NATIONAL LIBRARY OF MEDICINE Washington Founded 1836 U. S. Department of Health, Education, and Welfare Public Health Scrrice 242 r p, K " +s II--' •"■' < h: RESEARCHES INTO THE COMPARATIVE STRUCTURE OF THE LIVER, BY JOSEPH LEIDY, M. D. WITH THREE PLATES. [Extracted from the American Journal of the Medical Sciences Tor January, IMV; The following observations* on the comparative structure of the liver were made solely for my own satisfaction, but I have been induced to make them public from the impression that some of the facts presented may be new, or that, at least, the anatomy of the liver being presented in a somewhat new aspect, may be better understood than by former methods of description. It will be remarked that as respects the anatomy of the liver of verte- brated animals, the facts observed by me corroborate the very carefully made researches and accurate descriptions of the anatomy of the human liver by Mr. Kiernan. {Philosophical Transactions, London, lsW.) The constant presence of the liver throughout the whole of the vertebrate scries of animal life and the higher orders of the invertebrate series, is sufficient proof of the importance of the organ, in connection with the digestive apparatus. It is an extensive secreting surface collected into a small bulk, and in the vertebrata it is arranged in such a complex manner, that before the researches of Mr. Kiernan, anatomists entirely failed in obtaining any satisfactory knowledge relative to its arrangement. Before entering upon the details of the anatomy of the liver, it may be well to examine in a very general manner the nature of secretion and the orcans which possess this function. It is rather difficult to give a good general deliniiion of the term secretion, for there are many organic pro- . These observations were made with an excellent microscope manufactured by Oberhaeuser. ,1 I»ar». and belonging .o Dr. Wm. Schmode.of lb., c.y. 10 whose liberality 1 am indebted for its use during a long period. 2 Leidy on the Comparative Structure of the Liver. cesses which closely resemble it; such are respiration,* the renewal of epi- thelial surfaces, growth of hair, &c.; but to restrict it in some degree, it may be considered to be that process in an organic being by which certain organic cells, forming part of its composition, separate or form from the nutritive fluid, in contact with them upon one side through the intervention of a basement membrane, certain peculiar substances, which they give up upon the other side, to be appropriated to some other purpose in the organism, or to be cast ofTas useless, or which would be hurtful if retained. Particular organs devoted to this function are denominated glands, and the simplest form of a glandular body is a single organic cell resting upon a basement membrane, and having the function just spoken of. Such are probably the cells which secrete the irritating fluid in the Medusas, certain cellules in the Po/ygnstric animalculae, $c. An increase of such cells upon a plane surface is a little advance on the former state, and appears almost as simple;—of this character are the glandular patches found in the ali- mentary canal of insects, &c. The surface upon which the secreting cells are placed, being depressed below the general level, adds a little more to the complexity of the structure, and in this way the simple mucous follicles are formed,—the follicles of the provenlriculus of birds, follicles of Lieber- kuhn,&c. Should these depressions, which if deep are tubular, be divided towards the extremity, it increases the amount of secreting surface, and renders it more complex,—as in the compound follicles of the stomach, the sebaceous follicles of the skin, &c. If the divisions be subdivided, glands like the salivary, lachrymal, &c, are formed. If, instead of subdi- vision, the primitive divisions are much elongated, and convoluted to econo- mize space, we have a gland of which the testicle of the higher vertebrata is an example. If the tube be divided and subdivided to a great extent, and each ultimate division forms a reticulated mass of tubular structure, the most complex form of a gland is produced, as in the fully developed liver of most vertebrated animals. The relation of the channels which convey the nutritive fluid to and from glandular organs, of course adds cons.derably to their complexity. From the foregoing we perceive the complexity of a gland depends upon the extent and mode of arrangement of its secreting surface and the relation of the blood-vessels to it. The quantity of secreted matter depends upon the number and activity of the organic cells forming a secreting surface, and the quality appears not to depend upon the complexity of the gland, but upon the power of the individual cells ; for the same secretion produced in glands of the most in- tricate arnngement in higher animals, is produced in very simple structures in the lower animals, as instanced in the liver of the higher vertebrata, and that ofJimphioxus lanceolutus, the testicle of man and that oUulus, &c. • In respiration there is not only a substance derived from .he blood and given off externally, but a.so one derived from the exterior and given up to «he blood lo thai there ,s an interchange of substances which is not .he case in secretion Leidy on the Comparatirr Structure of the Liver. 3 In the lowest forms of vertebrated animal life, where no distinct digestive apparatus exists, we find no trace of biliary structure. When we rise a little in the scale, and arrive at animals with an internal digestive cavity, as the Polypi, it is probable that certain of the cells forming part of the parietes of the cavity, may possess the power of secreting a fluid analogous to bile. < 'ertain cellules, also, of the Polygastrica may be devoied to this purpose. The few short cu-ca of the digestive cavity of many Medusx may also be biliary in their nature. In the Txnoidea and Tremafoda, animals which have a ramiform digesi- ive apparatus, the coecal appendages have been considered as a rudimentary form of the organ. They, however, do not differ in structure, which is granular, from the main portion of the cavity, and, like the latter, receive part of the alimentary matter. The ctecal appendages of the digestive cavity of ihe Echinodermata are reported as representing the biliary struc- ture, a question, as in several of the former instances, which I have not yet examined. In many Annelida there are casca appended to the sides of the alimentary canal, which are lined by organic cells of small size, and probably secrete a biliary fluid, as in Hirudo, Arenicoia, &c. In the Mi/riapoda, emptying into the intestine, there are several long and delicate tubes of basement membrane lined by secreting cellules, in the Julidx averaging 012.1 millimetre in diameter, which are no doubt of this nature ; but as they do not differ in structure from corresponding tubes of insects, a more particular description of the latter will answer for the former. fleets —In insects the liver consists of a number of dstinct, white, yel- low or brown, filiform, tortuous tubes placed in close apposition with the sides of the alimentary canal and opening into it, generally hy separate orifices, frequently afier joining each other to form short trunks, as in Musca, Tabunus, &c, in the vicinity of the pyloric extremtty of the stomach. The smallest number, which is four, occurs in the fl.es (Ihptera); in the Lepidoptera there are s,x ; and in the Orlhoptera and Hymenoptera they are numerous. When few in number, they are very lot!,, somet.mes three or four tunes the length of the intestinal canal; when numerous the> are proportionately short, and more delicate. In some ms-cis they termi- nate in blind extremities, as in many Diplera, in others they unite ,n pa.rs t their farthest extremtty.so as to form loops as in the b^»-<0 When more intimately examined, these tubes (Plate I. F.g>. -. J, I. 5 0) a^ found to consist of a delicate tube of clear, transparent, amorphous bLement membmne, the inner surface of which is covered wnh secreting r I Fr -The thinness of the tube, the cells often project so as to g,re Granulated appearance when viewed by the nake ey, « ■« «£« fly, (.,/,„„, carnaria.) ,l»l. 1. Fig. 10 and generaHy "™* ** f« " tremitv, the sides of the tubes are so irregular that they appear as ,f mere.y foMfd.po- sea portions or lobes, which are invest, u \ bn.w.ii.u purPle. and bro»n. U -n he• ^ ,w ,ppBmB8, 6 Leidy on the Comparative Structure of the Liver. colour. In intimate structure it consists of numerous, small, irregular bodies, or lobules (lobules of Kiernan), corresponding to the lighter por- tions just mentioned, which are lobulated themselves and closely con- nected together by means of white and yellow fibrous tissue and the blood-vessels belonging to the organ, which correspond to the dark ground lines separating the lighter-coloured masses. The lobules are not regu- larly arranged side by side throughout the liver, but lie in all directions, principally, however, with their long diameters at right angles to the surfaces. When the vessels of a liver have been injected, and the organ then hardened in alcohol so that it may be rendered more consistent and its difference of structure more perceptible, and a section made at right angles to the surfaces of the organ, a view like Fig. 31, PI. 111., will be obtained. In such a section lobules will be observed to be cut in all directions; longitudinally, when they have a foliated appearance; ob- liquely or transversely when they have a more or less polygonal form depending upon the amount of mutual pressure at any part of the liver, being greatest in ihe interior, least near the surface. In their interior, sections of blood-vessels are seen, which belong to the hepatic veins ; and the vessels occupying the interspaces between them are branches of the hepatic artery and vena portarum. The lobules are composed of an inter- texture of biliary tubes, (pori biliari,) (PI. 111. Fig. 32,) and in the areola? or interspaces of the network the blood-vessels ramify and form amongst them- selves an intricate anastomosis, the whole being intimately connected to- gether by a combination of the white fibrous and yellow elastic tissue. In structure the biliary tubes (PI. II. Figs. 25, 28; PI. Ill Figs. 33, 34), correspond with those of the invertebraia, consisting of cylinders of base- ment membrane containing numerous secreting cells, and the only difference exists in the arrangement; the free tubes of the lower animals in the ver- tebrata becoming anastomosed or forming an inlertexture. The tubuli vary in size in an unimportant degree in different animals, and also in the same animal, being generally from two \o two and a half times the diameter of the secreting cells. The tubes of one lobule are distinct from those of the neighbouring lobuli, or only communicate indirectly by means of the trunks or hepatic ducts originating from the tubes and lying in the interspaces of the lobuli. The secreting cells (PI. II. Figs. 25, 20, 27, 20, 30 ; PI. III. Figs. 35,30,37) are irregularly angular, or polygonal in form from mutual pressure, and line the interior surface of the tubes. They vary in size in a moderate degree in different animals and also in the same animal, appear- ing to depend upon certain conditions of the animal and liver. The colour is light yellowish, or brownish when in mass, the other and darker colours of the liver appearing to depend upon the blood in the organ. They con- lain a'finely granular matter, oil globules, a granular nucleus, and a trans- parent nucleolus. The finely granular matter is the portion from which the colour of the r,.itf j / r - $ V < /IT fih m m ■o&».£ P^S's /£ t^jafr-t > * aeir^ '. ol Leidy A.'' S. £el. rsuu/a/7-y m .'&-■ 1 €> oO, .■-'J 3i° ~ 0 o => 0° / a ^ ^ *v & * ft ' ' ' **' ' i -, ■ % ^ '■ " - .-IS) '*$ V'?'V' ^ ^f'i-V S9 "#" ' I ' s deposited within the cells, producing what is called "fat liver/' The same condmoa of the liver is produced in drunkards, probably from the sumu at.on to nutrilion and the conversion of the alcoholic constituents mto lat. It u also produced in some other ways not well understood, as was Panted to ™ Llv in a case of Prof. S. Jackson's, in which the patient had laboured for a long time under disease of the liver and dy.pcp.ia. and died of hemor- 8 Leidy on the Comparative Structure of the Liver. rhage from the bowels consequent on extensive ulceration of the mucous membrane. The nucleus is generally central, frequently lateral, globular, and pretty uniform in size in the same animal. It is granular in structure and never contains oil globules ; generally, it is but indistinctly seen, excepting in fishes and reptiles, and frequently not at all, from the granular contents of the cell obscuring it, but is readily brought into view by the influence of acetic acid upon the hitter. Sometimes there are two nuclei instead of one. The nucleolus measures about -001 millimetre, is round in form, con- sistent, and transparent, and is situated in the centre of the nucleus. The interlobular trunks or commencement of the hepatic ducts, as they originate from the biliary tubes or pores, run in varied directions in regard to the lobules, and freely anastomose with each other, and by their con- vergence form trunks which take a general course at right angles to the sur- faces of the liver, and finally appear by several trunks externally beneath the liver. The blood-vessels (PI. III. Figs. 38,39) of the liver consist of two sets, the hepatic artery and vena portarum, wrhich convey the blood to it, and a third set, the hepatic veins, which conducts the effete blood from it into the general circulation again. The hepatic artery, much smaller than the vena portarum, appears to be appropriated to the nutrition and supply of oxygen to the tissues enter- ing into the composition of the liver; while the vena portarum is probably devoted to the conveyance of blood to the secreting cells, which appro- priate the peculiar fluid of the liver, or bile from it. These two vessels enter the liver at the place of exit of the hepatic ducts, and follow the same course inwards that the latter did in coming out. The artery in its passage supplies the ducts with branches and the vena portarum with vasa vasorum. In the intervals of the lobules they comport themselves very much in the manner of the interlobular ducts, and form an intricate net- work around the lobules, but whether the two sets of vessels anastomose I could not satisfactorily determine. They both send off numerous branches which enter the lobules at right angles to the length of the latter, and form an intricate plexus by turning through the interspaces of the biliary tubes. The vessels within the lobules freely communicate with each other and converge towards their interior, where they terminate in trunks, which run in ihe length of the lobules, and are the commencement of the hepatic veins. This free intercommunication of the three sets of vessels within the lobules has been fairly proved to me by a minute injection of a young liver, by Prof. Wm. E. Horner, of the University. All three of the ves- sels have been injected with red size, and it has penetrated beautifully, and a fine section from almost any part of it, beneath the microscope, has the appearance represented in Fig. 39. The commencing branches of the hepatic veins issue from the base of rut,j ■<-% .■■■i^'V-f:.- *-?'■•■;;..■.;■..; ■•<*■■•/■ ■»ViV-- -St*'- i- w &>*■ ,a.0co'. * '«J? .*$-. ^fVVVV>f N • '■•o.'Vfc '-■rV'-\>K n-;-rv^ ■««■.; » 0 o <° *ag< bet« " n Is nre is tilled with oil globule*. fPlaluvhullt.m eontacum), exhibiting 5, 6, Portion, of a biliary tube from the katydid (Platyphyllt ',he elongated cells and nuclei. magnified. The ' exhibiting the structure of c«eal tubes. 10 Leidy on the Comparative Structure of the Liver. Fig. 9, A ccecum or biliary tube of the cray fish (from an individual which was one inch in length and had been well fed for some time), highly magnified. At the bottom is seen a confused mass of cells with their nuclei and nucleoli, but at the upper part of the sac they are seen distinct, and polygonal from pressure. and filled with oil which renders them transparent. At the mouth or open extremity numerous detached cells and oil globules are perceived, which have been squeezed out by the pressure of a thin plate of glass. Figs. 10, 11, 12, 13, a, b, Exhibit the progressive change of the cells as they advance from the bottom of the tube. Fig. 14, Nuclei, from the secreting cells of the liver of the cray fish, highly magnified. " 15, Portion of the liver of the snail (Helix albolabris), moderately magnified, exhi- biting the arrangement of the lobules. " 16, A biliary ccecum from the liver of the snail, highly magnified. It shows the same structure as fig. 9. " 17, a, b, Two cells, from the bottom of a biliary ccecum of the snail, highly magnified. " 18, a, b, Two cells, more advanced, containing numerous very minute oil globules. " 19, a, b, c, Three cells, containing larger oil globules. " 20, A cell distended with oil. " 21, A cell containing nothing but six deep yellow consistent oil globules. " 22, A cell containing a hard yellow mass of fat. " 23, A cell ruptured and its contents escaping. " 24, Nuclei, from the cells of the liver of the snail, highly magnified. " 25, Portion of a biliary tube and secreting cells, from the liver of a lizard (Triton niger), highly magnified. " 26, Secreting cells, from the liver of an owl (Strix brachyotos), highly magnified. "27, " " from a duck (Anas acuta). " 28, Portion of a biliary tube of the rabbit (Lepus Americana*), highly magnified. " 29, 30, Secreting cells from the liver of the rabbit. " 31, Longitudinal section of human liver from the posterior part near the upper sur- face, magnified 3 diameters, from a preparation made by Prof. W. E. Horner of the University of Pennsylvania. The three sets of blood-vessels were in- jected with colouring matter and the preparation then preserved in alcohol. The blood-vessels represented in the drawing belong to the hepatic veins, and are seen at various parts coming from the interior of the lobules. The spaces between the lobules, which are filled with branches of the hepatic artery and vena portarum and hepatic ducts, have been purposely left white so as not to obscure the view of the lobules. u 32, Transverse section of a lobule of the human liver, taken from the same prepa- ration as fig. 31, highly magnified, and presenting to view the retietulated struc- ture of the biliary tubes. In the centre of the figure is seen the hepatic vein cut across and several small branches terminating in it. Where the injecting matter did not run freely, it is seen standing in dots along the course of the vessels. At the periphery are seen branches of the hepatic artery, vena porta- rum and hepatic duct. " 33, A small portion of fig. 33 more highly magnified. The secreting cells are seen within the tubes, and in the interspaces of the latter the fibrous tissue is repre- sented. " 34, Portion of a biliary tube, from a fresh human liver, very highly magnified. The secreting cells may be noticed to be polygonal from mutual pressure. B 35, Three secreting cells detached. 30, A secreting cell much more highly magnified, representing plainly the internal Leidy on the Comparative Structure of the Liver. 11 structure. The granular nucleus is central, and the imall Uxlics with thick block outlines represent oil globules. Fig. 37, Four seireting cells, from a human liver, in a diseased condition calW "fctty degeneration" or ''fat liver." The ^reut increase of oil globules is ob*crv»ble. " 38, Transverse section of a lobule, from the tnmc preparation b« 31 and 32, but not so highly magnified as 32, representing the relation of th,' three k< ts of blood- vessels with each other and with the biliary tubes. The anery is coloured red, the vein blue, and the vena portarum yellow. " 39 U< presents a longitudinal section, highly magnified, of a portion of n lobule, from an injected preparation, made by Dr. Horner, of the liver of a chiM about 6 years of age. It will bo perceived there is a free anastomosis of th*> three gets of vessels, which have all been successfully injected with re-d colour,i,g matter. Spaces arc also observablo through which the injection failed to p*»«. Observations on Ike existence of the Intermaxillary Bone in //<■ Embryo of the human subject. By Joseph Lbidt, M. D. The immortal Goethe, I believe, was the first to point out the existence of th, 08 iiiteruiaxillare in the human subject, but it has only been observed in an abnormal condition, or where there has been an arrest of development in con- nection with some cases of hare-lip; and the period of life in which it is found as a distinct piece, and its exact limits, have not yet been accurately determined. The' universality of the presence of the os intermaxillare in all animals l„ 1 ,\v man, its presence as a distinct piece in an abnormal condition in man, always defined by a lateral fissure which characterizes it as the incisive bone, and the uniform existence of a transverse fissure behind the incisive alveoli of the os maxillare supcrius of the human ftetus at birth, have led many anatomists to suspect its normal and independent existence in the embryotic condition of man at an earlier period than it has been sought for. As the negro in his aaatomical characters is not so far removed from the embryological condition as theW«'*e,it is to be presumed that the intermaxillary bone would remain longer distinct; and under such an impression 1 have several times desired medical students, from our Southern States, whose opportunities ol investigating the anatomy of the negro are frequent, to make this a subject oft inquiry. Such an opinion cannot be considered unworthy of attention, when it is recollected that Tschudi mentions the existence of a true os interi»rictah', as a constant condition, in certain branches of the aboriginal inhabitants of Peru, the Chinchas, Aymaras and Huancas. Recently having had an opportunity of examining several human enltyos in one of them I was fortunate enough to detect the intermaxillary bone aa a dis- tinct and independent piece. This embryo measured one inch and eleven lines from heel to vertex, and I presumed it to be about nine or ten weeks old. In it ossification had already advanced in the superior maxillary and inter- maxillary bones sufficiently to give them a determinate form, and their appear- ance, when magnified, is represented in the figures land 2, which were takeo from the specimens through the aid of the camera lucida. Fig. 1, Fig. 2. b— Fig. 1 represents the superior maxillary and intermaxillary bones, much mag- nified,' of a human embryo. The diawing was taken from the right side through the aid of the camera lucida, which reverses its position, a. superior maxillary bone; A. intermaxillary bone; c. line of articulation between the two bonesj d. palatine process; e. alveolar groove. Fig. 2 represents the antero-inferior surface of the separated intermaxillary bone, much magnified. (From the left side, but reversed by the camera.) a. ascend- ing or nasal process; 6. articulating surface tor the superior maxillary bone j c. incisor alveoli. The greatest breadth of the two bones in apposition is one line and two-thirds. the greatest height, being at the ascending or nasal process, is one line. The two pieces present a facial portion, consisting of the ascending or nasal process and part of the body of the bones; an alveolar ridge and groove and a palatine process projecting backward from the superior maxillary bone. They are easily separable at this period, and the articulation passes through the alveolar ridge at a point corresponding to the separation between the incisor alveoli and the canine alveolus, and extends transversely inwards behind the incisor alveoli, and verti- cally upwards, dividing the nasal process into two nearly equal portions. On the posterior surface of the nasal process the articulation is at the bottom of a comparatively deep and wide groove, which, however, does not appear to be part of the lachrymal canal, as the latter appears afterwards and external to the former groove. The preparations exhibiting these interesting points which prove the existence of the same law, throughout the animal kingdom, governing the forma- tion of the upper maxillary bones, I present for the inspection of the members of the Academy. In an embryonic skeleton in the Wistar Museum, measuring three and one- eighth inches in length, and purporting to be about nine weeks old, which, however, I think too young, the maxillo-intermaxillary articulation is still evident at the ascending process, but it does not divide the latter so equally, being more internal and inferior, apparently from a more rapid development of the nasal process of the true maxillary bone. Just above the alveolar ridge they are already anchylosed together. In another embryo, in the same museum, measuring three and one-fourth inches in length, the two bones have become firmly united, excepting behind the incisor alveoli-, but the line of original separation is readily traced out, from a greater degree of thinness and transparency along its course." The nasal process of the true maxillary bone has so much increased beyond the nasal process of the intermaxillary bone, that the latter no more ascends to the summit of the former, but is considerably inferior and internal. In the fetal skeleton, measuring five inches in length, all traces of the inter- articalation h*ve disappeared, except behind the incisor alveoli, which latter portion, as is well known, does not usually disappear until some time after birth, and in some instances is found in the adult cranium. NLM041390406