and Reiset, that although Avarm-blooded animals, when subjected to their ordinary regimen, usu- ally increase the amount of nitrogen in the atmosphere, yet that, when food is Avithheld, or the animals are fed upon a diet to which they are unaccustomed, an absorption of nitrogen takes place ; this being parti- cularly remarkable in the hibernating Mammalia, in which the gain in weight by the absorption of oxygen and nitrogen even exceeds the loss occasioned by the exhalation of carbon. 699. Having thus considered the changes produced by the Respira- tory function, in the air submitted to it, Ave have next to inquire into converse series of changes effected by it in the blood. The nature of these cannot be well stated with precision, as they have not yet been fully determined. It was formerly supposed, that the venous blood arrives at the lungs charged with carbon, and that this carbon is united with the oxygen of the air in the lungs themselves. Numerous facts, however, go to prove, that the blood comes to the lungs charged with carbonic acid; and that it gives out this ready formed, and receives oxygen in its stead. Thus it has been already shown, that there is a positive disappearance of oxygen, more of that element being withdrawn from the atmosphere, than is restored to it in the condition of carbonic acid ; so that Ave know that the surplus must be received into the blood. Further, cold-blooded animals may be made to breathe nitrogen or hydrogen for a sufficient length of time, to cause a large quantity of carbonic acid to be disengaged; and this must have been brought to the lungs ready formed, since no oxygen was present there to generate it. Lastly, it can be shown by experiment, that oxygen, carbonic acid, and nitrogen exist in a free state in blood, arterial as well as venous; but that the proportion of oxygen is greater in arterial than in venous blood, whilst that of carbonic acid is less. The following table expres- ses the percentage of each kind of gas in the two sorts of blood respec- tively, as deduced from the experiments of Magnus. Arterial Blood. Venous Blood. Carbonic acid,.....62-3 71-6 Oxygen.......23-2 15-3 Nitrogen,......14-5 131 Thus it appears that the quantity of nitrogen is very nearly the same in both, as would be anticipated from Avhat has been already stated in regard to its non-participation in the respiratory process ; whilst about one-third of the free oxygen of arterial blood disappears during its cir- culation in the systemic capillaries, to be replaced by an equivalent amount of carbonic acid; and a converse change takes place in the pul- monary capillaries, this additional portion of free carbonic acid being set free, and replaced by oxygen. 388 OF RESPIRATION. 700. Thus it is evident, that a part of the change effected in the Blood consists in an alteration in the proportion of the gases which always exist in it, either entirely free, or in a state of such loose com- bination that they can be removed by the air-pump. But it may be suspected, that a portion of the effect consists in the oxidation of the proteine of the fibrinous constituent; since the fibrine of arterial blood possesses properties that distinguish it from that of venous. It has been usually supposed that the hematosine of the red corpuscles under- goes a change under the influence of oxygen in the lungs, and a con- verse change in the systemic capillaries, Avhere it is subjected to the influence of carbonic acid; this change being indicated by the altera- tion in the color of the red corpuscles. The alteration in question, however, seems due rather to a physical than to a chemical change (§ 222); and we have no direct evidence, though much that it is indirect, of the special influence of the aeration of the blood upon the contents of the red corpuscles. It appears tolerably certain that a part of the oxygen imbibed in the lungs, is appropriated to the oxidation of the matters set free by the decomposition of the solid tissues; whilst another part enters into combination with fatty, saccharine, farina- ceous, and other matters, existing in the blood itself, and destined to be carried off in the form of carbonic acid and water, without ever enter- ing into the composition of the solid fabric. The relative amounts of carbonic acid formed in these two modes, vary in different animals and in different states of the same individual: for a man in a warm atmosphere, taking a moderate amount of exercise, may thus set free, by the waste of his muscular and other tissues, a sufficient quantity of car- bon for the maintenance of his animal heat by its union Avith oxygen ; but this is far from being sufficient, when a larger amount of heat must be evolved, to sustain the temperature of the body in a colder climate. 701. The blood parts in the lungs with a very large amount of mois- ture ; for the inspired air is always saturated with fluid, as soon as it reaches the air-cells; and, as it is heated at the same time to about 98°, it thus receiA'es a considerable addition, even if it were previously charged with as much as it could contain at a lower temperature. The total quantity of fluid thus disengaged will vary, therefore, with the amount previously contained in the atmosphere, being greater as this was less, and vice versd ; but the quantity that usually passes off seems to be from 16 to 20 ounces in the twenty-four hours. It cannot be doubted, that a great part of this water is a simple exhalation of that which has been absorbed; but, on the other hand, it seems probable that a portion of it may be actually formed in the system, by the union of a portion of the oxygen absorbed in the lungs, with the hydrogen of the combustible matters of the blood. In the various forms of saccharine and farinaceous aliments, the proportion of hydrogen and oxygen are such as Avould of themselves form water, when the carbon is withdrawn; but in oily and fatty matters, the proportion of oxygen is far too small thus to neutralize the hydrogen ; and it seems likely that by their oxidation in the blood, as by their combustion else- where, Avater is actually generated by the union of atmospheric oxygen ASPHYXIA—ITS CAUSES. 389 with their hydrogen, at the same time that carbonic acid is produced by its union with their carbon. 702. Along Avith the Avater thus extricated from the lungs, a certain amount of organic matter is set free. If the fluid be collected in a closed vessel, and be exposed to warmth, a very evident putrid odor is exhaled from it; and if the expired air be made to pass through sulphuric acid, that liquid is colored red. Every one knows that the breath itself possesses, occasionally in some persons, and constantly in others, a foetid taint; when this does not proceed from carious teeth, ulceration in the air-passages or lungs, or other similar causes, it must result from the excretion of the odorous matter, in combination with watery vapor, from the pulmonary surface. That this is the true account of it seems evident, from the analogous phenomenon of the exhalation of turpentine, camphor, alcohol, and other odorous sub- stances, Avhich have been introduced into the venous system, either by natural absorption, or by direct injection ; and also from the sudden- ness Avith Avhich the odor manifests itself, when the digestive apparatus is slightly disordered. -^ 5. Effects of Insufficiency, or Suspension, of the Aerating Process. 703. The change which the Blood undergoes, by being brought into relation with atmospheric air in the respiratory organs, is so important to life, that the entire suspension of it inevitably produces a fatal ter- mination, at no remote period; and if it be insufficiently performed, various disorders in the system are nearly sure to manifest themselves. The state Avhich is induced by the entire suspension of the aerating process, is termed Asphyxia; a word which literally means the absence of pulse, and Avould be applicable therefore to the stoppage of the cir- culation from any cause; though it is usually employed to designate the particular condition resulting from suspended respiration. Asphyxia may be produced in aquatic animals, as well as in those which breathe air, by cutting them oft" from the influence of the atmosphere ; for if a fish be placed in water from Avhich the air has been expelled by boiling, it is precisely in the condition of an air-breathing animal placed in a vacuum, since it has no power of obtaining oxygen by decomposing the Avater it inhabits, and is entirely dependent for the aeration of its blood, upon the air that is absorbed by the liquid. Again, if a fish be placed in water impregnated Avith carbonic acid, its death is nearly as instan- taneous as that of an air-breathing animal immersed in an atmosphere of that gas. 704. Asphyxia may result from a great variety of causes. Thus there may be a mechanical obstruction to the entrance of air through the trachea; as in hanging, strangulation, or drowning; or as in occlusion of the aperture of the glottis, by oedema of its lips, or by the presence of a foreign body in the larynx. Or, again, the passage may be perfectly free, and yet no air may enter, in consequence of some obstacle to the performance of the respiratory movements. This obstacle may be me- chanical ; as when a quantity of earth has fallen round the body, in such a manner as completely to prevent the distension of the chest and abdomen. Or it may result (and this is a most frequent occurrence) from torpidity 390 OF RESPIRATION. or complete inactivity of the ganglionic centre, which is concerned in the respiratory actions ; or from interruption to the transmission of its influence along the nervous trunks. Further, Avhen there is no obstacle to the free ingress or egress of air, Asphyxia may be produced by the want of oxygen in the atmosphere that is respired, or by the presence of car- bonic acid in too large an amount. And the presence of other gases, which exert a directly poisonous influence on the blood,—such as sulphu- retted hydrogen,—produces a state, which may be included under the same general description. 705. Now when, from any of these causes, the free exchange of car- bonic acid for oxygen in the pulmonary capillaries is checked, the first effect of the interruption appears to be, the stagnation of the blood in the pulmonary capillaries. This stagnation is evidently due, not to any deficiency of power in the heart; for that organ is not yet affected ; but to the insufficiency of the heart's power acting alone, to drive the blood through the pulmonary capillaries : the force which should be generated by chemical changes in them (§ 598), being deficient. The stagnation is not, hoAvever, complete at first; since the quantity of oxygen contained in the, lungs is sufficient to produce an imperfect arterialization of the blood; and the blood thus partially changed is transmitted to the left side of the heart, and is thence propelled to the system. Owing to its half-venous' condition, it cannot exert its usual stimulating influence on the tissues, especially the muscular and nervous ; and their powers are consequently weakened. For the same reason, it does not receive its usual auxiliary force in the systemic capillaries (§ 599) ; since the changes, which it ought to undergo in them, can only -be partially per- formed. 706. As the air included in the lungs loses more and more of its oxy- gen, and becomes more and more charged with carbonic acid, the aeration of the blood in the pulmonary capillaries becomes more and more im- perfect ; the quantity of blood which is allowed to return to the heart is gradually diminished, and its condition become more and more venous; and at last, the pulmonary circulation is altogether suspended. From the relation which the respiratory circulation bears to the systemic, in all the higher classes of animals, save Reptiles, it folloAvs that the sys- temic circulation must in like manner be brought to a stand. The venous blood accumulates in the pulmonary artery, in consequence of the ob- struction of its capillaries; it distends the right caAnties of the heart; and the accumulation extends to the venous system of the body in general, especially affecting those organs which naturally receive a large quantity of venous blood, such as the liver and spleen. The arterial system, on the other hand, is emptied in a corresponding degree; nearly all its blood having passed through the systemic capillaries; and no fresh supplies being received from the heart. From this deficiency, and from the venous character of the blood which the systemic arteries do contain, it results that the nervous and muscular systems lose their power; insensibility comes on, at first accompanied with irregular con- vulsive movements; but in a short time there is a total cessation of all movement except that of the heart; and the pulsations of that organ be- come feebler and feebler, until they cease altogether. The immediate cause of the cessation of the heart's action appears to be different on the ASPHYXIA—ITS PHENOMENA AND TREATMENT. 391 t two sides. Both are equally affected by the want of arterial blood in the capillaries of their own substance ; but the right side suffers from over distension, which produces a sort of paralysis of its muscular tissue; Avhilst the left side retains its contractility, but is not excited to con- traction, for want of the stimulus of arterial blood in its cavities. 707. In those warm-blooded animals which are not endowed Avith any special provision for enabling them to sustain life during the pro- longed suspension of the respiratory process, insensibility and loss of Aroluntary power almost invariably supervene within a minute and a half after the admission of air to the lungs has been entirely pre- vented ; though the respiratory efforts and convulsive actions, which are dependent upon the medulla oblongata and spinal cord, may con- tinue for a minute or two longer. The circulation generally comes to a complete stand Avithin ten minutes at farthest.—The chief exceptions are in the case of diving animals, which are provided with large arterial and venous reservoirs, that serve to maintain the circulation during a prolonged suspension of the respiratory process ; for the arterial plexuses being ordinarily filled, they afford a supply of aerated blood to the systemic capillaries, Avhen other blood is wanting; and the reser- voirs connected with the venous system, which were previously empty, receive this blood, and prevent it from exercising undue pressure on the heart. To such an extent is this provision carried in some ani- mals, that the Whale has been knoAvn to remain under water for an hour. Another exception exists in the case of hibernating Mammals, which are reduced for a time to the condition of cold-blooded animals, and which can, like the latter, sustain a prolonged suspension of the aerating process. And there is reason to believe that, in the state of Syncope or fainting,—in which the circulation is already reduced to a very low amount, in consequence of a partial failure in the heart's poAver, all the functions of the body being nearly suspended, and the demand for aeration being consequently very small,—the respiration may be suspended for a long period, even in the Human subject, with- out fatal results. Thus more than one case has been credibly recorded, in which recovery has taken place after complete submersion for more than three-quarters of an hour; and it is probable that, in these in- stances, a state of Syncope came on at the moment of the immersion, through the influence of mental emotion, or of concussion of the brain. 708. In the restoration of an animal from the state of Asphyxia, it is above all things of importance to renew the air in the lungs; for in this way the blood in the pulmonary capillaries will be aerated, the capillary circulation will be re-established, the right side of the heart will be relieved of its excessive load of venous blood, and the left side Avill receive the stimulus of a fresh supply of arterial blood ; so that, if its movements have not ceased altogether, it may be speedily restored to due activity. At the same time, the temperature of the body should be kept up by artificial warmth ; and the circulation in the skin should be excited by friction. Where no other means are at hand for intro- ducing pure air into the lungs (of which means the application of gal- vanism along the course of the phrenic nerve, so as to produce contraction of the diaphragm, Avill probably be the most effectual), the object may 392 OF RESPIRATION. be attained by forciby compressing the trunk on all sides, so as to empty the lungs as much as possible, and then allowing the chest to dilate again, by the elasticity of its walls. In this manner, a large proportion of the carbonic acid may be expelled, and a considerable proportion of fresh air introduced, in the course of a few minutes. If air be blown into the lungs by the bellows, great care must be taken to prevent the employment of too much force, which is likely to produce rupture of the air-cells. 709. Now when, from the more prolonged action of various causes that impede the due performance of the respiratory function, the aera- tion of the blood in the lungs is insufficient for health, though not such as to produce a complete stagnation of the movement, a variety of results may follow; of Avhich some, or others, will manifest themselves, according to the condition of the general system, and the peculiarities of the individual. Thus deficient respiration seems to favor the re- tention of fatty matter in the system ; and this not merely in the con- dition of Adipose tissue, which (unless it accumulate to excess) may be regarded as a healthy product; but also in the place of the normal components of other tissues, as the muscular and glandular, giving rise to the condition Avhich is termed "fatty degeneration."—Again, the due elaboration of the fibrine of the blood is undoubtedly prevented by an habitually-deficient respiration; and various diseases, Avhich result from the imperfect performance of this elaboration, consequently mani- fest themselves. The Scrofulous diathesis is thus frequently connected with an unusually small capacity of the chest.—Further, an habitual deficiency of respiration may impede, though it does not check, the cir- culation in the lungs ; and thus a tendency arises, in various pulmonary diseases, to an overloading of the pulmonary arteries, to a dilatation of the right cavities of the heart, and to a congestion of the venous system in general, as marked by lividity of the surface, by venous pul- sation, &c. This state may result, not merely from obstruction in the lungs themselves, but from deficiency of the respiratory movements, consequent upon torpidity of the medulla oblongata (as in apoplexy and narcotic poisoning) or upon partial interruption of the nervous circle requisite for all reflex movements. Thus Avhen the par vagum is divided, the number of respiratory movements is greatly diminished, and a partial stagnation of the blood in the lungs is the result. The same happens in certain forms of typhoid fever, in Avhich the respira- tory movements are preternaturally slow, in consequence of torpidity of the medulla oblongata. Noav in this state, an effusion of the watery part of the blood into the air-cells of the lungs (as in other cases of obstructed circulation) is very liable to occur; and when the lungs are thus loaded with fluid, the respiratory process is still more impeded, and the disorder has thus a tendency to increase itself. NATURE OF THE SECRETING PROCESS. 393 CHAPTER IX. OF SECRETION. 1. Of the secreting process in general; and of the instruments by which it is effected. 710. We have seen that, in the process of Nutrition, the circulating current not only deposits the materials, Avhich are required for the renovation of the solid tissues; but also takes back the substances, Avhich are produced by the decay of these, and which are destined to be throAvn oft' from the body. We have also seen, that it supplies the materials of certain fluids, which are separated from it to effect various purposes in the economy ; such as the Salivary and Gastric fluids, which have for their office to assist in the reduction of the food. Now the process by which the fluids of the latter kind are separated from the Blood, is precisely the same in character as that by which the products of decay are eliminated from it; and the structure of the organs con- cerned in the two is essentially the same. Hence both processes are commonly included under the general term Secretion, which simply denotes separation. Considered in its most general point of vieAv, this designation may be applied to every act, by which substances of any kind are separated from the blood. Thus the function of the cells, which are concerned in the elaboration of the organizable plasma, may be termed one of Secretion, because they draw from the blood a supply of Albumen, upon which their converting action is exercised; but as the product of their operation is returned to the blood again, and is employed for higher purposes in the economy, the process is usually termed Assimilation. In the same manner, the elaborating action of the Lymphatic Glands, with the Spleen, Thymus Gland, &c, is not usually termed Secretion ; since, although it is exercised upon matter draAvn from the blood, the product appears to-be delivered back into the circulating current, through the medium of the Lymphatic System. (chap, v.) With much more justice, however, the process of Respira- tion may be regarded as one of Secretion; for it consists, as we have seen, in the constant elimination of a substance from the blood, Avhich cannot be retained in it without the most injurious consequences. 711. There is an important difference in the characters of the prin- cipal products of the Secreting process, which is connected with the purposes that are to be answered by their separation. Some of these products are altogether different from the ordinary constituents of the animal fabric, and from the materials which the blood supplies for the nutrition of these. So different are they, that their presence in the circulating current has an injurious effect; and the great object of their separation is the maintenance of the purity of the blood. These poi- sons, for such they may be considered, are generated in the system by the decay and decomposition to which its several parts are liable; and they are just as noxious to it, as if they Avere absorbed from without. 394 OF SECRETION. We have seen that the retention of Carbonic acid in the blood for even a feAV minutes is fatal, both by putting a stop to the circulation, and by acting unfavorably upon the substance of some of the most impor- tant organs in the body. The same fatal result attends the retention of Urea and of Biliary matter, Avhich are among the other products of the decomposition of the tissues; but, although as certain, it is not so speedy, because the general circulation is not affected by the loss of secreting power on the part of the Kidneys and the Liver, and because the accumulation of the noxious matter is slower.—On the other hand, the ingredients that are met Avith in those secreted fluids, which are destined to answer some purpose in the economy, almost invariably bear a close correspondence with the ordinary materials of the blood. Thus in the Salivary, Gastric, Pancreatic, and Lachrymal fluids, the principal part of the solid matter consists of the saline and of the albu- minous constituents of the blood, the latter in a more or less altered condition. In Milk, again, we trace the ordinary constituents of the blood, with very little change. Thus it appears, that the separation of these fluids is not required so much to maintain the Blood in a state of purity, as to supply Avhat is needed for some subsequent operation in the economy ; and hence, if the secreting process be interrupted, in the case of any one of them, the suspension has usually no further effect, than that of disturbing the process to Avhich the fluid is usually subser- vient. If the secretion of Gastric fluid be checked, for example, under the influence of strong mental emotion, the Digestive operation is pre- vented from taking place. 712. The essential character of the true Secreting operation seems to consist,—not in the nature of the action itself, for this is identical with those of Assimilation and Nutrition, being, as we have seen (§ 239), a process of cell-groAvth,—but in the position in which the cells are developed, and the mode in Avhich the products of their action are afterwards disposed of. Thus the cells at the extremities of the intes- tinal Villi (§ 243), the cells of which the Adipose tissue is made up (§ 257), and the cells of which the greater part of the substance of the Liver is formed (§ 239), all have an attraction for fatty matter; and draw it from the neighboring fluids, at the expense of which they are developed, to store it up in their own cavities. But the cells of the first kind, when they have come to maturity, set free their contents, which are delivered to the absorbent vessels, to be introduced into the circulating current;—those of the second kind seem more permanent in their character, and retain their contents, so as to form part of the ordinary tissues of the body, until they are required to give them up for other purposes, when the matters which they have temporarily separated from the circulating current, are restored to it again without change;—and the cells of the third class, when they liberate their con- tents (which they are continually doing), cast them forth into the hepatic ducts, by which they are carried into the intestinal canal, whence a portion of them at least is directly conveyed out of the body. 713. It is, then, in the position of the Secreting cells,—which causes the product of their action to be delivered upon a free surface, commu- nicating, more or less directly, with an external outlet,—that their SIMPLEST FORM OF GLANDULAR STRUCTURE. 395 distinctive character depends. All the proper Secretions are thus either poured out upon the exterior of the body, or into cavities provided Avith orifices that lead to it. Thus we shall see that a considerable quantity of solid matter, and a very large quantity of fluid, of which it is desirable that the system should be freed, are carried off from the Cutaneous surface. Another most important secretion, containing a large quantity of solid matter, and serving also to regulate the quan- tity of fluid in the body,—namely, the Urinary,—is set free. by a channel expressly adapted to convey it directly out of the body. The same may be said of the Mammary Secretion, which is separated from the blood, not to preserve its purity nor to answer any purpose in the economy of the individual, but to afford nutriment to another being. And of the matters secreted by the very numerous glandulae situated in the walls of the Intestinal canal, a great part are obviously poured into it for no other purpose, than that they may be carried out of the body by the readiest channel. 714. The cells covering the simple membranes that form the free surfaces of the body, whether external or internal, are all entitled to be regarded as secreting cells, since they separate various products from the blood, which are not again to be returned to it. But the secreting action of some of these seems to have for its object the pro- tection of the surface; thus the Epidermic cells secrete a horny matter, by which density and firmness are imparted with the cuticle ; Avhilst by the epithelial cells of the Mucous membrane of the alimentary canal, and of other parts, their protective Mucus seems to be elaborated. But in general we find that special organs, termed Glands, are set apart for the production of the chief secretions; and we have now to consider the essential structure of these organs, and the mode of their operation.—A true Gland may be said to consist of a closely-packed collection of follicles, all of which open into a common channel, by Avhich the product of the glandular action is collected and delivered. The follicles contain the secreting cells in their cavities ; whilst their exterior is in contact with a network of blood-vessels, from which the cells draw the materials of their groAvth and development (Fig. 94). In any one of the higher animals, we may trace out a series of pro- gressive stages of complexity, in the various glands included within their fabric ; and in following any one of the glands that attain the highest degree of development (such as the Liver or Kidney), through the ascending scale ©f the Animal series, we should trace a very similar gradation from the simplest to the most complex form. 715. That there is nothing in the form or disposition of the compo- nents of the glandular structure, which can have any influence upon the character of the secretion it elaborates, is evident from the fact, that the very same product,—e. g., the Bile, or the Urine,—is found to issue from nearly every variety of secreting structure, as Ave trace it through the different croups of the Animal kingdom. The peculiar power by Avhich one orJPm separates from the blood the elements of the Bile, and another the elements of the Urine, whilst a third merely seems to draw off a certain amount of its albuminous and saline con- stituents, is obviously the attribute of the ultimate secreting cells, 396 OF SECRETION. which are the real agents in the secreting process (§ 239). Why one set of cells should secrete Bile, another Urea, and so on, we do not know; but we are equally ignorant of the reason, for which one set of cells converts itself into Bone, another into Muscle, and so on. This variety in the endowments of the cells, by whose aggregation and conversion the fabric of the higher Animals is made up, is a fact which we cannot explain, and which must be regarded (for the present, at least), as one of the "ultimate facts" of Physiological Science. 716. Passing by the extended membranous surfaces, and the protec- tive cells with Avhich they are covered, we find that the simplest form of a secreting organ is composed of an inversion of that surface into a series of follicles, which discharge their contents upon it by separate orifices. Of this we have an example in the gastric follicles, even in the higher animals; the apparatus for the secretion of the Gastric fluid never attaining any higher condition, than that of a series of distinct follicles, lodged in the walls of the stomach, and pouring their products into its cavity by separate apertures. In Fig. 107 is represented a a portion of the Ventriculus succenturiatus of a Falcon ; in which the simplest form of such follicles is seen. A somewhat more complex condition is seen in some of the Gastric follicles of the Human stomach (Fig. 80); the surface of each follicle being further extended by a sort of doubling upon itself, so as to form the commencement of secondary follicles, which open out of the cavity of the primary one.—Now a con- dition of this kind is common to all glands, in an early stage of their evolution; and in this stage, we meet Avith them, either by examining them in the lowest animals in Avhich they present themselves, or by looking to an early period of their embryonic development in the highest. Thus, for example, the Liver consists, in certain Polypes and in the lowest Mollusca, of a series of isolated follicles, lodged in the walls of the stomach, and pouring their product into its cavity by sepa- rate orifices; these follicles being recognised as constituting a biliary apparatus, by the color of their secretion. And in the Chick, at an Fig. 107. Fig. 108. Glandular follicles in ventriculus Origin of the Liver from the intestinal wall, in *• succenturiatus of Falcon. the embryo of the Fowl, on the fourth day of in- cubation :—a, heart; b, intestine; c, everted por- tion giving origin to liver ; d, liver ; e, portion of yolk-bag. jjL early period of incubation, the condition of the Liver is essentially the same with the preceding; for it consists of a cluster of isolated follicles not lodged in the Avails of the intestine, but clustered around a sort of DIFFERENT FORMS OF GLANDULAR STRUCTURE. 397 bud or diverticulum of the intestinal tube, which is the first condition of the hepatic duct, and into which they discharge themselves (Fig. 108). So, again, the Pancreas first presents itself in the condition of Fig. 109. Fig. 110. Rudimentary Pancreas from Cod;—a, Mammary Gland of Ornithorhyncus. pyloric extremity of stomach; 6, intes- tine. a group of prolonged follicles, or cseca, clustered round the commence- ment of the intestinal tube ; and this is its permanent condition in many Fishes (Fig. 109). And the Mammary Gland possesses an equally simple structure in the lowest of all the Mammalia (to which group it is restricted); namely, in the Ornithorhyncus (Fig. 110). 717. The next grade of complexity is seen, Avhere a cluster of the ultimate follicles open into one common duct, which discharges their product by a single outlet; a single gland often containing a number of such clusters, and having, therefore, several excretory ducts. A good example of such a condition, in which the clusters remain isolated from one another, is seen in the Meibomian glands of the eyelid (Fig. Ill); each of which consists of a double ro>v of follicles, set upon a long straight duct, that receives the products of their secreting action Fig. 111. Fig. 112. Meibomian glands of upper lid of Portion of Cowper's gland from Hedgehog; the new-born infant. follicles distended with air. and pours them out upon the edge of the eyelid. And of the more complex form, in Avhich a number of such clusters are bound together 398 OF SECRETION. in one glandular mass, we have an illustration in the accessory glands of the genital apparatus, in several animals, which discharge their secretion into the urethra by numerous outlets (Fig. 112); or in the Mammary glands of Mammalia in general, the ultimate follicles of which are clustered upon ducts that coalesce to a considerable extent, though continuing to form several distinct trunks even to their termi- nation. Such glands may be subdivided, therefore, into glandulse or Fig. 113. Lobule of Lachrymal Gland; from foetal sheep. lobules, that remain entirely distinct from each other (Fig. 113).—In the highest form of Gland, however, all the ducts unite; so as to form a single canal, which conveys away the products of the secreting action of the entire mass. This is the condition in which we find the Liver to exist, in most of the higher animals ; also the Pancreas, the Parotid Gland, and many others. In some of these cases, Ave may still sepa- rate the gland into numerous distinct lobules, which are clustered upon the excretory duct and its branches, like grapes upon a stalk; in others, however, the branches of the excretory duct do not confine themselves to ramifying, but inosculate, so as to form a network, which passes through the whole substance of the gland, and which connects together its different parts, so as to render the division into lobules less distinct. This seems to be the case in regard to the Liver of the higher Verte- brata (§ 723). 718. Whatever degree of complexity, however, prevails in the gene- ral arrangement of the elements of the Glands in higher animals, these Fig. 114. Fig. 115. Two follicles from the liver of Carcinus mcenas (Com- Ultimate follicles of Mammary gland with their mon Crab), with their contained secreting cells. secreting cells, a, a ;—b, b, the nuclei. elements are themselves everywhere the same; consisting of follicles that enclose the real secreting cells (Figs. 114 and 115). Noav from DIFFERENT FORMS OF GLANDULAR STRUCTURE. 399 the history of the development of Glands in general, it appears that the follicles may be- considered as parent-cells and that the secreting cells in the interior constitute a second generation, developed from the nuclei germinal spots on the walls of the first. Of such parent-cells, we have characteristic examples in the Peyerian glandulae of the intestinal canal (§ 450), and also in the Thyroid gland (§ 513) and Supra-Renal capsules (§ 510); and the most elaborate glands, in their earliest stage of development, present a similar condition. These closed cells become follicles, by opening at one extremity, either upon a neighboring free surface, or into a canal Avhich is prolonged from it. Thus the first rudi- ment of the Liver is formed by a thickening of the cellular mass in the walls of the alimentary canal, at the spot in which the hepatic duct is subsequently to discharge itself. This thickening increases, so as to form a projection upon the exterior of the canal; and soon after- wards the lining membrane dips down into it, so that a kind of caecum is formed, surrounded by a mass of cells, as shown in Fig. 108. The increase of the mass seems to take place by a continual new budding- forth of cells from its peripheral portion, which takes place to a consi- derable extent before the caecum in the interior begins to ramify. Gra- dually, however, the cells of the exterior become metamorphosed into fibrous tissue for the investment of the organ; those of the interior break down into ducts which form continuations of the principal canal; whilst those which occupy the intervening space, and Avhich form the bulk of the gland, seem to be developed into follicles, and to give origin to the proper secreting cells. As this is going on, the hepatic mass is gradually removed to a distance from the wall of the alimentary canal; and the caecum is narrowed and lengthened, so as to become a mere con- necting pedicle, forming, in fact, the main trunk of the hepatic duct.— The development of the Pancreas, Salivary glands, &c, seems to follow the same plan. 719. It has been pointed out by Prof. Goodsir, that the continued development and decay of the glandular structure,—in other Avords, the elaboration of its secretion,—may take place in two different modes. In one class of Glands, the parent-cell, having begun to develope new cells in its interior, gives way at one point, and bursts into the excretory duct, so as to become an open follicle, instead of a closed cell: its con- tained or secondary cells, in the progress of their own groAvth, draAv into themselves the matter to be eliminated from the blood, and, having attained their full term of life, burst or liquefy, so as to discharge their contents into the cavity of the follicle, whence they pass by its open orifice into the excretory duct: and a continual new production of secon- dary cells takes place from the germinal spot, or nucleus, at the extre- mity of the follicle, which is here a permanent structure. In this form of gland, we may frequently observe the secreting cells existing in vari- ous stages of development within a single follicle; their size increasing, and the character of their contents becoming more distinct in propor- tion to their distance from the germinal spot (which is at the blind termination of the follicle), and their consequent proximity to the outlet (Fig. 114). In some varieties of such glands, however, as in the greatly-prolonged follicles or tubuli uriniferi of the kidney, the produc- 400 OF SECRETION. tion of new cells does not take place from a single germinal spot at the extremity of the follicle, but from a number of points scattered through its entire length.—In the second type of Glandular structures, the parent-cell does not remain as a permanent follicle; but, having come to maturity and formed a connexion with the excretory duct, it dis- charges its entire contents into the latter, and then shrivels up and disappears, to be replaced by newly-developed follicles. In each parent- cell of a gland formed upon this type, we shall find all its secondary or secreting cells at nearly the same grade of development; but the diffe- rent parent-cells, of which the parenchyma of the gland is composed, are in very different stages of growth at any one period: some having dis- charged their contents and being in progress of disappearance, whilst others are just arriving at maturity and connecting themselves with the excretory duct; others exhibiting an earlier degree of development of the secondary cells ; others presenting the latter in their incipient con- dition ; whilst others are themselves just starting into existence, and as yet exhibit no traces of a secondary generation.—The former seems to be the usual type of the ordinary glands; the latter is chiefly, if not entirely, to be met with in the Spermatic glands. 2. Of the Liver and the Bile. 720. The LiArer is more rarely absent than any other Gland; being discoverable, under some form or other, in all but the very lowest mem- bers of the Animal kingdom. Its simple condition in the higher Po- lypes has been already noticed (§ 716); and it is met with, under an almost equally simple form, in the Starfish. As we ascend the scale, hoAvever, we find it assuming a much greater importance, and presenting a great increase in size. This is particularly the case in the Mollus- cous classes; and also in the Crustacea, a class which, in mode of respi- ration and in general habits, bears a great resemblance to the Mollusca. In nearly all such animals the Liver makes up a large proportion of the mass of the body. It usually consists of a series of large follicles, Fig. 116. Fig. 117. Lobule of Liver of Squilla Mantis; exterior. Lobule of Liver of Squilla Mantis, cut open. which branch out into smaller ones (Figs. 116 and 117), and of which several open into one excretory duct; but these ducts remain separate, and discharge their contents into the intestine by several distinct ori- COMPARATIVE STRUCTURE OF THE LIVER. 401 fices.—In Insects and other air-breathing Articulata, however, the Liver is much less developed; and its type remains much simpler. We usually find it consisting of a small number of caecal tubuli, which open separately into the intestinal canal, just below the stomach. These tubuli are often so long, as to pass several times from one extremity of the visceral cavity to the other, being doubled upon themselves; in other instances, we find that the principal tube or canal is beset with rows of short follicles, somewhat in the manner of Fig. 111. But they never cluster together so as to form a solid glandular mass. The low development of the liver in these animals, bears an evident relation with the high development of their respiratory apparatus; whilst, the respiration being comparatively feeble in the aquatic Mollusca and Crustacea, the development of the liver in those classes is enormous. 721. There is much difficulty in ascertaining the mode in which the elementary constituents of the Liver are arranged, in the fully-developed condition of that organ in the higher Vertebrata. At an early period of its development, as already remarked, it may be easily shown to con- sist, in the Fowl, of a series of distinct caeca, clustered round a projec- tion from the intestinal canal, and opening separately into it (Fig. 108); and it is a peculiarly interesting fact, that this very condition should exist as the permanent form of the Liver, in that curious little fish, the Amphioxus or Lancelot, which retains the embryonic type in so many parts of its conformation. In the Tadpole, again, the distinct caeca are very evident (Fig. 118); but here we see that the projection of the intestinal canal, instead of being a simple wide caecum, has become extended in length and contracted in diameter, at the same time di- viding and subdividing, so as to form an arborescent excretory duct, whose ramifications extend through the entire glandular mass. In this manner, then, is formed the complex system of hepatic ducts, which we find in the liver of the higher Vertebrata, branching out from the main trunk. But the mode in which the ultimate ramifications of these are arranged, and their relations with the secreting cells which make up the parenchyma of the gland, have not yet been fully elucidated. The fol- lowing are the principal facts, that have been ascertained on the sub- ject. 722. The entire Liver is made up of a vast number of minute lobules, of irregular form, but about the average size of a millet-seed. Each of these lobules contains the component elements of which the entire organ is made up; namely, branches of the hepatic artery and vein, branches of the portal vein, branches of the hepatic ducts, and secreting cells. The lobules are connected together in part by areolar tissue, but in great part by the anastomosis of the blood-vessels and hepatic ducts, which supply the adjoining lobules; indeed there is frequently no definite division of the glandular substance into lobules, other than that which is marked out by the arrangement of these canals (Figs. 119 and 121). The branches of the Hepatic Artery are principally distributed upon the Avails of the hepatic ducts, and upon the trunks and branches of the portal and hepatic veins, supplying them with their vasa vasorum ; also upon Glisson's capsule and its prolongations into the substance of the liver, -which prolongations form the greatest part of the connecting 402 OF SECRETION. structure that holds together the several elements. There is strong reason to believe, that the blood which the liver receives from the Fig. 118. Liver of Tadpole; showing distinct and free caecal terminations of the biliary ducts. hepatic artery is not destined to supply the materials for the biliary secretion, until it has become venous by travelling through the network, in which it is subservient to the nutrition of the tissues it permeates, as it is in other parts of the systemic capillary system.—The supply of blood, from which the materials of the biliary secretion are chiefly drawn, is afforded by the Vena Portar, which collects it as a Vein from the chylopoietic viscera, and which then subdivides as an Artery to dis- tribute it to the different parts of the Liver. Its branches proceed to Fig. 119. the capsules of the lobules, covering the whole external surface of the latter with their ramifications, and sending capillary twigs inwards, which converge towards the centre of each lobule (Fig. 119, 2, 2). As STRUCTURE OF THE LIVER. 403 the principal branches of these veins ramify in the spaces between the lobules, they are termed mfer-lobular veins.—On the other hand, the branches of the Hepatic Vein pass from the trunk to the centre of each lobule, from which they send out diverging capillary tAvigs (1, 1), to- wards the circumference; and these last, coming into connexion with the converging capillaries of the portal vein, establish a free capillary communication between the interior and exterior of each lobule. Thus the portal blood is first distributed to its exterior, then penetrates its substance, and then, after permeating the parenchymatous substance in numerous minutely-divided streams, is collected and carried off by the hepatic vein, of which a twig originates in the centre of each lobule. OAving to the peculiar position of the branches of the hepatic vein in the centre of each lobule, the lobules are appended to its main trunks almost in the manner of leaves upon a stem (Fig. 120). The precise relation of the capillaries of the hepatic artery with those of the portal Fig. 120. and venous systems has not yet been well ascertained; but there seems reason to believe, with Mr. Kiernan, that the arterial capillaries dis- charge themselves into the ultimate ramifications of the portal vein; and that thus the blood of the former, having become venous by trans- mission through the nutritive capillaries of the liver, mingles with the other venous blood collected by the venae portae, to supply the mate- rials of the secretory function, Avhich are eliminated from it during its passage into the hepatic vein. 723. The Hepatic Ducts also seem to form a plexus which surrounds the lobules, connecting them together, and sending branches towards the interior of each. The mode in which they terminate, hoAvever, and the precise relation in Avhich they stand to the hepatic cells, which form nearly the entire parenchyma of the Gland, has not yet been com- pletely elucidated. There seems reason to believe, hoAvever, that the tubular plexus extends throughout the substance of the lobule, filling up the entire space not occupied by the blood-vessels (its membranous Avail, however, being with difficulty traceable, owing to its extreme tenuity); and that the hepatic cells are contained Avithin it, as within the follicles or tubes of ordinary glands. These cells (which are easily Obtained in a separate condition by scraping a piece of fresh Liver) are of a flattened spheroidal or polygonal form ; and their diameter is usually from l-800th to l-1600th of an inch. Each cell presents a dis- 404 OF SECRETION. tinct nucleus; and it is usually around this, that the yellowish hue of the cell is the deepest. The cavity of the cell is chiefly occupied by biliary matter, much of which is in the condition of fine granular parti- cles too minute to be measured. In the midst of these, there are usually Horizontal section of two superficial lobules, showing the interlobular plexus of biliary ducts:—1,1, in- tralobular veins; 2, 2, trunks of biliary ducts, proceeding from the plexus which traverses the lobules; 3, nterlobular tissue; 4, parenchyma of the lobules. one or two large adipose globules, or five or six small ones (Fig. 122); but the amount of this fatty matter is liable to great variations (§ 754). The biliary matter which these cells contain, I22- marks them out as the real agents in the se- creting process ; this process consisting, it is evident, in the growth of the hepatic cells, Avhich, in the course of their development, eliminate from the blood the biliary matter, for which they have a special affinity. The Glandular ceiis of Liver -.-a, nucleus; mode in which the particles thus eliminated b, nucleolus; c, adipose particles. -i. i i • i i • are discharged into the hepatic ducts, to be by them conveyed to the intestine, cannot be understood, until the rela- tion between the secreting cells and the ultimate ramifications of the ducts shall have been more precisely determined. 724. The Bile which has been secreted by the hepatic cells, and Avhich has found its way into the ramifications of the hepatic ducts, may be directly conveyed by the trunk of the latter into the intestine, or it may regurgitate along the cystic ducts into the gall-bladder. It is probable that the secreting process is constantly going on; although, as in other cases, it may vary in its degree of activity at different times. When the process of digestion is taking place, and the small intestine is filled with chyme, there is probably an uninterrupted flow of bile into its cavity; but when the intestine is empty, the bile seems not to be admitted into it, but rather to flow back into the gall-bladder, in which it is stored up, as in a reservoir, until the time when it may be needed. In this reservoir it undergoes a certain degree of concen- tration, by the absorption of its watery part; and it also becomes mixed Avith a large proportion of mucus, which is secreted by the walls of the gall-bladder.—As the analyses of Bile have been chiefly made upon the fluid obtained from this receptacle, they probably over-esti- mate the proportion of solid matter contained in this secretion; Avhich COMPOSITION OF THE BILE. 405 is usually stated at from 8 to 9| per cent. Of this solid matter, about a tenth consists of alkaline and earthy salts, corresponding with those of the blood; whilst the remainder is made up of organic constituents. These are very readily decomposed, and enter into neAV combinations with the substances employed to separate them; so that different Chemists, by employing different means of analysis, have obtained re- sults which seem far from conformable. All are agreed, however, that the chief part of the solid ingredients of bile are allied to fat in compo- sition ; consisting of a very large proportion of carbon and hydrogen, and of a comparatively small amount of oxygen and azote. According to Prof. Liebig, the organic portion of ox-bile may be represented by the formula 76 Carbon, 6Q Hydrogen, 22 Oxygen, and 2 Nitrogen, with a considerable proportion of Sulphur. This substance, essentially corresponding with the bilic acid, choleic acid, bilin, picromel, &c, of different Chemists, seems to be a fatty acid (§ 261), united with soda, so as to constitute a soap. In healthy bile, the proportion of Cholesterine appears to be very small, and it is held in solution by the preceding ingredient; but in many disordered states, and especially in disease of the Gall-Bladder, this element is present in much larger amount; and it usually forms the principal, if not the sole ingredient in biliary con- cretions. It is a white crystallizable fatty matter, somewhat resem- bling spermaceti, free from taste and odor, and composed almost entirely of carbon and hydrogen ; its formula is 36 Carbon, 32 Hydro- gen, and 1 Oxygen.—The coloring matter of Bile is a substance distinct from the preceding; that of the Ox and other graminiA'orous animals appears to be identical, or nearly so, with the chlorophyll of the leaAres on which they feed ; but that of Human bile seems to possess different properties, and to be derived from the proper constituents of the blood. 725. Regarding the destination and purposes of this secretion in the Animal economy, the following may be considered as a tolerably com- plete summary; though it is impossible to speak with precision on some points, since the organic constituents of the Bile are liable to be so easily altered by various reagents, that they are with difficulty recog- nised.—A portion of the Bile unquestionably passes off, in Man and most other animals, with the faeces ; this portion, which includes the coloring matter, is probably that which would be most injurious, if retained in the blood, and is most purely excrementitious. In bilious diarrhoea, and under the influence of purgatives, especially mercurials, a large quantity of bile is discharged per anum, apparently almost unchanged. But in the healthy state, a portion, at least, of the soapy compound seems destined for reabsorption. Just as ox-gall is com- monly used to remove grease-spots, by its solvent power for fatty mat- ter, so does the bile seem to act in the living body, by rendering soluble the fatty matters of the fcod, and thus enabling them to be absorbed by the lacteals (§ 479). The fatty matter of the bile when reabsorbed with that of the neAvly-ingested food, is probably, like it, carried off by the respiratory process : but it is easily shoAvn, that the biliary matter cannot supply more than one-sixth or one-eighth of the amount of carbon eliminated from the lungs in the form of carbonic acid ; and 406 OF SECRETION. that it cannot be (as supposed by Liebig) the chief fuel of the process of combustion, which is kept up through the agency of those organs.— The secreting action of the Liver, by Avhich a certain product is entirely separated from the blood, constitutes, however, only a part of the action of that organ; since, as already shown (§ 493), the changes which it effects in the alimentary materials newly introduced into the current of the circulation, are at least equally important. A large part of the bulk of the Liver, in many of the lower animals, is made up of oleagi- nous matter; which appears to accumulate in the hepatic cells, giving them almost the character of fat-cells, in proportion as the respiratory function is inactive. Thus, the liver is very large and fatty in Mollusca and Crustacea; whilst, on the other hand, in Insects it is comparatively undeveloped. In Fishes, again, it is rich in oily matter, but in Mam- malia it is much less fatty in the state of health; whilst in the liver of Birds, scarcely any traces of fat are to be found. 726. The elements of the bile may be altogether supplied by the disintegration of the tissues; and this must certainly be the case, Avhen the amount of food taken is no more than enough to supply the waste of the system. We may regard it, then, as one office of the Liver, to remove from the blood such products of that disintegration, as are rich in carbon and hydrogen. It may be pretty certainly affirmed, however, that biliary matter does not pre-exist as such in the blood; but that its elements must be originally present there, under some more pernicious form. For it is found that the total suspension of the secreting action of the Liver, whereby the excrementitious matter is left to accumulate in the blood, has a much more prejudicial effect upon the system, than the reabsorption of Bile after it has been secreted, in consequence of an obstruction to its discharge through the ductus choledochus; so that it may be inferred that the noxious products of the disintegration of the tissues are transformed into comparatively innocent components of Bile, in the very act of secretion.—But there can be little doubt, that the Liver has also for its office, to draw off from the blood any superfluity which may exist in the non-azotized compounds derived from the food, beyond the amount that is requisite for the supply of the respiratory process, or that can be deposited as fat. For we conti- nually witness the results of habitual excess in the amount of such sub- stances, in producing that state of the system commonly termed bilious; of which all the symptoms are referable to the accumulation of the ele- ments of the bile in the blood, and the consequent deterioration in the purity of the circulating fluid. Where a tendency to such a state exists, proper means should be taken to stimulate the liver to increased activity; but the chief reliance should be placed on the avoidance of those articles of diet, which contain a large proportion of non-azotized matter, and on abstinence from superfluous nutriment of any de- scription. 3. Of the Kidneys and the Urine. 727. The Kidneys are perhaps the most purely excreting organs in the body ; their function being to separate from the blood certain matters STRUCTURE OF THE KIDNEY. 407 that would be injurious to it if retained, and these matters being des- tined to immediate and complete removal from the system. We have seen that, in the Lungs, the excretion of Carbonic acid is made subser- vient to the absorption of Oxygen ; and the separation of a fatty acid from the blood, which is effected by the Liver, is a means of introducing a new supply of fatty matter into the system. There is no ulterior purpose of this kind in the secreting action of the Kidney ; the product of which is invariably conveyed directly to an outlet, by which it may be discharged from the body. Some traces of Urinary organs may be de- tected in most of the higher Invertebrata; but it is in Fishes, that they first present a considerable development; and in ascending through the Vertebrated series, we find them rapidly increasing in the complexity of their organization, and in their functional importance, although their size and extent are not so great. In Fishes, the Kidneys very com- monly extend the whole length of the abdomen; and they consist of tufts of uniform-sized tubules, Avhich shoot out transversely at intervals from the long ureter, and which are connected together by a loose web Fig. 123. Embryo of Green Lizard;—a, heart; b, duplex aorta; c, vena cava; ^intes- tine ; e, liver;/, rudiment of AVolffian body;#, g, rudiments of extremities. of areolar tissue, that supports the network of vessels distributed upon their walls. This condition of the Urinary organs is very analogous to that of the Corpus Wolffianum or temporary kidney of the embryo of higher animals (Fig. 123,/). A similar condition is found in the true Kidney of higher animals at an early grade of development (as shown in Fig. 124); the tubuli uriniferi being short and straight. In their more advanced condition, however, they become long and convoluted; and the ramifications of the capillary vessels come into very close rela- tion with them (Fig. 125). It is in the higher Reptiles, that we first meet with the distinction between the cortical and medullary substance ; the former being the part in which the blood-vessels are most copiously distributed, and in which the tubuli have the most convoluted arrange- ment ; and the latter consisting chiefly of straight tubuli, converging towards the points at Avhich they discharge themselves into the ureter (Fig. 126). The bundles of tubuli and their vascular plexuses remain distinct, however, in Birds and in the lower Mammalia, so as to give to 408 OF SECRETION. the whole gland a lobulated character; but in the Human Kidney they come into closer contact; and the vascular connexion between the Fig. 125. ias5« Portion of Kidney from Coluber:—a, a, vascular trunk; b, 6, ureter j c, c, converging fasciculi of tubuli uriniferi. Pyramidal fasciculi of tubuli uriniferi of Bird, terminating in one of the branches of the ureter. plexuses of the different bundles is such, as to prevent any separation into distinct lobules. 728. The act of secretion appears to be effected, as in other Glands, by the epithelial cells lining the tubuli uriniferi; these cells drawing the materials of their development from the vascular plexus upon the exterior of these tubuli; and delivering them up, when they have com- pleted their own term of existence, to be carried off through the open Fig. 127. Fig. 128. 2 \M■ A section of the Kidney, surmounted by the supra-re- nal capsule; the swellings upon the surface mark the original constitution of the organ, as made up of dis- tinct lobes.—1. The supra renal capsule. 2. The vas- cular portion of the kidney. 3,3. Its tubular portion, consisting of cones. 4,4. Two of the papilla: project- ing into their corresponding calices. 5, 5, 5. The three infundibuli; the middle 5 is situated in the mouth of a calyx. 6. The pelvis. 7. The ureter. Portion of the Kidney of a new- born infant:—a, natural size; 1, 1, Corpora Malpighiana, as dispersed points in the cortical substance; 2, papilla; n, a smaller part magnified; 1, 1. Corpora Malpighiana; 2, 2. Tu- buli uriniferi. orifices of the tubuli. But the Kidney contains another apparatus, of a very peculiar description; which appears specially destined for the STRUCTURE OF THE KIDNEY. 409 separation of the superfluous fluid of the system. When a section of the Kidney is slightly magnified (Fig. 128, b), the cut surface is seen to be studded by a number of little dark points ; each one of which, when exa- mined under a higher magnifying power, is found to consist of a knot of minute blood-vessels, formed by the convolutions of thin-walled capilla- ries (Fig. 12!), m). It has been shown by Mr. Bowman, that each one of these knots is included in a flask-like capsular dilatation, connected with one of the tubuli uriniferi; several such capsules, it appears, being usually developed from the sides of each tubulus, like currants upon a stalk. Each of these vascular tufts (called Malpighian bodies, after their discoverer) is directly supplied by a branch of the renal artery (Fig. 129, af); which, upon piercing the capsule, subdivides into a group of capillaries ; and these, after forming the convoluted tuft, coa- lesce into a single efferent trunk (ef), which may be considered as re- presenting (in a small way) the vena portse. For the efferent trunks of the Malpighian bodies discharge their blood into the capillary plexus, ^Fig. 129. Distribution of the Renal vessels, from Kidney of Horse:—a, branch of Renal artery; af, afferent vessel; m, m, Malpighian tufts; ef, ef, efferent vessels; p, vascular plexus surrounding the tubes; st, straight tube; ct, convoluted tube. Magnified about 30 diameters. / which surrounds the tubuli uriniferi, and from which the solid matter of the urinary secretion is elaborated ; just as the vena portce supplies the capillary plexus, from which the biliary secretion is elaborated in the liver. In Reptiles (in Avhich, as in Fishes, the kidney is partly supplied by the hepatic portal system), the efferent vessels of the Mal- pighian bodies unite with branches of the portal vein to form the secre- ting plexus around the tubuli uriniferi; and even in Birds this arrange- ment still seems to prevail to a certain extent. Thus all the blood which the secreting plexus receives, has already passed, in each case, through a set of capillaries within or Avithout the organ ; those, namely, of the Malpighian bodies, or those of the parts supplying the hepatic portal system.—The special purpose of the Malpighian bodies appears to be, to alloAv of the transudation of the water of the blood, which is filtered off (so to speak) through the thin walls of their capillaries, and thus passes into the tubuli uriniferi. It is well known that the fluid and solid constituents of the urinary secretions bear no constant relation to each other; the amount of fluid depending mainly upon the degree 410 OF SECRETION. of fulness of the blood-vessels; whilst the amount of solid matter is governed, as we shall presently see, by the previous waste of the tissues. The quantity of fluid in the blood-vessels is governed by the relative amount that has been absorbed, and that Avhich has been exhaled from the skin ; so that the quantity to be drawn off by the Kidneys is^ in- creased, either by augmented absorption, or by diminished exhalation. The Malpighian bodies seem to act the part of a system of regulating valves ; permitting the transudation of only enough fluid to dissolve the solid matter, when there is no superfluity of water in the vessels ; but allowing the escape of an almost unlimited amount of it, when increased imbibition has rendered the vessels unusually turgid. 729. The average amount of Urine excreted in twenty-four hours, by adults Avho do not drink more than the Avants of nature require, is probably from 30 to 40 oz.; and its average specific gravity may be about 1020. The quantity of fluid is usually less, and the specific gravity of the secretion consequently greater, in summer than in Avin- ter, on account of the larger proportion of fluid exhaled by the skin during the former season. The quantity of solid matter has been found to vary, within the limits of ordinary health, from 3*6 to 6*7 per cent.; and the extent of variation in disease is doubtless much greater. About one-third of the solid matter is made up of alkaline and earthy salts; and the remainder is made up of organic compounds. The salts are partly those of the blood, which will not be separated during the transudation of the serum through the membranous walls of the Malpighian capilla- ries, although the albuminous matter is kept back (§ 196). But there is a much larger proportion of the alkaline and earthy phosphates in the urine, than is present in the blood; and this is liable to a further increase under circumstances to be presently alluded to.—The urine is normally acid, but the degree of its acidity has been shoAvn by Dr. Bence Jones to be continually changing, and to be considerably affected by food; being augmented by vegetable, and decreased by animal food. What the acid may be to which the acidity is due, is yet uncertain; possibly it is not always the same. 730. The organic compounds present in the Urinary secretion (in its healthy state at least) are undoubtedly the result of the waste or disin- tegration of the animal fabric; as Avell as (in certain cases) of the de- composition of constituents of the blood, which have never undergone conversion into organized tissue. Their unfitness to be retained within the system, is proved by the fatal results which speedily ensue when their elimination by the secreting process receives a check; and also by the crystalline form, in which the most characteristic of them pre- sent themselves,—such a form being altogether incompatible Avith the possession of plastic or organizable properties. Various well-defined compounds present themselves in the Urine of different classes of ani- mals ; and they are nearly all peculiarly rich in Nitrogen and deficient in Carbon, as compared with the Albuminous compounds. Thus, whilst the proportion of Nitrogen in Albumen is (by weight) as 1: 6-24 of the whole, it is in Urea as 1: 2*14, in Allantoin as 1: 2*21, in Kreatinine as 1: 2-69, in Uric acid as 1 : 3-00, and in Kreatine as 1 : 3-12. The only exception is in the case of Hippuric acid, which is discharged COMPOSITION OF THE URINE. 411 largely by herbivorous animals, and in which the proportion of Nitro- gen is as 1: 12-14. On the other hand, whilst the proportion of Car- bon in Albumen is as 1 : 1-80, it is in Kreatinine as 1 : 2-35, in Krea- tine as 1 : 2-73, in Uric Acid as 1 : 9-80, in Allantoin 1 : 3*87, and in Urea as 1 : 5*00. Here, again, Hippuric acid is exceptional; for its Carbon is as 1 : 1*57 of the whole, or in larger proportion than in Al- bumen.—We may say, then, that the characteristic components of the Urinary secretion are such products of the waste of the azotized tissues, as, from containing nitrogen in large proportion, are not adapted for elimination, either by the respiratory process, or by the biliary excre- tion. The only exception is in the case of Hippuric acid; and the large proportion of carbon and the small proportion of nitrogen contained in this substance, appear due to the great excess of non-azotized compounds in the food of the animals voiding it. 731. Of the compounds just enumerated, the most important, in Man, is that Avhich is named Urea. It exists in Urine in a state of perfect solution; and may be readily separated from it in the form of transparent colorless crystals, which have a faint and peculiar but not urinous odor. In its ultimate composition it is identical Avith Cyanate of Ammonia, being made up of 2 Carbon, 4 Hydrogen, 2 Nitrogen, and 2 Oxygen,—a formula much more simple than that of almost any other organic substance. The amount of Urea in the Urine is liable to very great variation, in accordance with the degree in which the disintegrating process has been taking place in the solid fabric; and also in confor- mity with the amount of azotized matter, which has been taken in as food. Supposing that the latter Avere so precisely adjusted to the wants of the system, as to supply only that Avhich is required for its mainte- nance, Ave might then measure the amount of previous waste, by the quantity of Urea present in the Urine. There can be no doubt as to the fact, that, other things being equal, the amount of Urea is greatly increased by any unusual exertion of the Muscular system; but such an increase cannot be invariably, or even usually, attributed to this cause; since it is equally certain, that any superfluity in the amount of azotized matter received into the blood, must be drawn off by the uri- nary excretion, and thus that an increase in the quantity of urea may be occasioned by an excessive use of proteine-compounds as articles of food. The average proportion of Urea, under ordinary circumstances as to diet and exercise, seems to be from 20 to 35 parts in 1000; but it may be raised to 45 parts by violent exercise, and to 53 parts by an exclusively animal diet; whilst it may fall as Ioav as from 12 to 15 parts, Avhen the diet is deficient in azotized matter. The total daily excretion of Urea in adult males seems to average about 430 grains, and that of females nearly 300 grains, but these averages may be Avidely departed from, on the side either of excess or diminution, according to the circumstances already noticed. It is interesting to observe, that children of eight years old excrete, on the average, half as much Urea as adults; -whilst, in very old persons, the quantity sinks to one-third, or even less. In proportion to their relative bulks, there- fore, children excrete at least tAvo or three times the quantity of urea that is set free by adults, and four or five times that which is excreted 412 OF SECRETION. by old persons,—a fact which corresponds with other indications of the far greater rapidity of interstitial change in the earlier periods of life, than in adult or advanced age. 732. There is an organic compound, nearly allied to Urea in compo- sition, but differing from it in its distinctly acid properties, and also in its comparative insolubility. This substance, termed Uric or Lithic Acid, forms but a small proportion of the solid matter of Human Urine in the state of health; but it is the chief element in the Urine of the lower Vertebrata; and its presence in too large a proportion is a fre- quent source of disease in Man. Its ultimate composition is 10 Car- bon, 4 Hydrogen, 4 Nitrogen, 6 Oxygen; it crystallizes in fine scales of a brilliant Avhite color and silky lustre; and it is so sparingly solu- ble in water, that at least 10,000 times its own Aveight of fluid is re- quired to dissolve it. In healthy Human urine, it is in a state of perfect solution, but it is precipitated by the addition of a small quantity of any acid, even the Carbonic: it is evident, therefore, that it is held in solu- tion by union with some base, and it seems probable that this base is ammonia. According to Dr. Bence Jones, the first precipitate thrown down by the addition of hydrochloric acid to ordinary urine, is urate of ammonia, which is less soluble in acid than in neutral or alkaline urine; and it is only after prolonged contact Avith the hydrochloric acid, that this salt is decomposed, and uric acid left behind. The solubility of urate of ammonia is much greater in warm than in cold urine ; and hence it frequently happens, that urine which is clear when voided, gives a pre- cipitate of urate of ammonia on cooling. 733. The proportion of Uric acid in healthy urine seldom rises above 1 part in 1000, and the quantity excreted daily is usually from 6 to 10 grains. The circumstances under Avhich it varies, however, have not been clearly determined. The absolute quantity in the urine bears no proportion to its acidity, nor is it indicated by the amount of deposit; for the acidity of the urine depends upon the presence of other acids; and a deposit of urate of ammonia may be due to an excess of acid, diminishing its solubility, rather than to an excess of the substance itself. Thus it may happen that a precipitate of urate of ammonia may be formed, when it is not present in any undue proportion, in con- sequence of the acid state of the urine ; Avhilst, on the other hand, there may be a large excess of urate of ammonia in the urine without any precipitate, if the urine should be alkaline. In disordered states of the system, there is often a great increase in the amount of uric acid in the urine; and there can be no doubt that this increase is partly controlla- ble by the reduction of the proportion of azotized matter in the food. In some of these cases, free uric acid is deposited, in consequence of the decomposition of the urate of ammonia by a large excess of acid in the urine. In attacks of gout, urate of soda is separated from the cir- culating blood, and is deposited in the tissues around the affected joints, forming the concretions termed "chalk'stones ;" and in this state of the system, uric acid may be detected in the blood. 734. There seems reason to believe that we are to regard Hippuric acid as a normal element of the Urine of Man; although it has been usually supposed to be restricted to the Herbivorous quadrupeds, Avhere COMPOSITION OF THE URINE. 413 it replaces Uric Acid. Its composition and properties are very diffe- rent from those of that substance. When pure, it forms long, transpa- rent, four-sided prisms; it is soluble in 400 parts of cold Avater, and dissolves readily at a boiling heat; and it has a strong acid reaction, with a bitterish taste. It is composed of 18 Carbon, 8 Hydrogen, 1 Nitrogen, and 5 Oxygen, with 1 equiv. of Water. When exposed to a high temperature, or subjected to the putrefactive process, it is partly converted into Benzoic acid ; and it is on the presence of the latter in putrefied Human Urine, that the belief in the existence of Hippuric acid in the same fluid when fresh, is chiefly grounded. It is a curious fact, that the administration of Benzoic acid causes the appearance of a large additional quantity of Hippuric acid in the Urine, so that its presence is then sufficiently evident; and this seems to be due to the union of the benzoic acid Avithin the system, either with glycocoll (§ 176), or Avith the elements which would have formed it; for one equivalent of benzoic acid, added to one of glycocoll, giA^es the precise equivalent of hippuric acid. It does not appear that, as once asserted, the adminis- tration of benzoic acid diminishes the quantity of uric acid normally present in the urine; but it seems to bring down the excess, where such exists, to about the normal quantity; and it may thus be employed with advantage in cases of uric acid gravel. 735. Much discussion has taken place, as to the normal presence of Lactic acid in the urine ; and the question cannot even now be regarded as completely determined. It is certain that the peculiar crystalline compound, procurable by treating the urine with zinc in solution, is not as Avas formerly maintained, a lactate of zinc; but that its composition is altogether different, as will be presently explained. But, on the other hand, it appears from the researches of Lehmann and others, that lactic acid is usually present in small quantity in healthy urine ; and that its quantity may be increased under such conditions, as either tend to augment the quantity of lactic acid in the blood, or to obstruct its elimination by the respiratory process. Thus an excess of farina- ceous food, which furnishes sugar and lactic acid faster than they can be throAvn off as carbonic acid and Avater; or an excess of exertion of the muscles, of whose disintegration lactic acid appears to be one of the results ; or pulmonary diseases, Avhich interfere with the normal aeration of the blood; all favor the appearance of lactic acid in the urine. It seems to be constant in herbivorous animals, and in patients suffering under chronic bronchitis, pulmonary emphysema, and similar disorders.—The urine of Man, more uniformly contains, however, tAvo substances termed Kreatine and Kreatinine; which seem to be the result of the degeneration of the muscles, as they may be obtained from the juice of raw flesh. The former of these, which is found in largest amount, is a neutral substance, crystallizing in long prisms, sparingly soluble in cold water, but dissolving readily in warm. By the action of strong acids, kreatine may be readily converted into kreatinine, which only differs from it in composition by containing two proportion- als less of the elements of water, but is a substance of very different chemical relations, having a strong alkaline reaction, and serving as a powerful organic base to acids. When long boiled with caustic baryta, 414 OF SECRETION. kreatinine is gradually resolved into urea ; and thus it would seem as if they hold an intermediate position between the components of the orga- nized tissues and the urea which may be considered as the ultimate pro- duct of their metamorphosis within the body. The peculiar crystalline compound just referred to, as having been formerly supposed to be lac- tate of zinc, has been shown to be really formed by a combination of zinc with a compound of kreatine and kreatinine. 736. Of the substances ranked under the head of Extractive Matters, little is definitely known; it appears, however, from recent researches, that they are peculiarly rich in carbon, and that they are liable to be greatly augmented, either by an excess of non-azotized matter in the food, or by any impediment to the action of the liver or lungs. A yellow extractive has been separated by Scherer, which he regards as proceeding from the final metamorphosis of the haematine of the blood ; and this seems nearly related to the purpurine, which sometimes gives a deep color to the sediment of urate of ammonia, and which is pecu- liarly liable to appear when the functional activity of the liver is below par. The ordinary coloring-matter of bile often presents itself in the urine in cases of jaundice: and, as Dr. Golding Bird has pointed out, there is a close similarity in composition between these three colored compounds, indicating a derivation from the same source. A sulphur extractive has also been obtained from the urine: in Avhich (as in the bile) there is a considerable proportion of free sulphur. 737. The Urine also contains a considerable amount of Saline matter, of which the acids as well as the bases are derived from the mineral kingdom; and the excretion of them, after they have served their pur- pose in the economy, appears to be one of the chief functions of the Kidney. Of these a part may find their way directly into the urine from the serum of the blood, when its Avater is being filtered off (so to speak) through the walls of the Malpighian capillaries; for although, from the peculiar properties of animal membranes (§ 196), the albumi- nous constituents of the serum are held back, the saline matter, which is in a state of perfect solution, must pass with the water. This is pro- bably the chief source of the large quantity of the muriates of soda and ammonia contained in the urine. But the Urinary secretion seems to be specially destined to eliminate the saline compounds, which are formed by the acidification of the Sulphur and Phosphorus, taken in with the proteine-compounds as food. These substances are united with Oxygen in the system, and are thus converted into Sulphuric and Phosphoric acids; which acids unite with alkaline bases, that were ingested in combination with Citric, Tartaric, Oxalic, and other organic acids; the latter undergoing decomposition within the system, and leaving the bases ready to unite with others. Such AYeakly-combined bases abound in the food of Herbivorous animals ; and their urine is almost invariably alkaline, the quantity of the Sulphuric and Phosphoric acids generated in the system not being sufficient to neutralize it. On the other hand, they are nearly absent in the food of the Carnivora; and their urine is therefore almost invariably acid, from the want of neutra- lization of the Sulphuric and Phosphoric acids. 738. The Alkaline Sulphates, whether taken in as such, or formed in COMPOSITION OF THE URINE. 415 the manner now described, are soluble enough to be always passed off in a fluid form; but this is not uniformly the case with the Phosphates, which are frequently deposited as sediments of a dead-Avhite aspect, sometimes crystalline, and sometimes wholly or partly amorphous. The crystalline sediment consists of the triple phosphate, or phosphate of ammonia and magnesia ; the amorphous contains an admixture of the phosphate of lime. The urine, Avhen these are deposited, is usually alkaline, sometimes very decidedly so ; and there is reason to think that, in many cases, this alkaline character, and the deposit of phosphatic sediments, are due to an alkaline secretion from the walls of the bladder and urinary passages, Avhich results from an irritable state of their mem- brane,—the urine, as secreted by the kidney, having its usual properties. That an alkaline condition of the urine, resulting from the presence of an unusual amount of bases, is capable of producing a phosphatic deposit, is shown by the simple experiment of adding ammonia to healthy urine, which will occasion a precipitation of the triple phosphate. 739. But there can be little doubt, that a frequent cause of the de- posit is excessive production of phosphate salts, arising from the in- creased Avaste or disintegration of Nervous matter, which takes place when it is in a state of unusual activity, either from intense thought, from prolonged exertion, or from continued anxiety. The general prin- ciples already set forth, in regard to the dependence of the functional activity of the Nervous Centres upon a supply of arterialized blood (§ 384), shoAV the probability that every act of theirs involves the oxy- genation of a certain quantity of nervous matter. In this oxygenation, phosphoric acid will be produced, from the large amount of phosphorus contained in the nervous matter; and this will unite in part with am- monia, Avhich is perhaps set free by the same metamorphosis, or is de- rived from other sources; and in part with bases derived from the food. The experience of every studious man must have shown him (if he make any observations on the matter at all) the frequent coincidence between the presence of phosphatic deposits in his urine, and an excess of men- tal labor ; and there are many instances on record, in which the peri- odical recurrence of the latter has been so invariably folloAved by the recurrence of the former, that no reasonable doubt can exist as to their mutual connexion. 740. It is very important for the successful treatment of those Uri- nary deposits, Avhich consist of the normal elements of the Urine,— namely, Lithic Acid, and the Phosphates,—that the leading facts al- ready stated should be borne continually in mind. In the first place, these sediments may depend upon the general condition of the fluid, and not upon any excess in the constituents of which they are com- posed ; thus a lithic deposit may result from the presence of an excess of some other acid in the urine ; and a phosphatic sediment may be produced by the excess of bases. In such cases, then, our treatment should be directed, not to diminish the quantity of the peculiar consti- tuents of the deposits, but to rectify the state of the Urine on which their precipitation depends. But, in the second place, the sediments may be present in such great amount, as to indicate that their consti- tuents are present in the urine to an excessive degree; and our treat- 416 OF SECRETION. ment must then be directed towards the diminution of the quantity pro- duced. Thus the tendency to lithic acid deposit may be frequently cured, by simply diminishing the quantity of azotized matter in the food; and the undue formation of the phosphates may be often kept in check by that mental repose, which is peculiarly required after long- continued and severe exercise of the intellectual faculties, or strong ex- citement of the feelings. 741. There is no doubt whatever, that the total suspension of the Urinary secretion is productive of rapidly-fatal results, from the accu- mulation of the elements of the secretion in the blood; and it would appear, that the tissue on Avhich their presence in the circulating fluid exerts the most injurious effects, is the Nervous. It is probable that Urea is the substance which is most directly concerned in producing the noxious influence; and we see an effort made by the system (so to speak) to get rid of it, in those cases in which a discharge of urinous fluid takes place by unusual channels, such as from the mucous mem- brane of the stomach, the mamma, the umbilicus, the nose, &c, when the usual secreting action of the Kidney has been suspended. Although the accounts of such cases have been treated with ridicule by some Physiologists, yet there seems no valid reason to discredit them, when it is borne in mind that, in persons who have died from the complete suspension of the secretion, effusions containing urea have been found in the serous cavities of the trunk, and in the ventricles of the brain. The poisonous influence of an accumulation of urea in the blood,when strongly exerted, produces in the first instance irregular or convulsive movements, which are dependent upon irritation of the Spinal system of nerves ; then loss of consciousness, depending upon the suspension of the powers of the Brain ; and lastly, complete suspension of the powers of the spinal system, so that the ordinary reflex actions cease, and life becomes extinct from the stoppage of the respiratory movements (§ 688). There is reason to believe, that many convulsive motions, for which no obvious cause can be assigned, have their origin in a disordered condi- tion of the blood, resulting from imperfect elimination of Urea; thus it has been ascertained that, in several cases of puerperal convulsions, urea was present in the blood; the functional power of the kidney being diminished by chronic disease. It is especially to be noticed, that most of the cases in which the urinary secretion is discharged through some irregular channel, occur in persons who have been sub- ject to those convulsive affections, which are commonly designated as hysterical; and that the discharge of a large quantity of urine through the natural channel, is often the termination of an hysterical paroxysm. It is desirable, therefore, that in all such obscure cases, the state of the urinary secretion should be carefully looked to. 4. Of the Cutaneous and Intestinal Glandulae. 742. The Glandulae Avhich are disposed in the substance of the Skin, and in the walls of the Intestinal canal, although individually minute, make up by their aggregation an excreting apparatus of no mean im- portance. . The Skin is the seat of two processes in particular ; one of CUTANEOUS GLANDULE. 417 which is destined to free the blood of a large quantity of fluid; and the other to draAv off a considerable amount of solid matter. To effect these processes, we meet with two distinct classes of glandulae in its substance; the Sudoriparous or sweat-glands; and the Sebaceous or oil- glands. They are both formed, however, upon the same simple plan; and can frequently be distinguished only by the nature of their secreted product. 743. The Sudoriparous or perspiratory glandulae form small oval or globular masses, situated just beneath the cutis, in almost every part of the surface of the body. Each is formed by the convolution of a single tube; which thence runs towards the surface as the efferent duct, making numerous spiral turns in its passage through the skin, and penetrating the epidermis rather obliquely, so that its orifice is covered by a sort of little valve of scarf-skin, which is lifted up as the fluid issues from it (Fig. 130). The convoluted knot, of which the gland Fig. 130. The anatomy of the Skin :—1. The Epidermis, showing the oblique laminae of which it is composed, and the imbricated disposition of the ridges upon its surface. 2. The Rete Mucosum, or deep layer of the epi- dermis. 3. Two of the quadrilateral papillary clumps, such as are seen in the palm of the hand or sole of the foot; they are composed of minute conical papillae. 4. The deep layer of the cutis, the Corium. 5. Adi- pose cells. 6. A Sudoriparous gland with its spiral duct, such as is seen in the palm of the hand or sole of the foot. 7. Another Sudoriparous gland with a straighter duct, such as is seen in the scalp. 8. Two hairs from the scalp, enclosed in their follicles; their relative depth in the skin is preserved. 9. A pair of Sebaceous glands, opening by short ducts into the follicle of the hair. consists, is copiously supplied with blood-vessels. On the palm of the hand, the sole of the foot, and the extremities of the fingers, the aper- tures of the perspiratory ducts are visible to the naked eye, being situ- ated at regular distances along the little ridges of sensory papillae, and giving to the latter the appearance of being crossed by transverse lines. According to Mr. Erasmus Wilson, as many as 3528 of these glandulae exist in a square inch of surface on the palm of the hand; and as every tube, when straightened out, is about a quarter of an inch in length, it folloAvs that in a square inch of skin from the palm of the hand, there exists a length of tube equal to 882 inches, or 73J feet. • The number 418 OF SECRETION. of glandulae in other parts of the skin is sometimes greater, but gene- rally less than this; and, according to Mr. Wilson, about 2800 may be taken as the average number of pores in each square inch throughout the body. Now the number of square inches of surface, in a man of ordinary stature, is about 2500; the number of pores, therefore, is seven millions ; and the number of inches of perspiratory tubing would thus be 1,750,000, or 145,833 feet, or 48,611 yards, or nearly 28 miles. 744. From this extensive system of glandulae, a secretion of watery fluid is continually taking place; and a considerable amount of solid matter also is draAvn off by the epithelium-cells that line the tubuli. Under ordinary circumstances, the fluid is carried off in the state of vapor, forming the insensible perspiration; and it is only when its amount is considerably increased, or when the surrounding air is already so loaded .with moisture as to be incapable of receiving more, that the fluid remains in the form of sensible perspiration upon the surface of the skin. It is difficult to estimate the proportion of solid matter con- tained in this secretion; partly on account of the great variations in the amount of fluid eliminated by the Sudoriparous glands, which are governed by the temperature of the skin ; and partly because the secre- tion can scarcely be collected for analysis free from the sebaceous and other matters which accumulate on the surface of the skin. According to Anselmino it varies from f- to If per cent.; and consists in part of lactic acid, to which the acid reaction and sour smell of the secretion are due; in part of a proteine-compound, which is probably furnished by the epithelium-cells that line the tubes; and in part of saline matters, directly proceeding from the serum of the blood. Urea has been re- cently detected in the perspiration of the inhabitants of warm climates. 745. The amount of fluid excreted from the skin is almost entirely dependent upon the temperature of the surrounding medium ; being in- creased with its rise, and diminished with its fall. The object of this variation is very evident; being the regulation of the temperature of the body. When the surface is exposed to a high degree of external heat, the increased amount of fluid set free from the perspiratory glands becomes the means of keeping down its own temperature; for this fluid is then carried off in a state of vapor, as fast as it is set free; and in its change of form, it withdraAvs a large quantity of caloric from the surface. But if the hot atmosphere be already loaded with vapor, this cooling poAver cannot be exerted; the temperature of the body is raised, and death supervenes, if the experiment be long continued. The cause of the increased secretion is probably to be looked for in the increased determination of blood to the skin, which takes place under the stimulus of heat.—The entire loss by Exhalation from the lungs and skin, during the twenty-four hours, seems to average a little above 2 lbs. In a warm dry atmosphere, however, it has been found to rise to as much as 5 lbs. whilst in a cold damp one, it may be lowered to If lb. Of this quantity, the pulmonary exhalation is usually somewhat less than one-third, and the cutaneous somewhat more than two-thirds; but when the quantity of fluid lost is unusually great, the increase must be chiefly in the Cuta- neous exhalation; since, as already pointed out (§ 701), the amount of CUTANEOUS EXHALATION. 419 exhalation from the lungs is not influenced by the external tempera- ture, but only by the degree in which the surrounding air is previously saturated with moisture. 746. The variations in the amount of fluid set free by Cutaneous and Pulmonary Exhalation, are counterbalanced by the regulating action of the Kidney; which allows a larger proportion of water to be strained off in a liquid state from the blood-vessels, as the Exhalation is less,—and vice versd. The Cutaneous and Urinary excretions seem to be vicarious, not merely in regard to the amount of fluid which they carry off from the blood, but also in respect to the solid matter which they eliminate from it. It appears that at least 100 grains of effete azotized matter are daily thrown off from the skin; and any cause which checks this excretion, must increase the labor of the Kidneys, or produce an accumulation of noxious matter in the blood. Hence attention to the functions of the skin, at all times a matter of great importance, is peculiarly required in the treatment of Urinary diseases ; and it will be often found that no means is so useful in removing the lithic acid deposit, as copious ablution and friction of the skin, com- bined with exercise. When the Exhalent action of the skin is com- pletely checked by the application of an' impermeable varnish, the effect is not (as might be anticipated) an elevation of the temperature of the body; on the contrary it is lowered, in consequence, as it would appear, of the interruption to the aeration of the blood through the skin, Avhich is a function of such importance in the lower animals (§ 671), and of no trifling account in Man; and in a short time, a fatal result ensues. A partial suppression by the same means gives rise to febrile symptoms, and to Albuminuria, or escape of the albuminous part of the liquor sanguinis into the urinary tubes, in consequence (it would appear) of the increased determination which then takes place towards the Kid- neys. These facts are interesting, as throwing light upon the febrile disturbance which accompanies those cutaneous diseases that affect the whole surface of the skin at once, and interfere Avith its functions; and as partly accounting also for the Albuminuria which frequently mani- fests itself during their progress, especially in Scarlatina. 747. The Skin is likewise furnished with numerous Sebaceous glands, which are distributed more or less closely over the whole surface of the body; being least abundant Avhere the Perspiratory glandulae are most numerous; and vice versd. They are altogether absent on the palms of the hands and the soles of the feet; and are particularly frequent in the skin of the face and in the scalp. They differ greatly in size and in de- gree of complexity; sometimes consisting of short straight follicles; some- times closely resembling the Sudoriparous glandulae, the tubes, however, being usually straighter and wider; and being sometimes much more complex in structure, consisting of a number of distinct sacculi clustered around the extremity of a common duct, into which they open, and form- ing little arborescent masses about the size of millet-seeds. In some situa- tions they acquire still greater complexity. Thus the Meibomian glan- dulae, which are found at the edges of the eyelids, and which secrete an unctuous matter for their lubrication, are long sacculi branching out at the sides (Fig. Ill); and the glandulae of the ear passage, which secrete 420 OF SECRETION. its cerumen or Avaxy matter, and which belong to the general Sebaceous system, are formed of long tubes, highly contorted, and copiously sup- plied with blood-vessels. In the hairy parts of the skin, we usually find a pair of Sebaceous follicles opening into the passage through which every hair ascends (Fig. 130, 9). The purpose of the sebaceous secretion is evidently to prevent the skin from being dried and cracked by the influence of the sun and air. It is much more abundant in those races of mankind which are formed to exist in warm climates, than in the races that naturally inhabit cold countries; and the former are accustomed to aid its preservative power, by lubricating their skin with vegetable oils of various kinds; which process they find to be of use, in protecting it from the scorching influence of the solar rays.—The Seba- ceous follicles are frequently the residence of a curious parasite, the Demodex folliculorum, which is stated by Mr. Erasmus Wilson to be present in great numbers in the skin of almost all inhabitants of large towns; the activity of their cutaneous glandular system being much checked by the want of free exposure to pure air, and by inert habits of life. 748. To what extent the Sebaceous secretion can be regarded as destined to free the Blood from deleterious matters, it may not, perhaps, be very easy to say; but with regard to the functions of the Skin taken altogether, as a channel for the elimination of morbific matters from the blood, it is probable that they have been much underrated; and that much more use might be made of it in the treatment of diseases,— especially of such as depend upon the presence of some morbific matter in the circulating current,—than is commonly thought advisable. We see that Nature frequently uses it for this purpose; a copious perspira- tion being often the turning-point or crisis of febrile diseases, removing the cause of the malady from the blood, and allowing the restorative powers free play. Again, certain forms of Rheumatism are charac- terized by copious acid perspirations; and instead of endeavoring to check these, we should rather encourage them, as the best means of freeing the blood from its undue accumulation of lactic acid. And it is recorded that in the "sweating sickness," which spread throughout Europe in the 16th century,, no remedies seemed of any avail but diaphoretics; which, aiding the powers of nature, concurred with them to purify the blood of its morbific matter. The hot-air bath, in some cases, and the Avet sheet (which, as used by the Hydropathists, is one of the most powerful of all diaphoretics), will be probably employed more extensively as theurapeutic agents, in proportion as the importance of acting on the Skin, as an extensive collection of glandulae, comes to be better understood. The absurdity of the "Hydropathic" treatment consists in its indiscriminate application to a great variety of diseases; no person who has watched its operation, can deny that it is a remedy of a most powerful kind; and if its agency be fairly tested, there is strong reason to belieA^e, that it will be found to be the most valuable curative means we possess for various specific diseases, Avhich depend upon the presence of" a definite "materies morbi" in the blood, especially Gout and chronic Rheumatism; as well as for that depressed state of the CUTANEOUS EXHALATION. 421 general system, Avhich results from the " wear and tear" of the bodily and mental powers. 749. The Mucous surface of the Alimentary Canal is furnished, like the skin, with a vast number of glandulae, varying in complexity, from the simple follicle, to a mass consisting of numerous lobules opening into a common excretory duct. The functions of these, as already pointed out, are equally various. The simple follicles appear destined, for the most part, to secrete the protective mucus, which intervenes between the membranous wall and the substances contained in the canal, and which serves to protect the former from the irritating action of the latter. The more complex follicles of the Stomach elaborate the Gastric fluid, which is the prime agent in the digestive process (§ 496). The still more elaborate glandulae of Brunner, situated in the walls of the duodenum, also seem to furnish a product which is con- cerned in the digestive operation (§ 480). But there is strong reason to believe, that the function of the Peyerian glandulae, which beset the walls of the lower part of the intestinal canal, is purely excretory; and that they are destined to eliminate putrescent matters from the blood, and to convey them, by the readiest channel, completely out of the body. That the putrescent elements of the faeces are not immediately derived from the food taken in, so much as from the secreting action of the intestinal glandulae, appears from this consideration ;—that faecal matter is still discharged, even in considerable quantities, long after the intestinal tube has been completely emptied of its alimentary con- tents. We see this in the course of many diseases, when food is not taken for many days, during which time the bowels are completely emptied of their previous contents by repeated evacuations ; and what- ever then passes, must be derived from the intestinal Avails themselves. Sometimes a copious flux of putrescent matter continues to take place spontaneously; Avhilst it is often produced by the agency of purgative medicine. The " colliquative diarrhoea," which frequently comes on at the close of exhausting diseases, and Avhich usually precedes death by starvation, appears to depend, not so much upon a disordered state of the intestinal glandulae, as upon the general disintegration of the solids of the body, which calls them into extraordinary activity, for the purpose of separating the decomposing matter. 750. Thus we perceive, that we have here, also, to watch for the indications of Nature; and that this extensive system of intestinal glandulae, being the principal channel for the elimination of putrescent matters from the blood, should be especially attended to, when there is reason to think-that such matters are present in too large an amount. Hence, when diarrhoea is already existing, we may often do more good by alloAving it to take its course, or even by increasing it by the agency of purgative medicines, than by attempting to check it, and thus causing the retention of the morbid matter in the circulating current. But, on the other hand, it is necessary to bear in mind the extreme irritability of the intestinal mucous membrane; and carefully to avoid exciting it, when it is already in excess, or when there is danger that it will supervene,—as in that form of Fever in Avhich there is a peculiar 422 OF SECRETION. liability to inflammation and ulceration of the walls of the alimentary canal, and of their contained glandulae. 5. General Summary of the Excreting Processes. 751. We have now passed in review the various processes, by Avhich the products of the disintegration of the animal tissues are carried off; and we have seen that the necessity for their removal is much more urgent than for their replacement. A cold-blooded animal may subsist for some weeks, or even months, without a fresh supply of food, the waste of its tissues being so small, if it remain in a state of rest, as to be quite compatible with the continuance of its life ; and a warm- blooded animal may live for many days or even weeks, provided that it has in its body a store of fat sufficient to keep up its heat by the com- bustive process. But in either case, if the exhalation of carbonic acid by the lungs, the elimination of biliary matter by the liver, the separa- tion of urea or uric acid by the kidneys, or the withdraAval of putrescent matter by the intestinal glandulae, be completely checked, a fatal result speedily ensues ;—more speedily in warm-blooded animals, than in those which cannot sustain a high independent temperature, on account of the greater proneness to decomposition in the bodies of the former, than in those of the latter;—and more speedily in the latter, when their bodies are kept at an elevated temperature by the warmth of the sur- rounding medium, that when the degree of heat is so low, that there is little proneness to spontaneous change in the substance of their bodies. 752. It may be taken as a general principle, in regard to the Ex- creting processes (including Respiration), that they have a threefold purpose ;—in the first place, to carry off the normal results of the waste or disintegration of the solid tissues, and of the decomposition of the fluids;—in the second place, to draAv off the superfluous alimentary matter, which though received into the circulating current, is not con- verted into solid tissue, in consequence of the want of demand for it; —and in the third place, to carry off the abnormal products, which occasionally result from irregular or morbid changes in the system. Thus by the Lungs are excreted a large amount of carbon, and some hydrogen, resulting from the disintegration of the tissues, especially the nervous and muscular; the same elements, in animals that take in a large proportion of farinaceous or oleaginous aliment, may be derived im- mediately from the food, without any previous conversion into solid tissue; and there can be little doubt that the respiratory function is also an impor- tant means of purifying the blood from various deleterious matters, either introduced from without (such as narcotic poisons), or generated within the body (such as the poison of fever).* And it is important to bear this last circumstance in mind ; since it enables us to understand how, if time be given, the system frees itself from such noxious sub- stances ; and points out the duty of the medical attendant to be rather that * There is strong reason to believe that, in many instances, a small amount of poi- sonous matter introduced from without, in the form of a contagion or miasm, may lead, by a process resembling fermentation, to the production of a large quantity of similar noxious substances in the animal fluids. GENERAL RELATIONS OF EXCRETING PROCESSES. 423 of supporting the powers of the body by judiciously-devised means, and of aiding the elimination of the morbid matter through the lungs and skin by a copious supply of pure air, than of interfering more actively to promote that which Nature is already effecting in the most advanta- geous manner. 753. In like manner, the Liver is charged with the separation of hydrocarbon, in a fluid form; for which a supply of oxygen is not requisite. This product is partly derived from the waste of the sys- tem ; but the arrangement of the biliary vessels leads to the belief, that part of it may be at once derived from crude matter, taken up by the mesenteric veins, and eliminated from them by the hepatic cells, with- out ever passing into the general circulation. And various facts seem to indicate, that the Liver is also destined to remove from the blood extraneous substances, which are noxious to it. Thus, in cases Avhere death has resulted from the prolonged introduction of the salts of Cop- per into the system, a considerable amount of that metal has been ob- tained from the substance of the gland. 754. It has been already pointed out (§ 726), that in those tribes of animals Avhose respiration is feeble, a considerable part of the mass of the liver is composed of fatty matter; and this condition may be in- duced, as a state of disease, in Avarm-blooded, energetically-respiring Birds and Mammals, by impediments to the due performance of the respiratory process. This is remarkably shown in the treatment of the geese which are to furnish the celebrated Strasburg pates. The unfor- tunate bird is closely confined at a high temperature; so that the respiration is reduced to its minimum amount, by the combined effects of warmth and muscular inac- tion ; and it is then crammed with maize, which contains a large amount of oily matter. The consequence is, that its liver soon enlarges, and becomes unusually fatty; its cells being gorged with oil-globules, instead of each con- Hepatic cells ... b b b a1 & , ' j ., . ,, i gorged with fat: taming no more than one or two : and it is then ready — a, atrophied for the epicureans Avho set so high a value on the pate pose^giobui'es341" de foie gras. A similar diseased condition of the liver frequently presents itself in Man, in connexion with chronic dis- orders of the respiratory organs, Avhich diminish the amount of hydro- carbon eliminated through their agency ; this " fatty liver" is peculiarly common in the advanced stages of Phthisis. It may arise, however, from a local disorder of nutrition, such as that which produces the fatty degeneration of other organs. Such a fatty degeneration may occur in the Kidney, for example, as a consequence of inflammation of its tissues. 755. With regard to the Kidneys, it has been already pointed out that they are the special emunctories of the azotized products of the decomposition of the tissues; and that they serve also to convey away the overplus of such earthy and alkaline salts, as are readily soluble. Moreover, it has been shown that the surplus proteine-compounds, which are not required for the nutrition of the system, must be excreted by their agency, after having been metamorphosed into urea. And we have now to notice, that other matters of an injurious charac- 424 OF SECRETION. ter, Avhether introduced from without, or generated within the system, are drawn off by the same channel. Thus the saline compounds, taken up by the absorbent process (§ 493), are for the most part set free through these organs; especially when their properties are such, as to excite the action of the kidneys in a peculiar degree. Thus, Prussiate of Potash has been detected in the urine, within one minute after it has been introduced into the stomach. It has been sometimes noticed that Iodide of Potassium, when administered as a medicine, is retained within the body for some days, producing extensive cutaneous erup- tions, or some other unusual consequence ; and that it then suddenly begins to pass off by the kidneys, and is excreted in very large quan- tities. Further, it has been shown by Dr. Letheby, that poisonous substances (such as arsenious acid), introduced into the system in small but frequently-repeated doses, may be carried out of the body with such rapidity as to be prevented from exerting their injurious effects, provided that diuretics be administered at the same time. The effect of the inhalation of the vapor of turpentine, even in a very diluted state, in speedily imparting to the urine the odor of violets, is an evidence that not merely the actual substances imbibed, but new and peculiar compounds to which they give rise, are thus eliminated by the Kidneys. 756. The most singular variations in the excretory function of the Kidneys are seen, however, when the Urine is charged with substances Avhich are not only foreign to it, but are altogether foreign to the r~healthy body. The most remarkable instance of this is seen in the disease termed Diabetes, in which a large quantity of Sugar is formed, either directly from the food, or by the disintegration of the solid / tissues; and in which this compound is eliminated by the Kidneys, imparting to the urine a saccharine taste. And another example of the same general fact is seen in the " oxalic diathesis," in which an unusual arrangement of the elements that usually form urea or uric acid, gives rise to a new and peculiar compound, oxalate of ammonia; this being drawn off by the kidneys, and being decomposed by the calcareous matter present in the urine, gives rise to a deposit of oxalate of lime. In the treatment of such diseases, our attention must be given, not so much to the secreting organ, as to the condition of the system at large, of which the character of the secreted product is the indication or exponent. 757. To what has already been stated in regard to the exhalant functions of the Lungs and Skin, it may be added that many states of disease are marked by an unusual odor emitted from the body; and there can be little doubt that the peculiar odorous matter is pre-formed in the blood,—as Ave know that the ordinary scent of any species (whether Man, Dog, Horse, Goat, &c.) may be set free from the blood of that species, by the addition of sulphuric acid. The existence of such odors, therefore, is not to be attributed to disordered function in the excreting organs; but to the formation of morbid products in the interior of the body, which these organs do their best to remove. The foetid breath, which frequently accompanies an attack of indigestion, is another instance of the power of the lungs to eliminate, not merely HEAT OF ANIMALS AND PLANTS. 425 Carbonic acid, but other products of the changes in composition which the food undergoes when introduced into the system. 758. The same remarks apply, and with yet greater force, to the Intestinal glandulae; whose function it is, not merely to remove the putrescent matter ordinarily formed by the disintegration of the tis- sues, or by the decomposition of unassimilated food, but also to draw off the still more offensive products of such changes as take place in disease. Thus there are conditions of the system, in which, Avithout any well-marked disorder, the faeces emit a peculiarly foetid odor ; and with these is almost always associated a depressed state of mind. Noav it can scarcely be doubted, that the real fault is here rather in the early part of the nutritive operations, than in the excretory func- tion ; and that the foetor of the contents of the intestine depends upon the undue formation of putrescent matter in the system, which, by taint- ing the blood, causes its action upon the brain to become unhealthy. The object of the physician will be here to eliminate the morbid pro- duct, by the moderate use of purgatives; and so to regulate the diet and regimen, as to correct the tendency to its formation.—An excessive foetor in the evacuations, as well as in the exhalations from the skin and lungs, is peculiarly characteristic of those very severe forms of typhus (noAV, happily, of comparatively rare occurrence), which are termed putrid fevers.^. Here the whole of the solids and fluids of the body appear to have an unusual tendency to decomposition, in conse- quence of the introduction of some morbid agent, which acts as a fer- ment ; and the system attempts to free itself from the products of that decomposition, by the various organs of excretion, particularly the Skin and Intestinal surface. 759. It is of great importance that the Student should form clear conceptions on this subject; and that he should not (as too often hap- pens), by directing his remedies to the mere symptoms or results of a disease, act in precise opposition to the natural tendency of the system to free itself from some unusual noxious matter, through those channels Avhich are ordinarily destined to carry off only the regular products of its disintegration. CHAPTER X. OF THE DEVELOPMENT OF HEAT, LIGHT, AND ELECTRICITY, IN THE ANIMAL BODY. 760. It has been shown, in an earlier part of this volume (chap, ii.), that all Vital actions require a certain amount of Heat for their per- formance ; and that there is a great variety amongst the different classes of Animals, both in regard to the degree of Heat which is most favorable to the several processes of their economy, and in regard to their own power of sustaining it, independently of oscillations in the 426 DEVELOPMENT OF HEAT, ETC. temperature of the surrounding medium. As a general rule, the In- vertebrated animals are cold-blooded; that is, they have little or no power of sustaining an independent temperature. The degree of energy of their vital actions entirely depends, therefore, upon the warmth they receive from the air or water they inhabit; they have no power of resisting the depressing influence of cold ; and they are gene- rally so organized, as to pass into a state of complete inaction or torpi- dity, when the temperature sinks below a, certain point,—after gradu- ally becoming more and more inert with every diminution in the heat of their bodies. The same is true, also, of most Fishes and Reptiles : but the animals of the former class, from the more equable temperature of the medium they inhabit, are not so liable to be reduced to inaction as the latter ; being usually so organized, as to retain their activity so long as the water around them continues liquid; and being actually imbedded in a frozen state, when the water around them is converted into ice, without the loss of their vitality. There are certain Fishes, hoAvever,—such as the Thunny, Sword-fish, and other large species of the Mackerel tribe,—which are able to sustain a temperature consi- derably above that of the sea they inhabit; thus in the Bonito, the heat of the body has been found to be 99°, Avhen the temperature of the surrounding sea was but 80J°. It is not probable, however, that the temperature of the body would be kept up to the same standard, if that of the sea should be considerably lowered ; but it Avould proba- bly remain at from 18° to 20° above the latter. And in like manner, it has been noticed that many of the more active Reptiles possess the power of sustaining the temperature of their bodies at 10° or 15° above that of the surrounding air. 761. The classes of animals which are especially endowed with the power of producing and maintaining heat, are Insects, Birds, and Mammalia. The remarkable variations which present themselves in the temperature of the first of these classes, and the connexion of these variations with the condition of the animals in regard to activity or repose, have already been sufficiently noticed (§ 123).—The tempera- ture of Birds is higher than that of any other class of animals; varying from 100° to 111° or 112°. The loAvest degree is found in some of the aquatic species, as the Gull, and in those which principally live on the ground, as the Fowl tribe ; and the highest in the birds of most active flight, as the SAvalloAV. The temperature of the Mammalia seems to range from about 96° to 104° ; that of Man has been observed as low as 96J°, and as high as 102°. The variations are dependent in part upon the temperature of the external air; but are influenced also by the general condition of the body as to repose or activity, the period of the day, the time that has elapsed since a meal, &c A someAvhat larger amount of caloric is generated during the day, than in the night; and the body is usually warmer, by a degree or tAvo, at noon, than at midnight. There is also a slight increase during the digestion of a meal; and exercise is a powerful means of raising the temperature.— The range of temperature is much greater in disease ; thus the thermo- meter has been seen to rise to 106° in Scarlatina and Typhus, and to HEAT OF ANIMALS AND PLANTS. 427 110?° in Tetanus; whilst it has fallen to 82° in Spasmodic Asthma, and to 77° in Cyanosis and Asiatic Cholera. 762. In searching for the conditions on which this production of heat within the Animal body is dependent, it is very important to bear in mind, that a similar generation of Caloric may be observed in the Vege- table kingdom. It appears from the most recent and exact experi- ments, that all living Plants are somewhat warmer than similar dead plants exposed to the same atmosphere; and that the elevation is the greatest in the leaves and young stems, in which the most active vital changes are taking place. But the most decided production of heat occurs in the flowering of certain Plants, such as the Arum, which have large fleshy receptacles, on which a great number of blossoms are crowded ; thus a thermometer placed in the centre of five spadixes of the Arum cordifolium has been seen to rise to 111°, and one placed in the midst of twelve spadixes has risen to 121°, whilst the temperature of the surrounding air was only 66°. In the germination of seeds, also, a great elevation of temperature occurs, which is rendered most evident by bringing together a number of seeds, as in the process of malting, so that the caloric is not dissipated as fast as it is generated; the ther- mometer, placed in the midst of a mass of seeds in active germination, has been seen to rise to 110°. 763. Thus it is evident that the chemical changes which are involved in the operations of Nutrition, are capable of setting free a large amount of heat; which, although ordinarily dissipated from the vegetating sur- face too speedily to manifest itself, becomes sensible enough, when this rapid loss is checked. If we further examine into the nature of the chemical changes which appear most concerned in this elevation of temperature, we find that they uniformly consist in the combination of the carbon of the plant with the oxygen of the atmosphere; so that a large quantity of carbonic acid is formed and set free, precisely in the manner of the Respiration of Animals. This process is so slowly per- formed, in the ordinary growth of Plants, that it is concealed (as it were) by the converse change,—the fixation of carbon from the carbonic acid of the atmosphere, under the influence of light (§ 83). But it takes place with extraordinary energy during flowering and germination; a large quantity of carbon being set free, by union with the oxygen of the air; and the starchy matter of the receptacle, or of the seed, being con- verted into sugar. Now it has been ascertained by careful experiments, that the amount of heat generated is in close relation with the amount of carbonic acid set free ; and that, if the formation of the latter be pre- vented, by placing the flower or the seed in nitrogen or hydrogen, no elevation of temperature takes place ; whilst, if the process be stimu- lated by pure oxygen, so that a larger proportion of carbonic acid is evolved, the elevation of temperature is more rapid and considerable than usual. 764. Upon examining into the conditions under which Caloric is gene- rated in the Animal body, Ave find them essentially the same. Wherever the temperature of the body is maintained at a regular standard, so as to be independent of variations in the Avarmth of the surrounding me- dium, we find a provision for exposing the blood most freely to the in- 428 ANIMAL HEAT. fluence of oxygen, and for extricating its carbonic acid ; thus in Birds and Mammals, the blood is distributed, in a minute capillary netAvork, on the walls of the pulmonary air-cells, the gaseous contents of Avhich are continually renewed; and in Insects, the air is carried into every part of the body by the ramifying tracheae. We constantly find a pro- portion between the amount of heat evolved, and that of carbonic acid generated; this is peculiarly evident in Insects, whose respiration and calorification vary so remarkably (§ 123); but it is also proved by comparing the amount of carbonic acid generated by warm-blooded ani- mals, Avhen the external temperature is low, and when more heat must be evolved to keep the temperature of their bodies up to its proper stan- dard, with that generated by the same animals in a warmer atmosphere, when the proper animal heat is diminished in amount (§ 691). 765. The sources of the Carbonic Acid thrown off by the lungs, have been already pointed out (chap, viii.) : it is partly deriAred from the metamorphosis of the tissues ; but partly, in all but purely carnivorous animals, more directly from the non-azotized portion of the food. The precise mode in which the carbon thus supplied is united with the oxy- gen derived from the atmosphere, is not yet known ; but it is certain that, in whatever manner the combination may take place, a certain measure of caloric must be generated. It appears, however, from various ex- periments, that the whole quantity of caloric generated by an animal in a given time, is greater than that which would be evolved by the com- bustion of the carbon, included in the carbonic acid evolved during the same time. Hence it is evident that other chemical processes occurring within the body are concerned in the maintenance of the temperature; and it is not difficult to point to some of these. It is probable, in the first place, that some of the Hydrogen of the food may be " burned off" by union with the oxygen of the atmosphere, so as to form part of the water which is exhaled from the lungs. Again, the sulphur and phos- phorus of the food are converted, by oxygenation, into sulphuric and phosphoric acids; in which process, heat must be generated. In the composition of urea, moreover, oxygen is present in much larger propor- tion, than it is in the proteine-compounds by the metamorphosis of which it is formed; so that in its production too, caloric will be generated. In fact it may be stated as a general truth, that the whole excess of the oxy- gen absorbed, over that Avhich is contained in the carbonic acid exhaled (§ 689), must be applied to purposes in the laboratory of the system, in which caloric will be disengaged. Still, the amount of Carbonic Acid exhaled must always be the measure of the chemical processes, by which heat is generated in the body; because it is itself the result of the chief of these processes (the union of carbon and oxygen), and because the surplus amount of oxygen which is absorbed, and which is applied to other purposes, is closely related to it. 766. The power of maintaining a high independent temperature is usually much less in young warm-blooded animals, than in adults. There are considerable variations in this respect, however, amongst different species; for where the young animal is born in such an ad- vanced condition, as to be thenceforth almost independent of parental assistance, it is capable of maintaining its own temperature; but where REGULATION OF HEAT IN MAN. 429 it is born in such a state, as to require to be supplied with food by the parent for some time, it is also more or less dependent upon the warmth imparted to it from the parental body. This is peculiarly the case with the young of the Human species, which is longer dependent upon parental aid, than that of any other animal. In the case of children born very prematurely, the careful sustenance of their heat is one of the points most to be attended to in rearing them; and even the most vigorous infants, born at the full time, are far from being able to keep up their proper standard without assistance, if exposed to a cool atmo- sphere. It has been ascertained that, during the first month of infant life, the mortality in winter is nearly double that of summer,—being 1-39 in January to 0*78 in July; and this striking difference cannot be attributed to any other cause, than the injurious influence of external cold, which the calorifying powers of the infant do not enable it to resist. As age advances, the power of generating heat increases, and the body becomes much more independent of external vicissitudes; so that, in adult life, the winter mortality is to that of summer, only as 1'05 to 0-91, or less than one-sixth more. In advanced age, the calori- fying power again diminishes; and this we should anticipate from the general torpor of the nutritive operations in old persons. Between 50 and 65 years of age, the relative winter and summer mortality are nearly as in the first month of infancy; and at 90 years, the average mortality of winter is much more than twice that of summer, being as 1-58 to 0-64. 767. It appears that there is a difference in calorifying power, not merely at different ages, but at different seasons: the amount of heat generated in summer not being sufficient, in many animals, to prevent the body from being cooled down by prolonged exposure to a tempera- ture, which is natural to them in winter. To what extent this is the case with Man, it is difficult to say. His constitution is distinguished by its power of adapting itself to circumstances; and he can live under extremes of temperature more wide than those, which most other ani- mals can endure (§ 113). Whether in the torrid zone, or in the arctic regions, he can maintain his healthy condition under favorable circum- stances; in each case his natural appetite leading him to the use of that kind and amount of food, which is best suited to the wants of his system. But the longer he has been habituated to a very warm or a very cold climate, the more difficult he at first finds it to live comforta- bly in one of an opposite character; as his constitution, having become adapted to one particular set of circumstances, requires time to accom- modate itself to an opposite one. 768. The means by which the heat of the body is prevented from rising above its normal standard, even in the midst of a very high tem- perature in the surrounding air, are of the most simple character. The excreting action of the skin is directly stimulated by the application of warmth to the surface; and the fluid which is poured forth, being im- mediately vaporized, converts a large quantity of sensible caloric into latent, and thus keeps down the temperature of the skin. By this pro- vision, the body may be exposed with impunity to dry air of 600° or more, so long as the supply of fluid is maintained. But it cannot long 430 ANIMAL LUMINOSITY. sustain exposure to air saturated with vapor, even though it may not be many degrees hotter than the body; because the cooling act of evapo- ration from the skin cannot then be carried on. 769. The evolution of Light is a very interesting phenomenon, chiefly witnessed among the lower animals, and usually supposed not to occur in any class above Fishes. It is particularly remarkable among the Radiata and inferior Mollusca. A large proportion of the Acalephce, or Jelly-fish tribe, possess the property of luminousness in a greater or less degree; and it is to small animals of this class, which sometimes multi- ply to an amazing extent, that the beautiful phenomenon of phosphores- cence of the sea is chiefly due. In the midst of the soft diffused light thus occasioned, brilliant stars, ribands, and globes of fire are frequently seen; these appearances being due to the luminosity of the larger species of the same tribe, or to that of other marine animals.—Some of the most remarkable examples of luminosity, in regard to the brilliancy of the light emitted, occur in the class of Insects. Here the emission is confined to one portion of the body, or to two or more isolated spots, instead of being diffused over a larger surface; and it is proportionally increased in intensity. The phenomenon of Animal Luminousness appears usually attributable to the formation of a peculiar secretion; which in many instances, continues to shine after removal from the animal, so long as it is exposed to the influence of oxygen: and it seems not unreasonable to believe that it depends upon a slow process of combustion, analogous to that which takes place when phosphorus is exposed to the air. There is a special provision in Insects, for convey- ing a large supply of air through the peculiar substance, which is depo- sited beneath the luminous spots; and the power which Glow-worms, Fire- flies, &c, possess, of suddenly extinguishing their light and as suddenly reneAving it, seems to depend upon their control over the air-aperture or spiracle by which air is admitted, the stoppage of the supply of air causing the immediate cessation of the luminousness, and its readmission occasioning a renewal of the process on which it depends.—It is proba- ble, however, that in certain cases the luminosity is rather of an electrical character. There are several of the smaller Annelida or marine Worms, which are brilliantly luminous when irritated; the luminosity having the character, however, of a succession of sparks, rather than of a steady glow. It appears from the experiments of M. Quatrefages, that this pecu- liar luminosity is the especial attribute of the muscular system; and that it is produced with every act of muscular contraction in these animals. 770. Although no such luminosity is commonly manifested in any of the higher Vertebrata, or in Man, yet there are well-authenticated cases, in which the phenomenon has presented itself in the living Human subject,*—luminous emanations from dead animal matter being of no unfrequent occurrence. In most of these cases, however, the indivi- duals exhibiting the luminosity had suffered from consumption, or some other wasting disease, and were near the close of their lives at the time; so that it is probable that a decomposition of the tissues was * See an account of several cases of the Evolution of Light in the Livin°- Human Subject, by Sir Henry Marsh, M.D., M.R.I.A., &c. ANIMAL ELECTRICITY.—ELECTRIC FISHES. 431 actually in progress, analogous to that which, when it occurs after death, imparts luminosity to the decaying body. One instance is recorded, in which a large cancerous sore of the breast emitted light enough to en- able the hands on a watch-dial to be distinctly seen Avhen it was held within a few inches of the ulcer; "here, too, decomposition Avas obviously going on, and the phosphorescent matter produced by it was exposed to the oxygenating action of the atmosphere. 771. Slight manifestations of free Electricity, or, in other words, dis- turbances of Electric equilibrium, are very frequent in living animals; and they are readily accounted for, when we bear in mind that nearly all chemical changes are attended with some alteration in the electric state of the bodies concerned; and when we consider the number and variety of such changes in the living animal body. When slight, how- ever, they can only be detected by refined means of observation ; and it is only when they are considerable, that they attract notice. The most remarkable examples of the evolution of free Electricity in Animals, are to be found in certain species of the class of Fishes; the best knoAvn of which are the Torpedo or Electric Ray, and the Gymnotus or Electric Eel. These possess organs, in which Electricity may be generated and accumulated in large quantities, and from which it may be discharged at will. The shock of a large and vigorous Gymnotus is sufficiently powerful to kill small animals, and to paralyse large ones, such as men and horses ; that of the Torpedo is less severe, but it is sufficient to be- numb the hand that touches it. 772. The electric organs of the Torpedo (which, from being found on European shores, has been the most studied) are of flattened shape, and occupy the front and sides of the body; forming two large masses, which extend backwards and outAvards from each side of the head. They are composed of tAvo layers of membrane separated by a consi- derable space ; and this space is divided by vertical partitions into hexagonal cells like those of a honeycomb, the ends of which are directed towards the two surfaces of the body. These cells, which are filled with a whitish soft pulp, somewhat resembling the substance of the brain, but containing more water, are again subdivided horizontally by membranous partitions; and all these partitions are profusely sup- plied with blood-vessels and nerves.—The electrical organs of the Gym- notus are essentially the same in structure ; but they differ in shape, in accordance with the conformation of the animal.—In these, and the other Electrical fishes, the electric organs are supplied Avith nerves of very great size, larger than any others in the same animals, and larger than any nerves in other animals of similar bulk. These nerves arise from a peculiar ganglionic enlargement of the Medulla Oblongata, termed the electric lobe, and seem chiefly analogous to the pneumogas- trics of other animals. 773. The following conditions appear to be essential to the manifes- tation of the Electric powers of these animals. Two parts of the body must be touched at the same time; and these two must be in different electrical states. The most energetic discharge is procured from the Torpedo, by touching its back and belly simultaneously; the electricity of the back being positive, and that of the belly negative. When tAvo 432 ELECTRIC FISHES.—ELECTRICITY OF MUSCLES. parts of the same surface, at an equal distance from the electric organ, are touched, no effect is produced, as they are equally charged with the same electricity ; but if one point be further from it than the other, a discharge occurs, the intensity of which is proportioned to the difference in the distance of the points from the electric organ. However much a Torpedo is irritated, no discharge can take place through a single point; but the fish makes an effort to bring the border of the other surface in contact with the offending body, through which a shock is then transmitted. This, indeed, is probably the usual way in which the discharge is effected.—The identity of animal with common Elec- tricity is proved, not merely by the similarity of the effects upon the feelings produced by the shock of both; but also by the fact that a spark may be obtained, and chemical decompositions effected, by the former, precisely as by the latter. 774. The power of the animal over the actions of its Electric organs, is dependent upon their connexion with the nervous centres. If all the nerve-trunks supplying the organ on one side be divided, the ani- mal's control over that organ will be destroyed ; but the power of the other may remain uninjured. If the nerves be partially divided on either or both sides, the power is retained by the portions of the organs which are still connected with the centres by the trunks that remain. Even slices of the organ, entirely separated from the body except by a nervous fibre, may exhibit electrical properties. Discharges may be produced, by irritating the part of the nervous centres from which the trunks proceed, so long as the latter are entire ; or by irritating the portions of the divided trunks which remain in connexion with the electric organs; or even by irritating portions of the electric organs themselves, when separated from the nervous centres.—In all these respects, there is a strong analogy between the action of the nerves on the Electric organs, and their action on the Muscles. The connexion of the organs specially appropriated to each of these actions with the Nervous system, the dependence of their functions upon the integrity of this connexion and upon the state of activity of the central organs, the influence of stimulation applied to the nervous centres or trunks, the results of ligature or section of the nerve, and the effects of poi- sonous agents, are all so remarkably analogous in the two cases, that it seems scarcely possible to doubt that the Nervous force is the agent Avhich is instrumental in producing both sets of phenomena. Still, how- ever, no proof whatever can be derived from this source, of the identity of nervous influence with any form of Electricity ; since all that can be legitimately inferred from it is, that Nerve-force acting through a parti- cular organic structure developes Electricity, in virtue of the correlation formerly explained (§ 396). 775. It is another interesting point of analogy between the action of Muscles, and that of the Electrical organs, that the former (as is now fully proved by the elaborate and exact researches of Matteucci) is attended with electric disturbance. In any fresh vigorous muscle, in a state of passive or tonic contraction, there is a continual electric current from the interior to the exterior, sufficient to excite the leg of a frog to energetic contraction, when its nerve is so applied to the MANIFESTATIONS OF ELECTRICITY. 433 muscle, as t.o receive the influence of this current. And a much more powerful current is produced, when the muscle is thrown, by a stimu- lus applied to its own nerve, into a state of energetic contraction. The explanation of the constant direction of the current, from the interior toAvards the exterior of the muscle, seems to be, that the changes con- nected with the nutrition and disintegration of the muscular tissue go on more energetically in its interior, than they do nearer its surface, where the proper muscular fibres are mingled with a large proportion of areolar and tendinous substance. 776. It was observed by Galvani that there exists in the Frog, during its whole life, a continual current of Electricity passing from its extremities towards its head ; and as no such current has been detect- ed in any other animal, it has been termed the courant propre, or pe- culiar current, of the Frog. It bears this curious analogy to the elec- tric discharges of Fishes,—that it is not manifested if the connexion be made between corresponding points of the opposite sides, but that it shows itself when the communication is made between points higher or lower in the body, whether on the same or on opposite sides.—There noAv seems reason to believe, however, from the observations of Mat- teucci, that this "proper current of the frog" is but a special case of the ordinary muscular current, depending upon the peculiar arrangement of the muscular and tendinous elements in this animal. Both currents are alike influenced by agents which affect the vitality of the muscle; and it is curious that poisoning with sulphuretted hydrogen should almost immediately put an end to each, although ordinary narcotic poisons have very little influence. 777. Manifestations of Electricity may be produced, in most animals having a soft fur, by rubbing the surface, especially in dry weather; this is a fact sufficiently well known in regard to the domestic Cat. Some individuals of the Human race exhibit spontaneous manifestations of electricity, which are occasionally of very remarkable power. There are persons, for instance, who scarcely ever pull off articles of dress that have been worn next their skin, without sparks and a crackling noise being produced, especially in dry weather. This is partly due, however, to the friction of these materials with the surface, and with each other. But the case of a lady has been recently put on record, who was for many months in an electric state so different from that of surrounding bodies, that, whenever she was but slightly insulated by a carpet or other feebly-conducting medium, sparks passed between her person and any object which she approached. When she was most favorably circumstanced, four sparks per minute would pass between her finger and the brass ball of a stove, at a distance of 1J inch. Va- rious experiments were tried, with the vieAv of ascertaining if the Elec- tricity Avas produced by the friction of articles of dress ; but no change in these seemed to modify its intensity. From the pain which accom- panied the passage of the sparks, this condition was a source of much discomfort to the subject of it. 28 434 OF GENERATION AND DEVELOPMENT. CHAPTER XI. .OF GENERATION AND DEVELOPMENT. 1. General View of the Nature of the Process. 778. There is no one of the functions of living beings, that distin- guishes them in a more striking and evident manner from the inert bodies which surround them, than the process of Generation. By this function, each race of Plants and Animals is perpetuated; whilst the individuals composing it successively disappear from the surface of the earth, by that death and decay which are the common lot of all. There are certain tribes, in which the death of the parent is necessary for the liberation of the germs from which a new race is to spring up. This is the case, for example, in some of the simplest Cellular Plants; in which every cell lives for itself alone, and performs its whole series of vital operations independently of the rest. But as, in more complex organisms, we find certain cells set apart for Absorption, others for Secretion, &c, so do we find a particular group of cells set apart for Reproduction; and these go through a series of changes peculiar to themselves, without interfering with the general life of the structure.— It is in the Vegetable kingdom, that the essential character of the Ge- nerative process can be best studied ; and we shall, in the first instance, therefore, inquire into the nature and import of the principal pheno- mena which it presents. 779. If we take as our starting-point the simple cell in which the in- dividuality of the lowest Algae seems to reside, Ave find that this cell, Fig. 132. Various stages of development of Bcematococcus binalis:—a, a, simple rounded cells; b, elongated cells, the endochrome preparing to divide; c, c, cells in which the division has taken place; d, cluster of four cells formed by a repetition of the same process. under the influence of light and warmth, and supplied with aliment, multiplies itself to an extent that almost seems unlimited; and this by a process of duplication exactly analogous to that which has been al- ready described (§ 212) as taking place in Cartilage;—the chief diffe- SIMPLEST FORMS OF GENERATIVE PROCESS. 435 rencc being, that in the latter the fission commences in the nucleus, each half of which seems to draw around it a portion of the contents of the cell, whilst in the former the fission shows itself at once in the entire endochrome, there being no distinct nucleus (Fig. 132). Now although the effect of this operation is to produce a great number of new cells, yet it cannot be truly considered as an act of Generation; for it is ob- viously analogous to that multiplication of the component cells, which takes place as a part of every process of development in the most com- plex organisms; the only difference being, that the new cells are here in great^ degree independent one of another, so as to be able to maintain their existence when isolated ; whilst among the higher tribes, there is so close a relation of mutual dependence between the component cells, that they cannot continue to live if separated from one another. And we shall hereafter see, that the early development of the embryonic Fig. 133. Successive stages of development of simpler Algse:—a, individual cells of Protococcus viridis; b, c, clusters formed by their multiplication; r, filament of Schizognium murale ; e, a similar filament, subdividing late- rally, which constitutes the early form of the Utvacece ; p, o, portions of the expanded thallus of Viva furfu- racea, formed by the continuance of the same process of transverse subdivision. mass, even in the highest Animals presents phenomena in all respects comparable to this multiplication of the simplest of Cellular Plants by successional subdivision (§ 805); all the descendants of the original cell, however, here remaining in mutual apposition, and concurring to make up what is commonly designated as a single individual, whilst in the 436 OF GENERATION AND DEVELOPMENT. simplest Plants, these cells, as their number is successively augmented by fission, arejnore and more widely separated from each other, and may disperse themselves over an extensive surface.—If, however, they should remain in connexion with each other, they may form clusters, or fronds (expanded leafy surfaces), according to the direction in which the subdivision takes place (Fig. 133); and this without the slightest departure from the original cellular type which is preserved throughout the structure, every part being exactly similar to every other. In these composite organisms, we usually find a provision, not merely for the extension of the original structure, but for the multiplication of indivi- duals, Avhich being still referable to the general type of cell-subdivision must be considered as a process of development, rather than of genera- tion. This consists in the emission, from the interior of certain of the cells, of broods of young cells formed in their interior; and these, in the loAver aquatic plants, are very commonly furnished with cilia, by the agency of which they are dispersed through the water, beginning to develope themselves into the likeness of the organisms from which they sprang, as soon as their movement has ceased. These " zoo- spores," as they are termed, must be regarded as the representatives of the gemmar or buds of higher Plants. The latter are usually developed in continuity with the stock from which they originate; but there are many instances in which they are spontaneously detached; and there are few cases in which they will not continue their existence under favorable circumstances, when artificially separated from it, as is practised in the operations of grafting, budding, &c. 780. The true Generative process, on the other hand, seems to con- sist, throughout the Vegetable kingdom, in the reunion of the contents of tAvo cells which have been separated in the process of development and multiplication, and in the production of a germ as the result of this reunion, which is usually very different in its characters and properties from either of the cells whose contents have contributed to form it. This process has been observed to take place in the Vegetable kingdom, under three principal forms, which seem to be characteristic of the lowest Cryptogamia, of the higher Cryptogamia, and of the Phanero- gamia, respectively.—The first of these presents itself in those simple Cellular Plants, in which, whether the cells remain in connexion or not, their endowments are all of the same nature. At a certain time of the year (it would seem) in each species, the cells approach one another in pairs, and their endochromes (or colored contents) are intermingled (Fig. 134, a), either by the rupture of both cells (1), or by the formation of a direct communication from the interior of one to that of the other, in Avhich case the union of the two endochromes may take place either in the connecting channel (2), or in one of the pairs of cells (3). Of this process, Avhich is known as Conjugation, the result is the formation of a body known as a Sporangium, which may be considered as the first product of the true generative process; and from this sporangium (which is a single cell, or a pair or cluster of cells) a " neAV generation" is de- veloped by the subsequent process of fission and multiplication, There is here no definite distinction of the sexes, the conjugating cells being apparently alike in their endowments; such a distinction is shadowed SIMPLEST FORMS OF GENERATIVE PROCESS. 437 forth, however, where the sporangium is developed within one of them. —The second form of the true generative process is seen even in the higher Algae; and, although the extent of its prevalence has not yet been clearly determined, it is probably common to the Liverworts, Mosses, and Ferns, it being in the last of these groups that it has been most satisfactorily made out. In conformity with the separation or specialization of organs which is characteristic of these Plants, we find that the Generative power is now limited to certain small parts of them, and that these produce two orders of cells, very distinct in their endow- ments, which may be called respectively "sperm-cells," and "germ- cells." It is from the latter that the new plant originates ; but this it can only do, when the fertilizing influence of the former has been con- veyed to it; and the provision for this purpose is very remarkable. The sperm-cells, developed within bodies termed antheridia, form in their interior, as their characteristic products, minute spirally-coiled filaments, usually furnished with cilia at one extremity, and bearing a very close resemblance to the spermatozoa of animals (§ 785). These, when liberated from the cells within which they were formed, possess a very active power of movement, in virtue of which they make their way to the germ-cells; and when they have impinged against these, there is reason to believe that they dissolve away, and that the product of their diffluence is absorbed into the germ-cells and mingles with the contents of the latter, the formation of a germ being the result of this intermix- ture (Fig. 134, b). Here, then, we have the distinction of sexes well Fig. 134. Diagram representing the three principal forms of the Generative process in Plants:—A, conjugation of inferior Cryptogamia; formation of the sporangium, 6, by admixture of the discharged endochromes of the parent-cells, a, a ; 2, production of the sporangium, 6, within a dilatation formed by the union of the two parent-cells; 3, production of the sporangium, b, by the passage of the endochrome of cell a, into that of cell a*, marking out a sexual difference.—b, fertilization of germ in higher Cryptogamia; a, sperm-cell discharg- ing its spiral filament, a*, germ-cell, against which one of these filaments is impinging, b, germ produced by their contact.—c, fertilization of germ in Phanerogamia; a, germ-cell, or pollen-grain, sending its pro- longed tube dawn the style, until it reaches a*, the germ-cell, enclosed in the ovule, the section of whose coats is shown at c; from the contact of the two, is produced the germ b. marked ; but both sperm-cells and germ-cells are usually developed in the same organism, and are alike the product of a single original germ. Throughout the Cryptogamic series, the fertilized germ appears to be 438 OF GENERATION AND DEVELOPMENT. thrown at once upon the world, and is dependent for its supply of food upon its own absorbing and assimilating powers; these enable it to mul- tiply itself by fission, sometimes to a vast extent; and thus an elaborate and complex organism (such as a Tree-fern) may be produced.—In the third form of the generative process, which is peculiar to Phanerogamia (or Flowering Plants), there is the same distinction between " sperm- cells" and " germ-cells;" but the mode in which the action of the former upon the latter is brought about, is very different. The " sperm-cell," which is known as the pollen-grain, and is developed in the anthers of the flower, does not here evolve self-moAring filaments, but, when it falls upon the apex of the style, puts forth long tubes, which insinuate them- selves down between its loosely-connected tissue, until they reach the ovary at its base. Here they meet with the ovules, which are in reality " germ-cells" imbedded in a mass of nutriment stored up by the parent; and the pollen-tube, entering the micropyle or foramen of the ovule, penetrates into such close approximation to the germ-cell contained within it, that its contents find a ready passage by endosmose into the latter (Fig. 134, c). Here, again, therefore, we have the same essential phenomenon,—the intermixture of the contents of the sperm-cell and of the germ-cell, as the condition for the development of the true germ. But this germ, still making its first appearance as a single cell within the ovule, is supplied with nutriment by its parent; and this not merely whilst the ovule remains in connexion with the organism which evolved it, but for some time subsequently, the store laid up around it in the seed being the material at the expense of which its early deve- lopment takes place. It is not, in fact, until its true leaves have been evolved and its root-fibres have penetrated the soil, which takes place in the act of germination, that it becomes capable of absorbing and assimilating nutriment for itself. As soon, however, as this takes place, the young plant becomes independent of further assistance; and all its subsequent growth is provided for by its own powers. In pro- cess of time, its generative apparatus is evolved; and here, too, Ave find, that the two sets of sexual organs are usually developed in the same organism, it being only a small proportion of Phanerogamia that are dioecious, i. e., that have the male or staminiferous flowers, and the female or pistilline, restricted to different individuals.* 781. The history of embryonic Development in Flowering Plants, presents some interesting points of correspondence with that of the higher Animals.—The germ that is developed within the germ-cell (here designated the "embryonic vesicle" of the ovule) as the product of the admixture of its contents with those of the sperm-cell (or pollen- grain), is itself a single cell; and the early history of its development closely resembles that which may be observed in all the inferior Plants. In the first place it subdivides into tAvo, each of these into two others, and so on ; its first nisus or tendency being to the production, not of the parts which are to be evolved into the stem, roots, leaves,"&c, of the perfect plant, but of a leaf-like expansion, which may be likened to the * For a more particular account of the recent discoveries, on which the above account of the Generation of Plants is based, see the Author's " Principles of General and Com- parative Physiology," chap, xviii., sect. 2. SIMPLEST FORMS OF GENERATIVE PROCESS. 439 frond of the Cryptogamia, and of which the function is only temporary. It is by this organ, the single or double cotyledon, that the nourishment provided in the ovule is absorbed and prepared for the development of the young Plant; the permanent fabric of which, even at the time of the maturity of the seed, forms but a small proportion of the entire embryonic structure. In the act of germination, however, the perma- nent portions are developed at the expense of the temporary, the plu- mula and radicle absorbing the nourishment which has been elaborated by the cotyledons ; and having fulfilled its transient purpose, and com- pleted its term of life, the first leaf-like expansion withers and dies. The tissues of the young Plant are at first of the simplest possible character; but as the organs characteristic of its adult condition are one after another put forth (always originating in peculiar groups of cells), so do Ave find that the spiral vessels, woody fibre, &c, charac- teristic of the higher organisms, gradually make their appearance.—Thus we see that even the highest Plants have to pass through conditions closely conformable to those which are permanently shown in the lower; and that the parts which are first formed are destined for only a tem- porary purpose, that of preparing nourishment for the evolution of more permanent structures. We shall find, in tracing the history of the de- velopment of the higher Animals, that exactly the same general fact may be observed, in even a more striking manner; the number of diffe- rent stages being greater, and a yet larger proportion of the parts first formed having a merely temporary purpose, and being destined to an early decay, as soon as the more permanent parts of the fabric shall have been evolved. 782. Among many of the lower Animals, a multiplication of indivi- duals takes place by a process that closely resembles the budding of Plants; this also must be regarded, not as a proper act of Generation, but as a modification of the ordinary Nutritive process. The same may be said of the powers of reparation, which every animal body possesses in a greater or less degree, but which are by far the most remarkable among the lower tribes ; for when an entire member is renewed (as in the Star-fish), or even the whole body is regenerated from a small frag- ment (which is the case in many Polypes), it is by a process exactly analogous to that Avhich is concerned in the reparation of the simplest wound in our own bodies, and which, as already explained (§ 636), is but a modification of the process that is constantly renewing, more or less rapidly, every portion of their fabric. Although the buds thus produced and separated are usually developed into the likeness of the parent stock, yet this is sometimes not the case, the stock possessing one form, and the bud another, which may be quite different; as when certain fixed composite Zoophytes bud off free-moving solitary Medusae, these last depositing ova, from which the Zoophytic type is regenerated. When, however, this phenomena, to which the name of " alterations of generations" has been given (erroneously, in the Author's opinion),* is carefully examined, it is found that the bud thus detached is really *For a discussion of this subject, see the Author's " Principles of General and Com- parative Physiology," chap, xviii., sect. 1. 440 OF GENERATION AND DEVELOPMENT. the generative apparatus of the parent stock,, furnished (it may be) with nutrient and locomotive organs of its OAvn; and that neither can be re- garded as a complete organism without the other. Thus, the Medusa contains the proper generative apparatus of the Zoophyte, which developes no other; and the " aggregate" Salpae that are budded- forth from a kind of stalk in the interior of the "solitary" form, must be regarded as altogether constituting its true generative apparatus, since it never produces any other. In all instances it will be found, that whatever may be the variations which present themselves in the entire history of any species, the immediate product of the true Gene- rative act is always the same. 783. This act, in Animals as in Plants, requires the concurrent action of two sets of organs, evolving "sperm-cells" and "germ-cells" respectively; and it is curious that these should present the closest approach to those of the higher Cryptogamia, rather than to those of Plants above or below them in the scale. The two sets of organs may be united in the same individual, as they are in most Plants ; and the ova may be fertilized from the seminal cells of the same being;—as happens in many Zoophytes and in some of the lowest tribes of Mol- lusca. Or, the two sets of organs being present in each individual, it may not be capable of self-impregnation; but, in the congress of two individuals, each impregnates, and is impregnated by, the other;—as may be observed in the Snail, and many of the higher Molluscs. Or the sexes may be altogether distinct; one individual possessing only the male or spermatic organs; and the other the female, or germ-nourishing apparatus ;—this is observed in the higher classes of the Radiated, Mollus- cous, and articulated sub-kingdoms and it is the case in all Vertebrata. 784. The earliest part of the history of Embryonic Development is nearly the same in all Animals ; for it consists in the multiplication of the single cell of Avhich the original germ is composed, until a cluster is formed, all the cells of which appear to be in all respects similar to one another. Each of these cells either takes into itself, or draAvs around it, a portion of the vitellus or yolk, which is the nutrient substance of the ovum; and thus either the whole of this vitellus, or a portion of it, is subdivided into a number of minute spherules, altogether constituting what is known as the "mulberry mass" (Fig. 135). The former seems to be the case, when the grade of development of the organism which is to be formed at the expense of the yolk is very low; whilst the latter plan is followed, when the yolk is destined to afford a prolonged suste- nance to the embryo, which attains a high degree of development Avhilst supported upon it alone. Thus among the Invertebrata generally, we find that the embryo comes forth from the egg in a very simple condi- tion, a large part of its structure having undergone but little change from the state of the "mulberry mass;" and in these, the Avhole yolk undergoes subdivision. The same is the case, too, in the Batrachian Reptiles, which issue from the egg in a form very different from that into which they are to be subsequently developed; and it is the case even with Mammalia, but for a very different reason, their embryonic structure, formed at the expense of the yolk, being destined to acquire additional material for its full development from a source altogether SIMPLEST FORMS OF GENERATIVE PROCESS. 441 different. In the highest Mollusca, however, as also in Fishes, Reptiles, and Birds, the portion of the yolk which undergoes subdivision is com- paratively small; and the great mass of the vitellus is destined to be Fig. 135. ABC D 09 E F G H Successive stages of segmentation in the vitellus of the Ovum of Ascaris acuminata :—a, ovum recently impregnated, the yolk-bag slightly separated from the enveloping membrane ; b, first fission into two halves; c, second fission, forming four segments; d, yolk, now divided into numerous segments; e, formation of "mulberry mass" by further segmentation; r, the mass of cells now beginning to show the form of the future worm; o, further progress of its evolution: h, the worm, formed by the conversion of the yolk-cells now nearly mature. subsequently absorbed into the substance of the germ, by a process analogous to that by Avhich the food of the adult is imbibed. Hence the portion of their yolk which undergoes subdivision, and helps to con- stitute the "mulberry mass," may be termed the "germ-yolk," whilst the remainder may be designated as the "food-yolk." 785. When the whole of the yolk is taken into the mulberry mass, the formation of the embryo is usually the result of the progressive metamorphosis of its parts; the cells of the surface being converted into the integument, and those of the inner part into the internal organs. This is the case, for example, in the Intestinal Worm, some of the stages in whose development are shown in Fig. 135. The embryonic con- dition of many of the organs is frequently retained, at the time when the young animal comes forth from the egg ; those parts only being completed which are necessary to enable it to obtain its nutriment.— Other organs are subsequently evolved, at the expense of the food thus introduced; and thus a complete change or metamorphosis may take place, in regard alike to external form and to internal structure, betAveen the larval and adult states. Of this phenomenon Ave have charac- teristic examples in the groups of Insects and Batrachia ; and although it was formerly considered exceptional, it is now knoAvn to be the or- dinary occurrence among the lower tribes of animals ; it being com- paratively rare for any of them to come forth from the egg under their adult forms. This change is sometimes obviously gradual, as in the progressive advance of the Tadpole into the condition of the Frog ; but it is sometimes apparently sudden, as when the Chrysalis skin is thrown off, and the perfect Insect comes forth. In the latter case, however, the change is really just as gradual as in the former; since the develop- ment of the organs characteristic of the perfect Insect is taking place during the Avhole of the Chrysalis period, to be displayed and brought 442 OF GENERATION AND DEVELOPMENT. into use at its termination. Thus the whole life of the Insect, up to its last change, may be regarded as one of prolonged embryonic develop- ment ; and the same may be said of that of the Frog, up to the time when its permanent organs are fully evolved.—No such ostensible me- tamorphosis takes place, however, in any of the animals which are pro- vided with a "food-yolk;" for this supplies that material for the con- tinued development of the embryo within the egg, which is elsewhere to be obtained out of it; and thus the embryo is supported, until it has nearly attained its adult condition, although far from having acquired its adult size. Noav in all these cases, it is very interesting to remark that the first nisus is towards an extension of the embryonic mass as a membranous expansion (evidently analogous to the cotyledon of the Flowering Plants, § 781) over the "food-yolk;" in this "germinal mem- brane," which forms a sort of temporary stomach, blood-vessels are de- veloped, which absorb the prepared nutriment and convey it to the per- manent portion of the embryonic structure; and when its function is completed, the store of aliment being exhausted, and the proper nutrient apparatus of the embryo being ready for action, we lose sight of it al- together. We shall find that a similar germinal membrane is formed in the Human ovum, although there is no " food-yolk ;" its formation being apparently requisite for ulterior purposes, and the portion of the mul- berry mass which gives origin to the permanent part of the embryonic structure being comparatively small. 2. Action of the Male. 786. The share in the Reproductive Function, which belongs to the Male Sex, essentially consists in the formation and liberation of the fertilizing bodies termed Spermatozoa. These are prepared within peculiar cells, as already described (§ 241) ; and the " sperm cells" are either scattered through the soft parenchyma of the body, as happens among some of the lowest animals ; or they are confined to certain parts of it, as in those a little more elevated in the scale; or they are formed within follicles or tubes, clustered together into ian organ of a glandular character, known as the Testis. Such an organ is found in all Insects and Mollusca ; as well as in Vertebrated Animals. In the first of these classes, it is formed on the general plan of their proper glands (§ 720); being usually composed of tubes, more or less elongated, and some- times terminating in enlarged follicles. In the Molluscs, on the other hand, it is almost invariably composed of clusters of follicles. In either case, the seminal cells are developed within the tubes or follicles, a9 are the ordinary secreting cells of the Liver or Kidney within the tubes or follicles of those glands; and their contents are discharged by an excretory duct, which terminates in an organ that conveys them out of the body, either emitting them into the surrounding Avater (as hap- pens with many Mollusca), or depositing them within the body of the female. It is curious that, in some of the loAvest Fishes, we should return to one of the simplest conditions of this organ,—a mass of vesicles, without any excretory duct. In these cases the secretion formed within the vesicles escapes, by their rupture, into the abdominal ACTION OF THE MALE. 443 Fig. 136. Anatomy of the Testis :— cavity; whence it passes out by openings that lead directly to the ex- terior.—The Testis in Man is formed, in every essential particular, upon the plan of the ordinary Glands. It consists of several distinct lobules, separated by processes of the fibrous envelope, or tunica albuginea, which pass down between them; and each lobule consists of a mass of convoluted tubuli seminiferi, through which blood-vessels are minutely distributed. The diameter of these tubuli is tolerably uniform; being, when they are not over-distended, from l-195th to l-170th of an inch. They form frequent anastomoses with each other; and on this account it is difficult to trace out their free or caecal extremities. The tubuli of each testis discharge their contents into an efferent duct, the Vas deferens; and by this the product is conveyed into the Vesiculi seminalis on each side, which, like the gall-bladder and urinary bladder, serves to store up the secretion until the proper time arrives for discharging it. The product of the action of the Testis consists of a fluid, through which the Sper- matozoa are diffused ; these last bodies being usually set free by the rupture of the seminal cells, before they leave the tubuli of the testis. It is difficult to determine the precise characters of the fluid portion \ \ ^l-^^t\^iZt\i of the secretion; as this is mingled with other se- s! 3. The lobuii testis. 4, 4. , , ', r ,1 t\ . . i i i/> Tne vasa recta. 5. The rete cretions (such as that or the Prostate gland, and ot testis. 6.The vasaefferentia, the mucous lining of the Vesiculae seminales and gInTehdlc^nS1uiLDldiaa»7amepr7' spermatic ducts), before it is emitted. And an ^rh^Tobtmfcftt exact analysis is not of much consequence; since epididymus. s. The body of ,, r i i , .i , ,i ■!• „ the epididymus. 9. The globus there can be no doubt that the peculiar powers of minor of the epididymus. 10. the fluid depend upon the Spermatozoa. It may TasecuTSmdaberrrnans. n" The be stated, however, that the Spermatic fluid has an alkaline reaction, and that it contains albumen, together with a peculiar animal principle termed Spermatine; and that it also includes saline matter, consisting chiefly of the muriates and phosphates, espe- cially the latter, which form crystals when the fluid has stood for some little time. 787. The Spermatozoa, or minute filamentous bodies set free by the rupture of the spermatic cells, are distinguished by their power of spontaneous movement, which occasioned them to be long regarded as proper Animalcules. It is now clear, hoAvever, from the history of their development, as Avell as from other considerations, that they cannot be justly regarded in this light; and that they are analogous to the repro- ductive particles of Plants, which, in many cases, exhibit a spontaneous motion of extraordinary activity, after they have been set free from the parent structure. The Human Spermatozoa consists of a little oval flattened "body," from the l-600th to the l-800th of a line in length; from Avhich proceeds a filiform "tail," gradually tapering to a very fine point, of l-50th or at most l-40th of a line in length. The whole is perfectly transparent; and nothing that can be called structure can be 444 OF GENERATION AND DEVELOPMENT. satisfactorily distinguished within it. The movements are principally excited by the undulations of the tail, which give a propulsive action to the body. They may continue for many hours after the emission of the fluid; and they are not checked by its admixture with other secre- tions, such as the urine, and the prostatic fluid. When the seminal fluid remains in contact with a living surface (as when deposited in the generative organs of the female), the Spermatozoa may retain their vitality for some days; and an instance has already been referred to (§ 240), in Avhich the later stages of the development of the Sperma- tozoa actually take place in this situation,—the seminal fluid emitted by the male (among many Crustacea) not containing any Spermatozoa completely formed, but numerous spermatic cells, which undergo the remainder of their development, and then rupture and set free their contents, within the oviducts of the female. 788. The power of procreation does not exist in the Human Male (except in rare cases) until the age of from 14 to 16 years; at which epoch, the sexual organs undergo a much-increased development; and the instinctive desire, which leads to the use of them, is awakened in the mind. From that time to an advanced age, the procreative power remains, in the healthy state of the system; unless it be exhausted by excessive use of it, or by too energetic a direction of the mental or corporeal powers to some other object. The formation of Seminal fluid being, like the proper acts of Secretion, very much influenced by con- ditions of the Nervous System, is increased by the continual direction of the mind toAvards objects which arouse the sexual propensity; and thus, if sexual intercourse be very frequent, a much larger quantity of the fluid will be produced than if it is more rarely emitted, although the amount discharged on each occasion will be less. The formation of this product is evidently a great tax upon the corporeal powers; and it is a well-knoAvn fact, that the highest degree of bodily and mental vigor is inconsistent with more than a very moderate indulgence in sexual intercourse; whilst nothing is more certain to reduce the powers, both of body and mind, than excess in this respect. 789. It may be stated as a general laAv, prevailing equally in the Vegetable and Animal kingdoms,—that the development of the indi- vidual, and the reproduction of the species, stand in an inverse ratio to each other. We have seen that, in many organized beings, the death of the parent is necessary to the production of a new generation; and even in numerous species of Insects it folloAvs very speedily upon the sexual intercourse. It is a curious fact, that Insects which usually die, the male almost immediately after the act of copulation, and the female very soon after the deposition of the eggs, may be kept alive for many Aveeks or even months, by simply preventing the copulation. And there can be no doubt that, in the Human race, early death is by no means an unfrequent result of the excessive or premature employ- ment of the genital organs; and Avhere this does not produce an immediately-fatal result, it lays the foundation of future debility, that contributes to produce any forms of disease to which there may be a constitutional predisposition, especially those of a Scrofulous nature. 790. The emission of the Spermatic fluid is an, act of a purely reflex ACTION OF THE FEMALE. 445 nature; the Will having no poAver either to effect or to restrain it. The stimulus is given by the friction of the surface of the Glans Penis against the rugous walls of the Vagina; the sensibility of the organ being at the same time much increased, by the determination of blood to it. The impression is at last sufficiently strong to produce, through the medium of the lower part of the Spinal cord (which is the gangli- onic centre of the circle of afferent and efferent nerves connected with this organ), a reflex contraction of the muscles surrounding the Vesi- culae seminales. These receptacles discharge their contents (which consist partly of the Spermatic fluid, and partly of a secretion of their OAvn) into the Urethra; and from this they are expelled, Avith some degree of force, and with a kind of spasmodic action, by its own Com- pressor muscles. Although the sensations concerned in this act are ordinarily most acutely pleasurable, yet there appears to be sufficient evidence that they are by no means essential to its performance; and that the impression conveyed to the Spinal cord may excite the con- traction of the Ejaculator muscles, like other reflex operations, without producing sensation (§ 394). 3. Action of the Female. 791. The share of the Female in the Generative act is greater than that of the Male; for she not only furnishes, in the "germ-cell," a product which is as essential as that supplied by the "sperm-cell" for the first formation of the germ; but she also supplies it with the mate- rials Avhich it requires for its development, up to the condition in which it can support its own life. The mode in which this is accomplished, is essentially the same Avith that in which the process is effected in Plants. In certain parts of the female structure are developed pecu- liar bodies termed ova ; Avhich contain, not merely the germ-cells, but in addition a store of nutriment adapted to supply the wants of the germ. The fertilizing influence finds its way into these; and the germs thus produced begin to grow at the expense of the material with which they are surrounded. This, as already pointed out, may enable the embryo to develope itself, without any further assistance (save a warm tempe- rature) into the form it is permanently to assume ; as in the case of Birds and Reptiles, Avhich do not come forth from the investments of the egg, until they have attained the form characteristic of the group to Avhich they belong. Or it may only serve for the early part of the process ; and one of two methods may then be employed to complete it;—either a new connexion is formed between the parent and the embryo, by which the former continues to supply the latter with nutri- ment, more directly from its blood,—as is the case with Mammalia,-^or the embryo issues from the egg, in a condition very unlike that which it is permanently to attain, but in a form Avhich enables it to acquire its own nourishment, and to pass through the latter stages of its evolution quite independently of any assistance from its parent: this is the case with a large proportion of the Invertebrata. 792. The Ova, like the seminal cells, are scattered through the soft parenchyma of the body, in animals of the lowest class; but they are 446 OF GENERATION AND DEVELOPMENT. more commonly developed in certain distinct portions of the fabric ; being sometimes formed in the midst of solid masses of areolar or cel- lular texture; whilst in other instances they are developed, like the spermatic cells, in the interior of tubes and vesicles resembling those of glands, and furnished with an excretory duct. The latter condition obtains in the greater proportion of the higher Invertebrated animals, and in some Fishes; but in the Vertebrated classes we return to the type which characterizes the egg-producing organs in many Zoophytes, —namely, the development of the egg in the midst of a mass of solid parenchyma, from which it gradually makes it way, to escape into the visceral cavity. The Ovarium of the Mammal, Bird, or Reptile, as well as that of most Fishes, differs entirely, therefore, from that of the In- vertebrata ; for the latter have all the essential characters of true glands; whilst the former are nothing else than masses of parenchyma, copiously supplied with blood-vessels, and having dispersed through their substance certain peculiar cells, termed ovisacs, within which the ova are developed. In order that the latter may be set free, not only must the ovisac itself burst (like parent-cells in general), but the pecu- liar tissue and the envelopes of the ovarium must likewise give way. When the ova thus escape into the abdominal cavity, they may lie there for some time, at last to be discharged through simple openings in its walls, as happens in those Fishes which have this form of ovarium : or they may be at once received into the trumpet-shaped expansion of tubes, that shall convey them to these orifices. These tubes are termed oviducts, in common with the excretory ducts of the glandular ovaria of Invertebrated animals; for their function is the same,—that of con- veying the ova to the outlet by which they are extruded from the body. Fig. 137. The Uterus with its appendages viewed on their anterior aspect. 1. The body of the uterus. 2. Its fundus. 3. Its cervix. 4. The os uteri. 5. The vagina; the number is placed on the posterior raphe or columna, from which the transverse rugae are seen passing off at each side. 6, 6. The broad ligament of the uterus. 7. A convexity of the broad ligament formed by the ovary. 8, 8. The round ligaments of the uterus. 9, 9. The Fallopian tubes. 10,10. The fimbriated extremities of the Fallopian tubes ; on the left side the mouth of the tube is turned forwards in order to show its ostium abdominale. 11. The ovary. 12. The utero-ova- rian ligament. 13. The Fallopio-ovarian ligament, upon which some small fimbriae are continued for a short distance. 14. The peritoneum of the anterior surface of the uterus. This membrane is removed on the left Bide, but on the right is continuous with the anterior layer of the broad ligament. They are represented in Mammalia by the Fallopian tubes, which are true oviducts, although they terminate in the uterus instead of proceed- ing directly to the outlet. And it is by the fimbriated extremities of the Fallopian tubes (Fig. 137, 10, 10), which apply themselves closely STRUCTURE AND DEVELOPMENT OF THE OVUM. 447 to the surface of the ovaries at the time of the discharge of the ova, that these are received and conveyed to the uterus, instead of being allowed (as in some of the lower animals) to fall into the abdominal cavity. 793. There are many cases among the lower classes, in which the ovum is retained within the oviducts, so that the young comes into the world alive; and there are few in Avhich, during this delay, it receives a direct supply of additional nourishment from the fluids of its parent. It is in the Mammalia, however, that we find the most remarkable and complete provision for this purpose. Still, the lowest division of this group approximates closely, in the type of its generative apparatus to the Oviparous Vertebrata; for the oviducts of the Monotremata remain distinct from each other, and terminate separately, in the uro-genital canal, each of them having first undergone dilatation into a uterine cavity, so that these animals have two completely distinct uteri. In the Marsupialia, there is a closer approximation of the two lateral sets. of organs on the median line; for the oviducts converge toAvards one another, and meet on the median line, but without coalescing; so that these animals have a true "double uterus," opening by two orifices into the vaginal canal,—a condition which is sometimes met with as a mal- formation in the Human female. The vaginal canal, however, is also double; which is less frequently observed in the Human species. The two preceding orders constitute the sub-class of Lmplacental Mammals; the development of their ova within the uteri being cut short at a period anterior to the formation of the placenta (§ 818).—As we ascend through the series of Placental Mammals, we, find the lateral coales- cence of the uterine dilatations of the Fallopian tubes becoming more and more complete. It first shows itself in the vagina, which is every- where single, although a trace of separation into two lateral halves is seen in the Mare, Ass, Cow, Pig, and Sloth, in which animals it is tra- versed, in the virgin state, by a narrow, vertical partition. In many of the Rodentia, the uterus still remains completely divided into two lateral hahres, opening into the vagina by separate orifices; whilst in others, these coalesce at their lower portion, forming a rudiment of the true "body" of the uterus of the Human female. This part increases in the more elevated Herbivora and Carnivora, at the expense of the lateral ununited portions, Avhich are now termed the "cornua;" but even in the lower Quadrumana, the uterus is somewhat cleft at its summit, and the "angles," into which the oviducts enter, form a considerable part of the whole organ. As we ascend through the Quadrumanous series towards Man, we find the "body" of the uterus increasing, and the "angles" diminishing in proportion, until the original division is completely lost sight of, except in the slight dilatation of the cavity at the points at which the Fallopian tubes enter it. 794. Having thus briefly noticed the most important characters of the organs provided for the original production and for the subsequent reception of the ova, we have now to inquire into the history of their development.—The essential structure of the ovule, or unfertilized egg, appears to be the same in all animals. It consists externally of a membranous sac, so termed, from the nature of its contents, the vitelline 448 OF GENERATION AND DEVELOPMENT. membrane, or yolk-bag. The vitellus, or yolk, consists chiefly of albu- men and oil-globules; and floating in this fluid is seen a cell of peculiar aspect, termed the germinal vesicle, upon the walls of which is a very distinct nucleus, termed the germinal spot.—The layer of albumen sur- rounding the yolk, and termed the white of the Bird's egg, together with the membrane Avhich envelopes this and forms the basis of the shell, are not added until after the ovum has left the ovarium. They are not present in the eggs of many of the loAver Invertebrata; these con- sisting merely of the parts which are formed within the ovarium. 795. The structure of the ovule in Mammals differs in no essential particular from that just described; but the yolk is much less in amount than in the ovules of Invertebrated animals; since only the very earliest stages of the development of the embryo are to take place at its expense. The vitelline membrane is of peculiar thickness and transparency; and as, when the ovum is compressed under the micro- • scope, it is seen as a broad transparent belt, it is commonly known as the zona pellucida. We shall find that the ovule, after leaving the ovarium and receiving the fertilizing influence, becomes enclosed, whilst passing through the Fallopian tube, with a layer of albuminous matter, which represents the white of the Bird's egg; and with an additional fibrous envelope, which corresponds with the membrane enveloping the latter. This fibrous membrane, termed the Chorion, afterwards be- comes subservient, however, to various important changes ; by means of which the ovum is again brought into connexion with the parent, to derive from the blood of the latter the materials requisite for the con- tinued development of the embryo. These changes will be described hereafter (§§ 811, 818). 796. The Ovisac of Mammalia forms the inner layer of what is termed the Graafian follicle, after the name of its discoverer; and in- stead of closely enveloping the ovulum, as it does in oviparous animals, it contains, in addition to it, a quantity of granular matter, consisting of cells arranged in membranous layers, together with fluid. In the immature ovisac, these cells occupy nearly the whole space betAveen the ovisac and the ovum, but they gradually dissolve aAvay, especially on the side nearest the surface of the ovary; and at the same time an al- buminous fluid is effused from the deeper part of the ovisac, which pushes before it the residual layer of cells that immediately sur- rounds the ovum (forming the discus proligerus), and thus carries it against the opposite wall. The outer layer of the Graffian follicle is formed by a thickening and condensation of the surrounding paren- chyma of the ovarium ; and it is quite distinct from the ovisac Avhich it envelopes. It is extremely vascular, and is evidently destined to afford to the contained structures the materials for their development, Avhich they receive and appropriate by their own powers of absorption and assimilation. 797. The Mammalian Ovarium may be seen, even in the foetal ani- mal, to contain immature ova, enclosed within their ovisacs ; and the several parts of the former may be clearly distinguished in those which are in the more advanced stages of development. It appears that, during the period of childhood, there is a continual rupture of the ovi- MENSTRUAL DISCHARGE. 449 sacs (or parent-cells), and discharge of ova, at the surface of the ova- rium ; but these ova never attain so high a degree of development, as to render them fit for impregnation. Their evolution takes place more completely, as well as more rapidly, at the period of puberty, when there is a greatly-increased determination of blood to the genital organs, and a correspondingly-augmented energy in their nutritive opera- tions. . At this epoch, th_ parenchyma of the ovarium is crowded with ovisacs ; which are still so minute, that in the Ox, according to Dr. Barry's computation, a cubic inch would contain 200 millions of them. Some of those nearest the surface, however, are continually attaining increased development; and a rupture of some of the Graafian follicles, and a discharge of ova prepared for impregnation, from the exterior of the ovarium, thenceforth take place, with more or less tendency to periodicity, during the whole time that the female is in a state of apti- tude for procreation. 798. In the Human female, the period of Puberty usually occurs be- tween the 13th and 16th years. The differences in the time of its advent partly depend upon individual constitution, and partly upon various ex- ternal circumstances, such as temperature, habits of life, &c. As a general rule, habitual exposure to a warm atmosphere, an inert life, sensual indulgence, and circumstances that excite the sexual feelings, favor the approach of Puberty; whilst a cold climate and hardy life retard it. The appearance of the Catamenial discharge usually takes place Avhilst the evolution of the genital organs is in progress ; and it is a decided indication, when it occurs, that the aptitude for procreation has been attained. It is not unfrequently delayed much longer, however; and its absence is by no means to be regarded as a proof of inability to con- ceive. The Catamenial fluid, as it proceeds from the lining membrane of the Uterus, seems to be nothing else than Blood; but in its passage through the vagina, this is deprived of its coagulating power by admix- ture Avith the vaginal mucus. The appearance of clots in the discharge may usually be regarded as an indication than an excess of blood is escaping from the uterine surface. In some cases of difficult Menstrua- tion, Avhich seem to depend upon a state of low inflammation in the Uterus, the fibrine has such a tendency to become organized, as to form shreds, or layers of false membrane, which sometimes plug up the os uteri. The healthy Menstrual secretion is remarkable for its very acid character.—It has been recently maintained that this periodical dis- charge of blood from the lining membrane of the uterus is dependent upon the ovarian oestrum ; but there seems adequate reason for the belief, that, although the two phenomena are usually consentaneous, yet that they are essentially independent; since, each occasionally recurs without being accompanied by the other. The catamenial dis- charge usually makes its appearance pretty regularly (save during preg- nancy and lactation) at intervals of 28 days ; but there are many females in Avhom its recurrence takes place with no less regularity at shorter or at longer intervals. The duration of the flow, too, is subject to great variations; for in some individuals it does not last above a day or two, whilst in others it continues a week or more. 799. This flux of blood from the lining membrane of the Uterus is 29 450 OF GENERATION AND DEVELOPMENT. not confined to the Human female, as was formerly supposed; but occurs in some of the lower Mammalia in the state of heat, or periodi- cal aptitude for procreation, at Avhich time the ovarium contains ova ready for impregnation. The chief peculiarity attending its appearance in the Human female, is its regular monthly return. In the natural condition of many of the lower Mammalia, as in Oviparous animals, the period of heat recurs at some one time of the year,—usually in the spring; or, in the smaller and more prolific species, from two to six times. And in those which have undergone a change by domesti- cation, the recurrence is usually irregular, depending upon various circumstances of regimen, temperature, &c. The general analogy between the Menstruation of the Human female and the Heat of the lower Mammalia,—consisting in the peculiar aptitude for impregnation which then exists in consequence of the maturation of ova in the ovarium, —cannot now be questioned; but it appears that in the Human female ova may be matured and impregnated at any part of the period which elapses between the occurrences of the Catamenial discharge; though it is certain that the aptitude for conception is much greater, during the few days which precede and follow the menstrual period, than at any intervening time. The duration of the period of aptitude for pro- creation, which is marked by the continued appearance of the Cata- menia, is more limited in Women than in Men; usually terminating at about the 45th year. It is sometimes prolonged, however, for ten or even fifteen years longer; but cases are rare, in which women above 50 years of age haAre borne children. There is usually no menstrual flow during pregnancy and lactation; in fact the cessation of the Cata- menia is usually one of the first signs indicating that conception has taken place. It is by no means uncommon, however, for them to appear once or twice subsequently to Conception; and their appearance during Lactation, especially if it be much prolonged, is still more fre- quent ; hence it might be inferred, that the continuance of Lactation may not prevent a fresh conception,—which is found to be true in practice. 800. We shall now take a brief survey of the changes which occur in the Ovulum, when it is being prepared for fecundation ; and of the principal features of its subsequent development.—Up to the period when the Ovule is nearly brought to maturity, it remains suspended in the centre of the cavity of the Ovisac; but it then begins to move towards that side of the Graafian follicle, which is nearest the surface of the ovarium. An important change is at the same time occurring in the Graafian follicle itself; for whilst the part with which the ovule comes in contact gradually thins away, the outer or vascular layer of the remainder, especially on that side most deeply imbedded in the ovary, becomes much increased in thickness; and a great increase takes place at that part, in the cellular layer that lines the ovisac, which presents a reddish glutinous aspect. This subsequently under- goes a still greater augmentation, and becomes more fleshy; projecting like a mass of granulations from the interior of the ovisac, and receiving blood-vessels which pass into it from the vascular membrane that surrounds it. At the same time, the wall of the Graafian follicle FERTILIZATION OF THE OVUM. 451 is thrown into wrinkles, which are directed towards the interior, so as to occasion the contraction of the cavity; and thus it comes to be entirely filled with the new growth, the centre of which is marked by a sort of stelliform cicatrix. This substance speedily becomes of a paler hue than at first, and is known from its color as the corpus luteum.—The escape of the ovule from the ovarium involves pro- cesses which are essentially the same, whether it be impregnated or not; but the subsequent changes differ in the two cases, so that the corpus luteum which accompanies the pregnant state is usually a much larger and more highly-organized body, than that which is found in the ovary of the unimpregnated female. This difference may be due in part to the absence, in the latter case, of that special determination of blood to the genital organs, which takes place in the former.—It is obvious, then, that the presence of a small and imper- fect corpus luteum in the ovary, merely indicates that an ovum has been matured and discharged, and affords no evidence of impreg- nation or sexual intercourse. The presence of a large and character- istic corpus luteum, on the other hand, may be regarded as affording undoubted evidence that impregnation has taken place. When fully formed, the corpus luteum may be rather more than half an inch in one of its diameters, and rather less in the other; it usually forms a projection on the surface of the ovary, and occupies from one-fourth to one-half of the whole area of its section. It is frequently, how- ever, much smaller than this; and on the whole it may be said that the presence of a corpus luteum as large as a full-sized pea, is toler- ably certain evidence of impregnation. After delivery, the size of the corpus luteum rapidly diminishes, and in a few months it ceases to be recognisable as such; the cicatrix by which the ovum has escaped, however, remains visible for some time longer. 801. The increase of size which is observable in the ovule that is being prepared for fecundation, is chiefly due to an augmentation in the substance of the Yolk ; and this also becomes more firm and granular than before. But the most curious change is that which takes place in the Germinal Vesicle; for this, although previously in the centre of the yolk, now moves up towards the side of it which is nearest the surface of the ovary, and becomes flattened against the yolk-bag. At the same time, it ceases to present its ordinary pellucidity and becomes obscure ; and this alteration appears to be due to the development of a brood of young cells in its interior. From the recent observations of Mr. New- port and others, it would seem that it then bursts and sets these free, so that they become diffused through the yolk ; and as this change may happen before fecundation, it must be regarded as being preparatory to it, or at any rate as being independent of it. 802. The ova thus matured and prepared for fecundation, are dis- charged from the surface of the ovary by the process already described, Avhether sexual intercourse take place or not; and being received into the Fallopian tubes, they are by them conducted towards the uterus. Their transmission may be effected by a kind of peristaltic movement Avhich is observed to take place in these tubes during the periods of heat; or, it may be, by the action of the cilia which line them; the di- 452 OF GENERATION AND DEVELOPMENT. rection of both movements being the same,—namely, from the ovaries towards the uterus. If in their course they should not receive the fer- tilizing influence, they appear soon to die and to disintegrate ; but if they should be impregnated by contact with the spermatic fluid, they almost immediately begin to undergo the first of those changes, which tend to the production of a new organism. 803. Much discussion has taken place, with regard to the exact point at which the fertilization of the ovulum takes place ; but this does not seem to be a matter of much consequence, as we find the order of the diffe- rent steps to vary considerably in different classes of animals. Thus in many aquatic Mollusca, and even in a large proportion of the class of Fishes, there is no act of copulation whatever ; but the spermatic fluid when emitted by the male, is diffused through the water, and fertilizes the ova which have been deposited by the female in his neighborhood. In the Frog, again, and in other Reptiles, the spermatic fluid is emitted upon the ova, at the time that they are being extruded by the female. In many Insects and Crustacea, in which a single congress often serves to fertilize many thousand eggs, the deposition of which occupies a period of several weeks or months, the spermatic fluid is received and stored up in a saccular dilatation of the oviduct of the female, which is termed the spermo-theca ; and in this manner it serves to impregnate the ova, as they are successively developed, and are conveyed to the outlet of the oviduct. In Birds, we find that ova are often set free from the ovarium in a state of full maturity, but without fertilization ; and that they re- ceive their additional layer of albumen and their shelly envelope, in passing down the oviduct, so as, at the time of their deposition, to differ in no obvious particular from fertile eggs. It is doubtful, in regard to Mammalia, whether the act of fertilization takes place before the ovum has been completely extricated from the ovisac, or subsequently to its finally quitting the ovarium and being received into the Fallopian tube. It is quite certain that the spermatozoa frequently, if not invariably, find their way to the surface of the ovary, being carried thither by their own spontaneous movements ; and it seems on the whole most pro- bable, that the fertilization of the ova usually takes place before they have entirely escaped from the ovisac, or whilst they are still in the commencement of the Fallopian tube. It is not unlikely that the place of the act of fecundation varies, according to the point at which the ovule and the seminal fluid first come into contact,—which may depend upon the degree of maturity of the ova at the period of copulation. 804. Everything indicates that the contact of the Spermatozoon with the Ovulum is the one thing needful in the act of fecundation; and there is strong reason to believe, from Mr. Newport's recent observations, that when this contact occurs, the spermatozoa undergo solution ; and that it is in the absorption of the product of that solution into the interior of the ovum (thus blending, as in Plants, the contents of the " sperm-cell" with those of the " germ-cell"), that the act of fecundation essentially consists. Availing himself of the agency of caustic potass, which has been found to be a powerful solvent of the spermatozoa, Mr. N. applied this agent to the ova at determinate periods after the application of these bodies, which he had separated from the liquor seminis by filtra- FERTILIZATION OF THE OVUM. 453 Fig. 138. tion ; and he found that when the interval of time between the one application and the other was only one or two seconds, only the early stages of the process took place, and no embryo was produced; when an interval of five seconds was allowed, very few embryos were produced from a large number of ova ; but when the interval was fifteen seconds or more, the proportion of embryos produced was much greater. Thus it seems obvious that time is an important element in the fertilizing process ; and that fertilization may be incompletely effected, for want of a sufficient penetration of the product of the diffluence of the Sperma- tozoa. How this product acts upon the contents of the ovum, however, and whether one or many of the cells set free by the rupture or solution of the germinal vesicle are fertilized by it, have not yet been ascertained. 805. The first change which can be observed to be consequent upon fecundation in the Mammalian ovum is the " segmentation" of the yolk; the entire mass of which, though previously compact and uniform, resolves itself, first into two, then into four, then into eight seg- ments (Fig. 138); each segment containing a transparent vesicle, which may be surmised to be a descendant of the original germ-cell. By a continuance of the same process, the whole cavity of the vitelline sac, or zona pel- lucida, becomes occupied by spherical parti- cles of yolk (each containing a pellucid par- ticle), the aggregation of which gives it a mulberry-like appearance; and by its fur- ther continuance, the component dells becom- ing more and more minute, the mass comes to present a uniform finely-granular aspect. At this stage it does not appear that the se- veral segments of the yolk hare a distinct enveloping membrane; but an envelope is now formed around each of them, converting it into a cell, of which the included particle forms the nucleus. This happens first to the peripheral portions of the mass; and as its cells are fully developed, they arrange them- selves at the surface of the yolk into a kind of membrane; at the same time assuming a pentagonal or hexagonal shape from mutual pressure, so as to resemble pavement epithe- lium (Fig. 139). As the globular masses of the interior are gradually converted into cells, ,i_ i ,.i p j I. Progressive stages in the segmenta- tney also pass to the surface and accumulate tion oftheviteiius of the Mammalian there; thus increasing the thickness of the SSZrtJSwKffi'rfSfiS envelope alreadv formed by the more super- into.two; c, further subdivision, pro- r J ... ', . r ducing numerous segments. ficial layer of cells, while the central part of the yolk remains, filled only with a clear fluid, which seems to be the product of the liquefaction of some of the interior spherules. By this process, the external part of the yolk is converted into a kind of secon- 454 OF GENERATION AND DEVELOPMENT. dary envelope, constituting the germinal membrane; and as this forms a complete sac enveloping the liquefied yolk of the interior, and as the Fig. 139. A B Latter stage in the segmentation of the yolk of the Mammalian Ovum;—at A is shown the " mulberry ma,ss" formed by the minute subdivision of the vitelline spheres; at B, a further increase has brought its surface into contact with the vitelline membrane, against which the spherules are flattened. whole structure of the future embryo originates in its substance, it has been termed by Bischoff the blastodermic vesicle. 806. The Blastodermic Vesicle very soon after its formation, pre- sents at one point an opaque roundish spot, which is produced by an accumulation of cells and nuclei of less transparency than elsewhere; and it is within this, which is termed the area germinativa, that all the structures of the permanent organism originate. When seen in section, this mass of cells presents the aspect shown at Fig. 140, c. The Ger- minal Membrane increases in extent and thickness by the production of new cells; and it subdivides into two layers, which, although both at first composed of cells, soon presents distinctive characters, and are concerned in very different ulterior operations. The outer one of these is commonly known as the serous layer, and the inner as the mucous; the division is at first most evident in the neighborhood of the area germinativa; but it soon extends from this point, and implicates nearly the whole of the germinal membrane. 807. The area germinativa soon loses the rounded form which it at first possessed, and becomes first oval, and then pear-shaped. Whilst Fig. 140. Plan of early Uterine Ovum. Within the external ring, or zona pellucida, are the blastodermic vesicle, a; the yolk, b; and the incipient embryo, c. this change is taking place in it, there gradually appears in its centre a clear space, termed the area pellucida ; and this is bounded externally GERMINAL MEMBRANE. 455 by a more opaque circle (whose opacity is due to the greater accumula- tion of cells and nuclei in that part), which subsequently becomes the area vasculosa. In the formation of these two spaces, both the serous and mucous layers of the germinal membrane seem to take their share; but the foundation of the vertebral column and nervous centres appears to be laid chiefly if not entirely in the serous layer; whilst the mucous is afterwards concerned more especially in the formation of the nutritive apparatus. Between these a third layer subsequently makes its ap- pearance ; which, as the first vessels of the embryonic structure are formed in it, is termed the vascular layer. 808. Thus the first development of the Mammalian embryo is into a sac, enclosing the store of nutriment that has been prepared for it,— in fact, a stomach; and we shall presently see, that it is by the agency of the walls of this sac, that the nutrient materials which it encloses are prepared for being appropriated to the development of the more permanent part of the fabric, which is to be evolved from the centre of the mulberry mass. But we may here stop to notice the interesting fact, that the development of the ovum in the lowest classes of animals may almost be said to cease at this point; the external layer of the germinal membrane remaining as the integument; the internal layer becoming the lining of the stomach; and the space occupied by the yolk forming the digestive cavity, into which an entrance or mouth is formed, by the thinning away of the germinal membrane at a certain point, round which tentaeula or prolonged lips are usually developed. This is the essential part of the history of development in the simpler Polypes; and we see how remarkably it corresponds with the history of development of the lower Cryptogamic plants, in which the first-formed membranous expansion, or primary frond, remains as the permanent leaf.—In the Mammalia, on the other hand, the greater part of the germinal membrane, and of the cavity which it forms, have a merely temporary purpose; being cast off, when they have performed their function, like the cotyledons of Flowering Plants. 809. During the time which is occupied by these important changes, the Ovum passes through the Fallopian tubes, and makes its way into the Uterus. During its transit through the Fallopian tubes, the Mam- malian ovum,—like the ovum of Birds in its passage through the ovi- duct,—receives an additional layer of albuminous matter secreted from the walls of the tube; and this is surrounded by a fibrous membrane, whose structure and mode of formation have been described on a former occasion (§§ 181,182). The outer layer of this envelope, in the egg of the Bird, is further consolidated by the deposition of particles of car- bonate of lime in its areolae; but it undergoes no higher organization. In the Mammal, however, this new envelope (termed the Chorion) is a formation of great importance; being the medium through which the whole subsequent nutrition of the embryo is derived. This is at first taken in by means of a number of villous processes, proceeding from the entire surface of the Chorion, and giving it a spongy or shaggy appearance; these processes (which are composed of nucleated cells) serve as absorbing radicles, which draw in the fluids afforded by the parent; and they thus make up for the early exhaustion of the small 456 OF GENERATION AND DEVELOPMENT. supply of nutritious matter stored up in the ovum itself. The con- tained embryo appropriates the fluid which is thus imbibed, by simple absorption through its surface; and thus it is nourished until a more special provision for its development comes into action. The structure of this organ, termed the Placenta, cannot be understood, until the concurrent changes in the lining membrane of the Uterus have been considered. 810. This membrane, in its natural condition, presents on its free surface the orifices of numerous cylindrical follicles ; which are arranged parallel to each other, and at right angles to the surface. In the spaces between these follicles, the blood-vessels form a dense capillary network. When impregnation takes place, this mucous membrane swells and be- comes lax; its capillaries increase in size; the follicles are developed into glandular cavities, and become turgid with a white epithelium ; and the interfollicular spaces are crowded with nucleated cells, Avhich fill up the meshes of the capillary network. In this peculiar condition, the uterine mucous membrane is termed the Decidua. At a later period, the decidua may be found to consist of two distinct layers; the decidua ver/z, lining the uterus ; and the decidua reflexa, covering the exterior of the ovum. Much discussion has taken place with regard to the mode in which the decidua reflexa originates, and the question cannot even now be considered as determined. The view recently put forth by Coste is, perhaps, as probable as any. He considers that the ovum on its entrance into the uterine cavity, is partly imbedded in its thick vas- cular lining membrane, and that this swells up around it, like the granulations around the pea in an issue; so that at last the OArum be- comes completely invested by the special envelope thus formed, which closes in around it, constituting the decidua reflexa, and which is at first not in contact with the decidua vera at any part save Avhere it has sprung from it. As the ovum increases in size, the decidua reflexa grows with it, and is thus gradually brought into contact with the decidua vera which lines the uterus, the cavity between them being obliterated; and at a later period, the two coalesce, so that they are no longer distinguishable from each other. 811. When the ovum has arrived in the Uterus, therefore, and the villous tufts of its chorion are developed, these come into contactj in the first instance, with the epithelial layer which intervenes between them and the vascular decidua. Through this cellular membrane, therefore, the ovum must derive its nutriment from the vascular sur- face ; and it cannot be deemed improbable, that its office is to draw from the subjacent vessels the materials which are to serve for the nutrition of the ovum, and to present it to the villous tufts of the chorion. Each of these, as already mentioned, is composed of an assemblage of nucleated cells, which are found in various stages of development; and the villus seems to elongate by the development of new cells from the germinal spot at its free extremity, whilst, like the spongiole of the plant, it draws in nutriment from the soil in which it is imbedded. On the other hand, the Decidua at this early period, appears to be actively employed in preparing nutriment for the embryo; for its cellular layer is so abundant, as to form abed into FORMATION OF DECIDUA, AND VILLI OF CHORION. 457 which the tufts of the chorion are received; whilst its follicles are enlarged into glandulae of sufficient size, to allow these villi (in some Mammals at least) to extend themselves into their interior.—In its earliest grade of development, as already remarked, the chorion and its villi contain no vessels; and the fluid drawn in by the tufts is communicated to the embryo, by the absorbing powers of its germinal membrane. But when the tufts are penetrated by blood-vessels, and their communication with the embryo becomes much more direct, the means by which they communicate with the parent are found to be essentially the same;—namely, a double layer of cells, one layer belonging to the foetal tuft, the other to the vascular maternal surface. (See § 819.) 812. We now return to the Embryo itself; the general history of whose development has been already traced, up to the period at which the cluster of cells in the Area Germinativa is about to give origin to the permanent structures of the foetus. The parts first formed in the embryo of Vertebrated animals, are such as most characteristically distinguish them from all others ;—namely, the Vertebral Column, and Spinal Cord. The first indication of these consists in the formation of what is termed the primitive trace, which is a shallow groove, lying between two oval ridges (Fig. 141, v), known as the laminar dorsales. The form of these is changed with that of the area pellucida; at first they are oval, then pear-shaped, and at last become of a violin-shape. At the same time they rise more and more from the surface of the Area pellucida, so as to form ridges of higher elevation, with a deeper groove between them; and the summits of these ridges tend to approach one another, and gradually unite, so as to convert the groove into a tube. It is within this, that the Cerebrospinal Axis is after- Ayards formed; the brain being developed in the anterior dilated por- tion, and the spinal cord in the posterior more contracted part. The Fig. 141. The germ and surrounding parts, from a more advanced Uterine Ovum:—6, blastoderma, or germinal membrane; a g, area germinativa; c, cephalic extremity of the germ ; v, first indications of vertebrae; q, cau- dal extremity. former remains unclosed much longer than the latter. The tube within which the neural axis is thus enclosed, and which is entirely composed of nucleated cells (Fig. 16), is termed the Chorda dorsalis ; and it retains its embryonic type, in many of the lower Fishes, which neArer possess a true vertebral column. The elements of the vertebrae 458 OF GENERATION AND DEVELOPMENT. are developed in a fibrous membrane, with which the chorda dorsalis subsequently becomes invested ; the neural arches being the parts first formed. 813. During the progress of this change, another very important one is taking place, which has reference to the nutrition of the embryo during its further development. This is the formation of vessels in the substance of the germinal membrane; which vessels serve to take up the nourishment supplied by the yolk, as well as that derived from the chorion externally, and to convey it through the tissues of the embryo. These Vessels are first seen in that part of the vascular lamina of the germinal membrane, which immediately surrounds the embryo; and they form a network, bounded by a circular channel, which is known under the name of the Vascular Area (Fig. 142). This gradually extends itself, until the vessels spread over the whole of the germinal membrane. The vessels are probably formed by the coalescence of the original cells of the layer; and the first blood-discs which they contain seem to originate in the nuclei of these cells. This network of vessels serves to receive the nutritious matter contained in the yolk-bag, and to convey it to the embryo ; but the act of absorption seems to be per- formed here as elsewhere, by cells, a layer of Avhich always intervenes between the vascular layer and the yolk itself. These cells probably correspond in function with those of the villi of the intestinal canal in the adult (§ 243); as the vessels of the yolk-bag, or temporary diges- Fig. 142. Vascular area of Fowl's egg, at the beginning of the third day of incubation; a, a, yolk; 6, 6, b, b, venous sinus bounding tbe area, c, aorta; d, punctum saliens, or incipient heart; e, e, area pellucida; /, /, arteries of the vascular area; g, g, veins; h, eye. tive cavity, represent those of the alimentary canal, to be afterwards developed from a portion of it. The vessels of the yolk-bag terminate in two large trunks, which enter the embryo at the point that after- wards becomes the umbilicus, and which are known as the Omphalo- Mesenteric, Meseraic, or Vitelline vessels (Fig. 146, q, r). The first movement of fluid takes place towards the embryo ; and this may be witnessed before any distinct heart is evolved. FORMATION OF VESSELS AND DIGESTIVE CAVITY. 459 814. The formation of the Heart takes place in a thickened portion of the Vascular layer; by the liquefaction of the interior of a mass of cells, of which the exterior constitute the first walls of the cavity. These gradually acquire firmness and consistency, and are endowed Avith a contractile power that enables them to execute regular pulsa- tions. In this early condition, the heart is known as the punctum saliens (d, Fig. 142). The first appearance of the heart in the Chick is at about the 27th hour; the time of its formation in Mammalia has not been distinctly ascertained. 815. Concurrently with the formation of the Vascular system, the production of the permanent Digestive cavity commences. This origi- nates in the separation of a small portion of the yolk-bag, lying imme- diately beneath the embryo, from the general cavity, in the following manner.—At about the 25th hour of incubation, in the Fowl's egg, the parts of the germinal membrane which lie beyond the extremities, and Fig. 143. Fig. 144. Diagram of Mammalian Ovum at later stage; the digestive cavity beginning to be separated from the jrolk-sac, and the amnion beginning to be formed:'— a, chorion; 6, yolk sac; c, embryo ;d and e, folds of the serous layer rising up to form the Amnion. SftffiKBS5^ The Amnion in process of formation, by the arching-over of the serous lamina:—a, the cho- rion ; 6, the yolk-bag, surrounded by serous and vascular laminae; c, the embryo: d, e, and/, ex- ternal and internal folds of the serous layer, forming the amnion; g, incipient allantois. which spread out from the sides of the embryo, are doubled in, so as to make a depression upon the yolk; and their folded edges gradually approach one another under the abdominal surface of the embryo, so as at last to meet and enclose a cavity, which is at first simple in its form, but which is subsequently rendered more complex by the prolongation and involution of its walls in various parts, so as to form the stomach and intestinal tube (Figs. 143, 144, 145). This digestive cavity com- municates for some time with the yolk-bag (from which it has been thus pinched off, as it were), by a wide opening, that is left by the imperfect meeting of the folds of the germinal membrane that constitute its walls. In the Mammalia, this orifice is gradually narroAved, and is at last com- pletely closed; and the yolk-bag, thus separated, is afterwards thrown off. It may be detected, however, upon the umbilical cord, up to a late period of pregnancy, and is known as the Umbilical vesicle (Fig. 146, t). In Birds, and other oviparous animals, the whole of the yolk-bag is 460 OF GENERATION AND DEVELOPMENT. ultimately drawn into the abdomen of the embryo; the former gradu- ally shrinking, as its contents are exhausted; and the latter enlarging, so as to receive it as a little pouch or appendage. In Fishes, the hatching of the egg very commonly takes place before the process has been completed; so that the little Fish swims about with the yolk-bag hanging from its body. 816. Whilst these processes are going on in the Vascular and Mu- cous layers of the germinal membrane, a remarkable change is taking place in that portion of the Serous lamina, which surrounds the Area pellucida. This rises up on either side in two folds (Fig. 143); and these gradually approach one another, at last meeting in the space between the general envelope and the embryo, so as to form an addi- tional investment to the latter. As each fold contains two layers of membrane, a double envelope is thus formed; of which the outer layer (Fig. 144, d, e, and Fig. 145, h) aftenvards adheres to the inner sur- face of the chorion, the original yolk-bag, or Zona pellucida, being now lost sight of; whilst the inner one (Fig. 144, /, /, and Fig. 145,/) remains as a distinct sac, to which the name of Amnion is given. This takes place during the third day in the Chick; but the period at which it occurs in the Human Ovum has not yet been clearly ascertained. 817. The embryo, like the adult, has. need of Respiration; in order that the carbonic acid set free in the Nutritive operations may be re- moved from its fluids. In Fishes, the surrounding water acts with suffi- cient power upon the vessels of the yolk-bag, to produce the required aeration, up to the time when the gills of the young animal are ready to come into play. But in the higher oviparous animals, Avhose deve- lopment proceeds further before they leave the egg, a more special pro- vision is made for the purpose. A bag, termed the allantois, sprouts (as it were) from the lower end of the body (Fig. 144, g); and gradu- ally enlarges, passing round the embryo (Fig. 145, g), so as in Birds Fig. 145. Diagram representing a Human Ovum in second month:—a, 1, smooth portion of chorion; a, 2, villous portion of chorion; k, k, elongated villi, beginning to collect into Placenta; b, yolk sac or umbilical vesicle; c, embryo; /, amnion (inner layer); g, allantois; h, outer layer of amnion, coalescing with chorion. almost completely to enclose it, intervening between the germinal mem- brane and the shell, and receiving the direct influence of the air that penetrates the latter. It is thus the temporary lung of the air-breathing RESPIRATION OF EMBRYO;—ALLANTOIS. 461 oviparous animal; and it serves for the aeration of its fluids, up to the time when it quits the egg. In the ovum of the Mammal, the chief office of the Allantois is to convey the vessels of the embryo to the cho- rion ; and its extent bears a pretty close correspondence with the extent of surface through which the chorion comes into vascular connexion with the decidua,—this extent varying considerably in the different orders of Mammalia. Thus in the Carnivora, whose placenta extends like a band around the whole ovum, the allantois also lines the whole inner surface of the chorion, except where the umbilical vesicle comes into contact with it. On the other hand, in Man and the Quadrumana, whose pla- centa is restricted to one spot, the allantois conveys the foetal vessels to one portion only of the chorion (Fig. 146); although, according to Coste, Fig. 146. Diagram of the Circulation in the Human Ovum, at the commencement of the formation of the Placenta; —a, venous sinus, receiving all the systemic veins; 6, right auricle; b', left auricle; c, right ventricle; d, bulbus aorticus, subdividing into e, e', e", branchial arteries; /, arterial trunk formed by their confluence ; g, vena azygos superior; h, confluence of the superior and inferior azygos; j, vena cava inferior; A;, vena azygos inferior; m, descending aorta; n, n, umbilical arteries proceeding from it; o, umbilical vein; q, omphalo-mesenterlc vein ; r, omphalo-mesenteric artery, distributed on the walls of the vitelline vesicle, t ; v, ductus venosus; y, vitelline duct; z, chorion. it completely surrounds the embryo. When these vessels have reached the Chorion, they ramify in its substance, and send filaments into its villi; and in proportion as these villi form that connexion with the uterine structure which has been already described (§ 811), do the ves- sels increase in size. They then pass directly from the foetus to the chorion; and the allantois, which is no longer of any use, ceases to present itself as a distinct sac. The lower part of it, however, pinched off (as it were) from the rest, remains as the Urinary bladder; and the 462 OF GENERATION AND DEVELOPMENT. Urachus or suspensory ligament of the latter represents the duct by which the Allantois was originally connected with the abdominal cavity. 818. The connexion which is thus formed between the Vascular sys- tem of the foetus and that of the parent, is the only one that exists in the lower Mammalia; which are thus properly designated as "nonpla- cental." Each villus of the Chorion contains a capillary loop; this is enclosed in a layer of cells; and this again in a lamina of basement- membrane ;—the whole forming the foetal tuft. This comes into contact with the cellular decidua, which lies upon the basement-membrane covering the vascular layer of the decidua. Now the Placenta is com- posed of these very elements, arranged in a more complex manner. It is formed by an extension of the vascular tufts of the Chorion at certain parts; and a corresponding adaptation, on the part of the Uterine struc- ture, to afford to these an increased supply of nutritious fluid. These specially-prolonged portions are scattered, in the Buminants and some other Mammalia, over the whole surface of the Chorion, forming what are termed the Cotyledons; but in the higher orders, and in Man, they are concentrated in one spot, forming the Placenta. In some of the lower tribes, the maternal and foetal portions of the placenta may be very easily separated; the former consisting of the thickened decidua; and the latter being composed of the prolonged and ramifying vascu- lar tufts of the Chorion, which dip down into it. But in the Human placenta, the two elements are mingled together through its whole substance. 819. The foetal portion of the Placenta consists of the branches of the umbilical vessels ; which subdivide at the point at which they enter the mass, and form, by their minute ramifications, a large part of its substance. Each villus contains a capillary vessel, which forms a series of loops, communicating with an artery on the one side, and with a vein on the other; but the same capillary may enter several villi, before re-entering a larger vessel. The vessels of the villi (Fig. 147, g) are covered, as in the chorion, by a layer of cells (/), enclosed in base- ment-membrane (e); but the foetal tuft thus formed is enclosed in a second series of envelopes (a, b, c), derived from the maternal portion of the placenta; a space (d) being left between the two, however, at the extremity of the tuft. The vascular tufts not unfrequently extend beyond the uterine surface of the placenta, and dip down into the ute- rine sinuses, where they are bathed in the maternal blood.—The ma- ternal portion of the Placenta may be regarded as a large sac, formed by a prolongation of the internal coat of the great uterine vessels. Against the foetal surface of this sac, the tufts just described may be said to push themselves, so as to dip down into it, carrying before them a portion of its thin wall, which constitutes a sheath to each tuft. In this manner, the whole interior of the placental cavity, is intersected by numerous tufts of foetal vessels, disposed in fringes, and bound down by the membrane that forms its proper wall; just as the intestines are covered and held in their places by the peritoneum. Now as this dila- tation of the uterine blood-vessels carries the decidua before it, every one of the vascular tufts that dips down into it will be covered with a STRUCTURE OF PLACENTA. 463 layer of the cellular structure of the latter (Fig. 148, and Fig. 149, e); and this will also form a part of all the bands that connect and tie Fig. 147. Fig. 148. Extremity of a Placental villus;—a, external mem- brane of the villus, continuous with the lining mem- brane of the vascular system of the mother; 6, external cells of the villus, belonging to the placental decidua; c, c, germinal centres of the external cells ; d, the space between the maternal and foetal portions of the villus; e, the internal membrane of the villus, continuous with the external membrane of the chorion; /, the internal cells of the villus, belonging to the chorion; g, the loop of umbilical vessels. Portion of the external mem- brane, with the external cells, of a Placental villus;—a, cells seen through the membrane; 6, cells seen from within the vil- lus ; e, cells seen in profile along the edge of the villus. down the tufts (Fig. 149, g). The blood is conveyed into the cavity of the placenta by the "curling arteries," so named from their peculiar course (Fig. 149, c), which proceed from the arteries of the uterus ; and Fig. 149. Diagram illustrating the arrangement of the Placental Decidua:—a, decidua in contact with the interior of the uterus; 6, venous sinus passing obliquely through it by a valvular opening; c, a curling artery passing in tbe same direction; d, tbe lining membrane of the maternal vascular system, passing in from the artery and vein, lining the bag of the placenta, and covering e, e, the foetal tufts passing on to them from their stems from the foetal side of the cavity, also by the terminal decidual bars f,f, from the uterine sido, and from one tuft to the other by the lateral bar, g; h, h, separated foetal tufts, showing the internal membrane and cells, which, with the loops of umbilical vessels, constitute the true foetal portion of the tufts. it is returned by large, short, straight trunks, which pass obliquely through the decidua (Fig. 149, b), and discharge their contents into the great uterine sinuses. 820. There is no more direct communication between the Mother^ and Foetus than this; all the observations which have been supposed to prove a direct vascular continuity, being certainly fallacious. The function of the Placenta is manifestly double. The foetal tufts draw, from the maternal blood, the materials which are required for the nutrition of the embryo,—these materials having been first selected 464 OF GENERATION AND DEVELOPMENT. and partly elaborated by the two sets of intervening cells; and in this character, the foetal tufts resemble the villi of the intestinal surface, which dip down into the fluids of the alimentary canal, and absorb the nutritive material which they furnish. But the Placenta also serves as a respiratory organ; aerating the blood of the foetus, by exposing it to the influence of the oxygenated blood of the Mother; and in this respect the foetal tufts bear a close correspondence with the gills of aquatic animals, bringing the blood into relation with a surrounding fluid medium containing oxygen, which is imbibed by the blood in exchange for the carbonic acid given off. And it is probable, too, that the Placenta is to be regarded as an excreting organ; serving for the removal, through the maternal blood, of excrementitious matter, whose continued circulation in the blood of the foetus would be preju- dicial to it. 821. The formation of the Human Placenta commences in the latter part of the second month of utero-gestation; during the third, it acquires its proper character; and it subsequently goes on increasing, in accordance with the growth of the ovum. The vessels of the Uterus undergo great enlargement throughout; but especially at the part to which the Placenta is attached; and the blood, in moving through them, produces a peculiar murmur, which is usually audible with dis- tinctness at an early period of pregnancy, and which may be regarded (Avhen due care is taken to avoid sources of fallacy) as one of its most unequivocal physical signs. The sound is most commonly heard near the situation of the Fallopian tube of the right side; and it corresponds with the pulse of the mother. * 822. It would be inconsistent with the character and objects of this Treatise, to follow in any detail, the history of the development of the Foetus, during its intra-uterine life ; and a general account of the evo- lution of most of the chief organs, has been given in connexion with that of their structure. The condition of the circulating apparatus, however, at the period of birth, deserves especial notice. A general account of the development of the simple pulsating trunk, which consti- tutes its first form, into the four-cavitied heart of the higher Vertebrata, and of the conversion of the single trunk proceeding from it, with its four pairs of branchial arches, into the aorta and pulmonary arteries, with their chief subdivisions, has been already given (§ 566). Up to the time of birth, however, the partition betAveen the Auricles is incom- plete ; a large aperture, the foramen ovale, existing in it. There is also a direct communication between the pulmonary artery and the aorta, by the ductus arteriosus; and another direct channel between the umbi- lical vein and the vena cava, by the ductus venosus. 823. The following is the course of the Circulation of the Blood in the Foetus. The fluid brought from the Placenta by the umbilical vein .(Fig. 150, 3), is partly conveyed at once to the vena cava ascendens, by means of the ductus venosus (5), and partly flows through two trunks (4, 4), that unite with the portal vein (7) returning the blood from the intestines, into the substance of the liver, thence to be returned to the vena cava by the hepatic vein. Having thus been transmitted through the two great depurating organs, the placenta and the liver, the blood that FCETAL CIRCULATION. 465 enters the vena cava is purely arterial in its character; but being mixed in the vessels Avith the venous blood that is returned from the trunk and lower extremities, it loses this character in some degree, by the time that it reaches the heart. In the right auricle, Avhich it then enters, it would also be mixed with the venous blood which is brought down from the head and upper extremities by the de- scending cava; were it not that a very curious provision exists to im- pede (if it does not entirely prevent) any further admixture. This con- sists in the arrangement of the Eusta- chian valve, which directs the arterial current (that flows upward through the ascending cava) into the left side of the heart, through the foramen ovale, whilst it directs the venous current (that is being returned by the descending cava) into the right ven- tricle. When the ventricles contract, the arterial blood contained in the left is propelled into the ascending Aorta, and supplies the branches that proceed to the head and upper ex- tremities, before it undergoes any further admixture ; Avhilst the venous blood contained in the right ventricle, is forced into the Pulmonary artery, and thence through the ductus arte- riosus (17), which is like a continua- tion of its trunk, into the descending aorta, mingling with the arterial cur- rent Avhich that vessel previously con- veyed, and thus supplying the trunk and lower extremities with a mixed fluid. A portion of this is conveyed, by the umbilical arteries, to the Pla- centa ; in which it undergoes the renovating influence of the maternal blood, and from which it is returned in a state of purity. 824. Hence the head and superior extremities, whose development is required to be in advance of that of the loAver, are supplied with blood nearly as pure as that Avhich returns from the placenta; whilst the rest Fig. 150. The Foetal Circulation:—1. The umbilical cord, consisting of the umbilical vein and two um- bilical arteries ; proceeding from the placenta (2). 3. The umbilical vein dividing into three branches; two (4, 4) to be distributed to the' liver; and one (5), the ductus venosus, which enters the inferior vena cava (6). 7. The portal vein, returning the blood from the intestines, and uniting with the right hepatic branch. 8. The right auricle ; the course of the bloood is denoted by the arrow, proceeding from 8, to 9. the left auricle. 10. The left ventricle ; the blood fol- lowing the arrow to the arch of the aorta (11), to be distributed to the branches given off by the arch to the head and upper extremities. The ar- rows 12 and 13, represent the return of the blood from the head and upper extremities through the jugular and subclavian veins, to the supe- rior vena cava (14), to the right auricle (8), and in the course of the arrow through the right ventricle (15), to the pulmonary artery (16). 17. The ductus arteriosus, which appears to be a proper continuation of the pulmonary artery; the offsets at each side are the right and left pulmonary artery cut off; these are of extremely small size as compared with the ductus arterio- sus. The ductus arteriosus joins the descending aorta (18, 18), which divides into the common iliacs, and these into the internal iliacs, which become the hypogastric arteries (19), and return the blood along the umbilical cord to the pla- centa ; while the other divisions, the external iliacs (20), are continued into the lower extremi- ties. The arrows at the terminations of these vessels mark the return of the venous blood by the veins to the Inferior cava. 466 OF GENERATION AND DEVELOPMENT. of the body receives a mixture of this, with what has previously cir- culated through the system. The Pulmonary arteries convey little or no blood through the lungs ; the current of blood, propelled from the right ventricle, passing directly omvards through the ductus arteriosus, into the aorta.—At birth, hoAvever, the course of the circulation under- goes great changes, that it may be adapted to the new mode, in which the infant is henceforth to obtain its nutrition and to carry on its respi- ration. As soon as the lungs are distended by the first inspiration, a portion of the blood of the pulmonary artery is diverted into them, and there undergoes aeration ; and, as this proportion increases, with the full activity of the lungs, the ductus arteriosus gradually shrinks, and its cavity finally becomes obliterated. At the same time, the foramen ovale is closed by a valvular fold ; and thus the direct communication between the two auricles is cut off. When these changes have been accom- plished, the circulation, which was before carried on upon the plan of that of the higher Reptiles (§ 563), becomes that of the complete warm- blooded animal; all the blood which has been returned in a venous state to the right side of the heart, being transmitted through the lungs, be- fore it can reach the left side, or be propelled from its arterial trunks.— It is by no means unfrequent, however, for some arrest of devolopment to prevent the completion of these changes ; and various malformations, involving an imperfect discharge of the circulating and respiratory functions, may hence result. 825. The average length of time, which elapses between Conception and Parturition, in the Human female, appears to be 280 days, or 40 weeks. There can be little doubt, however, that Gestation may be occasionally prolonged for one, two, or even three weeks, beyond that period ; such prolongation not being at all unfrequent amongst the lower animals ; and numerous well-authenticated instances of it, in the Human female, being upon record. Upon what circumstances this departure from the usual rule is dependent, has not yet been ascertained; but it is a remarkable circumstance, ascertained by the observations of cattle- breeders, that the male has an influence upon the length of gestation,— a large proportion of cows in calf by certain bulls exceeding the usual period, and a small proportion falling short of it. In such cases, we must attribute the prolongation of the period to some peculiarity in the embryo, derived from its male parent. 826. The shortest period at Avhich Gestation may terminate, consis- tently with the life of the child, has not yet been precisely ascertained; the difficulty of determining the precise date of conception being usually such, in this case as in the preceding, as to prevent the exact length of the Gestation from being known. Thus, the commencement of preg- nancy being fixed by the time of the cessation of the Catamenia, when there is no more definite guide, it is obvious that the act of Conception may have taken place during any part of the interval that has elapsed since the last monthly period; and thus a doubt may exist as to the length of the Gestation, to the extent of from one to three weeks. There are very satisfactory cases on record, in which, from the degree of de- velopment of the infant at birth, as well as from other circumstances, it might be certainly known not to haAre attained 26 or 27 weeks, or LENGTH OF GESTATION. 467 little more than six months; and in which, by careful treatment, the infant was reared in a condition of health and vigor. And there is reason to believe, that infants have lived for some time, and might pro- bably have been reared under better management, that were born as early as the 24th or 25th week. 827. The act of Parturition, by which the foetus is expelled from the Uterus, is accomplished in part by the contractile poAver of the Uterus itself; and in part by the combined operation of the various mus- cles, Avhich press upon the abdominal cavity, and which effect the expul- sion of the faeces and urine. No definite account can be given of the reasons, Avhy this change should take place at the period which has been mentioned as its usual date ; but we are as much in the dark in regard to other periodic phenomena of Animal life; and we must probably look for its source in the maturation of the placental structure, which prepares it for detachment (like the dropping-off of a ripe fruit), and in the complete evolution of the contractile tissue of the uterus, the con- tractions of which may be considered to commence spontaneously when it has attained a certain epoch in its growth, just as do those of the heart in the embryo (§ 814). For some days previously to the com- mencement of labor, there is usually a slow contraction of the fibres of the fundus and body of the uterus, and a yielding of those of the cervix; so that the child lies lower, and the size of the abdomen dimi- nishes. This slow contraction is probably not dependent upon any act of the nervous system; but upon the direct excitement of the contrac- tility of the muscular substance of the uterus. When labor properly commences, however, the Spinal system of nerves comes into play, and the uterine contractions are of a reflex nature. As before, hoAvever, the act of contraction is confined to the fundus and body of the uterus ; the fibres of the cervix uteri, and of the vagina, being in a state of re- laxation, Avhich alloAvs them to yield to the pressure of the child's head. In the first stage of labor, the Uterine contractions appear to be alone concerned; and it is not until the head cf the child is passing through the os uteri, and is entering the vagina, that the assistance of the ab- dominal muscles is called in. These act, in the first instance, as in ordinary expiration ; but their power is much increased by the voluntary retention of the breath, so that the whole of their contractile force may be applied to the expulsion of the foetus. In a later stage of labor, this retention of the breath becomes involuntary, during the accession of the " pains ;" and the expulsion of the foetus is commonly effected Avith considerable force, especially if the previous resistance has been con- siderable. 828. The same action Avhich expels the foetus, usually detaches the placenta; and if the uterus contract with sufficient force, after this has been thrown off, the orifices of the vessels Avhich communicated with it are so effectually closed, that little or no hemorrhage from that source takes place. When efficient contractions do not occur, they may fre- quently be excited by pressure upon the uterus itself; by the applica- tion of cold to the abdominal surface, to the extremities, and (in severe hemorrhage) to the entire body; or by the application of the child to the nipple, Avhich will frequently at once succeed in producing the de- 468 OF GENERATION AND DEVELOPMENT. sired effect. The efficacy of these means,—the latter in particular,— obviously depends upon the influence of the spinal cord and its nerves upon the muscular fibres of the uterus; the application of cold to the surface, or the irritation of the nipple, occasioning a reflex action in the uterus. But it is probable that this organ has also considerable poAver of contracting, independently of the nervous system; thus there are well-authenticated cases on record, in which the foetus has been expelled after the somatic death (§ 65) of the parent; Avhich must have been in consequence of the persistence of the independent contractility of the Uterus, and the relaxed state of all the parts through which the child had to make its exit. 829. The cause of the occasional occurrence of the parturient efforts at an unusually early period, is as little understood as that of their ordi- nary action. There are some individuals, in whom this regularly happens at a certain month; so that it seems to be an action natural to them. In many cases, however, it may be traced to some undue exertion of body, or mental excitement; and not unfrequently to a general constitu- tional irritability, Avhich renders the system liable to be derange d by very trifling causes. Premature labor is ahvays to be prevented, if possible, being injurious alike to both mother and child; and for this pre- vention we have chiefly to rely upon rest and tranquillity of mind and body, and upon the careful avoidance of all those exciting causes, which are liable to produce uterine contractions by their operation upon the nervous system ; whilst, at the same time, any measures which will in- vigorate the body, without stimulating it, should not be overlooked. 830. A peculiar preparation is made, in the females of the class Mam- malia, for the sustenance of the infant during a long period after birth. This consists in the secretion of a fluid, from the glands termed Mam- mary, which contains all the elements that are required for the develop- ment of the body of the infant, during the first year. These glands present themselves in an almost rudimentary state, in some of the non- placental animals of the class ; consisting only of 151- a feAV large follicles, which open separately upon the surface (Fig. 110). In the higher Mammalia, hoAvever, we find it composed of vast numbers of minute follicles, clustered together upon excre- tory ducts. The general arrangement of these, in the human subject, is seen in Fig. 151; and in Fig. 115, the character of the follicles them- selves, and of the secreting epithelial cells they Termination of portion of . , i 1 • i • p • miik-duct in foiiicics; from a contain, as seen under a much higher magnifying c^r1aeniS^ power, has been already shown. Each Mammary gland consists of a number of, glandulae, which are held together by areolar and fibrous tissue ; this arrangement may probably have reference to the mobility, which it is requisite that the different parts of the mass should possess, one upon the other, in con- sequence of its situation upon the pectoralis muscle. The ducts con- verge and unite together ; so as at last to form ten or twelve principal trunks, Avhich terminate in the nipple. At the base of the nipple, these tubes dilate into reservoirs, Avhich extend beneath the areola, and to MAMMARY GLAND. 469 some distance into the gland, when the breast is in a state of lactation. These, which are much larger in many of the loAver Mammalia than they are in the Human female, seem to have for their office to contain a store of milk, sufficient to supply the immediate wants of the child when it is first applied to the breast; so that it shall not be disappointed, but shall be induced to proceed with sucking, until the draught be oc- casioned (§ 836). 831. The Mammary gland may be detected at an early period of foetal existence, and it then presents no difference in the male and female; and it continues to grow, in each sex, in proportion to the body at large, up to the period of puberty. At that epoch, however, the gland begins to undergo a great enlargement in the female; and by the age of twenty, it attains its full size previous to lactation. Even then, how- ever, the milk-follicles cannot be injected from the tubes. During preg- nancy, the mammary glands receive a greatly-increased quantity of blood. This determination often commences very early: and produces a feeling of tenderness and distension, which is a valuable sign (where it occurs in conjunction with others) of conception having taken place. The vascularity of the gland continues to increase during pregnancy; and, at the time of parturition, its lobulated character can be distinctly felt. The follicles cannot be readily injected, however, until the gland is in a state of complete functional activity; i. e., during lactation.— The Mammary gland of the Male does not undergo this increase of deve- lopment, except under certain peculiar circumstances to be presently noticed (§ 836); and it remains a sort of miniature picture of that of the female, varying in diameter from that of a large pea to an inch or even two inches. 832. The Milk, secreted by the Mammary glands, consists of Water, holding in solution the peculiar Albuminous substance termed Caseine, and various Saline ingredients, together with (in most cases) a certain form of Sugar; and having Oleaginous globules suspended in it. These globules appear to be surrounded by a thin pellicle, which keeps them asunder, so long as the milk remains at rest.—The existence of these elements in ordinary Milk, as that of the Coav, is made apparent by the processes to which it is subjected in domestic economy. If it be allowed to stand for some time, exposed to the air, a large part of the oleagi- nous globules come to the surface, in consequence of their inferior spe- cific gravity; and thus is formed the cream, which includes also a con- siderable amount of caseine, with the sugar and salts of the milk. These may be partly separated by the continued agitation of the cream, as in the process of churning ; this, by rupturing the emrelopes of the oil-glo- bules, separates it into butter, formed by their aggregation, and butter- milk, containing the caseine, sugar, &c. A considerable quantity of caseine, however, is still entangled with the oleaginous matter; and this has a tendency to decompose, so as to render the butter rancid. It may be separated by keeping the butter melted at a temperature of 180°, when the caseine will fall to the bottom, leaving the butter pure and much less liable to change; an operation which is commonly knoAvn as the clarifying of butter.—The Milk, after the cream has been removed, still contains the greater part of its caseine and sugar. If it be kept 470 OF GENERATION AND DEVELOPMENT. long enough, a spontaneous change takes place in its composition; an incipient change in the caseine being the cause of the conversion of the sugar into lactic acid; and this coagulating the caseine, by precipitating it in small flakes. The same precipitation may be accomplished at any time by the agency of various acids, especially the acetic, which does not act upon Albumen; but Caseine cannot be coagulated, like albu- men, by the influence of heat alone. The most complete coagulation of Caseine is effected by the agency of the dried stomach of the calf, knoAvn as rennet; Avhich exerts so powerful an influence as to coagulate the caseine of 1,800 times its weight of milk. It is thus that, as in the making of cheese, the curd is separated from the whey ; the former consisting chiefly of the caseine ; whilst the latter contains a large proportion of the saline and saccharine matter, which entered into the original composition of the milk. These may be readily separated by evaporation. 833. The principal characters of Caseine have been already stated (§ 172).—The Oleaginous matter consists, like the fats in general, of the two substances, elaine and stearine; but it also contains another substance peculiar to it, which is termed butyrine. This last (to which the characteristic smell and taste of butter are due) is converted by saponification into three volatile acids, of strong animal odor, to which the names of butyric, capric, and caproic acids have been given. This change may be effected, at any period, by treating the butyrine with alkalies ; but it may also take place by spontaneous decomposition, which is favored by time and moderate warmth.—The Sugar of Milk is pecu- liar as containing nearly 12 per cent, of water: so that it may be con- sidered as really a hydrate of sugar. It is nearly identical in its com- position with starch ; and may, like it, be converted into true sugar by the agency of sulphuric acid. But it is chiefly remarkable for its proneness to conversion into lactic acid, under the influence of a fer- ment or decomposing azotized substance.—The Saline matter contained in Milk appears to be nearly identical with that of the blood; with a larger proportion, however, of the phosphates of lime and magnesia, which amount to 2 or 2J parts in 1000. These are held in solution chiefly by the Caseine, which has a remarkable power of combining Avith them. 834. Thus ordinary Milk contains the three classes of organic prin- ciples, which form the chief part of the food of animals,—namely, the albuminous, the saccharine, and the oleaginous; together with the mi- neral elements, which are required for the development and consolida- tion of the fabric of the infant. It would appear, however, that the combination of all these is not necessary; but rather has reference to the composition of the food on which the animal is destined to be after- wards supported. Thus it has been lately shown that, in the Carnivora, the milk contains no sugar; which principle is altogether wanting in the food of the adult. Amongst the different species of Herbivorous animals, the proportion of the several ingredients varies considerably; and it is also liable to considerable variation in accordance with the nature of the food, the amount of exercise taken by the animal that affords it, and other circumstances. Thus in the milk of the Coav, Goat, COMPOSITION OF MILK. 471 and Sheep, the average proportions of Caseine, Butter, and Sugar are nearly the same one with another, each amounting to from 3 to 5 per cent. In the milk of the Ass and Mare, on the other hand, the pro- portion of Caseine is under 2 per cent., the oleaginous constituents are scarcely traceable, whilst the sugar and allied substances rise to nearly 9 per cent. In the human female, the saccharine and oleaginous ele- ments are both present in large amount; whilst the Caseine forms a moderate proportion.—The proportion of saccharine and oleaginous elements appears to be considerably affected by the amount in which these are present in the food; and by the degree in which the quantity ingested is consumed by the respiratory process. Thus, a low external temperature, and out-door exercise, by increasing the production of carbonic acid from the lungs, occasion the consumption of the oleagi- nous and saccharine matters, which might otherwise pass into the milk, and thus diminish the amount of cream. On the other hand, exer- cise favors the secretion of caseine; which would seem to shoAV, that this ingredient is derived from the disintegration of the azotized tissues. Thus in Switzerland, the cattle which pasture in exposed situations, and which are obliged to use a great deal of muscular exertion, yield a very small quantity of butter, but an unusually large proportion of cheese; yet the same cattle, Avhen stall-fed, give a large quantity of butter, and very little cheese. 835. The Milk first secreted after parturition, knoAvn as the Colos- trum, is very different from ordinary milk, and possesses a strongly- purgative action, which is useful in clearing the bowels of the infant, from the various secretions Avhich have accumulated in them at birth, constituting the meconium. The Colostrum, when examined Avith the Microscope, is found to contain a multitude of large yellow granulated corpuscles ; each of which seems composed of a number of small grains aggregated together. The Colostric character is sometimes retained for some time after birth, and severely affects the health of the infant. This may happen without any peculiarity in the ordinary characters of the secretion, Avhich has all the appearance of healthy milk ; but the Microscope at once detects the difference, by the presence of the colos- tric corpuscles. 836. The formation of this Secretion is influenced by the Nervous system, to a greater degree, perhaps, than that of any other. The process may go on continuously, to a slight degree, during the Avhole period of lactation; but it is only in animals that have special reser- voirs for the purpose, that any accumulation of the fluid can take place. In the Human female, as Ave have seen, these are so minute as to hold but a trifling quantity of milk ; and the greater part of the secretion is actually formed whilst the child is at the breast. The irritation of the nipple produced by the act of suction, and the mental emotion connected with it, concur to produce an increased flow of blood into the gland, Avhich is known to Nurses as the draught; and thus the secretion is for the time greatly augmented. The draught may be produced simply by the emotional state of mind, as by the thought of the child when absent; and the irritation of the nipple may alone occasion it; but the tAvo influences usually act simultaneously. 472 OF GENERATION AND DEVELOPMENT. The most remarkable examples of the influence of such stimuli on the Mammary secretion, are those in which milk has been produced by girls and old women, and even by men, in quantity sufficient for the support of an infant. The application of the child to the nipple in order to tranquillize it, the irritation produced by its efforts at suction, and the strong desire to furnish milk, seem in the first instance to occasion an augmented nutrition of the gland, so that it becomes fit for the performance of its function ; and then to produce in it that state of functional activity, the result of which is the production of Milk. 837. It is not only in this way, that the Mammary secretion is influenced by the condition of the mind; for it is peculiarly liable to be affected as to quality, by the habitual state of the feelings, or even by their temporary excitement. Thus a fretful temper not only lessens the quantity of milk, but makes it thin and serous, and gives it an irritating quality ; and the same effect will be produced for a time by a fit of anger. Under the influence of grief or anxiety, the secretion is either checked altogether, or it is diminished in amount, and deteriorated in quality. The secretion is usually checked altogether by terror ; and under the influence of violent passion, it may be so changed in its characters, as to produce the most injurious and even fatal consequences to the infant. So many instances are noAV on record, in Avhich children, that have been suckled within a few minutes after the mothers have been in a state of violent rage or terror, have died suddenly in convul- sive attacks, that the occurrence can scarcely be set doAvn as a mere coincidence ; and certain as we are of the deleterious effects of less severe emotions upon the properties of the milk, it does not seem un- likely that, in these cases, the bland nutritious fluid should be converted into a poison of rapid and deadly operation. 838. Of the quantity of Milk ordinarily secreted by a good Nurse, it is impossible to form any definite idea ; as the amount which can be artificially drawn, affords no criterion of that which is ordinarily secreted at the time of the draught. The quantity Avhich can be squeezed from either breast at any one time, and, which, therefore, must have been contained in its tubes and reservoirs, is about two ounces. The amount secreted Avill depend upon several circumstances; such as the nature and amount of the ingesta; the state of bodily health; and the condi- tion of the mind. An adequate but not excessive supply of nutritious food, in which the farinaceous, oleaginous, and albuminous principles are duly blended; a vigorous but not plethoric constitution, regular habits, and moderate exercise, together with a cheerful and tranquil temper, altogether produce the most beneficial influence upon the secretion. It is seldom that stimulating liquors, which are so commonly indulged in, are anything but prejudicial; but the unmeasured con- demnation of them, in which some writers have indulged, is certainly injudicious ; as experience amply demonstrates the improvement in the condition both of mother and infant, Avhich occasionally results from the moderate employment of them.—In the administration of medicines to the mother, it is very desirable that the tendency of soluble saline substances to pass into the milk, and thus to affect the child, should be borne in mind. The vegetable substances used in medicine seem to GENERAL FUNCTIONS OF NERVOUS SYSTEM. 473 have much less disposition to pass off by this secretion; and they are consequently to be preferred during lactation. 839. From the close correspondence which exists between the ele- ments of the Milk and those of the Blood, it is evident that we cannot expect to trace the existence of the former, as such, in the circulating current. It is interesting, however, to remark, that a preparation appears to be taking place in the laboratory of the system, for the pro- duction of this secretion, long before the period of parturition. The Urine of pregnant Avomen almost invariably contains a peculiar sub- stance termed kiestine, which is nearly related to caseine, and which disappears from the urine as soon as lactation has fully commenced. It would seem, therefore, that a compound of this nature is in course of preparation during pregnancy; and that it is eliminated by the kidney, until the Mammary Gland is prepared for the active perform- ance of its functions.—That the kidney may relieve the system from the accumulation of other constituents of the mammary secretion, appears from a case recently put on record; in which the urine of a parturient female, who did not suckle her infant, was found to contain a considerable quantity of butyric acid, during several days. There can be no doubt that, in ordinary states of the system, this secretion can- not be required for the depuration of the blood, since it does not occur in the male at all, and is present in the female at particular times only. But these facts afford ground to believe that, when the process is going on, certain products are generated in the system, Avhich are not found there at other times. And it is quite certain that the sudden checking of the secretion, or the reabsorption of the fluid already poured out, occasioning an accumulation of these substances in the circulating cur- rent, may give rise to very injurious consequences. Some very curious instances are on record, in which a transference of the secreting power to some other surface has taken place under such circumstances; so as to relieve the system from the accumulation in question. CHAPTER XII. OF THE NERVOUS SYSTEM AND ITS ACTIONS. 1. General View of the operations, of which the Nervous System is the instrument. 840. We have now considered the entire series of those operations, Avhich make up the vegetative or organic life of the Animal; including those functions by Avhich the germ is prepared, by which it is nourished until it can be left to its OAvn powers, by which its continued development is effected until the fabric characteristic of the adult has been built up, and by Avhich the normal constitution is maintained through a length- ened period,—so long as the necessary materials are supplied, and no 474 OF THE NERVOUS SYSTEM AND ITS ACTIONS. check or hindrance is interposed, by external influences, to that regular sequence of changes, on which the continuance of its poAvers depends. In this survey it will have been perceived, that the essential parts of these operations are, in Animals as in Plants, completely independent of the influence of that which constitutes the peculiar endowment of Animals; namely, the Nervous System. a. The Reduction of the food in the Stomach, by the solvent power of the gastric fluid, is a purely chemical operation, with which the Ner- vous System has nothing whatever to do, excepting that it perhaps accelerates the process, by stimulating the Muscular coat of the stomach to that peculiar series of contractions, which keeps the contents of the cavity in continual movement, and favors the action of the solvent upon it. b. In the process of Absorption, by which the nutritive materials, with other substances, are introduced into the vessels, the Nervous Sys- tem has no participation ; this being a purely vegetative operation, partly dependent upon the simple physical conditions which produce Endos- mose, and partly on a process of cell-groAvth. c. The Assimilation of the neAV material, effected, as we have seen reason to believe, by another set of independent cells, can receive but little influence from the Nervous System, and is obA'iously capable of taking place without its aid. d. The Circulation of the Blood, again, though dependent in part upon the impulsive poAver of a Muscular organ, the heart, is not on that account brought into closer dependence upon the Nervous System; for we have seen that the contractions of the heart result from its own inherent poAvers, so as to continue after it has been completely detached from the body ; and that the capillary poiver, which is the chief agent in the movement of the blood in the loAver animals, and Avhich exerts an important subsidiary action in the higher, is the result of the exercise of certain affinities betAveen the blood and the surrounding tissues, in which the Nervous System can have no immediate concern. e. The act of Nutrition, in which every tissue draAvs from the circu- lating blood the materials for its own continued growth and develop- ment, and by which it incorporates these with its own substance, is but a continuance of the same kind of operation as that which takes place in the early development of the embryo, long anteriorly to the first appearance of the nervous system,—namely, a process of cell-develop- ment and metamorphosis, Avhich must be, from its very nature, indepen- dent of Nervous agency. /. The same may be said of the Secreting operation in general; for this essentially consists in the separation of certain products from the blood, by cells situated upon free surfaces; Avhich thus remove those products from the interior of the fabric. g. And the interchange of oxygen and carbonic acid, which takes place between the atmosphere and the venous blood, Avhen brought into mutual relation in the lungs, and which is the essential part of the func- tion of Respiration, is an operation of a merely physical character, with Avhich the Nervous system can have no direct concern. h. Finally, the development of the " sperm-germ-cells" in the one GENERAL FUNCTIONS OF NERVOUS SYSTEM. 475 sex, and of the ova containing germ-cells in the other, the subsequent fertilization of the latter by the former, and the changes consequent upon that act, together making up the function of Generation, may be all regarded as modifications of the ordinary Nutritive processes; and are effected, like these, by the inherent powers of the parts concerned in them, at the expense of the materials supplied by the blood, without any direct dependence upon the Nervous system. ^ 841. Still, although the various processes, which make up the essen- tial part of the nutritive operations, in Animals as in Plants, are no more dependent on any peculiar influence derived from a Nervous sys- tem, in the former, than they are in the latter, it must be evident, from the details already given, that there must be in Animals various acces- sory changes, which are requisite for the continuance of the former, and which can only be effected by the peculiar powers with which Animals are endoAved.—Thus, to commence with Digestion: this preliminary process, Avhich the nature of the food of the plant renders unnecessary for its maintenance, can only be accomplished by the introduction of the food into a cavity or sac, in which it may be submitted to the action of the solvent fluid. The operation of grasping and swallowing the food, wherever it is performed, is accomplished through the agency of the Nervous system ; and if it be checked by the loss of Nervous power, the Digestive process must cease for want of material.—So, again, although interchange of gaseous ingredients betAveen the atmosphere and the circulating fluid may take place with sufficient energy in Plants and the lower Animals, through the mere exposure of the general surface to the atmosphere, yet we find that, in all the higher Animals, certain move- ments are requisite, for the continual reneAval of the air or water Avhich are in contact with one side of the respiratory surface, and of the blood which is in relation with the other: for the direction of which move- ments a Nervous system is requisite.—In the excretory processes, more- over, the removal of the effete matters from the body can only be accomplished, in the higher Animals, by certain combined movements; the object of which is, to take up the products that are separated by the action of the proper secreting cells, and to carry them to the exte- rior of the body, there to be set free; and these combined movements can only be effected by the agency of the Nervous system.—Lastly, in the act of Reproduction, the arrangement of the sexual organs in Animals requires that a certain set of movements should be adapted to bring together the contents of the "sperm-cells" of the male, and of the "germ- cells" of the female; and also for the expulsion of the ovum from the body of the latter, in a state of more or less advanced development. For these movements a special arrangement is made, in the construction of the Nervous system, and in the application of its peculiar poAvers. 842. Thus we see that, although the Organic functions of the Animal are essentially independent of the Nervous System, this system affords the conditions which are requisite for their continued maintenance; being the instrument whereby the muscles are called into action for the performance of the various combined actions, that constitute the mecha- nism (so to speak) by which the Vegetative part of the fabric is com- bined with the Animal portion of the organism. We are not to suppose, 476 OF THE NERVOUS SYSTEM AND ITS ACTIONS. hoAvever, that every movement which takes place in the Animal body is dependent upon the Nervous System ; for we have seen that the Muscu- lar tissue may be employed to perform contractions excited by stimuli applied to itself, and that it may thus execute a set of movements in which the nervous system has no direct participation. And it is desirable that the Student should observe, that these are, in all instances, those most directly connected with the Vegetative functions, and, at the same time, those of the simplest and most straightforward character.—Thus, the peristaltic movement, by which the alimentary and foecal matters are propelled along the Intestinal tube, results from the direct excite- ment of the contractility of its muscular wralls, and is entirely indepen- dent of Nervous agency; and this movement is accomplished by the successive contraction of the different fasciculi surrounding the tube, which take up (as it were) each others' action (§ 352). So again, the successive contractions and dilatations of the cavities of the Heart, which perform so important a part in the Circulation of the blood, are the result of the properties inherent in that organ; the muscular fibres of which are excited to a peculiar rhythmical and consentaneous con- traction, by the flow of blood into the cavities when dilating. More- over, in the Excretory ducts of various glands, we find a Muscular coat, by which the fluids secreted in the glands are propelled towards their outlet on the exterior of the body, or on one of its free internal surfaces. 843. In these instances, then, we observe that the simple Contractility of Muscular structure, excited by direct stimulation, is applied to effect the movements most closely connected with the Organic functions. With the processes, therefore, which take place in the penetralia of the system, the Nervous System has no direct concern. Its office is to guard the portals for entrance and exit; and to fill those chambers, which admit the new materials from the external world; or to empty the receptacles, which collect from the interior of the system the effete matters that are to be cast out from it. And we find that, for these offices, the Nervous system is employed in its very simplest mode of operation ;—that which does not involve Sensation, Intelligence, Will, or even Instinct (in the proper sense of that term), but which may take place independently of all consciousness,—by the simple reflexion of an impression, conveyed to a ganglionic centre by one set of fibres proceeding towards it from the circumference, along another set which passes from it to the mus- cles, and calls them into operation (§ 394). This reflex function, there- fore, is the simplest application of the Nervous System in the Animal body. We shall presently see reason to believe, that a very large pro- portion of the movements of many of the lower animals are of this reflex character; and that they are not necessarily accompanied by sensation, although this may usually be aroused by the same cause which produces them. As we rise, however, in the scale of Animal existence, we find the reflex movements forming a smaller and smaller proportion of the whole; until, in Man, they constitute so limited a part of the entire series of movements of which the Nervous system is the agent, that their very existence has been overlooked. 844. But the main purpose of the Nervous System is to serve as the DEPENDENCE OF MENTAL ACTIONS UPON SENSATIONS. 477 instrument of the Psychical* powers, which are the distinguishing attribute of the Animal. It has been already pointed out, that the pos- session of Consciousness (or of the capability of receiving sensations), and the power of executing Spontaneous Movements (that is, movements which are not immediately dependent upon external stimuli), constitute the essential features in which the Animal differs from the Plant. All the other differences in structure, that respectively characterize these two classes of living beings, are subordinate to this one leading distinc- tion,—the presence of a Nervous system, and of its peculiar attributes in the one,—and its absence in the other. Noav when Ave attempt to analyze those peculiar attributes, we may resolve them, like the pro- perties of the material body, into different groups. We find that the first excitement of all mental changes, whether these involve the action of the feelings or of the reason, depends upon sensations; which are produced by impressions made upon the nerves of certain parts of the body, and are conveyed by these to a particular ganglionic centre, which is termed the sensorium,—being the part in which Sensation, or the capability of feeling external impressions, especially resides. 845. Noav there are numerous actions, especially among the lower Animals, which seem to be as far removed from the influence of the Will, and as little directed by Intelligence, as the Reflex movements themselves; but which, nevertheless, depend upon sensation for their excitement. The sensation may immediately direct the movement, and may call the muscular apparatus into action in such a manner, as, with- out any calculation of consequences, any intentional adaptation of means to ends, any exertion of the reason, or any employment of a discrimi- nating Will, to produce an action, or train of actions, as directly and obviously adapted to the Avell-being of the individual, as we have seen those of the reflex character to be. Of this we have an excellent example in the act of Sneezing; the purpose of Avhich is obviously to expel from the nasal passages those irritating matters, the sense of whose presence excites the complicated assemblage of muscular move- ments, concerned in the operation. This class of actions may be appro- priately termed the Consensual; and under it Ave may include most of those purely instinctive actions of the loAver animals, Avhich, being prompted by sensations, cannot be assigned to the reflex group. These seem to make up, with the reflex, nearly the whole of the Animal functions in many tribes; but they are found to be gradually brought under the domination of the Intelligence and Will, as we rise towards Man, in whom these faculties are most strongly developed, so as to keep the Consensual as well as the Reflex actions quite in subordination to the more elevated purposes of his existence.—Closely allied, however, to these, are the purely Emotional movements; in which the sensation excites a mental feeling or impulse, that reacts upon the muscular system without giving rise to any distinct idea, and consequently without having called the intellect and will into exercise. In fact, these emotional movements are often performed in opposition to the * This term, derived from the Greek -ji/;^, is used to designate the sensorial and mental endowments of Animals, in the most comprehensive acceptation of those terms. 478 OF THE NERVOUS SYSTEM AND ITS ACTIONS. strongest efforts of the will to restrain them; as when laughter is provoked by some ludicrous sight or sound, or by the remembrance of such, at an unseasonable time. It is probable, from the strong mani- festations of emotion exhibited by many of the lower animals, that some of the actions which Ave assemble under the general designation of instinctive, are to be referred to this group. 846. There are many sensations, however, which do not thus imme- diately give rise to muscular movements; their operation being rather that of stimulating to action the Intellectual powers. There can be little doubt that all Mental processes are dependent, in the first instance, upon Sensations, which serve to the Mind the same kind of purpose that food and air fulfil in the economy of the body. If we could imagine a being to come into the Avorld with its mental faculties fully prepared for action, but destitute of any power of receiving sensations, these faculties would never be aroused from the condition in Avhich they are in profound sleep; and the being must remain in a state of com- plete unconsciousness, because there is nothing of which it can be made conscious, no kind of idea which can be aroused within it. But after the mind has once been in active operation, the destruction of all future power of receiving sensations would not reduce it again to the inactive condition. For sensations are so stored up in the mind by the power of Memory, that they may give rise to ideas at any future time; and thus the mind may feed, as it were, upon the past. Now the ideas which are excited by sensations, and which are colored by the state of Feeling which accompanies them, become the subjects of Reasoning processes more or less complex, sometimes of the utmost brevity and simplicity, sometimes of the most refined and intricate nature. These reasoning processes, when they result in a determination to execute a particular movement, execute that movement by an act of Volition; the peculiar character of which is that it is the expression of a definite purpose, of a designed adaptation of means to ends, on the part of the individual performing it, instead of being the result of the mere blind, indiscriminating impulse, which seems to be the mainspring of the instinctive operations. It is in Man that we find the highest develop- ment of the reasoning faculties; but it is quite absurd to limit them to him, as some have done, since no impartial observer can doubt that many of the lower animals can execute reasoning processes, as complete in their way as those of Man, though much more limited in their scope. 847. Thus, then, wTe have to consider the Nervous system under four heads;—first, as the instrument of the Reflex actions;—second, as the instrument of the Consensual actions;—third, as the instrument of the Emotional actions;—and fourth, as the instrument of the Intellectual processes and of Voluntary movements. There is reason to believe that the Nervous Centre from which the muscles derive their impulse to contract is the same, whether the movement be prompted by an impres- sion which does not excite the consciousness, by a sensation, by an emotion, or by a volition; and that this instrument may be played upon, .so to speak, by other centres, which minister to these functions respec- tively. In order that the relations of the component parts of the Nervous apparatus may be better understood, it will be desirable to NERVOUS SYSTEM OF RADIATA. 479 take a brief survey of its comparative structure in the principal groups ot Animals ; and to inquire what actions may be justly attributed to its several divisions in each instance,—commencing with those in which the structure is the simplest, and the variety of actions the smallest; and passing on gradually to those in which the structure is increased in complexity by the addition of new and distinct parts, and in which the actions present a corresponding variety. 2. Comparative Structure and Action of the Nervous System. 848. From what has been already said (§ 373-9) of the characters ot the two elementary forms of the Nervous tissue, it is evident that no JNervous system can exist, in which both these forms should not be present. We look, therefore, for ganglia composed of the vesicular nervous substance, and serving as the centres of nervous power; and lor cords or trunks, composed of the tubular substance, and serving to communicate between the ganglia and the parts with which they are to be functionally connected. Now it is quite certain that, at present, no such Nervous apparatus can be detected in many of the lowest Animals; and some Physiologists have had recourse to the supposition of their possessing a "diffused" nervous system; that is, of their possessing nervous particles, in a separate form, incorporated, as it were, with their tissues. But we have seen that each tissue possesses its own pro- perties, and can perform its own actions independently of the rest;__ that even the contractility of Muscular fibre is by no means dependent upon the Nervous system, though usually called into play through its means;—and that the simplest office of a Nervous System is to pro- duce a muscular movement in respondence to a certain impression; which action requires that it should have an internuncial or commu- nicating poAver, only to be exercised (so far as we at present know) by continuous fibres. _ The apparent absence of a Nervous system is doubtless to be attributed, in many instances, to the general softness of the tissues of the body, which prevents it from being clearly made out among them. And it is to be remembered, that, on the principles already stated, we should expect to find it bearing a much smaller proportion to the entire structure, in the lowest Animals, whose functions are chiefly Vegetative,—than in the highest, in which the vegetative functions seem destined merely for the development of the Nervous and Muscular systems, and for the sustenance of their powers. 849. Among the Radiated classes, the parts of Avhose bodies are arranged in a circular manner around the mouth, and repeat each other more or less precisely, the Nenrous system presents a corresponding form. In the Star-fish, for example, Avhich is one of the highest of these animals, it forms a ring, which surrounds the mouth ; this ring consists of nervous cords, which form communications between the several ganglia, one of which is placed at the base of each ray. The number of these ganglia corresponds with that of the rays or arms • being five in the common Star-fish ; and from nine to fifteen, in the" species possessing those several numbers of members. The ganglia 480 OF THE NERVOUS SYSTEM AND ITS ACTIONS. appear to be all similar to one another in function, as they are in the distribution of their branches ; every one of them sending a large trunk along its OAvn ray, and tAvo small filaments to the organs in the central disk. The rays being all so similar in structure, as to be exact repeti- tions of each other, it would appear that none of the ganglia can have any controlling power over the rest. All the rays (in certain species) have at their extremities what seem to be very imperfect eyes ; and so far as these can aid in directing the movements of the animal, it ia obvious that they will do so towards all sides alike. Hence there is no one part, which corresponds to the head of higher animals ; and the ganglia of the nervous system, like the parts they supply, are but repe- titions of one another, and are capable of acting quite independently. Each would perform its own individual functions if separated from the rest; but, in the entire animal, their actions are all connected with each other by the circular cord, which passes from every one of the five gan- glia to those on either side of it. We shall find that, in Articulated and Vertebrated animals, there is a similar repetition of corresponding ganglia on the two sides of the median plane of the body; and that these are connected by transverse bands, analogous in function to the circular cord of the Star-fish. Moreover, we shall see a like repetition of ganglia, almost or precisely similar in function, in passing from one extremity of these animals to the other; and these ganglia are con- nected by longitudinal cords, whose function is in like manner commis- sural.—From the best judgment wre can form of the actions of the Star-fish, by comparing them with the corresponding actions of higher animals, we may fairly regard the greater number of them as simply re- flex ; being performed in direct respondence to external stimuli, the impression made by which is propagated to one or more of the ganglia, and excites in them a motor impulse. How far the movements of these animals are indicative of sensation, we have not the power of deter- mining ; but it may be safely affirmed, that they afford no indication of the exercise of reasoning faculties or of voluntary power. 850. Perhaps the simplest form of a Nervous system is that pre- sented by certain of the loAver Mollusca; for the body not here pos- sessing any repetition of similar parts, the nervous system is destitute of that multiplication of ganglia which we see in the Star-fish; whilst the limited nature of the animal powers involves a corresponding sim- plicity in the integral parts of their instrument. The animals, to which reference is here made, form the class Tunicata, Avhich is inter- mediate, in many respects, between the ordinary Molluscs and the Zoo- phytes. They consist essentially of an external membranous bag or tunic, within Avhich is a muscular envelope, and again Avithin this a respiratory sac, which may be considered as the -dilated pharynx of the animal. At the bottom of this last, is the entrance to the stomach; which, with the other viscera, lies at the lower end of the muscular sac. The external envelopes have two orifices ; a mouth, to admit water into the pharyngeal sac ; and an anal orifice, for the expulsion of the water which has served for respiration, and of that which has passed through the alimentary canal, together with the faecal matter, the ova, &c. A current of water is continually being drawn into the NERVOUS SYSTEM OF MOLLUSCA. 481 pharyngeal sac, by the action of the cilia that line it; and of this, a part is driven into the stomach, conveying to it the necessary supply of aliment in a very finely-divided state; Avhilst a part is destined merely for the aeration of the circulating fluid, and is transmitted more directly to the anal orifice, after having served that purpose. These animals are for the most part fixed to one spot, during all but the earliest period of their existence; and they give but little external manifestation of life, beyond the continual entrance and exit of the currents already adverted to, which being effected by ciliary action, is altogether independent of the nervous system (§ 234). When any substance is^ drawn in by the current, however, the entrance of which would be injurious, it excites a general contraction of the mantle or muscular envelope; and this causes a jet of water to issue from one or both orifices, which carries the offending body to a distance. And, in the same manner, if the exterior of the body be touched, the mantle suddenly and violently contracts, and expels the contents of the sac. 851. These are the only actions, so far as we know, which the Ner- vous system of these animals is destined to perform. They do not ex- hibit the least trace of eyes, or of other organs of special sense; and the only parts that appear peculiarly sensitive, are the small tentacula, or feelers, that guard the oral orifice. BetAAreen the tAvo apertures in the mantle, Ave find a solitary ganglion, which receives branches from both orifices, and sends others over the muscular sac. This, so far as we knoAv at present, constitutes the whole nervous system of the animal; and it is fully sufficient to account for the movements which have been described. For the impression produced by the contact of any hard substance with the tentacula, or with the general surface of the mantle, being conveyed by the afferent fibres of this ganglion, will excite in it a reflex motor impulse ; which, being transmitted to the muscular fibres of the contractile sac, as well as to those circular bands that surround the orifices and act as sphincters, will produce the movements in ques- tion. 852. In the Conchifera, or Molluscs inhabiting bivalve shells, there are invariably two ganglia, having different functions. The larger of these (Plate II., Fig. 1, e), corresponding to the single ganglion of the Tunicata, is situated towards the posterior end of the body (that is the end most distant from the mouth), in the neighborhood of the posterior muscle that draws the valves together; and its branches are distributed to that muscle, to the mantle, to the gills (d, d), and to the siphons (e, e)> by which the water is introduced and carried off. But we find another ganglion, or rather pair of ganglia (a, a), situated near the front of the .body, either upon the oesophagus, or at its sides; these ganglia are connected with the very sensitive tentacula which guard the mouth; and they may be regarded as presenting the first approach, both in position and functions, to the brain of higher animals. In the Oyster, and others of the lower Conchifera, which have no foot, these are the only principal ganglia: but in those having a foot,—which is a muscular tongue-like organ,—we find an additional ganglion (b) con- nected with it. This is the case in the Solen, or animal of the Razor- 31 482 OF THE NERVOUS SYSTEM AND ITS ACTIONS. shell; whose foot is a very powerful boring instrument, enabling it to penetrate deeply into the sand.—Here, then, we have three distinct kinds of ganglionic centres; every one of which may be doubled, or repeated on the tAvo sides of the body. First, the cephalic ganglia, a, a, which are probably the sole instruments of sensation and of the consen- sual movements; these are almost invariably double, being connected together by a transverse band, which arches over the oesophagus. Se- cond, the pedal ganglion, b, which is usually single, in conformity with the single character of the organ it supplies; but in one very rare Bivalve Mollusc, the foot is double, and the pedal ganglion is double also. Third, the respiratory ganglion, c, which frequently presents a form that indicates a partial dhrision into .two halves, corresponding with the repetition of the organs it supplies on the two sides of the body. Besides these principal centres, Ave meet with numerous smaller ones upon the nervous cords (/, /, and g, g,) which proceed from them to the different parts of the general muscular envelope or mantle. 853. Now it will be observed, that the two cephalic ganglia a, a, are connected with the pedal ganglion, b, by means of a pair of trunks pro- ceeding from the former to the latter; and that they are, in like man- ner, separately connected with the respiratory or branchial ganglion, c. There is good reason to believe, that the pedal and branchial ganglia minister to the purely reflex action of the organs they respectively sup- ply ; and that they would serve this purpose as well, if altogether cut off from connexion with the cephalic ganglia: whilst the cephalic, being the instruments of the actions which are called forth by sensation, exert a general control and direction over the movements of the animal, through the medium of the trunks by Avhich they communicate with the ganglia in immediate connexion with the muscular apparatus. It is difficult, however, to make satisfactory experiments upon this subject in these animals, their movements being for the most part slow and feeble, and their nervous system not readily accessible; and our idea of the re- spective functions of their ganglia is chiefly founded upon the distribu- tion of their nerves, and upon the analogous operations of the ganglia that correspond to them in other animals. 854. In ascending through the series of the Mollusca, we find the Nervous system increasing in complexity, in accordance with the gene- ral organization of the body: the addition of new organs of special sensation, and of new parts to be moved by muscles, involving the .addition of new ganglionic centres, Avhose functions are respectively adapted to these purposes. But we find no other multiplication of similar centres, than a doubling on the two sides of the body; excepting in a few cases, where the organs they supply are correspondingly mul- tiplied. We have a very characteristic example of this in the arms of ;the €uttle-fish, which are furnished with great numbers of contractile suckers, every one possessing a ganglion of its own. Here we can trace very clearly the distinction between the reflex actions of each individual sucker, depending upon the powers of its own ganglion; and the move- ment prompted by sensation which results from its connexion with the cephalic ganglia. The nervous trunk, which proceeds to each arm, may be distinctly divided into two tracts ; in one of which are contained NERVOUS SYSTEM OF MOLLUSCA. 483 the ganglia which peculiarly appertain to the suckers, and which are connected with them by distinct filaments; whilst in the other there is nothing but fibrous structure, forming a direct communication between these and the cephalic ganglia; so that each sucker has a separate rela- tion with a ganglion of its own, whilst all are alike connected with the cephalic ganglia, and are placed under their control. We see the results of this arrangement, in the modes in which the contractile power of the suckers may be called into operation. When the animal embraces any sub- stance with its arm (being directed to this action by its sight or other sensa- tion) it can bring all the suckers simultaneously to bear upon it; evidently by a voluntary or instinctive impulse transmitted along the connecting cords that proceed from the cephalic ganglia to the ganglia of the suckers. On the other hand, any individual sucker may be made to contract and attach itself by placing a substance in contact with it alone; and this action will take place equally well when the arm is separated from the body, or even in a small piece of the arm when recently severed from the rest,—thus proving that, when it is directly excited by an impres- sion made upon itself, it is a reflex act, quite independent of the cephalic ganglia, not involving sensation, and taking place through the medium of its OAvn ganglion alone. 855. In the Molluscous classes, generally speaking, the Nervous system bears but a small proportion to the whole mass of the body; and the part of it which ministers to the general movements of the fabric, is often small in proportion to those which serve some special purpose, such as the actions of respiration. This is what we should expect from the general inertness of their character, and from the small amount of muscular structure which they possess. On the other hand, in the Articulated classes, in which the locomotive apparatus is highly developed, and its actions of the most energetic kind, we find the Ner- vous system almost entirely subservient to this function. In its usual form, it consists of a chain of ganglia, connected by a double cord; commencing in the head, and passing backwards through the body (Plate II., Fig. 2). The ganglia, though they usually appear single, are really double; being composed of two equal halves, sometimes closely united on the median line, but occasionally remaining separate, like the cephalic ganglia of the Solen (Fig. 1, a, a), and being united together by a transA'erse commissural trunk. In like manner, the longitudinal cord, though really double (as seen in the upper part of Fig. 2), often appears to be single, in consequence of the close approximation of its lateral halves (as in the lower part of Fig. 2). In general we find a ganglion in each segment; giving off nerves to the muscles of the legs, as in Insects, Centipedes, &c; or to the muscles that move the rings of the body, where no extremities are developed, as in the leech, worm, &c. In the lower Vermiform (or worm-like) tribes, especially in the marine species, the number of segments is frequently very great, amounting even to several hundreds ; and the number of ganglia follows the same proportion. Whatever be their degree of multiplication, they seem but repetitions of one another; the functions of each segment being the same Avith those of the rest. The cephalic ganglia, however, are ahvays larger and more important; they are connected with the organs of 484 OF THE NERVOUS SYSTEM AND ITS ACTIONS. special sense ; and they evidently possess a power of directing and con- trolling the movements of the entire body; whilst the power of each ganglion of the trunk is confined to its OAvn segment.—The longitudinal ganglionic cord of Articulata occupies a position which seems at first sight altogether different from that of the nervous system of Vertebrated animals; being found in the neighborhood of the ventral or inferior surface of their bodies; instead of lying just beneath their dorsal or upper surface. There is reason, however, for regarding the whole of the body of these animals as having an inverted position ; so that they may be considered as really crawling upon their backs. On this view, their longitudinal nervous tract corresponds with the spinal cord of Vertebrata in position, as we shall find that it does in function. 856. We shall draw our chief illustrations of the structure of the nervous system in the Articulated series, from the class of Insects; in which it has been particularly examined. In these animals the number of segments never exceeds twelve (exclusive of the head), either in their larva, pupa, or imago states; and the total number of pairs of ganglia, therefore, never exceeds thirteen, including the cephalic ganglia. These, in the larva, are nearly equal in size, one to another (Plate II., Fig. 2, a, and 1-12); the functions of the different segments of the body being almost uniform ; and the development of the organs of special sense not being such as to involve any considerable predominance in the size of the cephalic ganglia. We observe, at the anterior extremity, the pair of cephalic ganglia (a); from which proceeds, on each side, a cord of communication to the first ganglion (1) of the trunk. This double cord, with the ganglia above and below, thus forms a ring which embraces the oesophagus; the cephalic ganglia being situated on the upper side of it, whilst the ganglionic column of the trunk lies beneath the alimentary canal along its whole length. In the Sphinx ligustri, or Privet Hawk- moth, the nervous system of whose larva is here represented, the last two segments of the body are drawn together as it were, into one; and instead of distinct 11th and 12th ganglia, we find but a single mass nearly double the size of the rest, and obviously formed of the elements that would have otherwise gone to form the two. 857. When the structure of the chain of ganglia is more particularly inquired into, it is found to consist of tAvo distinct tracts ; one of which is composed of nervous fibres only, and passes backwards from the cephalic ganglia, over the surface of all the ganglia of the trunk; whilst the other includes the ganglia themselves. Hence every part of the body has tAvo sets of nervous connexions ; a direct one with the ganglion of its own segment, and an indirect with the cephalic ganglia. Impressions made upon the afferent fibres, which proceed from any part of the body to the cephalic ganglia, become sensations when conveyed to the latter: Avhilst in respondence to these, the influence of sensa- tions received by the cephalic ganglia, and operating through them, harmonizes and directs the general movements of the body, by means of the communicating cords proceeding from them. For the reflex operations on the other hand, the ganglia of the ventral cord are suffi- cient; each one ministering to the actions of its own segment, and, to a certain extent also, to those of other segments. It has been ascer- NERVOUS SYSTEM OF ARTICULATA. 485 tained by the careful dissections of Mr. Newport, that of the fibres con- stituting the roots, by which the nerves are implanted in the ganglia, some pass into the vesicular matter of the ganglion, and, after coming into relation with its vesicular substance, pass out again on the same side (Fig. 152,/, k); whilst a second set, after traversing the vesicular Fig. 152. Portion of the ganglionic tract of Polydesmus maculatus:—6, inter-ganglionic cord; c, anterior nerves; d, posterior nerves; /. lc, fibres of reflex action; g, h, commissural fibres; i, longitudinal fibres, softened and en- larged, as they pass through ganglionic matter. matter, passes out by the trunks proceeding from the opposite side of the same ganglion ; and a third set runs along the portion of the cord which connects the ganglia of different segments, and enters the ner- vous trunks that issue from them, at a distance of one or more ganglia above or below. Thus it appears, that an impression conveyed by an afferent fibre to any ganglion, may excite a motion in the muscles of the same side of its OAvn segment; or in those of the opposite side; or in those of segments at a greater or less distance, according to the point at Avhich the efferent fibres leave the cord. 858. The general conformation of Articulated animals, and the arrangement of the parts of their nervous system, render them pecu- liarly favorable subjects for the study of the reflex actions ; some of the principal phenomena of which will now be described. If the head of a Centipede be cut off, whilst it is in motion, the body will continue to move onAvards by the action of its legs ; and the same will take place in the separate parts, if the body be divided into several distinct por- tions. After these actions have come to an end, they may be excited again, by irritating any part of the nervous centres, or the cut extre- mity of the nervous cord. The body is moved fonvards by the regular and successive action of the legs, as in the natural state; but its move- ments are always fonvards, never backAvards, and are only directed to one side Avhen the fonvard movement is checked by an interposed ob- stacle. Hence, although they might seem to indicate consciousness and a guiding will, they do not so in reality; for they are carried on, as it were, mechanically ; and shoAv no direction of object, no avoidance of danger. If the body be opposed in its progress by an obstacle of not more than half of its own height, it mounts over it, and moves directly onAvards as in its natural state ; but if the obstacle be equal to its own height its progress is arrested, and the cut extremity of the body re- 486 OF THE NERVOUS SYSTEM AND ITS ACTIONS. mains forced up against the opposing substance, the legs still continuing to move.—If, again, the nervous cord of a Centipede be divided in the middle of the trunk, so that the hinder legs are cut off from connexion with the cephalic ganglia, they will continue to move, but not in har- mony with those of the fore part of the body; being completely para- lysed, as far as the animal's controlling power is concerned; though still capable of performing reflex movements, by the influence of their own ganglia, which may thus continue to propel the body, in opposition to the determination of the animal itself.—The case is still more re- markable, when the nervous cord is not merely divided, but a portion of it is entirely removed from the middle of the trunk ; for the anterior legs still remain obedient to the animal's control; the legs of the seg- ments from which the nervous cord has been removed, are altogether motionless; whilst those of the posterior segments continue to act, through the reflex powers of their own ganglia, in a manner which shows that the animal has no power of checking or directing them. 859. The stimulus to the reflex movements of the legs, in the fore- going cases, appears to be given by the contact of the extremities with the solid surface on which they rest. In other cases, the appropriate impression can only be made by the contact of liquid; thus a Dytiscus (a kind of water-beetle) having had its cephalic ganglia removed, re- mained motionless, so long as it rested upon a dry surface; but when cast into water, it executed the usual swimming motions with great energy and rapidity, striking all its comrades to one side by its vio- lence, and persisting in these for more than half an hour. Other movements, again, may be excited through the respiratory surface. Thus, if the head of a Centipede be cut off, and, while it remains at rest, some irritating vapor (such as that of ammonia or muriatic acid) be caused to enter the air-tubes on one side of the trunk, the body will be immediately bent in the opposite direction, so as to withdraw itself as much as possible from the influence of the vapor; if the same irri- tation be then applied on the other side, the reverse movement will take place; and the body may be caused to bend in two or three diffe- rent curves, by bringing the irritating vapor into the neighborhood of different parts of either side. This movement is evidently a reflex one, and serves to withdraw the entrances of the air-tubes from the source of irritation; in the same manner as the acts of coughing and sneezing in the higher animals cause the expulsion, from their air-pas- sages, of solid, liquid, or gaseous irritating matters, which may have found their way into them. 860. From these and similar facts it appears, that the ordinary movements of the legs and wings of Articulated animals are of a reflex nature, and may be effected solely through the ganglia with which these organs are severally connected; whilst in the perfect being, they are harmonized, controlled, and directed by the instinctive guidance, which depends upon sensations acting through the cephalic ganglia and the fibres proceeding from them. There is strong reason to believe, that the operations to which these ganglia are subservient, are almost entirely of a consensual nature; being immediately prompted by sensa- tions, chiefly those of sight, and seldom involving any processes of a REFLEX AND CONSENSUAL ACTIONS OF INSECTS. 487 truly rational character. When we attentively consider the habits of these animals, we find that their actions, though evidently directed to the attainment of certain ends, are very far from being of the same spontaneous nature, or from possessing the same designed adaptation of means to ends, as those performed by ourselves, or by the more in- telligent Vertebrata, under like circumstances. We judge of this by their unvarying character, the different individuals of the same species executing precisely the same movements when the circumstances are the same; and by the very elaborate nature of the mental operations which would be required, in many instances, to arrive at the same results by an effort of reason. Of such we cannot have a more remark- able example, than is to be found in the operations of Bees, Wasps, and other social Insects; which construct habitations for themselves, upon a plan which the most enlightened human intelligence, working according to the most refined geometrical principles, could not surpass; but which yet do so without education communicated by their parents, or progressive attempts of their own, and with no trace of hesitation, confusion, or interruption,—the different individuals of a community all laboring effectively for one common purpose, because their instinctive or consensual impulses are the same. 861. It is interesting to remark that, in the change from the Larva to the perfect or Imago state of the Insect, the Cephalic ganglia undergo a great increase in size. (Plate II., Fig. 3, a, a.) This evidently has reference to the increased development of the organs of special sense in the latter; the eyes being much more perfectly formed; antennae and other appendages used for feeling being evolved; and rudimentary organs of hearing and smell being added. In respondence to the new sensations, which the animal must thus acquire, a great number of new instinctive actions are manifested; indeed it may be said, that the instincts of the perfect Insect have frequently nothing in common with those of the Larva. The latter have reference to the acquirement of food ; the former chiefly relate to the acts of reproduction, and to the provisions^requisite for the de- posit and protection of the eggs and the early nutrition of the young. —We find another important change in the nervous system of the adult or perfect Insect; namely, the concentration of the ganglionic matter of the ventral cord in the thoracic region (e,f); with the three segments of which, the three pairs of legs and the tAvo pairs of wings are connected. The nine segments of the abdomen, in the perfect Insect, give attachment to no organs of motions, and are seldom them- selves very movable; and we find that the ganglia which correspond with them have undergone no increase in size, but have rather dimi- .nished, and have sometimes almost completely disappeared. Where the last segment, however, is furnished with a particularly movable appendage, such as a sting, or an ovipositor, Ave always find a large ganglion in connexion with it. 862. These ganglia of the ventral cord evidently correspond in function with the pedal ganglion of the Mollusca; being so many repetitions of it; in accordance with the number of members. We have noAV to speak of a system of respiratory ganglia, which also are 488 OF THE NERVOUS SYSTEM AND ITS ACTIONS. repeated in like manner, in accordance Avith the condition of the respiratory apparatus ; this being diffused through the Avhole body, in most of the Articulata, instead of being restricted to one spot as in the Mollusca. The system of respiratory nerves consists of a chain of minute ganglia, lying upon the larger cord, and sending off its delicate nerves betAveen those that proceed from the ganglia of the latter, as seen in Fig. 2. These respiratory ganglia and their nerves are best seen in the thoracic portion of the cord, where the cords of communica- tion between the pedal ganglia diverge or separate from one another. And this is particularly the case in the Pupa state, when the whole cord is being shortened, and their divergence is increased. The tho- racic portion of the cord, in the Pupa of the Sphinx ligustri, is shown in Plate II., Fig. 4; where a, b, and c, represent the 2d, 3d, and 4th double ganglia of the ventral cord; d, d, the cords of connexion between them, here Avidely diverging laterally; and e, e, the small respiratory ganglia, which are connected with each other by delicate filaments that pass over the ganglia of the ventral cord, and Avhich send off lateral branches, that are distributed to the air-tubes and other parts of the respiratory apparatus, and communicate with those of the other system. 863. Besides the respiratory system of ganglia and nerves, there is in Insects, as in some Molluscs, a set of minute ganglia, which is espe- cially connected with the acts of mastication and SAvalloAving, its fila- ments being distributed to the muscles of the mouth and pharynx, and some of its ganglia being even found on the stomach, Avhere that organ is remarkable for its muscular powers. The number and arrangement of these ganglia vary considerably in different animals, even in those of the same group: but some traces of this distinct system, which is designated as the stomato-gastric, may always be found. One of the minute ganglia appertaining to it, and forming its anterior termination, is seen to lie on the median line, in front of the great cephalic ganglia, in Plate II., Fig. 3, c. From this a trunk passes backwards along the oesophagus; which may be likened to the oesophageal branches of the Par vagum in Vertebrata. Two other small ganglia communicating with this, are seen at d, d. 864. We are not without traces, moreover, among Invertebrated animals, of the Sympathetic system of the higher classes; though it is quite a mistake to compare the entire system of nerves and ganglia in the former, with the Sympathetic system of the latter, as was formerly done. The chief distribution of the branches of the Sympathetic of Vertebrata is upon the walls of the blood-vessels, and upon the muscular substance of the heart and alimentary canal; and it is by the passage of some of the filaments, from the system of minute ganglia just, pointed out, to the dorsal vessel, that we recognise it as combining the functions of the Sympathetic Avith those of the gastric and cardiac por- tions of the Par vagum. It will be remembered that there is a frequent inosculation between these two nerves, even in the highest animals. 865. Thus we have seen that, in Invertebrated animals, the Nervous System consists of a series of isolated ganglia, connected together by fibrous trunks. The number of these ganglia, and the variety of their NERVOUS SYSTEM OF INVERTEBRATA. 489 function, entirely depend upon the number and variety of the organs to be supplied. In the lowest Mollusca, the regulation of the ingress and egress of water seems almost the only function to be performed; and here we have but a single ganglion. In the Star-fish, we have five or more ganglia; but they are all repetitions one of another, and are ob- viously the centres of action to the several segments to which they respectively belong, neither having a predominance over the rest. And in the higher Mollusca, and in Articulata, we have a ganglion, or more commonly a pair of ganglia, situated at the anterior extremity of the body, connected with the organs of special sensation, and evidently ex- erting a dominant influence over the rest. In the lower Mollusca, we have but a single ganglion for general locomotion; but this is doubled laterally, and repeated longitudinally in the Articulata, in accordance with the multiplication of their locomotive organs, so as to form the ventral cord. In like manner, the Mollusca possess a single ganglionic centre for the respiratory movements; and this is repeated in every segment of the Articulata, forming a chain of respiratory ganglia, which regulates the actions of the extensively-diffused respiratory apparatus of these animals. The acts of mastication and deglutition, again, in both sub-kingdoms, are immediately dependent upon a distinct set of gan- glionic centres; which are connected, however, like the preceding, with the cephalic ganglia. And we have further seen, that, wherever special organs are developed, whose operations depend upon muscular contrac- tion, ganglionic centres are developed in immediate relation with them ; so as to enable them to act by their simple reflex poAver, as well as under the direction of the cephalic ganglia; as in the case of the suckers of the Cuttle-fish.—Now when we inquire into the relation of the cephalic ganglia of Invertebrata with the Brain of the higher Verte- brated animals, we find that these organs cannot be compared in their totality; for that the former are the representatives of a certain portion only, and that usually but a small one, of the latter. The cephalic ganglia of the Centipede, for example, receive nerve-trunks from the eyes, the antennae, and other sensory organs, and give off motor nerves to the different movable parts of the head; and the history of their development, which has been studied by Mr. Newport, shows that they may be considered as the coalesced ganglia of the four segments of which the anterior part of the head is composed ; whilst the first sub- oesophageal ganglia are formed by the coalescence of the four segments entering into the composition of the posterior part of the head. The increased bulk of the cephalic ganglia in the higher Articulata, and especially in the perfect Insect, is obviously for the most part dependent upon the increased development of the visual apparatus, for we find it everyAvhere proportional to it; and thus we may look upon them as mainly optic ganglia, serving to direct the actions of the animal through the sense of sight.—There is no part of these organs which can be con- sidered as superadded to the ganglionic masses which are the immediate centres of the cephalic nerves; consequently there is nothing which can be likened either to the Cerebrum or to the Cerebellum of Verte- brata. And the representative of these cephalic ganglia in the Verte- brated Encephalon, is that series of ganglionic centres at the base of 490 OF THE NERVOUS SYSTEM AND ITS ACTIONS. the brain, which constitute (as we shall presently find) its fundamental portion, and with which all the cephalic nerves are immediately con- nected. 866. When we direct our attention to the nervous system of the Vertebrated series, we perceive that it differs from that of the Inver- tebrated classes we have been considering, in two remarkable features. In these last it has seemed but as a mere appendage to the rest of the system, designed to bring its several parts into more advantageous rela- tion. On the other hand, in the Vertebrata, the Avhole structure ap- pears subservient to it, and designed but to carry its purposes into operation.—Again, in the Invertebrata, we do not find any special adaptation of the organs of support for the protection of the Nervous System ; for it is either enclosed, with the other soft parts of the body, in one general hard tegument, as in the Star-fish and other Echinoder- mata, and in Insects, Crustacea, and other Articulata; or it receives a still more imperfect protection, or even none at all, as in the Mollusca. Noav in the Vertebrata, we find a special and complex bony apparatus, adapted in the most perfect manner for the protection of the Nervous system ; and it is, in fact, the possession of a jointed spinal column, and of its cranial expansion, which best characterizes the group. 867. The Nervous System of Vertebrata is not merely remarkable for its high development, relatively to the remainder of the structure: it is also distinguished by the possession of parts, to which we have nothing analogous in the lower tribes; and by the mode in which these are concentrated and combined, so as to form one continuous mass, instead of consisting of a series of scattered ganglia.—The chief parts which are newly introduced (so to speak) in this sub-kingdom, are the Cerebral Hemispheres and Cerebellum ; of Avhich there are no traces whatever in the loAver Articulata and Mollusca, and but very uncertain representations in the highest. These are superimposed, as it were, upon the cephalic ganglia connected with the organs of special sense, and upon the cords that connect them with the first ganglion of the trunk.—Again, we find that the locomotive ganglia, which formed the long knotted cord of the Articulata, are united with the centres of the respiratory system, and with those of the stomato-gastric system, to form one continuous tract, which commences anteriorly from the gan- glia of special sense, and runs backwards* without interruption, in the canal of the Vertebral column, forming the spinal cord. This is a con- tinuous instead of an interrupted ganglionic mass; it is composed of two lateral halves, precisely similar to each other; and each of these consists of two parts, as distinct from each other as the two tracts in the ventral cord of the Articulata,—namely, a fibrous structure, which connects every part of it with the Encephalon (or collection of nervous masses within the cranium), and Avhich also serves to connect together the different parts of the cord itself,—and a vesicular portion, which forms the proper centre of the greater part, if not the whole, of the fibres entering into the roots of those nerves. The anterior portion of the Spinal cord, which is prolonged into the cranium, and comes into * When we speak of the Vertebrata generally, their bodies are of course supposed to be in a horizontal position,—not vertical, as in Man. NERVOUS SYSTEM OF VERTEBRATA. 491 immediate relation with the Encephalon, is termed the Medulla Oblon- gata. It is in this, that the centres of the respiratory and stomato- gastric nerves are found; the situation of these important ganglia within the cranium, being obviously destined to protect them from those inju- ■ ries, to Avhich the Spinal Cord itself is liable. 868. Thus, then, Ave are led to recognise in the Nervous system of .Vertebrata the following fundamental parts.—1. A system of ganglia subservient to the reflex actions of the organs of locomotion, and cor- responding with the chain of pedal or locomotive ganglia that makes up the chief part of the ventral cord of the Articulata; in this system, the gray or vesicular matter forms one continuous tract, which occupies the interior of the Spinal Cord.—2. A ganglionic centre for the move- ments of respiration, and another for those of mastication and degluti- tion ; these, with part of the preceding, make up the proper substance of the Medulla Oblongata.—3. A series of ganglia, in immediate con- nexion with the organs of Special Sense; these are situated within the cranium, at the anterior extremity of the Medulla Oblongata; and, in the lowest Vertebrata, they constitute by far the largest portion of the entire Encephalon.—4. The Cerebellum, Avhich is a sort of off-shoot from the upper extremity of the Medulla Oblongata, lying behind the preceding.—5. The Cerebral Hemispheres, a pair of ganglionic masses, which lie upon the ganglia of special sense, capping them over more or less completely, according to their relative development.—These last two organs exist in the lowest Vertebrata, as in Invertebrated animals generally, in quite a rudimentary state; but their development, rela- tively to other parts of the Encephalon, and to the entire bulk of the animal, increases as Ave ascend the scale; so that in Man and the higher Mammalia they constitute by far the largest portion of the Nervous centres, and are essential to the greater part of the operations of the Nervous system. The development of the Cerebral Hemispheres holds a close relation with the increase of the Lntelligence, and with the pre- dominance of the Will over the involuntary impulses. The increased size of the Cerebellum, on the other hand, seems connected with the necessity which exists, for the adjustment and combination of the loco- motive powers, when the variety in the movements performed by the animal is great, and a more perfect harmony is required among them. —A sketch of the mode in which these different parts are combined and arranged in the several classes of Vertebrata, and of their relative development in each, will aid us in the subsequent more detailed exami- nation of their functions. 869. In the class of Fishes, taken as a whole, the Encephalon bears a much smaller proportion to the Spinal Cord, than in the higher Ver- tebrata. In the curious Amphioxus, or Lancelot, there is no discover- able nenous mass anterior to the Medulla Oblongata; and wre have here, therefore, an animal regularly formed upon the plan Avhich occasionally presents itself as a monstrosity in Man,—namely, having the Spinal Cord and Medulla Oblongata for the Avhole of the nervous centres, and being anencephalous, or destitute of any proper encephalon. In some of the lowest Vermiform (Avorm-like) Fishes, such as the Lamprey, the cephalic masses are very little more developed in proportion to the 492 OF THE NERVOUS SYSTEM AND ITS ACTIONS. Spinal Cord, than are the cephalic ganglia of Insects in reference to their chain of ventral ganglia. But as the organs of special sense ac- quire a more complete evolution, we find the ganglia connected with them presenting a greatly-increased size. On opening the cranial caA'ity of a Fish, we usually observe four nervous masses (three of them in pairs) lying, one in front of the other, nearly in the same line with the Spinal cord. The first or most anterior of these are the Olfactoryt ganglia (Plate II., Figs. 5, 6, 7, a), or the ganglia of the nerves of smell; the nature of which is knoAvn, from their being situated at the origin of the Olfactory nerves. In the shark and some other Fishes, these are separated from the rest by peduncles or foot-stalks; a fact of much interest, as explaining the arrangement which we find in Man. What is commonly termed the trunk of his Olfactive nerve is really the commissure connecting the OlfactiA'e ganglion (known as the bulbous enlargement that lies upon the cribriform plate of the Ethmoid bone) with the other portions of his Encephalon: the proper fibres of the nerve being those which come off from this ganglion, in the numerous branches that proceed from it into the nasal cavity.—Behind the Olfac- tive ganglia is a pair of masses, b, b, of which the relative size varies greatly in different Fishes. Thus, in the Perch, whose Encephalon is here figured, their size is intermediate between that of the first and third pairs; being as much inferior to that of the third, as it is superior to that of the first. On the other hand, in the Shark and several other Fishes, they' are considerably larger than the succeeding pair. These second ganglia are commonly considered as the rudiments of the Cere- bral Hemispheres; but there seems reason for regarding them as being chiefly the representatives of the Corpora Striata; the existence of a cerebrum being only indicated by a thin layer of vesicular matter, which overlies the ventricle that is found in these bodies in the brains of Cartilaginous Fishes alone.—Behind them, and forming the third pair of ganglionic masses, c, c, are two large bodies, from Avhich the optic nerves arise; these evidently represent the Optic ganglia, which constitute the principal mass of the cephalic ganglia in Insects and the higher Mollusca, and with Avhich the Corpora Quadrigemina of higher Vertebrata partly correspond ; but they probably represent also the Thalami Optici of the brain of Man and the higher animals.—At the back of these, overlying the top of the spinal cord, is a single mass, d, the Cerebellum. This, also, varies greatly in its relative dimensions, being much more highly developed in the active and rapacious Sharks, than it is in Fishes of inferior muscular energy and variety of move- ment.—The Spinal Cord e, is divided at the top by a fissure, which is most wide and deep beneath the cerebellum, where there is a complete separation between its two halves. This opening corresponds to that, through which the oesophagus passes in the Invertebrata; but as the entire nervous mass of Vertebrated animals lies above the alimentary canal (or nearer the dorsal surface), it does not serve the same purpose in them; and in the higher classes, the fissure is almost entirely closed by the union of the tAvo halves on the median plane, the fourth ven- tricle, hoAvever, being a remnant of it. This cavity is partly seen in Fig. 7, Avhich is a vertical section of the brain whose upper and under NERVOUS CENTRES OF REPTILES. 493 surfaces are shown in Figs. 5 and 6.—In the lateral strands of the Medulla Oblongata, close to the fourth ventricle, there is a pair of gan- glionic centres (characterized by the presence of vesicular matter), in which the auditory nerve terminates ; and these are sometimes deA'eloped as distinct ganglionic enlargements. Other separate ganglia, sometimes of considerable size, are very commonly found at the origin of the Par Vagum.—It is curious to notice the very large comparative size of the Pineal gland (/), and of the Pituitary body (h), in this class; the func- tions of these organs are entirely unknown. ^ 870. The analogy of the Optic lobes of Fishes to the Corpora Quad- rigemina and Thalami Optici of the fully-formed brain of the higher Vertebrata, is not so complete as it is to certain parts which occupy their place at an earlier period. In the Human Embryo, at about the 6th , week, the Encephalon consists of a series of vesicles arranged in a line Fig. 153. Human embryo of sixth week, enlarged about three times:— a, vesicle of corpora quadrigemina; 6, vesi- cle of cerebral hemispheres; c, vesicle of thalami optici and third ventricle; d, vesicle for cerebellum and medulla oblongata; e, auditory vesicle; f, olfactory fossa; /(, liver; ** caudal extremity. with each other ; of which those that represent the cerebrum (b, Fig. 153) are the smallest, whilst that which represents the cerebellum (d) is the largest. BetAveen the cerebral and the cerebellic vesicles are two others (c, and a), of which the posterior one is the Optic ganglion, and answers to the Tubercula Quadrigemina; whilst the anterior contains the Third Ventricle, and corresponds in some degree to the Thalami Optici. This condition is precisely represented in the Lamprey; but in most Fishes, the optic ganglia, and the parts surrounding the third ventricle, form but one lobe ; so that the third ventricle seems hollowed out of the cptic ganglia, as shown in Fig. 7, c, (Plate II.) 871. The Encephalon of Reptiles does not show any considerable advance in its general structure, above that of the higher Fishes. The Cerebral Hemispheres (Plate II. Figs. 8, 9, 10, b), are always much larger than the Olfactive and Optic ganglia; and they generally cover- in the latter (c, c) in part, by their posterior extremities. The Cere- bellum is almost invariably of small proportionate dimensions; and this is especially the case in the Frog, in which it does not even cover-in the fourth ventricle. This low development of the Cerebellum in Reptiles, is what might be anticipated from the general inertness of those ani- mals, and the Avant of variety in their movements. The Spinal Cord is 494 * OF THE NERVOUS SYSTEM AND ITS ACTIONS. still very large, in proportion to the nervous masses contained in the skull; and, as we shall hereafter see, its poAver of keeping up the move- ments of the body, after it has been cut off from all connexion, with the brain, is very considerable.—We find that, in Reptiles, as in Fishes, the Spinal Cord may have a nearly uniform size from one extremity to the other, like the ventral cord of the lower Articulata; or it may present considerable enlargements at particular spots, like the ganglionic cord in the thoracic regions of Insects. This difference depends upon the degree of development of the special locomotive organs. Thus in the Eel and Serpent, whose movements are accomplished by the undulations of the entire trunk, and which are destitute of members, we find a uni- form development of ganglionic matter in the spinal cord. On the other hand, in the Flying-fish, in which the pectoral fins or anterior extremi- ties effect the greater part of the propulsion of the body, we find a great ganglionic enlargement of the Spinal cord, at the part in which the nerves of those members are connected: in the Frog, whose movements are chiefly effected by the posterior extremities, Ave find a similar enlarge- ment at the roots of the crural nerves : and in the Turtles and Lizards, the two pairs of whose members are nearly equal in function, and serve to effect the principal movements of the body, we find an anterior and posterior enlargement of the Spinal Cord, corresponding to the parts with which the nenes of these members are connected. 872. We find in Birds a considerable advance in the character of the Encephalon, towards that which it presents in Mammalia. The Cere- bral Hemispheres (Plate II., Figs. 11, 12, 13, b) are greatly increased in size; and they cover-in, not merely the olfactory ganglia, but in great part also, the optic ganglia. The former are of comparatively small size; the organ of smell in Birds not being much developed. The latter are very large, in conformity with the acuteness of sight which is highly characteristic of the class. The cerebellum is of large size, as we should expect from the number and complexity of the muscular movements performed by animals of this class; but it is still undivided into hemispheres. The Spinal Cord is still of considerable size in com- parison Avith the Encephalon; and it is much enlarged at the points whence the legs and wings originate. In the species which have the most energetic flight, such as the Swallow, the enlargement is the greatest where the nerves of the wings come off; but in those which, like the Ostrich, move principally by running on the ground, the poste- rior enlargement, from which the legs are supplied with nerves, is much the more considerable. 873. In the Mammalia we find the size and general development of the Encephalon presenting a gradual increase, as we ascend the series, from the non-placental Monotremes and Marsupials, towards Man. In the former, the Hemispheres exhibit no convolutions; and the great transverse commissure, or connecting band of fibrous structure, termed the corpus callosum, is deficient. As we rise through the true viviparous division of the class, we notice a gradually-increasing prolongation of the Cerebral Hemispheres backwards; so that first the optic ganglia, and then the cerebellum, are covered-in by them. The latter partly shows itself, however, in all but Man and the Quadrumana, when we NERVOUS CENTRES OF MAMMALIA—SPINAL CORD. 495 look at the brain from above downwards: as we see in the Encephalon of the Sheep (Plate II., Figs. 14, 15, d). The Cerebral hemispheres increase, not only in size ; but also in complexity of structure, both external and internal. Their exterior, instead of remaining smooth, is marked by convolutions ; which serve to extend very greatly the amount of surface over which blood-vessels can pass into the gray substance. Their internal structure becomes more complex, in the same proportion as their size and the depth of their convolutions increase; and in Man all these conditions present themselves in a far higher degree than in any other animal. ^ The number of commissural bands, connecting the two hemispheres with each other transversely, and uniting their anterior and posterior portions, is very greatly increased ; and in fact, a large proportion of their mass is composed, in Man, and the higher Mammalia, of fibres of this character.—In proportion to the increase of the Cere- bral hemispheres, there is a relative diminution in the size of the ganglia of special sense ; but their dimensions, as compared with the entire bulk of the animal, are by no means reduced, but are even increased. The Olfactive ganglia (Fig. 14, a) are always readily discoverable; being separated from the remainder of the encephalic masses by a peduncle on each side. The Optic ganglia (Fig. 15, c), on the other hand, are so completely covered-in by the Hemispheres, that it is only when the latter are turned aside that we can discern them. They differ in exter- nal aspect from the optic ganglia of Birds and the lower Vertebrata; being divided by a transverse furrow into anterior and posterior emi- nences, whence they are known as the Corpora Quadrigemina. The Auditory ganglia are lodged in the substance of the Medulla Oblongata, forming the ''gray nuclei" of the strands termed the "posterior pyra-, mids;" and similar nuclei in the "restiform bodies" are the ganglionic centres of the Glosso-pharyngeal nerves, and perhaps minister to the sense of Taste. Besides these, however, are the tAvo large bodies termed the Corpora Striata and Thalami Optici, which have been commonly considered as appendages of the Cerebrum, but which must undoubtedly be regarded as independent of it, and as themselves constituting gan- glionic centres, whose development bears no constant proportion to that of the Cerebrum. From the peculiar relation presently to be described (§ 901), Avhich these bodies bear on the one hand to the Spinal Cord, and on the other to the rest of the Encephalon, there seems strong reason to believe that they together constitute the ganglionic centre of the sense of Touch, and of the motions which are automatically prompted by it.—The Cerebellum is chiefly remarkable for the development of its lateral parts or hemispheres, and for the intricate arrangement of the gray and Avhite matter in them (Fig. 15, d); the central portion, some- times called the vermiform process, is relatively less developed than in the lower Vertebrata, where it forms the entire organ.—The Spinal Cord is much reduced in size, when compared with other parts of the nervous centres; the motions of the animals of this class being more dependent upon their will or guided by their sensations ; and the simply reflex actions bearing a much smaller proportion to the rest. The development of ganglionic enlargements, in accordance with the presence or absence of high locomotive powers in the extremities, follows the same rule as in the preceding classes. 496 OF THE NERVOUS SYSTEM AND ITS ACTIONS. 3. Functions of the Spinal Cord and its Nerves. 874. In commencing our more detailed examination into the func- tions of the different parts of the Nervous system in Vertebrated ani- mals, it seems best to commence with the Spinal Cord ; this being the portion whose presence is most essential to the continuance of life. As already mentioned, Infants are sometimes born Avithout any Cerebrum or Cerebellum ; and such have existed for several hours or even days, breathing, crying, sucking, and performing various other movements. The Cerebrum and Cerebellum have been experimentally removed from Birds and young Mammalia, thus reducing these beings to a similar condition ; and all their vital operations have, nevertheless, been so regularly performed, as to enable them to live for weeks, or even months. In the Amphioxus, as already remarked, we have an example of a completely-formed adult animal, in which no rudiment of a Cere- brum or Cerebellum can be detected. And in ordinary profound sleep, or in apoplexy, the functions of these organs are so completely sus- pended, that the animal is, in all essential particulars, in the same con-' dition for a time as if destitute of them. It is possible, indeed, to re- duce a vertebrated animal to the condition (so far as its nervous system is concerned) of an Ascidian Mollusc (§ 850) ; for it may continue to exist for some time, when not merely the Cerebrum and Cerebellum haA'e been removed from above, but when nearly the whole Spinal Cord has been removed from below,—that part only of the latter being left, Avhich is the centre of the respiratory actions, and which corresponds to the single ganglion of the Tunicata. On the other hand, no animal can exist by its Encephalon alone, the Spinal Cord being destroyed or re- moved ; for the reflex actions of the latter are so essential to the con- tinuance of its respiration, and consequently of its circulation, that if they be suspended (by the destruction of the portion of the cord Avhich is concerned in them), all the organic functions must soon cease. 875. Although the Spinal Cord was formerly regarded as little else than a bundle of nerves proceeding from the Brain, yet its true rank, as a distinct centre of nervous poAver, is noAV universally admitted. That the actions prompted by it, when these do not originate in one of the higher centres, are of a purely reflex nature,—consisting in the excitement of muscular movements in respondence to external impres- sions, Avithout the necessary intervention of sensation,—appears to be a necessary inference from the facts that have been brought to light by experiment and observation. Experiments on the nature of this func- tion are best made upon cold-blooded animals ; as their general functions are less disturbed by the effects of severe injuries of the nervous sys- tem, than are those of Birds and Mammals. When the Cerebrum has been removed, or its functions have been suspended by a severe blow upon the head, a variety of motions may be excited by their appropriate stimuli. Thus, if the edge of the eyelid be touched with a straw, the lid immediately closes. If a candle be brought near the eye, the pupil contracts. If liquid be poured into the mouth, or a solid substance be pushed within the grasp of the muscles of deglutition, it is swallowed. REFLEX FUNCTION OF THE SPINAL CORD. 497 If the foot be pinched, or burned with a lighted taper, it is withdrawn; and (if the animal experimented on be a Frog) the animal will leap away, as if to escape from the source of irritation. If the cloaca be irritated with a probe, the hind-legs will endeavor to push it away. 876. Now the performance of these, as well as of other movements, many of them most remarkably adapted to an evident purpose, might be supposed to indicate, that sensations are called up by the impres- sions ; and that the animal can not only feel, but can voluntarily direct its movements, so as to get rid of the irritation which annoys it. But such an inference Avould be inconsistent with other facts.—In the first place, the motions performed by an animal under such circumstances are never spontaneous, but are ahvays excited by a stimulus of some kind. Thus, a decapitated Frog, after the first violent convulsive movements occasioned by the operation have passed away, remains at rest until it is touched : and then the leg, or its whole body may be thrown into sudden action, Avhich immediately subsides again. In the same manner, the act of swallowing is not performed, except when it is excited by the contact of food or liquid ; and even the respiratory movements, spontaneous as they seem to be, would not continue, unless they were continually re-excited by the presence of venous blood in the vessels. These movements are necessarily linked with the stimulus that excites them ; that is, the same stimulus will always produce the same movement, when the condition of the body is the same. Hence it is evident, that the judgment and will are not concerned in pro- ducing them ; and that the adaptiveness of the movements is no proof of the existence of consciousness and discrimination in the being that executes them,—the adaptation being made for the being, by the pecu- liar structure of its nervous apparatus', which causes a certain move- ment to be executed in respondence to a given impression,—not by it. An animal thus circumstanced may be not unaptly compared to an au- tomaton ; in which particular movements, adapted to produce a given effect, are produced by touching certain springs. Here the adaptation was in the mind of the maker or designer of the automaton ; and so it evidently is, in regard to the reflex or consensual movements of animals, as well as with respect to the various operations of their nutritive sys- tem, over Avhich they have no control, yet Avhich concur most admira- bly to a common end. 877. Again, we find that such movements may be performed, not only Avhen the Brain has been removed, the Spinal cord remaining en- tire, but also when the Spinal cord has been itself cut across, so as to be divided into two or more portions, each of them completely isolated from each other, and from other parts of the nervous centres. Thus, if the head of a Frog be cut off, and its spinal cord be divided in the middle of the back, so that its fore-legs remain connected with the upper part, and its hind-legs with the loAver, each pair of members may be excited to movement by a stimulus applied to itself; but the two pairs will not exhibit any consentaneous motions, as they will do when the spinal cord is undivided. Or, if the Spinal cord be cut across, without the removal of the Brain, the lower limbs may be excited to move- ment, by an appropriate stimulus, though they are completely paralysed 498 OF THE NERVOUS SYSTEM AND ITS ACTIONS. to the will ; whilst the upper remains under the control of the animal, as completely as before. Now it is not conceivable that, in this last case, sensation and volition should exist in that portion of the spinal cord, which remains connected with the nerves of the posterior extremi- ties, but which is cut off from the brain. For if it were so, there must be two distinct centres in the same animals, the attributes of the brain not being affected ; and, by dividing the spinal cord into two or more segments, Ave might thus create in the body of one animal two or more distinct centres of sensation, independent of that which still holds its proper place in the Encephalon. To say that tAvo or more distinct centres of sensation are present in such a case, would really be in effect the same as saying, that there are two or more distinct minds in one body,—which is manifestly absurd. 878. But the best proofs of the limitation of the endowments of the Spinal Cord, are derived from the phenomena presented by the Human subject, in cases where that organ has suffered injury, by disease or accident, in the middle of the back. We find that, when this injury has been severe enough to produce the effect of a complete division of the Cord, there is not only a total want of voluntary control over the lower extremities, but a complete absence of sensation also,—the individual not being in the least conscious of any impression made upon them. When the lower segment of the Cord remains sound, and its nervous connexions with the limbs are unimpaired, distinct reflex movements may be excited in the limbs, by stimuli directly applied to them, as, for instance, by pinching the skin, tickling the sole of the foot, or applying a hot plate to its surface ; and this without the least sensation, on the part of the patient, either of the cause of the moA7ement, or of the move- ment itself. This fact, taken in connexion with the preceding experi- ments, both upon Vertebrated and Articulated animals, distinctly proves that Sensation is not a necessary link in the chain of reflex actions; but that all which is required is an afferent fibre, capable of receiving the impression made upon the surface, and of conveying it to the centre; a ganglionic centre, composed of vesicular nervous substance, into which the afferent fibre passes; and an efferent fibre, capable of transmitting the motor impulse, from the ganglionic centre, to the muscle which is to be thrown into contraction. 879. These conditions are realized in the Spinal Cord. We may have reflex actions excited through any one isolated segment of it, as through a single ganglion of the ventral cord of Articulata ; but they are then confined to the part supplied by the nerves of that segment. Thus, if the spinal cord of a Frog be divided just above the origin of the crural nerves, the hind-legs may be throAvn into reflex contraction by various stimuli applied to themselves ; but the fore-legs will exhibit no movement of this kind. But Avhen the brain has been removed, and the Spinal Cord is left entire, movements may be excited in distant parts, as, for example, in the fore-legs, by any powerful irritation of the posterior extremities, and vice versd. This is particularly well seen in the convulsive movements, Avhich take place in certain disordered states of the nervous system; a slight local irritation being sufficient to throAV almost any muscle of the body into a state of energetic action (§ 885). REFLEX FUNCTION OF THE SPINAL CORD. 499 And a similar state may be artificially induced, by applying Strychnine (in solution) to the Spinal Cord of a decapitated Frog. ^ 880. The minute anatomy of the Spinal Cord is a subject of great difficulty; and our notions of the course of the fibres within it are rather founded upon physiological phenomena, and upon the more evident structure of the ventral column in Articulata, than upon what can be clearly demonstrated in Vertebrated animals. The roots of the Spinal nerves are all distinctly separable into an anterior and a posterior fasci- culus ; and it is certain that these fasciculi have entirely opposite func- tions. If they be laid bare, and the anterior fasciculus of any spinal nerve be touched, violent contractions are immediately seen in the mus- cles supplied by that nerve; these contractions are as strongly manifested if the anterior roots be divided, and their separated end be irritated; whilst no such result follows, whatever amount of irritation be applied to the ends still in connexion with the cord. Notwithstanding these violent movements, the animal shows little or no sign of pain. On the other hand, if the posterior roots be irritated, the animal gives signs of acute pain, and no vigorous muscular contractions are produced. The move- ments Avhich are Avitnessed are evidently of a reflex nature, being called forth through the anterior roots; as is proved by their cessation when these are divided. Further, if the posterior roots be divided, and the separated ends be irritated, no effect Avhatever is produced; no move- ment is excited; and no sensation is occasioned; but if the ends still in connexion with the cord be irritated, the animal shows signs of pain as before.—Hence it is evident, that the posterior roots are made up of afferent fibres, that is, of the fibres Avhich convey impression towards the nervous centres; which impressions, if confined to the cord itself, excite reflex actions; whilst, if conveyed to the brain, they produce sen- sations. On the other hand it is equally evident, that the anterior roots are composed of efferent or motor fibres, Avhich serve to comrey to the muscles the motor impulses originating in the nervous centres; these impulses may be occasioned by the reflex action of the Spinal cord ; or they may descend from the Brain, wThere they have been generated by a Consensual or Emotional impulse, or by an act of the Will. 881. The Spinal Cord is a completely double tract; being composed of two distinct halves, united together on the median plane by nume- rous commissural fibres. This union is much closer in Man and the Mammalia, than it is in the lower Vertebrata; but the division is still marked externally, by a deep fissure on the anterior surface of the cord, and by a shalloAver one on its posterior aspect. Its surface is traversed, moreover, by two furrows on each side; so that each half is divided into three columns, the anterior, lateral, and posterior. The anterior roots of the spinal nerves join the Cord for the most part along the line of the anterior furrow; and the posterior along the line of the posterior furrow; so that the middle or lateral column lies between them, the anterior column being altogether in front of them, and the posterior column behind them. When a transverse section of the Cord is made, it is seen to contain, on each side, a crescentic patch of gray or vesicu- lar substance; the points of each crescent are directed towards the anterior and posterior furrows of its own side respectively; whilst the convexities of the two crescents approach one another near the median 500 OF THE NERVOUS SYSTEM AND ITS ACTION?. plane, and are connected by a transverse tract of gray matter. The remainder of the cord is made up of white or tubular substance, the course of whose fibres is for the most part longitudinal. The posterior peak of the crescentic patch of gray matter approaches very closely to the bottom of the posterior furrow; Avhilst the anterior peak does not come into nearly the same degree of proximity with the bottom of the anterior furroAv. Hence, it is considered by some, that the lateral or middle columns of the cord, being much less completely isolated from the anterior columns than they are from the posterior, should be associated with the former, under the name of antero-lateral columns. 882. Upon tracing the roots of the nerves into the substance of the Cord, the connexion of a part of their fibres with its gray or vesicular substance is easily made evident. Of these fibres, therefore, it serves as the proper ganglionic centre. There is reason to believe, both from anatomical investigation, and from physiological phenomena, that, as in the Articulata (§ 857), a part of the afferent or excitor fibres, after traversing the gray substance, pass out on the same side as the efferent or motor; whilst another portion crosses to the opposite side, and forms part of its efferent trunks.—It is pretty certain that other fibres of the roots become continuous with the longitudinal fibres that form the white strands of the Spinal Cord; but it is by no means certain, on that account, that they pass on to the Brain ; and, in fact, there is adequate evidence that if any of the fibres thus establish a direct communication between the Encephalic centres and the spinal nerve-trunks, their pro- portion must be very small, the chief part of the longitudinal strands of the Cord being apparently made up of commissural fibres which esta- blish an intimate connexion betAveen its different segments, as in Insects. This will appear from the facts to be next stated. The thickness of the Spinal Cord differs considerably at its different parts. Thus in the cervical region, there is an enlargement corresponding with the origins of the nerves that form the brachial plexus; this enlargement is partly caused by an increase in the amount of gray matter; but the amount of fibrous structure also, is much greater than at the upper part of the cervical region. On the other hand, there is a still greater enlarge- ment of the cord in the lumbar region, at the part Avhence the nerves of the loAver extremities arise ; and this enlargement is caused by the great increase in the amount both of the gray matter and of the white at that point. It may be easily shoAvn by direct measurement, that the fibrous strands of the upper cervical region would not by any means serve to carry omvard to the brain those of the lumbar region alone, much less with the addition of other fibres proceeding from all the inter- mediate nerves. Further, if the fibrous strands Avere for the most part (as formerly supposed) directly continuous betAveen the brain and the roots of the spinal nerves, the white portion of the Spinal Cord, in such animals as Serpents, in which it has no ganglionic enlargements, should progressively diminish in diameter with every pair of nerves into which it sends fibres, from its cephalic to its caudal extremity; this, however, is by no means the case, the Spinal cord of Serpents being remarkable for its uniform diameter throughout.—It is obvious, then, that if any of the longitudinal fibres of the cord should thus pass REFLEX FUNCTION OF THE SPINAL CORD. 501 direct from the Encephalon to the nerve-roots, their proportion must be very small. And Ave shall see that all the phenomena which have been supposed to indicate an immediate communication between the brain and the nerves, admit of another and more satisfactory explanation. 883. It was supposed by Sir C. Bell (who was the first to determine the relative functions of the two roots of the spinal nerves in Verte- brated animals), that the anterior columns of the spinal cord have a function corresponding to that of the anterior roots of the spinal nerves; and the posterior columns with the posterior roots. But from the diffi- culty of tracing the connexion between the longitudinal fibres of the cord and any portion of the roots, it is at present impossible to say hoAv far there is any anatomical reason for the assumption of this cor- respondence; and it is quite certain that the physiological facts at present known, from observation of the effects of disease or injury upon different tracts of the spinal cord, do not bear out the supposition. As to what the precise functions of the several columns are, however, it is not easy to form any other conjecture, that shall be consistent with all the phenomena at present knoAvn. 884. Of the particular Reflex actions to which the Spinal Cord (using that term in its limited sense, as excluding the Medulla Oblongata) is subservient, those most connected with the organic functions have already been noticed. They are chiefly of an expulsive kind; being destined to force out the contents of various cavities of the body. Thus the ordinary acts of defecation and urination, the ejaculatio seminis and parturition, are all reflex actions, over Avhich the will has a greater or less degree of control; being able to keep the two former ones in check, so long as the stimulus is not very violent, and being also capable of effecting them by itself; but having no control over the two latter, either by way of acceleration or prevention, when once the stimulus by which they are excited has come into full action.—The movements of the posterior extremities are among the most remarkable of those, Avhich seem due to the action of the proper Spinal Cord. It has been already noticed, that these may be excited, eATen in Man, when the spinal cord has been severed in the middle Avithout injury to its lower segment; and it is remarkable, that gentle stimuli, applied to the skin of the sole Qf the foot, appear the most capable of producing them. We haAre seen Iioav completely, in the lower animals, the acts of progression may be sustained, by the repeated stimulus of the contact of the ground, or of fluid, without any influence from the cephalic ganglia; the power of these being limited, it would seem, to the control and direction of them. And there is strong reason to believe that so far as the ordinary acts of locomotion are concerned, the movements of the inferior extremities in Man may be performed on the same plan,'being continued by the reflex influence of the successive impulses of 'the feet upqn the ground, when once set in action by the will, whilst Ave are Avalking steadily on- Avards,—the mind b^ing at the same time occupied by some train of thought, which engrosses its whole attention. Theirs are< fcAv persons, to Avhom it has not occasionally happened that, on awakingt (as it were) from their revery, they have found themselves in a place very different from that to which they had intended going; and even Avhen tYe Ton- 502 OF THE NERVOUS SYSTEM AND ITS ACTIONS. sciousness is sufficiently on the alert to allow sensations to guide, direct, and control the motions of the limbs, their actions appear to be per- formed without the agency of the will, which may be entirely concen- trated upon some interior mental operation. It is certain that, in Birds, the movements of flight may be performed after the removal of the Cerebrum. 885. There are many irregular or abnormal reflex actions, performed ■'it through the instrumentality of the Spinal Cord, the study of which is of the highest importance to the Medical Man. It is probable that all Con- vulsive movements are produced through its agency and that of the Medulla Oblongata; for it has been found, by repeated experiments, that these movements are never produced by injuries of the Cerebral hemispheres.—Convulsive movements may be of three kinds. 1. They may be simply reflex ; being the natural result of some extraordinary irritation. 2. They may be simply centric ; depending upon a peculiar condition of the ganglionic centre of the Spinal Cord, which occasions muscular movements without any stimulation. This may be dependent upon an abnormal state of the Blood. We know that it may be pro- duced by the introduction of certain poisons (as strychnia) into the cir- culation; and it is probable that morbid matters generated within the body may have the same effect. 3. They may depend upon the com- bined action of both principles; the nervous centres being in a very irritable state, Avhich causes very slight irritations (such as would other- wise be inoperative) to excite violent reflex or convulsive movements. This last is by far the most common cause of the convulsive actions, that occur in various diseased conditions of the system. Thus, convul- sions are not unfrequent in children, during the period of teething; being produced by the irritation, which results from the pressure of the tooth, as it rises against the unyielding gum. In this case, the stimulus would scarcely be sufficient to produce the violent result, were it not for a peculiarly excitable state of the Spinal Cord, brought about by various causes. In like manner, when such an excitable state exists, convulsions may be occasioned by the presence of intestinal worms, of irritating substances, or even simply of undigested matters, in the ali- mentary canal; and will cease as soon as they are cleared out, in the same manner as the convulsions of teething may often be at once checked, by the free lancing of the gums. 886. The influence of the condition of the Spinal Cord itself, is peculiarly seen in the convulsive diseases termed Hydrophobia, Teta- nus, Epilepsy, and Hysteria.—In the first of these, not only the Spinal Cord, but the Medulla Oblongata, and the ganglia of Special Sense, are involved; their peculiar condition being the result, it would appear, of the introduction of a poison into the blood. It is most remarkable that the Cerebrum should so completely escape its influence. When the state of intense excitability in these centres, is once established, the slighest stimulus is sufficient to bring about convulsive movements of the utmost violence. It is characteristic of this complaint, that the stimuli most effectual in exciting the movements, are those Avhich act through the nerves of special sense; thus the sight or the sound of water AV'ill bring on the paroxysm; and any attempt to taste it increases HYDROPHOBIA ; TETANUS ; EPILEPSY ; HYSTERIA. 503 the severity of the convulsions.—In tetanus there appears to be a simi- larly excitable state of the Spinal Cord and Medulla Oblongata, not involving the ganglia of special sense. This may be the result of causes altogether internal, as in the idiopathic form of the disease; in Avhich the condition exactly resembles that, which may be artificially induced by the administration of Strychnine, or by its application to the cord. Or it may be first occasioned by some local irritation, as that of a lacerated wound; the irritation of the injured nerve being propagated to the nervous centres, and establishing the excitable state in them. When the complaint has once established itself, the removal of the original cause of irritation (as by the amputation of the injured' limb) is seldom of any avail; since the slightest impressions upon al- most any part of the body, are sufficient to excite the tetanic spasm.— In like manner, Epilepsy, which consists in convulsive actions with temporary suspension of the functions of the Encephalon, may result from the irritation of local causes, like the convulsions of teething; and may, like them, cease when-the sources of irritation are removed. But when it becomes confirmed, it seems to involve a disorder of the nervous centres, which no local treatment can influence. 887. These and other forms of Convulsive disorder, when productive of a fatal result, usually act by suspending the respiratory movements; the muscles which effect these being fixed by the spasms, so that the air cannot pass either in or out, and suffocation takes place as completely as if the entrance to the air-passages were closed. It is remarkable that every one of them may be imitated by Hysteria ; a state of the nervous system, in which there is a peculiar excitability, but in Avhich there is no such fixed tendency to irregular action, as Avould indicate any positive disease,—one form of convulsion often taking the place of another, at short intervals, with the most wonderful variety. It Avill often be found, that the convulsions may be immediately traced to some local irritation ; thus they are particularly liable to occur at the catamenial periods, especially if the menstrual flux be deficient; but it does not seem improbable, that here too the presence of morbid matters in the blood has much to do with the development of that peculiar excitability, Avhich gives to slight local irritations such a powerful agency. 888. The statement that the Spinal Nerves arise by double roots, is not without exception as regards some, Avhich arise from its cranial prolongation, and which are distributed to the parts of the head and neck. The first spinal nerve, or sub-occipital (the 10th pair of Willis) not unfrequently arises by a single set of roots, from the anterior por- tion of the cord ; and it is then purely motor, except in virtue of its inosculation with other nerves. The Hypoglossal (9th pair of Willis) appears to be also a purely motor nerve; arising by one set of roots, and beino- distributed entirely to the muscles of the tongue, which organ derives its sensibility from other nerves. The Glossopharyn- geal usually arises from a single set of roots, and these correspond with the posterior roots of the spinal nerves ; in some animals, howrever, and occasionally in man, there is a distinct anterior root, and the nerve acquires direct motor functions. It may in some respects be 504 OF THE NERVOUS SYSTEM AND ITS ACTIONS. considered as making up, with the preceding, an ordinary spinal nerve. The Spinal Accessory, again, appears to be chiefly or entirely a motor nerve at its origin; and in like manner the Pneumogastric, or Par Vagum, seems at its roots to correspond with the posterior roots of the ordinary spinal nerves, and to execute functions analogous to theirs ; but these two nerves exchange fibres, so that each acquires in part the endoAvments of the other. The Facial nerve (or portio dura of the 7th), which is the nerve that supplies the muscles of the head in general, arises by a single root, and is exclusively motor in its properties,— except in branches which have received sensory filaments by inoscula- tion with other nerves. The same is the case, also, with the Motor Nerves of the Orbit? (the 6th, 4th, and 3d, of Willis), which arise by single roots, and which have no sensory endowments but those which they obtain by inosculation with the Fifth pair.—On the other hand, the Fifth pair arises by a double root; that which corresponds to the anterior or motor root of the spinal nerves is very small, however, and only enters the third division of the nerve, which supplies the muscles concerned in mastication; the other root, corresponding with the pos- terior roots of the spinal nerves, is of large size, and its branches are distributed to the face and head, endoAving them with sensibility. Thus the sensory division of the fifth pair, being distributed, not merely to the same parts with its motor division, but also to the parts which de- rive their motor endoAvments from the Facial nerve, and from the nerves of the orbit, may be regarded as making up, together Avith all of them, one ordinary Spinal nerve. 4. Functions of the Medulla Oblongata. 889. This portion of the nervous centres, as already stated, does not differ in any essential particular from the Spinal Cord, of which it may be considered as a cranial prolongation. But the arrangement of its constituent parts is peculiar; for whilst it is the medium by which the various strands of the Spinal Cord are connected with the different portions of the Encephalon, it is also remarkable as being the gan- glionic centre, concerned in the maintenance of the action of respi- ration, and in the ingestion of food. Four principal strands of nervous matter may be distinguished anatomically, in each of its lateral halves ; these are, anteriorly, the Anterior Pyramids; next, the Olivary bodies ; next the Restiform bodies; and lastly, the Posterior Pyramids. It Avill be presently seen, hoAvever, that the physiological relations of these strands, as indicated by their connexions with the Encephalon above, with the Spinal Cord below, and with the nerves that have their contres in them, are very different from what their mere relative posi- tions Avould indicate.—The gray or vesicular substance in this part no longer holds the same relation to the white, that it possesses in the Spinal Cord; but is principally aggregated in three pairs of gangli- onic centres, of Avhich the anterior forms the nucleus of the Olivary body, the lateral of the Restiform, and the posterior of the Posterior Pyramidal. 890. The Anterior Pyramids consist entirely of fibrous structure, STRUCTURE OF THE MEDULLA OBLONGATA. 505 and establish a communication betAveen the motor tract at the base of the Encephalon (which is chiefly derived from the Corpora Striata) and the anterior and antero-lateral columns of the Spinal Cord. They have also a connexion with the Cerebellum. A large part of its fibres decussate, those that proceed from the right hemisphere passing into the left side of the cord, and those from the left hemisphere into the right side of the cord; an arrangement which fully explains the fact, that in Hemiplegia, the paralytic affection of the body is on the side opposite to that of the lesion of the brain. A small proportion of the fibres of the anterior pyramids does not decussate; and this passes doAvn, with fibres from the olivary columns, into the anterior columns of the cord; whilst the decussating fibres dip more deeply away from the anterior surface of the cord, and connect themselves rather with its lateral or middle columns. 891. The fibrous portion of the Olivary body is connected above with the Motor tract, with the Corpora Quadrigemina, and with the Cere- / bellum, and below with the anterior columns of the Spinal Cord.—The vesicular nucleus of the Olivary body, on the other hand, which is knoAvn as the corpus dentatum., seems to be especially connected with the origins of the nerves concerned in the regulation of the movements of the tongue; thus we find that anteriorly a portion of the roots of the hypoglossal, which is the motor nerve of the tongue, issue from it; whilst posteriorly, a portion of the roots of the glossopharyngeal, which is one of the sensory nerves of the tongue, seem to terminate in it. 892. The fibres of the Restiform columns are continuous above with those of the hemispheres of the Cerebellum ; and beloAV they pass, Avithout decussation, chiefly into the posterior columns of the Spinal Cord, a band of arciform fibres, however, crossing over to the anterior and lateral columns on each side. The ganglia imbedded in these columns, hoAvever, seem to possess a completely independent function ; being the centres of the Pneumogastric nerves, which are the chief ex- citors of the Respiratory movements, as Avell as of a portion of the Glosso-pharyngeal nerves. 893. The Posterior Pyramids are two small strands of fibrous struc- ture, lying between the t\vo restiform bodies, and occupying the portion of the Medulla Oblongata on either side of the posterior median furrow. They may be traced upAvards into the Thalami Optici, and downwards into the posterior columns and the posterior part of the lateral. They undergo a decussation in their upward course ; but it is not certain Avhether this decussation involves all their fibres.—The gray nuclei of the Posterior Pyramids, which are situated immediately beneath the fourth ventricle, are the ganglionic centres of the Auditory nerves, or the proper Auditory ganglia. 894. When Ave consider these various lines of communication simply in their Physiological relations, as establishing connexions between the Encephalon above and the Spinal Cord beloAV, it will be convenient first to notice and -put aside the Cerebellar. Of these there are tAvo sets ; the principal forming the Restiform bodies, which connect the Cerebellum with the posterior columns of the Spinal cord; whilst there 506 OF THE NERVOUS SYSTEM AND ITS ACTIONS. is another division, Avhich comes into connexion through the Olivary and Pyramidal bodies, with the anterior and antero-lateral columns.— The remaining fibres, which constitute what are improperly called the Crura Cerebri, may be considered as forming two principal tracts, the sensory and the motor; these being distinguished as such by the cha- racter of the nerves which arise in their course. The sensory tract passes upwards from the posterior columns of the Spinal Cord, and the posterior part of the lateral, to the Thalami Optici; it is obviously con- tinuous below with the tract in which the posterior roots of the Spinal nerves terminate, and in its upward course it receives the large or sen- sory root of the Fifth pair; whilst passing through the Pons Varolii, it undergoes a partial decussation. On the other hand, the motor tract may be regarded as descending from the Corpora Striata and Tuber- cula Quadrigemina into the anterior and antero-lateral columns of the Spinal Cord ; in its course it gives off the roots of all the motor nerves usually considered as cranial: and the greater part of its fibres undergo decussation beloAV the Pons Varolii.—The functions of the Medulla Oblongata are, therefore, of a double character;—to bring the higher parts of the Encephalon into connexion with the Spinal Cord and the Nerves that issue from it; and to serve as a centre for the reflex movements, performed through the nerves that issue from it. In both respects it corresponds precisely with any segment of the Spinal Cord itself; and there is no reason to believe, that it possesses any other or more special endowments. The importance, however, of the reflex acts of Respiration and Deglutition, over Avhich it presides, causes this por- tion of the Medulla to be the one, whose integrity is most essential to the preservation of life; and therefore it seems to possess a character more distinctive than it really has. Fig. 154. • Dissection of the Medulla Oblongata, to show the connexions of its several strands:—A, corpus striatum; b, thalamus opticus; c, d, corpora quadrigemina; e, commissure connecting them with the cerebellum; P, corpora restiformia; P, r, pons varolii; it, st, sensory tract; mt, mt, motor tract; g, olivary tract; p, pyra- midal tract; og, olivary ganglion ; op, optic nerve; 3i», root of the third pair (motor); 5s, sensory root of the fifth pair. 895. The chief excitor nerve of the Respiratory movements, as already stated (§§ 685-687) is the afferent portions of the Par Vagum : but the STRUCTURE AND FUNCTIONS OF THE MEDULLA OBLONGATA. 507 afferent portion of the Fifth pair is also a powerful excitor; and the afferent portions of all the spinal nerves, conveying impressions from the general surface of the body, are also capable of contributing to the excitement necessary for the production of the movement.—The chief motor nerves are the phrenic and intercostals; which, though issuing from the Cord at a considerable space lower doAvn, probably originate in the Medulla Oblongata. The motor portions of several other spinal nerves are also partly concerned; as are also the Facial nerve, the motor portion of the Par Vagum, and the Spinal Accessory. The ordinary movements of Respiration involve little action of any motor nerves but the Phrenic and Intercostal; and it is only when an excess of the stimulus (produced, for example, by too long a suspension of the aerating process) excites extraordinary movements, that the nerves last enumerated are called into action. 896. The acts of Prehension of food Avith lips, and of Mastication, though usually effected by voluntary power in the adult, seem to be capable of taking place as a part of the reflex operation of the Medulla Oblongata, in the Infant, as in the loAver animals. This is particularly evident in the prehension of the nipple by the lips of the infant, and the act of suction which the contact of that body (or of any resembling it) seems to excite. The experiments provided for us by nature, in the production of anencephalous monstrosities, fully prove that the integrity of the nervous connexion of the lips and respiratory organs Avith the Medulla Oblongata, is alone sufficient for the performance of this action; and experiments upon young animals, from which the brain has been removed, establish the same fact. Thus Mr. Grainger found that, upon introducing his finger, moistened with milk, or with sugar and water, between the lips of a puppy thus mutilated, the act of suction was excited; and not merely the act of suction itself, but other movements having a relation to it; for as the puppy lay on its side, sucking the finger, it pushed out its feet, in the same manner as young pigs exert theirs in compressing the soav's dugs. This action seems akin to many of those, by Avhich the lower animals take in their food; and Ave may thus recognise in the Medulla Oblongata a distinct centre of reflex action for the reception and deglutition of aliment, analogous to the stomato-gastric ganglia of Invertebrated animals. 897. In the movements of Deglutition, which, as formerly explained (§ 453), are purely reflex, the chief excitor is undoubtedly the afferent portion of the Glosso-pharyngeal nerve. It is found that, if the trunk of this nerve, or its pharyngeal (but not its lingual) branches, be pinched, pricked, or otherwise irritated, whilst still in connexion with the Medulla Oblongata, the movements concerned in the act of swallowing are excited. The same occurs if, when the trunk of the Glosso-pharyn- geal has been divided, the cut extremity in connexion with the Medulla Oblongata is irritated; but little or no muscular contraction is produced by irritation of the separated extremity; whence it is apparent, that the Glosso-pharyngeal has little or no direct motor poAver, but acts as an excitor. In this it appears to be assisted by the branches of the Fifth pair distributed upon the fauces; and probably, also, by the branches of the superior laryngeal distributed upon the Pharynx. The 508 OF THE NERVOUS SYSTEM AND ITS ACTIONS. motor influence, which is generated in respondence to the stimulus thus conveyed, appears to act chiefly through the branches of the Par Vagum, Avhich are distributed to most of the muscles concerned in SAvallowing; but the Facial, the Hypoglossal, the motor portion of the Fifth, and perhaps also the motor portions of some of the Cervical nerves, are also concerned in the movement, and may effect it, though with difficulty, after the pharyngeal branches of the Par Vagum have been divided. 898. In the propulsion of the food doAvn the Oesophagus, to which the glosso-pharyngeal nerve does not extend, the muscular contraction, so far as it is of a reflex nature (§ 455), must depend upon the oesopha- geal branches of the Par Vagum alone; their afferent portion being the excitor, and their motor portion giving the requisite stimulus to the muscles. The same must be the case in regard to the muscular contrac- tions of the cardiac and pyloric sphincters, and of the Avails of the sto- mach, so far as regards their dependence upon the nervous system at all; but the degree of this is doubtful. 899. There are other reflex actions of the Medulla Oblongata, con- nected with the regulation of the aperture of the Glottis; these, which are effected through the superior and inferior laryngeal branches of the Par Vagum, will be better noticed, Avhen the actions of the Larynx are under consideration (§ 976).—In like manner the reflex action concerned in the regulation of the aperture of the Pupil, will be more conveniently noticed in the sketch to be hereafter given of the Physiology of Vision (§ 969). 5. Functions of the Sensory Ganglia. 900. All the nerves of Sensation, both general and special, may be traced into a series of ganglionic masses lying at the base of the brain; Avhich seem to constitute their OAvn particular centres. Thus Ave have seen in Fishes, the Olfactive, Optic, and Auditory ganglia, marked out as such, by the termination of the nerves proceeding from the organs of smell, sight, and hearing, in these masses respectively. These ganglia bear an evident correspondence with the cephalic ganglia of the Inverte- brata; which must chiefly, hoAvever, be regarded as optic ganglia, since the development of the eyes far surpasses that of the other organs of special sense. On the other hand, they find their representatives in certain organs at the base of the brain, in Man and the higher Mamma- lia ; which, though small in proportion to the whole Encephalon, are capable of being clearly marked out as the ganglionic centres of the several nerves of sense.—Thus, anteriorly, Ave have the Olfactive gan- glia, in what are commonly termed the bulbous expansions of the Olfac- tive nerve; Avhich, however, are real ganglia, containing gray or vesicular substance; and their separation from the general mass of the Encepha- lon, by the peduncles or footstalks commonly termed the trunks of the olfactory nerves, finds its analogy in many species of Fish (§869). The ganglionic nature of these masses is more evident in many of the lower Mammalia, in Avhich the organ of smell is highly developed, than it is in Man, whose olfactive poAvers are comparatively moderate.—At some distance behind these, we have the representatives of the Optic Ganglia, THALAMI OPTICI, AND CORPORA STRIATA. 509 m the Tubercula Quadrigemina, to which the principal part of the roots of the Optic nerve may be traced. Although these bodies are so small in Man, as to be apparently insignificant, yet they are relatively larger, and form a more evidently-important part of the encephalon, in many of the lower Mammalia; though still presenting the same general aspect.—The Auditory ganglia seldom form distinct lobes or projec- tions ; but are usually lodged in the substance of the Medulla Oblon- gata. lheir_ real character is most evident in certain Fishes, as the Carp; in which we find the Auditory Nerve having as distinct a gan- glionic centre as the Optic. In higher animals, however, we are able to trace the Auditory nerve into a small mass of gray matter, which lies on each side of the Fourth Ventricle; and although this is lodged in the midst of parts whose function is altogether different, yet there seems no reason for doubting that it has a character of its own, and that it is really the ganglion of the auditory nerve.—We are not able to fix upon any such mass of gray matter, as the distinct Gustatory ganglion; nor Fig. 155. Diagram of the relation of the Sensori-motor tract at the Vase of the Brain, to the Cerebrum, as seen in horizontal section:—olf, olfactive ganglia; opt, optic ganglia; aud, auditory ganglia; cs, corpora striata; thai, thalami optici; a, a, olfactive nerves; b, b, optic nerves; c, c, auditory nerves. is it necessary to attempt to do so ; for, as we shall see hereafter, there is strong reason to regard the sense of Taste as only a refined kind of Touch, combined with the sense of Smell. 901. At the base of the Cerebral Hemispheres, Ave find tAvo gan- glionic masses on either side; through which all the fibres pass that connect the Hemispheres with the Medulla Oblongata. These are the Corpora Striata, and Thalami Optici. Upon tracing fonvards the tract of motor fibres that ascend from the Anterior Pyramids, we find it passing chiefly into the Corpora Striata; whilst if we follow the 510 OF THE NERVOUS SYSTEM AND ITS ACTIONS. Sensory Column that ascends from the Posterior Pyramids, Ave shall find it to enter the Thalami Optici. These bodies have been usually considered as mere appendages to the Cerebrum ; but the fact that they are independent centres of action is fully established by the presence of a large quantity of vesicular matter in their substance; and there is now a sufficiently large amount of evidence, both anatomical and physio- logical, to render it probable that the fibres Avhich seem to pass through them from the Crura Cerebri, and then to radiate towards the periphery of the Cerebral Hemispheres, do not do so in reality, but that these ganglionic masses receive, on the one hand, the fibres that ascend to them from the Medulla Oblongata, and, on the other, are the point of departure of a new set, passing to the proper Cerebrum. Looking to the connexion of the Thalami Optici with the sensory tract, it may be regarded as not improbable that Ave may consider them as the ganglionic centres of common sensation; standing in the same relation to the sen- sory nerves, that converge from various parts of the body towards the Encephalon, as do the Optic and other ganglia to their nerves of special sensation. And as these last give origin to motor fibres, so may Ave regard the ganglionic matter of the Corpora Striata (Avhich are in close connexion with the Thalami) as probably sharing in the same function; giving origin to the motor fibres, which produce the respondent con- sensual movements; just as the anterior peak of gray matter in the Spinal Cord gives exit to the motor filaments, which effect the reflex movements excited through the afferent fibres forming part of the pos- terior roots. 902. The functions of this series of ganglia may be more certainly determined by the aid of Comparative Anatomy, than by experimental mutilations. Reverting to the class of Fishes, we find that it there constitutes, with the Cerebellum, nearly the entire Encephalon ; scarcely a rudiment of the true Cerebellum being discoverable in that group.* And when Ave descend to the Im^ertebrata, Ave find the cephalic masses entirely to consist of the ganglionic centres of the nerves of sense and motion. There can scarcely be a reasonable doubt, that these Cephalic ganglia are the seat of consciousness and the sources of those movements which are directed by sensation, in such animals as present this low type of nervous organization; and there is no adequate reason for the belief that the superaddition of the Cerebral Hemispheres in the Verte- brated series alters the endoAvments of the Sensory Ganglia on which they are superimposed; on the contrary, we everywhere see that the addition of neAV ganglionic centres, as instruments of new functions, leaves those which were previously existing in the discharge of their original duties. Hence Ave should be led to regard them as the centres of consciousness, even in Man, each pair of ganglionic centres minister- ing to that peculiar kind of sensation for Avhich its nerves and the organs they supply are set apart; thus we should consider the Optic ganglia to be the seat of Visual sensations, the Auditory to be the seat of the sense of hearing, and so on. And we should also consider them as the instru- * The ganglionic masses, commonly designated as the Cerebral lobes or hemispheres, must be really likened in great part (as already stated \ 869) to the Corpora Striata. FUNCTIONS OF THE SENSORY GANGLIA. 511 ments whereby sensations, of whatever kind, either originate or direct Automatic movements. 903. So far as the results of experiments can be relied on, they afford a confirmation of these views, by showing that sensory impressions can be felt, and that automatic movements of a higher kind than the simply reflex can be called into play after the removal of the Cerebral Hemispheres, provided that these ganglia be left intact. Thus, if a Bird be thus mutilated, it maintains its equilibrium, and recovers it when it has been disturbed; if pushed, it walks ; if throAvn into the air, it flies. A pigeon deprived of its cerebrum has been observed to seek out the light parts of a partially-illuminated room in which it was con- fined, and to avoid objects that lay in its Avay; and at night, when sleeping Avith closed eyes and its head under its Aving, it raised its head and opened its eyes upon the slightest noise. So, again, the removal or destruction of one pair of these Sensory centres appears to involve the loss of the particular sense to which it ministers; and frequently, also, to occasion such a disturbance in the ordinary movements of the animal, as to show the importance of these centres in regulating them. Such experiments have been chiefly made upon the Optic ganglia, or Corpora Quadrigemina, the partial loss of which on one side produces temporary blindness in the eye of the opposite side, and partial loss of muscular poAver on the opposite side of the body; and the removal of a larger portion, or the complete extirpation of it, occasions permanent blindness and immobility of the pupil, and temporary muscular weak- ness, on the opposite side. This temporary disorder of the muscular system sometimes manifests itself in a tendency to move on the axis, as if the animal Avere giddy; and sometimes in irregular convulsive move- ments. Here, then, we have proof of the necessity of the integrity of this ganglionic centre for the possession of the sense of vision; and Ave have further proof that the ganglion is connected with the muscular apparatus, by motor nerves issuing from it. The reason why the eye of the opposite side is affected is to be found in the decussation of the optic nerves, a point to be immediately adverted to (§ 907). The in- fluence of the operation on the muscles of the opposite side of the body is at once understood from the fact of the decussation of the motor fibres in the anterior pyramids (§ 890). Similar disturbances of move- ment have been produced by injuries to the organs of sense themselves, or to the nerves connecting them Avith the sensorial centres. Thus, if one of the eyes of a pigeon be blindfolded, or its humors be evacuated, vertiginous motions ensue ; and section of one of the semicircular canals of the ear in pigeons and rabbits has been found to occasion constant efforts to move in the plane of that canal, thus confirming the belief that the function of these canals is to indicate the direction of sounds (§ 952). 904. Notwithstanding that, in Man, the high development of Lntel- ligcnce and the exercise of the Will, supersede in great degree the operations of Instinct, Ave still find that there are in ourselves certain movements which can be distinguished as neither voluntary nor simply reflex, and which are examples of the method of operation that seems to be the chief source of the actions of the lowTer Vertebrata, as of the 512 OF THE NERVOUS SYSTEM AND ITS ACTIONS. Invertebrated classes in general. These movements are as automatic and involuntary as are the ordinary reflex actions, but differ from them in requiring that the impressions which originate them should he felt as sensations ; and hence they are conveniently designated as consensual. As examples of this group, we may advert to the start upon a loud and unexpected sound; the sudden closure of the eyes to a dazzling light, or on the approach of bodies that might injure them, Avhich has been observed to take place even in cases in which the eyelids could not be voluntarily closed; the act of sneezing excited by an irritation of the nostril, and sometimes also by a dazzling light; the semi-convulsive movements and the laughter called forth by tickling; and the vomiting occasioned by the sight or the smell of a loathsome object. So, again, the act of yawning is ordinarily called forth by certain uneasy sensa- tions within ourselves, but also by the sight or hearing of the act as performed by another. Various phenomena of disease exhibit the powerful influence of sensations in producing automatic motions. As instances of this kind, we may refer to the effects of the sight or the sound of liquids, or of the slightest currents of air, in exciting the Hydrophobic paroxysm; whilst in many Hysteric subjects the sight of a paroxysm in another individual is the most certain means of its induc- tion in themselves. The most remarkable examples, hoAvever, of auto- matic movements depending upon sensations, are those which we come to perform habitually, and, as we commonly say, mechanically, Avhen the attention and the voluntary effort are directed in quite a different channel. Thus the man who is walking through the streets in a com- plete revery, unravelling some knotty subject, or working out a mathe- matical problem, not only performs the movements of progression, which may be simply reflex, with great regularity, but also directs these in a manner which plainly indicates the guidance of sensations. Thus, he will avoid obstacles in the line of his path, and he Avill folloAV the course Avhich he has been accustomed to take, although he may have intended to pass along some very different route; and it is not until his attention is recalled to his situation, that his train of thought suffers the least intermission, or that his will is brought to bear upon his motions. 905. We may trace the agency of the Sensory Ganglia, however, in the Human subject, not merely in their direct and independent opera- tion upon the muscular system, but also in the manner in Avhich they participate in all Voluntary actions. The existence of a Sensation of some kind, in connexion with a Muscular exertion, seems essential to the continuance of the latter. Our ordinary movements are guided by what is termed the Muscular Sense ; that is, by a feeling of the con- dition of the muscle, that comes to us through its OAvn sensory nerves. How necessary this is to the exercise of muscular power may be best judged of from cases in Avhich it has been lost. Thus, a woman Avho had suffered complete loss of sensation in one arm, but who retained its motor power, found that she could not support her infant upon it, without constantly looking at the child; and that, if she Avere to remove her eyes for a moment, the child would fall in spite of her knowledge that her infant wras resting upon her arm, and of her desire to sustain it. Here, the muscular sense being entirely deficient, the sense of DECUSSATION OF THE OPTIC NERVE. 513 vision supplied Avhat Avas required, so long as it was exercised upon the object; but as soon as this guiding influence wras withdrawn, the strong- est will could not sustain the muscular contraction. Again, in the production of vocal sounds, the nice adjustment of the muscles of the larynx, which is requisite to produce determinate tones, can only be effected in obedience to a mental conception of the tone to be uttered; and this conception cannot be formed, unless the sense of hearing has previously brought similar tones to the mind. Hence it is that persons who are born deaf are also dumb. They may have no malformation of the organs of speech; but they are incapable of uttering distinct vocal sounds or musical tones, because they have not the guiding conception, or recalled sensation, of the nature of these. By long training, and by efforts directed by the muscular sense of the larynx itself, some persons thus circumstanced have acquired the power of speech ; but the want of sufficiently definite control over the vocal muscles is always very evident in their use of the organ. 906. Quitting noAV the functions of the Sensory Ganglia, we have briefly to notice certain peculiarities in the characters of the Nerves to which they serve as the centres. And of these peculiarities, there is one of a very remarkable nature, which is common to the three nerves of special sense,—namely, the Olfactive, Optic, and Auditory ;—that they are not in the least degree endowed Avith common sensibility; so that they may be cut, stretched, pinched, &c, without producing the least pain. Consequently, the ordinary sensibility of the surfaces they supply is entirely due to the branches of the Fifth pair, which are dis- tributed upon them ; and Ave may have a loss of either the general or special sensibility of any of the organs of sense, without the other being affected, save indirectly. Again, we do not find that irritation of these nerves produces any other purely reflex movements, than such as are connected with the operations of the organs of sense, in which they respectively originate. Thus the Olfactory nerve cannot, by any irrita- tion, be made to excite a reflex movement; the only reflex action that can be excited by irritating the Optic nerve, is contraction of the Pupil; and the regulation of the tension of the Membrana Tympani (if, as is probable, this is effected by the motor poAver of the Facial nerve, excited by impressions made upon the organ of sense), appears to be the only reflex action to which the Auditory nerve can minister. 907. There is a further peculiarity, of a very marked kind, attend- ing the course of the Optic nerves; this is the crossing or decussation Avhich they undergo, more or less completely, whilst proceeding from their ganglia to the eyes. In some of the lower animals, in which the two eyes (from their lateral position) have entirely different spheres of vision, the decussation is complete ; the whole of the fibres from the right Optic ganglion passing into the left eye, and vice versd. This is the case for example, with most of the Osseous Fishes (as the cod, halibut, &c); and also, in great part at least, with Birds. In the Human subject, however, and in animals which, like him, have the two eyes looking in the same direction, the decussation seems less complete ; but there is a very remarkable arrangement of the fibres, which seems destined to bring the two eyes into peculiarly consentaneous action. 514 OF THE NERVOUS SYSTEM AND ITS ACTIONS. The posterior border of the Optic Chiasma is formed exclusively of co?n- missural fibres, which pass from one optic ganglion to the other, with- out entering the real optic nerve. Again, the anterior border of the chiasma is composed of fibres, which seem, in like manner, to act as a commissure between the two retinar; passing from one to the other, without any connexion with the optic ganglia. The tract which lies between the two borders, and occupies the middle of the chiasma, is the true optic nerve; and in this it Avould appear that a portion of the fibres decussates, whilst another portion passes directly from each Op- tic ganglion into the corresponding eye. The fibres which proceed from the ganglia to the retinae, and constitute the proper optic nerves, may be distinguished into an internal and external tract. Of these, the external, on each side, passes directly onwards to the eye of that side; whilst the internal crosses over to the eye of the opposite side. The dis- tribution of these two sets of fibres in the retina of each side respectively, is such that, according to Mr. Mayo, the fibres from either optic gan- glion will be distributed to its own side of both eyes;—the right optic ganglion being thus exclusively connected with the outer part of the retina of the right eye, and with the inner part of the retina of the left eye; and the left optic ganglion being, in like manner, connected ex- clusively with the outer side of the left retina, and with the inner side of the right. Now as either side of the eye receives the images of objects, which are on the other side of its axis, it follows, if this account of their distribution be correct, that in Man, as in the lower animals, each ganglion receives the sensations of objects situated on the opposite sides of the body. The purposes of this decussation may be, to bring the visual impressions, which are so important in directing the move- ments of the body, into proper harmony with the motor apparatus ; so that, the decussation of the motor fibres in the pyramids being accom- panied by a decussation of the optic nerves, the same effect is produced as if neither decussated,—which last is the case with Invertebrated animals in general. 6. Functions of the Cerebellum. 908. Much discussion has taken place, of late years, respecting the uses of the Cerebellum ; and many experiments have been made to determine them. That it is in some way connected with the powers of motion, might be inferred from its connexion with the antero-lateral columns of the Spinal Cord, as well as with the posterior ; and the com- parative size of the organ, in different orders of Vertebrated animals, gives us some indication of what the nature of its functions may be. For Ave find its degree of development corresponding pretty closely with the variety and energy of the muscular movements which are habitually exe- cuted by the species ; the organ being the largest in those animals Avhich require the combined effort of a great variety of muscles to maintain their usual position, or to execute their ordinary movements; whilst it is the smallest in those which require no muscular exertion for the one pur- pose, and little combination of different actions for the other. Thus in animals that habitually rest and move upon four legs, there is com- FUNCTIONS OF THE CEREBELLUM. 515 paratively little occasion for any organ to combine and harmonize the actions of their several muscles ; and in these, the Cerebellum is usually small. But among the more active predaceous Fishes (as the Shark), Birds of the most powerful and varied flight (as the SAvallow), and such Mammals as can maintain the erect position, and can use their extremities for other purposes than support and motion,—we find the Cerebellum of much greater size, relatively to the remainder of the Encephalon. There is a marked advance in this respect, as we ascend through the series of Quadrumanous animals ; from the Baboons, which usually walk on all-fours, to the semi-erect Apes, which often stand and move on their hind-legs only. The greatest development of the Cere- bellum is found in Man; who surpasses all other animals in the number and variety of the combinations of muscular movement which his ordinary actions involve, as well as of those which he is capable, by practice, of learning to execute. 909. From experiments upon all classes of Vertebrated animals, it has been found that, when the Cerebellum is removed, the power of walking, springing, flying, standing, or maintaining the equilibrium of the body, is destroyed. It does not seem that the animal has in any degree lost the voluntary power over its individual muscles; but it can- not combine their actions for any general movements of the body. The reflex movements, such as those of respiration, remain unimpaired. When an animal thus mutilated is laid on its back, it cannot recover its former posture; but it moves its limbs, or flutters its wings, and evi- dently is not in a state of stupor. When placed in the erect position, it staggers and falls like a drunken man; not, hoAvever, without making efforts to maintain its balance. Phrenologists, who attribute a different function to the Cerebellum have attempted to put aside these results, on the ground that the severity of the operation is alone sufficient to produce them ; but, as we shall presently see, many animals may be subjected to a much more severe operation, the removal of the Cerebral hemispheres, without the loss of the power of combining and harmoniz- ing the muscular actions, provided the Cerebellum be left uninjured.— Thus, then, the idea of the functions of the Cerebellum, which we derive from Comparative Anatomy, seems fully borne out by the results of ex- periment ; and it is also consistent Avith the indications which may be drawn from the observations of Pathological phenomena. When the Cerebellum is affected with chronic disease, the motor function is seldom destroyed ; but the same kind of want of combining power jsIioavs itself, as when the organ has been purposely mutilated. Some kind of lesion of the motor function is invariably to be obsen'ed ; whilst the mental powers may or may not be affected,—probably according to the influ- ence of the disease in the Cerebellum, upon other parts. ^ The same absence of any direct connexion with the Psychical poAvers, is shoAvn in the fact, that inflammation of the membranes covering it, if confined to the Cerebellum, does not produce delirium. Sudden effusions of blood into its substance may produce apoplexy or paralysis ; but this may occur as a consequence of effusions into any part of the Encephalon, and does not indicate that the Cerebellum has anything to do with the mental functions, or with the power of the Will over the muscles. 516 OF THE NERVOUS SYSTEM AND ITS ACTIONS. 910. There is another doctrine, however, in regard to the functions of the Cerebellum, first propounded by Gall; which ought not to be altogether passed by. According to the system of Phrenologists, the Cerebellum is the organ of the sexual instinct; and its connexion with the motor function is limited to the performance of the movements, to which that instinct leads. This doctrine derives no support, however, from the fact* supplied by Comparative Anatomy; for there is a com- plete want of correspondence between the size of the Cerebellum in different animals, and the power of their sexual instinct.—Again, although Pathology has been appealed to, as showing a decided con- nexion between disease of the Cerebellum and affection of the Genital organs (manifesting itself in priapism, turgescence of the testes, seminal emissions, &c), yet it appears, on a careful examination of evidence, that such a sympathy is comparatively ^rare, not being displayed in more than one out of every seventeen cases of Cerebellic disease. And where it is manifested, it is explicable quite readily by the known fact, that this kind of excitement of the genital organs may be produced by excitement of the spinal cord and medulla oblongata.—Little or no light has been thrown on this question by experiment. It was asserted by Gall, that the Cerebellum is very small in castrated animals; but this assertion has been met by the most positive counter-statements on the part of Leuret, who has shown that the average weight of the Cere- bellum (both absolutely and in proportion to the weight of the entire encephalon) is even greater in Geldings, than in Stallions or Mares.—It is asserted, however, that the results of observation in Man lead to a positive conclusion, that the size of the Cerebellum is a measure of the intensity of the sexual instinct in the individual. This assertion has been met by the counter-statement of others, that no such relation exists. There are, of course, very great difficulties in regard to the collection of accurate information on this subject; and the question must be at present regarded as sub judice. 911. It may be added, that the idea of a special connexion between the sexual instinct and the Cerebellum, is not inconsistent with the view of its function previously stated ; and it would seem to derive some confirmation from the fact, that an unusual amount of muscular exer- tion appears to have a peculiar tendency to depress the sexual passion, even whilst it increases the general vigor of the system. If the Cere- bellum be really connected with both kinds of functions, it does not seem unlikely that the excessive employment of it upon one, should diminish its energy in regard to the other. Further, it seems not improbable, that the Lobes of the Cerebellum are the parts specially concerned in the regulation of the muscular movements ; whilst the central portion (constituting the Vermiform process in Man, but form- ing the entire cerebellum of many of the lower Vertebrata, such as the Frog) may contain the centre of the sexual sensations, and may thus be the instrument of the consensual actions to Avhich they give rise. 7. Functions of the Cerebrum. 912. The view which has been taken of the Comparative structure of the Nervous system, in different animals, leads to the conclusion, that RELATIVE DEVELOPMENT OF THE CEREBELLUM. 517 the Cerebral Hemispheres are far from being the essential parts of the apparatus they were formerly imagined to be ; and that they are, on the contrary, superadded organs, of which we find no distinct represen- tatives in the Invertebrata, and of which the first appearance (in the class of Fishes) exhibits them in the light of appendages, destined to perform some special function peculiar to Vertebrated animals. The results of the removal of the Cerebral Hemispheres, in animals to which the shock of the operation does not prove immediately fatal, fully con- firms this view; and must appear extraordinary to those, who have been accustomed to regard these organs as the centre of all energy. Not only Reptiles, but Birds and Mammalia, if their physical wants be supplied, may survive the removal of the whole Cerebrum for weeks, or even months. If the entire mass be taken away at once, the operation is usually fatal; but if it be removed by successive slices, the shock is less severe, and the depression it produces in the organic functions is soon recovered from. It is difficult to substantiate the existence of actual sensation in animals thus circumstanced; but their movements appear to be of a higher kind, as already remarked (§ 903), than those resulting from mere reflex action. Thus they will eat food, when it is put into their mouths ; although they do not go to seek it. If violently aroused, the animal has all the manner of one waking from sleep; and it manifests about the same degree of consciousness as a sleeping Man, whose torpor is not too profound to prevent his suffering from an uneasy position, and who moves himself to amend it. In both cases, the movements are consensual only, and do not indicate any voluntary power ; and we may well believe that, in the former case as in the latter, though felt they are not remembered; an active state of the Cerebrum being essential to memory, though not to sensations, which simply excite certain actions.—When the Cerebral Hemispheres are being removed, slice by slice, it is noticed that injuries of these organs neither occasion any signs of pain, nor give rise to convulsive move- ments. Even the Thalami and Corpora Striata may be wounded, without the excitement of convulsions; Avhilst, if the incisions involve the Tubercula Quadrigemina, convulsions uniformly occur. It has been often observed in Man, that, when it has been necessary to separate protruded portions of the brain from the healthy part, no sensation was produced, even though the mind Avas perfectly clear at the time. Hence it would appear that neither is the Cerebrum itself the centre of sensa- tion, nor is it so connected with that centre, as to be able to convey to it sensory impressions of an ordinary kind. This is analogous to the condition of the nerves of special sense, as already remarked. That no irritation of the cerebral substance should excite convulsive movements, is a very remarkable circumstance; and it seems to indicate, that the changes which mental operations produce in the cerebral fibres, cannot be imitated, as changes in other motor fibres may be, by physical im- pressions. . 913. As already stated, the relative amount of Lntelhgence in diffe- rent animals bears so close a correspondence with the relative size and development of the Cerebral Hemispheres, that it can scarcely be ques- tioned that these constitute the organ of the Reasoning faculties, and 518 OF THE NERVOUS SYSTEM AND ITS ACTIONS. issue the mandates by which the Will calls the muscles into action. It must be borne in mind, however, that size is not by any means the only indication of their comparative development. As we advance from the lower to the higher Vertebrata, we observe a marked advance in the complexity of the structure of the Cerebrum. Its surface becomes marked by convolutions, that greatly increase the area over which blood-vessels can enter it from the surrounding membranes; and in pro- portion to the increase in the number and depth of these, do we find an increase in the thickness of the layer of gray matter, which is the source of all the powers of the organ. The arrangement of the white or fibrous tissue, which forms the interior of the mass, also increases in com- plexity ; and as we ascend even from the lower Mammalia up to Man, we trace a 'marked increase in the number of the fibres, which esta- blish communication between different parts of the organ. It is, in fact, not merely from the different parts of the gray matter which forms the surface of the hemispheres, that these commissural fibres arise ; but also from those isolated portions of vesicular substance, which are found in different parts of their interior; and an extremely complex system is thus formed, which is still but very imperfectly understood. 914. The most important group of commissural fibres, is that which connects the Sensory with the Hemispheric Ganglia; that is, which radiates from the Thalami Optici and Corpora Striata, to the stratum of gray matter which forms the convoluted surface of the Cerebrum. These fibres constitute, in fact, the principal part of the white sub- stance of the brain ; the remainder being made up by the commissures to be presently described, and by commissural fibres which (it is pro- bable) connect the different parts of the Cerebral surface with each other. It was formerly supposed (and is still maintained by many Anatomists), that the radiating fibres which may be traced to the Cor- pora Striata and Thalami Optici, pass through these bodies, and are continuous with the Crura Cerebri, and consequently with the sensory and motor tracts of the Medulla Oblongata. But Avhen the small size of the Crura Cerebri is compared with the relatively enormous bulk of the radiating fibres, it is obvious that the former can only contain but a very small proportion of the latter; and as no absolute continuity has been traced, it appears more conformable to Anatomical and Phy- siological probability, to believe that the fibres of the Crura Cerebri pass no further upwards than the Sensory Ganglia, and that the radiating fibres take a fresh departure from these bodies, to pass to- wards the surface of the Cerebrum.—Thus, then, we should be led to regard the Spinal Cord, Medulla Oblongata, and chain of Sensory Ganglia, as precisely representing the entire Nervous System of Insects, the character of whose action is essentially automatic ; and to consider the Cerebrum as an organ superadded at its summit, receiving all its incitement to action from impressions transmitted to it through the Sensory Ganglia, and carrying into effect its volitional determinations and emotional impulses, not (as formerly supposed) by immediately ex- citing muscular movements through nervous communications passing direct from the convoluted surface of the Cerebrum, but by playing downwards upon the Automatic apparatus by Avhich its mandates are COMMISSURES OF THE CEREBRAL HEMISPHERES. 519 carried into effect (Figs. 155, 156). Of this view we shall presently find that there is strong physiological evidence. Fig. 156. Diagram of the mutual relations of the principal Encephalic centres, as shown in a vertical section :—a, Cerebrum; b, Cerebellum; c, Sensori-motor tract, including the Olfactive ganglion olf the Optic opt, and the Auditory aud, with the Thalami Optici thai, and the Corpora Striata cs; r>, Medulla Oblongata; e, Spinal Cord:—a, olfactive nerve; 6,optic; c, auditory; d, pneumogastric; e, hypoglossal; /, spinal: fibres of the medullary substance of the cerebrum are shown, connecting its ganglionic surface with the sensori-motor tract. 915. The two Hemispheres are united on the median line by several transverse commissures; of which the Corpus Callosum is the most im- portant. This consists of a mass of fibres very closely interlaced to- gether ; which may be traced into the substance of the hemispheres on each side, particularly at their lower part, Avhere they are connected with the thalami optici and corpora striata. It is difficult, if not impos- sible, to trace its fibres any further; but there can be little doubt that they radiate, with the fibres proceeding from the bodies just named, to the different parts of the surface of the hemispheres. This commissure is altogether absent in Fish, Reptiles, and Birds; and it is partially or completely wanting in the Mammalia with least perfect brain, as the Rodents and Marsupials.—The other transverse commissures rather belong to the Sensory Ganglia than to the Cerebral hemispheres. Thus the anterior commissure particularly unites the Corpora Striata of the two sides; but many of its fibres pass through those organs, and radiate towards the convolutions of the hemispheres, especially those of the middle lobe. This commissure is particularly large in those Marsu- pials, in which the corpus callosum is deficient.—The posterior com- missure is a band of fibres which connects the Optic Thalami; crossing over from the posterior extremity of one to that of the other.—Besides these, there are other groups of fibres, which seem to have similar com- missural functions, but which are intermingled with vesicular substance. 520 OF THE NERVOUS SYSTEM AND ITS ACTIONS. Such are the soft commissure, which also extends between the Thalami; the Pons Tarini, Avhich extends between the two crura or peduncles of the Cerebrum; and the Tuber Cinereum, which seems to unite the optic tracts with the thalami, the corpus callosum, the fornix, &c, and to be a common point of meeting for several distinct groups of fibres. 916. The anterior and posterior parts of the hemispheres, moreover, are connected by longitudinal Commissures, of which some lie above, and some below, the corpus callosum ; and of these, also, a part belong to the Sensory Ganglia. Above the transverse fibres of the corpus callosum, there is a longitudinal tract on each side of the median line, which serves to connect the convolutions of the anterior and posterior lobes of the brain.—And above this, again, is the superior longitudinal commissure, which is formed by the fibrous matter of the great convolu- tion nearest the median plane on the upper surface of the brain, and which connects the convolutions of the anterior and middle lobe with those of the posterior.—Beneath the great transverse commissure, we find the most extensive of all the longitudinal commissures, namely, the Fornix. This is connected in front with the optic thalami, the mammil- lary bodies, the tuber cinereum, &c; and behind, it spreads its fibres over the hippocampi (major and minor), which are nothing else than peculiar convolutions that project into the posterior and descending cornua of the lateral ventricles. The fourth longitudinal cotnmissure is the tarnia semicircularis, which forms part of the same system of fibres with the fornix; connecting the corpus mammilare and thalamus opticus with the middle lobe of the cerebral hemisphere. If, as Dr. Todd has remarked, we could take away the corpus callosum, the gray matter of the internal convolution, and the ventricular prominence of the optic thalami, then all these commissures would fall together, and become united as one and the same series of longitudinal fibres.—It is curious that there should be no direct communication betAveen the Cerebral hemispheres and the Cerebellum ; the only commissural band between them being the processus a cerebello ad testes, which passes onwards, through the Tubercula Quadrigemina, to the Thalamus Opticus on each side. This would seem to confirm the idea of the complete distinctness of their functions. 917. The Cerebrum appears to be the instrument of all those psychi- cal operations, which are superadded, in Man and the higher Verte- brata, to mere sensations. The impressions which are merely felt in the sensorium, give rise, Avhen they pass upAvards into the Cerebrum, to Ldeas, which then become the material (so to speak) of all the higher mental processes. These processes may be ranked under two dis- tinct heads, namely, the Emotional and the Intelligential; the former being most intimately connected with the sensations Avhich prompt them, whilst the latter are commonly of a much more abstract character. The Emotions may, in fact, be considered as feelings of pleasure or pain associated with particular classes of ideas; and it is this association which gives them the character of the moving or active powers of the mind, and which makes them, either directly or indirectly, the springs of the greater part of our actions. When strongly excited, the Emo- tions may produce movements which the Will may not be able to re- FUNCTIONS OF THE CEREBRUM. 521 strain ; as when Ave burst into laughter at some ludicrous image presented to the mind, either by a present sensation or by an act of the Memory or Imagination, notwithstanding the strongest inducements presented by "time, place, and circumstance" to a preservation of our gravity. The distinctness of the character of Emotional and Volitional move- ments is further evident from this, that cases of paralysis not unfre- quently occur (especially in the facial nerves, through which most of the muscles of "expression" are excited to action), in which the muscles are obedient to one class of impulses, Avhile the other exerts no power over them. Thus, in one instance, the muscles of one side of the face were palsied in such a manner, that the patient could not voluntarily close his eye, nor draw his mouth towards that side ; yet when any ludicrous circumstance caused him to laugh, their usual play was manifested in the expression of his countenance. And in another case, the muscles were obedient to the will; but when the individual laughed or cried under the influence of an emotion, it was only on one side of his face. To these may be added another case, in which the right arm was com- pletely palsied, so that the individual had not the least voluntary power over it; yet it was violently agitated, whenever he met a friend whom he desired to greet.—The influence of an undue tendency to Emotional excitement, is remarkably seen in what are ordinarily termed Hysterical states of the system; in which violent convulsive paroxysms are fre- quently brought on by the most trivial causes, if these should call the passions or affections of the mind into undue activity. There can be no doubt that many of the peculiar actions performed by the subject of what is termed Mesmeric influence, are the result of a condition of this nature. There appears to be, in such persons, a proneness to activity of the consensual and emotional parts of the nervous centres, Avhich manifests itself most strongly when the control of the Avill is withdrawn; and thus very slight impressions produce very powerful involuntary movements,—especially when this response is favored by the strong desire, on the part of the patient, to exhibit any particular manifestation that is known to be expected by the bystanders. 918. It has been supposed by some, that the Emotional movements of Man and the higher animals may be ranked in the same category with Lnstinctive actions of the lower; and that the Desires of the former are comparable to the instinctive Propensities of the latter. But this comparison is erroneous; for what Ave term propensities (among the lower animals) are nothing else than tendencies to perform particular movements in respondence to particular sensations, without any idea of the purpose of the movement or of the object AA'hich has excited it; whilst an Emotion involves an idea of the object which has called it up, and a Desire ■ involves a conception of the object to be obtained.—The imitative actions afford a good example of the difference betAveen a propensity and a desire. The former is manifested in such imitative movements as are purely consensual; the sensation, Avhich is the mainspring of the action in each case, exciting a respondent automatic movement, as when we yawn involuntarily from seeing or hearing the action performed by another or as when children learn undesignedly to perform many of the movements Avhich they Avitness in adults. This propensity to in- 522 OF THE NERVOUS SYSTEM AND ITS ACTIONS. voluntary imitation is much stronger in some individuals than in others. On the other hand, imitative actions may be voluntarily performed, as the result of a desire to execute them, which involves a distinct idea of the object; and the moving force of this desire is derived from the pleasure which the individual derives from the performance, and Avhich he finds either in the act itself, or in the enjoyment which it affords to others, or in its prospective benefits (pecuniary or otherwise) to himself. Thus we see that the Mind (properly so called) is concerned in all Emo- tional actions; whilst there is no evidence of the participation of any higher N attribute than sensation in the purely Instinctive acts; and even this is not a requisite link in the chain, by which many of the movements are excited, that are usually grouped together under that designation. 919. Again, the Emotions may be excited by operations of the Mind itself, as well as by sensations immediately received from without. Thus, involuntary laughter may result from a ludicrous idea, called up by some train of association, and having no obvious connexion with the sensation which first set this process in operation ; and the various move- ments of the face and person by which Actors endeavor to express strong emotions, are most effectual in conveying their meaning, when they result from the actual working of the emotions in the mind of the performer, who has by an effort of the will, identified himself (so to speak) with the character he personates. A still more remarkable case is that in which paroxysms of Hysterical convulsion, in themselves beyond the power of the Will to excite or control, are brought on by a voluntary effort; this being exerted, not in the attempts to perform the move- ments, but in "getting up," so to speak, the state of feeling, from which, AAThen it is once excited, the movements spontaneously flow. In all these instances, and others of like nature, it would seem as if the agency of the Cerebrum produced the same condition in the Automatic centres, as that which is more directly excited by sensations received through their own afferent nerves.—But on the other hand, the Emo- tions, by their influence on the Reasoning processes, are largely con- cerned in many actions which are strictly voluntary; in fact it may be questioned Avhether there are any of our actions, the power necessary for whose performance is not derived, directly or indirectly, from emotional states of mind; all our motives to any kind of exertion being found, if care- fully analysed, to have reference to pleasure to be derived, or pain to be avoided, either in the very performance of the action, or in the conse- quences Avhich our reasoning processes connect with it. And it will be found that the difference betAveen those persons who are said to act from feeling, and those who are said to be guided by reason, is not precisely what these terms imply ; for the actions of both are equally determined by the motives supplied by emotional states ; and the difference rather • lies in this, that one class act on their first impulses without considering the consequences, whilst the other calculate the remoter results, and weigh the future pain against the present pleasure, the ultimate enjoy- ment against the immediate distress.—The Emotional states are pecu- liarly liable to be influenced by the condition of the corporeal system ; thus a very slight depravation of the blood may produce an irresistible tendency to take a gloomy view of everything to which the mind may FUNCTIONS OF THE CEREBRUM. 523 be directed, and especially of all that relates to the individual; whilst that condition of perfect health which is derived from wholesome re- creation, fresh air, active exercise, &c, is almost always accompanied Avith a degree of cheerfulness and elasticity, which occasions even real evils to be but comparatively little felt. 920. When we turn our attention to the Intelligential actions of which the Cerebrum appears (in our present state of being) to be the exclusive instrument, we perceive that the attribute by which they are distinguished both from the Instinctive and Emotional, is their inten- tional or purposive performance, in accordance with the mental concep- tion of the object to be attained, and the intellectual belief as to the most advantageous means of accomplishing it. The decision thus formed by the Reasoning processes, is put into operation by the Will; and thus it is the characteristic of a Voluntary act, that it is designed by the individual to answer a certain purpose which is distinctly present to the mind.—Now when we come to analyze the faculties concerned in this class of operations, we find that the one most closely related to the simple Sensorial powers already treated of, and at the same time most essential to all the "higher operations, is Memory. This faculty is one of those first awakened in the opening mind of the Infant; and we find traces of it in animals that seem to be otherwise guided by pure Instinct. It obviously affords the first steps towards the exercise of the reasoning poAvers ; since no experience can be gained without it, and the foundation of all intelligent adaptation of means to ends lies in the application of the knoAvledge which has been acquired and stored up in the mind. There is strong reason to believe that this attribute belongs to the Cerebrum exclusively; no impression made upon the Sensorial centres being ever remembered, unless they are registered (as it were) in this organ. And further, there is evidence that no impression of this kind once made upon the Cerebrum is ever entirely lost, in the normal state; although disease or accident will sometimes occasion a complete destruction of the memory, or wrill obliterate the remembrance of a particular class of objects or of ideas. All memory, hoAvever, seems to depend upon the principle of Sugges- tion ; one idea being linked with another, or with a particular sensa- tion, in such a manner as to be called up by its recurrence ; and a period of many years frequently intervening, without that combination of circumstances presenting itself Avhich is requisite to arouse the dor- mant impression of some early event. Sometimes this combination occurs in Dreaming, Delirium, or Insanity, three states which agree in this, that the Will has no control over the current of thought; and ideas are thus recalled, of which the mind in a state of healthy activity ■has no remembrance. 921. It is upon the ideas aroused in the mind by Sensorial changes, or recalled by Conception, or evolved by the process of Reflection (in Avhich the mind perceives its OAvn operations, and traces relations amongst its objects of thought), or generated by the Lmagination (which really acts, however, rather by combining into new forms, than by creating altogether de novo), that all acts of Reasoning are based. These consist, for the most part, in the aggregation and collocation of 524 OF THE NERVOUS SYSTEM AND ITS ACTIONS. ideas, the decomposition of complex ideas into more simple ones, and the combination of simple ideas into general expressions; in which are exercised the faculty of Comparison, by which the relations and con- nexions of ideas are perceived, that of Abstraction, by which we fix our attention on any particular qualities of the object of our thought, and isolate it from the rest, and that of Generalization, by which we grasp in our minds some definite notions in regard to the general relations of those objects. These are the processes chiefly concerned in the simple acquirement of KnoAvledge, Avith which class of opera- tions the Emotional part of our nature has very little participation, save as furnishing the desire which may be the necessary incitement to the exertion of the intellect. A certain measure of intellectual activity seems natural to Man, provided that the development of the mind has taken place under favorable circumstances; and our highest pleasures are connected with the healthful and almost spontaneous exer- cise of its faculties. 922. But the Will possesses a determining power over the mental as Avell as over the bodily operations; and it is, in fact, this determin- ing power which is the source of the self-control that characterizes the well-regulated mind of Man, and distinguishes him alike from the madman and the brute. The regulation of our conduct consists in the application of our reasoning poAvers to the circumstances of our con- dition, and in the due regulation of those emotional tendencies, which (as already pointed out, § 919) are the moving springs of our actions. However powerful these tendencies may be, there can be no doubt, that we possess within ourselves the means of checking them, by withdrawing our minds by a voluntary effort from the thoughts which they suggest, as well as by calling forth opposing influences within us, so that the decision which is finally arrived at is something very different from that which the first "balance of motives" Avould have produced. It is the deficiency or entire loss of this power of self- control that usually constitutes the first step in the development of Insanity; for this state generally consists, not so much in a perversion of the reasoning processes, as in a disorder of the emotional state, which causes the patient to dwell upon particular trains of thought, until his feelings in relation to them become exaggerated or perverted; and at last intellectual delusions arise, from the habit of vieAving every- thing that comes before the mind through a distorted medium, and from the substitution of the patient's morbid imaginings for real occur- rences. In what is now termed impulsive Insanity, there is intellectual perversion ; but a desire of some kind is so powerfully excited, that the Will cannot control it. And every phase may be witnessed between a state of this kind, which renders the individual an irresponsible agent, and that mental condition in which the individual, though originally fully able to control himself, habitually gives Avay to his passions, and thus, by their continual indulgence, at last alloAvs them to become the domi- nant powers of his mind. 923. Although the Will has been usually regarded as directly deter- mining those muscular movements Avhich are usually distinguished as Voluntary, through the intermediation of fibres originating in the cere- FUNCTIONS OF THE CEREBRUM. 525 bral convolutions and proceeding to the muscles, yet a careful analysis of the process fully bears out the idea already put forwards, that the Will really operates through the Automatic apparatus, exciting parti- cular groups of muscular actions, just as they would be called forth by sensations directly excited by external objects. For it has been shown that the Craniospinal Axis (consisting of the Sensory Ganglia, Me- dulla Oblongata, and Spinal Cord) receives all the sensory nerves, and gives origin to all the motor; and that the fibres which pass between the Cerebral convolutions and the Sensory Ganglia, probably serve to bring these centres into mutual relation, and are not continuous with those of any nerves, either sensory or motor. And we might expect, therefore that the addition of a cerebrum to this automatic apparatus would have the effect of supplying a new stimulus to movement, which, whilst proceeding from mental operations, should still act through the same mechanism as that already provided for the reflex and consensual movements. Now, when we attentively consider the nature of what we are accustomed to call voluntary action, we perceive that the agency of the Will is limited to the determination of the result; and that it has nothing to do with the selection and co-ordination of the individual movements, by which that result is brought about. If it were other- wise, we should be dependent upon our anatomical knowledge, for our power of performing even the simplest movements of the body. Again, there are very few cases in which we can single out any indivfdual muscle, and put it into action independently of others; and the cases in which we can do so, are those in which a single muscle is concerned in producing the result, as in the elevation of the eyelid; and we then really single out the muscle by "willing" the result. Thus, then, how- ever startling the position may at first appear, Ave have a right to affirm that the Will cannot exert any direct or immediate power over the muscles; but that its determinations are carried into effect through an intermediate mechanism, which, without any further guidance on our own part, selects and combines the particular muscles whose contractions are requisite to produce the desired movement. We have seen that the Sensorial centres play (so to speak) upon the Cerebrum, sending to it impressions of a kind fitted to call forth its peculiar activity as an instru- ment of purely mental operations ; and in return, the Cerebrum appears to play downwards upon the motor portion of the automatic apparatus, sending to it volitional impulses Avhich excite its motorial activity. And thus Ave see that the very same action may be excited through an impres- sion conveyed to the centres of the whole system through some one or more nerves of the external senses, or through the fibres converging to them from the cerebral convolutions, Avhich have been not unaptly called " the nerves of the internal senses ;" and may hence be automatic in the first case, and voluntary in the second. For example, in the act of Coughing, we have the very same combination and succession of diverse but mutually related actions, whether the operation be excited by the presence of an irritating particle in the air-passages, or be per- formed as the consequence of a voluntary effort. And a little attention to his own consciousness will satisfy the reader, that as regards the selection and co-ordination of the movements which are concerned, the 526 OF THE NERVOUS SYSTEM AND ITS ACTIONS. Intelligence and Will are no more concerned in the one case than in the other.—And hence it follows, that all the movements which are per- formed by the instrumentality of the Cerebro-spinal system of ganglia and nerves, are in their essential nature automatic; and that their cha- racter as Reflex, Instinctive, Emotional, or Voluntary, is entirely dependent on the nature and seat of the impulses which respectively originate them. 924. There are various conditions, some of them natural, others morbid, in which the distinctness of the functions of the Cerebral Hemi- spheres is well marked. Thus in profound sleep they seem to be entirely dormant; the Spinal Cord and Medulla Oblongata, by which the necessary reflex actions are carried on, being alone in a state of activity. In this condition, the Sensory ganglia also appear to be in a torpid state; but in less profound sleep, actions are often performed, which may be referred to the consensual group,—being such as the sen- sation would immediately prompt, without any reflection, and not being remembered in the waking state. Thus we turn in our beds, under the influence of an uneasy sensation ; or we give some sign of recognition when our names are called. The first of these appears to be a purely consensual movement, being as automatic as if it Avere a reflex action; the other seems to have become as automatic by the influence of habit, and to belong to that class of secondarily automatic actions, in which the movement, though at first directed by the will, has become, after very frequent performance, so closely associated with the guiding sug- gestion, as to be called forth by it alone (§ 904).—In the Coma of Apo- plexy, Narcotic Poisoning, &c, we witness the same gradations as in ordinary sleep. When it is least profound, it seems to affect the Cere- bral hemispheres alone ; the Sensory Ganglia being still, in some degree, open to the reception of impressions. When complete, hoAvever, none but reflex actions can be excited; and if it advance to a fatal termina- tion, it does so by the supervention of the same state of torpidity in the Medulla Oblongata, whereby the respiratory movements are brought to a close. These movements do not cease until the power of deglutition has been lost, and until the eye ceases to close, when the edge of the lid is irritated; but when this is the case, a fatal termination may be apprehended, as it is thus shown that the torpor is extending to the Spinal system of Nerves.—In the condition of Dreaming, it would seem as if the Cerebrum were partially active; a train of thought being sug- gested, frequently by sensations from without; which is carried on with- out any controlling or directing power on the part of the Mind; and which is not corrected, or is only modified in a limited degree, by the knowledge acquired by experience. This condition is still more remark- able in Somnambulism, or (as it has been better termed) Sleep-walking; in which the dreams are not only acted, but may be often acted on with the utmost facility,—a suggestion conveyed through any of the senses excepting sight (which is usually in abeyance) being apprehended and followed-up Avith the utmost readiness, and, in like manner, with little or no correction from experience. Between this condition, and that of ordinary dreaming, on the one hand, and that of complete insensibility on the other, there is every shade of variety; which is presented FUNCTIONS OF THE SYMPATHETIC SYSTEM. 527 by different individuals, or by the same individual at different times. The Cerebellum, in the Sleep-walking state, seems to be frequently in a condition of peculiar activity; a remarkable power of balancing and combining the movements of the body, being often exhibited. 925. On the other hand, there may be an undue exaltation or a per- version of mental activity, without any affection of the sensorial appa- ratus. This is well seen, for example, in the first stage of Alcoholic excitement, and in that of Mania, Phrenitis, and other disorders in which the Cerebral Hemispheres are especially affected. Frequently, as in the case of Alcohol, Opium, Haschish, &c, we may directly attri- bute the morbid action of the Cerebrum to the presence of a poison in the blood which permeates it; and there is strong reason to believe that many other forms of delirium are partly due to a perverted state of that fluid. On the other hand, there can be no doubt that an extreme depression of intellectual power, as well as of the emotional state, is often to be attributed to a depravation of the blood; a slight accumula- tion of bile being very prone to occasion this state in some individuals, and an entire change being effected by a mild dose of mercurial prepa- rations, which, by eliminating the bile, restores the circulating fluid to its proper purity. And it may be fairly suspected, that the foul atmo- sphere in the midst of which a large class of our population habitually lives, has the effect, by keeping their blood charged Avith noxious mat- ters, of so perverting the actions of the brain, that neither the intellec- tual powers nor the moral sense can be duly exercised; and thus it may be anticipated that Sanitary Reform will largely benefit not merely the corporeal but the mental and moral health of those, Avhose position is at present one of fearful degradation from the want of it. 8. Functions of the Sympathetic System. 926. The Cerebro-Spinal apparatus, of which the several parts have now been described, is not the only system of ganglia and nerve-trunks, that is contained within the body of a Vertebrated animal. There is another system, having its own set of centres, and its own distribution of branches, characterized also by a peculiarity in the nature of the nervous fibres of which its trunks are composed, and communicating at numerous points with the preceding. It will be remembered that, in front of the vertebral column, there is a series of ganglia on each side ; communicating, on the one hand, Avith the spinal nerves, as they issue from the vertebral canal; and also connecting themselves with the two large Semilunar ganglia, which lie amidst the abdominal viscera; as well as Avith a series of ganglia, that is found near the base of the heart. In the head, also, there are numerous scattered ganglia, which evidently belong to the same system ; having several communications with the cephalic nerves ; and being also connected Avith the chain of ganglia in the neck. The branches proceeding from this series of ganglia are dis- tributed, not to the skin and muscles (like those of the cerebro-spinal system), but to the organs of digestion and secretion, to the heart and lungs, and particularly to the Avails of the blood-vessels, on Avhich they form a plexus whose branches probably accompany their minutest rami- 528 OF THE NERVOUS SYSTEM AND ITS ACTIONS. fications. The peculiar connexion of this system of nerves with the organs of vegetative life, has caused it to receive the designation of the Nervous System of Organic Life; the Cerebro-Spinal system being termed the Nervous System of Animal Life. It is also not unfrequently termed the ganglionic system; on account of the separation of its centres into scattered ganglia, which forms a striking contrast to the concentra- tion that is so evident in the Cerebro-spinal system. But this term is objectionable, as leading to a supposed analogy between this system and the general nervous system of Invertebrata, whose centres are equally scattered;—an analogy which is completely erroneous, since as we have seen, this last is chiefly the representative of the Cerebro-Spinal system of Vertebrated animals. The term Sympathetic is perhaps the best; although it must not be supposed that this system of nerves is the instrument of by any means all the sympathies, which manifest them- selves between different organs. 927. The sympathetic system contains both classes of nervous fibres; —the ordinary white tubular fibres, all of which are probably derived from the Cerebro-Spinal system ; and the gray or gelatinous fibres, part of which seem to belong to itself (§ 375). Thus we may consider each system as intermingling itself with the other ;—'the Cerebro-Spinal sys- tem transmitting some of its fibres, both motor and sensory, into the Sympathetic;—whilst the Sympathetic is represented in the Cerebro- Spinal system, by certain fibres and collections of vesicular matter of its own. The trunks that proceed from the Semilunar ganglia, are almost entirely composed of gray or organic fibres; whence it is evident that these ganglia are to be regarded as the true centres of the Sympa- thetic system. On the other hand, the trunks which issue from the chain of spinal ganglia, contain a large admixture of white or tubular fibres. 928. The Sympathetic nerves possess a certain degree of power of exciting Muscular contractions, in the various parts to which they are distributed. Thus by irritating them, immediately after the death of an animal, contractions may be excited in any part of the alimentary canal, from the pharynx to the rectum, according to the trunks which are irritated,—in the heart, after its ordinary movements have ceased, —in the aorta, vena cava, and thoracic duct,—in the ductus choledochus, uterus, fallopian tubes, vas deferens, and vesiculse seminales. But the very same contractions may be excited, by irritating the roots of those Spinal nerves, from which the Sympathetic trunks receive their white fibres; and there is, consequently, strong reason to believe that the motor power of the latter is entirely dependent upon the Cerebro-spinal system. Whatever sensory endowments the Sympathetic trunks pos- sess, are probably to be referred to the same connexion. In the ordi- nary condition of the body, these are not manifested. The parts ex- clusively supplied by Sympathetic trunks do not appear to be in the least degree sensible; and no sign of pain is given when the Sympa- thetic trunks themselves are irritated. But in certain diseased conditions of those organs, violent pains are felt in them; and these pains can only be produced through the medium of fibres communicating with the sensorium through the spinal nerves. GENERAL AND SPECIAL SENSATIONS. 529 929. It is difficult to speak with any precision, as to the functions of the Sympathetic system. There is much reason to believe, how- ever,^ that it constitutes the channel through which the passions and emotions of the mind affect the Organic functions ; and this especially through its power of regulating the calibre of the arteries. We have examples of the influence of these states upon the Circulation, in the palpitation of the heart which is produced by an agitated state of feel- ing ; in the Syncope, or suspension of the heart's action, which some- times comes on from a sudden shock; in the acts of blushing and turning pale, which consists in the dilatation or contraction of the small arteries ; in the sudden increase of the salivary, lachrymal, and mam- mary secretions, under the influence of particular states of mind, which increase is probably due to the temporary dilatation of the arteries that supply the glands, as in the act of blushing; and in many other phe- nomena. It is probable that the Sympathetic system not only thus brings the Organic functions into relation with the Animal, but that it also tends to harmonize the former with each other, so as to bring the various acts of secretion, nutrition, &c, into mutual conformity. For whilst the quantity of a secreted product, or the amount of tissue gene- rated in a part, may be affected by an increase or diminution in the calibre of the vessels supplying it, the quality of the secretion, or the character of the tissue, may be likewise affected (there seems valid reason to believe) by that Nervous force, whose relations to the Physical and Chemical, as well as to all other Vital forces, are so intimate (§ 396). CHAPTER XIII. OF SENSATION, GENERAL AND SPECIAL. 1. Of Sensation in general. 930. All beings of a truly Animal Nature possess (there is good reason to believe), a consciousness of their own existence, first derived from a feeling of some of the corporeal changes taking place within themselves; and also a greater or less amount of sensibility to the con- dition of external things. This consciousness of what is taking place within and around the individual, is all derived from impressions made upon its afferent nervous fibres ; which, being conveyed by them to the central sensorium, are there felt (§ 390). Of the mode in which the impression, hitherto a change of a physical character, is there made to act upon the mind, we are absolutely ignorant; we only know the fact. Although we commonly refer our various sensations to the parts at which the impressions are made,—as, for instance, when we say that we have a pain in the hand, or an ache in the leg,—we really use incorrect language; for, though we may refer our sensations to the 530 OF SENSATION IN GENERAL. parts where the impression is first made on the nerves, they are really felt in the brain. This is evident from two facts;—first, that if the nervous communication between the part and the brain be interrupted, no impressions, however violent, can make themselves felt; and second, that if the trunk of the nerve be irritated or pinched, anywhere in its course, the pain which is felt is referred, not to the point injured, but to the surface to Avhich these nerves are distributed. Hence the well- known fact that, for some time after the amputation of a limb, the patient feels pains, which he refers to the fingers or toes that have been removed; this continues until the irritation of the cut extremities of the nervous trunks has subsided. 931. It would seem probable that, among the lower tribes of Ani- mals, there exists no other kind of sensibility, than that termed general or common ; which pervades, in a greater or less degree, nearly every part of the bodies of the higher. It is by this, that we feel those im- pressions, made upon our bodies by the objects around us, which pro- duce the various modifications of pain, the sense of contact or resistance, the sense of variations of temperature, and others of a similar character. From what was formerly stated (§ 403) of the dependence of the im- pressibility of the sensory nerves, upon the activity of the circulation in the neighborhood of their extremities, it is obvious that no parts destitute of blood-vessels can receive such impressions, or (in common language) can possess sensibility. Accordingly Ave find that the hair, nails, teeth, cartilages, and other parts that are altogether extra-vascu- lar, are themselves destitute of sensibility; although certain parts con- nected with them, such as the bulb of the hair, or the vascular membrane lining the pulp-cavity of the tooth, may be acutely sensitive. Again, in tendons, ligaments, fibro-cartilages, bones, &c, whose substance con- tains very few vessels, there is but a very Ioav amount of sensibility. On the other hand, the skin and other parts, which are peculiarly adapted to receive such impressions, are extremely vascular; and it is interesting to observe, that some of the tissues just mentioned become acutely sensible, when new vessels form in them in consequence of diseased action. It does not necessarily folloAV, hoAvever, that parts should be sensible in a degree proportional to the amount of blood they may contain; for this blood may be sent to them for other purposes, and they may contain but a small number of sensory nerves. Thus, although it is a condition necessary to the action of Muscles, that they should be copiously supplied Avith blood (§ 359), they are by no means acutely sensible; and, in like manner, Glands, which receive a large amount of blood for their peculiar purposes, are far from possessing a high degree of sensibility. 932. But besides the general or common sensibility, which is dif- fused over the greater part of the body, in most animals, there are cer- tain parts, which are endowed with the property of receiving impres- sions of a peculiar or special kind, such as sounds or odors, that would have no influence on the rest; and the sensations which these excite, being of a kind very different from those already mentioned, arouse ideas in our minds, which we should never have gained without them. Thus, although we can acquire a knowledge of the shape and position GENERAL AND SPECIAL SENSATIONS. 531 of objects by the touch, we could form no notion of their color with- out sight, of their sounds without hearing, or of their odors without Bmell. The nerves Avhich convey these special impressions, as already mentioned, are not able to receive those of a common kind; thus the eye, however well fitted for seeing, would not feel the touch of the finger, if it were not supplied by branches from the Fifth pair, as well as by the Optic. Nor can the different nerves of special sensation be affected by impressions that are adapted to operate on others; thus the ear cannot distinguish the slightest difference between a luminous and a dark object; nor could the eye distinguish a sounding body from a silent one, except when the vibrations can be seen. But Electricity possesses the remarkable power, when transmitted along the several nerves of special sense, of exciting the sensations peculiar to each; and thus, by proper management, this single agent may be made to produce flashes of light, distinct sounds, a phosphoric odor, a peculiar taste, and a pricking feeling, in the same individual, at one time. Each kind of sensation may also be excited, however, by mechanical irritation of the nerve which is subservient to it.—The feeling of pain may be in- duced by impressions made upon the nerves of special sense, as Avell as upon those of feeling, if these impressions be too violent or excessive. Thus the dazzling of the eye by a strong light, and still more, the action of a moderate light in an irritable state of the retina,—sudden loud sounds, or even sounds of moderate intensity but of peculiar harsh- ness,—powerful odors, or even such as are agreeable in moderation,— produce feelings of uneasiness, which may be properly called painful, even though they are different from those excited through the nerves of common sensation. 933. As a general rule, it may be stated, that the violent excite- ment of any sensation is disagreeable; even when the same sensation, experienced in a moderate degree, may be a source of extreme pleasure. But the question of degree is relative rather than absolute: that is, a sensation may be felt as extremely violent by one individual, whilst another, who is more accustomed to sensations of the same kind, is not disagreeably affected by it. Thus our sensations of heat and cold are entirely governed by the previous condition of the parts affected; as is shown by the well-known experiment of putting one hand in hot water, the other in cold, and then transferring them both to tepid water,— which will seem cool to the one hand and warm to the other. The same is the case in regard to light and sound, smell and taste. A per- son going out of a totally dark room, into one moderately bright, is for the time painfully impressed by the light, but soon becomes habituated to it; whilst another, who enters it from a room brilliantly illuminated, will consider it dark and gloomy. . 934. The intensity with Avhich sensations are felt, therefore, depends upon the degree of change which they produce in the sensorium. The more frequent the recurrence of any particular sensation, the more does the system become adapted to it, and the less change does it pro- duce. It is, therefore, perceived in a less and less degree, and at last it ceases to excite attention. The stoppage of a constantly-recurring sensation, however, will produce a change, which makes as strong an 532 OF SENSATION IN GENERAL. impression on the system as its first commencement; thus there are persons, who have become so habituated to the sound of a waterfall or even of a forge-hammer, that they cannot sleep anywhere but in its vicinity; and it is well known that, when a person has gone to sleep under the influence of some continuous or frequently-recurring sound (such as the voice of a reader, the dropping of water, the tread of a sentinel, &c), the cessation of the sound will cause his awaking. 935. The acuteness of particular sensations is influenced in a re- markable degree by the attention they receive from the mind. If the mind be entirely inactive, as in profound sleep, no sensation whatever is produced by very feeble impressions; on the other hand, when the mind is from any cause strongly directed upon them, impressions very feeble in themselves produce sensations of even painful acuteness. It is in this manner, that the habit of attending to sensations of any par- ticular class increases their vividness; so that they are at once per- ceived by an individual on the watch for them, when they do not ex- cite the observation of others. We may even, by a strong effort, direct the mind into one particular channel, so as to receive only those sensa- tions which have reference to it, and to be unconscious quoad all others. Thus, the application of the mind to some particular train of thought may prevent our being conscious of anything that is going on around or within us,—the conversation of friends,—the striking of the clock,— the calls of hunger, &c. This abstraction may be altogether voluntary; and the possession of the power of thus withdrawing the mind at will from the influence of external disturbing causes, and of fixing it upon any particular train of ideas, is an extremely valuable one. But it may also be involuntary, and may be a source of inconvenience from its tendency to recur at improper times,—producing the habitual state which is known as absence of mind or reverie. 936. It is desirable that we should make a distinction, between the sensations themselves, and the ideas which are the immediate results of those sensations, when they are perceived by the mind. These ideas relate to the cause of the sensation, or the object by which the impres- sion is made. Thus, the formation of the picture of an object, upon the retina, produces a certain impression upon the optic nerve; which, being conveyed to the sensorium, excites a corresponding sensation, with which, in all ordinary cases, we immediately connect an idea of the nature of the object. So closely, indeed, is this idea usually re- lated to the sensation, that we are not in the habit of making a distinc- tion between them. Thus, I may say at this moment, " I see a book on the table before me;" the fact being, that I am conscious of a certain picture, which conveys to my mind the ideas of a book and of a table, and of their relative positions; these ideas being (in Man) the result of experience and associations,—in fact, originating in the immediate application of the knowledge we have previously acquired, that a cer- tain object, whose picture we see, is a book, another object a table, and so on. We are liable to be deceived on this assumption; as when, by a clever imitation, a picture on a plane surface is made to represent an object in relief, so perfectly as at once to excite the idea of the latter, INTUITIVE AND ACQUIRED PERCEPTIONS. 533 —which may not be corrected, until we have ascertained by the touch, the flatness of the real object. 937. This production of ideas, by the agency of sensations, is a pro- cess altogether mental, and dependent upon the laws of Mind. We find that some of these perceptions or elementary notions are intuitive: that is, they are prior to all experience, and are as necessarily connected with the sensation which produces them, as reflex movements are with the impression that excites them. This seems to be the case, for example, with regard to erect vision. There is no reason whatever to think, that either infants or any of the lower animals see objects in an inverted position, until they have corrected their notion by the touch; for there is no reason why the inverted picture on the retina should give rise to the idea of the inversion of the object. The picture is so received by the mind, as to convey to us an idea of the position of external objects, which harmonizes with the ideas we derive through the touch; and whilst we are in such complete ignorance of the manner in which the mind becomes conscious of the sensation at all, we need not feel any difficulty about the mode in which this conformity is effected. But in Man, as already stated, the attaching definite ideas to certain groups of lines, colors, &c, with respect to the objects they represent, is a subsequent process, in Avhich experience and memory are essentially concerned; as Ave see particularly well, in cases presently to be referred to, in which the sense of sight has been acquired comparatively late in life, and in which the mode of using it, and of connecting the sensa- tions received through it with those received through the touch, has had to be learned, by a long-continued training. The elementary notions thus formed,—which may, by long habit, present themselves as immediately and unquestionably, as if they were intuitive,—are termed acquired perceptions. 938. It is probable that, among the lower animals, the proportion of intuitive perceptions is much greater than in Man; whilst, on the other hand, his power of acquiring perceptions is much greater than theirs. So that, whilst the young of the lower animals very soon becomes pos- sessed of all the knowledge which is necessary for the acquirement of its food, the construction of its habitation, &c, its range is very limited, and it is incapable of attaching any ideas to a great variety of objects, of which the Human mind takes cognizance. This correspondence between the acquired perceptions of Man, and the intuitive perceptions of many of the loAver animals, is strikingly evident in regard to the power of measuring distance. This is acquired very gradually by the Human infant, or by a person who has first obtained the faculty of sight later in life ; but it is obviously possessed by many of the lower animals, to Avhose maintenance it is essential, immediately upon their "entrance into the world. Thus, a Fly-catcher, immediately after its exit from the egg, has been known to peck at and capture an insect,— an action which requires a very exact appreciation of distance, as well as a power of precisely regulating the muscular movements in accordance with it. 534 OF SENSATION. 2. Of the Sense of Touch. 939. By the sense of Touch is usually understood that modification of the common sensibility of the body, of which the surface of the skin is the especial seat, but which exists also in some of its internal reflexions. In some animals, as in Man, nearly the whole exterior of the body is endowed with it, in no inconsiderable degree; whilst in others, as the greater number of Mammalia, most Birds, Reptiles, and Fishes, and a large proportion of the Invertebrata, the greater part of the body is so covered with hairs, scales, bony or horny plates, shells of various kinds, complete horny envelopes, &c, as to be nearly insensible; and the faculty is restricted to particular portions of the surface, or to organs projecting from it, which often possess a peculiarly high degree of this endowment. Even in Man, the acuteness of the sensibility of the cutaneous surface varies greatly in different parts, being greatest at the extremities of the fingers and in the lips, and least in the skin of the trunk, arm, and thigh. Thus the two points of a pair of compasses (rendered blunt by bits of cork) can be separately distinguished by the point of the middle finger, when approximated so closely as l-3d of a line; whilst they require to be opened so widely as 30 lines from each other, to be separately distinguished, when pressed upon the skin over the spine, or upon that of the middle of the arm or thigh. 940. The impressions that produce the sense of touch are received through the sensory papillar, with which the surface of the true Skin is beset,—more or less closely, according to the part of it that is examined. These papillee are minute elevations, which enclose loops of capillary vessels (Fig. 157), and branches of the sensory nerves. With regard Fig. 157. Capillary network at margin of lips. to the precise course of the latter, there is some uncertainty; but it is probable, from analogy, that the representation given of them by Gerber (Fig. 158) is in the main correct; and that each loop of the Sensory nerve is connected with one or more vesicular foci, on some change in which the formation of the sensory impression is immediately dependent (§ 382). It is peculiar to the sense of Touch, and to that of Taste (which is closely related to it) that the impression must be made by the contact of the object itself with the sensory surface, and not through any intermediate agency. The only exception to this is in regard to the sense of Temperature, which seems to be in many respects different from ordinary touch; here the proximity of the warm SENSE OF TOUCH. 535 or cold body is sufficient, the impressions being made after the manner of those of odors, sounds, &c. It is worth remarking, with reference to the question of the special nature of the sensory fibres, which are the Fig. 158. Distribution of the tactile nerves at the extremity of the Human Thumb, as seen in a thin perpendicular section of the skin. channel of these impressions, that no mechanical irritation of the nerves of common sensation ever seems to excite sensations of heat or cold; these being apparently almost as distinct from the sense of contact as they are from that of light or sound. 941. The only idea communicated to our minds, when this sense is exercised in its simplest form, is that of resistance; and we cannot acquire a notion of the size or shape of an object, or of the nature of its surface, through this sense alone, unless we move the object over our own sensory organ, or pass the latter over the former. By the various degrees of resistance which we then encounter, we form our estimate of the hardness or softness of the body. By the impressions made upon our sensory papillae, when they are passed over its surface, we form our idea of its smoothness or roughness. But it is through the muscular sense which renders us cognizant of the relative position of the fingers, the amount of movement the hand has performed in passing over the object, and of other impressions of like nature, that we acquire our notions of the size and figure of the object; and hence we perceive that the sense of touch, without the power of giving motion to the tactile organ, would haA'e been of comparatively little use. It is chiefly in the variety of movements of which the hand of Man is capable,—thus con- ducive as they are, not merely to his prehensile powers, but to the exercise of his sensory endowments,—that it is superior to that of any other animal; and it cannot be doubted that this affords us a very important means of acquiring information in regard to the external world, and especially of correcting many vague and fallacious notions which Ave should derive from the sense of Sight, if used alone. On the other hand, it must be evident that our knowledge would have but a very limited range, if this sense were the only medium through which we'could acquire ideas. Of this Ave have the clearest evidence in the very imperfect development of the mental powers in those unfortunate persons who have suffered under the deprivation of sight and hearing from their birth, and who have been consequently cut off from the most direct means of profiting by the knoAvledge possessed by their fellow- beings, through want of poAver to use the organs of speech. It is only 536 OF SENSATION. where such individuals have fallen under the care of judicious and perse- vering instructors, that their mental powers have been called into their due activity, or that any ideas have been awakened, beyond those imme- diately connected with the gratification of the animal wants, or with painful or pleasurable sensations. Thus a mind quite capable of being aroused to activity and enjoyment, may remain in a condition nearly allied to that of idiocy, simply for want of the sensations requisite to produce ideas of a higher and more abstract character than those de- rived through the sense of Touch, Taste, and Smell. 942. For the exercise of the sense of Temperature, the integrity of the sensory apparatus contained in the Skin appears to be requisite; for it has been ascertained by the recent experiments of Prof. Weber, that if the integuments be removed, the application of hot or cold bodies only causes pain, their elevation or depression of temperature not being perceived ; and the same is the case when hot or cold bodies are applied to the nerve-trunks. It is worthy of note that there are many cases on record in which the sense of Temperature has been lost, while the ordinary tactile sense remained; and the former is sometimes preserved when there is a complete loss of every other kind of sensi- bility. So again we find that the subjective sensations of temperature (that is, sensations which originate from changes in the body itself, not from external impressions) are frequently excited quite independently of the tactual sensations; a person being sensible of heat or of chilli- ness in some part of his body, without any real alteration of its tempera- ture, and without any corresponding affection of the tactual sensations. It is curious that the intensity of the sensation of temperature should depend, not merely upon the relative degree of heat to which the part is exposed (§ 933), but also upon the extent of the surface over which it is applied ; a weaker impression made on a larger surface seeming more powerful than a stronger impression made on a small surface. Thus, if the forefinger of one hand be immersed in water at 104°, and the Avhole of the other hand be plunged in water at 102°, the cooler water will be thought the warmer; whence the well-known fact that water in which a finger can be held without discomfort, will produce a scalding sensation when the entire hand is immersed in it. 3. Of the Sense of Taste. 943. The sense of Taste, like that of Touch, is excited by the direct contact of particular substances with certain parts of the body; but it is of a much more refined nature than touch, inasmuch as it communi- cates to us a knowledge of properties which that sense would not reveal to us. All substances, however, do not make an impression on the organ of Taste. Some have a strong savor, others a slight one, and others are altogether insipid. The cause of these differences is not altogether understood; but it may be remarked, that, in general, bodies which cannot be dissolved in water, alcohol, &c, and which thus cannot be presented to the gustative papillae in a state of solution, have no taste. This sense has for its chief purpose to direct animals in their choice of food; hence its organ is always placed at the entrance to the SENSE OF TASTE. 537 digestive canal. In higher animals, the Tongue is the principal seat of it; but other parts of the mouth are also capable of receiving the impression of certain savors. The mucous membrane which covers the tongue is Copiously supplied with papillae, of various forms and sizes. Those of simplest structure closely resemble the cutaneous papilla ; but there are others, which resemble clusters of such papillae, each being composed of a fasciculus of looped capillaries (Fig. 159), with a bundle Fig. 159. Capillary network of fungiform papillae of the Tongue. of nerve-fibres, whose precise mode of termination it has not yet been found possible to ascertain. These fungiform papillae, which are covered with a very thin epithelium, are probably the special instru- ments of the sense of taste ; for the exercise of which it seems probable that the sapid substances must penetrate, in solution, to the interior of the papilla. When these papillae are called into action by the contact of substances having a strong savor, they not unfrequently become very turgid, by a distension of their vessels analogoas to that which occurs in erection ; and they rise up from the surface of the mucous membrane, so as to produce a decided roughness of its surface. The conical papillae, on the other hand, are furnished with thick epithelial investments, which are sometimes prolonged into filamentous appen- dages ; and, looking to their higher development among other animals, and the offices to Avhich they are there subservient, it seems probable that their functions are purely mechanical, and that they serve especially to cleanse the teeth from adhering particles. The nerve-fibres can be seen to form distinct loops in their interior, at some distance from the apex. 944. There has been much discrepancy of opinion as to the nerve which is specially concerned in the sense of Taste. The tongue of Man is supplied by two sensory nerves : the lingual branch of the Fifth pair; and the Glosso-pharyngeal. The former chiefly supplies the upper sur- face of the front of the tongue, and is copiously distributed to the papillae near the tip. The latter is mostly distributed upon the mucous surface of the Fauces, and upon the back of the tongue ; but it sends a branch fonvards, beneath the lateral margin on eaeh side, which supplies the edges and inferior surface of the tip of the tongue, and inosculates with the preceding. There is reason to believe, from experiment, that the gustative sensibility of the tongue is not destroyed by section of either of these nerves; though the operation produces a total or partial loss of sensibility over certain parts of the surface. There seems good reason to conclude, that the lingual branch of the Fifth pair is the 538 OF SENSATION. nerve, through which the sense of Taste, as well as that of Touch, is exercised, in the parts of the tongue to which it is specially distributed, —which are those that possess both senses in the most acute degree; and that the Glosso-pharyngeal is subservient to the same functions in the parts supplied by it, being probably the exclusive channel, also, through wdiich the impressions made by disagreeable substances taken into the mouth, are propagated to the Medulla Oblongata, so as to pro- duce nausea and excite efforts to vomit. The latter nerve is also, as we have seen, the principal channel of the impressions that give rise to the reflex act of swallowing ; with which the fifth pair is concerned in a much inferior degree (§ 897). 945. A considerable part of the impression produced by many sub- stances taken into the mouth, is received through the sense of Smell, rather than through that of Taste. Of this, any one may easily satisfy himself, by closing the nostrils, and breathing through the mouth only, whilst holding in his mouth, or even rubbing between his tongue and his palate, some aromatic substance ; its taste is then scarcely recognised, although it is immediately perceived when the nasal passages are re- opened, and its effluvia are drawn into them. There are many sub- stances, however, which have no aromatic or volatile character; and whose taste, though not in the least dependent upon the action of the nose, is nevertheless of a poAverful character. Some of these produce, by irritating the mucous membrane, a sense of pungency, allied to that which the same substances (mustard, for instance) will produce, when applied to the skin for a sufficient length of time, especially if the Epi- dermis have been removed. Such sensations, therefore, are evidently of the same kind Avith those of Touch, differing from them only in the degree of sensibility of the organ through which they are received. But there are others which produce sensations entirely different from any that can be received through the skin, and which are properly distin- guished, therefore, as gustative ; such are common Salt, which may be considered as a type of the saline taste, Sugar, the type of the saccharine, Quinine of the bitter, and Tannin of the astringent, and Citric acid of the sour. All such substances, therefore, are said to possess sapid pro- perties, exciting distinctive tastes, quite irrespectively of any aromatic or odoriferous properties which they may also possess, as well as of their stimulating action on the skin. 4. Of the Sense of Smell. 946. Certain bodies possess the property of exciting sensations of a peculiar nature, which cannot be perceived by the organs of taste or touch, but which seem to depend upon the diffusion of the particles of the substance through* the surrounding air, in a state of extreme minute- ness. As the solubility of a substance in liquid seems a necessary con- dition of its exciting the sense of Taste, so does its volatility, or tendency to a vaporous state, appear requisite for its having Odorous properties. Most volatile substances are more or less odorous : whilst those which do not readily transform themselves into vapor, usually possess little or no fragrance in the liquid or solid state, but acquire strong odorous SENSE OF SMELL. 539 properties, as soon as they are converted into vapor,—by the aid of heat for example. There are some solid substances, which possess very strong odorous properties, without losing weight in any appreciable degree by the diffusion of their particles through the air. This is the case, for example, with Musk; a grain of which has been kept freely exposed to the air of a room, whose doors and windows were constantly open, for a period of ten years ; during which time the air, thus con- tinually changed, was completely impregnated with the odor of musk; and yet, at the end of that time, the particle was not found to have per- ceptibly diminished in weight. We can only attribute this result to the extreme minuteness of the division of the odorous particles of this sub- stance. There are other odorous solids, such as Camphor, which rapidly lose weight by the loss of particles from their surface, when freely exposed to the air. 947. The conditions of the sense of Smell are very simple. The Ol- factory nerve is minutely distributed over the Schneiderian membrane, which is itself highly vascular. The arrangement of the ultimate fibres of this nerve has not been ascertained. The Schneiderian membrane is kept constantly but moderately moist, by a mucous secretion from its surface ; and this condition is essential to the acute perception of odors. If the mucous surface be too dry, as happens Avhen the fifth pair is Fig. 160. septum nasi. paralysed, the sensation is blunted, or even destroyed; and the same effect is produced by the presence of too copious a secretion, as when 540 OF SENSATION. we are suffering under an ordinary cold.—The highest part of the nasal fossae appears to be that, in which there is the most acute sensibility to odors; and hence it is, that, when we snuff the air, so as to direct it into this portion of the cavity, we perceive delicate odors, which would otherwise have escaped us. The acuteness of the sense of Smell depends in no small degree, upon the extent of surface exposed by the membrane lining the nasal cavity; and in this respect Man is far surpassed by many of the lower Mammalia, especially the Ruminants, which are warned by its means of the proximity of their enemies. The habit of attention to sensory impressions of this class, however, very much heightens their acuteness: hence in those who suffer under blindness and deafness conjointly, it is usually the principal means by which indivi- duals are distinguished, and the presence of strangers recognised; and there are cases, in which individuals in a state of somnambulism have exhibited a degree of acuteness of smell, quite comparable to that which is characteristic of Deer, Antelopes, &c. 948. Besides ministering to the sense of Smell, by stimulating the secreting powers of its surface, the Fifth pair has another very impor- tant function,—that of endowing the interior of the nose with common sensibility, and thus receiving the impression produced by acrid or pun- gent substances, which act upon it in the same way as they do upon the tongue. Such substances are felt, by the irritation they produce, rather than smell; and the sensation they occasion gives rise to the consensual act of sneezing, by which a violent blast of air is directed through the nasal passages, in such a manner as to clear them of the irritating mat- ter, whether solid (as snuff), fluid or gaseous. Hence this action may be excited by the contact of an irritant with the Schneiderian mem- brane, after the olfactory nerve has been divided, if the branches of the fifth pair be entire; whilst it does not take place when the fifth pair is paralysed, even though the sense of smell is retained. 5. Of the Sense of Hearing. 949. By this sense we become acquainted with the sounds produced by bodies in a certain state of vibration ; the vibrations being propagated through the surrounding medium, by the corresponding waves or undu- lations Avhich they produce in it. Although air is the usual medium through which sound is propagated, yet liquids or solids may answer the same purpose. On the other hand, no sound can be propagated through a perfect vacuum.—It is a fact of much importance, in regard to the action of the Organ of Hearing, that sonorous vibrations which have been excited, and are being transmitted, in a medium of one kind, are not imparted with the same readiness to others. The following conclu- sions have been drawn from experimental inquiries on this subject. I. Vibrations excited in solid bodies, may be transmitted to water without much loss of their intensity; although not with the same readi- ness that they would be communicated to another solid. II. On the other hand, vibrations excited in water lose something of their intensity in being propagated to solids; but they are returned, as it were, by these solids to the liquid, so that the sound is more loudly SENSE OF HEARING. 541 heard in the neighborhood of these bodies, than it would otherwise have been. III. The sonorous vibrations are much more weakened in the trans- mission of solids to air; and those of air make but little impression on solids. IV. Sonorous vibrations in water are transmitted but feebly to air; and those which are taking place in air are with difficulty communicated to water ; but the communication is rendered more easy by the inter- vention of a membrane extended between them. The application of these conclusions, in the Physiology of Hearing, will be presently apparent. 950. It is on the Auditory nerve (commonly termed the Portio Mollis of the 7th pair), that the sonorous undulations make their im- pression ; but we invariably find, that this impression is made through the medium of a liquid, contained in a cavity, on the walls of which the ultimate branches of this nerve are distributed. The simplest form of the organ of Hearing, such as we find in Cephalopods and in certain Fishes, consists merely of a cavity excavated in the solid framework of the head; which cavity is filled with liquid, and lined by a membrane on Avhich the auditory nerve is distributed. These animals are inhabi- tants of the water ; and the sonorous vibrations excited in this medium being communicated to the solid parts of the head, will be by them again transmitted to the contained fluid, without much diminution of their intensity ; according to principles I. and II.—In certain Crustacea, however, whose organ of hearing is contained in the base of the anten- nae, as well as in most Fishes, we find the auditory cavity or vestibule no longer entirely closed; but having an aperture on its external side, which is covered in by a membrane. Here the vibrations of the liquid within the cavity will be more directly excited by those of the sur- rounding medium, for if this be water, it will propagate its undulations into the cavity, with little interruption from the membrane stretched across its mouth ; whilst, if it be air, the interposition of this very mem- brane will greatly assist in the transmission of the vibrations to the water of the auditory cavity, according to principle IV. In most of the animals which have the organ of hearing constructed upon this simple plan, the force of the vibrations of the fluid within the cavity is increased by several minute stony concretions (termed otolithes), which are sus- pended in it. These act according to principle II. Some traces of them are found in the higher animals ; in which they are for the most part superseded, however, by an apparatus better adapted to augment the intensity of the sonorous vibrations. 951. This apparatus consists, in all Vertebrated animals above the inferior Reptiles, of the tympanum or drum, with its membrane and chain of bones; together Avith, in the mammalia, the external ear; which is adapted to direct itself, more or less completely, towards the point from which the sonorous vibrations proceed, and to give them a degree of preliminary concentration. The tympanic apparatus is interposed betAveen the external ear and the membrane covering the foramen ovale, which is the entrance to the real auditory cavity; and its purpose is evidently, to receive the sonorous vibrations from the air, 542 OF SENSATION. and to transmit them to that membrane, in such a manner that the vibrations thus excited in the latter may be much more powerful, than they would be if the air acted immediately upon it, as in the loAver Vertebrata.—The usual condition of the Membrana Tympani appears to be rather lax; and, when in this condition, it vibrates in accordance with grave or deep tones. By the action of the tensor tympani it may i; j| be tightened, so as to vibrate in accordance with sharper or higher | tones ; but it will then be less able to receive the impressions of deeper ; sounds. This state we may easily induce artificially, by holding the breath, and forcing air from the throat into the Eustachian tube, so as to make the membrane bulge out by pressure from within ; or by ex- hausting the cavity by an effort at inspiration, with the mouth and nostrils closed, which will cause the membrane to be pressed inwards by the external air. In either case, the hearing is immediately found to be imperfect; but the deficiency relates only to grave sounds, acute ones being heard even more plainly than before. There is a different limit to the acuteness of the sounds, of which the ear can naturally take cognizance, in different persons. If the sound be so high in pitch, that the membrana tympani cannot vibrate in unison with it, the individual will not hear it, although it may be loud; and it ha3 been noticed, that certain individuals cannot hear the very shrill tones pro- duced by particular Insects, or even Birds, which are distinctly audible to others. 952. Not only do we find the tympanic apparatus superadded, in the higher forms of the organ of Hearing, but also the Semicircular Canals, and the Cochlea.—The former exist in all Vertebrata, save the loAvest Fishes ; and in nearly every case, they are three in number, and lie in three different planes. Hence it has been supposed, with some probability, that they assist in producing the idea of the direction of sounds. The Cochlea does not exist at all in Fishes; and in Reptiles its condition is quite rudimentary. In Birds, this cavity is more com- pletely formed, though the passage is nearly straight instead of spiral; of its real character, however, there can be no doubt, from its being divided, like the cochlea of Man, by a membranous partition, on which the ramifications of the auditory nerve are spread out. This appendage has been supposed to be the organ that enables us to judge of the pitch of sounds; an idea which derives some confirmation from the correspondence between the development of the cochlea in different animals, and the variety in the pitch (or length of the scale) of the sounds which it is important that they should hear distinctly, espe- cially the voices of their own kind.—That the Vestibule with the passages proceeding from it, constitutes the true organ of hearing, even in Man, is evident from the fact, that when (as not unfrequently happens) the tympanic apparatus has been entirely destroyed by disease, so as to reduce the organ to the condition of that in Avhich no such apparatus exists, the faculty of Hearing is by no means abolished, although it is deadened. 953. The faculty of Hearing, like other senses, may be very much increased in acuteness by cultivation; but this improvement depends rather upon the habit of attention to the faintest impressions made SENSE OF HEARING. 543 upon the organ, than upon any change in the organ itself. This habit may be cultivated in regard to sounds of some one particular class; all others, being heard as by an ordinary person. Thus, the watchful North American Indian recognises footsteps, and can even distinguish Fig. 161. A diagram of the Ear:—p. The pina. t. The tympanum. I. The labyrinth. 1. The upper part of the helix. 2. The antihelix. 3. The tragus. 4. The antitragus. 5. The lobulus. 6. The concha. 7. The upper part of the fossa scaphoidea. 8. The meatus. 9. The membrana tympani, divided by the section. 10. The three little bones, crossing the area of the tympanum, malleus, incus, and stapes: the foot of the stapes blocks up the fenestra ovalis upon the inner wall of the tympanum. 11. The promontory. 12. The fenestra rotunda; the dark opening above the ossicula leads into the mastoid cells. 13. The Eustachian tube; the little canal upon this tube contains the tensor tympani muscle in its passage to the tympanum. 1-1. The vestibule. 15. The three semicircular canals, horizontal, perpendicular, and oblique. 16. The ampullae upon tbe perpendicular and horizontal canals. 17. The cochlea. 18. A depression between the convexities of the two tubuli which communicate with the tympanum and vestibule: the one is the scala tympani, terminating at 12; the other is the scala vestibuli. between the tread of friends and foes ; whilst his white companion, who has lived among the busy hum of cities, is unconscious of the slightest sound. Yet the latter may be a musician, capable of dis- tinguishing the tones of all the different instruments in a large orches- tra, of following any one of them through the part which it performs, and of detecting the least discord in the blended effects of the Avhole,— effects which would be to the unsophisticated Indian but an indistinct mass of sound. In the same manner, a person who has lived much in the country, is able to distinguish the note of every species of bird that lends its voice to the general chorus of nature; whilst the inhabi- tant of a town hears only a confused assemblage of shrill sounds, which may impart to him a disagreeable rather than a pleasurable sen- sation. 954. In all continued sounds or tones, there are several points to be attended to. In the first place, we take cognizance of their pitch; which depends upon the number of vibrations in a given time,—the high notes being produced by the most rapid vibrations, and the low notes by the sloAvest. The ear can appreciate tones produced by 24,000 544 OF SENSATION. impulses per second, the pitch of which is about four octaves above the highest F of the piano-forte. On the other hand, no sequence of vibra- tions fewer than 7 or 8 in a second, can produce a continuous tone, because the impression left by each impulse has passed away, before the next succeeds ; and there is consequently nothing more than a succession of distinct beats. The strength or loudness of musical tones depends (other things being equal) on the force and extent of the vibrations, com- municated by the sounding body to the medium which propagates them. This will diminish, however, with distance, which softens loud tones by lowering the intensity of the undulations, as a consequence of their more extensive diffusion. The causes of the difference in the timbre, or quality of musical tones,—such, for instance, as those which exist between the tones of a flute, a violin, a trumpet, and a human voice, all sounding a note of the same pitch, are unknown: but they pro- bably depend upon differences of form in the undulations. Our ideas of the direction and distance of sounds, are for the most part formed by habit. Of the former we probably judge in great degree, by the rela- tive intensity of the impressions received by the two ears; though we may form some notion of it by a single ear, if the idea just stated as to the use of the semicircular canals (§ 952), be correct.—Of the dis- tance of the sounding body, we judge by the intensity of the sound, comparing it with that which we know the same body to produce when nearer to us. The Ear may be deceived in this respect as well as the eye; thus the effect of a full band at a distance may be given by the subdued tones of a concealed orchestra close by us; and the Ventrilo- quist produces his deception, by imitating as closely as possible, not the sounds themselves, but the manner in which they would strike our ears. 6. Of the Sense of Sight. 955. By the faculty of Sight we are made acquainted in the first place, with the existence of Light; and by the medium of that agent we take cognizance of the form, size, color, position, &c, of bodies that transmit or reflect it. As to the mode in which luminous impres- sions are propagated through space, philosophers are at present unde- termined ; and the question is of no physiological importance, since all are agreed as to the laws which regulate their transmission. These laws, which will be found at large in any Treatise on Natural Philoso- phy,* may be briefly stated as follows. I. Light travels in straight lines, so long as the medium through Avhich it passes is of uniform density. II. When the rays of light pass from a rarer medium into a denser one, they are refracted towards a line drawn perpendicular to the sur- face they are entering. III. When the rays of light pass from a denser medium into a rarer one, they are refracted from the perpendicular. IV. When rays proceeding from the several points of a luminous object, at a distance, fall upon a double convex lens, they are brought * See Dr. Golding Bird's Manual, Chap. XXII. SENSE OF SIGHT. 545 to a focus upon the other side of it; in such a manner that an inverted picture of the object is formed upon a screen, placed in the proper posi- tion to receive it. Thus in Fig. 162, a b is the object, and e f the Fig. 162. lens ; the rays issuing from the two extremities and the centre of the object, are brought to a corresponding focus at a less distance on the other side of it, so as to form a distinct picture; but as the rays from A are brought to a focus at d, and those from B at c, the picture will be inverted. V. The further the object is removed from the lens, the nearer will the picture be brought to it, and the smaller will it be. VI. If the screen be not held precisely in the focus of the lens, but a little nearer, or further off, the picture will be indistinct; for the rays which form it will either not have met, or they will have crossed each other. 956. The Eye, in its most perfect form—such as it possesses in Man and the higher animals,—is an optical instrument of wonderful com- pleteness ; designed to form an exact picture of surrounding objects upon the Retina or expanded surface of the Optic nerve, by which the impression is conveyed to the brain. The rays of light, which diverge from the several points of any object, and fall upon the front of the cornea, are refracted by its convex surface, whilst passing through it into the eye, and are made to converge slightly. They are brought more closely together by the crystalline lens, which they reach after passing through the pupil; and its refracting influence, together with that produced by the vitreous humor, is such as to cause the rays, that issued from each point, to meet in a focus on the retina. In this manner, a complete inverted image is formed, as shown in Fig. 163; which represents a vertical section of the eye, and the general course Fig. 163. of the rays in its interior. As in the preceding figure, the rays which issue from the point A are brought to a focus at D; whilst those diverg- ing from b are made to converge upon the retina at c.—The Retina, which is itself so thin as to be nearly transparent, is spread over the 546 OF SENSATION. layer of black pigment, which lines the choroid coat. The purpose of this is evidently to absorb the rays of light that form the picture, imme- diately after they have passed through the retina ; in this manner, they are prevented from being reflected from one part of the interior of the globe to another; which would cause great confusion and indistinctness in the picture. Hence it is that, in those albino individuals (both of the Human race, and among the loAver animals), in whose eyes this pigment is deficient, vision is extremely imperfect, except in a very feeble light; for the vascularity of the choroid and iris is such as to give to these membranes a bright red hue, which enables them power- fully to reflect the light that reaches the interior of the eye, when they are not prevented from doing so by the interposition of the pigmentary layer. 957. The Eye is so constructed, as to avoid certain errors and de- fects, to which all ordinary optical instruments are liable. One of these imperfections, termed spherical aberration, results from the fact, that the rays of light, passing through a convex lens whose curvature is circular, are not all brought to their proper foci, those which have passed through the exterior of the lens being made to converge sooner than those Avhich have traversed its central portion. The result of this imperfection is, that the image is deficient in clearness, unless only the central part of the lens be employed.—The other source of imper- fection is what is termed chromatic aberration ; and it results from the unequal degree in Avhich the differently-colored rays are refracted, so that they are brought to a focus at different points. The violet rays, being the most refrangible, are soonest brought to a focus; and the red being the least refrangible, have their focus at the greatest distance from the lens. Hence it is impossible to obtain an image by an ordi- nary lens, in which the colors of the object are accurately repre- sented ; for the foci of its differently-colored portions will be different; and its white rays will be decomposed, so that the outlines will be sur- rounded by colored fringes.—The Optician is enabled to correct the effects of these aberrations, by combining lenses of different densities and curvatures; so arranged as to correct each other's errors, without neutralizing the refractive power. Thi3 is precisely the plan adopted in the construction of the Eye; which, when perfectly formed, and in a healthy state, forms an accurate picture of the object upon the retina, free from either spherical or chromatic aberration. This is effected by the combination of humors of different densities, having curvatures precisely adapted to the required purpose. 958. There are certain variations, hoAvever, in the conformation of the eye which diminish the perfection of its result. Thus the Cornea may be too convex, and the whole refractive powrer too great; so that the image of an object at a moderate distance is formed in front of the retina, instead of upon it. When this is the case, a distinct image can only be formed, by bringing the object nearer to the eye; the effect of which will be, to throw the picture further back. Such an eye is said to be myopic, or short-sighted; and its imperfection may be corrected by placing a concave lens in front of the cornea, of a curvature adapted to neutralize Avhat is superfluous in the convexity of the latter.—On OF THE EVE AS AN OPTICAL INSTRUMENT. 547 the other hand, if the cornea be too flat, and the refractive power of the humors be too low, the com'ergent rays proceeding from an object at a moderate distance will not meet upon the retina, but behind it (if they were allowed to pass on); consequently the picture is indistinct; and it can only be made clear, either by withdrawing the object to a greater distance, which will bring the focus of the eye nearer to its front, or by interposing a convex lens to increase the refractive power of the eye. Such a condition is termed presbyopic (from its being common in aged persons), or long-sighted. It may proceed to such an extent, that not even the removal of the object to any distance can permit the forma- tion of a distinct picture; so that the assistance of a convex lens must be obtained even to see remote objects clearly; though a less degree of convexity will be required, than for the clear vision of nearer objects. This state is particularly well marked after the operation for cataract; for the removal of the crystalline lens so greatly diminishes the refrac- tive poAver of the eye, as to render necessary the assistance of convex lenses of high curvature. 959. The power, by which a healthy, well-formed eye can accommo- date itself to the distinct vision of objects at varying distances, is a very remarkable one ; and its rationale is not yet properly understood. Ac- cording to the laws already stated (§ 955, V. and VI.) the picture of a near object can only be distinct when formed more remotely from the lens than the picture of a distant object. Consequently when the eye, that has been looking at a distant object, and has seen it clearly, is turned to a near object, a distinct picture of the latter cannot be formed without some alteration, either in the distance between the refractive surfaces and the retina, or in the curvature of the former. It seems most probable that, in the Human eye, this adjustment is chiefly effected by the automatic contraction of the ciliary muscle; which, by drawing the lens nearer to the iris, will thus increase its distance from the retina. But this may not be the sole change. 960. The various humors and containing membranes of the Eye, thus answer the purpose of a most delicate and self-adjusting Optical Fig. 164. Distribution of Capillaries in vascular layer of Retina. instrument; the sole part, which is immediately concerned in the act of sensation, being the Retina, or net-like expansion of the Optic nerve, 518 OF SENSATION. which lies between the black pigment and the vitreous humor. It is in this structure, that the presence of cells at the peripheral as Avell as the central extremities of the afferent nerves (§ 381), may be most clearly demonstrated. They can scarcely be distinguished, in many animals, from the cells of the vesicular matter of the brain ; and, like the latter, they lie in the midst of a plexus of capillary blood-vessels (Fig. 164), which supplies the materials requisite for their groAvth and activity. For the maintenance of the due nutrition of this organ it is requisite that it should be occasionally called into use. If its functional power be destroyed by opacity of the anterior portion of the eye, the nutrition of the retina and optic nerve suffers to such a degree that these parts cease, after a time, to exhibit their characteristic structure ; thus showing that the general rules already stated (chap, vii.) in regard to the connexion between the functional activity and the due nutrition of tissues and organs, hold good with respect to the Nervous structure.— The fibres of the Optic nerve when they diverge to form the Retina, lose their tubular structure; their central axis only being continued, in the form of gray fibres (§ 375), and some of these becoming directly conti- nuous with the caudate vesicles (§ 378). 961. The picture of external objects, which is formed upon the Retina, closely resembles that which we see in a Camera Obscura. It represents the outlines, colors, lights and shades, and relative posi- tions, of the objects before us ; but these do not necessarily convey to the mind the knowledge of their real forms, characters, or distances. The perception of the latter, as already remarked (§ 936), is a mental process; and it may be intuitive or acquired,—the latter, it would seem, being the general condition of the function in Man, the former in the lower animals. The Infant is educating his perceptive powers, long before any indications present themselves, of the exercise of higher mental faculties. By the combination, especially, of the sensations of sight and touch, he is learning to judge of the surfaces of objects as they feel, by the appearance they present,—to form an idea of their distance, by the mode in which his eyes are directed towards them,— and to estimate their size, by combining the notions obtained through the picture on the retina, with those he acquires by the movement of his hands over their different parts. A simple illustration will show, how closely the ideas excited by the two sets of sensations, are blended in our minds. The idea of smoothness is one which has reference to the touch; and yet it constantly occurs to us, on looking at a surface which reflects light in a particular manner. On the other hand, the idea of polish is essentially visual, having reference to the reflection of light from the surface of the object; and yet it would occur to us from the sensa- tions conveyed through the touch, even in the dark. 962. That this sort of combination is not intuitive in Man, but is the result of experience, is evident from the numerous observations made upon those who had acquired the sense of Sight for the first time, after long familiarity with the characters of objects as perceived through the Touch. Thus a boy of four years old, upon whom the operation for con- genital cataract had been very successfully performed, continued to find his way about his father's house, rather by feeling with his hands, SENSE OF VISION. 549 as he had been formerly accustomed to do, than by his newly-acquired sense of Sight; being evidently perplexed, rather than assisted by the sensations which he derived through it. But when learning a new locality, he employed his sight and evidently perceiA'ed the increase of facility which he derived from it. Among the many interesting particulars recorded of the youth on whom Cheselden operated with equal success, it is mentioned that, although perfectly familiar with a dog and a cat by feeling them, and quite able to distinguish between them by his sight, it was long before he associated his visual with his tactual sensations, so as to be able to name either animal by sight alone.—The question was put by the celebrated Locke, whether a person born blind, who was able by his touch to distinguish a cube from a sphere, would, on suddenly obtaining his sight, be able to recognise each by the latter sense; the reply was given in the negative; and the experience of the cases just referred to, as well as of many others, fully justifies such an answer. 963. Still there are, even in Man, certain intuitive perceptions, which afford great assistance in the formation of ideas regarding ex- ternal objects through the visual sense. And the first of these is the power by which we recognise their erect position, notwithstanding the inversion of the image upon the retina. This is certainly not a matter of experience; nor is it capable of explanation (as some have thought) by a reference to the direction in which the rays fall upon the retina. It is the mind which rectifies the inversion ; and as already remarked, it is just as difficult to understand how the inverted image on the retina should be taken cognizance of by the mind at all, as it is to comprehend hoAV it should be thus rectified. In fact, there is no real connexion whatever, between the inversion of the image upon the retina, and that wrong perception of external objects, which some have thought to be its necessary consequence. Any distortion of the picture, giving a wrong vieAv of the relative positions of the objects represented, Avould be attended with a different result.—The same may be said of the cause of the singleness of the sensation perceived by the mind, although an image is formed upon the retina of each eye, of those objects at least, which lie in the field of vision that is common to both. This blending of the pictures, formed upon the two retinae, into a single perception, appears to be, in part at least, the effect of habit. For Avhen the images do not fall upon those parts of the two retinae Avhich are accustomed to act together, double vision is the result. Thus if, when looking steadily at an object, we press one of the eyeballs sideways with the finger, we see two representations of the object; and the same thing frequently occurs, as a result of an affection of the nerves or muscles of one or both eyes (as in ordinary strabismus or squinting), or from some change in the nervous centres, as in various disorders of the Encephalon, and in in- toxication. If this condition should be permanent, howeA'er, we usually find that the individual becomes accustomed to the double images, or rather ceases to perceive that they are double; probably because the mind becomes habituated to receive the impressions from the tAvo parts of the retinse Avhich now act together. And if, after the double vision has passed away, the conformity of the tAvo eyes be restored (as by the 550 OF SENSATION. operation for the cure of squinting), there is double vision for some little time, although the two parts of the retinas, which originally acted toge- ther, are now brought into their pristine position. 964. But the images thus combined are far from being identical; and one of the most remarkable of all our intuitive perceptions, is that by which they are reconciled and combined, and are caused to give rise to an idea that differs essentially from either image. No near object can be seen by the two eyes in the same manner; of this the reader may easily convince himself, by holding up a thin book, in such a position that its back shall be in a line with the nose, and at a moderate distance from it; and by looking at the book, first with one eye, and then with the other. He will find that he gains a different view of the object with each eye, when used separately; so that if he were to represent it, as he actually sees it under these circumstances, he would have two per- spective delineations differing from one another, because drawn from different points. But on looking at the object Avith the two eyes con- jointly, there is no confusion between these pictures; nor does the mind dwell upon either of them singly; but the union of the two intuitively gives us the idea of a solid projecting body,—such an idea as we could . only have otherwise acquired by the exercise of the sense of touch. That this is really the case, has been proved by experiments with a very ingenious instrument, the Stereoscope, invented by Prof. Wheatstone; which is so contrived, as to bring to the two eyes, by reflection from mirrors, two different pictures, such as would be accurate representa- tions of a solid object, as seen by the tAvo eyes respectively. When the arrangement is such, as to bring the images of these pictures to those parts of the retinas, which would have been occupied by the images of the solid (supposing that to have been before the eyes), the mind will perceive, not one or other of the single representations of the object, nor a confused union of the two, but a body projecting in relief, the exact counterpart of that from which the drawings were made.—Thus the combination of the two pictures, and the perception of an object different from either of them, is effected by a mental process of an in- stinctive kind, of the nature of which we know nothing further. 965. When two pictures, representing dissimilar objects, are pro- jected upon the retinae of the two eyes by means of the Stereoscope, the result is a curious one. The mind perceives only one of them, the other being completely excluded for a time; but it commonly happens that, after one has been seen for a short period, the other begins to attract attention and takes its place, the first entirely disappearing; so that there is no confusion or intermingling of images, except at the moment of change. The Will may determine, to a certain extent, which object shall be seen ; but not entirely; for if one picture be more illuminated than the other, it will be seen during a larger proportion of the time.—An interesting variation of this experiment may be made, without the aid of the Stereoscope, by holding a piece of blue glass before one eye, and a piece of yellow glass before the other. The re- sult will be, not that everything will be seen of a green color, but that the surrounding objects will be seen alternately blue and yellow; or sometimes the field of vision will be blue spotted with yellow, alterna- ESTIMATE OF DISTANCE BY VISION. 551 ting with yellow spotted with blue. Thus, when we have two dissimilar objects before the eyes, our attention cannot be kept upon either, to the exclusion of the other, but is alternately and involuntarily directed, either in part or completely, to one and the other. 966. Our ideas of the distance of near objects is evidently acquired from experience; and is suggested by the muscular sensations which are produced by the contraction of the adductor muscles of the eyes. When we direct our eyes towards a near object, a certain degree of con- vergence takes place between their axes; the degree increasing as the distance between the object and the eyes diminishes; and vice versd. We instinctively interpret the sensations thus produced, in such a manner as to be able to compare, with great accuracy, the relative dis- tances of two objects that are not remote from the eyes. This intuition, however, is evidently one of the acquired kind; as may be seen by watching the actions of an infant, or of a person who has recently be- come possessed of Vision. When an object is held before the eyes, and an attempt is made to grasp it, the manner, in which the attempt is made clearly shows, that there is• no power of forming a preciseidea of its situation, such as that which exists in many of the lower animals from their first entrance into the world (§ 938). The impressions made upon the eyes have to be corrected by those received through the touch, be- fore the power of judging of distance is acquired. How much this power depends upon the conjoint use of both eyes, is evident from the difficulty with which any actions, that require an exact appreciation of distance, are performed by those Avho have lost the sight of one eye, until they have acquired neAV modes of judging of it. 967. In regard to remote objects, we have not the same guide; since the convergence of the eyes, in viewing them, is so slight that the axes are virtually parallel. Our judgment of their distance is chiefly founded upon their apparent size, if their actual size be known to us; and also upon the extent of ground, Avhich we see to intervene between ourselves and the object. But if we do not know their actual size, and are so situated that we cannot estimate the intervening space, we form our iudgment chiefly from the greater or less distinctness of their color and outline. Hence our idea of it will be very much affected by varying states of the atmosphere; a slight haziness increasing the apparent dis- tance; whilst a peculiarly clear state of the air will seem to cause re- mote objects to approach much more closely. This want of conver- gence between the axes of the two eyes, has the further effect of causing the pictures upon the two retinae to be nearly identical; and conse- quently the idea of projection is not so strongly excited ; nor are we able to distinguish with the same certainty between a well-painted picture, in which the lights and shades are preserved, and the objects themselves m968e Our notion of the size of an object is closely connected with that of its distance. It is founded upon the dimensions of the picture nroiected on the retina; and the dimensions of this picture will vary, according to the laws of optics (§ 955) inversely as the distance,— beincr fo? example, twice as great when the object is vieAved at the distance of one foot as when it is carried to the distance of two feet. When we know the relative distances of two objects, the estima- 552 OF SENSATION. tion of their real comparative sizes from their apparent sizes is easily effected by a simple process of mind; but this is not the case, when we only guess at their distances ; and our estimate of the size of objects, even moderately remote, is as much affected by states of the atmosphere as that of their distance,—the one being, in fact, proportional to the other. Thus a slight mist, which gives the idea of increased distance, will also augment the apparent size; because in order that an object two miles off, should produce a picture upon the retina of the same extent with that made by an object one mile off, it must have double the dimensions. It is evident that our perception of the size of objects must be acquired by experience, in the same manner as that of their distance has been shown to be. 969. We have now to consider briefly some other phenomena of Vision, in which the acts of mind, that have been just alluded to, do not participate.—The contraction of the Pupil, under the stimulus of light, seems to be effected by a sphincter muscle, which surrounds the orifice, and which is put in action by a branch of the Third pair of nerves. This is an action with which the will has nothing to do ; and it takes place entirely without our consciousness. Although it is due to the stimulus of light, yet there is reason to believe that the conscious- ness of the presence of light is not requisite; and that it is, therefore, a purely reflex action. The Optic nerve seems to be the channel through which the impression is conveyed to the nervous centres; whilst the Third pair is that through which the motor impulse is con- veyed to the iris; but there is some ground for the idea that the Fifth pair may in some degree convey the requisite stimulus, when the optic nerve has been divided. That the dilatation of the pupil is a muscular action, appears probable from the fact that the radiating fibres of the iris are of the same character with the circular; both sets constituting, in Man, a peculiar variety of the non-striated form of muscular tissue. Through what nervous channel, however, the stimulus to this action is conveyed, has not yet been clearly made out. The contraction of the pupil is evidently designed to exclude from the interior of the eye, such an amount of light as would be injurious to it; whilst its dilatation in opposite circumstances admits the greatest possible number of rays. There is a contraction of the pupils, however, Avhich takes place without any change in the amount of light. This occurs when the two eyes are made to converge strongly upon any object brought very near them; and its purpose appears to be, to prevent rays from entering the eye at such a wide angle, as would render it impossible for them to be all brought to their proper foci, and would thus produce an indistinct image. 970. In the use of the Eye, like that of the Ear, there is a tendency to blend into one continuous image a succession of luminous impressions made at short intervals; upon which fact depend a number of* curious optical illusions. The length of the greatest interval that can elapse without an interruption of the presence of the image (in other words the duration of the visual impression), may be measured by causing a luminous object to whirl round, and by ascertaining the longest period that may be allowed for each revolution, consistently with the complete- OF THE VOICE AND SPEECH. 553 ness of the circle of light thus formed. By experiments of this kind, the time has been found to vary, in different individuals, or in different states of the same individual, from about l-4th to l-10th of a second : that is, the impression must be repeated from four to ten times in each second to insure the continuousness of the image. 971. The impressions of variety of color, are produced by the diffe- rently-colored rays, which objects reflect or transmit to the eye. It is curious that some persons, whose sight is perfectly good for forms, distances, &c, are unable to discriminate colors. This curious affection has received the name of Daltonism; from the circumstance that the celebrated Dalton Avas an example of it. There are numerous modifi- cations of it; the want of power to discriminate color being total in some; whilst in others it extends only to certain shades of color, or to the complementary colors. 972. When the retina has been exposed for some time to a strong impression of some particular kind, it seems less susceptible of feebler impressions of the same kind ; thus if we look at any brightly-luminous object, and then turn our eyes upon a sheet of paper, we shall perceive a dark spot upon it: the portion of the retina, which had received the brighter image not being affected by the fainter one. Again, when the eyes have received a strong impression from a colored object, the spot Avhich is seen Avhen the eyes are directed upon a white surface ex- hibits the complementary color; for the retina has been so strongly affected in the part that originally received the image, by its vivid hue, that it does not perceive the fainter hue of the same kind in the object to which it is then turned, and it is impressed only by the remaining rays forming the complementary colors. This explanation applies to the phenomena of the colored shadoAvs which are often seen at sunset, and of those which may be seen in a room whose light enters through colored glass or drapery. For if the prevailing light be of one color, —orange or red for instance,—the eye will not take cognizance of that color in the faint light of the shadoAvs ; and Avill see only its comple- ment, blue or green. If the shadoAv be viewed through a tube, in such a manner that the general colored ground is excluded, it presents the ordinary tint. CHAPTER XIV. OF THE VOICE AND SPEECH. 973. There is one particular application of Muscular power in Man, which deserves special consideration, as being that by which he effects his most complete and intimate communication with his felloAvs ;—that, namely, by which his organ of Voice is put into action. In all air- breathing Vertebrata, the production of sound depends upon the pas- sage of air through a certain portion of the respiratory tubes, which is 554 OF THE VOICE AND SPEECH. so constructed as to set it in vibration, as it passes forth from the lungs.—In Reptiles, the vibrating apparatus is situated at the point, where the trachea opens into the front of the pharynx; it is of very simple construction, however, being only composed of a slit bounded by two contractile lips; and few of the animals of this class can pro- duce any other sound than a hiss, which, owing to the great capacity of their lungs, is often very much prolonged.—In Birds, the situation of the vocal organ is very different. The trachea opens into the front of the pharynx as in Reptiles, by a mere slit; the borders of which have no other movement than that of approaching one another, so as to close the aperture when necessary. This appears to be the instru- ment for regulating the ingress and egress of air, in conformity Avith the wants of the respiratory function. The vocal larynx of Birds is situated at the lower extremity of the trachea, just Avhere it subdivides into the bronchial tubes; and it is of very complex construction, espe- cially in the singing birds.—In Mammalia, on the other hand, the vocal organ and the regulator of the respiration are united in one larynx, which is situated at the top of the trachea. There are few, if any, of this class, Avhich have not some vocal sound; but the variety and expressiveness which can be given to it, .differ considerably in the several orders; being by far the greatest in Man, who alone, there is reason to believe, has the power of producing articulate sounds, or pro- per language. 974. The Larynx is built up as it were, upon the Cricoid cartilage (Fig. 165, x w r u), Avhich surmounts the trachea, and which might be Fig. 165. i Bird's-eye view of Larynx from above, after Willis:— g e h, the thyroid cartilage, embracing the ring of the cricoid r u x w, and turning upon the axis x z, which passes through the lower horns ; n f, n f, the ary- tenoid cartilages, connected by the arytenoideus transversus; t v, t v, the vocal ligaments; n x, the right cricoarytenoideus lateralis (the left being removed); v k f, the left thyro-arytenoideus (the right being re- moved) ; n I, n I, the crico-ary tenoidei postici; b b, the crico-arytenoid ligaments. considered as its highest ring modified in form, its depth from above downwards being much greater posteriorly than anteriorly. This is embraced, as it were, by the thyroid cartilage (g eh); which is arti- REGULATION OF THE APERTURE OF THE GLOTTIS. 555 culated to the sides of the Cricoid by its lower horns, round the extre- mities of which it may be considered to rotate, as on a pivot. In this manner, the front of the Thyroid cartilage may be lifted up, or de- pressed, by the muscles which act upon it; whilst the position of its posterior part is but little changed. Upon the upper surface of the back of the Cricoid cartilage, are seated the two small Arytenoid carti- lages (n f) ; these are so tied to the cricoid by a bundle of strong liga- ments (b b), as to have a sort of rotation upon an articulating surface, which enables them to be approximated or separated from each other, —their inner edges being nearly parallel in the first case, but slanting away from each other in the second. To the summit of these cartilages are attached the Chordse vocales, or vocal ligaments (t u), composed of yellow fibrous or elastic tissue. These stretch across to the front of the Thyroid cartilage; and it is upon their condition and relative situ- ation, that the absence or the production of vocal tones, and all their modifications of pitch, depend. They are rendered tense by the de- pression of the front of the Thyroid cartilage, and relaxed by its ele- vation ; by which action the pitch of the tones is regulated. But for the production of any vocal tones Avhatever, they must be brought into a nearly parallel condition, by the mutual approximation of the points of the arytenoid cartilages to which they are attached ; whilst in the intervals of vocalization, these are separated, and the rima glottidis, or fissure between the chordae vocales, assumes the form of a narrow V, with its point directed backwards. 975. Thus there are two sets of movements concerned in the act of vocalization;—the regulation of the relative position of the Vocal Cords, Avhich is effected by the movements of the Arytenoid cartilages; —and the regulation of their tension, which is determined by the movements of the Thyroid cartilage. The Arytenoid cartilages are made to diverge from one another by means of the Crico-arytenoidei postici of the two sides (n I, N I), which proceed from their outer cor- ners and turn somewhat round the edge of the Cricoid, to be attached to the lower part of its back ; their action is to draw the outer corners of the Arytenoid cartilages outwards and downwards, so that the points to which the vocal ligaments are attached are separated from one another, and the rima glottidis is thrown open. The action of these muscles is antagonized by that of the Arytenoideus transversus, which draAvs together the Arytenoid cartilages ; and by that of the Crico-arytenoidei laterales of the two sides (n x), which run forwards and doAvnwards from the outer corners of the Arytenoid cartilages, and tend by their contraction to bring together their anterior points, to which the Vocal ligaments are attached.—The depression of the front of the Thyroid cartilage, and the consequent tension of the Vocal liga- ments, is occasioned by the conjoint action of the Crico-thyroidei of the two sides, Avhich occasions the Thyroid and Cricoid cartilages to rotate, the one upon the other, at the articulation formed by the infe- rior cornua of the former; and this action will be assisted by the Sterno-thyroidei, which tend to depress the front of the Thyroid carti- lage by pulling from a fixed point below. On the other hand, the elevation of the front of the Thyroid cartilage, and the relaxation of 556 OF THE VOICE AND SPEECH. the Vocal ligaments are affected by the contraction of the Thyro-ary- tenoidei of the two sides (v k f), whose attachments are the same as those of the Vocal ligaments themselves ; and this is aided by the Thyro- hyoidei, which will tend to draw up the front of the Thyroid cartilage, acting from a fixed point above. 976. The muscles which govern the aperture of the glottis,—those namely, which separate and bring together the arytenoid cartilages, and thus widen or contract the space between the posterior extremities of the vocal ligaments,—have important functions in connexion with the Respiratory actions in general; standing as guards, so to speak, at the entrance of the lungs. We can entirely close the glottis through their means, by an effort of the Will, either during inspiration or expi- ration ; and it is a spasmodic movement of this sort, which is concerned in the acts of Coughing and Sneezing, the purpose of Avhich is to expel by a sudden and powerful blast of air, any irritating substances, whether solid, liquid, or gaseous, Avhich have found their way into the air-passages. These muscles appear to be under the sole direction of the inferior or recurrent laryngeal nerve ; which seems to possess ex- clusively motor endowments. When this nerve is divided, on each side, or when the par vagum is divided above its origin, the muscles of the larynx (with the exception of the crico-thyroid) are paralysed; and the aperture of the glottis may remain open, or may be entirely closed, ac- cording to the manner in which its lips are affected by the currents of air in ingress or egress. It is found that, under such circumstances, tranquil respiration may be carried on; but that any violent ingress or egress of air will tend to drive the lips of the glottis (these being in a state of complete relaxation) into apposition with each other, so as com- pletely to close the aperture. The character of the superior laryngeal nerve appears to be almost exclusively afferent; no muscle, except the crico-thyroid, being thrown into contraction when it is irritated; whilst, on the other hand, if it be divided, neither the act of coughing, nor any reflex respiratory movement whatever, can be excited, by irritating the lining membrane of the larynx. 977. During the ordinary acts of inspiration and expiration, the Chordae vocales appear to be widely separated from each other, and to be in a state of the freest possible relaxation. In order to produce a vocal sound, they must be made to approach one another, and their inner faces must be brought into parallelism; both of which ends are accomplished by the rotation of the Arytenoid cartilages; whilst, at the same time, they must be put into a certain degree of tension, by the depression of the Thyroid cartilage. Both of these movements take place consentaneously, and are mutually adapted to each other; the vocal ligaments being approximated, and the rima glottidis consequently narroAved, at the same time that their tension is increased. There is a certain aperture, which is favorable to the production of each tone, although the pitch itself is governed by the tension of the Vocal Cords; and it is, perhaps, to a want of consent betAveen the tAvo, that the pecu- liarly discordant nature of some voices, which appear incapable of pro- ducing a distinct musical tone, is due. 978. It has been fully proved, by the researches of Willis, Miiller, REGULATION OF THE PITCH OF VOCAL SOUNDS. 557 and others, that the action of the Vocal ligaments, in the production of sound, bears no resemblance to that of vibrating strings; and that it is not comparable to that of the mouth-piece of the flute-yipes of the Or- gan; but that it is, in all essential particulars, the same with that of the reeds of the Hautboy or Clarionet, or the tongues of the Accordion or Concertina. All the phenomena attending the production of Musical tones are fully explicable on this hypothesis; except the production of falsetto notes, which has not yet been clearly accounted for.—The power Avhich the Will possesses, of determining, with the most perfect precision, the exact degree of tension which these ligaments shall re- ceive, is extremely remarkable. Their average length in the Male, in the state of repose, is estimated by Miiller at about 73-100ths of an inch ) whilst, in the state of greatest tension, it is about 93-100ths ; the whole difference, therefore, is not above 20-100ths, or one-fifth of an inch. In the female glottis, their average dimensions are about 51- lOOths, and 63-100ths, respectively; so that the difference is here only 12-100ths, or less than one-eighth of an inch. Now the natural com- pass of the voice, in most persons who have cultivated the vocal organ, may be stated at about two octaves, or 24 semitones. Within each semitone, a singer of ordinary capability could produce at least ten dis- tinct intervals; so that, for the total number of intervals, 240 is a very moderate estimate. There must, therefore, be at least 240 different states of tension of the vocal cords, every one of which can be at once determined by the will, when a distinct conception exists of the tone to be produced (§ 905); and, as the whole variation in their length is not more than one-fifth of an inch, even in Man, the variation required, to pass from one interval to another, will not be more than l-1200th of an inch.—And yet this estimate is much below that, which might be truly made from the performance of a practised vocalist. The cele- brated Madame Mara is said to have been able to sound 50 different intervals betAveen each semitone; the compass of her voice was at least 40 semitones, so that the total number of intervals was 2000. The extreme variation in the length of the vocal cords, even taking the larger scale of the Male larynx, not being more than one-fifth of an inch, it may be said that she was able to determine the contractions of her vocal muscles to the ten-thousandth of an inch. 979. It is on account of the greater length of the Vocal cords, that the pitch of the voice is much lower in Man than in Woman : but this difference does not arise until the end of the period of childhood,—the size of the larynx being about the same in the Boy and Girl, up to the age of 14 or 15 years, but then undergoing a rapid increase in the former, whilst it remains nearly stationary in the latter. Hence it is that Boys, as well as Girls and Women, sing treble ; whilst Men sing tenor, which is about an octave lower than the treble; or bass, which is several notes lower still. The cause of the variation in the timbre or quality in different voices is not certainly known; but it appears to be due, in part, to differences in the degree of flexibility and smoothness in the cartilages of the larynx. In Women and children, these carti- lages are usually soft and flexible, and the voice is clear and smooth ; Avhilst in men, and in Avomen whose voices have a masculine roughness, 558 OF THE VOICE AND SPEECH. the cartilages are harder, and are sometimes almost completely ossified. The loudness of the voice depends in part upon the force with which the air is expelled from the lungs ; but the variations in this respect which exist among different individuals, seem partly due to the degree in which its resonance is increased by the vibration of the other parts of the larynx, and of the neighboring cavities. In the HoAvling Monkeys of America, there are several pouches opening from the larynx, which seem destined to increase the volume of tone that issues from it;—one of these is excavated in the substance of the hyoid bone itself. Although these Monkeys are of inconsiderable size, yet their voices are louder than the roaring of lions, and are distinctly audible at the distance of two miles; and when a number of them are congregated together, the effect is terrific. % 980. The vocal sounds produced by the action of the larynx are of very different characters; and may be distinguished into the cry, the song, and the ordinary or acquired voice. The cry is generally a sharp sound, having little modulation or accuracy of pitch, and being usually disagreeable in its timbre or quality. It is that by which animals express their unpleasing emotions, especially pain or terror; and the Human infant, like many of the lower animals, can utter no other sound.—In song, by the regulation of the vocal cords, definite and sus- tained musical tones are produced, Avhich can be changed or modulated at the will of the individual. Different species of Birds have their respective songs ; which are partly instinctive, and partly acquired by education. In Man, the poAver of song is entirely acquired ; but some individuals possess a much greater facility in acquiring it than others, —this superiority appearing to depend upon their more precise con- ception of the tones to be sounded, as well as their more ready imitation, —besides differences in the construction of the larynx itself. The larynx of an accomplished vocalist, obedient to the expression of the emotions, as well as to the dictates of the will, may be said to be the most perfect musical instrument ever constructed.—The voice is a sound more resembling the cry, in regard to the absence of any sustained musical tone; but it differs from the cry, both in the quality of its tone, and in the modulation of which it is capable by the Avill. In ordinary conversation, the voice passes through a great variety of musical tones, in the course of a single sentence, or even a single word, sliding imperceptibly from one to another; and it is when we attempt to fix it definitely to a certain pitch, that we change it from the speak- ing to the singing tone. 981. The poAver of producing articulate sounds, from the combination of which Speech results, is altogether independent of the Larynx ; being due to the action of the muscles of the mouth, tongue and palate. Distinctly-articulate sounds may be produced without any vocal or laryngeal tone, as when we whisper; and it has been experimentally shoAvn, that the only condition necessary for this mode of speech is the propulsion of a current of air through the mouth, from back to front. On the other hand, Ave may have the most perfect laryngeal tone without any articulation; as in the production of musical sounds, not connected with words. But in ordinary speech, the laryngeal tone is modified by ARTICULATE SPEECH—VOAVELS AND CONSONANTS. 559 the various organs which intervene between the larynx and the os externum. The simplest of these modifications is that by which the Vowel sounds are produced,—these sounds being continuous tones, modified by the form of the aperture through which they pass out. Thus, let the reader open his mouth to the widest dimensions, depress the tongue, and raise the velum palati, so as to make the exit of air as free as possible; on then making a vocal sound, he will find that this has the character of the vowel a in ah. On the other hand, if he draw together the lips, still keeping the tongue depressed, he will pass to the sound represented in the English language by oo, in the Continental languages by u. By attention to the production of other vowel sounds, it Avill be found that they are capable of being formed by similar modi- fications in the form of the buccal cavity and the size of the buccal orifice; and that they are capable of being sustained for any length of time. There is an exception, however, in regard to the sound of the English i, as in fine, which is, in reality, a diphthongal sound, produced in the act of transition from a peculiar indefinite murmur to the sound of the long e, which takes its place AA'hen Ave attempt to continue it. The short voAvel sounds, moreover, such as a in fat, e in met, o in pot, &c, are not capable of being perfectly prolonged, as they require, for their true enunciation, to be immediately followed by a consonant.—A tolerably good artificial imitation of VoAvel sounds has been effected by means of a reed-pipe representing the larynx, surmounted by an India- rubber ball, Avith an orifice, representing the cavity and orifice of the mouth. By modifying the form of the ball, the different vowels could be sounded during the action of the reed. 982. In the production of the sounds termed Consonants, the breath suffers a more or less complete interruption, in its passage through the parts anterior to the larynx. The most natural primary division of these sounds is into those which require a total stoppage of the breath at the moment previous to their being pronounced, and which, there- fore, cannot be prolonged; and those in pronouncing which the inter- ruption is partial, and Avhich can, like the vowel sounds, be prolonged ad libitum. The former have received the designation of explosive con- sonants ; the latter are termed continuous. In pronouncing any conso- nants of the explosive class, the posterior nares are completely closed; and the whole current of air is directed through the mouth. This may be checked by the approximation of the lips, as in pronouncing b and p; by the approximation of the point of the tongue to the front of the palate, as in pronouncing d and t; or by the approximation of the middle of the tongue to the arch of the palate, as in pronouncing the hard g or k. The difference betAveen b, d, and#, on the one hand, andp, t, and k, on the other, depends simply upon the greater extent of the meeting surfaces in the former case than in the latter. In sounding some of the con- tinuous consonants, the air is not allowed to pass through the nose; but the interruption in the mouth is incomplete; this is the case with v and /, s and z. In others, the posterior nares are not closed, and the air has a nearly free passage, either through the nose alone, as in m and n, or through the nose and mouth conjointly, as in I and r. The sound of h is a mere aspiration, caused by an increased force of breath; and that 560 OF THE VOICE AND SPEECH. of the guttural ch, as it exists in Welsh, Gaelic, and most Continental languages, is an aspiration modified by the elevation of the tongue, which causes a slight obstruction to the air, and an increased resonance in the back of the mouth. 983. The study of the mode in Avhich the different Consonants are produced, is of particular importance to those who labor under defec- tive speech, especially that difficulty which is known as Stammering. This very annoying impediment is occasioned by a want of proper control over the muscles concerned in Articulation; which, instead of obeying the Will, are sometimes affected with an involuntary or spas- modic action, that interrupts the pronunciation of particular words,— just as, in Chorea, the muscles of the limbs are interrupted by spas- modic twitchings, in the performance of any voluntary movement. In fact, persons affected with general Chorea frequently stammer; show- ing that ordinary Stammering may be considered as a kind of local Chorea. The analogy between the two states is further indicated by the corresponding influence of excited Emotions in aggravating both. It is in the pronunciation of the consonants of the explosive class, that the stammerer usually experiences the greatest difficulty; for the total interruption to the breath, which they occasion, is frequently con- tinued involuntarily ;* so that either the expiration is entirely checked, the whole frame being frequently thrown into the most distressing semi- convulsive movements, or the sound comes out in jerks. Sometimes, however, the spasmodic action occurs in the pronunciation of vowels and continuous consonants ; the stammerer prolonging his expiration, with- out being able to check it. 984. The best method of curing this defect (where there is no mal- formation of the organs of speech, but merely a want of poAver to use them aright), is to study the particular difficulty under which the indi- vidual labors; and then to cause him to practise systematically the various movements concerned in the production of the sounds in ques- tion, at first separately, and afterwards in combination,—until he feels' that his voluntary control over the muscles is complete. The patient would at first do well to practise sentences, from which the explosive consonants are omitted; his chief difficulty, arising from the spasmodic suspension of the expiratory movement, being thus avoided. Having mastered these, he may pass on to others, in Avhich the difficult letters are sparingly introduced; and may finally accustom himself to the use of ordinary language. One of the chief points to be aimed at, is to make the patient feel that he has command over his muscles of articula- tion ; and this is best done, by gradually leading him from that which he can do, to that which he fears to attempt. * The interruption of the expiratory movement in Stammering, is usually stated to take place in the glottis ; but the Author is satisfied that, in all ordinary cases at least, it is in that condition of the mouth, which is preparatory to the pronunciation of one of the explosive consonants. INDEX. N.B. The numbers refer to the Paragraphs. Aberration corrected in eye, 957. Absorbent Cells, 243-245. Vessels, 489. Absorption, from alimentary canal, 489, 490; by lacteals, 243, 244; by blood-vessels, 491-493. from general and pulmonary surfaces, 522, 523. interstitial, by lymphatics, 502, 503; by blood-vessels, 502, 503. Adipose tissue, 257-263, 423-425. Age, influence of, on pulse, 579; on nutrition, 622-627. Air-cells of lungs, 676-679. Albumen, 167-171. conversion of, into fibrine, 519. Albuminous compounds of Plants, 174. Albuminuria, 533, 746. Alga?, development and generation of, 779, 780. • Aliment, sources of demand for, 406-415. effect of variations in supply of, 416- 426. relative value of different kinds of, 427-441. necessity for mixture in, 437. Allantois, formation of, 817. Amnion, formation of, 816. Amphioxus, see Lancelot. Anterior Pyramids, 890. Apoplexy, 688, 924. Area germinativa, 806. pellucida, 807. vasculosa, 551, 813. Areolar tissue, 194-196, 205. Arteries, movement of Blood in, 582-588. elasticity of, 583; tonicity of, 584; contractility of, 585, 586; pulsa- tion of, 583, 584; uniform capacity of, 587; anastomosis of, 588. Articulata, circulation in, 552,553; respi- ration in, 657-661; nervous system in, 856-863. Articulate speech, 981, 982. Asphyxia, 628, 703-709. Assimilating cells, 514, 519. Assimilation, 519; by the liver, 493. Asthma, 678. Atrophy, 619, 620. Attention, effects of, 935. Auditory ganglia, 900. nerve, 950. Azotized Compounds in Plants, 174,428, 429. in Animals, 167-179, 428, 429. destination of, in food, 429, 433. Basement membrane, 206-209. Batrachia, respiration of, 670, 671. Bile, composition and properties of, 724-726. uses of, in digestion, 476-479. Birds, circulation in, 564, 565 ; respiration in, 672-674; lymphatic system in, 500; nerv- ous centres of, 872; heat of, 761. Blastodermic vesicle, 805, 806. Blastema, organizable, 213. Blood, composition of, 525-528; uses of seve- ral constituents of, 529,530; changes of, in disease, 531-534. corpuscles of, white, 214; red, 215-223. coagulation of, 535. buffy coat of, 536, 537. rate of movement of, 577. influence of respiration on, 699-702. Blushing, 603. Bone, structure and composition of, 287-299 j development of, 300-306; regeneration of, 307-309. Brunner's glands, 450, 480. Buffy coat of blood, 536, 537. Butyric acid, 430, 833. Ccecum, secondary digestion in, 481. Canaliculi of Bone, 290. Cancelli of Bone, 289. Cancer-cells, 212, 255. Capillaries, movement of Blood in, 589-604 ; variations in its rate, 597. variations in size of, 595, 603; influence of nerves upon, 603, 604. independent force generated in, 598-600. Carbotiic acid, decomposition of, by Plants, 79-87; necessity for excre- tion of, 641; sources of, in Animal bodies, 642-648. mode of its extrication, 649- 652; amount set free, 691- 698. Cartilage, 264-273 ; multiplication of cells of, 212; "ossification of, 300-303. Caseine, 172, 832. Catamenia, 798, 799. 36 562 INDEX. Cells, vegetable, general history of, 26-42. animal, general history of, 210, 211; production of, 212, 213. isolated, various forms of, 214-216. the immediate agents in Organic func- tions, 246, 247. union of, 248-254. coalescence of, 252-254. changes of form in, 255. Cementum, 319. Centipede, experiments on, 858, 859. Cerebellum, 867-869. functions of, 908-911. Cerebric acid, 383. Cerebrum, 867-873. functions of, 912-925. Chlorosis, state of blood in, 219, 532, 537. Cholesterine, 724. Chondrine, 177. Chorda dorsalis, 211, 251, 812. Chord Be vocales, 974-979. Chorea, 983. Chorion, 795, 809, 817, 818. Chyle, composition and properties of, 515-519. corpuscles of, 214, 518, 519. Chyme, 473, 476. Cilia, 234, 235. Cineritious substance, 379. Circulation, 538,539; in Plants, 540-548 ; in lowest Animals, 549, 550; in Echi- nodermata, 552; in Articulata, 552, 553; in Mollusca, 555-557; in Fishes, 558-560; in Reptiles, 561-563; in Birds and Mammals, 564, 565. in early embrvo, 551, 554, 566; in foetus at birth, 823, 824. Coagulation, of Albumen, 168, 169; of Blood, 535; of Caseine, 172; of Chyle, 518; of Fibrine, 180-187. Cochlea, 952. Cold,degree of, sustainable by Plants, 110,111. degree of, sustainable by Animals, 136. Colostrum, 835. Colors, perception of, 971. Commissures of brain, 913-916. Complementary colors, 972. Conchifera, nervous system of, 852, 853. Concussion, 581. Congestion, arterial, 601, 602. venous, 609, 610. Consensual actions, 903-905. Consonants, 982. Contractility of Muscle, 347. of Vegetable tissues, 345, 346. Convulsive actions, 885. Coral, 277. Cornea, 274. Corpora Malpighiana of the Spleen, 506; of the*Kidney, 728. Corpora Quadrigemina, 873, 900, 902. Striata, 901. Wolffiana, 727. Corpus Callosum, 915. Luteum, 800. Corpuscles of blood, red, 215-223; white, 214. of Chyle and Lymph, 214. Correlation of Forces, 53-61. Cortical substance of brain, 380. Cranium, circulation in, 611. Crura cerebri, 894. Crustacea, geographical distribution of, 133 ; shells of, 286; respiration of, 658. Crusta petrosa, 319. Cryptogamia, generation of, 780. Crystalline lens, 275. Cuttle-fish, nervous cords in arms of, 854. Daltonism, 971. Death, somatic, 65, 68, 69, 628, 629. molecular, 66, 67. Decidua, 810,811. Decussation, of Anterior Pyramids, 890; of Posterior Pyramids, 893 ; of Optic Nerves, 907. Defecation, 462, 463. Deglutition, 453, 454, 897. Dentine, 311-316. Determination of blood, 601. Development, early history of, in Plants, 781; in Animals, 784, 785; see Embryo. Developmental process, influence of heat on, 124-127. Diffusion, mutual, of gases, 650. Digestion, organs of, 442-450. nature of the process of, 471, 472. Disintegration of tissues, 617; of Muscular tissue, 361; of Nervous tissue, 381. Distances, estimate of, 966, 967. Doris, gills of, 651, 656. Dormant Vitality, 43-46. Double vision, 963. Draper, Prof., his views on the capillary cir- culation, 545-548, 598, 599. Dreaming, 924. Duration of pregnancy, 825, 826. of impressions on Ear, 956. of impressions on Eye, 970. Dytiscus, experiments on, 859. Ear, structure of, 950-952. Echinodermata, shells of, 278, 279; circula- tion in, 552. Electricity, development of, in Animals, 771- 777; in Torpedo and Gymno- tus, 771-774 ; in Muscles, 775; in the Frog, 776 ; in higher ani- mals, 777. influence of, on organized bodies, 142; on Vegetation, 143, 144, effects of shocks of, 145 ; influ- ence of, on Animals, 146-148; on Muscles, 351; on nerves, 396, 932. Embryo, early development of, 805, 808; for- mation of vertebral column in, 812; forma- tion of vessels in, 813; formation of heart in, 814; formation of digestive cavity in, 815 ; circulation in, 551, 554, 556. Emotional movements, 917, 919. Emotions, influence of, on hunger, 483; on salivary secretion, 467; on heart's action, 580; on capillary circulation, 603; on mam- mary secretion, 836, 837. Enamel, 318. Endosmose, 491, 492. Entozoa, circulation in, 549, 550. Epidermis, 224-228. Epilepsy, 886. Epithelium, 231-239. Erect vision, 963. Exhalation of water, from lungs, 701; from cutaneous surface, 743-746. of organic matter, 702, 746. Excreting processes, general review of, 751- 759. 1 Eye, structure of, 956-960. INDEX. 563 Facial nerve, 888. Fat, 257-263, 423-425. Fatty liver, 754. Fecundation of ovum, 803, 804. Ferments, action of, on blood, 534. Fertilization of ovum, 803, 804. Fibre, white, 189, 191. yellow, 190, 192. Fibres, formation of, from cells, 193. Fibrillation, 183, 213. Fibrine, coagulation of, 180-187. composition of, 178, 179. production of, 519. Fibrous membranes, 188. Fibrous tissues, simple, 188-193. Fibro-Cartilage, 188, 269, 272. Fifth Pair, 686, 888. Fishes, lymphatic system in, 499 ; circulation in, 558-560 ; respiration in, 663-667; heat of, 760; electricity of, 771-774; nervous centres in, 869, 870. Foetus, circulation in, 822-824. Follicles of glands, 238, 714-719. Follicles of Lieberkiihn, 449. Food, see Aliment. Force, abstract nature of, 18, 71. Forces, Physical, see Physical Forces. Vital, see Vital Forces; their relation to the Physical, 61. Gall-bladder, 478. Ganglion, 380. Gangrene, 634. Gases, mutual diffusion of, 650. Gastric fluid, properties and actions of, 468- 472. conditions of its secretion, 474, 475. Gastric follicles, 469. Gelatinous nerve-fibres, 375. Gelatine, 175, 176, 264, 298; uses of, as food, 429. Gemmation, in Plants, 779; in Animals, 782. Generation, essential character of the pro- cess, 780-783; action of the male in, 786- 790; action of the female in, 791-804. Geographical distribution of Animals, 133. distribution of Plants, 102-106. Germ-cells, 240, 242, 780, 783. Germinal membrane, 805. spot, 794. vesicle, 794. Gestation, duration of, 825, 826. Gills, structure of, 651, 655, 656, 663. Glands, essential parts of, 238, 714-719. Globuline, 221. Glosso-pharyngeal nerve, 888, 897. Glottis, regulation of aperture of, 976. Glycerine, 260. Glycocoll, 176, 734. Gout, 422,615. Graafian Vesicle, 796. Granulation, 637. Gravity, influence of, on venous circulation, 609, 610. Gray nerve-fibres, 375. Gymnotus, 771-774. Haematine, 221,222. Hair, 328-330. Haversian canals, 293-297. Hearing, sense of, 949-954. Heart, action of, 568-570; sounds of, 571- 575; propulsive power of, 576-578 ; frequency of contractions of, 579. power of, independent of nervous agency, 580; influenced by mental emotions, 580 ; by state of nervous system, 581. first development of, in embryo, 554, 566,814. Heat, amount of, developed in Insects, 123 ; in Fishes, 760; in Birds, 761; in Mammals, 761; in Plants, 762. development of, chiefly dependent on production of carbonic acid, 763, 764 ; but partly on other oxidizing pro- cesses, 765; inferior in young ani- mals, 766. of body, kept down by perspiration, 745,768. its influence upon vital activity in gene- ral, 97,98; upon Vegetation, 99-111; upon Animal life, 112-141. degree of, sustainable by Animals, 138- 141; by Plants, 108. Hemispheres, Cerebral, 912-916. Hippuric acid, 730, 734. Hunger, sense of, 483-485. Hybernation, 120, 121. Hydra, stomach of, 443. Hydrophobia, 886, 908. Hypertrophy, 617, 618. Hypo-glossal nerve, 888. Hysteria, 741,887, 908. Inanition, Chossat's experiments on, 117. Incubation, heat supplied in, 125-127. Inflammation, nature of the process, 631,632 ; state of the blood in, 531, 536. Inorganic substances in food, 438-441. Insalivation, 446, 451, 452. Insects, circulation in, 552; respiration in, 659, 660 ; nervous system of, 856-864; re- flex actions of, 858-860; instinctive actions of, 860, 861; heat of, 123. Instinctive actions of Man, 904. Intelligence, 913, 920. Intercellular substance, 248-253. Intestinal canal, structure of, 447-450; move- ments of, 460, 461. Iris, movements of, 969. Irritability of Muscles, 348-363. Kidneys, structure of, 727, 728; action of. 729-741,755. Kreatine, 735. Kreatinine, 735. Lacteals, 494, 496, 499. Lactic acid, in gastric fluid, 470; in urine, 735. Lacunae of Bone, 290. Laminae dorsales, 812. •> Lancelot, 215, 251, 560, 869. Laryngeal nerves, 976. Larynx, structure and actions of, 974, 979. Lead-palsy, 614. Leucin, 170, 176. Light, laws of transmission and refraction of, 955. influence of, on vegetation, 79-92; on growth and development of animals, 93-96. emission of, by animals, 769; by man. 770. 564 INDEX. Life, idea of, 49 ; conditions of, 70; duration of, 25. Lime, in Animal body, 438, 440, 441. Lithic acid, 732, 733. diathesis, 422, 733. Liver, structure of, 720-723; assimilating action of, 493; secreting action of, 476-479, 724-726. Luminousness, animal, 769, 770. Lungs, structure of, 676-679. Lymph,composition and properties of, 515,520. Lymphatics, 498-503. Male, action of, in reproduction, 786-790. Malignant diseases, 640. Malpighian bodies, of Kidney, 728; of Spleen, 506. Mammalta, lymphatic system in, 500; cir- culation in, 564, 565; respiration in, 674- 676; heat evolved in, 761; nervous centres of, 873; ovisac of, 795. Mammary glands, 830, 831. Margarine, 259. Mastication, act of, 451, 452, 896. Medulla Oblongata, structure of, 889-893; functions of, 894-899. Memory, 924. Menstrual secretion, 798,799. Mesenteric glands, 496. Metamorphosis of animals, 407, 784, 785 ; in- fluence of heat upon, 127. Milk, 436; composition and properties of, 832-834. circumstances influencing secretion of, 836-839. Mineral ingredients of food, 438-441. Moisture, proportion of, in different parts of the body, 149-152 ; necessity of, for growth of Plants and Animals, 153-157; effects of withdrawal of, 158-161. Mollusca, circulating system of, 555-557; respiratory organs of, 654-656; nervous system of, 850-854. Mucous membrane, 199-204. Mucus, 237, 464. Mulberry mass, 784, 805. Muscles, contractility of, 345-371 ; irritability of, 348-363 ; tonicity of, 364-366 ; rigor mortis of, 367-369; pecu- liar force of, 370; heat evolved by, 371; electricity evolved by, 775. energy of, dependent on supply of blood, 358-360. disintegration of, 361. Muscular fibre, striated, 332-336 ; non- striated, 337; development of, 338, 339; vessels and nerves of, 340, 341. Muscular sense, 904. Myolemma, 333. Myopia, 958. Nail, 226. Nervous System, general view of actions of, 840-848. Nervous System, in Radiata, 849; in Tuni- cata, 850, 851 ; in Bivalve Mollusca, 852, 853; in higher Mollusca, 854; in Articu- lata, 855-863 ; in Vertebrata, 867, 868; in Fishes, 869, 870; in Reptiles, 871 ; in Birds, 872; in Mammals, 873. Nervous Tissue, 372-405 ; fibrous, 373-377; vesicular, 378, 379 ; arrangement of ele- ments of, in nervous centres, 380; in peri- phery, 381, 382; chemical composition of, 383; disintegration and renewal of, 384- 389 ; conditions of activity of, 390-404. Neurilemma, 373. Nucleus, 211. Nutrition, 612-615. activity of, dependent on func- tional activity of parts, 616-627. Oesophagus, passage of food along, 455, 898. Oily compounds in food, 430, 432, 435; im- portance of, 530. Oleine, 259. Oleo-phosphoric acid, 383. Olfactive lobes, 8G9-873, 900. Olfactory nerve, 906, 946, 947. Olivary bodies, 891. Omphalo-mesenteric vessels, 813. Optic lobes, 869-873, 900. Optic nerves, 906, 907, 960. Orbit, motor nerves of the, 888. Organized structures, general characters of, 2-15. Osseous tissue, see Bone. Ossification, 300-303. Otolithes, 950. Ova of animals, 791-795. Ovarium, 793-797. Ovisac, 792,796. Oxygen, necessity for, in animal body, 649 ; mode of introduction of, 650-652. Pancreatic secretion, 480. Papilla?, sensory, of skin, 382,940 ; of tongue, 943. Parturition, 827-829. Par Vagum, 459, 487, 580, 685, 686, 888,895, 897-899. Pedal ganglia in Mollusca, 852, 853. in Articulata, 857, 862. Pepsine, 470, 471. Perception, nature of, 936, 937. Perceptions, tactual, 941; visual, 961-968. Peristaltic movement, 352, 460. Perspiration, 743-746. Peyer's glands, 450, 749. Phosphate of lime, in food, 438-441 ; in bone, 298. Phosphatic deposits, 386, 738-740. Phosphorus, in animal body, 438, 439. Phanerogamia, generation and development of, 780, 781. Physical Forces, 19-23, 52; correlation of, 53-55; relations of, to Vital, 61-63; their influence on vital action, 73-78. Pigment-cells, 229, 230. Placenta, structure of, 818-820. Placental tufts, 245, 818. Plants, heat of, 762; circulation in, 540-548 ; respiration in, 84, 641, 642; reproduction in, 778-781. Pneumogastric nerve, see Par Vagum. Polypes, digestive process in, 443, 444. Posterior Pyramids, 893. Pregnancy, duration of, 825, 826. Prehension of food, 896. Presbyopia, 958. Primary membrane, 206-209. Primitive trace, 812. Proteine, 171. Puberty in male, 788 ; in female, 798. Pulp of hair, 328, 330. INDEX. 565 Pulp of teeth, 310, 313, 314. Pulsations of heart, 579. Pulse, in arteries, 583, 584; respiratory, in veins, 607. Pupil, changes in diameter of, 969. Purpurine, 736. Pus, 632-637. Pyramids, anterior, 890. posterior, 893. Radiata, Nervous system of, 849. Receptaculum Chyli, 497. Red corpuscles of blood, characters of, 215, 216; development of, 217-220. Reduction of food, provisions for, 445. Reflex actions, nature of, 392-396. of Articulata, 858-860; of Mollusca, 850, 851, 854 ; of Vertebrata, 875-879, 884. Regeneration of parts, influence of heat upon, 129; of nerve, 389. Reproductive cells, 240-242. Reptiles, circulation in, 562, 563; lymphatic system in, 499; respiration in, 668-671; nervous centres in, 871. Respiration, nature of the process, 641; sources of demand for it, 642-648; mode of its per- formance, 649-652. organs *of, in lowest animals, 653 ; in Mollusca, 654-656 ; in Annelida, 657; in Crus- tacea, 658; in Insects, 659, 660 ; in Spiders, 661 ; in v Fishes, 663-667; in Rep- tiles, 668-671 ; in Birds, 672-674; in Mammalia and Man, 675-678. movements of, 679-688. chemical phenomena of, 689- 702. insufficient, effects of, 703- 709. Respiratory nerves, in Insects, 862; in Mol- luscs, 850-853; in Vertebrata, 684-688, 895. Respiratory pulse, 607. Restiform bodies, 892. Retina, general structure of, 960. Rigor Mortis, 367-369. Ruminating stomach, 457. Saccharine compounds in food, 430-434, 493. Salivary glands, 465. secretion, 446, 466, 467. Satiety, sense of, 486, Sebaceous follicles, 747, 748. Secreting cells, 238, 239, 712-714. Secretion, nature of the process, 710-713. effects of suppression of, 711. Segmentation of vitellus, 805. Selecting power of individual parts, 612-615. Semicircular canals, 952. Sensation, 389, 390, 930, 931; nerves of, 389, 390, 900, 901; general and special, 932. Sensations, regulation ot muscular movement by, 902-904. Sensorium, 390. . Sensory Ganglia, 900, 901; functions of, 902 -905. Sensory nerves, 906, 907. Serous membranes, 197, 198. Shell, of Echinodermata, 278, 279 ; of Mol- lusca, 280-285 ; of Articulata, 284-286. Sight, sense of, 955-972. Single vision with two eyes, 963. Size of objects, estimate of, 968. Skin, 199, 205, 742-748, 940. ' Sleep, 924. Smell, sense of, 946-948. Sneezing, 948. Solen, nervous system of, 852, 853. Somnambulism, 924. Sounds, propagation of, 949; qualities of, 954. Sounds of heart, 571-575. Speech, 981, 982. Sperm-cells, 240, 241, 780, 783. Spermatic fluid, 786, 787; emission of, 790. Spermatozoa, 240, 786, 787; use of, in fecun- dation, 803, 804. Sphinx ligustri, nervous system of, 856, 857. Spiders, respiratory organs of, 661. Spinal Cord, 867, 868; its independence of the Brain, 874-879 ; structure of, 880-883 ; reflex actions of, 884 ; disordered states of. 885-887. Spinal accessory nerve, 580, 888. Spinal nerves, origin of, 880, 882. peculiar, 888. Spiracles of Insects, 659. Spleen, structure of, 506 ; uses of, 507-509. Stammering, 983, 984. Starchy compounds in food, 430-434. Star-fish, nervous system of, 849. Starvation, Chossat's experiments on, 117. Stearine, 259. Stereoscope, 964, 965. Stomach, 447, 448; movements of, 458, 459. in Ruminants, 457. Stomato-gastric nerves of Invertebrata, 863. of Vertebrata, 896. Strabismus, 963. Suction, act of, 896. Sudoriparous glandulae, 743, 744. Supra-renal capsules, 510. Sympathetic System, in Man, functions of, 926-929. traces of, among Invertebrata, 864. Syncope, 581,628. Synovial membranes, 197, 198. Tadpole, respiration of, 670; metamorphosis of, 670. Ttiste, nerves of, 944. sense of, 945. Teeth, structure and development of, 310- 327. Temperature, sense of, 933, 942. Testis, structure of, 786. Tetanus, 886. Thalami Optici, 901. Thirst, 488. Thoracic duct, 497. Thymus Gland, 511,512. Thyroid Gland, 513. Tickling, 907. Tongue, papilla? of, 943. Tonicity of arteries, 365, 584; of muscle, 364 -366. Torpedo, electricity of, 771-774. Torpidity, induced by cold, 97, 98, 136; by want of moisture, 158-161. Touch, sense of, 939-942. Tubercula quadrigemina, 869, 870-873,900. 566 INDEX. Tubercular diathesis, 626, 638, 639. Tunicata, nervous system of, 850, 851. Tympanum, 951. Tyrosin, 170. Ulceration, 633. Umbilical vessels, 818. Umbilical vesicle, 815. Urea, 73Q, 731. Uric acid, 732, 733. Urine, composition and properties of, 729- 740; effects of suppression of, 741. Uterus, comparative structure of, 793 ; partu- rient action of, 827. Vascular area, 551, 813, 814. Vegetable kingdom, office of, 15. Vegetation, influence of Light upon, 79-92; influence of Heat upon, 98-107; influence of Electricity upon, 143-145. Veins, movement of blood in, 605-610; pul- sation in, 607, 608. Venous congestion, 609, 610. Villi of mucous membrane, 243, 492 ; of yolk- bag, 244; of placenta. 245. Vital Actions, 16-18, 47-51. Vital Forces, 17, 24, 52; their relations to each other, 58-60; to the Physical forces, 61-63. Vital Stimuli, 61. Vitellus, 784, 794; segmentation of, 805. Vitreous body of eye, 276. Vocal ligaments, 974-979. Voice, production of, 973-979. Voluntary movements, nature of, 923. Vowel sounds, production of, 981. White fibrous tissue, 189, 191. Worm tribes, circulation in, 552; respiration in, 657. > Yellow fibrous tissue,«190, 192. Yolk, see Vitellus. 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The high reputation of the author, on both sides of the Atlantic, is a sufficient guarantee for the completeness and accuracy of any work to which his name is prefixed; but this volume comes with the additional recommendation that it is the one on which the author has bestowed the greatest care, and on which he is desirous to rest his reputation. Two years have been devoted to the pre- paration of this edition, which has been thoroughly remoulded and rewritten, so as, in fact, to constitute a new work. The amount of alterations and addi- tions may be understood from the fact that of the ten hundred and eighty pages of the text, but one hundred and fifty belong to the previous edition. Con- taining, as it does, the results of years devoted to study and observation, it may be regarded as a complete exposition of the most advanced state of knowledge in this rapidly-progressive branch of science, and as a storehouse of facts and principles in all departments of Physiology, such as perhaps no man but its author could have accumulated and classified. In every point of mechanical execution, and profuseness of beauty of illus- tration, the Publishers risk nothing in saying that it will be found all that the most fastidious taste could desire. " A truly magnificent work. In itself a perfect physiological study."—Ranlcing's Abstract, July 21,1851. '■ It is impracticable for us to give an analysis of the varied contents of this most useful volume. Its pro- duction has been a labor of love with its author, and has subjected him to much thought, and to no little toil, without the expectation of pecuniary profit. It is to be hoped, however, that so much ability, zeal, and industry, may meet with their reward, and that future editions may remunerate him for productive exertions so beneficial in their results to others. We may remark, in conclusion, that the work is beautifully gotten up in the English fashion, and that the illustrations are in the first style of art."—Medical Examiner. "This work stands without its fellow. It is one few men in Europe could have undertaken; it is one no man, we believe, could have brought to so successful an issue as Dr. Carpenter. It required for its produc- tion a physiologist at once deeply read in the labors of others, capable of taking a general, critical, and unprejudiced view of those labors, and of combining the varied, heterogeneous materials at his disposal, so as to form an harmonious whole. " We feel that this abstract can give the reader hut a very imperfect idea of the fulness of this work, and no idea of its unity, of the admirable manner in which material has been brought, from the most various sources, to conduce to its completeness, of the lucidity of the reasoning it contains, or of the clearness of language in which the whole is clothed. Not the profession only, but the scientific world at large, must feel deeply indebted to Dr. Carpenter for this great work. It must, indeed, add largely even to his high reputation."—Medical Times. '• This is a hook without a rival; and Dr. Carpenter has laid the medical profession, as well as all who study and who love natural history, under deep and lasting obligations by its production. Our limits forbid us from indulging in comment or criticism; and we can only heartily commend the work to our readers."— London Journal of Medicine. " The present edition contains upwards of one thousand pages, in close type, and contains a mass of infor- mation not to be easily found, even in a well furnished library. Physiology, Zoology, Botany, and Micro- scopy, all lend their'aid to the elucidation of the laws of life and development; and the style is such as to interest the reader, and to fix his attention upon the particular subject to which he has occasion to refer. We must also observe that the beautiful and accurate illustrations which accompany this edition (exceed- ing three hundred in number) make plain to the eye that in which description would fail, and materially aid the author in familiarizing his readers with the results of numerous microscopical observations. " It is our opinion that whether for reference or study in the subject to which it specially refers, no better book than Dr. Carpenter's 'Principles of Physiology, General and Comparative,' can be placed in the hands of student or practitioner. It would also he a valuable addition to the library of every well-educated man, although not a member of the profession."—London Medical Gazette. " I recommend to your perusal a work recently published by Dr. Carpenter. It has this advantage, it is very much up to the present state of knowledge on the subject. It is written in a clear style, and is well illustrated."—Professor Sharpey's Introductory Lecture. " In Dr. Carpenter's work will be found the best exposition we possess of all that is furnished by compa- rative anatomy to our knowledge of the nervous system, as well as to the more general principles of life and organization."—Dr. Holland's Medical Notes and Reflections. '■ See Dr. Carpenter's ' Principles of General and Comparative Physiology.' —a work which makes me proud to think he was once my pupil."—Dr. EUiotson's Physiology. BLANCHARD & LEA'S MEDICAL AND SURGICAL PUBLICATIOXS. TO THE MEDICAL PROFESSION. In the present catalogue we have affixed prices to our publications, in obedience to the repeated requests of numerous members of the profession. While books, like all other articles, must necessarily vary somewhat in cost throughout the ex- tended territories of this country, yet our publications will generally be furnished at these rates by booksellers throughout the Union, who can readily and speedily procure any which they may not have on hand. To accommodate those physicians who have not convenient access to bookstores, or who cannot order through merchants visiting the larger cities, we will forward our works by mail,/ree of postage, on receipt of the printed prices in current funds or postage stamps. As our business is wholesale, and we open accounts with book- sellers only, the amount must in every case, without exception, accompany the order, and we can assume no risks of the mail, either on the money or the books; and as we sell only our own publications, we can supply no others. Physicians will, therefore, see the convenience and advantage of making their purchases, when- ever practicable, from the nearest bookseller. We can only add that no exertions are spared to merit a continuance of the gratifying confidence hitherto manifested by the profession in all works bearing our imprint. BLANCHARD & LEA. Philadelphia, February, 1859. *„* We have just ready a new Illustrated Catalogue of our Medical and Scientific Publications, forming an octavo pamphlet of 80 large pages, containing specimens of illustrations, notices of the medical press, &c. &c. It has been pre- pared without regard to expense, and will be found one of the handsomest specimens of typographical execution as yet presented in this country. Copies will be sent to any address, by mail, free of postage, on receipt of nine cents in stamps. Catalogues of our numerous publications in miscellaneous and educational litera- ture forwarded on application. TWO MEDICAL PERIODICALS, FREE OF POSTAGE, FOR FIVE DOLLARS PER ANNUM. THE AMERICAN JOURNAL OF THE MEDICAL SCIENCES, subject to postage, when not paid for in advance, - - - - - - -$5 00 THE MEDICAL NEWS AND LIBRARY, invariably in advance, - - 1 00 or, both periodicals furnished, free of postage, for Five Dollars remitted in advance. THE AMERICAN JOURNAL OF THE MEDICAL SCIENCES, Edited by ISAAC HAYS, M. D., is published Quarterly, on the first of January, April, July, and October. Each number contains at least two hundred and eighty large octavo pages, handsomely and appropriately illustrated, wherever necessary. It has now been issued regularly for more than thirty-five year*, and it has been under the control of the present editor for more than a quarter of a century. Throughout this long period, it has maintained its position in the highest rank of medical periodicals both at home and abroad, and has received the cordial support of the entire profession in this country. Its list of Collaborators will be found to contain a large number of the most distinguished names of the pro fession in every section of the United States, rendering the department devoted to ORIGINAL COMMUNICATIONS full of varied and important matter, of great interest to all practitioners. As the aim of the Journal, however, is to combine the advantages presented by all the different varieties of periodicals, in its iy 2 BLANCHARD & LEA'S MEDICAL REVIEW DEPARTMENT will be found extended and impartial reviews of all important new works, presenting subjects of novelty and interest, together with very numerous BIBLIOGRAPHICAL NOTICES, including nearly all the medical publications of the day, both in this country and Great Britain, with a choice selection of the more important continental works. This is followed by the QUARTERLY SUMMARY, being a very full and complete abstract, methodically arranged, of the IMPROVEMENTS AND DISCOVERIES IN THE MEDICAL SCIENCES. This department of the Journal, so important to the practising physician, is the object of especial care on the part of the editor. It is classified and arranged under different heads, thus facilitating the researches of the reader in pursuit of particular subjects, and will be found to present a very full and accurate digest of all observations, discoveries, and inventions recorded in every branch of medical science. The very extensive arrangements of the publishers are such as to afford to the editor complete materials for this purpose, as he not only regularly receives ALL THE AMERICAN MEDICAL AND SCIENTIFIC PERIODICALS, but also twenty or thirty of the more important Journals issued in Great Britain and on the Conti. nent, thus enabling him to present in a convenient compass a thorough and complete abstract of everything interesting or important to the physician occurring in any part of the civilized world. To their old subscribers, many of whom have been on their list for twenty or thirty years, the publishers feel that no promises for the future are necessary; but those who may desire for the first time to subscribe, can rest assured that no exertion will be spared to maintain the Journal in the high position which it has occupied for so long a period. By reference to the terms it will be seen that, in addition to this large amount of valuable and practical information on every branch of medical science, the subscriber, by paying in advance, becomes entitled, without further charge, to THE MEDICAL NEWS AND LIBRARY, a monthly periodical of thirty-two large octavo pages. Its "News Department" presents the current information of the day, while the " Library Department" is devoted to presenting stand- ard works on various branches of medicine. Within a few years, subscribers have thus received, without expense, the following works which have passed through its columns:— WATSON'S LECTURES ON THE PRACTICE OF PHYSIC. BRODIE'S CLINICAL LECTURES ON SURGERY. TODD AND BOWMAN'S PHYSIOLOGICAL ANATOMY AND PHYSIOLOGY OF MAN. WEST'S LECTURES ON THE DISEASES OF INFANCY AND CHILDHOOD. MALGAIGNE'S OPERATIVE SURGERY, with wood-cuts. SIMON'S LECTURES ON GENERAL PATHOLOGY. BENNETT ON PULMONARY TUBERCULOSIS, with wood-cuts, WEST ON ULCERATION OF THE OS UTERI, BROWN ON THE SURGICAL DISEASES OF FEMALES, with wood-cuts, and WEST ON DISEASES OF WOMEN, Part I. (See p. 32.) While in the number for January, 1858, is commenced a new and highly important work, HAB^RSHON ON DISEASES OF THE ALIMENTARY CANAL. It will be seen that this treatise covers the whole ground of affections of the Digestive Organ?, which furnish so very large a portion of the daily practice of the physician. The author's position in Guy's Hospital, and the fact that the work has only just appeared in London, are sufficient guar- antee that it is up to the hour, and presents the most advanced condition of this department of medi- cal science, while its thorough practical character is manifested by the great number of cases on which it is founded, no less than one hundred and sixty-three being carefully analyzed in all their details, throughout its pages. It is therefore hoped that it will be found to fully maintain the valu- able practical character of the works heretofore presented to subscribers through this medium. It will thus be seen that for the small sum of FIVE DOLLARS, paid in advance, the subscriber will obtain a Quarterly and a Monthly periodical, EMBRACING NEARLY SIXTEEN HUNDRED LARGE OCTAVO PAGES, mailed to any part of the United States, free of postage. These very favorable terms are now presented by the publishers with the view of removing ail difficulties and objections to a full and extended circulation of the Medical Journal to the office of every member of the profession throughout the United States. The rapid extension of mail facili- ties will now place the numbers before subscribers with a certainty and dispatch not heretofore attainable; while by the system now proposed, every subscriber throughout the Union is placed upon an equal footing, at the very reasonable price of Five Dollars for two periodicals, without further expense. Those subscribers who do not pay m advance will bear in mind that their subscription of Five Dollars will entitle them to the Journal only, without the News, and that they will be at the expense ' of their own postage on the receipt of each number. The advantage of a remittance when order- ing the Journal will thus be apparent. As the Medical News and Library is in no case sent without advance payment, its subscribers will always receive it free of postage. Remittances of subscriptions can be mailed at our risk, when a certificate is taken from the Post- master that the money is duly inclosed and forwarded. Address BLANCHARD & LEA, Philadelphia. AND SCIENTIFIC PUBLICATIONS. 3 ALLEN (J. M.), M. D., Professor of Anatomy in the Pennsylvania Medical College, Ac. THE PRACTICAL ANATOMIST; or, The Student's Guide in the Dissecting- ROOM. With 266 illustrations. In one handsome roval 12mo. volume, of over 600 pages, lea- ther. $2 25. 3 In the arrangement of this work, the author has endeavored to present a complete, and thorough course of dissections in a clearer and more available form for practical use, than has as yet been accomplished. The chapters follow each other in the order in which dissections are usually con- ducled in this country, and as each region is taken up, every detail regarding it is fully described and illustrated, so that the student is not interrupted in his labors, by the necessity of referring from one portion of the volume to another. However valuable may be the " Dissector's Guides" which we, of late, have had occasion to notice, we feel confident that the work of Dr. Allen is superior to any of them. We believe with the author, that none is so fully illustrated as this, and the nrrangement of the work is such as to facilitate the labors of the student in acquiring a thorough practical knowledge of Anatomy. We most cordi- Blly recommend it to their attention.—Western Lan- cet, Dec. 1856. We believe it to be one of the most useful works upon the subject ever written. It is handsomely illustrated, well printed, and will be found of con- venient size for use in the dissecting-room.—Med. Examiner, Dec. 1856. From Prof. J. S. Davis, University of Va. I am not acquainted with any 'work that attains so fully the object which it proposes. From C. P. Fanner, M. D., Demonstrator, Uni- versity of Michigan. I have examined the work briefly, but even this examination has convinced me that it is an excellent guide for the Dissector. Its illustrations are beau- tiful, and more than I have seen in a work of this kind. I shall take great pleasure in recommending it to my classes as the text-book of the dissecting- room. ANALYTICAL COMPENDIUM OF MEDICAL SCIENCE, containing Anatomy, Physiology, Surgery, Midwifery, Chemistry, Materia Medica, Therapeutics, and Practice of Medicine. By John Neill, M. D., and F. G-. Smith, M. D. New and enlarged edition, one thick volume royal 12mo. of over 1000 pages, with 374 illustrations. $3 00. OF See Neill, p. 24. ABEL (F. A.), F. C.S. AND C. L. BLOXAM. HANDBOOK OF CHEMISTRY, Theoretical, Practical, and Technical; with a Recommendatory Preface by Dr. Hofmann. In one large octavo volume, extra cloth, of 662 with illustrations. $3 25. ASHWELL (SAMUEL), M.D., Obstetric Physician and Lecturer to Guy's Hospital, London. A PRACTICAL TREATISE ON THE DISEASES PECULIAR TO WOMEN Illustrated by Cases derived from Hospital and Private Practice. Third American, from the Third and revised London edition. In one octavo volume, extra cloth, of 528 pages. $3 00. The most useful practical work on the subject in the English language. — Boston Med. and Surg. Journal. The most able, and certainly the most standard and practical, work on female diseases that we have yet seen.—Medico-Chirurgical Review. ARNOTT (NEILL), M. D. ELEMENTS OF PHYSICS; or Natural Philosophy, General and Medical. Written for universal use, in plain or non-technical language. A new edition, by Isaac Hays, M. D. Complete in one octavo volume, leather, of 484 pages, with about two hundred illustra- tions. $2 50. _____________ BUDD (GEORGE), M. D., F. R. S., Professor of Medicine in King's College, London. ON DISEASES OF THE LIVER. Third American, from the third and enlarged London edition. In one very handsome octavo volume, extra cloth, with four beauti- fully colored plates, and numerous wood-cuts. pp. 500. $3 00. (Just Issued.) Has fairly established for itself a place among the classical medical literature of England.—British and Foreign Medico-Chir. Review, July, 1857. Dr. Budd's Treatise on Diseases of the Liver is now a standard work in Medical literature, and dur- ing the intervals which have elapsed between the successive editions, the author has incorporated into the text the most striking novelties which have cha- racterized the recent progress of hepatic physiology and pathology; so that although the size of the book is not perceptibly changed, the history of liver dis- eases is made more complete, and is kept upon a level with the progress of modern science. It is the best work on Diseases of the Liver in any language.__ London Med. Times and Gazette, June 27, 1857. This work, now the standard book of reference on the diseases of which it treats, has been carefully revised, and many new illustrations of the views of the learned author added in the present edition.__ Dublin Quarterly Journal, Aug. 1S57. BY THE SAME AUTHOR. ON THE ORGANIC DISEASES AND FUNCTIONAL DISORDERS OF THE STOMACH. In one neat octavo volume, extra cloth. $1 50. From the high position occupied by Dr. Budd as a teacher, a writer, and a practitioner, it is almost needless to state that the present book may be con- Rultedwith great advantage. It is written in an easy style, the subjects are well arranged, and the practi- cal precepts, both of diagnosis and treatment, denote the character of a thoughtful and experienced phy- sician.—London Med. Times and Gazette. 4 BLANCHARD & LEA'S MEDICAL BUCKNILL (J. C), M. D., Medical Superintendent of the Devon County Lunatic Asylum; and DANIEL H. TUKE, M. D., Visiting Medical Officer to the York Retreat. A MANUAL OF PSYCHOLOGICAL MEDICINE; containing the History, Nosology, Description, Statistics, Diagnosis, Pathology, and Treatment of INSANITY. With a Plate. In one handsome octavo volume, of 536 pages. $3 00. (Now Ready, July, 1858.) The increase of mental disease in its various forms, and the difficult questions to which it is constantly giving rise, render the subject one of daily enhanced interest, requiring on the part of the physician a constantly greater familiarity with this, the most perplexing branch of his profes- sion. At the same time there has been for some years no work accessible in this country, present- ing the results of recent investigations in the Diagnosis and Prognosis of Insanity, and the greatly improved methods of treatment which have done so much in alleviating the condition or restoring the health of the insane. To fill this vacancy the publishers present this volume, assured that the distinguished reputation and experience of the authors will entitle it at once to the confidence of both student and practitioner. Its scope may be gathered from the declaration of the authors that "their aim has been to supply a text book which may serve as a guide in the acquisition of *ueh knowledge, sufficiently elementary to be adapted to the wants of the student, and sufficiently modern in its views and explicit in its teaching to suffice for the demands of the practitioner." BENNETT (J. HUGHES), M.D., F. R. S. E., Professor of Clinical Medicine in the University of Edinburgh, &c. THE PATHOLOGY AND TREATMENT OF PULMONARY TUBERCU- LOSIS, and on the Local Medication of Pharyngeal and Laryngeal Diseases frequently mistaken for or associated with, Phthisis. One vol. 8vo.,extra cloth, with wood-cuts. pp. 130. $1 25. BENNETT (HENRY), M. D. A PRACTICAL TREATISE ON INFLAMMATION OF THE UTERUS, ITS CERVIX AND APPENDAGES, and on its connection with Uterine Disease. New and much enlarged edition, preparing by the author for publication in 1859. We are firmly of opinion that in proportion as a Knowledge of uterine, diseases becomes more appre- ciated, this work will be proportionably established as a text-book in the profession.—The Lancet. When, a few years back, the first edition of the present work was published, the subject was one al- most entirely unknown to the obstetrical celebrities of the day ; and even now we have reason to know that the bulk of the profession are not fully alive to the importance and frequency of the disease of which it takes cognizance. The present edition is so much enlarged, altered, and improved, that it can scarcely be considered the same work.—Dr. Ranking's Ab- stract. BY THE SAME AUTHOR. A REVIEW OF THE PRESENT STATE OF UTERINE PATHOLOGY. 8vo., 75 pages, flexible cloth, 50 cents. BIRD (GOLDING), A. M., M. D., Sec. URINARY DEPOSITS: THEIR DIAGNOSIS, PATHOLOGY, AND THERAPEUTICAL INDICATIONS. A new and enlarged American, from a late improved London edition. With over sixty illustrations. In one royal 12mo. vol, extra cloth, pp.372. $130. It can scarcely be necessary for us to say anything uf the merits of this well-known Treatise, which so admirably brings into practical application the re- sults of those microscopical and chemical researches regarding the physiology and pathology of the uri- aary secretion, which have contributed so much to the increase of our diagnostic powers, and to the extension and satisfactoryemploymentof our thera- peutic resources. In the preparation of this new edition of his work, it is obvious that Dr. Golding Bird has spared no pains to render it a faithful repre- sentation of the present state of scientific knowledge on thesubject it embraces.— The British and Foreign Medico-Chirurgical Review. BOWMAN (JOHN EJ, M.D. PRACTICAL HANDBOOK OF MEDICAL CHEMISTRY. Second Ame- rican, from the third and revised English Edition. In one neat volume, royal 12mo., extra cloth, with numerous illustrations, pp. 288. $1 25. BY THE SAME AUTHOR. INTRODUCTION TO PRACTICAL CHEMISTRY, INCLUDING ANA- LYSIS. Second American, from the second and revised London edition. With numerous illus- trations. In one neat vol., royal 12mo., extra cloth, pp. 350. $1 25. BEALE ON THE LAWS OF HEALTH IN RE- LATION TO MIND AND BODY. A Series of Letters from an old Practitioner to a Patient. In one volume, royal 12mo., extra cloth, pp. 296. 80 cents. tfUSHNAN'S PHYSIOLOGY OF ANIMAL AND VEGETABLE LIFE ; a Popular Treatise on the Functions and Phenomena of Organic Life. In one handsome royal ISmo. volume, extra cloth, with over 100 illustrations, pp.234. 80 cents. BUCKLER ON THE ETIOLOGY, PATHOLOGY, AND TREATMENT OF FIBRO-BRONCHI- TIS AND RHEUMATIC PNEUMONIA. In one 8vo. volume, extra cloth, pp.150. SI 25. BLOOD AND URINE (MANUALS ON). BY JOHN WILLIAM GRIFFITH, G. OWEN REESE, AND ALFRED MARKWICK. One thick volume, royal 12mo., extra cloth, with plates, pp. 460. SI 25. BRODIE'S CLINICAL LECTURES ON SUR- GERY. 1 vol. 8vo. cloth. 350 pp. SI 25. AND SCIENTIFIC PUBLICATIONS. 5 BARCLAY (A. W.) M. D., Assistant Physician to St. George's Hospital, &c. A MANUAL OF MEDICAL DIAGNOSIS; being an Analysis of the Signs and Symptoms of Disease. In one neat octavo volume, extra cloth, of 424 pages. $2 00. (A new work, now ready.) Of works exclusively devoted to this important deficiency, is the object of Dr. Barclay's Manual. branch, our profession has at command, compara- j The task of composing such a work is neither an tively, but few, and, therefore, in the publication of easy nor a light one; but Dr. Barclay has performed the present work, Messrs. Blanchard k Lea have ! it in a manner which meets our most unqualified conferred a great favor upon us. Dr. Barclay, from approbation. He is no mere theorist; he knows his having occupied, for ajonj* period, the position of work thoroughly, and in attempting to perform it, has not exceeded his powers.—British Med. Journal, Medical Registrar at St. George's Hospital, pos sessed advantages for correct observation and reli- able conclusions, as to the significance of symptoms, which have fallen to the lot of but few, either in his own or any other country. He has carefully systematized the results of his observation of over twelve thousand patients, and by his diligence and judicious classification, the profession has been presented with the most convenient and reliable work on the subject of Diagnosis that it has been our good fortune ever to examine; we can, there- fore, say of Dr. Barclay's work, that, from his sys- tematic manner of arrangement, his work is one of the best works " for reference" in the daily emer- gencies of the practitioner, with which we are ac- quainted ; but, at the same time, we would recom- mend our readers, especially the younger ones, to rend thoroughly and study diligently the whole work, and the ''emergencies" will not occur so often.— Southern Med. and Surg. Journ., March, 1858. To give this information, to supply this admitted Dec. 5, 1857. We venture to predict that the work will be de- servedly popular, and soon become, like Watson^s Practice, an indispensable necessity to the practi- tioner.—iV. A. Med. Journal, April, 1S58. An inestimable work of reference for the young practitioner and student.—Nashville Med. Journal, May, 1858. We hope the volume will have an extensive cir- culation, not among students of medicine only, but practitioners also. They will never regret a faith- ful study of its pages.—Cincinnati Lancet, Mar. '58. This Manual of Medical Diagnosis is one of the most scientific, useful, and instructive works of its kind that we have ever read, and Dr. Barclay has done good service to medical science in collecting, arranging, and analyzing the signs and symptoms of so many diseases. — N. J. Med. and Surg. Re- porter, March, 1858. BARLOW (GEORGE H.), M. D. Physician to Guy's Hospital, London, &c. A MANUAL OF THE PRACTICE OF MEDICINE. With Additions by D. F. Condie, M. D., author of "A Practical Treatise on Diseases of Children," &c. In one hand- some octavo volume, leather, of over 600 pages. $2 75. We recommend Dr. Barlow's Manual in the warm- est manner as a most valuable vade-mecum. We have had frequent occasion to consult it, and have found it clear, concise, practical, and sound. It is eminently a practical work, containing all that is essential, and avoiding useless theoretical discus- sion. The work supplies what has been for some time wanting, a manual of practice based upon mo- dern discoveries in pathology and rational views of treatment of disease. It is especially intended for the use of students and junior practitioners, but it will be found hardly less useful to the experienced physician. The American editor has added to the work three chapters—on Cholera Infantum, Yellow Fever, and Cerebro-spinal Meningitis. These addi- tions, the two first of which are indispensable to a work on practice destined for the profession in this country, are executed with great judgment and fi- delity, by Dr. Condie, who has also succeeded hap- pily in imitating the conciseness and clearness of style which are such agreeable characteristics of the original book.—Boston Med. and Surg. Journal. BARTLETT (ELISHA), M. D. THE HISTORY, DIAGNOSIS, AND TREATMENT OF THE FEVERS OF THE UNITED STATES. A new and revised edition. By Alonzo Clark, M. D., Prof. of Pathology and Practical Medicine in the N. Y. College of Physicians and Surgeons, &c. In one octavo volume, of six hundred pages, extra cloth. Price $3 00. It is the best work on fevers which has emanated i logy. His annotations add much to the interest of from the American press, and the present editor has the work, and have brought it well up to the condi- carefully availed himself of all information exist- ing upon the subject in the Old and New World, so that the doctrines advanced are brought down to the latest date in the progress of this department of Medical Science.—London Med. Times and Gazette, May 2, 1857. This excellent monograph on febrile disease, has stood deservedly high since its first publication. It will be seen that it has now reached its fourth edi- tion under the supervision of Prof. A. Clark, a gen- tleman who, from the nature of his studies and pur- suits, is well calculated to appreciate and discuss the many intricate and difficult questions in patho- tion of the science as it exists at the present day in regard to this class of diseases.—Southern Med. and Surg. Journal, Mar. 1857. It is a work of great practical value and interest, containing much that is new relative to the several diseases of which it treats, and, with the additions of the editor, is fully up to the times. The distinct- ive features of the different forms of fever are plainly and forcibly portrayed, and the lines of demarcation carefully and accurately drawn, and to the Ameri- can practitioner is a more valuable and safe guide than any work on fever extant.—Ohio Med. and Surg. Journal, May, 1857. BROWN (ISAAC BAKER), Surgeon-Accoucheur to St. Mary's Hospital, &c. ON SOME DISEASES OF WOMEN ADMITTING OF SURGICAL TREAT- MENT. With handsome illustrations. One vol. 8vo., extra cloth, pp. 276. $ I 60. Mr Brown has earned for himself a high reputa- tion in the operative treatment of sundry diseases and injuries to which females are peculiarly subject. and merit the careful attention of every surgeon- accoucheur.—Association Journal. We have no hesitation in recommending this book We can truly say of his workthat it is an important, ^ the carefu, attention of all ,nr °who make addition to obstetrical lit«»t"f-. "'£7 | female complaints a part of their study and practice. suggestions and contrivances which Mr Brown de- _DubHn q\iarterly JoumaL * p Bcribes, exhibit much practical sagacity and skill, | "* i) BLANCHARD & LEA'S MEDICAL CARPENTER (WILLIAM B.), M. D., F. R. S., &c, Examiner in Physiology and Comparative Anatomy in the University of London. PRINCIPLES OF HUMAN PHYSIOLOGY; with their chief applications to Psychology, Pathology, Therapeutics, Hygiene, and Forensic Medicine. A new American, from the last and revised London edition. With nearly three hundred illustrations. Edited, with addi- tions, by Francis Gurney Smith, M. D., Professor of the Institutes of Medicine in the Pennsyl- vania Medical College, &c. In one very large and beautiful octavo volume, of about nine hundred large pages, handsomely printed and strongly bound in leather, with raised bands. $4 25. In the preparation of this new edition, the author has spared no labor to render it, as heretofore, a complete and lucid exposition of the most advanced condition of its important subject. The amount of the additions required to effect this object thoroughly, joined to the former large size ol the volume, presenting objections arising from the unwieldy bulk of the work, he has omitted all those portions not bearing directly upon Human Physiology, designing to incorporate them in his forthcoming Treatise on General Physiology. As a full and accurate text-book on the Phy- i-iology of Man, the work in its present condition therefore presents even greater claims upon the student and physician than those which have heretofore won for it the very wide and distin- guished favor which it has so long enjoyed. The additions of Prof. Smith will be found to supply whatever may have been wanting to the American student, while the introduction of many new illustrations, and the most careful mechanical execution, render the volume one of the most at- tractive as yet issued. For upwards of thirteen years Dr. Carpenter's work has been considered by the profession gene- rally, both in this country and England, as the most valuable compendium on the subject of physiology in our language. This distinction it owes to the high attainments and unwearied industry of its accom- plished author. The present edition (which, like the last American one, was prepared by the author him- self), is the result of such extensive revision, that it may almost be considered a new work. We need hardly say, in concluding this brief notice, that while the work is indispensable to every student of medi- cine in this country, it will amply repay the practi- tioner for its perusal by the interest and value of its contents.—Boston Med. and Surg. Journal. This is a standard work—the text-book used by all medical students who read the English language. It has passed through several editions in order to keep pace with the rapidly growing science of Phy- siology. Nothing need be said in its praise, for its merits are universally known ; we have nothing to say of its defects, for they only appear where the science of which it treats is incomplete.—Western Lancet. The most complete exposition of physiology which any language can at present give.—Brit, and For. Med.-Chirurg. Review. The greatest, the most reliable, and the best book on the subject which we know of in the English language.—Stethoscope. This book should not only be read but thoroughly Btudied by every member of the profession. None are too wise or old, to be benefited thereby. But especially to the younger class would we cordially commend it as best fitted of any work in the English language to qualify them for the reception and com- prehension of those truths which are daily being de- veloped in physiology.—Medical Counsellor. Without pretending to it, it is an encyclopedia of the subject, accurate and complete in all respects— a truthful reflection of the advanced state at which the science has now arrived.—Dublin Quarterly Journal of Medical Science. A truly magnificent work—in itself a perfect phy- siological study.—Ranking's Abstract. This work stands without its fellow. It is one few men in Europe could have undertaken; it is one To eulogize this great work would be superfluous We should observe, however, that in this edition the author has remodelled a large portion of the former, and the editor has added much matter of in- terest, especially in the form of illustrations. We may confidently recommend it as the most complete work on Human Physiology in our language.— Southern Med. and Surg. Journal, December, 1855, The most complete work on the science in our language.—Am. Med. Journal. The most complete work now extant in our lan- guage.—N. O. Med. Register. The best text-book in the language on this ex- tensive subject.—London Med. Times. A complete cyclopaedia of this branch of science. —N. Y. Med. Times. The profession of this country, and perhaps also of Europe, have anxiously and for some time awaited the announcement of this new edition of Carpenter's Human Physiology. His former editions have for many years been almost the only text-book on Phy- siology in all our medical schools, and its circula- tion among the profession has been unsurpassed by any work in any department of medical science. It is quite unnecessary for us to speak of thii work as its merits would justify. The mere an- nouncement of its appearance will afford the highest pleasure to every student of Physiology, while its perusal will be of infinite service in advancing physiological science.—Ohio Med. and Surg. Journ, no man, we believe, could have brought to so suc- cessful an issue as Dr. Carpenter. It required for its production a physiologist at once deeply read in the labors of others, capable of taking a general, critical, and unprejudiced view of those labors, and of combining the varied, heterogeneous materials at his disposal, so as to form an harmonious whole. We feel that this abstract can give the reader a very imperfect idea of the fulness of this work, and no idea of its unity, of the admirable manner in which material has been brought, from the most various sources, to conduce to its completeness, of the lucid- ity of the reasoning it contains, or of the clearness of language in which the whole is clothed. Not the profession only, but the scientific world at large, must feel deeply indebted to Dr. Carpenter for this freat work. It must, indeed, add largely even to is high reputation.—Medical Times. BY the same author. (Lately Issued.) PRINCIPLES OF COMPARATIVE PHYSIOLOGY. New American, from the Fourth and Revised London edition. In one large and handsome octavo volume, with ovet three hundred beautiful illustrations, pp. 752. Extra cloth, $4 80; leather, raised bands, f 5 25. The delay which has existed in the appearance of this work has been caused by the very thorough revision and remodelling which it has undergone at the hands of the author, and the large number of new illustrations which have been prepared for it. It will, therefore, be found almost a new work, and fully up to the day in every department of the subject, rendering it a reliable text-book for all students engaged in this branch of science. Every effort has been made to render its typo- graphical finish and mechanical execution worthy of its exalted reputation, and creditable to the mechanical arts of this country. AND SCIENTIFIC PUBLICATIONS. 7 CARPENTER (WILLIAM B.)f M. D., F. R. S., Examiner in Physiology and Comparative Anatomy in the University of London. THE MICROSCOPE AND ITS REVELATIONS. With an Appendix con- taining the Applications of the Microscope to Clinical Medicine, &c. By F. G. Smith, M. D. Illustrated by four hundred and thirty-four beautiful engravings on wood. In one large and very handsome octavo volume, of 724 pages, extra cloth, $4 00; leather, $4 50. Dr. Carpenter's position as a microscopist and physiologist, and his great experience as a teacher eminently qualify him to produce what has long been wanted—a good text-book on the practical use of the microscope. In the present volume his object has been, as stated in his Preface, " to combine, within a moderate compass, that information with regard to the use of his ' tools,' which is most essential to the working microscopist, with such an account of the objects best fitted for Uis study, as might qualify him to comprehend what he observes, and might thus prepare him to benefit science, whilst expanding and refreshing his own mind." That he has succeeded in accom- plishing this, no one acquainted with his previous labors can doubt. The great importance of the microscope as a means of diagnosis, and the number of microsco- pists who are also physicians, have induced the American publishers, with the author's approval, to add an Appendix, carefully prepared by Professor Smith, on the applications of the instrument to clinical medicine, together with an account of American Microscopes, their modifications and accessories. This portion of the work is illustrated with nearly one hundred wood-cuts, and, it is hoped, will adapt the volume more particularly to the use of the American student. Every care has been taken in the mechanical execution of the work, which is confidently pre- sented as in no respect inferior to the choicest productions of the London press. The mode in which the author has executed his intentions may be gathered from the following condensed synopsis of the CONTENTS. Introduction—History of the Microscope. Chap. I. Optical Principles of the Microscope. Chap. II. Construction of the Microscope. Chap. III. Accessory Apparatus. Chap. IV. Management of the Microscope Chap. V. Preparation, Mounting, and Collection of Objects. Chap. VI. Microscopic Forms of Vegetable Life—Protophytes. Chap. VII. Higher Cryptoga- mia. Chap. VIII. Phanerogamic Plants. Chap. IX. Microscopic Forms of Animal Life—Pro- tozoa—Animalcules. Chap. X. Foraminifera, Polycystina, and Sponges. Chap. XI. Zoophytes. Chap. XII. Echinodermata. Chap. XIII. Polyzoa and Compound Tunicata. Chap. XIV. Molluscous Animals Generally. Chap. XV. Annulosa. Chap. XVI. Crustacea. Chap. XVII. Insects and Arachnida. Chap. XVIII. Vertebrated Animals. Chap. XIX. Applications of the Microscope to Geology. Chap. XX. Inorganic or Mineral Kingdom—Polarization. Appendix. Microscope as a means of Diagnosis—Injections—Microscopes of American Manufacture. Those who are acquainted with Dr. Carpenter's previous writings on Animal and Vegetable Physio- fogy, will fully understand how vast a store of know- ledge he is able to bring to bear upon so comprehen- sive a subject as the revelations of the microscope ; and even those who have no previous acquaintance with the construction or uses of this instrument, will find abundance of information conveyed in clear and simple language.—Med. Times and Gazette. Although originally not intended as a strictly medical work, the additions by Prof. Smith give it a positive claim upon the profession, for which we doubt not he will receive their sincere thanks. In- deed, we know not where the student of medicine will find such a complete and satisfactory collection of microscopic facts bearing upon physiology and practical medicine as is contained in Prof. Smith's appendix; and this of itself, it seems to us, is fully worth the cost of the volume.—Louisville Medical Review, Nov. 1856. BY THE SAME AUTHOR. ELEMENTS (OR MANUAL) OF PHYSIOLOGY, INCLUDING PHYSIO- LOGICAL ANATOMY. Second American, from a new and revised London edition. With one hundred and ninety illustrations. In one very handsome octavo volume, leather, pp. 566. $3 00. In publishing the first edition of this work, its title was altered from that of the London volume, by the substitution of the word " Elements" for that of " Manual," and with the author's sanction the title of " Elements" is still retained as being more expressive of the scope of the treatise. To say that it is the best manual of Physiology Those who have occasion for an elementary trea- now before the public, would not do sufficient justice tise on Physiology, cannot do better than to possess to the author.—Buffalo Medical Journal. themselves ofthe manual of Dr. Carpenter .—Medical In his former works it would seem that he had Examiner. exhausted the subjectof Physiology. In the present, The best and most complete expose of modern heeives theessence, as it were, ofthe whole.—N. Y. Physiology, in one volume extant in the English Journal of Medicine. language.—St. Louis Medical Journal. BY the same author. (Preparing.) PRINCIPLES OF GENERAL PHYSIOLOGY, INCLUDING ORGANIC CHEMISTRY AND HISTOLOGY". With a General Sketch of the Vegetable and Animal Kingdom. In one large and very handsome octavo volume, with several hundred illustrations. The subject of general physiology having been omitted in the last editions ot the author's " Com- parative Physiology" and "Human Physiology," he has undertaken to prepare a volume which shall present it more thoroughly and fully than has yet been attempted, and which may be regarded tis an introduction to his other works. BY THE SAME AUTHOR. A PRIZE ESSAY ON THE USE OF ALCOHOLIC LIQUORS IN HEALTH AND DISEASE. New edition, with a Preface by D. F. Condie, M. D., and explanations of scientific words. In one neat 12mo. volume, extra cloth, pp. 178. 50 cents. s BLANCHARD & LEA'S MEDICAL CONDIE (D. F.), M. D., &.C. A PRACTICAL TREATISE ON THE DISEASES OF CHILDREN. Fifth edition, revised and augmented. In one large volume, 8vo., leather, of over 750 pages. $3 25. (Now Ready, December, 1858.) In presenting a new and revised edition cf this favorite work, the publishers have only to state that the author ha* endeavored to render it in every respect "a complete and faithful exposition of the pathology and therapeutics of the maladies incident to the earlier stages of existence—a full and exact account ofthe diseases of infancy and childhood." To accomplish this he has subjected the whole work to a careful and thorough revision, rewriting a considerable portion, and adding several new chapters. In this manner it is hoped that any deficiencies which may have previously existed have been supplied, that the recent labors of practitioners and observers have been tho- roughly incorporated, and that in every point the work will be found to maintain the high reputation it has enjoyed as a complete and thoroughly practical book of reierence in infantile affections. A few notices of previous editions are subjoined. We pronounced the first edition to be the best work on the diseases of children in the English language, and, notwithstanding all that has been published, we still regard it in that light.—Medical Examiner. Dr. Condie's scholarship, acumen, industry, and practical sense are manifested in this, as in all his numerous contributions to science.—Dr. Holmes's Report to the American Medical Association. Taken as a whole, in our judgment. Dr. Condie's Treatise is the one from the perusal of which the practitioner in this country will rise with the great- est satisfaction.—Western Journal of Medicine and Surgery. One of the best works upon the Diseases of Chil- aren in the English language.—Western Lancet. We feel assured from actual experience that no physician's library can be complete without a copy of this work.—N. Y. Journal of Medicine. A veritable pediatric encyclopaedia, and an honoi to American medical literature.—Ohio Medical and Surgical Journal. We feel persuaded that the American medical pro- fession will soon regard it not only as a very good, but as the vert best "Practical Treatise on the Diseases of Children."—American Medical Journal. In the department of infantile therapeutics, the work of Dr. Condie is considered one of the best which has been published in the English language. —The Stethoscope. The value of works by native authors on the dis- eases which the physician is called upon to combat, will be appreciated by all; and the work of Dr. Con- die has gained for itself the character of a safe guide for students, and a useful work for consultation by those engaged in practice.—N. Y. Med. Times. This is the fourth edition of this deservedly popu- lar treatise. During the interval since the last edi- tion, it has been subjected to a thorough revision by the author; and all new observations in the pathology and therapeutics of children have been includea in the present volume. As we said btfore, we do not know of a better book on diseases of chil- dren, and to a large part of its recommendations we yield an unhesitating concurrence.—Buffalo Med. Journal. Perhaps the most full and complete work now be- fore the profession ofthe United States; indeed, we may say in the English language. It is vastly supe- rior to most of its predecessors.—Transylvania Med. Journal, CHRISTISON (ROBERT), M. D., V. P. R. S. E., &c. A DISPENSATORY; or, Commentary on the Pharmacopoeias of Great Britain and the United States; comprising the Natural History, Description, Chemistry, Pharmacy, Ac- tions, Uses, and Doses of the Articles of the Materia Medica. Second edition, revised and im- proved, with a Supplement containing the most important New Remedies. With copious Addi- tions, and two hundred and thirteen large wood-engravings. By R. Eglesfeld Griffith, M. D. In one very large and handsome octavo volume, leather, raised bands, of over 1000 pages. $3 50. COOPER (BRANSBY B ) F R. S. LECTURES ON THE PRINCIPLES AND ' PRACTICE OF SURGERY- In one very large octavo volume, extra cloth, of 750 pages. $3 00. COOPER ON DISLOCATIONS AND FRAC- TURES OF THE JOINTS —Edited by Bransby B. Cooper, F. R. S., &c. With additional Ob- servations by Prof. J. C. Warren. A new Ame- rican edition. In one handsome octavo volume, extra cloth, of about 500 pages, with numerous illustrations on wood. $3 25. COOPER ON THE ANATOMY AND DISEASES OF THE BREAST, with twenty-five Miscellane- ous and Surgical Papers. One large volume, im- perial 8vo., extra cloth, with 252 figures, on 36 plates. $2 50. COOPER ON THE STRUCTURE AND DIS- EASES OF THE TESTIS, AND ON THE THYMUS GLAND. One vol. imperial 8vo., ex- tra cloth, with 177 figures on 29 plates. $2 00. COPLAND ON THE CAUSES, NATURE, AND TREATMENT OF PALSY AND APOPLEXY. In one volume, royal 12mo., extra cloth, pp. 326. 80 cents. CLYMER ON FEVERS; THEIR DIAGNOSIS, PATHOLOGY, AND TREATMENT In one octavo volume, leather, of 600 pages. $1 50. COLOMBAT DE L'ISERE ON THE DISEASES OF FEMALES, and on the special Hygiene of their Sex. Translated, with many Notes and Ad- ditions, by C. D. Meigs, M.D. Second edition, revised and improved. In one large volume, oc- tavo, leather, with numerous wood-cuts. pp. 720. $3 50. CARSON (JOSEPH), M.D., Professor of Materia Medica and Pharmacy in the University of Pennsylvania. SYNOPSIS OF THE COURSE OF LECTURES ON MATERIA MEDICA AND PHARMACY, delivered in the University of Pennsylvania. Second and revised edi- tion. In one very neat octavo volume, extra cloth, of 208 pages. $1 50. CURLING (T. B.), F. R.S., Surgeon to the London Hospital, President of the Hunterian Society, &c. A PRACTICAL TREATISE ON DISEASES OF THE TESTIS, SPERMA- TIC CORD, AND SCROTUM. Second American, from the second and enlarged English edi- tion. In one handsome octavo volume, extra cloth, with numerous illustrations, pp. 420. $2 00. AND SCIENTIFIC PUBLICATIONS. CHURCHILL (FLEETWOOD), M. D., M. R. I. A. ON THE THEORY AND PRACTICE OF MIDWIFERY. Edited, with Notes and Additions, by D. Francis Condie, M. D., author of a "Practical Treatise on the Diseases of Children," &c. With 139 illustrations. In one very handsome octavo volume, leather, pp.510. $3 00. To bestow praise on a book that has received such marked approbation would be superfluous. We need only say, therefore, that if the first edition was thought worthy of a favorable reception by the medical public, we can confidently affirm that this will be found much more so. The lecturer, the practitioner, and the student, may all have recourse to its pages, and derive from their perusal much in- terest and instruction in everything relating to theo- retical and practical midwifery.—Dublin Quarterly Journal of Medical Science. A work of very great merit, and such as we can confidently recommend to the study of every obste- tric practitioner.—London Medical Gazette. This is certainly the most perfect system extant. It is the best adapted for the purposes of a text- book, and that which he whose necessities confine him to one book, should select in preference to all others.—Southern Medical and Surgical Journal. The most popular -work on midwifery ever issued from the American press.—Charleston Med. Journal. Were we reduced to the necessity of having but one work on midwifery, and permitted to choose, we would unhesitatingly take Churchill.—Western Med. and Surg. Journal. It is impossible to conceive a more useful and elegant manual than Dr. Churchill's Practice of Midwifery.—Provincial Medical Journal. Certainly, in our opinion, the very best work on the subject which exists.—N. Y. Annalist. BY THE SAME AUTHOR. (Just Issued.) ON THE DISEASES OF INFANTS AND CHILDREN. Second American Edition, revised and enlarged by the author. Edited, with Notes, by W. V. Keating, M. D. In one large and handsome volume, extra cloth, of over 700 pages. $3 00, or in leather, $3 25. In preparing this work a second time for the American profession, the author has spared no labor in giving it a very thorough revision, introducing several new chapters, and rewriting others, while every portion of the volume has been subjected to a severe scrutiny. The efforts of the American editor have been directed to supplying such information relative to matters peculiar to this country as might have escaped the attention of the author, and the whole may, there- fore, be safely pronounced one of the most complete works on the subject accessible to the Ame- rican Profession. By an alteration in the size of the page, these very extensive additions have been accommodated without unduly increasing the size of the work. contribution for the illustration of its topics. The material thus derived has been used with consummate skill, and the result has been a work creditablealike -N. A. Medico-Chir. No work holds a higher position, or is more de- serving of being placed in the hands of the tyro, the advanced student, or the practitioner.—Medical Examiner. Previous editions, under the editorial supervision of Prof R. M. Huston, have been received with marked favor, and they deserved it; but this, re- printed from a very late Dublin edition, carefully revised and brought up by the author to the present time, does present an unusually accurate and able exposition of every important particular embraced in the department of midwifery. * * The clearness, directness, and precision of its teachings, together with the great amount of statistical research which its text exhibits, have served to place it already in the foremost rank of works in this department of re- medial science.—N. O. Med. and Surg. Journal. In our opinion, it forms one of the best if not the very best text-book and epitome of obstetric science which we at present possess in the English lan- guage.—Monthly Journal of Medical Science. The clearness and precision of style in which it is written, and the great amount of statistical research which it contains, have served to place it in the first rank of works in this departmentof medical science. — N. Y. Journal of Medicine. Few treatises will be found better adapted as a text-book for the student, or as a manual for the frequent consultation of the young practitioner.— American Medical Journal. This work contains a vast amount of interesting matter, which is so well arianged and so curtly worded that the book may be regarded as an ency- clopaedia of information upon the subject of which it treats. It is certainly also a monument of Dr. Churchill's untiring industry, inasmuch as there is not a single work upon the diseases of children with which we are acquainted that is not fully referred to and quoted from in its pages, and scarcely a con- tribution of the least importance to any British or Foreign Medical Journal, for some years past, which is not duly noticed.—London Lancet, Feb. 20, lt>58. Availing himself of every fresh source of informa- tion, Dr. Churchill endeavored, with his accustomed industry and perseverance, to bring his work up to the present state of medical knowledge in all the subjects of which it treats; and in this endeavor he has, we feel bound to say, been eminently success- ful. Besides the addition of more than one hundred and thirty pages of matter, we observe that some entirely new and important chapters are introduced, viz : on paralysis, syphilis, phthisis, sclerema, &c. &c. As the work now stands, it is, we believe, the most comprehensive in the English language upon the diseases incident to early life.—Dublin Quarterly Journal, Feb. 1858. It brings before the reader an amount of informa- tion not comprised in any similar production in the language. The amount of labor consumed upon its production can only be conceived by those who have been similarly occupied, every work of note pub- lished witnin the last twenty-five years in the dit- ferent languages of Europe having been laid under I and Surgical Journal BY THE SAME AUTHOR. ESSAYS ON THE PUERPERAL FEVER, AND OTHER DISEASES PE- CULIAR TO WOMEN. Selected from the writings of British Authors previous to the close of the Eighteenth Century. In one neat octavo volume, extra cloth, of about 450 pages. $2 50. to the author and his country. Review, May, 18^8. After this meagre, and we know, very imperfect notice of Dr. Churchill's work, we shall conclude by saying, that it is one that cannot fail from its co- piousness, extensive research, and general accuracy, to exalt still higher the reputation of the author in this country. The American reader will beoarticu- larly pleased to find that Dr. Churchill has done full justice throughout his work to the various American authors on this subject. The names of Dewees, Eberle, Condie, and Stewart, occur on nearly every page, and these authors are constantly referred toby the author in terms ofthe highest praise, and with the most liberal courtesy.—The Medical Examiner. We recommend the work of Dr. Churchill most cordially, both to students and practitioners, as a valuable and;reliable guide in the treatment of the dis- eases of children.—Am. Journ.of the Med. Sciences. We know of no work on this department of Prac- tical Medicine which presents so candid and unpre- judiced a statement or posting up of our actual knowledge as this.— N. Y. Journal of Medicine. Its claims to merit both as a scientific and practi- cal work, are of the highest order. Whilst we would not elevate it above every other treatise on the same subject, we certainly believe that very few are equal to it, and none superior.—Southern Med. 10 BLANCHARD & LEA'S MEDICAL CHURCHILL (FLEETWOOD), M. D., M. R. I. A., &c. ON THE DISEASES OF WOMEN; including those of Pregnancy and Child- bed. A new American edition, revised by the Author. With Notes and Additions, bv D Fran- cis Condie, M. D., author of "A Practical Treatise on the Diseases of Children." With nume- rous illustrations. In one large and handsome octavo volume, leather, of 76S pages. (Just Issued, 1857.) $3 00. This edition of Dr. Churchill's very popular treatise may almost be termed a new work, so thoroughly has he revised it in every portion. It will be found greatly enlarged, and thoroughly brought up to the most recent condition of the subject, while the very handsome series of illustra- tions introduced, representing such pathological conditions as can be accurately portrayed, present a novel feature, and afFord valuable assistance to the young practitioner. Such additions as ap- peared desirable for the American student have been made by the editor, Dr. Condie, while a marked improvement in the mechanical execution keeps pace with the advance in all other respects which the volume has undergone, while the price has been kept at the former very moderate rate. extent that Dr. Churchill does. His, indeed, is the only thorough treatise we know of on the subject; and it may be commended to practitioners and stu- It comprises, unquestionably, one of the most ex- act and comprehensive expositions of the present state of medical knowledge in respect to the diseases of women that has yet been published.—Am. Journ. Med. Sciences, July, 1857. We hail with much pleasure the volume before us, thoroughly revised, corrected, and brought up to the latest date, by Dr. Churchill himself, and rendered still more valuable by notes, from the ex- perienced and able pen of Dr. D. F. Condie, of Phil- adelphia.— Southern Med. and Surg. Journal, Oct. 1657. This work is the most reliable -which we possess on this subject; and is deservedly popular with the profession.—Charleston Med. Journal, July, 1857. Dr. Churchill's treatise on the Diseases of Women is, perhaps, the most popular of his works with the profession in this country. It has been very gene- rally received both as a text-book and manual of practice. The present edition has undergone the most elaborate revision, and additions of an import- ant character have been made, to render it a com- plete exponent of the present state of our knowledge of these diseases.—N. Y. Journ. of Med., Sept. 1857. We now regretfully take leave of Dr. Churchill's book. Had our typographical limits permitted, we should gladly have borrowed more from its richly stored pages. In conclusion, we heartily recom- mend it to the profession, and would at the same time express our firm conviction that itwill not only add to the reputation of its author, but will pTove a work of great and extensive utility to obstetric practitioners.—Dublin Medical Press. We know of no author who deserves that appro- bation, on "the diseases of females," to the same dents as a masterpiece in its particular department. The former editions of this work have been com- mended strongly in this journal, and they have won their way to an extended, and a well-deseTved popu- larity. This fifth edition, before us, is well calcu- lated to maintain Dr. Churchill's high reputation. It was revised and enlarged by the author, for his American publishers, and it seems to us that there is scarcely any species of desirable information on itB subjects that may not be found in this work.—Tht Western Journal of Medicine and Surgery. We are gratified to announce a new and revised edition of Dr. Churchill's valuable work on the dis- eases of females We have ever regarded it as one of the very best works on the subjects embraced within its scope, in the English language; and the present edition, enlarged and revised by the author, renders it still more entitled to the confidence of the profession. The valuable notes of Prof. Huston have been retained, and contribute, in no small de- gree, to enhance the value of the work. It is a source of congratulation that the publishers have permitted the author to be, in this instance, his own editor, thus securing all the revision which an author alone is capable of making.—The Western Lancet. As a comprehensive manual for students, or a work of reference for practitioners, we only speak with common justice when we say that it surpasses any other that has ever issued on the same sub- ject from the British press.—The Dublin Quarterly Journal. DICKSON (S. H.), M. D., Professor of Practice of Medicine in the Jefferson Medical College, Philadelphia. ELEMENTS OF MEDICINE; a Compendious View of Pathology and Thera- peutics, or the History and Treatment of Diseases. In one large and handsome octavo volume, of 750 pages, leather. $3 75. As an American text-book on the Practice of Medicine for the student, and as a condensed work of reference for the practitioner, this volume will have strong claims on the attention ofthe profession. Few physicians have had wider opportunities than the author for observation and experience, and few perhaps have used them better. As the result of a life of study and practice, therefore, the present volume will doubtless be received with the welcome it deserves. This book is eminently what it professes to be; a distinguished merit in these days. Designed for " Teachers and Students of Medicine," and admira- bly suited to their wantB, we think it will be received, on its own merits, with a hearty welcome.—Boston Med. and Surg. Journal. Indited by one of the most accomplished writers of our country, as well as by one who has long held a high position among teachers and practitioners of medicine, this work is entitled to patronage and careful study. The learned author has endeavored to condense in this volume most of the practical matter contained in his former productions, so as to adapt it to the use of those who have not time to devote to moreextensiveworks.—SouthernMed. and Surg. Journal. Prof. Dickson's work supplies, to a great extent, a desideratum long felt in American medicine.—N. O. Med. and Surg. Journal, Estimating this work according to the purpose for which it is designed, we must think highly of its merits, and we have no hesitation in predicting for it a favorable reception by both students and teachers. Not professing to be a complete and comprehensive treatise, it will not be found full in detail, nor filled with discussions of theories and opinions, but em- bracing all that is essential in theory and practice, it is admirably adapted to the wants of the American student. Avoiding all that is uncertain, it presents more clearly to the mind of the reader that which is established and verified by experience. The varied and extensive reading of the author is conspicuously apparent, and all the recent improvements and dis- coveries in therapeutics and pathology are chroni- cled in its pages.—Charleston Med. Journal. In the first part of the work the subject of gene- ral pathology is presented in outline, giving a btau- tiful picture of its distinguishing features, and throughout the succeeding chapters we find that he has kept scrupulously within the bounds of sound reasoning and legitimate deduction. Upon the whole, we do not hesitate to pronounce it a superior work in its class, and that Dr. Dickson merits a place in the first rank of American writers.—Western Lancet. AND SCIENTIFIC PUBLICATIONS. 11 DRUITT (ROBERT), M.R. C.8., &.C. THE PRINCIPLES AND PRACTICE OF MODERN SURGERY. Edited by F. W. Sargent, M. D., author of " Minor Surgery," &c. Illustrated with one hundred and ninety-three wood-engravings. In one very handsomely printed octavo volume, leather, of 576 large pages. $3 00. Dr. Druitt's researches into the literature of his ; ments of theoretical and practical Surgery—that it subject have been not only extensive, but well di- is found to contain reliable and authentic informa- rected ; the most discordant authors are fairly and ! tion on the nature and treatment of nearly all surgi- impartially quoted, and, while due credit is given | cal affections—is a sufficient reason for the liberal to each, their respective merits are weighed with patronage it has obtained. The editor, Dr. F. W. an unprejudiced hand. The grain of wheat is pre- i Sargent, has contributed much to enhance the value of the work, by such American improvements as are calculated more perfectly to adapt it to our own views and practice in this country. It abounds everywhere with spirited and life-like illustrations, which to the young surgeon, especially, are of no minor consideration. Every medical man frequently needs just such a work as this, for immediate refer- ence in moments of sudden emergency, when he has not time to consult more elaborate treatises.—The Ohio Medical and Surgical Journal. No work, in our opinion, equals it in presenting so much valuable surgical matter in so small a sompass.—St. Louis Med. and Surgical Journal. served, and the chaff is unmercifully stripped off. The arrangement is simple and philosophical, and the style, though clear and interesting, is so precise, that the book contains more information condensed into a few words than any other surgical work with which we are acquainted.—London Medical Times mnd Gazette. Druitt's Surgery is too well known to the Ameri- can medical profession to require its announcement anywhere. Probably no work of the kind has ever been more cordially received and extensively circu- lated than this. The fact that it comprehends in a comparatively small compass, all the essential ele- DALTON, JR. (J. C), M. D. Professor of Physiology in the College of Physicians, New York. A TREATISE ON HUMAN PHYSIOLOGY, designed for the use of Students and Practitioners of Medicine. With two hundred and fifty-four illustrations on wood. In one very beautiful octavo volume, of over 600 pages, extra cloth, $1 00; leather, raised bands, $4 25. (Now ready, Jan. 1859.) The object of the author has been to present a condensed view of the present condition of his subject, divested of mere theoretical views and hypothetical reasonings, but comprehending all important details which may be received as definitely settled. His long experience as an investi- gator and as a teacher has given him peculiar advantages in this, and he has endeavored wherever practicable to show the means by which results have been reached, so as to afford the student the means of pursuing original research, as well as a complete text-book of the science in its most ad- vanced condition. Of the numerous illustrations, all are original with the exception of eleven, so that the whole possesses a completeness and authority not otherwise attainable, and in the me- chanical execution every care has been taken to present one of the handsomest volumes as yet produced by the American press. To our mind, fulfils in a most admirable manner I pressed into a reasonable compass, embracing like- the objects contemplated by the author. The broad | wise the results of recent laborers in this department field of physiology has been traversed with discri- of our science.—Med. and Surg. Reporter, Jan. 29, mination, and its most valuable acquisitions com- | 1859. DUNGLISON, FORBES, TWEEDIE, AND CONOLLY. THE CYCLOPiEDIA OF PRACTICAL MEDICINE: comprising Treatises on the Nature and Treatment of Diseases, Materia Medica, and Therapeutics, Diseases of Women and Children, Medical Jurisprudence, &c. &c. In four large super-royal octavo volumes, of 3254 double-columned pages, strongly and handsomely bound, with raised bands. $12 00. * * This work contains no less than four hundred and eighteen distinct treatises, contributed by sixty-eight distinguished physicians, rendering it a complete library of reference for the country practitioner The most complete work on Practical Medicine extant; or, at least, in our language.—Buffalo Medical and Surgical Journal. For reference, it is above all price to every prac- titioner.—Western Lancet. One of the most valuable medical publications of the day__as a work of reference it is invaluable- Western Journal of Medicine and Surgery. It has been to us, both as learner and teacher, a work for ready and frequent reference, one in which modern English medicine is exhibited in the most advantageous light.—Medical Examiner. We rejoice that this work is to be placed within the reach of the profession in this country, it being unquestionably one of very great value to the prac- titioner. This estimate of it has not been formed from a hasty examination, but after an intimate ac- quaintance derived from frequent consultation of it during the past nine or ten years. The editors are practitioners of established reputation, and the list of contributors embraces many of the most eminen professors and teachers of London, Edinburgh, Dub- lin, and Glasgow. It is, indeed, the great merit of this work that the principal articles have been fur- nished by practitioners who have not only devoted especial attention to the diseases about which they have written, but have also enjoyed opportunities for an extensive practical acquaintance with them, and whose reputation carries the assurance of their competency justly to appreciate the opinions of others, while it stamps their own doctrines with high and just authority.—American Medical Journ. DEWEES'S COMPREHENSIVE SYSTEM OF MIDWIFERY. Illustrated by occasional cases and many ensravings. Twelfth edition, with the author's last improvements and corrections In one octavo volume, extra cloth, of 600 pages. S3 20. DEWEES'S TREATISE ON THE PHYSICAL AND MEDICAL TREATMENT OF CHILD- REN. The last edition. In one volume, octavo, extra cloth, 548 pages. $2 80. DEWEES'S TREATISE ON THE DISEASES OF FEMALES. Tenth edition. In one volume, octavo xtra cloth, 532 pages, with plates. S3 00. DANA ON ZOOPHYTES AND CORALS. In one volume, imperial quarto, extra cloth, with wood- cuts. $15 00. Also, AN ATLAS, in one volume, imperial folio, with sixty-one magnificent colored plates. Bound in half morocco. 830 00. DE LA BECHE'S GEOLOGICAL OHSERVER. In one very large and handsome octavo volume, ex- tra cloth, of 700 pages, with 300 wood-cuts. $4 00. FRICK ON RENAL AFFECTIONS; their Diag- nosis and Pathology. With illustrations. One volume, royal 12mo., extra cloth. 75 cents. 12 BLANCHARD <& LEA'S MEDICAL DUNGLISON (ROBLEY), M.D., Professor of Institutes of Medicine in the Jefferson Medical College, Philadelphia. NEW AND ENLARGED EDITION, Now Ready, 1858. MEDICAL LEXICON; a Dictionary of Medical Science, containing a concise Explanation ofthe various Subjects and Terms of Anatomy, Physiology, Pathology, Hygiene, Therapeutics. Pharmacology, Pharmacy, Surgery, Obstetrics, Medical Jurisprudence, Dentistry, &c. Notices of Climate and of Mineral Waters; Formulae for Officinal, Empirical, and Dietetic Preparations, &c. With French and other Synonymes. Fifteenth edition, revised and very greatly enlarged. In one very large and handsome octavo volume, of 992 double-columned pages, in small type; strongly bound in leather, with raised bands. Price $4 00. Especial care has been devoted in the preparation of this edition to render it in every respect worthy a continuance of the very remarkable favor which it has hitherto enjoyed. The rapid sale of Fifteen large editions, and the constantly increasing demand, show that it is regarded by the profession as the standard authority. Stimulated by this fact, the author has endeavored in the present revision to introduce whatever might be necessary " to make it a satisfactory and desira- ble—if not indispensable—lexicon, in which the student may search without disappointment for every term that has been legitimated in the nomenclature of the science." To accomplish this, large additions have been found requisite, and the extent of the author's labors may be estimated from the fact that about Six Thousand subjects and terms have been introduced throughout, ren- dering the whole number of definitions about Sixty Thousand, to accommodate which, the num- ber of pages has been increased by nearly a hundred, notwithstanding an enlargement in the size ofthe pase. The medical press, both in this country and in England, has pronounced the work in- dispensable to all medical students and practitioners, and the present improved edition will not lose that enviable reputation. The publishers have endeavored to render the mechanical execution worthy of a volume of such universal use in daily reference. The greatest care hns been exercised to obtain the typographical accuracy so necessary in a work of the kind. By the small but exceedingly clear type employed, an immense amount of matter is condensed in its thousand ample pages, while the binding will be found strong and durable. With all these improvements and enlargements, the price has been kept at the former very moderate rate, placing it within the reach of all. This work, the appearance of the fifteenth edition of which, it has become our duty and pleasure to announce, is perhaps the most stupendous monument of labor and erudition in medical literature. One would hardly suppose after constant use of the pre- ceding editions, where we have never failed to find a sufficiently full explanation of ever) medical term, that in this edition " about six thousand subjects and terms have been added,''' with a careful revision and correction of the entire work. It is only neces- sary to announce the advent of this edition to make it occupy the place ofthe preceding one on the table of every medical man, as it is without doubt the best and most comprehensive work of the kind which has ever appeared.—Buffalo Med. Journ., Jan. 1858. The work is a monument of patient research, skilful judgment, and vast physical labor, that will perpetuate the name of the author more effectually than any possible device of stone or metal. Dr, Dunglison deserves the thanks not only of the Ame- rican profession, but of the whole medical world.— North Am. Medico-Chir. Review, Jan. 1858. A Medical Dictionary better adapted for the wants of the profession than any other with which we are acquainted, and of a character which places it far above comparison and competition.—Am. Journ. Med. Sciences, Jan. 1858. We need only say, that the addition of 6,000 new terms, with their accompanying definitions, may be said to constitute a new work, by itself. We have examined the Dictionary attentively, and are most happy to pronounce it unrivalled of its kind. The erudition displayed, and the extraordinary industry which must have been demanded, in its preparation and perfection, redound to the lasting credit of its author, and have furnished us with a volume indis- pensable at the present day, to all who would find themselves au niveau with the highest standards of medical information.—Boston Medical and Surgical Journal, Dec. 31, 1857. Good lexicons and encyclopedic works generally, are the most labor-saving contrivances which lite- rary men enjoy; and the labor which is required to produce them in the perfect manner of this example is something appalling to contemplate. The author tells us in his preface that he has added about six thousand terms and subjects to this edition, which, before, was considered universally as the best work of the kind in any language.—Silliman's Journal, March, 1858. He has razed his gigantic structure to the founda- tions, and remodelled and reconstructed the entire pile. No less than six thousand additional subjects and terms are illustrated and analyzed in this new edition, swelling the grand aggregate to beyond sjxty thousand ! Thus is placed before the profes- sion a complete and thorough exponent of medical terminology, without rival or possibility of rivalry. —Nashville Journ. of Med. and Surg., Jan. 1858. It is universally acknowledged, we believe, that this work is incomparably the best and most com- plete Medical Lexicon in the English language. The amount of labor which the distinguished author has bestowed upon it is truly wonderful, and the learning and research displayed in its preparation are equally remarkable. Comment and commenda- tion are unnecessary, as no one at the present day thinks of purchasing any other Medical Dictionary than this.—St. Louis Med. and Surg. Journ., Jan. 1858. It is the foundation stone of a good medical libra- ry, and should always be included in the first list of books purchased by the medical student.—Am. Med. Monthly, Jan. 1858. A very perfect work of the kind, undoubtedly the most perfect in the English language.—Med. and Surg. Reporter, Jan. 1858. It is now emphatically the Medical Dictionary of the English language, and for it there is no substi- tute.—N. H. Med. Journ., Jan. 1858. It is scarcely necessary to remark that any medi- cal library wanting a copy of Dunglison's Lexicon must be imperfect.—Cin. Lancet, Jan. 1858. We have ever considered it the best authority pub- lished, and the present edition we may safely say has no equal in the world.—Peninsular Med. Journal, Jan.1858. The most complete authority on the subject to be found in any language.— Va.Med. Journal, Feb. '58. BY THE SAME AUTHOR. THE PKACTICE OP MEDICINE. A Treatise on Special Pathology and The- rapeutics. Third Edition. In two large octavo volumes, leather, of 1,500 pages. $6 25. AND SCIENTIFIC PUBLICATIONS. 13 DUNGLISON (ROBLEY), M.D., Professor of Institutes of Medicine in the Jefferson Medical College, Philadelphia. HUMAN PHYSIOLOGY. Eighth edition. Thoroughly revised and exten- sively modified and enlarged, with five hundred and thirty-two illustrations. In two large and handsomely printed octavo volumes, leather, of about 1500 pages. $7 00. In revising this work for its eighth appearance, the author has spared no labor to render it worthy a continuance ofthe very great favor which has been extended to it by the profession. The whole contents have been rearranged, and to a great extent remodelled ; the investigations which of late years have been so numerous and so important, have been carefully examined and incorporated, and the work in every respect has been brought up to a level with the present state of the subject. The object of the author has been to render it a concise but comprehensive treatise, containing the whole body of physiological science, to which the student and man of science can at all times refer with the certainty of finding whatever they are in search of, fully presented in all its aspects; and on no former edition has the author bestowed more labor to secure this result. We believe that it can truly be said, no more com- plete repertory of tacts upon the subject treated, can anywhere be found. The author lias, moreover, that enviable tact at description and that facility and ease of expression which render him peculiarly acceptable to the casual, or the studious reader This faculty, so requisite in setting forth many erraver and less attractive subjects, lends additional charms to one always fascinating.—Boston Med. and Surg. Journal, Sept. 1856. The best work of the kind in the English lan- guage.—Silliman's Journal. The present edition the author has made a perfect mirror of the science as it is at the present hour. As a work upon physiology proper, the science of the functions performed by the body, the student will find it all he wishes.—Nashville Journ. of Med. Sept. 1856. That he has succeeded, most admirably succeeded in his purpose, is apparent from the appearance of The most complete and satisfactory system of j an eighth edition. It is now the great encyclopaedia Physiology in the English language.—Amer. Med on the subject, and worthy of a place in every phy- Journal. [ sician's library.— Western Lancet, Sept. 1856. BY the same author. (Now Ready.) GENERAL THERAPEUTICS AND MATERIA MEDICA; adapted for a Medical Text-book. With Indexes of Remedies and of Diseases and their Remedies. Sixth Edition, revised and improved. With one hundred and ninety-three illustrations. In two large and handsomely printed octavo vols., leather, of about 1100 pages. $6 00. From the Author's Preface. " Another edition of this work being called for, the author has subjected it to a thorough and careful revision. It has been gratifying to him that it has been found so extensively useful by those for whom it was especially intended, as to require that a.sizth edition should be issued in so short a time afier the publication of a fifth. Grateful for the favorable reception of the work by the profession, he has bestowed on the preparation of the present edition all those cares which were demanded by the former editions, and has spared no pains to render it a faithful epitome of General Therapeutics and Materia Medica. The copious Indexes of Remedies and of Diseases and their Remedies can- not fail, the author conceives, to add materially to the value of the work." This work is too widely and too favorably known to require more than the assurance that the author has revised it with his customary industry, introducing whatever has been found necessary to bring it on a level with the most advanced condition ofthe subject. The number of illustration's has been somewhat enlarged, and the mechanical execution of the volumes will be found to have undergone a decided improvement. The work will, we have little doubt, be bought and read by the majority of medical students; its size, arrangement, and reliability recommend it to all; no one, we venture to preilict, will study it without profit, and there are few to whom it. will not be in some measure useful as a work of refer- ence. The young practitioner, more especially, will find the copious indexes appendid to this edition of great assistance in the selection and pieparation of suitable formulas.— Charleston Med. Journ. and Re- view, Jan. 1858. In announcing a new edition of Dr. Dunglison's General Ttierapeutics and Materia Medica, we have no words of commendation to bestow upon a work whose merits have been heretofore so often and so justly extolled. It must not be supposed, however, that the present is a mere reprint of the previous edition: the character of the author for laborious research, judicious analysis, and clearness of ex- pression, is fully sustained by the numerous addi- tions he has made to the work, and the careful re- Vision to which he has subjected the whole.—JV. A. Medico-Chir. Review, Jan. 1858. by the same author. (.4 new Edition.) NEW REMEDIES, WITH FORMULAE FOR THEIR PREPARATION AND ADMINISTRATION. Seventh edition, with extensive Additions. In one very large octavo volume, leather, of 770 pages. $3 75. Another edition of the " New Remedies" having been called for, the author has endeavored to add everything of moment that has appeared since the publication of the last edition. The articles treated of in the former editions will be found to have undergone considerable ex- pansion in this, in order that the author might be enabled to introduce, as far as practicable, the results of the subsequent experience of others, as well as of his own observation and reflection ; and to make the work still more deserving of the extended circulation with which the preceding editions have been favored by the profession. By an enlargement of the page, the numerous addi- tions have been incorporated without greatly increasing the bulk of the volume.—Preface. The great learning of the author, and his remark- able industry in pushing his researches into every source whence information is derivable,have enabled him to throw together an extensive mass of facts and statements, accompanied by full reference to authorities; which last feature renders the work practically valuable to investigators who desire tp examine the original papers.—The American Journal of Pharmacy. One of the most useful of the author's works.— Southern Medical and Surgical Journal. This elaborate and useful volume should be found in every medical library, for as a book of re- ference, for physicians, it is unsurpassed by any other work in existence, and the double index for diseases and for remedies, will be found greatly to enhance its value.—New York Med. Gazette. 14 BLANCHARD & LEA'S MEDICAL ERICHSEN (JOHN), Professor of Surgery in University College, London, &c. THE SCIENCE AND ART OF SURGERY; being a Treatise on Surgical Injuries, Diseases, and Operations. New and improved American, from the second enlarged and carefully revised London edition. Illustrated with over four hundred engravings on wood. In one large and handsome octavo volume, cf one thousand closely printed pages, leather, raised bands. $4 50. (Now Ready, January, 1859.) The very distinguished favor with which this work has been received on both sides ofthe Atlan- tic has stimulated the author to render it even more worthy of the position which it has so rapidly attained as a standard authority. Every portion has been carefully revised, numerous additions have been made, and the most watchful care has been exercised to render it a complete exponent ofthe most advanced condition of surgical science. In this manner the work has been enlarged by about a hundred pages, while the series of engravings has been increased by more than a hundred, rendering it one of the most thoroughly illustrated volumes before the profession. The additions of the author having rendered unnecessary most of the notes of the former American editor, but little has been added in this country; some few notes and occasional illustrations have, however, been introduced to elucidate American modes of practice. It is, in our humble judgment, decidedly the best Dook of the kind in the English language. Strange that just such books are notoftener produced by pub- lic teachers of surgery in this country and Great Britain. Indeed, it is a matter of great astonishment, hut no less true than astonishing, that ofthe many works on surgery republished in this country within the last fifteen or twenty years as text-books for medical students, this is the only one that even ap- proximates to the fulfilment of the peculiar wants of youngmen just entenngupon the study of this branch of the profession.— Western Jour .of Med. and Surgery. Its value is greatly enhanced by a very copious well-arranged index. We regard this as one of the most valuable contributions to modern surgery. To one entering his novitiate of practice, we regard it the most serviceable guide which he can consult. He will find a fulness of detail leading him through every step ofthe operation, and not deserting him until the final issue ofthe case is decided. For the same rea- son we recommend it to those whose routine of prac- tice lies in such parts of the country that they must rarely encounter cases requiring surgical manage- ment.—Stethoscope. Embracing, as will be perceived, the whole surgi- cal domain, and each division of itself almost com- plete and perfect, each chapterfull and explicit, each subject faithfully exhibited, we can only express oui estimate of it in the aggregate. We consider it an excellent contribution to surgery, as probably the best single volume now extant on the subject, and with great pleasure we add it to onr text-books.— Nashville Journal of Medicine and Surgery. Prof. Erichsen's work, for its size, has not been surpassed; his nine hundred and eight pages, pro- fusely illustrated, are rich in physiological, patholo- gical, and operative suggestions, doctrines, details, and processes; and will prove a reliable resource for information, both to physician and surgeon, in the hour of peril.—JV. 0. Med. and Surg. Journal. We are acquainted with no other work wherein so much good sense, sound principle, and practical inferences, stamp every page.—American Lancet. ELLIS (BENJAMIN), M.D. THE MEDICAL FORMULARY: being a Collection of Prescriptions, derived from the writings and practice of many of the most eminent physicians of America and Europe. Together with the usual Dietetic Preparations and Antidotes for Poisons. To which is added an Appendix, on the Endermic use of Medicines, and on the use of Ether and Chloroform. The whole accompanied with a few brief Pharmaceutic and Medical Observations. Tenth edition, revised and much extended by Robert P. Thomas, M. D., Professor of Materia Medica in the Philadelphia College of Pharmacy. In one neat octavo volume, extra cloth, of 296 pages. $1 75. FOWNES (GEORGE), PH. D., &c. ELEMENTARY CHEMISTRY; Theoretical and Practical. With numerous illustrations. Edited, with Additions, by Robert Bridges, M. D. In one large royal 12mo. In leather, $1 50; extra cloth, $ 1 35. The work of Dr. Fownes has long been before the public, and its merits have been fully appreci- ated as the best text-book on chemistry now in existence. We do not, of course, place it in a rank superior to the works of Brande, Graham, Turner, Gregory, or Gmelin, but we say that, as a work for students, it is preferable to any of them.—Lon- don Journal of Medicine. A work well adapted to the wants of the student It is an excellent exposition of the chief doctrines and facts of modern chemistry. The size of the work, and still more the condensed yet perspicuous style in which it is written, absolve it from the charges very properly urged against most manuals termed popular.—Edinburgh Journal of Medical Science. volume, of over 550 pages, with 181 wood-cuts We know of no better text-book, especially in the difficult department of organic chemistry, upon which it is particularly full and satisfactory. We would recommend it to preceptors as a capital " office book" for their students who are beginners in Chemistry. It is copiously illustrated with ex- cellent wood-cuts, and altogether admirably "got up."—N. J. Medical Reporter. A standard manual, which has long enjoyed the reputation of embodying much knowledge in a small space. The author has achieved the difficult task of condensation with masterly tact. His book is con- cise without being dry, and brief without being too dogmatical or general.—Virginia Med. and Surgical Journal. FISKE FUND PRIZE ESSAYS. THE EFFECTS OF CLIMATE ON TUBERCULOUS DISEASE. By Edwin Lee, M. R. C. S., London, and THE INFLUENCE OF PREGNANCY ON THE DEVELOP- MENT OF TUBERCLES. By Edward Warren, M. D., of Edenton, N. C. Together in one neat octavo volume, extra cloth. $1 00. (Just Ready.) FERGUSSON (WILLIAM), F. R. S., Professor of Surgery in King's College, London, &c. A SYSTEM OF PRACTICAL SURGERY. Fourth American, from the third and enlarged London edition. In one large and beautifully printed octavo volume, of about 700 pages, with 393 handsome illustrations, leather. $3 00. No work was ever written which more nearly I The addition of many new pages makes this work comprehended the necessities of the student and | more than ever indispensable to the student and prac- practitioner, and was more carefully arranged to titioner.—Ranking's Abstract. that single purpose than this.—N. Y. Med. Journal. | AND SCIENTIFIC PUBLICATIONS. 15 FLINT (AUSTIN), M. D., Professor of the Theory and Practice of Medicine in the University of Louisville, &c. (An Important New Work.) PHYSICAL EXPLORATION AND DIAGNOSIS OF DISEASES AFFECT- ING THE RESPIRATORY ORGANS. In one large and handsome octavo volume, extra cloth, 636 pages. $3 00. We can only state our general impression of the high value of this work, and cordially recommend it to all. We regard it, in point both of arrangement and of the marked ability of its treatment of the sub- jects, as destined to take the first rank in works of this class. So far asour information extends, it has at present no equal. To the practitioner, as well as the student, it will be invaluable in clearing up the diagnosis of doubtful cases, and in shedding light upon difficult phenomena.—Buffalo Med. Journal. This is the most elaborate work devoted exclu- sively to the physical exploration of diseases of the lungs, with which we are acquainted in the English language.—Boston Med. and Surg. Journal. A work of original observation of the highest merit. We recommend the treatise to every one who wishes to become a correct auscultator. Based to a very large extent upon eases numerically examined, it carries the evidence of careful study and discrimina- tion upon every pasje. It does credit to the autnor, and, through him, to the profession in this country. It is, what we cannot call every book upon auscul- tation, a readable book.—Am. Jour. Med. Sciences. NOW COMPLETE, GRAHAM (THOMAS), F. R. S., THE ELEMENTS OF INORGANIC CHEMISTRY, including the Applica- tions ofthe Science in the Arts. New and much enlarged edition, by Henry Watts and Robert Bridges, M. D. Complete in one large and handsome octavo volume, of over 800 very large pages, with two hundred and thirty-two wood-cuts, extra cloth. $4 00. **,£ Part II., completing the work from p. 431 to end, with Index, Title Matter, &c, may be had separate, cloth backs and paper sides. Price $2 50. The long delay which has intervened since the appearance of the first portion of this work, has rendered necessary an Appendix, embodying the numerous and important investigations and dis- coveries of the last few years in the subjects contained in Part I. This occupies a large portion of Part II., and will be found to present a complete abstract of the most recent researches in the general principles of the science, as well as all details necessary to bring the whole work thoroughly up to the present time in all departments of Inorganic Chemistry. Gentlemen desirous of completing their copies of the work are requested to apply for Part II. without delay. It will be sent by mail, prepaid, on receipt of the amount, $2 50. From Prof. Wolcott Gibbs, N. Y. Free Academy. May 25, 1658. The work is an admirable one in all respects,and its republication here cannot fail to exert a positive influence upon the progress of science in this country. From Prof. E. N. Horsford. Harvard College, April 27, 185S. It has, in its earlier and less perfect editions, been familiar to me, and the excellence of its plan and the clearness and completeness of its discussions, have long been my admiration. It is a very acceptable addition to the library of standard boohs of every chemical student. No reader of English works on this science can afford to be without this edition of Prof. Graham's Elements.— Silliman's Journal, March, 1858. From Prof. J. L. Crawcour, New Orleans School of Medicine, May 9, 1858. It is beyond all questicn the best systematic work on Chemistry in the English language, and I am gratified to find that an American edition at a mo- derate price has been issued, so as to place it within the means of students. It will be the only text-book 1 shall now recommend to my class. GRIFFITH (ROBERT E.), M. D., &.c. A UNIVERSAL FORMULARY, containing the methods of Preparing and Ad- ministering Officinal and other Medicines. The whole adapted to Physicians and Pharmaceu- tists. Second Edition, thoroughly revised, with numerous additions, by Robert P. Thomas, M. D. Professor of Materia Medica in the Philadelphia College of Pharmacy. In one large and handsome octavo volume, extra cloth, of 650 pages, double columns. " $3 25. 00; or bound in sheep, It was a work requiring much perseverance, and when published was looked upon as by far the best work of its kind that had issued from the American press. Prof Thomas has certainly "improved," as well as added to this Formulary, and has rendered it additionally deserving of the confidence of pharma- ceutists and physicians.—Am. Journal of Pharmacy. We are happy to announce a new and improved edition of this, one ofthe most valuable and useful works that have emanated from an American pen. It would do credit to any country, and will be found of daily usefulness to practitioners of medicine; it is better adapted to their purposes than the dispensato- ries.— Southern Med. and Surg. Journal. It is one ofthe most useful books a country practi- nistering medicines that can be desired by the physi- cian and pharmaceutist.— Western Lancet. The amountof useful, every-day matter.for a prac ticing physician, is really immense.—Boston Med. and Surg. Journal. We predict a great sale for this work, and we espe- cially recommend it to all medical teachers.—Rich- mond Stethoscope. This edition of Dr. Griffith's work has been grea'ly improved by the revision and ample additions of Dr. Thomas, and is now, we believe, one of the most complete works of its kind in any language. The additions amount to about seventy pages, and no effort has been spared to include in them all the re- cent improvements which have been published in tionercan possibly have in his possession.—Medical medical journals, and systematic treatises. A work Chronicle. of this kind appears to us indispensable 1o the physi- This is a work of six hundred and fifty-one pages, cian, and there is none we can more cordially recom- embracin«- all on the subject of preparing and admi- mend.—N. Y. Journal of Medicine. GLUGE (GOTTLIEB), M. D., Professor of Physiology and Pathological Anatomy in the University of Brussels, &c. AN ATLAS OF PATHOLOGICAL HISTOLOGY. Translated, with Notes and Additions, by Joseph Leidy, M. D., Professor of Anatomy in the University of Pennsylva- nia. In one volume, very large imperial quarto, extra cloth, with 320 figures, plain and colored, on twelve copperplates. $5 00. 16 BLANCHARD & LEA'S MEDICAL GROSS (SAMUEL D.), M. D., Professor of Surgery in the Jefferson Medical College of Philadelphia, tec. New Edition (Now Ready.) ELEMENTS OF PATHOLOGICAL ANATOMY. Third edition, thoroughly revised and greatly improved. In one large and very handsome octavo volume, with about three hundred and fifty beautiful illustrations, of which a large number are from original drawings. Price in extra cloth, $4 75; leather, raised bands, $5 25. The very rapid advances in the Science of Pathological Anatomy during the last few years have rendered essential a thorough modification of this work, with a view of making it a correct expo- nent of the present state of the subject. The very careful manner in which this task has been executed, and the amount of alteration which it has undergone, have enabled the author to say that '' with the many changes and improvements now introduced, the work may be regarded almost as a new treatise," while the efforts of the author have been seconded as regards the mechanical execution of the volume, rendering it one of the handsomest productions of the American press. A very large number of new and beautiful original illustrations have been introduced, and the work, it is hoped, will fully maintain the reputation hitherto enjoyed by it of a complete and practical ex- position of its difficult and important subject. We most sincerely congratulate the author on the We have been favorably impressed with the gene- successful manner in which he has accomplished his ral manner in which Dr. Gross has executed his task proposed object. His book is most admirably cal- of affording a comprehensive digest of the present culated to fill up a blank which has long been felt to state ofthe literature of Pathological Anatomy, and exist in this department of medical literature, and as such must become very widely circulated amongst all-classes of the profession. — Dublin Quarterly Journ. of Med. Science, Nov. 1857. have much pleasure in recommending his work to our readers, as we believe one well deserving ot diligent perusal and careful study.—Montreal Med. Chron., Sept. 1857. BY THE SAME AUTHOR. A PRACTICAL TREATISE ON THE DISEASES, INJURIES, AND MALFORMATIONS OF THE URINARY BLADDER, THE PROSTATE GLAND, AND THE URETHRA. Second Edition, revised and much enlarged, with one hundred and eighty- four illustrations. In one large and very handsome octavo volume, of over nine hundred pages. m leather, raised bands, $5 25; extra cloth, $4 75. A volume replete with truths and principles ofthe utmost value in the investigation of these diseases.— American Medical Journal. On the appearance of the first edition of this work, the leading English medical review predicted that it would have a " permanent place in the literature of surgery worthy to rank with the best works of the present age." This prediction has been amply ful- filled. Dr. Gross's treatise has been found to sup- ply completely the want which has been felt ever since the elevation of surgery to the rank of a science, of a good practical treatise on the diseases of the bladder and its accessory organs. Philosophical in its design, methodical in its arrangement, ample and sound in its practical details, it may in truth be said to leave scarcely anything to be desired on so im- portant a subject, and with the additions and modi- fications resulting from future discoveries and im- provements, it will probably remain one of the most valuable works on this subject so long as the science of medicine shall exist.—Boston Med. and Surg. Journal. Whoever will peruse the vast amount of valuable practical information it contains, and which we have been unable even to notice, will, we think, agree with us, that there is no work in the English language which can make any just pretensions to be its equal.—iV. Y. Journal of Medicine. BY THE SAME AUTHOR. A PRACTICAL TREATISE ON FOREIGN BODIES IN THE AIR-PAS- SAGES. In one handsome octavo volume, extra cloth, with illustrations, pp. 468. $2 75, BY THE SAME AUTHOR. (In press.) A SYSTEM OF SURGERY j Diagnostic, Pathological, Therapeutic, and Opera- tive. With very numerous engravings on wood. In two large octavo volumes. GIBSON (WILLIAM), M. D., Late Professor of Surgery in the University of Pennsylvania, &c INSTITUTES AND PRACTICE OF SURGERY; being Outlines of a Course of Lectures. Eighth edition, improved and altered. With thirty-four pla'es. In two handsome octavo volumes, containing about 1000 pages, leather, raised bands. $6 50. GRAY (HENRY), F. R. S., Lecturer on Anatomy at St. George's Hospital, London, &c. ANATOMY, DESCRIPTIVE AND SURGICAL. The Drawings by H. V. Carter, M. D., late Demonstrator on Anatomy at St. George's Hospital; the Dissections jointly by the Author and Dr. Carter. In one magnificent imperial octavo volume, with 363 large and elaborate engravings on wood. (At press.) The author has endeavored in this work to cover a more extended range of subjects than is customary in the ordinary text-books, by giving not only the details necessary for the student, but also ihe application of those details in the practice of medicine and surgery, thus rendering it not only a guide for the learner, but an admirable work of reference for the active practitioner. The engravings form a special feature in the work, many of them being the size of nature, nearly all original, and having the names of the various parts printed on the body of the cut, in place of figures of reference with descriptions at the foot. They thus form a complete and splendid series, which will greatlv assist the student ia obtaining a clear idea of Anatomy, and will also serve to refresh the memory of those who may find in the exigencies of practice the necessity of recalling the de- tails of tne dissecting room. GARDNER'S MEDICAL CHEMISTRY, for the use of Students and the Profession. In one royal l'2mo. vol., ex. cloth, pp. 396, with illustrations. 81 00. HUGHES' CLINICAL INTRODUCTION TO THE PRACTICE OF AUSCULTATION AND OTHER MODES OF PHYSICAL DIAGNOSIS, IN DISEASES OF THE LUNGS AND HEART. Second American, from the second London edition. 1 vol. royal l'imo., ex. cloth, pp. 304. $1 00. AND SCIENTIFIC PUBLICATIONS. 17 HOBLYN (RICHARD D.), M. D. A DICTIONARY OF THE TERMS USED IN MEDICINE AND THE COLLATERAL SCIENCES. A new American edition. Revised, with numerous Additions, by Isaac Hays, M. D., editor of the " American Journal ofthe Medical Sciences." In one large royal 12mo. volume, leather, of over 500 double columned pages. $1 50. If the frequency with which we have referred to this volume since its reception from the publisher, two or three weeks ago, be any criterion for the future, the binding will goon have to be renewed, even with cureful handling. We find that Dr. Hays has done the profession great service by his careful and industrious labors. The Dictionary has thus become eminently suited to our medical brethren in this country. The additions by Dr. Hays are in brackets, and we believe there is not a single page bnt bears these insignia; in every instance which we have thus far noticed, the additions are really needed and ex- ceedingly valuable. We heartily commend the work to all who wish to be au courant in medical termi- nology.— Boston Med. and Surg. Journal. To both practitioner and student, we recommend this dictionary as being convenient in size, accurate in definition, and sufficiently full and complete for ordinary consultation.—Charleston Med. Journ. and Review. Hoblyn has always been a favorite dictionary, and in its present enlarged and improved form will give greater satisfaction than ever. The American editor, Dr. Hays, has made many very valuable additions. —N. J. Med. Reporter. To supply the want of the medical reader arising from this cause, we know of no dictionary better arranged and adapted than the one bearing the above title. It is not encumbered with the obsolete terms of a bygone age, but it contains all that are now in use ; embracing every department of medical science down to the very latest date. The volume is of a convenient size to be used by the medical student, and yet large enough to make a respectable appear- ance in the library of a physician.—Western Lancet. Hoblyn's Dictionary has long been a favorite with us. It is the best book of definitions we have, and ought always to be upon the student's table.— Southern Med. and Surg. Journal. HUNTER (JOHN). (See "Ricord," page 26, for Ricord's edition of Hunter on Venereal.) HOLLAND (SIR HENRY), BART., M.D..F. R. S., Physician in Ordinary to the Queen of England, &c. MEDICAL NOTES AND REFLECTIONS. From the third London edition. In one handsome octavo volume, extra cloth, $3 00. HABERSHON (S. O.), M. D., Assistant Physician to and Lecturer on Materia Medica and Therapeutics at Guy's Hospital, &c. PATHOLOGICAL AND PRACTICAL OBSERVATIONS ON DISEASES OF THE ALIMENTARY CANAL, CESOPHAGUS, STOMACH, O/ECUM, AND INTES- TINES. With illustrations on wood. In one handsome octavo volume. (Publishing in the Medical News and Library for 1858 and 1859.) HORNER (WILLIAM E.), M. D., Professor of Anatomy in the University of Pennsylvania. SPECIAL ANATOMY AND HISTOLOGY. Eighth edition. Extensively revised and modified. In two large octavo volumes, extra cloth, of more than one thousand pages, handsomely printed, with over three hundred illustrations. $6 00. ANATOMICAL ATLAS. page 28. BY THE SAME AUTHOR. By Professors Horner and Smith. See Smith, HAMILTON (FRANK H.), M. D., Professor of Surgery, in Buffalo Medical College, &c. A COMPLETE TREATISE ON FRACTURES AND DISLOCATIONS. one handsome octavo volume, with several hundred illustrations. (Preparing.) In JONES (T. WHARTON), F. R. S., Professor of Ophthalmic Medicine and Surgery in University College, London, &c. THE PRINCIPLES AND PRACTICE OF OPHTHALMIC MEDICINE AND SURGERY. With one hundred and ten illustrations. Second American from the second and revised London edition, with additions by Edward Hartshorne, M. D., Surgeon to Wills' Hospital, &c. In one large, handsome royal 12mo. volume, extra cloth, of 500 pages. $1 50. We are confident that the reader will find, on perusal, that the execution ofthe work amply fulfils the promise of the pre/ace, and sustains, in every point, the already high reputation of the author as an ophthalmic surgeon as well as a physiologist and pathologist. The book is evidently the result of much labor and research, iind has been written with the greatest care and attention; it possesses that best quality which a general work, like a sys- tem or manual can show, viz : the quality of having all the materials whencesoever derived, so thorough- ly wrought up, and digested in the author's mind, as to come forth with the freshness and impressive- ness of an original production. We entertain little doubt that this book will become what its author hoped it might become, a manual for daily reference and consultation by the student and the general prac- titioner. The work is marked by that correctness, clearness, and precision of style which distinguish all the productions of the learned author.—British and For. Med. Review. 18 BLANCHARD & LEA'S MEDICAL JONES (C. HANDFIELD), F. R. S., &. EDWARD H. SIEVEKING, M.D., Assistant Physicians and Lecturers in St. Mary's Hospital, London. A MANUAL OF PATHOLOGICAL ANATOMY. First American Edition, Revised. With three hundred and ninety-seven handsome wood engravings. In one large and beautiful octavo volume of nearly 750 pages, leather. $3 75. As a concise text-book, containing, in a condensed form, a complete outline of what is known in the domain of Pathological Anatomy, it is perhaps the best work in the English language. Its great merit consists in its completeness and brevity, and in this respect it supplies a great desideratum in our lite- rature. Heretofore the student of pathology was obliged to glean from a great number of monographs, and the field was so extensive that but few cultivated it with any degree of success. As a simple work of reference, therefore, it is of great value to the student of pathological anatomy, and should be in every physician's library.—Western Lancet. In offering the above titled work to the public, the authors have not attempted to intrude new views on their professional brethren, but simply to lay before them, what has long been wanted, an outline of the present condition of pathological anatomy. In this they have been completely successful. The work is one of the best compilations which we have ever perused.—Charleston Medical Journal and Review. We urge upon our readers and the profession gene- rally the importance of informing themselves in re- gard to modern views of pathology, and recommend to them to procure the work before us as the best means of obtaining this information.—Stethoscope. From the casual examination we have given we are inclined to regard it as a text-book, plain, ra- tional, and intelligible, such a book as the practical man needs for daily reference. For this reason it will be likely tp be largely useful, as it suits itself to those busy men who have little time for minute investigation, and prefer a summary to an elaborate treatise.—Buffalo Medical Journal. KIRKES (WILLIAM SENHOUSE), M. D., Demonstrator of Morbid Anatomy at St. Bartholomew's Hospital, &c. A MANUAL OF PHYSIOLOGY. A new American, from the third and improved London edition. With two hundred illustrations. In one large and handsome royal 12mo. volume, leather, pp. 586. $2 00. (Just Issued, 1857.) In again passing this work through his hands, the author has endeavored to render it a correct exposition of the present condition of the science, making such alterations and additions as have been dictated by further experience, or as the progress of investigation has rendered desirable. In every point of mechanical execution the publishers have sought to make it superior to former edi- tions, and at the very low price at which it is offered, it will be found one of the handsomest and cheapest volumes before the profession. In making these improvements, care has been exercised not unduly to increase its size, thus maintaining its distinctive characteristic of presenting within a moderate compass a clear and con- nected view of its subjects, sufficient for the wants of the student. One of the very best handbooks of Physiology we possess—presenting just such an outline of the sci- This is a new and very much improved edition of Dr. Kirkes' well-known Handbook of Physiology. Originally constructed on the basis of the admirable treatise of Muller, it has in successive editions de- veloped itself into an almost original work, though no change has been made in the plan or arrangement. It combines conciseness with completeness, and is, therefore, admirably adapted for consultation by the busy practitioner.—Dublin Quarterly Journal, Feb. 1857. Its excellence is in its compactness, its clearness, and its carefully cited authorities. It is the most convenient of text-books. These gentlemen, Messrs. Kirkes and Paget, have really an immense talent for silence, which is not so common or so cheap as prat- ing people fancy. They have the gift of telling us what we want to know, without thinking it neces- sary to tell us all they know.—Boston Med. and Surg. Journal, May 14, 1857. ence, comprising an account of its leading facts and generally admitted principles, as the student requires during his attendance upon a course of lectures, or for reference whilst preparing for examination.— Am. Medical Journal. We need only say, that, without entering into dis- cussions of unsettled questions, it contains all the recent improvements in this department of medical science. For the student beginning this study, and the practitioner who has but leisure to refresh his memory, this book is invaluable, as it contains all that it is important to know, without special details, which are read with interest only by those who would make a specialty, or desire to possess a criti- cal knowledge of the subject.—Charleston Medical Journal. KNAPP'S TECHNOLOGY ; or, Chemistry applied to the Arts and to Manufactures. Edited, with numerous Notes and Additions, by Dr. Edmund Ronalds and Dr. Thomas Richardson. First American edition, with Notes and Additions, by Prof. Walter R. Johnson. In two handsome octavo volumes, extra cloth, with about 500 wood- engravings. $6 00. LUDLOW (J. L.), M. D. A MANUAL OF EXAMINATIONS upon Anatomy, Physiology, Surgery, Practice of Medicine, Obstetrics, Materia Medica, Chemistry, Pharmacy, and Therapeutics. To which is added a Medical Formulary. Designed for Students of Medicine throughout the United States. Third edition, thoroughly revised and greatly extended and enlarged. With three hundred and seventy illustrations. In one large and handsome royal 12mo. volume, leather, of over 800 closely printed pages. (Just Issued.) $2 50. The great popularity of this volume, and the numerous demands for it during the two years in which it has been out of print, have induced the author in its revision to spare no pains to render it a correct and accurate digest ofthe most recent condition of all the branches of medical science. In many respects it may, therefore, be regarded rather as a new book than a new edition, an entire section on Physiology having been added, as also one on Organic Chemistry, and many portions having been rewritten. A very complete series of illustrations has been introduced, and every care ha.s been taken in the mechanical execution to render it a convenient and satisfactory book for study or reference. The arrangement of the volume in the form of question and answer renders it especially suited for the office examination of students and for those preparing for graduation. We know of no better companion for the student crammed into his head by the various professors to during the hours spent in the lecture room, or to re- whom he is compelled to listen.—Western Lancet, fresh, at a glance, his memory of the various topics May, 1857. AND SCIENTIFIC PUBLICATIONS. 19 LEHMANN (C. G.) PHYSIOLOGICAL CHEMISTRY. Translated from the second edition by George E. Day, M. D., F. R. S., &c, edited by R. E. Rogers, M. D., Professor of Chemistry in the Medical Department of the University of Pennsylvania, with illustrations selected from Funke's Atlas of Physiological Chemistry, and an Appendix of plates. Complete in two large and handsome octavo volumes, extra cloth, containing 1200 pages, with nearly two hundred illus- trations. $6 00. This great work, universally acknowledged as the most complete and authoritative exposition of the principles and details of Zoochemistry, in its passage through the press, has received from Professor Rogers such care as was necessary to present it in a correct and reliable form. To such a work additions were deemed superfluous, but several years having elapsed between the appear- ance in Germany of the first and last volume, the latter contained a supplement, embodying nume- rous corrections and additions resulting from the advance ofthe science. These have all been incor- porated in the text in their appropriate places, while the subjects have been still further elucidated by the insertion of illustrations from the Atlas of Dr. OttoFunke. With the view of supplying the student with the means of convenient comparison, a large number of wood-cuts, from works on kindred subjects, have also been added in the form of an Appendix of Plates. The work is, therefore, pre- sented as in every way worthy the attention of all who desire to be familiar with the modern facts and doctrines of Physiological Science. The most important contribution as yet made to it treats.—Edinburgh Monthly Journal of Medical Science. Already well known and appreciated by the scien- tific world, Professor Lehmann's great work re- quires no laudatory sentences, as, under a new garb, it is now presented to us. The little space at our command would ill suffice to set forth even a small portion of its excellences.—Boston Med. and Surg. Journal, Dec. 1855. Physiological Chemistry.—Am. Journal Med. Sci tnces, Jan. 1856. The present volumes belong to the small class of medical literature which comprises elaborate works of the highest orderof merit.—Montreal Med. Chron- icle, Jan. 1856. The work of Lehmann stands unrivalled as the most comprehensive book of reference and informa- tion extant on every branch of the subject on which BY THE SAME AUTHOR. (Just Issued.) MANUAL OF CHEMICAL PHYSIOLOGY. Translated from the German, with Notes and Additions, by J. Cheston Morris, M. D., with an Introductory Essay on Vital Force, by Samuel Jackson, M. D., Professor of the Institutes of Medicine in the University of Pennsylvania. With illustrations on wood. In one very handsome octavo volume, extra cloth, of 336 pages. $2 25. From Prof. Jackson's Introductory Essay. In adopting the handbook of Dr. Lehmann as a manual of Organic Chemistry for the use of the students of the University, and in recommending his original work of Physiological Chemistry for their more mature studies, the high value of his researches, and the great weight of his autho- rity in that important department of medical science are fully recognized. LAWRENCE (W.), F. R. S., &.C. A TREATISE ON DISEASES OF THE EYE. A new edition, edited, with numerous additions, and 243 illustrations, by Isaac Hays, M. D., Surgeon to Will's Hospi- tal, &c. In one very large and handsome octavo volume, of 950 pages, strongly bound in leather ....'«U ..n.'tt^.l UnJa ur schools that its im- portance deserves. Our larger works are also very defective in their teaching on these small practical points. This little book ^ill supply the void which all must feel who have not studied its pages.—West- ern Lancet, March, 1856/ SKEY'S OPERATIVE SURGERY. In one very handsome octavo volume, extra cloth, of over 650 pages, with about one hundred wood-cuts. $3 25. STANLEY'S TREATISE ON DISEASES OF THE BONES. Inone volume, octavo, extra cloth, 286 pages. $1 50. SOLLY ON THE HUMAN BRAIN; its Structure, Physiology, and Diseases. From the Second and / much enlarged London edition. In one octavo volume, extra cloth, of 500 pages, with 1-20 wood- cuts. $2 00. / SIMON'S GENERAL PATHOLOGY, as conduc- ive to the Establishment of Rational Principles for the prevention and Cure of Disease. In one neat octavo volume, extra cloth, of 212 pages. $1 25. TANNER (T. H.), M. D., Physician to the Hospital for Women, &c. A MANUAL OF CLINICAL MEDICINE AND PHYSICAL DIAGNOSIS. To which is added The Code of Ethics of the American Medical Association. Second American Edition. In one neat volume, small 12mo., extra cloth, 87| cents. The work is an honor to its writer, and must ob-ttioners,it has only to be seen, to win for itself a tain a wide circulation by its intrinsic merit alone/1 place upon the shelves of every medical library. Suited alike to the wants of students and practf- |—Boston Med and Surg. Journal. Now Complete. TODD (ROBERT BENTLEY), M. D., F. R. S., Professor of Physiology in King's College, London; and WILLIAM BOWMAN, F. R. S., Demonstrator of Anatomy in King's College, London. THE PHYSIOLOGICAL ANATOMY AND PHYSIOLOGY OF MAN. With about three hundred large and beautiful illustrations on wood. Complete in one large octavo volume, of 950 pages, leather. Price $4 50. 1^° Gentlemen who have received portions of this work, as published in the " Medical News and Library," can now complete their copies, if immediate application be made. It will be fur- nished as follows, free by mail, in paper covers, with cloth backs. Parts I., II., III. (pp. 25 to 552), $2 50. Part IV. (pp. 553 to end, with Title, Preface, Contents, &c), $2 00. Or, Part IV., Section II. (pp. 725 to end, with Title, Preface, Contents, &c), $1 25. One of the very best books ever issued from any A magnificent contribution to British medicine, and the American physician who shall fail to peruse it, will have failed to read one of the most instruc- tive books of the nineteenth century.—N. O. Med. and Surg. Journal, Sept. 1857. It is more concise than Carpenter's Principles, and more modern than the accessible edition of Mailer's Elements; its details are brief, but sufficient; its descriptions vivid; its illustrations exact and copi- ous ; and its language terse and perspicuous.— Charleston Med. Journal, July, 1857. We know of no work on the subject of physiology so well adapted to the wants of the medical student. Its completion has been thus long delayed, that the authors might secure accuracy by personal observa- tion.—St. Louis Med. and Surg. Journal, Sept. '57. medical press. We think it indispensable to every reading medica) man, and it may, with all propriety, and with the utmost advantage be made a text-book by any student who would tnoroughly comprehend the groundwork of medicine.—N. O. Med. Newst June, 1857. Our notice, though it conveys but a very feeble and imperfect idea of the magnitude and importance of the work now under consideration, already tran- scends our limits ; and, with the indulgtnce of our readers, and the hope that they will peruse the book for themselves, as we feel we can with confidence recommend it, we leave it in their hands for them to judge of its merits.—The Northwestern Med. and Surg. Journal, Oct. 1857. TODD (R. B.), M. D., F. R. S., &c. CLINICAL LECTURES ON CERTAIN DISEASES OF THE URINARY ORGANS AND ON DROPSIES. In one octavo volume. (Just Issued, 1857.) $1 50. AND SCIENTIFIC PUBLICATIONS. 29 TAYLOR (ALFRED S.), M. D., F. R. S., Lecturer on Medical Jurisprudence and Chemistry in Guy's Hospital. MEDICAL JURISPRUDENCE. Fourth American Edition. With Notes and References to American Decisions, by Edward Hartshorne, M. D. In one large octavo volume, leather, of over seven hundred pages. $3 00. This standard work has lately received a very thorough revision at the hands ofthe author, who has introduced whatever was necessary to render it complete and satisfactory in carrying out the objects in view. The editor has likewise used every exertion to make it equally thorough with regard to all matter;- relating to the practice of this country. In doing this, he has carefully ex- amined all that has appeared on the subject since the publication ofthe last edition, and has incorpo- rated all the new information thus presented. The work has thus been considerably increased in size, notwithstanding which, it has been kept at its former very moderate price, and in every respect it will be found worthy of a continuance of the remarkable favor which has carried it through so many editions on both sides of the Atlantic. A few notices of the former editions are appended. we do not hesitate to affirm that after having once commenced its perusal, few could be prevailed upon to desist before completing it. In the last London edition, all the newly observed and accurately re- corded facts have been inserted, including much that is recent of Chemical, Microscopical, and Patholo- gical research, besides papers on numerous subjects never before published .-Charleston Medical Journal and Review. It is not excess of praise to say that the volume before us is the very best treatise extant on Medical Jurisprudence. In saying this, we do not wish to be understood as detracting from the merits of the excellent works of Beck, Ryan, Traill, Guy, and others; but in interest and value we think it must be conceded that Taylor is superior to anything that has preceded it.—N. W. Medical and Surg. Journal. We know of no work on Medical Jurisprudence which contains in the same space anything like the same amount of valuable matter.—N. Y. Journal of Medicine. No work upon the subject can be put into the hands of students either of law or medicine which will engage them more closely or profitably; and none could be offered to the busy practitioner of either calling, for the purpose of casual or hasty reference, that would be more likely to afford the aid desired. We therefore recommend it as the best and safest manual for daily use.—American Journal oj Medical Sciences. This work of Dr. Taylor's is generally acknow- ledged to be one of the ablest extant on the subject of medical jurisprudence. It is certainly one of the most attractive books that we have met with; sup- plying so much both to interest and instruct, that by the same author. (Nearly Ready.) ON POISONS, IN RELATION TO MEDICAL JURISPRUDENCE AND MEDICINE. Second American, from a second and revised London edition. In one large octavo volume. The length of time which has elapsed since the first appearance of this work, has wrought so great a change in the subject, as to reauire a very thorough revision to adapt the volume to the present wants of the profession. ThelraRid advance of Chemistry has introduced into we many new substances which may become fajal through accident, carelessness, or design—while at the same time it has likewise designated new and more exact modes of counteracting or detecting those previously treated of. Mr. Taylor's position as the leading medical jurist of England, has during this period conferred on him extraordinary advantages in acquiring experience in all that relates to this department, nearly all cases of moment being referred to him for examination, as an expert whose testimony is generally accepted as final. The results of his labors, therefore, as gathered together in this volume, carefully weighed and sifted, and presented in the clear and intelligible style for which he is noted, may be received as an acknowledged authoritj-, and as a guide to be followed with implicit confidence. WILSON (MARRIS), M. D. ON DISEASES OF THE VESICUL.E SEMINALES. page 19. WILLIAMS (C. J. B.i, M. D., F. R. S., Professor of Clinical Medicine in University College, London, &c. PRINCIPLES OF MEDICINE. An Elementaiy View of the Causes, Nature, Treatment, Diagnosis, and Prognosis of Disease; with brief remarks on Hygienics, or the pre- servation of health. A new American, from the third and revised London edition. In one octavo volume, leather, of about 500 pages. $2 50. (Just Issued.) The very recent and thorough revision which this work has enjoyed at the hands of the author has brought it so completely up to the present state ofthe subject that in reproducing it no additions have been found necessary. The success which the work has heretofore met shows that its im- portance has been appreciated, and in its present form it will be found eminently worthy a continu- ance of the same favor, possessing as it does the strongest claims to the attention of the medical student and practitioner, from the admirable manner in which the various inquiries in the different branches of pathology are investigated, combined and generalized by an experienced practical phy- sician, and directly applied to the investigation and treatment of disease. See "Lallemand," We find that the deeply-interesting matter and Btyle of this book have so far fascinated us, that we have unconsciously hung upon its pages, not too long, indeed, for our own profit, but longer than re- viewers can be permitted to indulge. AVe leave the further analysis to the student and practitioner. Our judgment ((f the work has already been sufficiently exDresscd. It is a judgment of almost unqualified nraise The work is not of a controversial, but of a didictic character; and as such we hail it, and recommend it for a text-book, guide, and constant companion to every practitioner and every student who wishes to extricate himself from the well-worn ruts of empiricism, and to base his practice of medi- cine upon principles.—London Lancet. A text-book to which no other in our language is comparable.—Charleston Medical Journal. No work has ever achieved or maintained a more deserved reputation.— Va. Med. and Surg. Journal. WHITEHEAD ON THE CAUSES AND TREAT- I MBNT OF ABORTION AND STERILITY. | Second American Edition. In one volume, octa- vo, extra cloth, pp. 308. $1 75. 30 BLANCHARD & LEA'S MEDICAL New and much enlarged edition—(Now Ready, Oct. 1858.) WATSON (THOMAS), M.D., &c. Late Physician to the Middlesex Hospital, &c. LECTURES ON THE PRINCIPLES AND PRACTICE OF PHYSIC. Delivered at King's College, London. A new American, from the last revised and enlarged English edition, with Additions, by D. Francis Condie, M. D., author of" A Practical Treatise on the Diseases of Children," &c. With one hundred and eighty.five illustrations on wood. In one very large and handsome volume, imperial octavo, of over 1200 closely printed pages in small type ; the whole strongly bound in leather, with raised bands. Price $4 25. The publishers feel that they are rendering a service to the American profession in presenting at so very moderate a price this vast body of sound practical information. Whether as a guide for the student entering on a course of instruction, or as a book of reference for daily consultation by the practitioner, " Watson's Practice" has long been regarded as second to none; the soundness and fulness of its teachings, the breadth and liberality of its views, and the easy and flowing style in which it is written having won for it the position of a general favorite. That this high reputa- tion might be fully maintained, the author has subjected it to a thorough revision; every portion has been examined with the aid of the most recent researches in pathology, and the results of modern investigations in both theoretical and practical subjects have been carefully weighed and embodied throughout its pages. The watchful scrutiny of the editor has likewise introduced whatever possesses immediate importance to the American physician in relation to diseases inci- dent to our climate which are little known in England, as well as those points in which experience here has led to different modes of practice ; and he has also added largely to the series of illustra- tions, believing that in this manner valuable assistance may be conveyed to the student in elucidat- ing the text. The work will, therefore, be found thoroughly on a level with the most advanced state of medical science on both sides of the Atlantic. The additions which the work has received are shown by the fact that notwithstanding an en- largement in the size of the page, more than two hundred additional pages have been necessary to accommodate the two large volumes of the London edition (which sells at ten dollars), within the compass of a single volume, and in its present form it contains the matter of at least three ordinary octavos. Believing it to be a work which should lie on the table of every physician, and be in the hands of every student, the publishers have put it at a price within the reach of all, making it one of the cheapest books as yet presented to the American profession, while at the same time the beauty of its mechanical execution renders it an exceedingly attractive volume. It would appear almost superfluous to adduce commendatory notices of a work which has so long been established in the position of a standard authority as "Watson's Practice." A few ex- tracts are, however, subjoined from reviews of the new and improved edition. The fourth edition now appears, so carefully re- vised, as to add considerably to the value of a book already acknowledged, wherever the English lan- guage is read, to be beyond all comparison the best The lecturer's skill, his wisdom,his learning,are equalled by the ease of his graceful diction, his elo- quence, and the far higher qualities of candor, of courtesy, «f modesty, and of generous appreciation s> stematic work on the Principles and Practice of | of merit in others. May he long remain to instruct Physic in the whole range of medical literature. Every lecture contains proof of the extreme anxiety ofthe author to keep pace with the advancing know- ledge of the day, and to bring the results of the labors, not only of physicians, but of chemists and histologists, before his readers, wherever they can be turned to useful account. And this is done with such a cordial appreciation of the merit due to the industrious observer, such a generous desire to en- courage younger and rising men, and such a candid acknowledgment of his own obligations to them, that one scarcely knows whether to admire most the pure, simple, forcible English—the vast amount of useful practical information condensed into the Lectures—or the manly, kind-hearted, unassuming character of the lecturer shining through his work. —London Med. Times and Gazette, Oct. 31, 1857. Thus these admirable volumes come before the profession in their fourth edition, abounding in those distinguished attributes of moderation, judgment, erudite cultivation, clearness, and eloquence, with which they were from the first invested, but yet richer than before in the results of more prolonged observation, and in the able appreciation of the latest advances in pathology and medicine by one of the most profound medical thinkers of the day.— London Lancet, Nov. 14, 1857. us, and to enjoy, in the glorious sunset of his de- clining years, the honors, the confidence and love gained during his useful life.—N. A. Med -Chir. Review, July, 1858. Watson's unrivalled, perhaps unapproachable work on Practice—the copious additions made to which (the fourth edition) have given it all the no- velty and much of the interest of a new book.— Charleston Med. Journal, July, 1858. Lecturers, practitioners, and students of medicine will equally hail the reappearance of the work of Dr. Watson in the form of anew—a fourth—edition. We merely do justice to our own feelings, and, we are sure, of the whole profession, if we thank him for having, in the trouble and tuTmoil of a large practice, made leisure to supply the hiatus caused by the exhaustion of the publisher's stock of the third edition, which has been severely felt for the last three years. For Dr. Watson has not merely caused the lectures to be reprinted, but scattered through the whole work we find additions or altera- tions which prove that the author has in every way sought to bring up his teaching to the level of che most recent acquisitions in science.—Brit, and For. Medico-Chir. Review, Jan. 1858. WHAT TO OBSERVE AT THE BEDSIDE AND AFTER DEATH, IN MEDICAL CASES. Published under the authority of the London Society for Medical Observation. A new American, from the second and revised London edition. In one very handsome volume, royal 12mo., extra cloth. $1 00. To the observer who prefers accuracy to blunders j One of the finest aids to a young practitioner w« and precision to carelessness, this little book is in- have ever seen.—Peninsular Journal of Medicine. valuable.—N. H. Journal of Medicine. I WILDE (W. R.), Surgeon to St. Mark's Ophthalmic and Aural Hospital, Dublin. AURAL SURGERY, AND THE NATURE AND TREATMENT OF DIS- EASES OF THE EAR. In one handsome octavo volume, extra cloth, of 476 pages, with illustrations. $2 80. AND SCIENTIFIC PUBLICATIONS. 31 New and much enlarged edition—(Now Ready, Oct. 1858.) WILSON (ERASMUS), F. R. S., A SYSTEM OF HUMAN ANATOMY, General and Special. A new and re- vised American, from the last and enlarged Engli>h Edition. Edited by W. H. Gobrecht, M. D., Professor of Anatomy in the Philadelphia Medical College, &c. Illustrated with three hundred and ninety-seven engravings on wood. In one large and exquisitely printed octavo volume, of over 600 large pages; leather. $3 25. The publishers trust that the well earned reputation so long enjoyed by this work willjbe more than maintained bythe present edition. Besides a very thorough revision by the author, it has been most carefully examined by the editor, and the efforts of both have been directed to introducing everything which increased experience in its use has suggested as desirable to render it a complete text-book for those seeking to obtain or to renew an acquaintance with Human Anatomy. The amount of additions which it has thus received may be estimated from the fact that the present edition contains over one-fourth more matter than the last, rendering a smaller type and an enlarged page requisite to keep the volume within a convenient size. The author has not only thus added largely to the work, but he has also made alterations throughout, wherever there appeared the opportunity of improving the arrangement or style, so as to present every fact in its most appro- priate manner, and to render the whole as clear and intelligible as possible. The editor has exercised the utmost caution to obtain entire accuracy in the text, and has largely increased the number of illustrations, of which there are about one hundred and fifty more in this edition than in the last, thus bringing distinctly before tbe eye ofthe student everything of interest or importance. The publishers have felt that neither care nor expense should be spared to render the external finish ofthe volume worthy of the universal favor with which it has been received bythe American profession, and they have endeavored consequently to produce in its mechanical execution an im- provement corresponding with that which the text has enjoyed. It will therefore be found one of the handsomest specimens of typography as yet produced in this country, and in all respects suited to the office table of the practitioner, notwithstanding the exceedingly low price at which it has been placed. A few notices of former editions are subjoined. This is probably the prettiest medical book ever j evident. Let students, by all means examine the published, and we believe that its intrinsic merits are in keeping with its exterior advantages, having examined it sufficiently to satisfy us that it may be recommended to the student as no less distinguished by its accuracy and clearness of description than by its typographical elegance. The wood-cuts are ex- quisite.—British and Foreign Medical Review. , An elegant edition of one of the most useful and accurate systems of anatomical science which has been issued from the press. The illustrations are really beautiful. In its style the work is extremely concise and intelligible. No one can possibly take up this volume without being struck with the great beauty of its mechanical execution, and the clear- ness of the descriptions which it contains is equally BY THE SAME AUTHOR. (Just Issued.) ON DISEASES OF THE SKIN. Fourth and enlarged American, from the last and improved London edition. In one large octavo volume, of 650 pages, extra cloth, $2 75. The writings of Wilson, upon diseases of the skin, claims of this work on their notice, before they pur- chase a text-book of the vitally important science which this volume so fully and easily unfolds.— Lancet. In every respect, this work, as an anatomical guide for the student who seeks to obtain know- ledge which he has not yet acquired, and for the practitioner who wishes to keep up that which he find-", gradually fading from his mind, merits our warmest and most decided praise.—Med. Gazette. We regard it as the best system now extant for students.—Western Lancet. It therefore receives our highest commendation.— Southern Med. and Surg. Journal. are by fir the most scientific and practical that have ever been presented to the medical world on this subject. The present edition is a great improve- ment on all its predecessors. To dwell upon all the greut merits and high claims of the work before us, seriatim, would indeed be an agreeable service; it would be a mental homage which we could freely offer, but we should thus occupy an undue amount of space in this Journal. We will, howiver. look at some of the more salient points with which it abounds, and which make itincompuraoiy superior in excellence to all other treatises on the subject of der- matology. No mere speculative views are allowed a place in this volume, which, without a doubt, will, for a very long period, be acknowledged as the chief standard work on dermatology. The principles of an enlightened and rational therapeia are introduced on every appropriate occasion. The general prac- titioner and surgeon who, peradventure, may have for years regarded cutaneous maladies as scarcely worthy their attention, because, forsooth, they are not fatal in their tendency; or who, if they have attempted their cure, have followed the blind guid- ance of empiricism, will almost assuredly be roused to a new and becoming interest in this department of practice, through the inspiring agency of this book.—Am. Jour. Med. Science, Oct. 1857. ALSO, NOW READY, A SERIES OF PLATES ILLUSTRATING WILSON ON DISEASES OF THE SKIN ; consisting of nineteen beautifully executed plates, of which twelve are exquisitely colored, presenting the Normal Anatomy and Pathology of the Skin, and containing accurate re- presentations of about one hundred varieties of disease, most of them the size of nature. Price in cloth $4 25. In beauty of drawing and accuracy and finish of coloring these plates will be found equal to anything of the kind as yet issued in this country. One of the best specimens of colored lithographic illustrations that have ever been published in this country. Tlie representations of diseases of the skin, even to the most minute shade of coloring, are remarkably accurate, giving the student or practi- tioner a very correct idea of the disease he is study- ing. We know of no work so well adapted to the wants of the general practitioner as Wilson's, with the accompanying plates. — Med. and Surg. Re- porter, -May, 1*56. We have already expressed our high appreciation of Mr. Wilson's treatise on Diseases of the Skin. The plates are comprised in a separate volume, which we counsel all those who possess the text to purchase. It is a beautiful specimen of color print- ing, and the repiesentations of the various forms of skin disease are as faithful as is possible in plates of the size.—Boston Med. and Surg. Journal, April 8, IboS. 32 BLANCHARD & LEAS MEDICAL PUBLICATIONS. WILSON (EftASMUSI, M. D., F. R. S. Lecturer on Anatomy, London. THE DISSECTOR'S MANUAL; or, Practical and Surgical Anatomy. Third American, from the last revised and enlarged English edition. Modified and rearranged, by William Hunt, M. D., Demonstrator of Anatomy in the University of Pennsylvania. In one large and handsome royal 12mo. volume, leather, of 582 pages, with 154 illustrations. $2 00. The modifications and additions which this work has received in passing recently through tbe author's hands, is sufficiently indicated by the fact that it is enlarged by more than one hundred pages, notwithstanding that it is printed in smaller type, and with a greatly enlarged page. BY THE SAME AUTHOR. ON CONSTITUTIONAL AND HEREDITARY SYPHILIS, AND ON SYPHILITIC ERUPTIONS. In one small octavo volume, extra cloth, beautifully printed, with four exquisite colored plates, presenting more than thirty varieties of syphilitic eruptions. $2 25. BY THE SAME AUTHOR. HEALTHY SKIN; A Popular Treatise on the Skin and Hair, their Preserva- tion and Management. Second American, from the fourth London edition. One neat volume, royal 12mo.; extra cloth, of about 300 pages, with numerous illustrations. $1 00; paper cover, 75 cents. (Now Complete.) WEST (CHARLES), M. D., Accoucheur to and Lecturer on Midwifery at St. Bartholomew's Hospital, Physician to the Hospital for Sick Children, &c. LECTURES ON THE DISEASES OF WOMEN. Now complete in one hand- some octavo volume, extra cloth, of about 500 pages ; price $2 50. Also, for sale separate, Part II, being pp. 309 to end, with Index, Title matter, &c, 8vo., cloth, price $1. EP Copies done up in paper covers, for mailing, will be sent, free of postage, to any address within the United States on receipt of One Dollar in current funds or postage stamps. Subscribers to the "Medical News and Library" who received the first portion of this work as published in 1856 and 1857, should lose no time in securing the completion. Part I will no longer be sold separate. As the first part of this work formed a complete treatise on the Diseases of the Uterus, so Part II is complete in itself as a text-book on the affections of the Uterine Appendages, the Ovaries, Vagina, Bladder, and External Organs. It will be found fully to maintain the high character ac- quired by the preceding portion immediately on its appearance, and the whole will constitute a reliable text-book on this interesting and difficult branch of practice. A few notices of Part I are added. painstaking, practical physician is apparent on every page.—N. Y. Journal of Medicine, March, 1858. We know of no treatise of the kind so complete and yet so compact.—Chicago Med. Journal, Janu- ary, 1858. A fairer, more honest, more earnest, and more re- liable investigator of the many diseases of women and children is not to be found in any country.— Southern Med. and Surg. Journal, January 1853. We gladly recommend his Lectures as in the high- est degree instructive to all who are interested in obstetric practice.—London Lancet. We have to say of it, briefly and decidedly, that it is the best work on the subject in any language; and that it stamps Dr. West as the facile princeps of British obstetric authors.—Edinb. Med. Journ. As a writer, Dr. West stands, in our opinion, se- cond only to Watson, the " Macaulay of Medicine;" he possesses that happy faculty of clothing instruc- tion in easy garments; combining pleasure with profit, he leads his pupils, in spite of the ancient proverb, along a royal road to learning. His work is one which will not satisfy the extreme on either side, but it is one that will please the great ma- jority who are seeking truth, and one that will con- vince the student that he has committed himself to a candid, safe, and valuable guide. We anticipate with pleasure the appearance of the second part of the work, which, if it equals this part, will com- plete one of our very best volumes upon diseases of females.—N. A. Med.-Chirurg. Review, July, 1858. We must now conclude this hastily written sketch with the confident assurance to our readers that the work will well repay perusal. The conscientious, BY THE SAME AUTHOR. (Just Issued.) LECTURES ON THE DISEASES OF INFANCY AND CHILDHOOD. Second American, from the Second and Enlarged London edition. In one volume, octavo, extra cloth, of nearly five hundred pages. $2 00. We take leave of Dr. West with great respect for ligation by this able, thorough, and finished work his attainments, a due appreciation of his acute upon a subject which almost daily taxes to the ut- powers of observation, and a deep sense of obliga- most the skill of the general practitioner. He haa tion for this valuable contribution to our profes- with singular felicity threaded his way through ail sional literature. His book is undoubtedly in many the tortuous labyrinths of the difficult subject he has respects the best we possess on diseases of children, undertaken to elucidate, and has in many of the Dublin Quarterly Journal of Medical Science. darkest corners left a light, which will never be Dr. West haB placed the profession under deep ob- extinguished.—IVosAirfH* Medical Journal. BY THE SAME AUTHOR. AN ENQUIRY INTO THE PATHOLOGICAL IMPORTANCE OF ULCER- ATION OF THE OS UTERI. In one neat octavo volume, extra cloth. $1 00. YOUATT (WILLIAM), V. S. THE DOG. Edited by E. J. Lewis, M. D. With numerous and beautiful illustrations In one very handsome volume, crown Svo., crimson cloth, gilt. $ 1 25. ) '-^tVU^/ .*£ m m ,,;•■ :»m«£.V. ':■'-' **&■■■■■■ ••.,..■