inaiiuinal LIBKAKY UF MEDICINE NATIONAL LIBRARY OF MEDK 'V 5r *fr\ C^ 3Nioia3w jo Asvaan ivnoiivn 3NiDia3w jo Aavaan ivnoiivn 3nidio3w jo Aavaan wno / 811 IVNOIIVN * ^m X x DF MEDICINE NATIONAL LIBRARY OF MEDICINE NATIONAL LIBRARY OF MEDICINE NATIONAL LIBRARY OF MEDIC an ivnoiivn 3NiDia3w jo Aavaan ivnoiivn 3noi"-"- 3NiDia3w jo Aavaan ivnoiivn snidiosw jo Aavaan ivnoiivn snidiqsw jo Aavaan ivnoij NATIONAL LIBRARY OF MEDICINE NATIONAL LIBRARY OF MEDICINE NATIONAL LIBRARY OF MEDIC K/ S 3Nma3w jo Aavaan ivnoiivn snidiqsw jo Aavaa-* '"-~" ^NiDiasw jo Aavaan ivnoi. h E^hbSSDO WIN NATIONAL LIBRARY ( . _____ "> /mt^S!- / m\ I < X ! /^ NLM005594634 \ J= ICINE NATIONAL I \ <«1 IOI1VN 3NIDIQ3W DUE DEC I m 3NiDia3w jo Aavaan wnoiivn snidiqsw jo Aavaan ivnoiivn LAST DATE ATIONAL library of medicine jiDiasw jo Aavaan wnoiivn ATIONAL LIBRARY OF MEDICINE NOI1VN 3NOIQ3W A/ viS^I ^S V% J JK a^ ^idiqsw jo Aavaan wnoiivn DICINE NATIONAL LIBRARY OF MEDICINE NATIONAL LIBRARY OF MEDICINE NOIiVN 3NI3I03W jo Aavaan wnoiivn CXix%i ^ / NATIONAL LIBRARY OF MEDICINE NATIONAL LIBRARY OF MEDICINE 5 "V s | Xv-5^ r r~^-' SNioiasw jo Aavaan wnoiivn NATIONAL LIBRARY OF MEDICINE "" NATIONAL LIBRARY OF MEDICINE rSNioicnw jo Aavaan wnoiivn SNiDiasw jo Aavaan wnoiivn • PSYCHIATRY A CLINICAL TREATISE ON Diseases of the Fore-Brain BASED UPON A STUDY OF ITS STRUCTURE, FUNCTIONS, AND NUTRITION THEODOR meynert, M.D. PROFESSOR OF NERVOUS DISEASES AND CHIEF OF THE PSYCHIATRIC CLINIC IN VIENNA TRANSLATED (UNDER AUTHORITY OF THE AUTHOR) BY B. SACHS. M.D. INSTRUCTOR IN DISEASES OF THE MIND AND NERVOUS SYSTEM IN THE NEW YORK POLYCLINIC PART I. The Anatomy, Physiology, and Chemistry of the Brain NEW YORK AND LONDON G. P. PUTNAM'S SONS (T|jc Jlmciurbocher |lrrss 1885 fc IAgijs r COPYRIGHT BY G. P. PUTNAM'S SONS 1885 Press of G. P. Putnam's Sons New York TRANSLATOR'S PREFACE. I HAVE no apologies to offer for presenting this American edition of Professor Meynert's " Psychiatry" to the English medical public. It is a scientific treatise on diseases of the mind by the one best fitted to write such a treatise. To most medical men Meynert is known as the great brain-anatomist. This book may serve incidentally as a text-book on the anatomy of the brain ; but it is not merely that. I would direct particular atten- tion to the subsequent chapters of this treatise, from which the students of psychiatry, of physiology, and of psychology may gather much information and much food for reflection. For the shortcomings of my translation, I crave the indul- gence of the reader. I am quite certain that those best acquainted with the original will not underrate the difficulties of the task, and will be most lenient in passing judgment upon my errors. That there are such, I do not doubt. It has been my aim to furnish a readable translation of this treatise, and to accomplish this, the attempt at a literal translation had to be abandoned. I trust, however, that I have in no instance departed from the sense of the original. In regard to the encephalic nomenclature employed in the first section of this volume, I would say that I have coined but very few new terms; and that I have either used such terms as are familiar to all English students of cerebral anatomy, or have retained the Latin terms used by the author. The latter was done with the view of avoiding a conflict between the text and the author's plates. In order to make this volume complete in itself, I have pre- pared a special index to the subject-matter therein discussed. A similar index will be added to Part II. My special thanks are gladly given to my friend, Dr. M. Allen Starr, of this city, for his kind assistance in revising proof; and iii IV Psychiatry. to the publishers, Messrs. G. P. Putnam's Sons, for their generous efforts to make this translation worthy of the original. If the work now given to the public will advance the cause of psychiatry in this country and in England, I shall feel that I have in some small measure acknowledged the debt of gratitude I owe to my former master, the author. B. Sachs, M.D. 28 East Sixty-second Street, New York, October 12, 1885. AUTHOR'S PREFACE. The reader will find no other definition of " Psychiatry " in this book but the one given on the title-page: " Clinical Treatise on Diseases of the Fore-Brain." The historical term psychiatry, i. e. " treatment of the soul," implies more than we can accom- plish, and transcends the bounds of accurate scientific investiga- tion. Were I to give a functional designation to the morbid affections of the fore-brain, I would choose the term " Diseases of the Mind." And on this term I would insist in order to avoid the common fallacy that it is permissible to regard the contents of cortical memories as paled sensory images, although we acknowledge the origin of these memories from external sensory stimuli. We shall show in this book that the fore-brain can neither give rise to hallucinatory phenomena, nor that its func- tional manifestations, the so-called " memories," are possessed of the slightest sensory qualities. It would be better, therefore, to speak of memory-symbols. In our memory of the most glaring sunlight, of the most intense roll of thunder, there is not as much as the billionth part of the light of a glow-worm, or of the sound produced by the falling of a hair upon water. But is there any other word in our language with which to designate phenomena devoid of all sensory qualities, but the word " psychical "? This most conspicuous fact regarding the .functional activity of the fore-brain accentuates the difference between the abstract and material character of our concepts. The latter is lacking altogether, and is purely a matter of external perception. But the centres for such perception are in the basal portions of the brain, and not in the/^r^-brain. The entire fore-brain I designate generally as cortex, for the conducting elements of the fore-brain __the axial fibres—are prolongations of the cortical cells, and, therefore, part of the cortex proper. Those divisions of the brain which effect sensory perceptions, without the aid of the fore- brain, and which are concerned particularly with the execution of VI Psychiatry. reflex movements (in the broadest sense of that term), are desig- nated, by way of antithesis, as subcortical centres. This antithesis will be evident also in the different degrees of excitation (of these different centres), and will furnish us with the most important clue to the understanding of the activity of the cerebral mechan- ism under morbid conditions of the mind. I have not, and never had, the slightest inclination for making books. To this effort I was impelled by the conviction that there was need of a scientific treatise on mental diseases in spite of the present large literature on the subject. The least doubt as to the correctness of any views expounded in this book induced me to stop, and to interrupt work until I had satisfied myself of the correctness of those views by scientific investigation and reflec- tion. This will explain why so many years have elapsed from the time the book was begun (in 1877) to the date of its publication. The title-page refers to the fundamental studies indispensable to an understanding of the clinical manifestations of mental diseases. My intention of elucidating diseases of the fore-brain in this way is based upon the conviction that our knowledge of them should be obtained, as all sound clinical knowledge is acquired, by a study of the structure, the function, and the nutrition of each organ. Hitherto the science of psychiatry has been too largely subjective. Many of its teachings, which are not based upon the studies just referred to, had better be forgotten. Naturally enough a knowledge of clinical phenomena precedes in time the knowledge of the fundamental facts underlying these phenomena. Morbid symptoms are not recognized by their scientific sub- stratum, but in studying this substratum we are actuated by the desire to fathom the phenomena of disease. It follows, therefore, that the first half' of the present work is suggested by the second, clinical, half; and the subjects discussed in these fundamental chapters are chosen with a view to the thorough understanding of the clinical facts. These chapters constitute an integral part of this clinical treatise, and are not an adjunct to it. This, we take it, is a novel and legitimate feature of the book. The number of clinical cases upon which my views rest is not only sufficient to base original conclusions upon, but by reason of their large numbers, these cases seem to me to be absolutely con- vincing. In 1875 I was fortunate enough to be able to change my clinic at the Vienna Provincial Insane Asylum, where there was but 1 Part I. of the translation. Author s Preface. vn a slight change in the number and character of the inmates, for the Psychiatric Clinic of the Vienna General Hospital. My present "clinic" is the only State Insane Asylum of Austria, though its quarters are not in keeping with its importance. Fourteen to sixteen hundred patients are received annually into this clinic, and of these, only those who are inhabitants of Lower Austria are soon dismissed and assigned to other institutions. It was the study of such a vast number of cases which convinced me that many cases could not be properly classified according to the old rules and within the artificial types of mental diseases. The many variations from these types appeared not only to favor, but actually to compel, the establishment of a natural system of classification. During the interval that elapsed between the inception and completion of my book, I have seen reason to depart in some respects from the account here given of the anatomy of the brain. I have not adopted any new method, but have elaborated with greater care the old cleavage (Abfaserung) method of my prede- cessors,—a laborious method, which has been unduly crowded out by the study of brain-sections. The former method supplies us not only with a key to the complexities of brain-sections, but enables us also to extend our knowledge of the minute anatomy of the brain beyond the information we can obtain from micro- scopical sections. From the notes to be appended to the end of this work,1 the reader will gather wherein my views have been necessarily modified or supplemented by the investigations of other authors and my latest researches. The most important results which I have recently arrived at are in regard to the cortical and ganglionic fibres of origin of th.e pes pedunculi (crusta), and the connection of these fibres with the pyramids. My dissent from the adoption of a common system of cranial measurements, to which I agreed, at Ranke's suggestion, is explained by the fact that the chapter 2 on pathological craniology was written as early as 1880. In view of the necessity of starting from anatomical facts, I have endeavored, in every case, not only to give due weight to the structure of the brain as the fundamental basis for the various forms of disease, but have endeavored, with the same end in view, to insist upon and to explain every visible symptom ex- hibited by the patient. This refers to the consideration of move- 1 In Part II. of this edition. viii Psychiatry. ments of expression as well, which have not been properly utilized hitherto as aids to diagnosis. Thus in spite of the frequent smiles to be observed in stuporous patients during maniacal moods, stuporous insanity has been classified under melancholia. Dissatisfied with the statistical method, which laid inordinate stress upon hereditary predisposition to disease, I have considered predisposition as a form of disease and not as a condition antece- dent to it. I have above all referred to the anatomical peculiari- ties constituting predisposition. I was not content, as others have been, to accept the mystical conception of heredity, but have in- sisted on the anatomical peculiarities in patients which constitute predisposition. And these peculiarities were inferred not only from external appearances, but also from a due consideration of all abnormal proportions of the body. In an article, published as early as 1878, I showed that the nutrition and the excitability of the brain must be regarded as depending upon the reciprocal re- lation existing between the weights of the brain and of the heart. The important investigations of Benecke and Thoma on the size and weight of the different organs of the body appear to me to supply a firm anatomical foundation for the doctrine of predispo- sition to disease. As regards the theory of predisposition, and more particu- larly the doctrine of heredity, which has been carried to the ex- treme of assuming the existence of innate ideas, and which, in clinical medicine, has led to the erroneous theory of moral insan- ity, I have deemed it necessary to criticise, in its proper connec- tions, Darwin's theory of the inheritance of acquired faculties, as has been done before me by other German authors, among them DuBois Reymond and Weissman. It is taking altogether too simple a view of things, to regard morality as one of man's talents, and as a definite psychical property which is present in some persons and lacking in others. Indeed, there is great truth in Weissman's observation : " Talents do not depend upon the pos- session of any special portion of the brain ; there is nothing simple about them, but they are combinations of many and widely different psychical faculties." Just as the much-abused doctrine of hereditary predisposition to disease casts too great a suspicion upon the- limits of mental health, so there is possibly the danger that this same suspicion might be increased by the consideration of those errors of organi- zation which contitute the basis of predisposition. But thinking Author s Preface. IX physicians will avoid this danger, for they will distinguish be- tween those who are possibly "called" to disease, and that for- tunately smaller number of persons who are, in the saddest sense of the term, " chosen " for disease. Thus the limits of relative health, as they are generally conceived, will not be narrowed down by abstract theorizing to an intangible line. I trust that the dialectic efforts in this direction will bear fruit in the domain of forensic medicine, and it would be gratifying to me to know that I had contributed somewhat to this end. Vienna, Easter, 1884. Theodor Meynert. CONTENTS. PAGE Structure and Architecture of the Brain . . . i The Minute Anatomy of the Brain ..... 56 Anatomical Corollaries and Physiology of Cerebral Architecture ........ 138 The Nutrition of the Brain ...... 213 Appendix—Machanism of Expression . . . . .271 Index............279 STRUCTURE AND ARCHITECTURE OF THE BRAIN. SURFACES OF THE BRAIN. UNTIL recently, it was commonly believed that no vestiges of the vertebrate brain were to be found in ampJiioxns lancco- latus. Its spinal cord, instead of dilating above into a rudi- mentary encephalon, was thought to terminate in conical fashion. Rohony, however, has demonstrated the existence of a primary cerebral vesicle in the amphioxus as well. The possession of a brain and spinal cord may, therefore, be said to be charac- teristic of all vertebrates. The prosencephalon (the fore-brain), the largest division of the human brain, diminishes so rapidly in volume as we descend in the animal series that even among the simpler mammalian forms it does not in every case exceed in mass the other parts of the brain. In the evolution of the brain from the medullary tubes, the prosenceph- alon is represented by secondary subordinate ap- pendices of the primary cerebral vesicle. These vesicles of the hemispheres (so-called) are sym- metrically disposed, and lie one at each side of the median axis of the primitive brain structure. The arrow in Fig. I points to the aperture leading from the anterior cerebral vesicle to the laterally situated vesicle of the hemisphere—the fore-brain. The names given by Carl V. Baer to the various parts of the brain are most ap- propriate. We distinguish (Fig. i) the transition from the spinal cord into the after-brain' (me- dulla oblongata) ; next the hind-brain (H. cere- bellum), which the roof of the fourth ventricle joins to the posterior wall of the ventricle of the after-brain. The mid-brain (Jl. corpora quad- V Ion Prosencepha- (fore-brain ; Vorderhirn). Z. Thai am encephalon ('Tween-brain; Zwis- chenhirn). Y. Fora- men between median and r. lateral ventri- cle. M. Mesenceph- alon (M id-b rain; Mitlelhim). H. Ep- encephalon. (Hind- brain ; Hinterhirn.) N. Metencephalon. (After-brain; Nachh- irn.) (Reichert.) 1 The terms After-, Hind-, Mid-, Inter-, and Fore-brain are respectively synony- mous with Met-, Ep-, Mes-, Thalam-, and Pros-encephalon.—S. 2 Psychiatry. Fig. Frontal As- of the inter-brain) rigcmina) forms the summit of this structure. A part of the pro- sencephalic vesicle lies between the mid-brain and the vesicle of the hemisphere. This part is termed the inter-brain, and corre- sponds to the region of the optic thalamus (Z.). The vesicles of the hemispheres develop into the fore-brain (V.). Fig. 2 is intended to show what an insignificant part of the original brain mass the prosencephalon is. (We descend from the mid- and inter-brain -to the fore-brain as by a series of steps.) In consequence of a sharp bend of the cerebral axis on an ideal trans- verse diameter, the after-brain appears below as a pect of a Fcetal posterior branch of this curved arc. A horizontal RXeic\e\t.a^ter section of the brain (Fig. 3.)' shows the originally V. Prosenceph- lens-shaped convex outer surface of the fore-brain bamencepha- (Fig- 3. v0> at a later stage of development. This Ion. M. Mesen- originally convex surface of the fore-brain (which at cephalon. N. ° : Metencephalon. this stage overlaps about halt shows, midway between its anterior and pos- terior end, a trough-shaped depression. This depression is the first indication of the fossa Sylvii (S.). It is to be noticed that the walls of the vesicles of the hemispheres have increased in thickness toward the median line at the ex- pense of the lateral ventricles {a. p.); as a result of this growth we find the ganglionic region, which, later on, divides into the caudate and lenticular nuclei protruding between the anterior and posterior horns. Corresponding to this thickening of sub- stance toward the median line, the outer surface remains relatively undeveloped. On the other hand, the anterior and posterior portions of the rudimentary hemispheres do not form any ganglionic masses on their inner surface, but develop mainly into cor- tical and medullary substance, deposited upon their outer surfaces. The intervening portion (S.) sinks in, and thus forms the Sylvian fossa. The surface which presents in this fossa is the island of Reil. Horizontal Section of Fcetal Brain ; After His. VV. Fore-brain. a. Anterior horn. p. Pos- terior horn of the lateral ventricle. S. Fossa Sylvii, the outer surface of the ganglionic region of the fore-brain. Z. Inter-brain. m. 3d Ventricle. Be- hind this, a part of the mid-brain, surrounding the Aq. Sylvii.' The posterior boundary of the mid - brain is ex- plained by reference to the pari eta flexure (Fig. 4.) 1 This figure must be inverted when compared with Fig. 5, p. 4. Convex Surface of the Brain. 3 At this level the student may note how in the fore-brain the anterior (a.) and pos- terior (p.) horns of the lateral ventricles open widely into the hollow of the primary cerebral vesicle. At this point the median wall of the prosencephalon encircles this opening. The outer wall of the prosencephalon has given rise mid-way to the fossa Sylvii. The inner wall closes in upon the cerebral ventricle. The fornix ascending from the cornu ammonis between S. and Z., regions corresponding to the corpus striatum and the thalamus opticus, constitutes the posterior portion of the median wall. The superior convex arch of the fornix is flattened down, but the descend- ing portion of the fornix is enclosed within the front wall of the anterior cerebral vesicle. The fornix bounds the hollow separating the fore- and inter-brain, which cavity is rendered cleft-shaped by the inward growth of the thalamus, and forever after remains broadest at its anterior end (foramen Monroi). The upper wall of the thalamencephalon consists simply of the membranous roof of the third ventricle, which passes to the edge of an arch-shaped constriction arising from the upper and anterior wall of the thalamencephalon. This constriction repre- sents the fimbria of the fornix. The middle choroid plexus of the upper wall is continued through the foramen Monroi into the plexus of the lateral ventricles. The fornix in reality limits the extent of the fore-brain. As soon as the septum and the corpus callosum are developed, the gyrus fornicatus ap- View of the Convex pears to be a limiting formation, or at least a secondary free Surface of a fcetal margin of the cerebral cortex. An examination of Fig. 3 shows Brain. that the outer cerebral wall growing from the fossa Sylvii tow- p S 1 7* aid the median line gives rise to the ganglia of the prosen- Thalamenceph. M. cephalon and encroaches upon the annularopening in the median Mesenceph. H. wall and thus fills in the once copious hollow of the ventricle. Epenceph. N. Met- ' . , ,. , • ,• enceph. Beneath the The prosencephalon continues to extend in a postenordirec- ixo^[^ portion of the tion at later stages of development ; we find, therefore, on hori- prosencephalon lies zontal sections, that the corpus striatum and thalamus opticus the olfactory lobe. are juxtaposed from the outside inwardly instead of lying one (Lettering as in , Fig. I.J behind the other. The succession of the three cerebral vesicles is marked by several flexions on the axis of the original medullary tube (Fig. 4). The cervical flexure (convex posteriorly) marks the transition from the spinal cord to the metencephalon ; the frontal flexure (convex anteriorly), the transition from the metencephalon to the epenceph- alon ; and beneath the mesencephalon the parietal flexure is formed. The chorda dorsalis terminates in the sinus formed by the last-named flexure. A.—THE PROSENCEPHALON (FORE-BRAIN). convex surface. The prosencephalon describes a curve about the Sylvian fossa which was formed by the insufficient increase in thickness of the 4 Psychiatry. outer surface. The upper portion of this curve is the frontal end of the prosencephalon, the lower portion is its temporal endf while its summit represents the occipital region. On account of the insertion of the parietal region between the frontal and occipital portions of the brain, the upper arm of the arc is the longer one. The island of Reil Fig. 4, S., is connected with a protuberance (the olfactory lobe) situ- ated on the lower aspect of the frontal portion of the fore-brain. Behind the ol- factory lobe, the thalamen- cephalon is visible in the shape of a basal protuber- ance which forms the region of the infundibulum on the anterior border of the parie- tal flexure. Let us now proceed to examine with the aid of sections of the adult brain, the island of Reil—the floor of the Sylvian fossa. A horizontal longitudinal sec- tion (Fig. 5), as well as a ver- tical frontal cross-section (Fig. 6) of the brain will serve our purpose. A nar- row passage, the Sylvian fissure (FS.), screens from view the sac-like dilatation of the fossa Sylvii in the furthest recess of which lies the island of Reil (J.). The outer walls of the Sylvian Horizontal Longitudinal Section from the Brain of Cercocebus Griseoviridis. F. Frontal end. O. Occipital end, of fore-brain. A. Cornu ammonis. R. Cortex. M. Medullary r^„„„ , u:„i, substance. L Island of Reil. FS. Fissura Svl- f°SSa> whlch are wanting ill Fi< 4, are formed by the thickening of on the vii. V. Anterior horn. Vp. Posterior horn of the lateral ventricle (prosencephalic cavity). T. "~\. " , Corp. callosum. S. Septum. Nc. Na. Nucleus Continued caudatus and L.1, L.a Nucleus lenticularis (the cerebral Substance ganglia of the fore-brain). Cp.e. Capsula externa. i.e., The medullary substance immediatelyadjoin- Convex surface of the pros ing the lenticular nucleus. To the outside of the capsule lies the claustrum, and next to this the thin medullary substance of the island.- Ca. encephalic arch, until the edges of this arch approach Convex Surface of the Brain. 5 Commissura anterior. Th. Thalamus opticus (thalamencephalon). Th'. Pulvinar of the opt. thalamus. Cm. Commissura media. Qu. Mesen- cephalon—(Corp. quadrigemina). Bs. Brach- ium superius (of the upper corp. bigemina). Aq. Aquceduct Sylvii. Gi. Corp. genie, internum. so nearly to each other as to leave nothing but a nar- row fissure (FS.), between them. The prosencephalic Ge. Corp. genie, ext. T'. Tegmentum, lying in arch surrounding the fossa front of the gray substance surrounding the aq. _ _ __ . °. Sylvii developed its sub- stance mainly in a lateral direction toward the cranial walls, whereas the region of Sylvii. Cp.i. Anterior portion of the inner capsule lying between the caudate and lenticular nu- clei. P. Posterior portion of the inner capsule between optic thalamus and lenticular nucleus,— the origin of the fibres of the cruscerbri. Om. Projection system from occipital lobe to pulvinar corp. genie, int. and ext., the inf. corp. bigemina f_he fossa S. itself increased in (Bi). and to the pedunculus. ., . , ■ , , 1 • „ „ v ; v thickness by developing gan- glia in a median direction toward the cerebral ventricle(Figs.5 and6). The fissure of Sylvius is not a simple passage. It is bounded not only by two protruding lips, the operculum above and the upper temporal convolution be- low (Fig. 6 external to J.), which cover in Burdach's upper and lower fissures, but also by a front and rear wall lying on either side of the island and lapping over the front and rear fissures of Bur- dach, which are thus vertical branches of the upper fissure. These clefts are reached by the ascending anterior and posterior branches of the Sylvian fissure, ramus adscendens anterior et pos- terior fissurce Sylvii. The anteri- or ascending limb lies behind the orbital surface of the frontal lobe, the posterior in front of the base of the first temporal convolution. The two ascending divisions of the Sylvian fissure bound the operculum. In man the island of Reil generally possesses six con- Fig. 6. Frontal Cross-Section from the Brain of Cercocebus Griseoviridis. J. Island of Reil, covered/by the oper- culum and the first temporal convolution. Gf. Gyrus fornicatus. Tb. Corp. Callo- sum. Nc. Nucl. caud. Ve. Lateral ventri- cle. VIII. Median ventricle. Th. Thalam. opt. Fd. Descending, Fa. Ascending fornix. St.i. Lower pedicle of the thala- mus from the ansa peduncularis. II. Chi- asma. II.' A parallel bundle of fibres. Ca. Radiating fibres from . . A. Amygdala. ^^. »xoU.a.u.s ..„._., *.~.. volutions diverging superiorly in temporal region to anterior commissure the shaoe of a fan. These COnvo- L-!« L-3' L-3 Nucl- lenticularis. ans. ansa " " . of lenticular nucleus. Ce. Capsula external lutions are continuous above with CI. Claustrum. Tp. Temporal lobe. 6 Psychiatry. the convolutions of the operculum, behind and below with the superior surface of the first temporal convolution (Huschke). Naturally enough these convolutions cannot be made out unless the entire brain-axis, together with the island of Reil, be dis- sected out from the surrounding mantle; the eye then rests upon the median surfaces of the hemispheres, and upon the inner surface of the convex hemispherical arch. The ascending divisions of the Sylvian fissure are not as well marked on the convex surface of the human brain, and still less on that of the monkey, as they are in the brains of the carnivora (Fig. 7). Among the last named this peculiarity of configura- tion is most distinct (Fig. 7). In the brain of the bear we External Aspect of a Bear's Brain. Fr. Occ. Tm. Frontal, occipital, and temporal end. Olf. Olfactory lobe. Unc. Uncus. Cbl. Cerebellum. Obi. Medulla oblongata. <#.§., |la. $lp. Sylvian fissure, anterior and posterior ascending limbs. C. Central fissure. SI.1 Parallel fissure. §l.a Interparietal fissure. A. Region of the anterior, P. Region of the posterior, central convolution. Arc. I., Arc. II. Lower and upper parietal arches. L.1, L.a Inferior parietal convolutions. L.3 Superior parietal lobe. L.1, L.2, L.3 Temporal convolu- tions. Fus. Gyrus fusiformis. can easily distinguish : a blunt lower end of the operculum (near if. 15.), a highly developed a?itcrior ascending (|la.), and a long pos- terior ascending, division (iljj.) of the Sylvian fissure. Similarly both the ascending branches are exquisitely distinct on the basal surface of the brain of Mustela (Fig. 10, right). The Sylvian fissure, which, in this instance screens from view an exceedingly rudimentary island, lies immediately adjoining to the lobus olfacto- rius (Ge.) throughout the entire length of the operculum, as was the case with the human brain represented in Fig. 4. In the rudi- mentary olfactory lobe of adult man, this external olfactory con- Convex Surface of the Brain. 7 volution (Fig. 15 Ge., Fig. 18) is replaced by the external white medullary strand of the trigonum olfactorium. In addition to the fossa Sylvii which is present in all mamma- lian brains, and is often the only fossa present, we must now con- sider the typical fissures of more highly developed brains ; these sulci mark out the various regions, or, at least, the convolutions of the convex surface of the fore-brain, and serve as our guides over this area. Apart from its morphological value, a thorough knowledge of these fissures and convolutions is absolutely indis- pensable to a proper appreciation of the physiological experiments performed on the animal brain. If the " type " has been made out clearly enough to enable us to determine the identical regions of the mammalian and the human brain, then the important conclusions from physiologi- cal experiment and from pathological anatomy may safely be compared one with the other. We find typical fissures in the human fcetal brain, from the sixth month of its development on. I have not had the opportunity of making any detailed inde- pendent study of the human fcetal brain. My description is based upon a study of the monkey's brain. In some slight degree I am justified in taking such a course by the saying of v. Bischoff, that the monkey brain is not a miniature model of the human brain, but that the for- rn-rrenrpcenrq arrested Fr. rrontai ena. mer represents arrested 0ccipital end. gu inferior frontal stages in the fcetal de- Tm. Fig 8. Brain of Hamadryas. Frontal end. Tm. Temporal end. Occ, gl.a Rudi- mentary superior frontal fissure. p(£. Anterior radial .° f 1 1 , fissure (sulc. praecentralis). rC Ecker's interparietal VClopment Ot tne lat- fissure (postenor radial fissure). |.S. Fissura Sylvii. ter. In the brain of $P- Ramus ascendens posterior. SI.1 First temporal ' 1 . . ^ fiss. (sulcus parallel.) §. occ. i. External occipital fissure the monkey tliere IS a (primate ape fissure), arc. occ. Occipital lobe. (The verv distinct median vertical fissure might answer to the interoccipital fissure \ . ' of man, and the horizontal fissure might mark Ecker's radiating furrow (£.), gyrus occipitalis, arc. II. Superior parietal arch. cnlm* rentralis or fis- arc- L Inf' pariet. arch. L.1, L.2, L.3 First, second, SU1CUS cen iraub, ui us and third temporal convoiutions. gp. 0. Sulcus praeoc- SUre of Rolando. The cipitalis. Cbl. Cerebellum. Obi. Med. oblongata. 8 Psychiatry. radiating, primary fissures diverge toward the upper border of the hemispherical arch, and converge toward its lower border, i. e., toward the operculum. In front of the sulcus Rolando, mark- ing the posterior boundary of the frontal lobe, lies the ante- rior median furrow, the sulcus prcecentralis, " transverse frontal fissure," (Fig. 8, pd). This sulcus praecentralis combines with the straight inferior frontal fissure (SI.1) to form a cross-bow. Convexity of the Human Brain. J. Island of Reil. -shaped, deep fissures, which converge toward the gyrus fornicatus, of which the superior is called sulcus occipitalis internus ; the inferior, sulcus calcarinus or sulcus hippocampi. The latter is one of the most constant fissures of the primate " type "; it can be discovered in the smooth- est primate brains which have not yet developed either the exter- Median Surface of the Brain. 23 nal occipital fissure or the sulcus centralis. It receives its name from the calcar avis (pes hippocampi minor); for, in conse- quence of the thinness of the mesial wall of the posterior horn, the calcar avis is formed by the convex protrusion of the reverse (Kehrseite) of this fissure upon the inner wall of the posterior cornu, just as the next following eminentia collateralis Meckelii is the converse of the next external fissure (S.O.T.F.). The sulcus calcarinus is in reality (--shaped (Fig. 14, £c). In the human brain the longitudinal fissure of the h- is frequently bent at an angle open posteriorly. In addition to the occipital fissure of the sulcus calcarinus, the median surface presents two other fissures—Ecker's occipitotemporal fissures. But these fissures do not be- long to that vertical portion of the median surface consid- ered hitherto, which lies next to the falx cerebri; for we must remember that the me- dian surface lies in two planes meeting at an obtuse angle. The longer vertical division, extending from the frontal end to the extremity of the oc- ciput, rests upon the falx cer- ebri ; the oblique horizontal division, extending merely from the vertex of the occi- put to the temporal end, rests upon the tentorium cerebelli. The superior occipito-temporal fissures, together with the sulcus calcarinus, bound a convolution which, tapering anteriorly and uniting with the uncinate convolution, bears a strong resemblance to the tongue of a dog; whence the name, gyrus glossiformis. The superior and inferior occipito-temporal fissures enclose a con- volution broadest at its middle part, the gyrus fusiformis. The latter is succeeded by the marginal convolution of the temporal lobe, which belongs to the convex surface as well (L.3). At the anterior margin of the temporal lobe the three convo- lutions of the convex surface unite to form a transverse-arched Fig. 14. Median Surface of the Primate Brain. Fr., Tm. Front., temp. end. L.occ Occipital lobe. an. Calloso-marginal con- volution. S.occ. Internal occipital fissure. St. Sulcus calcarinus. L.s Third frontal con- volution. Gf. Gyrus fornicatus. Qu. Lobus quadratus. Cun. Cuneus. Gl. Gyrus glossi- formis. Fus. Gyr. fusiform. L.3, L.2, L.J Temporal convolutions. A. Amygdala, unc Uncus. Trb. Corp. callosum (trabs). Sp. Septum pellucidum. Th. Optic thalamus. Ca. Anterior commissure. 24 Psychiatry. anastomosis, the cortical portion of which connects with the uncus (Fig. 14, Tra.). A few words must here be added in reference to the cornu ammonis, a formation complicating the temporal end of the gyrus fornicatus. The cortical substance of the gyrus fornicatus, which terminates with a free margin over the corpus callosum, presents on cross-section, as does every convolution, an [\ surrounding the medullary substance. Burdach applied the term cingulum to that portion of the gyrus fornicatus extending as far as beneath the splenium corp. callosi. At the temporal end of the gyrus forni- catus, in the gyrus uncinatus, the cortical substance does not ter- minate with a free margin until the middle portion of the convo- lution has undergone an c/vshaped involution (lamina convolutd), and the anterior portion has been divided up wave-like ; the wave- tops facing the inferior cornu are the so-called digitations. The C/>shaped involution allows the margin to become quite free, to rise above the medullary substance, and to appear from out of the involution-folds as the fascia dentata, the notched surface of which is due to constricting vessels. The cornu ammonis possesses four adjacent longitudinal emi- nences: 1. The gyrus uncinatus, the core of the cornu ammonis subiculum. 2. The Fascia Dentata Tarini (Fig. 13). The Fasciola Cinerea (Arnold) is a continuation of the fascia dentata on the lower surface of the great commissure, commissural fibres having perforated part of the median cortical substance some distance behind the septum. The cornu ammonis, unusually large in car- nivora, rodents, etc., lies adjacent to a considerable portion of the inferior surface of the corpus callosum, and extends far to the front. 3. The Fimbria (Sus.), the projection bundle of the cornu ammonis. Additions from other parts of the gyrus fornicatus transform this into the descending crus of the fornix (Fd.). 4. The Alveus (Fi.), whose white substance, spread over its own gray cortex, over the stratum convolutum, is continued into the cord-shaped fimbria. The alveus represents the ventricular surface of the cornu ammonis, which surface protrudes as though it were embossed from the inner wall of the inferior horn. The name, alveus, is quite appropriate, for the stratum convolutum is surrounded trough-like by this medullary layer, which is curved, with its convex surface toward the ventricle. The surface of the subiculum, together with the fascia dentata, forms a continuous cortical layer, which is folded CO-shaped from right to left. The surface of the cortex is hid in Basal Surface of the Brain. 25 the depths of this scroll. The alveus corresponds to the white substance of the convolutions, and consequently faces the ven- tricles. The uncus also is a complex formation. Its apex or pos- terior portion alone contains the forward part of the free median cortical margin, which projects in a transverse direction above the subiculum cornu ammonis, and yet is a part of it. The appear- ance of a longitudinal anterior flexion of the uncus, distinct from the cornu ammonis, corresponds to the convexity of the amygdala; this irregular gray mass lies in front of the cornu ammonis, and covers over the fissure which otherwise would have been evident on the brain surface at its point of flexion. B.—GANGLIA OF THE PROSENCEPHALON, THALAM- (DI-) ENCEPHALON, MESENCEPHALON, AND MET- ENCEPHALON. The cerebral cortex and its white substance betray, as it were, their morphological independence by the constriction round about the ganglia (Fig. 4, g. Fig. 15), and by the ease with which both the island of Reil and its medullary substance can be dissected out from the external surface of the large ganglia. This anatom- ical independence is in accord with the physiological indepen- dence of the fore-brain from the rest of the brain-substance. In order to demonstrate the entire natural surfaces of the cerebral ganglia and the brain-axis, it is necessary to remove, by a curved incision (passing around the island, on the floor of the Sylvian fossa), the hemispheric arch, which corresponds to the cerebral mantle of Burdach, in contradistinction to the nucleus of the brain (which includes the island, ganglia, and brain-axis). The basilar aspect of the nucleus, or rather of the brain-axis (Fig. 15), presents a free surface in front of the tractus opticus, in the form of the lamina anterior perforata ; the last-named structure is separated on each side from the island by the external olfactory convolution, before it joins the gyrus uncinatus. The convolu- tions of the island converge toward this external medullary strip of the trigonum olfactorium, and form generally a single convolution,—the peduncle of the island,—while they diverge above toward the operculum, dividing into five or seven convolu- tions. To the inside of the island, and the outside of the caudate nucleus, extends the plane of the incision which separated the trunk from the cerebral mantle, as though the former were the 26 Psychiatry. nucleus of the latter. Just-as the hemisphere describes an arc with a frontal, a temporal, and an occipital end, so we shall find that the collected fibres of the corona radiata describe a similar curve at their entrance into the cerebral trunk. A section like that repre- Fig. 15. Basilar Surface of the Brain-Axis and the Cerebellum. Fr., Tm., Occ. Section through the medullary substance, radiating from the frontal, temporal, and occipital portions of the cerebral mantle into the brain-nucleus. Js. Island of R. Ge. Med. substance upon the external olfactory convolution, pa. Lam- ina perforata anterior. II. Optic nerve ; the optic tract is divided posteriorly into an internal and external division. J. Infundibulum. M. Corp. mammillare. L. Lamina perforata posterior. Pd. Pedunculus (crus) cerebri. T. Tractus transversus pedunculi. V. Pons Varolii. P. Pyramis. O. Olive. R. Restiform body. M. Medulla spinalis. III. Oculomotor nerve. IV. N. trochlearis. V. Trigeminal nerve. VI. N. abdu- cens. VII. Facial nerve. VIII. Auditory nerve. IX. N. glossopharyngeus. X. N. vagus. XL N. recurrens Willissii (accessory nerve). XII. Hypoglossal nerve. Qu. Inferior surface of the lobus quadratus, superior lobe, of the cerebellar hemisphere. p.p. Lobus posterior inferior (semilunaris inferior), i.i.i. Lobus inferior (lobusgracilis et biventer). A. Amygdala. Fl. Floculus. Vr. Inferior vermiform process. sented in Fig. 15 will present, therefore, a frontal and temporal end (F., Tm.), and between the two a parietal and occipital sur- face (Figs. 16, 17, P. Occ). The marked convexity of the trunk, Basal Surface of the Brain. 2 7 causing it to protrude to the outside from under the insular con- volutions at this section of the corona radiata, is due to the large size of the nucleus lenticularis, and furthermore, the temporal portion of this nucleus projects below the cerebral trunk, a fact not easily appreciated on the hardened specimen, owing to the slight contrast of colors. The caudate nucleus passes from the fore- brain to the basal surface, and there appears as the lamina per- forata anterior. The inter- and mid-brain are continued to the base of the cerebrum through the tractus opticus, which, together with the optic chasm, constitutes the posterior boundary of the fore-brain. The infundibulum represents a basal continuation of the cere- bral gray substance surrounding the cavity of the thalamen- cephalon, and the corpus candicans must be regarded as medul- lary substance of the thalamencephalon, extending from the anterior tubercle of the thalamus to the basilar surface of the brain (Fig. 13, m. a.). The pedunculus (Pd.) cerebri, which is bound from the fere- brain to the spinal cord, covers superficially and laterally the anterior circumference of the mesencephalon. The median lamina perforata posterior forms the median suture (raphe) of the two halves of the tegmentum, which connects the mes- and thalamen- cephalon with the spinal cord. The pedunculus cerebri is crossed in the median line by the third pair of nerves (III. oculomotorius). The left half of the pedunculus cerebri is crossed by a tract (the tractus transversus pedunculi—Gudden) which does not always lie as superficially as here indicated, and connects the anterior end of the peduncle with the superior corp. bigemina. The pons Varolii constitutes the basilar surface (Kehrseite) of the epen- cephalon. From the pons there emerge laterally the trigeminal nerves, and inferiorly the abducens, facial, and acoustic nerves. Below the pons Varolii and between the pyramids we come upon the anterior furrow of the cerebral axis, which is well developed even between the crura. The floor of the anterior cleft is analogous to the lamina perforata posterior lying between the paired peduncles, for its floor reveals the median surface of the raphe of the posterior division of the oblongata. And in the oblongata also the raphe divides the prolongations of the tegmen- tum. The oblongata can be divided into a superior and an infe- rior portion. The superior division extends to the inferior mar- gin of the olivary bodies, the inferior half to the last decussating 28 Psychiatry. bundle of the pyramids. A longitudinal fissure divides each olivary body toward the median line from the pyramids, and toward the outer side from the region of the funiculus lateralis. On closer examination the olivary bodies do not appear smooth, but striated, owing to the passage over them of oblique, slightly de- scending bundles (stratum zonale) which originate in the corpus restiforme. As a rule the stratum zonale covers all the longitu- dinal fibres coursing back of the pyramids. If there be a super- ficial longitudinal bundle of fibres to the inner side of the olivary body,! this is termed fasciculus internus; if to the outside, fasciculus externus. These fasciculi may be nothing more than pyramid-fibres which the stratum zonale has separated from the main body of the pyramids. Behind the funiculus lateralis of the oblongata lies its most posterior longitudinal elevation, the corpus restiforme of the cerebellum (Fig. 15, R.). The basal surface of the human isthmus presents many striking points of con- trast with the brains of lower mammals, and even with the brains of primates. Entire structures seem to appear, or to disappear in man. On the whole, the pedunculus cerebri in animals is thin and possessed of so few nerve-tracts that the convexity of the tegmentum lying behind it causes it to bulge out. The pons appears low ; next to the pyramids rectangular formations are visible, which are made up of transverse fibres (corpus trapezoides). The olivary body, common to man, is wanting. I have shown that all these differences of structure depend upon a single factor, upon the greater or lesser development of the fore-brain. The smallness of the prosencephalon in mammals causes a diminution in the thickness of the peduncles, in the height of the pons, and in the size of the pyramids. Similarly the appearance of the trapezoid body and the disappearance of the olivary structures are due to the smallness of the fore-brain. If a mammal, whose basilar brain surface is as unlike that of man as is the brain of mustela (Fig. 10,p. 9), could develop mightier hemispheres, then its pes pedunculi would increase in size and in the number of its nerve-bundles, for the simple reason that the pes pedunculi to a great extent is the continuation of the medullary substance of the hemisphere. Furthermore, inasmuch as the pyramid, the prolongation of the pesped. gets but a part of the latter's fibres, while a large part of peduncular fibres reach the cerebellum by way of the brachium-pontis, we shall find that the number of bundles in the pons is dependent in the first instance upon the number in the pes pedunculi, and in the second instance upon the greater development of the medullary substance of the hemispheres, and upon the cerebral cortex whence the white fibres emanate. The more highly developed these cerebral masses, the greater the extent of surface from which the pons-fibres can originate. In man the pons is so excessively developed longitudinally that the deeper transverse bundles are completely covered. In mammals the pons is so short that the course of the pyramid is usually prolonged anteriorly and remains uncovered. For this rea.son transverse bundles (corp. trapez.) come into view next to the pyramids ; in man these bundles are screened from view by the lower bun- dles of the pons. The corpus trapezoides is, therefore, not wanting in the human brain ; it is simply invisible on the basilar surface. Nor is the olivary body really missing in animals. It lies behind the pyramids ; from this position it has been dis- lodged by the thickness of the pyramids in man, pushed to the side, and made visible Basal Surface of the Brain. 29 on the basilar surface. We infer from such facts as these, and from a study of the ganglia, that the cerebral structure is governed by a law which establishes a harmonic dependence between the formation of the brain-axis and the development of the functionally highest organ— the fore-brain. (Details to be found in the " Mittheilun- gen der anthropologischen Gesellschaft," Vienna, 1870.) The peculiarities of the human brain might not inappropriately be said to be "mentalized " (durchgeistigt). The lower forms of primates possess brains whose trunk shows evident transitional stages between the lower and higher forms ; for in- stance, the visible co-existence of olivary and trapezoid bodies (Fig. 11, p. n, Rh. O.). In the highest carnivora (bear) the brain presents the higher transitory stages of brain- structure. So early an observer as Stannius knew of the olivary bodies of water- mammals. The metencephalon, the oblongata, is apparently longer in animals owing to the shortness of the pons, and, with the exception of the pyramids, more highly developed than in man. At the level of the entrance of the brachium pontis,1 in front of the seventh and eighth pairs of nerves, the cerebellum presents the sulcus magnus horizontalis. From the basilar surface the lobus quadratus (Qu, Fig. 15) can be seen projecting above this furrow. This furrow divides deeply the convolutions of the pos- terior (subtentorial) surface of the cerebellum from those of the basilar surface. For our purpose it will be sufficient to give Henle's simplified account of the basilar lobes. From before backward we have the following hemispheric lobes: 1. The floc- culus. 2. The amygdala. 3. The inferior lobes (i. i. i.). 4- The posterior inferior lobe (pp.). The median lobules: 1. The nod- ulus. 2. The uvula. 3. The pyramid. 4. The posterior vermis. These are not basilar formations, but are covered by the oblon- gata. The pros- and thalam-encephalon possess surfaces which look from below into the cerebral ventricles, and consequently can be viewed from above (Fig. 16). A bundle of fibres belonging to the corpus striatum corre- sponds to the stria cornea (stria terminalis), which appears to be the line of demarcation between the fore- and inter-brain. The fibres of this bundle from the temporal lobe pass into the nucleus caudatus along the entire length of the latter. The surface of the stria cornea is formed by a fold of the ependyma, underneath which a large vein collects the lateral branches from the surface of the nucleus caudatus, and conducts them to the vena magna Galeni, which is lodged in the membranous covering of the corp. quadrig. The cavity of the thalamencephalon (ventriculus III.) forms a circle around the commissura media. If the mem- 1 Processus cerebelli ad pontem. 30 Psychiatry. branous roof of the 3d ventricle were left intact, and the aquae- ductus Sylvii properly closed, a cast of the third ventricle could be taken. This cast of the third ventricle would resemble a ring whose opening would correspond to the median commissure. Interiorly the ring would present a projection—the cast of the Fig. 16. Superior and Posterior Aspect of Brain-Axis, and of the Cerebellum. J. Island. F., P., Occ. Frontal, parietal, and occipital portions of the projection- system (radiating fibres from the cerebral mantle). Nc. Caudate nucleus. Str. Stria cornea. L. Linea aspera. Ca. Anterior commissure. Fx. Descending crus of the fornix. V. III. Ventricle. Cm. Median commissure. Th. Optic thalamus. Tb. a. Its anterior tubercle. Tb. m. Its median tubercle (elevation). Pulv. Pulvinar. con. Conarium, from the sides of which the plastic longitudinal prominences of the habenu- lae originate. Bs. Superior bigeminal body. Bi. Inferior bigeminal body. brs. Superior arm. bri. Inferior arm of the corpus quadrigeminum. CI. Central lobule of the superior vermiform process. M. Its mons. Fc Folium cacuminis (the top layer of the superior vermiform process). Al. Aire of the central lobule. Qu. Lobus quadratus (Burdach), superior lobe (Henle). Sl.s. Superior posterior lobe (lobus semilunaris superior). Sl.i. Lobus semilunaris inferior—(posterior lobe—Henle). infundibulum. Behind the caudate nucleus and the stria, the anterior commissure and the descending crus of the fornix, we come upon the thalamus opticus. Its surface is on the whole wedge-shaped, with a blunt anterior edge, and a convex, broad- faceted ending at the pulvinar. Its median surface is composed of Dorsal Surface of the Brain. 31 gray substance (central cavity gray), for the united cavities of the primary medullary tube, from the primary anterior cerebral vesicle to the end of the spinal cord, are lined with gray substance. This narrow gray, cleft-shaped covering of the diencephalon passes in the mid-brain into the aquaeductus Sylvii. The surface of the optic thalamus presents several promi- nences. The best defined is the habcnula—the pineal peduncle (between brs. and F. Fig. 17). The surface of the habenula, which is connected on both sides with the pineal region, receives white substance, as does the entire surface of the optic thalamus, from the stratum zonale. The stratum zonale of the habenula has the appearance of being torn off anteriorly after the removal of the adherent membranous covering of the ventricle from the ependyma. From this the false inference was made that there was a connection between the fornix and the habenula. The other three prominences are: 2. The anterior nucleus (Genu anterius Gratiolet) which tapers tail-like backwards and outwards tuberculum anterius (Burdach), (Fig. 16, Tb. a.). 3. The median protuberance (tuberculum medium) which arises simply from the demarcation of the tuberculum anterius and the flattening of the thalamus behind the habenula. 4. The pulvinar, which in the human subject presents the shape of a posterior free eminence. In man the ganglia of the mesencephalon, the corpora quadri- gemina, encroach considerably upon the thalamencephalon. The corpora quadrigemina comprise the superior and inferior corpora bigemina. If the pineal body (Fig. 17, con. cp.) be lifted from the back, we come upon a flat triangular space with its base above, situated in the middle of the superior corp. bigem., which receives the posterior surface of the pineal bod}-, and bends over into the posterior commissure. We notice also a groove, open posteriorly, between the upper surface of the corp. quadrigemina and the lower surface of the conarium which latter rests upon the former. This is the reverse of the convexity of the posterior commissure (convex anteriorly). This commissure, with its convexity looking anteriorly, is, therefore, a transverse curved medullary lamina, and not a funicular formation, as it would seem to be from an inspec- tion of the anterior surface. The superior corpus bigeminum is connected with an apparently columnar medullary border, which, concave posteriorly throughout its entire length, courses between the pulvinar and the corp. genie, internum. This is the brachium corporis bigemini superius (Fig. 17, brs.). The superior terminus 32 Psychiatry. of the corpus geniculatum internum also tapers toward the corp. bigeminum superius, combining with the latter under the brachium corp. big. sup. mentioned above. The arm of the superior corp. quadrigem. forms the boundary between the mes- and thalam- encephalon. The superior half of the corp. bigeminum inferius is connected with a flat, white medullary structure appearing from under the internal genu (brachium corporis bigemini inferius, Superior and Posterior Surfaces of the Prosencephalon of the Lobus Candicis (Stamm- lappen), of the Thalam-, Mes-, and Met-encephalon. J. Island. P., Occ. Parietal and occipital divisions of the projection systems from the cortex. Nc. Caudatus nucleus. St. Stria cornea. Th. Opticus thalamus. Pulv. Pulvinar. Ge. External geniculate body. Tr. Optic tract, Gi. Internal geniculate body. V. Third ventricle. Hb. Habenula conarii. con. Conarium. cp. Posterior surface of the posterior commissure. Fr. Frenulum, ve. Superior medullary velum. Bs., Bi. Corpus bigeminum superius et inferius. brs., bri. Brachium superius et in- ferius corporis quadrigemini. Ls., Li. Lemniscus superior et inferior (upper and lower fillet). 5. Locus cceruleus. £- Auditory nucleus. 7. 6. Common nucleus of the 6th and 7th nerves. 8. Ascending root of the 8th nerve (Engel) ; auditory rod (Bergmann). Br. Brachium pontis. R. Restiform body. Pr. Processus cerebelli ad cerebrum. VIII. Striae medullares nervi acustici transversae (auditory striae). 12. Region of the hypoglossal nucleus. 10. Ala cinerea, Arnold (nucleus of the vagus.— Stilling), ob. Obex : cun. Fasciculus, cuneatus Gr. Fasciculus gracilis. RL. Tuber- culum cinercum, Rolando. L. Lateral column. Note.—This drawing reproduces the exact position of the cerebral trunk within the cranial cavity, taking into account the parietal flexure. Fig. 17 bri.). This brachium passes into the white surface of the inferior corp. bigem. The superior corp. bigem. presents a gray surface, more especially, however, in animals. Below the brachium corp. bigem. superius lie the spindle-shaped corpus geniculatum internum (Fig. 17, gi.), and the club-shaped corp. geniculatum externum, growing smaller as it nears the tractus opticus Dorsal Surface of the Brain. 3 3 (Figs. 15 and 17, ge.). The transition of the external corpus geniculatum into the optic tract is the only part that is visible. A distinct demarcating furrow lies at the boundary between the corp. genie, int. (gi.) and the inner ribbon-shaped stria of the tractus (Fig. 15). This band appears (at least below the surface) to pass into the thalamus in front of the corp. geniculatum inter- num. If the size of any brain-formation increases with an increase in the size of correlated parts, we may maintain that the cor- pora geniculata will increase with a large growth of the corp. quadrigemina ; and, on the other hand, that in these very animals disappearance of the pulvinar causes a diminution in the size of the optic thalamus. I was the first to observe, on longi- tudinal sections through the brain of a new-born cat, that the largely developed corp. genie, externum ascends beyond the optic thalamus. Forel has given a more detailed description of this. Accordingly, the development of the corpora geniculata is dependent upon the mesencephalon. Between the posterior corp. bigemina a paired-conical column is formed, which, broadening below, passes as frenulum into the velum medullare superius (Fig. 17, ve.), and is continued through this into the medullary substance of the vermis superior. The velum m. superius is covered by the lingula, the foremost convolutions of the vermis superior. Stilling discovered the delicate bands of the lingula which lie immediately upon the processus cerebelli ad cerebrum (Fig. 17, to the right of Ling). The posterior surface of the after-brain would come into view below the mid-brain as a freed portion of the brain-axis, were it not covered by the cerebellum (hind-brain), (Fig. 16), which must be severed at its connecting arms, if a view of said posterior surface is to be obtained. On the posterior subtentorial surface of the cerebellum the mons (Fig. 16, M.), the third and largest division of the superior vermiform process, covers the central lobe, and this in turn the lingula of the medullary velum. In a similar fashion the lateral divisions of the lingula (frenula lingulae) are covered by the lat- eral portions of the central lobe—by the alae, and these in turn are concealed on each side beneath the hemispheres of the mons, the lobis quadratus (lobulus superior of Henle), which projects ante- riorly beyond the alae (Fig. 15, 16, Qu.). The great horizontal fissure divides the posterior lobes of the upper and lower surfaces of the cerebellum, the inferior post, lobe projecting in an occipital 34 Psychiatry. direction beyond the former (Fig. 16). In spite of the manifold numerical relations and the still more manifold modes of connec- tion between the median and lateral cerebellar covolutions (which Malacarne counted and Stilling studied most thoroughly), an in- crease of lateral convolutions seems to correspond to an increase of median convolutions; and yet there is one exception to this superficial agreement. At the bottom of the great horizontal fissure all the convolutions of the upper semilunar lobe (superior posterior lobe) are united into a single layer of vermiform con- volutions, in the folium cacuminis (Fig. 16, Sl.s. Fc). After dissecting away the cerebellum, we can examine the pos- terior surface of the brain-axis. The middle of the region between corp. quadrigemina and the entrance of the processus cerebelli into the cerebellum is taken up by the velum medullare (Fig. 17, to the right), developed from the frenulum. The velum medullare and the frenulum constitute the genuine processus cerebelli ad corpus quadrigeminum. At the superior margin of the valvula cerebri, the IV. nerve is seen to emerge (Fig. 17, IV.). The lateral mar- gins of the velum verge upon two powerful flat bands, the Binde- arme; these were falsely termed processus ccrebelli ad corpus quadrigeminum, and correctly described by Stilling as the processus cerebelli ad cerebrum) The processus ad cerebrum begin not quite at the corp. quadrig. and terminate at the frenulum lingulae (Fig. 17, right), in the direction of the cerebellum. Superficial bundles of fibres emerging from the velum medullare and twining around these processus ad cerebrum, separate them from the corp. quad- rig. ; these bundles pass outward and' over the processus cerebelli and disappear in the pons. Leveille" was the first to make a drawing of these bundles. I have designated these bundles as lemniscus inferior (Fig. 17, Li.), or the cerebellar bundles of the fillet. The superior fillet, the lemniscus Reilii (laqueus, Fig. 17, Ls.), covers the proces. ad cerebrum like a three-cornered cloth, lying external to the inferior fillet. The superior fillet ex- tends from the corp. quadrigemina to the superior margin of the pons. Toward the base of the brain, on the posterior surface of the trunk, the posterior aspects of the pes pedunculi and of the pons are visible. But the fillets, the processus ccrebelli ad cere- brum, and the velum medullare inferius, constitute a series of structures which reach the surface behind the pes pedunculi 1 Superior peduncles of the cerebellum.—S. The Brain-Axis. 35 as integral parts of the tegmentum. In the cerebellum the con- tinuation of the velum medullare and of the processus ad cerebrum passes with a free medullary surface behind the rhomboid fossa and there forms its roof—the tectum fossce rtiomboidalis. The central gray substance (Hohlengrau) of the mid-brain, sur- rounding the aquceductus Sylvii, dilates beneath the velum medul- lare into a semicircular canal with furrowed floor. The divergence (cordward) of the processus cerebelli ad cerebrum allows the central gray substance to expand most in breadth between the points of emergence of the two auditory nerves. At this level the corpora restiformia, which appear to be in direct connection with the funiculi graciles approach the central gray to form its lateral boundaries. These structures, converging symmetrically at the lower end of the central gray substance, cause the latter to deepen at the same time that it tapers to a point. Inferiorly the fossa is continued in funnel-shaped fashion into the central canal of the after-brain. This portion of the central gray substance has been termed fossa rhomboidea, from its being enclosed between the angles formed by the meeting of the processus ad cerebrum above, and the pedunculi cerebelli beloAV. The fossa rhomboidea has important relations to the origin of cerebral nerves. The same median furrow which was noticed on the ventral wall of the aquaeductus Sylvii divides the fossa rhomboidea into symmetrical halves. The cmincntice teretes, descending from the aquaed. Sylvii on each side of the median furrow, enlarge into an oval area just above the exit of the eighth nerve (Fig. 17, 6,7). To this area, which is intersected by the central fibres of the sixth and seventh pairs of nerves, various names have been given, viz.: abducens-facialis nucleus—Stilling and Clarke ; abducens nu- cleus—Deiter; facial nucleus—Schroder. To the outer side of the superior half of this prominence we find a long bluish groove, the fossa cosrulca. The bluish color is due, according to the laws of refraction through cloudy media, to the black cells lying immediately below the transparent ependyma. I have shown that in this groove one of the roots of the fifth nerve takes its origin (Fig. 17, 5). Below (caudad) the fossa cosrulca, the eminentia teres is bounded laterally by a rhomboid elevation (region of the viii. nucleus), extending at its broadest part quite to the median line. Striae medullares transversae—the superficial posterior roots of the viii. nerve—may divide this rhomboid into a superior 36 Psychiatry. and an inferior triangle (Fig. 17, viii., 8, 8). The acoustic nuclei and the joint nucleus of the sixth and seventh nerves are frequently separated from one another by several obliquely ascending striae medullares (Engel, Bergmann, " Klangstab ") ; occasionally the eighth nucleus is covered by these strice (Fig. 17, 8). The inner margin of this elevation bears on the lower part of the rhomboid fossa to the outer side. But this does not increase the breadth of the eminentia teres, for its outer margin converges below tow- ard the median furrow, to bound a median triangular elevation, which represents the region of the hypoglossal nerve (Fig. 17, 12). There are two reasons for this narrowing (below) of the eminentia teres : First, the converging fasciculi graciles crowd in upon the fossa rhomboidea at its point of junction with the canalis centralis ; and, secondly, the triangular nucleus of thetenth nerve, enlarging from above downward, edges its way in between the origin of the eighth nerve and the eminentia teres (12). This is the posterior nucleus of the vagus, which a grayish color (ala cinerea—Arnold) distinguishes from the whiter triangular area of the twelfth nerve, which area is covered by transverse fibres of the vagus. A sling-shaped commissure, pointed below, often causes a columnar elevation of the ala cinerea. At the lower angle of the rhomboid fossa, the obex (Fig. 17, Ob.) effects a connection with the membranous roof of the metencephalon and the choroid plexus, similar to the passage of the fimbria into the membranous roof of the diencephalon. This obex, a vestige of the fcetal posterior roof of the rhomboid fossa, remains attached to the latter when it is separated (by dis- section) from its surrounding parts. Let us pass now to the consideration of the nerve-tracts to be found on the posterior aspect of the after-brain. The corpus res- tiforme1 emerges from the cerebellum at about the level at which the posterior surface of the processus cerebelli ad cerebrum dis- appears within that organ (Fig. 17, R). Above the decussation of the pyramids, which begins a short distance below the formation of the central canal, the corpus restiforme appears to divide into the funiculus cuneatus and f. gracilis (Fig. I7,cun., Gr.). And yet the latter are separated from the corpus restiforme by a shallow transverse furrow, which proves that they cannot arise from a division of the corp. restiforme of the same side. The origin of the funiculi graciles et cuneati is marked by gray substance within these plastic, conical protuberances. 1 Pedunculus cerebelli ad medullam oblongatam. Architecture of the Brain. 37 Both these tracts of the oblongata are continued into the pos- terior columns of the spinal cord. In the cervical spinal cord the triangular column of Goll must be regarded as the prolongation of the funiculus gracilis. These posterior columns of the oblongata do not abut, as is the case in the spinal cord, upon the lateral columns (Fig. 17, L.). The superior beginnings of a caput cornu posterioris, of the gela- tinous substance of the gray spinal-cord nucleus, forces its way to the surface between the lateral and posterior fasciculi of the after-brain. This spindle-shaped mass—the tuberculum cinereum Rolando (Fig. 17, RL.)—is the nucleus of origin of the ascending root of the fifth nerve, which is equivalent to a posterior root of the spinal cord. The posterior roots of the spinal cord also pos- sess ascending fibres. The root of the fifth nerve covers the sub- stantia gelatinosa with a thin film of fibres, which do not alto- gether hide the gray nucleus (tuberculum cinereum) below. On the anterior basilar surface of the trunk the inferior margin of the olivary body marks the boundary between the upper and lower halves of the oblongata, which are more unlike each other in structure than are the superior half of the oblongata and the inferior half of the pons. On the posterior surface of the isthmus, the boundary of the lower half of the after-brain is marked best by the superior margin of the tuberculum cinereum Rolando. The plastic prominences of the lower half of the oblongata are in due succession from the anterior to the posterior median furrow of the trunk, as follows : 1. The pyramids bounded by the roots of the hypoglossal nerve (Fig. 15, P.); 2. Funiculus anterior; 3. Funiculus lateralis (lateral column),—these are separated by the anterior roots of the uppermost cervical nerves (Fig. 15, 1, 2,); 4. Tuberculum cinereum Rolando (Fig. 17, RL.); 5. Fasciculus cuneatus (Fig, 17, cun.); 6. Funiculus gracilis (Gr.). The final transition of the medulla oblongata into the spinal cord is effected at the level of the origin of the third cervical nerve. REMARKS ON THE ARCHITECTURE OF THE BRAIN. The structure of the cerebral cortex, like that of a crystal, can be studied best from its cleavage-surface. The cortical sub- stance contains two distinct sets of fibres. First, we observe that the cleavage-surface of each convolution presents radiating fibres which are quite distinct from radiating blood-vessels ; that this' 3 8 Psychiatry. system of cortical fibres passes into the medullary substance of the convolutions, and from there into that mass of internal white substance (in each hemisphere) which is known as the centrum semiovale of Vieussens. These radiating fibres extend ap- parently quite to the surface of the hemisphere ; such at least we should judge to be the casejrom an inspection of the many torn sections of convolutions in figures 18, 20, 21. The cortex exhibits on the convexity of each convolution the shape of an inverted [\, which is changed in the next adjoining fissure to an upright U (top and bottom of the cortical wave'). Cleavage of the cortex from the top of the inverted [\ opens up the sub- stance of the hemisphere in a direction parallel to the radial fibres. Secondly, in separating the cortical substance of any con- volution from its adjoining white substance, we discover another set of medullary fibres taking a very different direction from those previously mentioned ; these are the arciform fibres, or fibrae propriae of the cortex. If we attempt to dissect away the gray substance alone from the bottom of a fissure, and to remove the U- shaped mass, we obtain a smooth surface which it would be quite impossible to get at the summit of a convolution, where the cor- tical substance is continuous with the radiating fibres. This might lead one to suppose that no radial fibres start inward from the bottom of a fissure ; but in reality this appearance is due simply to the preponderance here of arciform fibres. The depressed surface of a cortical wave can be easily dissected out as from a smooth medullary groove, which on closer inspection is seen to consist of U-shaped medullary fibres. These valleys and fissures of the outermost medullary substance of the cortex, which when devoid of the cortical gray are left wider than the fissures between the convolutions, resemble the half of a gun-bar- rel constructed of wire rings. The smooth enucleation of the cor- tex is possible in the longest fissures and in the shortest insular formation ; and can be equally well effected at every depth, and whether the fissures be between primary, secondary, or tertiary convolutions. (Figs. 18-21 give a faithful representation of the convolution valleys and their vertical U-shaped cleavage sections. Some of them are marked As—association fibres.) These valleys and their U-shaped fibres are enclosed on both sides by radiating bundles, but this mode of dissection does not afford an insight The original reads : " Wellenberg und Wellenthal der Rinde."__S. Association Fibres of the Brain. 39 into the fine network of fibres by means of which radiating fibres pass also into the narrow cortex at the bottom of a fissure (which Henle incorrectly denies). On-the other hand, there is every evidence here of the part the U-shaped fibres take in the formation of the summit of a convolution. The U-shaped bundles of the cortex do not necessarily extend simply from one convolution to the one next adjoining, but they may skip one, two, three, or an entire series of convolutions, and may thus join con- volutions which are united among themselves to a convolution ly- ing at some distance from these. The shortest fibrce proprice lie nearest to the cortex ; the longest at the greatest depth, and are separated from the cortex by other intervening fibrae propriae, the length of which increases gradatim from the surface inward. The fibrce proprice or arciform fibres, must therefore be divided into short and long ones ; this division answering to a difference in shape, for the shortest fibres alone present the U-shape, which results from their close adaptation to the walls of a convolution depression. While the long bundles take entirely different direc- tions, determined by the curvature of the surface of the fore- brain, and by the longer or shorter distance these fasciculi have to travel. The fact of greatest importance concerning the fibrae pro- priae is that they begin and end in the "cortex. The direct opposite of these are those bundles of radiating fibres which take their origin in the cortex but end in some peripheral gray substance, nearer to the nuclei of the cerebral nerves, say, in one of the ganglionic masses at the base of the brain. These are not fibrce propria of the cortex ; in their course they project the cortical surface upon every imaginary or artificial plane situated below the fore-brain ; the higher the plane the more complete will this projection be. I propose, therefore, to call these bundles of fibres which origi- nate (but do not end) in the cortex, projection-fibres. This name is justifiable even if we were to consider the course of these fibres in a reverse direction. Let us disregard for the moment all but the complete nervous organization of man. The impressions of the body are conveyed to the brain by the ramifications of all the nerves and their terminal organs; mutatis mutandis we may argue that the cerebral cortex is the surface upon which the entire body is projected by means of these nerves. It is a difficult matter to make an appropriate subdivision 40 Psychiatry. of these intra-cortical fasciculi. The masses of fibrae propriae, which take an arched course, are distinctly limited on their concave surface, but their convex surfaces are connected indirectly by means of graded tangential fibres from the convexity of shorter arches with the investitures of the convolution depressions. Keeping these restrictions in mind, we may distinguish and shall. describe the following special formations : a. The medullary sub- Fig. 18. Dissection of the Cortex and the M edullary Substance of the Median Surface of the Brain. Fr., Tp., Occ. Frontal, temp., and occip. region. Tr. Corp. callosum (trabs cerebri), c. c. Cingulum. As. Fibrae propriae (association fibres). R. Cortex, bi. Fasciculus basalis internus (Burdach). Li. Fasciculus longitudinalis inferior. Olf. Olfactory lobe. La. Lamina perforata anterior, ca. Anterior commissure, unc Uncus. Sp. Septum pellucidum. Th. Optic thalamus, fd. Descending fornix. m. Corpus mammillare. fa. Ascending fornix. Q. Corp. quadrigemina. A. Aquaeductus Sylvii. Pv. Pulvinar thalami. Gi. Internal geniculate body. T. Teg- mentum. Pd. Pes pedunculi cerebri. St.i. Stilus intern, thalami optici. Lp. Posterior longitudinal fasciculus. Above pco., descent of post, commissure, co. Conarium. stance of the cingulum (Burdach); b. Fasciculus arcuatus (Bur- dach, Arnold); and c. Fasciculus uncinatus. The cingulum (Fig. 18, c.) surrounds the corpus callosum. Above it lies a broad groove, due to the removal of the calloso- marginal fissure. The medullary investment of this fissure is contiguous with the cingulum throughout its course, as an inspec- tion of the frontal end clearly shows. The same is the case with Association Fibres. 41 the fasciculi proprii of the gyrus fornicatus which cover the cingulum, and is true also of the superior frontal and parietal con- volutions surrounding this gyrus. Arnold recognizes a fusion of the gyrus uncinatus with the convolution of the cingulum ; if we grant this we must also allow a similar continuity of tissue to exist with the superficial medullary bundles of the marginal convolution. The lowest fasciculus of the cingulum adjoining the corp. callos., the nerve of Lancisi, whose relation to the medullary substance of the cortex was alluded to on p. 21 connects the cornu ammonis with the olfactory lobe by a circuitous route. The cingulum accompanies, as it were, the calloso-marginal de- pression ; it is the means of uniting successively all the indirect (short) and direct (long) cortical connections ; and after effecting this its fibres pass underneath the splenium of the corp. callos. and join (superiorly) the fasciculi proprii., on which the deep depression of the occipital fissure of the sulcus calcarinus with its medullary arches had rested. These medullary arches connect the cingulum with the gyrus glossiformis. The posterior half of the upper occipito-temporal fissure exhibits after removal of the cortex a depression formed by arches which pass from the gyrus glossiformis into the gyrus fusi- formis. In the anterior half of the same depression we find fibrce proprice connecting the uncinate with the fusiform convolution. In this instance the upper branch of the U-shaped bundles, de- scending from above, attains to a considerable length, for their fibres originate in the medullary substance of the cingulum. The fasciculus basalis internus of Burdach, which a process of lateral exfoliation has reduced to a narrow medullary layer, lies in the axis of the gyrus glossiformis (Fig. 18, bi.), tends toward the gyrus uncinatus, and seems to consist of naught but arched bundles. The medullary band of the gyrus fusiformis, the lower longitudi- nal fasciculus, so-called because it can be separated into long- extended arched bundles, contains fasciculi which take either a longitudinal or transverse direction, and which, with the various- sized fellow-bundles of the third temporal convolution, form the floor of the depression in which the inferior occipito-temporal convolution is bedded. Later on we shall see, too, that the Hnllnrv snr the convolution. 57 The Medullary substance of 58 Psychiatry. constitute in man but -fa-\ ; in smaller apes, \-\\ in the dog, \; in the cat, \\ in the bat, \; in the calf and deer, \ of the en- tire breadth of cortex. Their absolute breadth of 0.25 mm. in man is exceeded by 0.4 in the calf, and 0.5 mm. of this substance in the deer. This gray substance between the nervous elements appears under the microscope to be made up of densely crowded dark spots surrounded by tissue of a brighter color. It contains dis- tinct connective-tissue elements, with a network of fibrils from the processes of these elements, and a second network of fibres derived from the processes of the nerve-cells. I learned as much as early as 1870 from preparations hardened in bichromate of potash, stained with carmine, dehydrated, and rendered transparent by oil of cloves (Strieker's " Handbook of Histology "). Further details were recorded by Jastrowitz1 and Boll2 as a result of their methodi- cal researches, and it was shown to be a far more complicated structure. The following minute description of cortical gray substance applies to all gray sub- stance, wherever found, The cortical surface borders upon the pia mater. There is no epicortical lymphatic space, as was proved by careful injections 3 of the lymphatics of the pia mater. The gray substance has a sort of limiting membrane, consisting of connective-tissue cells with numerous processes, which are quite the same as the con- nective-tissue cells in the gray substance. Various influences conspire to affect the direction of these elements among the different tissue-structures, in consequence of which we find the processes from the outer cell-terminus spread flatly over the cortical surface, while the pointed processes starting from the other end of the cells, take a radiating course through the cortex. These processes, according to Lowe, are 0.125 mm. long and ramified, and are not unlike the cone-fibres of the retina. The outermost stratum of the cortex is, therefore, an epithelial arrangement of connective-tissue cells, whose individual form is best represented to the mind by the leaf of a palm-fan the stalk of which is at right angles to the leaf surface. Fleischl described as the outer- most cortical layer a fenestrated cuticula, which is rendered visible by staining the cortical surface with silver; this cuticula Boll considered to be an albuminous mem- brane in the valleys between the very delicate prominences on the cortex caused by the flat connective-tissue cells. Boll's "fusiform"4 cells are found, in lesser numbers and without the palm-fan shape, everywhere in the gray substance. These glia cells (neuro-glia), vary in shape (according to Jastrowitz), from spindle-shaped cells— resembling those of the tendons—to such as are surrounded by numerous fine (according to Boll) non-ramified processes. The size of the cells varies between 4-17 ^ (Jastrowitz). The body of the cell is at times considerably developed, and then again it disappears 1 Arch, fur Psychiatrie und Nervenkrankheiten, vol. II., p. 389 ; vol. III., p. 162, 1870-71. '* The same, vol. IV., p. 1, 1874. 3 Experiments of Boll and Golgi. 4 " Pinselzellen," " pencil-cells " would be more literal.—S. s fl = -j-gVff millimetre—S. The Cortex. 59 between the processes. From specimens obtained by teasing the preparations Boll described the protoplasm around the nucleus of the glia cells as a granular substance without well-defined boundaries, lying between the extremely delicate pencil-shaped fibres. The second form of non-nervous elements of the gray substance are the glia- nuclei, which are connected with the granular basement substance. They are more numerous in the newly-born and densest in the cortical layers. Their protoplasm can- not be separated from the granular basement substance. In the embryonic chick Boll could not observe that early stage in which distinct embryonic cells hold the ground occupied later on by the continuous basement substance, but he could notice that the granulated substance, consisting of a protoplasmic formation, uniformly studded with spherical dots, soon changes its appearance. This change is effected by a twig-like grouping of adherent granules, which is particularly distinct in the vicinity of the double-contoured nuclei, and converges slightly toward them. This resembles a formative activity of the cells. "On the strength of these investigations it would, perhaps, be fair to assume that in spite of the confluence of the cell-protoplasm, and the fusion of the cells proper, there exist virtually and physiologically separate cell units." The nuclei once densely crowded move far apart in consequence of the proliferation of the ground substance (increase of the granula—Boll). The granulated substance is absolutely independent of the nerve fiWrils, or of connective-tissue fibrils, but simply remains attached to these fibrils after teasing them (Jastrowitz, Boll). The granulated basement substance is highly albuminous. Boll explains this'by saying that the basement substance is developed out of primary embryonic connective-tissue cells, and that its quantity of albumen is similar to the granular albuminous material which is formed everywhere with the development of the connective tissue in and next to the nerve fibrils. This albuminous material is stored sparsely in some and liberally in other connective-tissue formations during the entire period of life. Jastrowitz ascribes to the granulated substance in the cortex the function of isolating nerve fibres, for in the cerebral medullary substance the axis-cylinders are separated from one another by the same white substance before the latter is transformed into the medullary sheath of the axis-cylinder. The views of Besser, Arndt, Rindfleisch, and Henle have been disproved by the investigations of Jastrowitz and Boll. As I stated in 1867, the nervous elements of the gray sub- stance, form regular concentric layers, the arrangement of these elements being modified according to the locality in which they occur. The commonest type of cortical stratification is found in all the convolutions of the convexity. A transverse section of the cor tex presents on the whole a uniformly gray appearance. In the middle of some of the broader convolutions we find a less grayish zone, due to loss of pigment. On transparent sections of these convolutions (magnified 100 diameters), we can distinguish five layers, the first being immediately beneath the pia mater. The first layer is made up chiefly of the basis substance and its connective-tissue elements. The latter are most numerous near the surface of the cortex. A few irregular, angular cells are seen scattered throughout this layer. This is the neuroglia layer. The second layer is sharply defined on its outer margin, and consists of 60 Psychiatry. densely crowded pyramidal-shaped bodies, which turn their apices toward the surface, and measure about 10 ju. in height. The internal boundary is also well marked, owing not so much to a change in the size of these nervous elements, as to the lesser den- sity of distribution. This is the layer of small pyramids. In the succeeding third layer, a distinct columnar arrangement of the nerve-cells, which are quite separate from one another in a trans- verse direction, may be noted. This is due largely to the aggre- gation of nerve fibres from the basilar aspect of the small pyramids. These nerve fibres increase inwardly, and push their way in between groups of pyramids. The pyramids, while in- creasing in calibre, attain to a height of 40 jj. or even 60 //. according to the width of the convolutions. In addition to the ramified apex-process, and from four to seven ramifying lateral base-processes, there is one median base-process (Fig. 24, lowest row of layer 3), which runs in a diametrically opposite direction, yet parallel to the apex-process, toward the medullary substance. The nuclei of the pyramids are miniature pyramids themselves, their angles often extending into the processes of the pyramids. This is the layer of large pyramids, or at least the layer which contains these. A sudden jump as regards calibre is made from these layers to the fourth, composed of multiform elements, chiefly of a rounded- angular nature. This is called the "granule-like" or granular formation. Between this layer and the medullary substance of the convolu- tion, we find a fifth layer not definitely bounded, and consisting near its outer margin of rather large but short pyramids. The further we proceed toward the white substance, the more spindle-shaped nerve cells of about 30 7«w. we find ; these spindle-shaped cells, sending processes toward the granule-layer, present the appearance of vertically compressed pyramids. But they never show any thing in the least akin to a median base-process. The medullary substance constitutes the sixth layer of these con- volutions. This layer contains a fair number of spindle-shaped nerve cells, taking the same direction which those of the fifth layer do. The axis-cylinder, as well as the medullary sheath, varies very much in size, from the exceedingly delicate to the dimensions of spinal-cord fibres. The central fibres lack the mem- brane of Schwann and Ranvier's internodia (Boll), which are char- acteristic features of the medullary substance in peripheral nerves. The Cortex. 61 The white substance is studded with granules (cubic cells, Boll), which in their arrangement (though normally interrupted in the adult brain) imitate the course which nerve fibres take. In the entire cortex there are but three forms of nerve corpuscles: (i) the pyramidal form, (2) the granule (mixed) form of small nerve cells, and (3) the spindle-shaped form. In the drawing (Fig. 24) the first formation is found in the second and third layers; the granules, in the fourth ; the spindle-shaped cells, in the fifth layer. In Fig. 25 we find the pyramids in the second layer; the granular elements in the second, fifth, and seventh layers ; the spindle-shaped bodies in the eighth layer. The pyrami- dal and spindle-shaped bodies differ from each other chiefly in regard to position. The longitudinal axis of the pyramidal bodies (parallel to each other) stands vertical to the cortical surface, while the longitudinal axis of the spindle-shaped cells lies parallel to the surface df the cortex. Throughout the central nervous organ a morphological law evidently operates, by reason of which the formative activity exercises an influence over the direction of the nerve cells, making the direction of their longitudinal axes parallel to that system of fibres which originates from them. Let us recall the various directions which the different systems of cortical fibres take, and which were demonstrated by the prepa- rations represented in Figs. 18 and 19. Obviously enough the direction of the pyramids (vertical to the cortical surface) is parallel to the projection-system fibres, and, on the other hand, the fibrae propriae (of the association-system), which do not conduct from or to the cortex, but from cortex to cortex, run parallel to the surface of the convolutions, just as in the above figures (24 and 25) the spindle-shaped cells lie parallel to that surface. Although the spindle-shaped cells are by no means bi-polar cells, and have un- doubted lateral processes as well, yet the latter run parallel to the surface, and, as they are turned away from the medullary substance of the convolution, can form no im- mediate connection with the projection-systems. We can obtain double proof of the parallelism existing be- tween the axes of spindle-shaped elements and the fibrae propriae by comparing a section of the cortex taken from the summit of a convolution with one (represented in Figs. 24 and 25) taken from the margin of a fissure. The fibrae propriae intersect'at the mar- gin of a sulcus with the projection-fibres (page 38), but at the apex of the convolution both systems of fibres run parallel to 62 Psychiatry. one another; just so the spindle-shaped bodies of the fifth cortical layer, bordering upon a fissure, intersect with the pyramids of the second and third layer, but at the summit of a convolution the former lie vertical (radial) to the surface of the cortex and parallel to the axes of the pyramids. Henle is wrong in stating that the pyramids are absent in the cortical substance bordering upon a fissure. The cortical nervous elements can be distinctly made out in the chick from the third day of development on, and very shortly after this the direction of their axes is apparent (Boll). In the human embryo both these nerve-cells and their processes show not later than the fourth month. At this period there is very little basis substance separating them ; as they are very dense at this early stage it is probable that they do not go on increasing in number. Their own growth and that of the basis substance seem to be part and parcel of the growth of the cortex. Besser, Arndt, and Henle have expressed different views on this subject. All cortical nerve corpuscles are composed of a sheathless pro- toplasmic substance, finely, but not always equally, granulated.1 According to Reinisch and Max Schulze, the protoplasm of the nerve cells has a striated appearance, which Boll has proved, in specimens stained with hyperosmic acid, to be common also to the pyramids of the cortex. The striations of the ganglionic processes have been familiar to us for some time. These pro- cesses are equivalent to the axis-cylinders of nerve fibres. Because of these minute fibrils, the ganglionic process and the axis- cylinder are supposed to be not morphological elements, but aggregations (fasciculi) of minute fibrils; these fibrils alone are present in the invertebrata, and for that reason are con- sidered the elementary nerve fibres. These processes and axis- cylinders emanate from the ganglion cells, in the protoplasm of which they appear to be scattered like the hairs of a brush ; they pass from one process to the other, or tend toward the nucleus, which seems to be surrounded by such fibrils. These fibrils cross each other, interlace closely with one another, and thus lose their individuality. Max Schulze was, perhaps, the first to suggest that a ganglionic cell might possibly, from the arrange- ment of its fibrils, represent a plexus, and that the protoplasm was simply traversed by afferent and efferent fibrils. 1 This granulation causes a pigmentation of the protoplasm, and distribution. The Cortex. 63 Other authors have endeavored to prove that the nerve fibrils do in reality terminate in the granular basis substance, where, according to Rindfleisch,1 the " fibrillary " and the " granular " structures are fused. Very soon afterward, Gerlach,2 after politely saying that he would confirm this view, contradicts it, by showing (on potassium and gold chloride preparations) that the ganglionic pro- cesses, after ramifying minutely, send terminal fibrils into a general network. Prior to this, Jastrowitz stoutly maintained that when- ever he detected fibrils covered by granular substance, the fibrillary elements extended beyond the granular. Boll declared that Rind- fleisch's hyperosmic-acid method was the very one best adapted to prove the independence of the finest fibrils within the granular substance. Those who attempt to divest the nerve cells of their importance as centres, and who trace the anatomical origin of nerves to a dif- fuse substance, such as the granular portion of the gray substance, render it impossible to establish a relation between the anatomical structure and physiological function of these parts. Strangely enough, the ganglion cell with its ramified processes would be in direct contrast to the spider in its web. The active agent would be, not in the spinning body but in the meshes of the web. Such a view of the anatomical structure of the nerve cell could un- doubtedly be entertained, but it would then be impossible to con- ceive of any physiological agent discriminating between two such masses as the ganglion cell and the intercellular substance. Even if we were for the moment to lay aside all justifiable doubts as to the striated appearance of the ganglionic bodies, the striae being de- ceptive, due perhaps to a mere folding, or to the changed tension of a body separated from its surroundings, to which its ramifications had attached it; discarding such doubts, I say, we can still find good anatomical reasons which force us to regard the nerve corpuscles of the cortex, the pyramids for example, as independent elements within the network of ganglion processes. 1. From the time of the existence of formative cells in the an- terior cerebral vesicle, the nerve cells are independent of the forma- tion of the basis substance, and, as we learn from comparative anatomy, the former is independent, throughout life, of the develop- ment of the latter; while, according to Boll, the fused pro- toplasm of the basis substance with its nuclei represents the remnants of other cells. 1 Centralblatt fur die medicinischen Wissenschaften, 1S72, page 77. ' Idem, 1S72, page 273. 64 Psychiatry. 2. The processes start in the ganglionic cell, and are originally nothing but a prolongation of the cell protoplasm, granular and non-striated, but undergoing later on numerous ramifications (Boll, /. c, " Entwicklung des Huhnchens " ). The cell processes perma- nently retain their protoplasmic character, not only in continuity with the protoplasm of the ganglion, but also after the interrup- tion of the granules at considerable distances from the ganglion, in the midst of the fibrillary striation (Max Schulze). From the very start the ganglion cell is an independent centre; the processes and their ramifications are portions of the ganglionic body. All their parts together form the protoplasmic individual. The striation would lie entirely within the latter, between one process and another. 3. The axis-cylinder develops out of special spindle-shaped cells in the medullary region (Boll), the protoplasm of which is con- tinued into the gray substance of the cortex, and there splits up into very delicate fibrils; on the hypothesis of Gerlach and Boll these most minute fibrils connect with the finest ramifications of the ganglionic processes. There would then be a secondary connec- tion merely between the processes of the latter and the largest aggregate of axis-cylinder fibrils. Ganglionic cells and axis-cylin- ders in their connections would be parts of two, not of one proto- plasmic body. This independence of the ganglionic cell as an anatomical centre does not admit of a doubt as to its importance as a functional centre. The composition of the axis-cylinder from the fibrillary elements of Schulze would not prove the isolation of these fibrils, for the axis-cylinder is not properly isolated until surrounded by a medul- lary sheath. Originally gray substance similar to the granulated basis substance of the cortex separates the axis-cylinders in the medullary layer from one another (Jastrowitz, Boll, Flechsig). From this Jastrowitz draws the correct inference that in default of a connection between this basis substance and the fibrillary elements (cf above) we must look upon the granular substance as the true isolating mass between the fibrils. Isolation is therefore carried further in the cortical than in the medullary substance, and yet the isolating mass retains its embryonic character, for the granulated basis substance does not become medullated. The anatomical structure of a functional centre demands that there should be no isolation in the centre itself. The striation of the ganglionic processes does no more prove the isolation of its The Cortex. 65 fibrils than it did in the case of the axis-cylinder. And then again the connection of the striated ganglionic processes with the proto- plasm of the cortical pyramids effects the realization of the transi- tion from the " fibrillar " to the " granular " where isolation ceases. Beyond this point the doctrine of cortical centres does not venture This is certainly not a novel point of view, but fortunately one that suffices for physiological purposes. We have no reason to grow sceptical on this point until we shall have succeeded in isolating Schulze's finest ganglionic fibrils from the surround- ing protoplasm, as has been done in the granulated basis sub- stance. The development of the axis-cylinder from the cortical nerve- elements, and the relation * of the former to the pyramids are as follows : The pyramids send median base processes (Meynert), which need not ramify (Koschewnikow), directly from their base into the medullary substance. Moreover, the pointed apex process extend- ing from the third layer to the most external layers of cortical ele- ments divides into a multitude of terminal ramifications, the finest spurs of.which return loop-shaped toward the white sub- stance (Boll) (?). These delicate filaments collect and form stouter filaments ; these in turn give rise to the myeline, which is, therefore, in the first instance, a product of the pointed proces- ses. Now this myeline is deposited next to those axis-cylinders which are formed by the median-base processes (Gerlach). The lateral processes could, in the same way, participate in the origin of the white substance. The gray substance, however, is traversed simply by nerve fibrils which enter into no combination with it. In regard to the grouping of ganglion cells, it is very evident that the dimensions of the pyramidal cells are directly proportion- ate'to the distance they are separated from the external cortical layers; that they are small externally, and steadily increase in size as far as the granular layer. The apex process splits up not only into a number of terminal ramifications, but gives off lateral branches nearer to the protoplasm, which (fibrils) take part in the formation of the so-called fibrillary network. Even Gerlach's preparations, which I have been permitted to examine, do not enable one to detect any other internodia but the ganglionic cells, so that the minute fibrils seem to form a compact 'According to Gerlach's staining method : potass, and gold chloride I; 10,000 water. 66 Psychiatry. mass rather than a net. The greater the cortical distance along which the apex process contributes branches to this network of fibres, the more fibrils this process will contain, and the stronger this process will necessarily be. Inasmuch as these ramified branches are continued in some way or other into the protoplasm of the pyramid, it naturally follows that stronger processes, or an increasing number of branches, will engender larger pyramids. I need scarcely add that the smaller pyramidal or other nerve cells, at a distance from the cortical surface ramify in the region im- mediately surrounding them—i. e., at a greater distance from the cortical surface. Stout and short, or long and thin, pointed processes do not occur. The external layer of pyramids, whose pointed processes soon enter the fibrillary net, is composed naturally enough of smaller elements. These pyramids are not wide apart either, for their lateral processes have but a narrow area over which to distribute their ramifications. This refutes the opinions of those who hold that the small pyramids are sen- sory, and the large pyramids motor in function ; it also disproves the theory of Wundt, that the large pyramids are the old, and the small pyramids the young cells. The pyramidal and spindle-shaped cells contain either oval, pyramidal, or spindle-shaped nuclei. The latter forms are found to be quite as numerous in brains that have never been hardened; for this reason Boll's view, that they are artificial products, seems untenable. These peculiar configurations of the nucleus seem to me to support the opinion of Beale, that the cell nucleus is sur- rounded by an optically denser protoplasm, which is differen- tiated from the external layers of protoplasm. The greater density of the nuclear portion of the protoplasm screens the oval nucleus, but it is penetrated by the glittering of the nucleolus. Owing to this dense protoplasmic layer the outlines of angular or pointed nerve corpuscles, together with their prolongations, seem crowded into the processes. No morbid change and no physiological experiment give us any reason to hope that we shall be able to explain the difference in form of cortical elements in closely neighboring layers. Morphological interpretation is the only method which can come to our rescue. The nerve corpuscles of the gray anterior horns of the spinal cord, of the central nuclei of the hypoglossal, facial, and abducens nerves, and as far up- wards as the oculomotomerve, all show long, slender cell-forms with numerous processes. These processes seem to arise with a broad base from the body of the cell. The same peculiarities of configuration which we observe in those nerve cells which are con- nected with centrifugal nerve tracts, are found in the cortical pyramids, and there can be explained only by the similarity in the distribution of these bodies. Gerlach has The Cortex. 67 compared the median base process with those spinal cell processes which enter the anterior roots. The granules of the fourth cortical layer, which are distinguished from the free nuclei by their size and protoplasm, from spider- and spindle-shaped cells by the distinct boundary of the protoplasm, and by a lesser number of stout processes, may be likened to those branched ganglionic cells which occur in centres connected with centripetally conducting tracts, as in the substantia gelatinosa of the fifth nerve and in the posterior horns of the spinal cord ; these granules maybe likened, also, to the inter- nal granular layer of the retina, and to the smaller elements of the granular layers of the olfactory lobe. The axis-cylinders of the fibres of the convolutions and of the entire fore-brain are developed at a very early embryonic stage. In the chick they develop from spindle-shaped cells on the fourth to sixth day after impregnation (Boll). Simultaneously with these elongated formative cells, chains of round elements appear which change into spider-cells, granules, and ganglion cells, which Boll supposes (perhaps correctly) to be diffused over the entire medullary substance of the hemispheres. Jastrowitz thought them erratic. These chains of connective-tissue elements, separating as growth continues, lie between bundles of fifty to sixty fibrils. The axis-cylinders are separated by a gray, granu- lar, albuminous basis substance, like that of the cortex. The fibres in the fore-brain do not develop medullary sheaths until later on, and for that reason are separated by gray substance only until long afterbirth (Jastrowitz),and in some parts up to the age of nine months (Flechsig). This gray substance may even appear darker than the cortex (Jastrowitz), though later on it becomes differentiated as the white substance. The transition of fun- damental gray substance into medullary sheaths is preceded by a fatty infiltration of the latter, and by the appearance of granular cells possessed of amoeboid movements (Boll); but later on their fat-granules disappear, and these cells are lodged between the medullary fibres as connective-tissue cells. Granular cells are present generally in the medullary substance from the fifth month of intra-uterine life to the fifth month post partum. A remnant of intrafibrillary gray substance is per- petuated. The star-shaped connective-tissue cells (Meynert), called also spider-cells (Jastrowitz) and fusiform-cells (Boll), form, after the complete development of white medullary substance, an interstitial fibrillary network of nerve fibres and bundles of the fore-brain, varying from fibres of minutest dimensions to fibres of respectable size. These cells seem to be fixed every- where to the medullary substance, while the free nuclei, present 68 Psychiatry. in varying numbers, might be classed as remnants of the gray medullary substance. The cortex of the outermost part of the occipital extremity, and of the calcarine fissure (to which the vertical line gives a H shape, and the occipital extremity a distinct wall), can be divided macro- scopically into three well-defined layers: (i) an external gray layer ; (2) a median, distinctly-limited white layer; and (3) an in- ner gray layer. Microscopically we can discern eight layers. The eight-layer cortex is distinguished from the four-layer cortex : I. By the reduction of the layer of pyramids to a narrow concentric layer (Fig. 25, 2), within which the size of the pyramids varies less than in the five-layer type. According to explanations given above, this is due to the lesser distance of the inner- most pyramidal strata from the neuroglia layer. 2. By three granular layers, which constitute the third, fifth, and seventh layers. The granules are curiously intermingled with small py- ramidal and small spindle-shaped cells. Owing to the narrow- ness of the medullary substance and the consequent greater parallelism of the stratum propriwn of this portion of the cortex, its eighth layer, answering to the fifth layer of the first type, gives us valuable information regarding the spindle-cells, which are larger than th6se spindle-cells to be found interspersed in other layers. 3. By the less densely packed elements of the third, fourth, and sixth layers, comprising chiefly small pyramids and transverse spindle-cells. Both these layers are fitly called intergranular layers. These intergranular layers contain at considerable distances apart either single pyramids of astounding size, or groups of two and three such pyramids. These are the largest ganglion cells to be met with in any part of the cere- brum. The pyramidal cells appear upon the inside of the granu- lar layer; their long pointed processes traverse the more external layers, and pass through the pyramids of the second intergranu- lar layer—through two granular strata—through an intergranular layer and a portion of the pyramidal layer. The gigantic dimen- sions of these solitary pyramids can be explained on the principles before alluded to in regard to the progressive increase in the size of these pyramids. The presence of large numbers of small pyramids, especially in the second and third granular layers, as well as in the layer of spindle-cells, would lead us to believe that these pointed processes and their ramifications terminate in that part of the gray network of fibres which does not belong to The Sulcus Calcarinus. 69 the most external layers of the cortex. The three granular and the intergranular layers con- sist really of mixed elementary forms, among which the gran- ular ganglionic cells prepond- erate. The connection of the pyramids with the previously developed delicate radial med- ullary bundles of the cortex is easily recognized. The small fusiform (spindle) cells of these layers are contiguous beyond a doubt with transverse nerve fibres, which are by no means numerous enough, however, to constitute, as Kolliker would have it, the white substance of the intermediate strand in the cortex of the sulcus calcarinus. To explain satisfactorily this white, intermediate strand, we must, because of its well-defined boundary, have recourse to the lack of pigment in the barren intergranular layers, for the nerve corpuscles evidently carry the pigment which is respon- sible for the deep tint of the gray substance. Besides, the effect of lack of pigment is intensified by radiating medul- lary bundles which are not prom- inent in other internal layers —in the seventh and eighth for instance,—where the impression gained from numerous pigment cells overshadows all else. 7o Psychiatry. Similar circumstances conspire to bring about a diffuse, paler tint of the middle portion of a section of the cortex taken from the central convolutions. Betz has stated that the anterior central convolution contained groups of particularly large pyramids, which he thought were the circumscript motor centres which Hitzig, on the strength ©f his physiological experiments, relegated exclusively to the anterior central convolution of the brain in dogs and monkeys. Apart from the mistake Ilitzig made, and which we corrected above, in establishing the homologon in carnivora of Ihe an- terior central convolution, it has been proved that the size of the pyramids depends upon their distance from the cortical surface. The largest pyramids will, therefore, be found in the broadest cortical region ; but the broadest cortex is that of both central convolutions. The third layer, the equivalent of the third layer in Fig. 24, is very broad, and contains larger pyramids, which are naturally farther apart than their fel- lows in the dense groups of small pyramids. For this reason we observe an increasing lack of pigment near the surface of the cortex, due to a gradual diminution in the size of the pyramids. And on the other hand, these pyramids do not move asunder by reason of their size alone, but their regular distribution is interfered with by the steady increase inward in the circumferences of radiating bundles of fibres, which at the same time intensify the paler tint of the third layer. The largest pyramids appear, therefore, to be arranged in small groups at some distance from one another. It would be wrong to argue from this that these large pyramids have a different signification from the smaller ones. Luys is in a great measure responsible for this mistake. Betz appears to me not to have made a discovery, but to have failed to appreciate the gen- eral relations which obtain in the primary disposition of the cortical elements. The gyrus uncinatus and the cornu ammonis, which represents a continuous involution of the former's cortex, are peculiarly con- structed, and show but a meagre variety of cortical nerve-forms. The cornu ammonis contains principally cortical elements of the pyramid type. Essentially the same is true of the lobi olfactorii, joined to the gyrus fornicatus by frontal and temporal connec- tions (olfactory convolutions). The cap of the olfactory lobe, the bulbus olfactorius (Fig. 10, Olf.) exhibits a special kind of cor- tical stratification. Nearest to the surface lies the origin of the olfactory nerve, which passes through the lamina cribrosa to the Schneiderian membrane. These nerves take their origin in glomeruli which contain capillary loops (Schlingen) and small nerve cells (Meynert). The glomeruli olfactorii are transformed in mammals (Clarke), and in the lower vertebrates (Leydig), with the aid of gray sub- stance, into round masses (stratum glomerulosum), between which granular nerve cells are collected and heaped up. 1. This dis- tention of the glomeruli in animals is a single instance of the proliferation, independent of the number of nerve fibres and cells, of amorphous connective-tissue substance in animals, in contra- distinction to its reduction in the gray substance of man (page 56). 2. Toward the inner side of the stratum glomerulosum the The Claustrum and Amygdala. 71 olfactory lobe contains the gelatinous (connective-tissue) layer of Clarke, which, like the cerebral cortex, possesses on the outside small nerve cells, which are changed into larger, angular, elongated nerve cells as we proceed inward. 3. Closely packed granular layers succeed the innermost largest nerve cells as they do in the cerebral cortex; these granules are divided into strata by the medullated substance parallel to the surface of the olfactory lobe. Animals whose olfactory lobe is hollowed out by a diverticulum from the lateral ventricle exhibit an ependymal layer with columnar, ciliated endothelium. 4. Through complications a fourth type of cortical substance is created, represented in the cortex surrounding the fossa Sylvii, where a gray lamina, the claustrum, runs parallel to the cortex of the island of Reil (Fig. 6, CI.), of the operculum (Fig. 5, between J and L), of the first temporal lobe, and, continuing beyond the anterior fissure, parallel to the cortex of the orbital surface (Fig. 26, CL.). The claustrum is developed out of the formation Sagittal Section through the Brain of Hamadryas, near the Island of Reil. Fr. Tm. Occ. Frontal, temporal ends and occipital region of the fore-brain. I. Island (should have been placed over the entrance to the Sylvian fossa), op. Opercu- lum, j. Upper margin of the island. CL. Claustrum. A. Amygdala. Cd. Anterior commissure. Am. Cornu ammonis. Ci. Inferior horn of the lateral ventricle. Cp. Posterior horn of the lateral ventricle. NL. Lenticular nucleus. NC. Temporal end of the caudate nucleus, nc. Hindermost portion of this nucleus. Between the two the stria cornea. Ce. Medullary capsule of the external geniculate body. of spindle-shaped cells of the fifth cortical layer; the elements of this layer and of the claustrum resemble each other, not only in size and shape, but also in being parallel to the surface of the cortex. The claustrum, which lines the inner surface of the island, adapts itself to the fan-shaped form of the latter, the handle of the fan pointing downward (basis insulae). 72 Psychiatry. The margin of this trilateral fan is bent outward on all three sides. The claustrum is continued forward beyond the anterior fissure into the posterior margin of the orbital surface of the frontal lobe, upward into the operculum, and below, but to a lesser extent, into the first temporal convolution. This fan- shaped formation is furrowed or rather folded to correspond to the rise and fall of the island-convolutions. The pedicle (handle) of this fan-shaped formation, a ball-shaped body—the amygdala, —represents at the same time the most anterior portion of the uncus lying to the inner side of, and behind, the base of the island. The amygdala (Figs. 6, 26, 30, 32, A.) is composed of the same elements as the claustrum. Between these two bodies (claustrum and amygdala) there are small transitional heaps or strips of gray substance, which cannot be definitely classed as belonging either to the claustrum or to the amygdala, but together with these two formations constitute a continuous system of gray substance (Fig. 26, CL., A.). Amygdala and claustrum are separated from each other by the anterior commissure in those frontal (transverse vertical) sections of the brain, on which the lateral portion of the anterior commis- sure, descending to the temporal lobe, is seen to pass between them (Fig. 6, Ca., CL, A.). The claustrum is a formation of gray substance. The nerve cells of this formation are present also in the white substance of the island, as well as in the external capsule between nucleus lenticularis and claustrum, and in all these structures these cells have their axes turned in the same direction. I wish to call attention again to the connection existing between the claustral investment of the Sylvian fissure, and the association- bundles of that region. (Compare Fig. 19 with Fig. 26.) The claustrum is not to be ranked among the ganglia, as it communicates with the fibrae propriae of the cortex. The cere- bral ganglia are connected with the cortical projection-bundles only. Before passing to the study of sections of the cerebral ganglia and the projection-system I must stop to refer again to the med- ullary formation of the anterior commissure. Like the corpus callosum this commissure represents a system of fibrae proprise con- necting both hemispheres,and uniting with those commissural fibres which join both olfactory lobes, themselves parts of the hemispheres. According to the very general account given of it in Fig. 20, the anterior commissure consists apparently of a round (on section, Commissura Anterior. 73 oblong) bundle of fibres, spreading out principally in the tem- poral and occipital lobes. But a more thorough examination of the anterior commissure (on sections) shows that a not incon- siderable number of its bundles originate in the olfactory lobe, and pass upward from the posterior margin of the trigonum olfac- Fig. 27. Transparent Frontal Section through the Cerebral Ganglia of Man. Anterior Commissure. J Island of Reil and its medullary substance. CI. Claustrum. Ce. External capsule. L.1, L.2 First and second divisions of the lenticular nucleus. Nc, Nc. Ventricular and basilar portions of the caudate nucleus. Ca. Anterior commissure. Olf. Olfactory bundles passing to anterior commissure. Orb. Orbital surface. torium through the substance of the lamina perforata anterior. And yet these bundles are not sufficient to account for the size of the anterior commissure (Fig. 27, Ca., Olf.). We must, therefore, in- fer that the anterior commissure contains bundles also which con- nect the occipital and temporal lobes, but stand in no sort of relation to the olfactory lobe. In man and animals this contribution to the 74 Psychiatry. anterior commissure from the olfactory lobe comprises most of the medullary substance of these lobes. The relative position of the bundles entering the commissura anterior from the hemispheres and from the' olfactory lobes is such, that the bundles coming from the latter join the commissure below, and those from the hemispheres form an acute angle in the frontal plane with the Fig. 28. Frontal Transverse Section through the Brain of a Dog. Anterior Commissure. R. Cerebral cortex. F. Gyrus fornicatus. M. Medullary substance of the fore-brain. Sp. Septum lucidum. Ye. Cerebral ventricle. Nc. Nucleus cau- datus. L.1 Cortex of the island of Reil. Mi. Medullary substance of the island of Reil. Ce. External capsule. L. Lenticular nucleus. CI. Claustrum. Ca. Fibres from the hemispheres to the anterior commissure. Olf. Olfactory division of the anterior commissure. upper portion of the commissure. Examined from this point of view, we observe that in the brain of a dog the superior—the hemispherical—portion of the anterior commissure (Fig. 28) 'The outer lower L on the figure should be J. (Engraver's mistake.)—S. Ganglia of the Brain. 75 is smaller than the inferior portion coming from the olfac- tory lobe. Consequently, the hemispherical bundles of the an- terior commissure, in animals having large olfactory lobes, would by no means cover in amount the wealth of com- missural fibres. For that reason the bundles of the anterior commissure must serve to unite both olfactory lobes. Further- more, it is apparent to the naked eye that fasciculi of the anterior commissure cross each other after the fashion of a twisted rope, and that these bundles are asymmetrically distributed in both halves of the anterior commissure. The latter undoubtedly possesses decussating bundles which connect each olfactory lobe with the temporal and occipital lobe of the opposite side. It is equally certain, however, and this we learn from longitudinal sections, that the entire medullary substance of the olfactory lobe does not pass into the anterior commissure, but that a part of this medullary substance forms into bundles which lie beneath the commissure and enter the basilar portion of the nucleus caudatus, i. e., the mass between the lamina perforata anterior and the commissura anterior. The course of these olfactory bundles within and outside of the anterior commissure establishes a forma- tion similar to the optic chiasma as interpreted by Joh. Miiller. As early as 1S61 Clarke described the cerebral cortex, and more perticularly the eight-layer type of the occipital lobe. Clarke differed from us in enumerating but six layers; for he does not separate the pyramidal, granular, and spindle-shaped cells at their proper boundaries. Clarke found that the cortical structure varied very much as regards the width of the several layers, and described varieties of type which may be found in one and the same convolution. Careful study of the structure of all convolutions will not permit me to establish any other types but those I have described above. GANGLIA OF THE FORE- AND INTER-BRAIN. The ganglionic masses of the brain, which are traversed by a large proportion of the projection-system, must be regarded as brain-forms (Organbilder), the contours of which are determined in the brain by the course of the projection-fibres. This is exhibited very clearly on gold-preparations in which the gray matter is not stained. THE GANGLIA .OF THE FORE-BRAIN (PROSENCEPHALON). the caudate and lenticular nuclei, really form but one mass, the continuity of which is interrupted here and there by the projec- tion-system. The projection-fibres surround the lenticular nucleus in the form of the internal and external capsule. 76 Psychiatry. In regard to the confluence of these masses we must note: I. That in the frontal lobe the large ganglionic masses of the fore-brain are divided into layers by the medullary lamellae from the internal capsule ; the more anteriorly they are situated, the broader will be the bridges of gray substance in this fore-brain ganglion between the successive layers of white fibres. Thus Gratiolet distinguishes between the corpus striatum intraventricu- lare (nucl. caudatus) and corpus striatum extraventriculare (nucl. lenticularis). Forward of the inter-brain {diencephalon) the inter- nal capsule lies solely between these two ganglia (Fig. 5, Cpi.; Fig. 27, Ci.). The sections of the projection-bundles belonging to these mas- ses contain not only fibres passing to the fore-brain ganglion, but also radiating fibres which travel from the frontal cortex to the anterior pedicle of the thalamus. (Compare Fig. 5 Cpi. with Fig. 21 f. Th.). In the diencephalon the corona radiata sends forth, in addition to the inner and external capsule, an oblique group of bundles lying one behind the other and collectively passing under the caudate nucleus to the stratum zonale of the optic thalamus. In the region of the inter-brain the caudate nucleus lies no longer to the inside of the internal capsule and next the lenticular nucleus, but on the former and above the latter (Figs. 6, 29—Nc., Th„ Ci.). 2. On longitudinal sections [Fig. 30, Pi. (n. c.)] the internal capsule exhibits a number of spindle-shaped masses of gray sub- stance, which connect the caudate nucleus with the more exteri- orly located portions of the lenticular nucleus. 3. On the upper wall of the inferior horn in the temporal lobe there is a complete fusion of the temporal extensions of both these ganglia (Fig. 26). In the direction of the occipital lobe the caudate and lenticular nuclei are separated by medullated bundles which pass from the white substance of the hemispheres to the capsule of the external geniculate body (Fig 26, NL., NC, Ce.). The nucleus caudatus constitutes, in reality, the internal margin of the lenticular nucleus, or, better still, of its lateral gray division. The lenticular nucleus is a wedge-shaped body ; its lateral di- vision, but imperfectly separated from the nucleus caudatus by medullary substance, forms the base of this wedge, looking for- ward and outward (Figs. 5, 6, 19, 23, 26, 30, 32, 34, 36—NL). The nucleus caudatus bears a certain definite quantitative relation to the lenticular nucleus : anteriorly, where the surface of the lentic- Nucleus Caudatus. 77 ular wedge is broadest, the nucleus caudatus develops its club- shaped head, while it diminishes posteriorly, and finally dwindles down into the cauda, just as the lateral division of the nucleus lenticularis is continued by means of a narrow, defective (notched) band to its equally meagre temporal continuation. As the arch-shaped portion of the fore-brain ganglion, the caudate nucleus surrounds with its head the lenticular nucleus as far as the basilar surface, and does by no means originate in the cerebral ventricle. By curving spirally about that part of the internal capsule which contains the anterior pedicle of the optic thalamus, the nucleus caudatus makes its way from the base of lenticularis (Fig. 28, Nc. L.). We have said that as far as minute structure and conn 78 Psychiatry. are concerned, these ganglionic masses must be regarded as one ; and yet quantitatively these two ganglia, caudate and lenticular nuclei, are entirely independent of one another. Man possesses the largest lenticular nucleus, but his corpus striatum (caudate nucleus) is not proportionately larger than that of mammals. A similar difference is to be noted between the brains of the monkey and the dog. Frontal sections exhibit the greater development of the len- ticular nucleus in man and the monkey, whereas the ganglion in the dog is inferior (quantitatively) to the caudate nucleus with its head extending around the internal capsule (compare Figs. 6 and 27 with Fig. 28). The arciform shape of the nucleus caudatus is the inevi- table result of the similar shape of the hemispherical arch around the fossa Sylvii. This arch sends out as its corona radiata an arch- shaped group of projection fibres, which, as they enter the fore- brain ganglion, separate the upper portion—the caudate nucleus— from the rest of the hemispheric ganglion (Fig. 21). The lenticu- lar nucleus is clearly a wedge, the longest axis of which forms an acute angle, open anteriorly, with the longitudinal axis of the hemispheres. The base of this wedge-shaped body lies to the outside (and to the front), its sharp edge (Schneide) toward the median surface; for more bundles enter it from the cerebrum than leave its median portion to pass into the crus cerebri. The fasciculi of the corona radiata diminish in number as they pass through the lenticular nucleus, on their way to the crus. The wedge occupies an oblique position, and for that reason presents the same appearance on longitudinal as on transverse sections. The sharp edge is turned chiefly to the back, the base to the front. This wedge-shape, which has been accounted for above, is complicated by the creation of three or four concentric divisions. In the inner divisions, the medullary bundles passing through this ganglion are more closely packed ; first, because there is less gray substance, and secondly, because not only their own bundles, but bundles also from the outer divisions, pass through these segments. These internal members of the lenticular nucleus seem, therefore, paler and more medullated (globipallidi) than the external division in which the gray substance preponderates. The vertical medullary partitions between the division of this ganglion are termed the lamince medullares nuclei lenticularis. The greater development anteriorly of the gray substance in Nucleus Lenticularis. 79 the nucleus lenticularis and its satellite, the nucleus caudatus, pro- vides a greater number of ganglionic cells to effect the connection with the superior member of the projection-system. For this reason the fibres of the corona radiata entering the posterior mem- bers must be far less in number; and the paucity of fibres will be more noticeable among those coming from the occipital than among those from the temporal lobes. The smooth anterior and exter- nal surface of the lenticular nucleus does not receive any fibres from the cortex, for but few bundles of the external capsule touch Fig. 30. Sagittal Longitudinal Section through the Brain of Hamadryas. Fr. Tm. Occ. As in previous sections. CI. Claustrum. A. Amygdala. Am. Cornu ammonis. S. calc. Calcarine fissure. D1F. Fossa Sylvii. cp. Posterior horn of the ventricle ; above it the caudate nucleus, behind it the greater commissure, ci. Inferior horn. ca. Anterior commissure, (n.c.) Caudate nucleus, as islands of gray substance in the corona radiata. nl. Lenticular nucleus. Pulv. Optic thalamus, gi. Internal geniculate body. ge. External geniculate body. II. Optic tract. Pi. PJ. Projection-bundles from the cortex. Pd. Pes pedunculi. Tg. Region of teg- mentum (index line passes behind and above it), brs. Brachium of the superior bigeminal body. (In the wood-cut the semicircular bundles of the latter are not dis- tinctly defined, nor is the index line properly placed.) the lenticular nucleus superficially and cross it in a direction parallel to its outer surface. The finely ramifying radiatingbundles of the first division, which collect into larger bundles, are dis- tinguished from fibres of the corona radiata by the fact that the latter are invariably largest nearest to the cortex, and do not increase in the opposite direction toward the crus cerebri. It can be demonstrated from sections in any direction that the lenticular nucleus receives its cortical fibres—the upper division of the pro- jection-system—on its upper surface facing the internal capsule. The cortical radiations of the capsula interna take in part a 8o Psychiatry. radial (?) course, and in part they pass through the laminae medullares, which are richly endowed with ganglionic cells. That the radial fibres are fibres of the corona radiata can be stated positively only of the upper layers of the lenticular nucleus. The nearer we get to the basilar surface, the more probable it is that these radiating fibres issue from the ganglion itself. It is quite easy to see that the radiating fibres of the globus pallidus are connected with the cortex through the laminae medullares, which pass vertically through the lenticular nucleus to the base, and that the radiations originate in the ganglionic cells of the cortex. The origin of the radial, branching bundles of the external layer of the lenticular nucleus is much more mysterious. I be- lieve that these fibres also originate in the first (outermost) lamina medullaris, and run in an opposite direction toward the external capsule ; that, moreover, after they have joined the gray substance of the lenticular nucleus near the external capsule, they turn about and are thus continued, together with other radiating white fibres of the globus pallidus, into the crus (Figs. 30, 32, 34, 36). In regard to the nucletis caudatus we must take notice, apart from its connection with the temporal lobe through the stria cornea, of the return of cortical fibres of the internal capsule, after an interruption through the cells of the ganglion, into bundles of the crus. There are no radial cortical fasciculi of the nucleus caudatus, but we find branching medullary fibres near the surface of the nucleus caudatus similar to those that pass through the internal capsule into the outer gray member of the nucleus lenticularis. The fibres of the second section of the projection-system, which originate centrally in the lenticular nucleus, and pass peripherally below the fore-brain into the crus cerebri, and indirectly into the anterior nerve-roots, and the equivalent cerebral nerve-roots, have a double origin in this ganglion. These fibres take (a) a vertical and (b) a transverse course. (a) From the middle divisions of the lenticular nucleus, the uppermost layer, probably (?) the continuation of those radial fibres of the external capsule which pass through the lenticular nucleus without the intervention of the lamince medullares, enters the pes pedunculi in front of those fibres of the crus cerebri which de- scend directly from the cortex (Fig. 33, 4). In the pes pedunculi, however, said fibres do not retain their anterior, superficial posi- tion, but cross those cortical bundles of the pes pedunculi which take a vertical course into the pons, in such a manner as to extend from the base backward into the stratum intermedium of Nucleus Lenticularis. 81 the crus cerebri. The stratum intermedium of the crus includes the substance of Soemmering. [Fig. 36 exhibits the continua- tion of the bundles of the lenticular nucleus, marked 4, Fig. 33. These bundles are those extending to JS below the chiasma (II.) and crossing the general run of fibres in the crus cerebri.] (b) The transverse bundles of the crus cerebri issuing from the lenticular nucleus run, in part, parallel to the smooth basilar sur- face of that ganglion, and constitute the ansa nuclei lenticularis (Fig. 23, left side). The departure of this ansa from the medullated ganglionic meshes of the lamina' medullares is well shown on allci frontal sections of the nucleus lenticularis; winding around the base of the internal capsule, the formation pushes its way from the inferior surface of the lenticular nucleus upward, forming on its course the innermost bundles of the crus. The base of the internal capsule (its anterior layers) passes into the base—the pes—of the crus cerebri. The fact that the bundles of the ansa nuclei lenticu- laris run upward, or more properly speaking backward, can be best and easily demonstrated by the through the red nucleus (NR)." fl.p J . . T Posterior longitudinal bundles. SS Cleavage method (Abfaserung). In stratum intermedium with Soemmer this way we Can discern laterally ing's substance ans. 1. Ansa lenticularis. a_y j jT Qptlc tract g 0 b Basilar optic the exit of the ansa lent, from the ganglion. Z. Stratum zonale of optic u *. f^Q1Qnt^„1nmn thalamus (cut across obliquely at lower gray substance of the lenticular nu- z ^ st>. ^tilus imemus "Jin Jrnal pedi. cleus; in a median direction the cle) of optic thalamus. Ca. Anterior 11 /• .1 *„4-„ commissure. development of the ansa into the innermost fasciculi of the crus, which surround the bundles of the pes, and pass behind the pes into the stratum intermedium (Figs. 6, 23, 31—ans. lent., SS.). In addition to the superficial stratum of transverse peduncular fibres of the lenticular nucleus, we find also a deeper system of peduncular fibres which, coming in successive layers from the laminae medullares, interlace as trans- verse laminae with the vertical laminae of the internal capsule (Fig. 36, above II.). The immediate cortical laminae (Rinden- f 1- p. g.o.b. Fig. 31- Sagittal Section through the Human Brain. Q. Superior bigeminal body. Cp. Posterior commissure, f. rfx. Fasciculus retroflexus. rfx. The same, passing 1. p. SS. 82 Psychiatry. blatter) of the pes pedunculi yield this area of interlacing fibres. The necessity of having these fibres interlaced is explained by the fact that the lenticular nucleus lies (in longitudinal sections) in front of, and the stratum intermedium of the cerebral peduncle behind, the cortical bundles of the internal capsule (Fig. 36). The tractus opticus constitutes the ideal boundary between the internal capsule and the crus. The pes pedunculi is made up of vertical laminae, which interlace at the nucleus lenticularis and continue throughout its length (Fig. 34). The medullary substance of the lenticular nucleus separates (probably not completely) from the crus cerebri above its entrance into the pons, dividing into laminae and crossing the crus posteriorly in the direction of Soem- mering's substance. In the stratum intermedium the medullary substance of the lenticular nucleus takes a downward course into the posterior division of the pons. The fasciculi of the pedun- cular stratum intermedium consequently increase on their down- ward course. In the upper half of the crus cerebri they are mere scattered bundles; in the lower half, however, they form compact transverse masses (Figs. 40 and 41). The stratum intermedium of the crus cerebri with which the prosencephalic ganglion is connected, extends below the thalamus, which is separated from this stratum by the lower half of the discus lent if or mis (Figs. 36 and 55). A marked increase in the thickness of the stratum intermedium does not take place until the mid-brain is reached. Consequently we may argue that the majority of the crus fibres from the lenticular nucleus enter the stratum intermedium below the discus lentiformis (Fig. 55, ss.). On this head consult pp. 51 and 53. CAPSULA INTERNA. Before giving a description of the diencephalic ganglion we must stop to glance at the capsula interna of the lenticular nucleus—-that complicated collection of fibres from the different divisions of the projection-system which separate the ganglion of the fore-brain from the thalamus opticus—the ganglion of the inter-brain. Cephalad the internal capsule connects with a large portion of the corona radiata, caudad it is continuous with the crus cerebri. The extent of the internal capsule cephalad is that of the ganglia, caudad it is bounded by the optic tract. The minute structure of the internal capsule can be studied best from sec- tions of a child's brain, which have been immersed in a solution of Capsula Interna. 83 palladium chloride 1 :1,000, before they are stained with potassium and gold chloride I : 10,000. Such preparations should present com- plete longitudinal sections of all the ganglia, and in addition they should be either sagittal or oblique toward the fore and outer part. The internal capsule is a conglomeration of medullated fibres from the hemisphere and the crura cerebri, and also of fibres from the cerebellum. We can discern five different orders of fibres in the internal capsule. 1. A portion of the internal capsule passes from the cortex to Sagittal Longitudinal Section through Brain of Hamadryas Passing through a Plane between that of Fig. 26 and of Fig. 30. Fr. Tm. Occ. As in previous sections. A. Amygdala, above it the claustrum. ri. Inferior horn of the ventricle. Cp. Posterior horn of the ventricle. NL. Lenticu- lar nucleus ; under its temporal prolongation, the caudate nucleus ; below this, the optic tract. (The line from II is incorrectly drawn.) Ge. External geniculate body. Am. Cornu ammonis. (The index line is too short.) Pulv. Pulvinar. brs. Ought to point to the foremost semicircle of bundles in the optic thalamus (the brachium corp. bigem. superius). S. cal. Should point to the calcarine fissure. Pd. Bundles of the crus from the internal capsule. Pj. Projection bundles, co. Anterior commissure. the lenticular nucleus (Fig. 5, 6, 30, 32). These fasciculi course along the superior surface of this nucleus, and on entering it, split up its superior margin. They do not enter the external segment of this nucleus in a body, but insinuate themselves in part (?) as lamiiuc medullares between the various divisions of the nucleus lenticu- laris (Fig. 34, 1). 2. Another set of fasciculi from the frontal cortex pass through the white substance of the hemisphere and finally 84 Psychiatry. enter the optic thalamus, constituting a seriatim defoliated mass of white substance along the inferior surface of the thalamus. The medullated lines seen on longitudinal sections are the edges of the medullary laminae succeeding one another from above downward, and alternating layer by layer with gray substance. This formation is the anterior pedicle of the thalamus, stilus ante- rior thalami optici (Figs. 5, 29, 33, 36, 55). From this group some fasciculi are contributed also to the stratum zonale. 3. Other fasciculi appear to connect the nucleus caudatus with the crus cerebri (Fig. 29). These bundles intersect with those of Fig- 33- Sagittal Longitudinal Section through the Brain of a New-Born Infant. (Preparation Stained with Potassium and Gold Chloride.) Th. Optic thalamus. C. gen. i. Internal geniculate body. n. 1. Caudate nucleus, n. L. Lenticular nucleus. A. Fasciculus Arnoldi. P. P. Pes pedunculi. 2. Bundles from the stilus anterior of the thalamus. 4. Cortical fasciculi and direct lenticular fasciculi. 5. Fasciculi from the internal capsule to the nucleus ruber. L. ltf. Should point to the black lenticular plate in the angle between 4 and 5. the stilus anterior thai, opt., lying to the outside of the main mass of the latter formation, and covering the inner portion of the lenticular nucleus. 4. The pes pedunculi contains descending medullated fibres coming from two masses of gray substance. The anterior of these laminated medullary masses issues from the nucleus lenticu- laris; the posterior and stronger mass, from the medullary sub- stance of the hemispheres—i. e., from the cortex. The latter might fitly be termed the " anterior cortical laminae " of the crus (laminae corticales antcriorcs pedunculi). If we choose to speak of Capsula Interna. 85 a single bundle of fibres in this connection, it should receive the name fasciculus of Arnold (Fig. 33, 4). The posterior cortical lamina? of the crus, the bundles of Tiirck, have been exhib- ited in Fig. 22 (Tm.), and Fig. 5 (Om., P.). 5. The fifth and most posterior medullary division (of the internal capsule), the basilar portion of which is covered by the laminae of the fasciculus of Arnold and by the discus lentiformis, does not merge into the pes pedunculi, but passes over this on its way to the red nucleus of the tegmentum. This bundle of fibres, as seen in longitudinal sections (Figs. 33 and 36, 5), repre- Transparent Vertical Cross-Section through the Human Brain, Including the Optic Thalamus. Gk. Stratum zonale adjoining the habenula conarii. P. g. h. Fasciculus retro- flexus, or peduncle of the habenula. 2. Bundles from the corona radiata to the thalamus. Between 2 and P.g.h., the semicircular boundary of the thalamic cone, lamina viedul- laris. N.c. Caudate nucleus. Cr. Corona radiata. Ci. Internal capsule. 1. Radi- ating bundles entering the lenticular nucleus (N.l.) To the right of the lenticular nucleus bundles from the corona radiata pass into the external capsule. N.R. Red nucleus with lateral radiating fibres from internal capsule. C. lent. Lenticular body. S.S. Median portion of the substantia nigra. Pp. Pes pedunculi. In it ascending bundles are visible, which bend downward into the stratum intermedium. sents the margin of a widely spread fan-shaped formation (Fig. 23). On cross-sections these fan-shaped bundles are also distinctly con- nected with the nucleus ruber (Fig. 34, to the outside of NR.). After leaving the cortex these fasciculi become entangled in the complexities of the corona radiata. The discus lentiformis is connected with a radiating arch of cortical fasciculi, which are in turn interwoven with the rest of the internal capsule. These fasciculi assert their individuality 86 Psychiatry. near their point of insertion, where they issue singly from this confusion of bundles. An innermost bundle of the internal cap- sule (Forel) is part of this radiating formation. Cordward, exter- nal fasciculi leave the discus lentiformis, and pass through the tegmentum into the brachium corporis bigemini inferius. In accordance with previous statements of mine, the pes pedunculi is traversed on the inner three quarters of its area by bundles destined only for the substantia nigra of Soemmering (Fig. 34). Its external quarter alone, into which the fasciculi of Tiirck are continued, contains bundles of fibres which pass from the upper surface of the discus lentiformis into the tractus opticus (fasciculi nervi optici of Stilling). The discus lentiformis is an isolated ganglion which communi- cates neither with the red nucleus of the tegmentum, nor with the substantia nigra of Soemmering. All the projection-bundles from the occipital lobe are part of the corona radiata, and not of the internal capsule. Among these bundles there are (1) those which lie to the outer side on the crus cerebri, and which I was the first to describe as bundles of Tiirck; (2) those which adjoin the former toward the inside, and pass through the brachium of the inferior bigeminal body into that body itself ; (3) those which enter the internal geniculate body ; (4) those which enter the external geniculate body ; (5) those destined for the superior bigeminal body; (6, and lastly) those which enter the pulvinar of the thalamus (Fig. 5, Om. p., br. s.y ge. i., ge. a., br. i., pulv.). OPTIC THALAMUS. The shape of the optic thalamus, and the medullary mark- ings within it, are influenced (1) by the medullary substance of the hemispheres ; (2) by fasciculi of the optic tract ; (3) by the origin of the tegmentum of the crus cerebri in the substance of the optic thalamus. Medullary layers make a well-defined ganglion of the optic thalamus. Its most distinct connection is with the tegmentum by means of direct medullary fibres, which thus make the optic thalamus one of the ganglia from which the spinal cord recruits its fibres. Its surface, facing the lateral ventricle, is covered by the stratum zonale, which extends from the stria cornea to the inner margin of the habenula—to the middle ventricle. Beneath the nucleus caudatus bundles enter the stratum zonale (1) from Thalamus Opticus. 87 the frontal lobe as superficial layers of the anterior pedicle (Figs. 5, 21, 29, 54, 55) ; (2) from the parietal lobe (Figs. 6, 34); (3) from the temporal and occipital lobes (Fig. 21); (4) from the retina, passing to the inside of the posterior temporal fasciculi through the tractus opticus; (5) from the ansa peduncularis along the anterior and inner edge of the stratum zonale (Figs. 21, 31) ; (6) from the ascending pillar of the fornix. These fasciculi of varied origin, covering the thalamus super- ficially before entering its depths, intersect at various points. They contain ganglion cells (Boll) before they are merged with the sub- stance of the ganglion proper. The outlying temporal fasciculi pass over the occipital and parietal fasciculi from behind, and cover them as far as the outer margin of the thalamus. Further- more, those bundles which come from the ansa peduncularis inter- sect with bundles from the frontal lobes to the stratum zonale (Fig. 36). Bundles of fibres from the ascending pillar of the fornix pass obliquely from the base of the brain through the interior of the thalamus in order to reach the stratum zonale (Figs. 6, 31, 55). The surface of the thalamus descends vertically from the inner margin of the stratum zonale to the third ventricle, and seems to consist of gray substance. But this gray substance is not part of the thalamus ; for it ex- tends below the optic thalamus into the infundibulum. It is part of that gray investment of the primitive medullary tube which be- longs to the primary anterior vesicle, which later on develops into the thalamencephalon ; the upper wall of the thalamencephalon retains a membranous covering. This central gray substance, which surrounds inferiorly the basal ganglion of the tractus opti- cus (Fig. 31), touches externally upon the inner wall of the stratum zonale of the thalamus. The stilus internus, derived from the ansa peduncularis, forms the median surface proper of the thalamus, and the (median) most superficial fasciculi of the stilus internus take an arciform course backward before entering the posterior commissure. The thalamic stilus anterior of the ansa peduncularis takes its origin in the temporal lobe (Fig. 6), and in all probability also in the external capsule. The optic thalamus has an outer boundary of medullary sub- stance ; for the internal capsule of the lenticular nucleus may at the same time be considered the external capsule of the thalamus (Figs. 5, 6) : and as soon as traces of the lenticular nucleus in the parietal region are lost, a continuation of medullary substance behind the 88 Psychiatry. internal capsule surrounds the thalamus on its outer side (Fig. 37). The stilus anterior encircles the thalamus on its lower side (Fig. 35). Like the stratum zonale, the stilus anterior is a layer of white substance superimposed directly upon the thalamus ; in entering the thalamus it divides into laminae, and may therefore be con- sidered an integral part of that ganglion. In that part of the thalamus which belongs to the parietal region, a narrow layer of substance forms what might, roughly speaking, be termed a claustrum. (This is the " lattice-like layer " of Arnold.) This formation has a twofold origin: (1) This outer narrow vertical layer of the ganglion (Figs. 30, 32) is bounded by fasciculi from the temporal lobe which ascend with the gray substance of the thalamus to the inner side of the transverse laminae of the parietal region, and in a direction parallel to the external surface of the ganglion ; (2) radiating fibres uniting the hemispheres with the corpus geniculatum externum, and, taking a vertical course, intersect in the external gray substance of the thalamus with transverse medullary laminae from the parietal region, which, on cross-section, present the appearance of transverse fibres. Being surrounded by a superior and inferior and by an exter- nal and internal medullary wall, the thalamus appears wedge- shaped, both on frontal and on sagittal sections. On anterior frontal sections (Fig. 6) the wedge turns its back upward and its edge downward ; in sagittal planes, it is low anteriorly and high posteriorly (Figs. 29, 36); and on transverse horizontal sections (Fig. 5), it is narrow anteriorly and broad posteriorly. The longitudinal axis of the thalamus is bent at its posterior end to the outer side, a consequence of the median intrusion of the mid- brain (Figs. 16 and 17); from among the structures of the mid-brain, the corpora quadrigemina press but little, while their appendages, the corpora geniculata, pressing from the inner and lower side, crowd in upon the optic thalamus (Figs. 5, 38, 30, 32). The formations surrounding the optic thalamus have shed con- siderable light upon the superior division of its projection-system between cortex and ganglion. We must now study: I. The course of the hemispheric fasciculi in the interior of the thalamus. II. The origin of the tegmentum in the thalamus. III. The basilar structures which support the thalamus and which, together with the bundles of the tegmentum, present a special sectional area. I. The hemispheric (white) substance in the interior of the Thalamus Opticus. 89 optic thalamus, the upper member of its projection-system, enters it from the surrounding white layer, and determines the arrange- ment of the gray substance by dividing up into laminae, which can be distinguished on gross macroscopical preparations. The medul- lary systems referred to above furnish distinct boundaries for two gray nuclei of the thalamus. a. The crus adsccndens fornicis divides above, and, before it is lost in gray substance, forms the boundary of the anterior nucleus, tuberculum anterius, nucleus caudatus thalami. The stratum zonale, dividing as it were into an upper and lower lamina, partici- pates in this boundary, through the mediation of a transverse layer of fibres from the medullary substance of the frontal lobe (Fig. 38, left a.). b. That portion of the stratum zonale which is derived from the ansa peduncularis bounds, and passes through the ganglion of the habenula conarii. The small nerve-cells of this ganglion resemble in form and density of arrangement those of the conarium. Both are united to bundles of the posterior commissure. c. Viewed on a cross-section the stilus internus (int. pedicle) of the optic thalamus (derived from the ansa peduncularis) is seen to spread, brush-like, at the outer side of the gray substance of the III. ventricle, over the entire width of the anterior half of the thalamus to the inside and outside of the crus fornicis (Fig 6, st. i. from Th. to Tp.). In the anterior third of the optic thalamus, and on frontal sections, we can discern a group of delicate bundles coursing upwards from the internal capsule over the lenticular nucleus into the deeper layers of the thalamus (Schnopfhagen). In the thalamus these bundles run parallel to those bundles of the ansa peduncularis which pass under the nucleus lenticularis. The former layer intersects from below with the radiating fibres arising from the stilus anterior of the thalamus, d. The stilus anterior (anterior pedicle) of the thalamus enters this ganglion, in which it is successively denuded of the various laminae of anterior medullary capsule (Figs. 5—Cp. i., 33, 36—2,38 —a, 5 5). These laminae inflate themselves so as to present a surface parallel to the convexity of the thalamus, and close like onion leaves in upon the medullary capsule. Layer by layer the medullary sub- stance alternates with the gray substance, the spindle-shaped nerve- cells of which invariably have their axes directed parallel to their medullary laminae. The medullary substance of the stilus anterior crosses in the anterior third of the thalamus the radiations belong- 9o Psychiatry. ing to the internal pedicle ; further back it lies more to the outer side. An inner (median) portion of the thalamus enters the pos- terior commissure ; the greater portion assumes a funnel shape, and passes directly into the tegmentum. The laminae of the anterior pedicle have their convexities concentrically arranged, conse- quently they, and the intervening gray substance, appear trun- cated on longitudinal and sagittal sections, and present the appear- ance of arches convex anteriorly, and pushed one within the other (Fig. 55), as seen in cross-sections. e. The radiating fibres from the parietal lobe give rise in the posterior half of the optic thalamus to transverse laminae alter- nating layer by layer with gray substances (Fig. 34). f. The relations of the occipital and temporal lobes to the thalamus were demonstrated in a previous section by means of preparations obtained by the cleavage method. (Fig. 21, p. 47.) Its relations to the optic radiations were commented upon above, (see p. 86; also pp. 93, 94.) II. Other medullary fibres in the thalamus opticus indicate the origin of the tegmentum of the crus cerebri. 1. On cross- sections, passing immediately in front of the corpora quadri- gemina through the widest part of the wedge-shaped formation of the ganglion habenulae (Figs. 16 and 17), we see a considerable group of fibres derived from various parts of the stratum zonale, and possibly also from the posterior commissure. These fibres pass through the swelling of the fifth-order-fasciculi of the internal capsule, through the nucleus ruber of the tegmentun. Longi- tudinal sections through this group clearly exhibit the passage of these fibres into the tegmentum. As the anterior innermost bundle of tegmental fibres, they either curve around the outer side of the third nerve, or are inter-woven with the latter and are then continued into the fasciculus longitudinalis of the posterior division of the pons. Through the intervention of ganglion cells, these fibres are in all likelihood continued into the oblongata, ultimately into the spinal cord. To this group of fibres the name pedunculus ganglii habenulce, or fasciculus retroficxus, has been given. Forel suggested " fasciculus of Meynert." (Figs. 31, 34, and 35.) 2. Above the nucleus ruber another nucleus is exhibited, which seems to unite toward the median line with the thal- amus opticus, but is distinctly limited to the outer and lower side. Appearances would lead one to suppose that arciform Origin of Tegmentum. 91 medullated fibres lying in the frontal plane had joined the cross- sections of those fasciculi which surround the nucleus ruber (Fig. 34). This medullated arch is surrounded on the outside by a few striated, irregular, concentric medullary lines, striae med- ullares thalami optici. But these medullary fasciculi do by no means course within the frontal planes alone. Schnopfhagen discovered on gold preparations that these striae consist of short fragments of fibres and of cross-sections of fibres. On lon- gitudinal sections of gold preparations (Fig. 36) these striae are seen to be made up of segments of fibres from various directions, some of which turn about to enter the tegmentum of the crus. On superficial examination the innermost lamina medullaris forms a medullary vesicle, which encloses a semicircular area of thalamic gray substance isolated from the remainder of the thal- amus on all but the inner side. This enclosed gray substance does attain to the dignity of a centre in the optic thalamus, for it is not traversed by the same longitudinal fasciculi which contain radiating fibres of the anterior pedicle, and in- vades much larger areas of the thalamic substance than the small semicircular area bounded by the laminae medullares after their interruption by ganglion cells of the thalamus. These stria medullares with ganglion-cells interspersed in between are simply slender continuations of the anterior pedicle. Cross-sections passed through the brain trunk at various levels present the same deceptive appearance of the innermost lamina medullaris taking a transverse arciform course to the tegmentum. This deception must be ascribed toe. number of concentric arches diminishing in size in a downward direction, of which one would fit within the other. At the level of the posterior commis- sure, this arciform formation has disappeared and a cross-section P.cm. Fig. 35- Sagittal Longitudinal Section through the Brain of Lemur. C.Qu. Corp. quadrigemina. Cp. Posterior commissure passing into the tegmentum. P.gl.h. Fasciculus retro- flexus of the tegmentum. III. Oculo- motorius nerve in front of the pons and behind the corpus mamillare—Cm. P. cm. I^S tegmental fasciculi. Th. Optic thalamus. st. inf. Stilus internus of thalamus, fd. Descending pillar of the fornix and anterior commissure. II. Optic chiasm. 92 Psychiatry. of bundles has usurped its place on the outer side of the nucleus ruber. From this we may infer that longitudinal fibres contribute to this origin of the tegmentum by uniting downward in funnel- shaped fashion. The laminae medullares of the optic thalamus are, therefore, composed not of transverse fasciculi, but of longitudinal bundles, which are juxtaposed in such a way as to give rise to an arciform formation on cross-sections. The laminae medullares seem to form a dense medullary mass, and to be the extension of long-drawn fasciculi: but this is a deceptive appearance, due probably to the C.ext. Fig, 36. Sagittal Oblique section through an Infant Brain. (Stained with Potassium and Gold Chloride.) This section through the thalamus and lenticular nucleus of the right side runs obliquely in an internal and posterior direction, and would cross the median line if continued below the corp. quadrigemina. Qu. Corp. quadrigemina A. Aqueductus Sylvii. Th. Bundles of tegmentum from the thalamus. R.K. Lateral layers of the red nucleus. Lm. Laminse medul- lares in optic thalamus. 2. Interlacing of bundles destined for the stratum zonale and for the thalamus in the internal capsule. Nl. Lenticular nucleus. C.ext. Its external capsule. Plbl. Superior peduncle of the cerebellum above the point of decussation. P.P. Pes pedunculi. J.S. Stratum intermedium. L.ltf. Lentiform disc. II. Optic tract. Ans.n.l. Ansa lenticularis, which on its way to the estratum intermedium is interwoven with the fasciculus of Arnold belonging to the internal capsule, y. Island. The layers as far as the internal capsule are incorrectly repre- sented in the wood-cut. N. B.—The apparent anterior end of the discus lentiformis represents the radiation of the posterior longitudinal fasciculus. The same is true of the dark crescent of the optic tract. introduction of transverse radiating fibres from the parietal lobe. These radiating fibres are interwoven with the juxtaposed, teg- mental fasciculi from the corona radiata and pass in the direction toward the third ventricle. By filling up the gaps left by the other Lamince Medullares. 93 order of fibres, these two distinct groups (the lam. med. and the fasciculi from the parietal lobe) form a compact mass of medullary substance. Longitudinal oblique sections through the thalamus exhibit, in addition, concentric arched formations, which are con- vex anteriorly and open posteriorly, thus contrasting strongly with the appearance of the striae medullares on longitudinal sections. But the former do not form compact medullary masses ; they are the result of the defoliation of the stilus anterior in the optic thalamus, the resulting laminae having a concentric arrangement, and presenting their convex surfaces toward the Frontal Transverse Section through the Human Thalamus. Nc. Caudate nucleus. Mh. Medullary substance of the hemispheres. Pd. Pes pedunculi. Z. Stratum Zonale of thalamus. Cm. Median commissure. Fa. As- cending pillar of fornix. Fd. Descending pillar of fornix. Lp. Cross-section and radiation of the posterior longitudinal fasciculus. Fc. Tuber cinereum. third ventricle. These laminae belong to an anterior region of the thalamus, in which they and the connecting fasciculi change off with layers of gray substance, in which they end and originate anew (Fig. 33). This entire system of nerve bundles, which enters the thalamus as its stilus anterior, and emerges from it as an outer division of the tegmental fasciculi, forms two stratified cones with ad- joining bases, situated about midway between the ends of the thalamus. The anterior cone is by far the larger one of the two ; the posterior cone is hollow, and exhibits a cleft-like open- ing toward the median line. The innermost median area of the 94 Psychiatry. thalamus is occupied by the system of fibres composing the internal pedicle, which connects with the posterior commissure and runs back of the fasciculus rctrofiexus. This accounts for the formation of the median cleft of the tegmental cone in the thalamus. At the same time the external concentric laminae which surround }he lamina medul- lares, are too low to rise in concentric fashion above the upper segment of the latter. There are several reasons for the lowness of these parts : First, the height of the brain- trunk diminishes in the direction from the thalamus to the tegmentum; and, secondly, we find that the fasciculi of the tegmentum, originating in an antero-posterior direction, take, in keeping with the parietal flexure a course from above downward toward the pons. To the tegmentum of the crus cerebri still other fibres are added, which emanate from the posterior commissure, lie below the fasciculus retroflexus, and in passing to the front run parallel to the " fasciculus of Meynert," 1 but later on curve downward toward the medulla (Fig. 35, Cp.). The commissura media, between the optic thalami, resembles the posterior commissure in structure ; for by the cleavage method we can prove that bundles pass through this commissure also, which unite the stilus interior of one side with the optic thalamus of the opposite side (Fritsch, Hollander). In the thalamus these bundles seem to ascend a short distance in an oblique longitudinal direction. A stout anterior bundle of fibres from the tractus opticus con- nects the retina with the thalamus. This bundle ascends between the corpus geniculatum externum and the pes pedunculi, in front of the pulvinar, and has its radiating fibres enter the thalamus on its inner aspect. These radiating fibres are situated at some dis- tance from the fibres connecting the tractus opticus and the discus lentiformis, and ought not, therefore, to be confounded with the latter group of fibres. III. The ganglia of the fore- and inter-brain are surrounded by medullary formations, which start from the base of the brain and take a course about parallel to that of the tractus opticus. These constitute the substantia innominata of Reil. This med- ullary layer lies in front of the tractus opticus, and in the depths of the brain structure it separates the optic tract from the pes pedunculi. After removing the optic tract (Fig. 23), we observe that this basal zone undermines the lamina per- forata anterior, and passes through the nucleus caudatus as it 1 Forel suggested this designation. Girdle Formations. 95 did through the anterior commissure, only in a more basilar direction. There are morphological reasons for the formation of this girdle. Its bundles pass from the outermost region of the brain-axis (princi- pally from the corona radiata of the temporal lobe, and from the external capsule), over the crus cerebri to the median points of decussation (median and posterior commissures. Schnopfhagen's decussation). v As regards the origin of this medullary substance of the ansa peduncularis (Gratiolet), from the temporal lobe, the interweaving of bundles of the ansa peduncularis with the fibres radiating into the pulvinar, and with the outermost bundles of the pes pedunculi (Fig. 23, Tm.), presents the appearance as though the outer fibres of the crus travelled in girdle-like fashion below the posterior margin of the lamina perforata anterior to the inner surface of the crus (Fig. 23, in front of Pd.). This is easily ex- plained, for the radiating fibres from the temporal lobe into the brain-trunk either turn backward to enter the thalamus and crus cerebri, or forward to enter the ansa peduncularis; but fibres run- ning in both directions, from identical points of the sectional area of the temporal corona radiata, interweave with one another. This network of fibres, and the course of these bundles in oppo- site directions, can be well exhibited on cleavage preparations (Fritsch, Hollander). The following formations go to make up the basilar girdles of the brain-trunk : 1. The tractus opticus, which not only sends decussating fibres through the chiasma into the optic nerve of the opposite side, but also transmits non-decussated posterior fibres into the optic nerve of the same side. Part of the tractus opticus adjoins the tuber cinereum, and here its upper surface is connected with Wag- ner's ganglion opticum basale (Fig. 31, g.o.b.). The chiasma is sus- pended as it were on this ganglion, for the latter stands out entirely free in a basilar direction. The ganglion transmits non- decussated fibres to the optic nerve. 2. In front of the chiasma we find the pedunculus scptipellucidi (as the superficial layer of the Iqminaperforata anterior) passing beneath the nucleus caudatus and to the outer side. Between this superficial layer of fibres and parallel deeper layers in the caudate nucleus, which, as was noticed above, arise from the ansa peduncularis, we discern a special stratum of dense, transversely directed ganglion cells, which are parallel to said fibres, and can be traced as far as the external capsule. They constitute a flat, sharply 96 Psychiatry. defined, special ganglionic formation, which equals in area the lamina perforata anterior. This ganglion ansalpeduncularis has its cells run parallel to the ansa peduncularis and is intersected by bundles coming from the latter. 3. Immediately above the optic tract lies the commissura inferior (Fig. 6, II.), the median portion of which is lodged in the convexity of the tuber cinereum, while its recurrent lateral portions lying to the outer side of, and immediately adjoining, the crus cerebri, about which they form a belt, run deeper in as- cending. 4. The ansa peduncularis. We can gain no insight into the general structure of the brain-trunk without keeping in mind preparations obtained by the cleavage method. From such preparations we can understand at a glance the stratification of the ansa peduncularis ; and transparent transverse sections will corroborate in excellent manner the facts brought out by these preparations. The posterior surface of the ansa peduncularis projects gutter-like over the commissura optica (Fig. 22 to the right and next to the cut end of the optic tract). The ansa peduncularis is composed of three layers: the internal pedicle of the optic thalamus, the posterior medullary lamina of the tegmen- tum (so-called fasciculus longitudinalis posterior), and the ansa lenticularis; all three layers taking entirely different courses. a. The stilus internus forms the most ventrad layer of this formation (ansa ped.). It was fully described on pp. 88 and 89. There is this, however, to be added, that part of its bundles cross the median line through the median commissure (Fritsch, Hollander). b. The posterior longitudinal bundle (fasciculus longitudinalis posterior) lies, as viewed from the base, above the stilus internus, and under the ansa lenticularis. To the outer side the three layers of the ansa ped. are super-imposed upon and cover one another. In the median direction they are distinctly separable, for the stilus interior extends least, and the ansa lenticularis extends farthest, backward. Within the ansa the posterior longi- tudinal bundle is distinguished by a more grayish tint. This applies to gross preparations, from which we may learn also that this bundle is softer than the other two layers. On their course backward, parallel to the third ventricle, the stilus internus and the fasciculus longitudinalis posterior diverge to a distance equal to the height of the aquceductus, for the stilus internus passes be- hind the aquaeduct into the posterior commissure, and the poste- Radiations of Post. Long. Fasciculus. 97 rior longitudinal fasciculus enters the gray substance in front of the aquaeduct. These bundles, emanating from the ansa pedun- cularis, are joined by others from the tuber cinereum, which lie close to the inner surface of the third ventricle, or the infundibu- lum, respectively, while, on the other hand, the commissura infe- rior lies closer to the outer surface of the tuber cinereum. The bundles in question were demonstrated also by Schnopfhagen on frontal sections of preparations which had been stained with gold. Turning back, at least a portion of these bundles enter Schnop- fhagen's commissure, which lies in the posterior wall of the third ventricle above the red nucleus and behind the corp. mammillaria. Below this commissure they course over the nucleus ruber (Fig. 31), and through the mid-brain (Figs. 40 and 41), probably as the innermost bundles of the cross-section. This portion of the pos- terior long, fasciculus, derived from the central gray substance, seems solely to cover the ascending crus of the fornix ; in order to enable the latter to enter the thalamic ganglion, the posterior longitudinal bundle divides into two laminae for a certain distance of its course. The expression fasciculus longitudinalis posterior applies only to the lower portion of this medullary formation, with which every one has been made familiar through Stilling's investigations, and which has been described as lying in front of the aquaeductus Sylvii in the mid-brain, and in front of the gray substance of the pons (Figs. 40, 41, 42, 43—L.). That portion of this system, which I described more than twelve years ago as a connecting link between the ansa peduncularis and the posterior longitudinal fasciculus (Figs. 31 from rfx to the basal fl.p.), is merely the inner margin of the powerful radiation of the posterior fasciculus, which covers the entire posterior surface of the peduncular system. The familiar leaf-shaped tapering (to the outside) form of the fasciculus longitudinalis posterior is nothing more than the lower, probably imperfect, extension of this medullary radiation. The radiation of the posterior fasciculus represents a projec- tion-system passing from the medullary substance of the hemi- spheres (from the cerebral cortex) into the central gray substance, which forms, as it were, at one and the same time, the floor of the ganglia and the roof of the crus cerebri. All the radiating fibres converge from the central surface toward the inner, thicker margin of this radiating system, along which they take a sagittal course downward. Fibres taking such 98 Psychiatry. a direction can be seen below the thalamus, on frontal sections also (Fig. 37, Lp; cut made in the year 1866). This fact was recognized later on by Forel. The radiations of the posterior longitudinal fasciculus, from the frontal lobe of the cerebrum, pass through the internal cap- sule into the medullary substance above and in front of the dis- cus lentiformis, which is part of the radiation of the posterior fas- ciculus (Fig. 33, the short, interwoven bundles of the fasciculus of Arnold). The radiations from the temporal lobe bend from below upward with those of the external capsule in the ansa peduncularis, and take the same course as the other fibres under the thalamus opticus (Fig. 31). In the ansa peduncularis these radiating fibres do not form a mere strand, but a broad, stout lamina, the outermost bundles of which lie so far distant from the median bundles that they ascend to the outside far above the tractus opticus (comp. Fig. 31, fl.p. and Fig. 29, where the ascent to the inner capsule is not marked above II.). That portion arising above the optic tract surrounds the inner capsule with its concave surface turned outward, so that the internal capsule is seen to interrupt on sagittal sections the course of the anterior segment of radiating fibres into that seg- ment which passes over the crus. No part of the tegmentum of the crus extends as high up in the brain-trunk as does the fasciculus posterior. At the highest point the radiations lie in front of the pes pedunculi (Fig. 31 fl.p.), then in front of the stratum intermedium, the bundles of which connect at the outer side with the nucleus len- ticularis ; after that this system of fibres rises above the discus lentiformis,1 and lower down the posterior fasciculus is pushed far off from the pes pedunculi by the nucleus ruber of the tegmen- tum. In order to cross the processus cerebelli ad cerebrum, the fasciculus longitudinalis ascends from the median side of the prae- peduncular (Wilder) radiations to that process itself. Over the discus lentiformis, on the contrary, the radiating fibres of the pos- terior bundles were covered by the radiations of the processus ad cerebrum (Forel), the latter lying close under the thalamus as the fifth layer of the internal capsule (Fig. 33, 5). In this figure the discus lentiformis unites with the radiations of the fasciculus longitudinalis posterior. Figures 54 and 55 exhibit the radiations of the longitudinal bundles as dark cross-sections of medullary sub- 1 In Fig. 31 it should fill out the space NR., fl.p. and of the bundles SS. Mesencephalon. 99 stance in front and above the discus lentiformis Lb. and L.lf. The nerve-bundles passing over these belong to the radiation of the processus cerebelli ad cerebrum. (c) The ansa lenticularis has been described above, p. 81. C—THE MID-BRAIN (MESENCEPHALON). Four symmetrically situated ganglia belong to each half of the mid-brain region, viz.: I. One half of the upper, 2. one half of the lower, corpus bigeminum ; 3. the external, and 4. the internal, Fig. 38. Horizontal Transverse Section through the Brain of a Dog—the Right Half nearer the Base than the Left. Fr Frontal lobe. P. Parietal lobe. occ. Occipital lobe. Fr. Corpus callosum. S Septum pellucidum. F. Descending pillar of the fornix, a. Stilus anterior of the thalamus Nc. Caudate nucleus. Th. Optic thalamus on its median surface ; next to the eray substance of the third ventricle are bundles in the habenula and to the out- side of it which enter the decussation of the posterior commissure in front of a frag- ment of the pineal gland, and are connected with the inner margin of the stratum zonale ; the median choroid plexus has been torn off from the latter, ge. External geniculate body. II. Optic tract. Bs., Bi. Superior and in enor bigeminal body Br Brachium of the superior bigeminal pair. c. Transverse bundles in the roof of the aqueduct, adjoining the posterior commissure. Aq. Sylvian aqueduct. corpus geniculatum. In mammals we must speak of upper and lower corpora geniculata (Forel). Both parts of the corpora quadrigemina unite at the one end with the hemispheres, and at the other end with the retina, which must be considered homolo- gous to the bulbus olfactorius. There is this difference, however, IOO Psychiatry. the bulbus olfactorius connects directly with the cortex; while the retina is united first to the corpus bigeminum. From the retina fibres of the optic tract enter internodal masses, which are joined to the cerebral cortex by portions of the so-called brachia of the corpora quadrigemina. i. Connections with the tractus opticus : The connection of the tractusopticus with the corpus geniculatum externum (Figs. 26, 30, 32, 38) is very evident. This ganglion is not a homo- geneous mass, but consists of gray laminae alternating with white substance. Properly speaking, the corpus genicu- latum externum is a folded gray membrane which does not, like the gray substance of the retina, unroll its entire surface, but remains enclosed in a medullary capsule. All the fibres of this capsule are not connected with cells of the corpora geniculata. The superior corpus bigeminum is joined to the corp. genie, ex- tern, by bundles of fibres which in man constitute the posterior portion of the supe- rior brachium, or its well- developed posterior margin, and are covered by the pul- vinar. In mammals whose pulvinar is scantily developed, these very same fasciculi lie exposed on the surface ; here, moreover, the corpus genicu- latum externum lies upon the The radiating fibres from the corp. genie. r9 Fig. 39- Oblique Section through the Brain of a Lion (Stained with Potassium and Gold Chloride). Qu. s. Superior pair of bigeminal bodies. R. Gray substance of its surface. Qu.i. Ob- lique section of the inferior bigeminal body. Rd. Radiating bundles from the corp. quadri- gemina to the gray substance surrounding the Sylvian aqueduct. L. Formation of the lem- niscus (the line of the upper L should point to the decussation). A. Sylvian aqueduct. Rph. Raphe. P. Pes pedunculi. Tg. Tegmentum of the crus.—The presence of external (dark) lemniscal fibres, derived from higher levels of the superior bigeminal body, is due to the ob- liquity of the section. optic thalamus (Forel). extern, into the corp. bigem. super, pass into the superficial layer of neuroglia; this layer of gray substance is more developed in the mammalian brain, on account of its greater (typical) wealth of con- Tractus Opticus. 101 nective tissue (Fig. 39, R., cortical layer, Forel). The fasciculi bi- gemino-genicular cs of the upper bigeminal body surround on both sides its oblique oval ganglia, which are flattened anteriorly by lying upon the stratum lemnisci. For this reason these ganglia present on cross-section the appearance of plano-convex lenses. The axes of the superior bigeminal body intersect close behind this ganglion and diverge anteriorly, leaving the posterior commissure exposed between them (Fig. 17.) In comparison with the distinctly trans- verse direction of the lemniscal layer, the fasciculi bigem.-geniculares, which enter along the entire upper margin of the superior corpus bigeminum, might be considered to be longitudinal fasciculi of the latter (Fig. 39). This is particularly true of those bundles which are situated on the inner border of this ganglion. The corpora quadrigemina are connected also with the corpus geniculatum internum, which is united distinctly (1) to the corpus bigeminum superius and (2) to the corpus bigeminum inferius, through bundles which reach it from the brachium corporis quadrigemini inferius. Radiations from the tractus opticus into the corp. geniculat. in- ternum travel by way of the discus lentiformis, with which the tractus is connected (Stilling). The discus is united toward the rear, by a medullated pedicle, to the brachium corporis quadrigemini inferius, situated on sagittal sections in front, and on frontal sections to the inner side, of the thalamus. At the same time it is unquestionably true that the internal geniculate body effects a connection between the inferior brachium of the quadrigeminal body and the superior bigeminal body; but its connections with the corpus bigeminum inferius are equally distinct, so that the optic tract may well appear to be connected by means of the discus lentiformis with the inferior bigeminal body. The trans- verse position of the bundles which surround the ganglion of the lower corpus bigeminum does not enable us to distinguish between the indirect radiations from the retina and the cortical fasciculi of the inferior brachium, as we can do with the longitudinal optic radiations in the upper bigeminal body. A superficial inspection of these parts would lead us to suppose that the tractus opticus is connected directly with the corpus geniculatum internum ; but this is not borne out by a more careful examination. It will be well to divide the radiations from the nervus and tractus opticus into four component parts : /. The anterior radia- tion into the ganglion basale. Ascending fasciculi unite this ganglion 102 Psychiatry. with other regions of the central gray substance. 2. The upper radiation, entering the discus lentiformis which lies above the corpora geniculata. This discus lentiformis enters apparently into connection with the radiation of the posterior longitudinal fascicu- lus (?). j. The outer radiation, or radiation into the corpus genicula- tum externum. 4. The internal radiation into the thalamus, which contributes, on the inner side and to the front of the corpus genicula- tum externum, to the formation of the latter's capsule ; and which, after the fashion of the corona radiata, enters the thalamus from the outer side in front of the pulvinar. That these bundles, after having been interrupted in the thalamus, are continued toward the corpus quadrigeminum, has not been proved. 2. There is a twofold, powerful connection between the corpora quadrigemina and the cerebral cortex. a. The corpora quadrigemina receive radiating fibres direct from the cortex (upper member of their projection-systems). In the superior bigeminal body, cortical radiations are found in the course of the arm of this upper corpus bigeminum, constituting its deeper anterior layers, which are covered by the connection between this said brachium and the corpus geniculatum externum (Fig. 38, left, between T. and P.). The brachium of the superior bigeminal body covers the corpora geniculata, and a wedge-shaped mass of thalamic substance between them ; on frontal sections this cuneus thalami optici intergenicularis appears triangular, The bundles of its brachium connect the cerebral cortex with the inferior bigeminal body. b. The superior and inferior corpora bigemina also receive in- direct fibres from the cortex, inasmuch as their annexed ganglia, the corpora geniculata, are favored with special cortical radiations. The cortical fibres entering the corpus geniculatum internum may possibly also enter the corpus bigem. superius, for (as was stated above) there is a distinct medullary connection between these ganglia. The connections of the external corpus geniculatum with the cerebral cortex radiate from above as a stout medullary mass into this ganglion. While coursing along the outer surface of the thalamus, this radiation contributes to the formation of the lattice stratum of the thalamus (vide p. 88 and Fig. 32, above Ge). The frenulum, or, as it should be termed, the processus cerebelli ad corpus quadrigeminum, unites both bigeminal bodies to the cerebellum. The peripheral connections of the corpora quadrigemina lead Mid-Brain—Cortex Cerebri. 103 (1) directly into the central gray substance surrounding the aquae- ductus Sylvii ; from the fascicular substance of the upper corpus bigeminum, or from its gray substance, numerous fine, radiating bundles are developed, which, on gold preparations, seem to be the main constituents of that area of the corpora quadrigemina lying to the outer side of the lemniscal layer; these penetrate the strat. lemnisci, and can be traced still further into the gray wall of the aqueduct than is exhibited on Fig. 39 Rd. The terminal masses of the optic tract are connected in this way with the cen- tral gray substance, in which fine medullary fibres, belonging to the origin of the third nerve, circumscribe a nucleus confluent with the nucleus of the fourth nerve (Fig. 39, Rd. ; Fig. 40, III.). 3. The corpora quadrigemina are connected indirectly with the central gray substance, for by giving rise to the lemniscus they constitute the central origin of a portion of the spinal cord. The bundles of the lemniscus enter the ganglia from deeper layers of the brachium corporis bigemini superius, and of the brachium inferius. The former portions, after broadening on their way downward into the median base of a triangle, attain the median line, which they cross, and then enter into a deeper stratum of the lemniscus of the opposite side. The latter division of the lemniscus forms a triangular group of bundles with an upper median base, which lie immediately below the brachium anterius, which in turn is spread most in a median direction. The bundles of the lemniscus arrange themselves in such a way on their downward course that those which lie uppermost at the decussa- tion are pushed furthest toward the raphe. The superior bigeminal body exhibits (if light is allowed to fall through) a concentric stratification corresponding (1) to its layer of neuroglia; (2) to a layer of vertical fasciculi from the corp. geniculatum externum ; (3) to a radial layer interwoven with these fasciculi ; (4) to the transverse layer of the lemniscus (Fig. 39)- The anterior surface of the lemniscus receives a number of fasciculi, the origin of which is a matter of conjecture ; their par- ticipation in the decussation of the lemniscus behind the aqueduct is very improbable also, for they would have to cross back again in front of the aqueduct. These bundles surround the aquaeduc- tus Sylvii, and lie next and to the one side of the lemniscus, of which Forel thinks they form a part, and which he terms fasciculi decussations anticce. They form a convex swelling along the 104 PsycJiiatry. anterior margin of the aqueduct (Fig. 40, Krz B.), due to their inter- weaving with longitudinal bundles in which I recognize the cross- section of the posterior commissure. These bundles appear to me to spring from that cluster of cells which Jacubowitsch and Virchow likened to the cells of the sympathetic, and which, at the same time, furnish the central origin of the descending root of the fifth nerve (Fig. 41, 5). I have termed these capsular bundles of the aqueduct quintus- Fig. 40. Transverse Section through the Region of the Superior Pair of Quadrigeminal Bodies in a Plane Vertical to the Longitudinal Axis of the Brain. Big. s. Superior bigeminal body. A. Sylvian aqueduct. Lms. Lemniscus (fillet) arising from the decussation behind the aqueduct. Bri. Inner protuberance and fasciculi of its brachium. Krz. B. Anterior cross-bundles of the corpus quadrigemi- num. x. Points to the region of decussation. Th. Bundles from the thalamus to the Tegmentum. L. Posterior longitudinal fasciculus. T. gris. Central cavity-gray. R. Ill, III. Nucleus and central roots of the oculo-motorius. 3 L.P. Root of the third nerve and lamina perforata posterior. R. Raphe. P.P. Pes pedunculi. S.S. Stra- tum intermedium and Soemmering's substance. R.K. Red nucleus of the tegmentum. columns; perhaps the term fasciculi marginales aquoeducti would be better (unbefangener). On the cross-sections they are seen to rise from a narrow margin parallel to the aqueduct, to spread in rays in order to interweave with the bundles of the posterior commissure mentioned above, then to attain their point of decus- sation in the median line by forming a narrow band between the posterior longitudinal bundle and the nucleus ruber (Fig. 40, L. and R.K.). From the point of decussation they encircle the Mid-Brain. !Oo nucleus ruber on the inner side, and are then lost to the front and to the outer side of this nucleus. The corpus bigeminum inferius is simpler in structure than the superior bigeminal body. The fasciculi of the brachium corp. quadrigem. inferius, which run under and past the corpus genicu latum internum, give the former a white surface, traverse the gan- glion, and bound it below giving it the shape of a biconvex lens. After decussating in the roof of the aquaeductus, the bundles of the Fig. 41. Cross-Section through the Inferior Pair of Bigeminal Bodies in Man. Big. i. Inferior bigeminal body. x. Crossed transition of the inferior brachium of the corp. bigem. into the lemniscus. A. Aqueduct (the outlines of which have been unnecessarily indented and prolonged anteriorly). L. Posterior longitudinal fasciculus. IV. Nucleus of fourth nerve (the line should point further outward to the oval space containing small cells). 4. Root of fourth nerve. 5. Descending trigeminal root. P. Cbl. Dcs. Decussation of the superior peduncles of the cerebellum. Th. Thalamic fasciculi of the tegmentum. Lms. Lemniscus. Lp. Posterior perforated lamina. S.S. Stratum intermedium, and Soemmering's substance. P.P. Pes pedunculi. above brachium enter the lower division of the quadrigeminal por- tion of the lemniscus (Fig. 41, Fig. 56, L. II.). The cross-section of the posterior decussation of the lemniscus extends on sagittal sections through the middle of the inferior corpus bigeminum only as far as the middle of its entire length, and stops short at the beginning of the frenulum. The mid-brain (mesencephalon), like the posterior half of the io6 Psychiatry. inter-brain (thalamencephalon), shows four distinct layers: I. Posteriorly, the ganglion, Big. s., Big. i; 2. The cross-section of the tegmentum (Figs. 40 and 41, Th., Lms., L., R.K., P. Cbl.); 3. The stratum intermedium (S.S. ; in Fig. 41 the arrow is too short). 4. The pes pedunculi (P.P.). Above its point of decussation the superior peduncle (Binde- arm) traverses the tegmentum in front of (ventrad) the upper bigeminal body ; during decussation this cerebellar peduncle crosses the tegmentum in front of the lower bigeminal body. Ventrad of the lower bigeminal bodies the fasciculi of the tegmentum are crowded from the inner; side outward, owing to the considerable space occupied by the decussation. The sec- tions of the medullary portion of the stratum intermedium do not extend as far to the outside, but further toward the median line, than the sections of the medullary fibres of the lemniscal layer (Fig. 41). The composition of the tegmentum remains quite unchanged in the upper cross-sections of the pons, ventrad of the entire corpus quadrigeminum ; with this exception, however, that the fasciculi of the processus cerebelli ad cerebrum emerge again from the decussation, proceed to the outer side, and form that posterior portion of the system of the superior peduncle, which comes from the region of decussation and enters the cerebellum (Figs. 22, 42). The superior peduncle, as it reaches the external surface, appears club-shaped on cross-section (Figs. 42-45). In the cross- sections of the pons, at levels below the corp. quadrigeminum, this peduncle is powerfully developed. On its outer surface lies the lemniscus, which consists of bundles that on their downward course appear dark, like cross-sections, and of other bundles, the course of which, both to the inner and outer side of the former group, happens to coincide with the plane of the section. The outermost of these flowing bundles might Droperly be considered as part of the origin of the lemniscus from the lower corpus bigeminum ; the inner group of bundles is undoubtedly part of the inferior lemniscus, which ascends from the vermis superior of the cerebellum, through the valvula cerebri, to a position above the processus ad cerebrum, and is transformed in the medulla spinalis into a fasciculus of the lateral column, immediately in front of the posterior cornu (Figs. 23 and 54, LS). The cross- sections of the thalamic bundles seem to be surrounded by the Oculomotor Nucleus. 107 sup. peduncles, and deserve to be termed the thalamic area of the posterior division of the pons. Before giving an account of our present knowledge of the course the fibres take from the brain-trunk to the spinal cord, we will take note of the relations of the gray substance to the nerve- nuclei, down to the point at which the eighth nerve emerges from the brain substance. On gold preparations from the infantile brain (Figs. 43-46) the gray substance seems to be a barren base- ment substance. On sections through the brain-trunk at the level of the upper corpus bigeminum, the aquaeductus Sylvii was seen to be sur- rounded by a gray substance; and the gray substance of its anterior wall could well be compared with the anterior horn of the spinal cord, into which this gray substance is ultimately pro- longed. The nuclei of the third nerve (Fig. 40, III.) are bounded by fibres which reach the central gray substance after passing from the crus cerebri of the opposite side through the raphe, and which in describing delicate, posteriorly convex arches, surround and pene- trate an oblique-oval group of nerve-cells. From this nucleus, which remains confluent with the rest of the gray substance in spite of its well-defined boundary lines, stout root-bundles of the third nerve emerge (Fig. 40, 3), which turn to the front and are slightly concave toward the raphe; these roots either surround the red nucleus of the tegmentum, or lie on its inner side. The innermost of those bundles which pass into the raphe approach close to the aquaeductus, between the nuclei of the third nerves, and there divide into a number of diverging fibres. It is as difficult to trace these to their final termination, as it is to follow up those radiating bundles which seem to approach them from the corpus quadrigeminum. Cross-sections of descending root-fibres of the fifth nerve lie immediately next to those mar- ginal bundles (p. 104) which enter the decussation in the tegmen- tum, in front of the aquaeductus Sylvii. These descending roots -pring from clusters of vesicular cells. The shape of these cells, which possess distinct sheaths, we said above could be compared to the shape of the sympathetic ganglion cells. They take no part in the gray reticulum of fibres surrounding the aquaeduc- tus, into which optic fibres, and the prolongations of third nerve- cells enter. On cross-sections, through the plane of the lower bigeminal 108 Psychiatry. body (Fig. 41, L., IV.), the area occupied by the nucleus of the third nerve seems imbedded in a posterior groove of the posterior longitudinal bundle (nucleus of the fourth nerve). From this nucleus bundles pass to the outside, and along the margin of the aquaeductus Sylvii, which take an obliquely descending course from the anterior periphery of this nucleus to its posterior periph- ery. In a part of this course they necessarily on cross-section present the appearance of circular formations at the lateral margins S.cin. Fig. 42. Cross-Section through the Pons, at the Level of the Exit of the Fourth Nerve. x. Decussation of the fourth-nerve roots, in the velum medullare. 4, 4. Fourth- nerve roots. 5. Descending trigeminal nucleus bordering anteriorly on the posterior longitudinal fasciculus, Th. P.Clb. Superior peduncle of cerebellum. S.fecz. Its fasciculi lying one above the other. Dcs. Decussation of the peduncles. Lms. Lemniscus. St. Lms. Lemniscal layer, including the stratum intermedium. F.trv. p. Transverse bundles of pons. Ped. Peduncular fasciculi. S.cin. Gray floor of fourth ventricle (substantiacinerea). of this central gray substance. These ring-like fasciculi are situ- ated to the inside also of the trigeminal root in the gray substance surrounding the aqueduct. Finally they decussate and enter the valvula cerebri, immediately below the corp. quadrig., from which they emerge free (Fig. 42, 4, 4). New additional bundles from the round cells in the region of the lower bigeminal body render the crescent of the fifth nerve more distinct, which at the level of the emergence of the fourth nerve forms the outermost part of the anterior boundary of the aqueduct. The posterior longitu- Substantia Fcrruginosa. 109 dinal bundles abut upon the median portion of the aqueduct. The velum medullare forms its posterior boundary. At this level a new formation of cells is discovered within the gray substance— i. e., the substantia fcrruginosa (Fig. 43, S. foa.). This is con- nected with bundles which run as far as the raphe in a trans- verse direction parallel to the gray substance. Near the raphe large nerve cells are liberally interspersed between, and inter- Sbsl.cir fppl. Fig. 43- Potassium and Gold-Chloride Preparation from Brain of the New-Born. Cross-Section through the Pons at the Level of the Emergence of the Small Trigemi- nal Root (the Right Half of the Section Representing a Higher Level than the Left). Cbl Cerebellum. v. IV. Fourth ventricle. sbst. cir. Gray floor of fourth ventricle nucl. m. Nucleus of the small (inner) fifth-nerve root. S. gl. Gelatinous sub stance with (sm ) bundles of the large fifth-nerve root. Lms. Lemniscal layer and stra- mmTnermedSm. Br. p. Bundles from the brachium pontis fp la. Anterior longitu- dinal bundles of pons. fppl. Interwoven transverse bundles of pons. fppr. Deep dinal bundles £pfnf «ons \ Laree fifth-nerve root. Th. Thalamic area of posterior £So^ /d- Descending fifth-nerve root (The d^rclublhaped section of the superior cerebellar penduncles are not lettered.) woven with, the fibres of the posterior longitudinal bundle. A transverse commissure or decussation apparently establishes a connection between these cells. It is certain that the dark cells under the locus cceruleus on the floor of the fourth ventricle (Fig. Atf connect in this way with fibres of the raphe, which are derived from the anterior division of the pons and lend a medullary I IO Psychiatry. appearance to its posterior division. These fibres may be looked upon as originating in the pyramidal tract. The substantia ferruginea is a powerful mass extending from the lower corpus bigeminum nearly as far as the lower half of the pons (Stilling). At its periphery it is connected with the innermost bundles of the large root of the fifth nerve. The nucleus of origin of the small fifth-nerve root is situated pos- teriorly from, and to the inner side of, the large fifth-nerve root (Fig. 43, nucl. mi). From this smaller root arise the inner bundles of the entire fifth-nerve root (Fig. 43, sm.). The central nuclei of that massive portion of the large trigeminal root, which originates and emerges at the same level, presents a clustered arrangement of minute heaps of small-celled gray gelatinous substances (Fig. 43, S. gl.). At the level at which the entire trigeminal nerve emerges (Figs. 43 and 44—5) we distinguish from within outwards: (1) an inner large-celled trigeminal nucleus, with the small root; (2) bundles passing from within outward, and between the cross-section of posterior longitudinal bundles, which bundles are derived from the subst. ferruginea of the opposite side, and belong to the descend- ing roots; (3) the upper descending trigeminal root (the familiar small crescent—Figs. 43—rd, 44, r.dest.); (4) the bundles of the ascending trigeminal root, which evidently originate from a nucleus situated below the emergence of the fifth nerve—this nucleus extending to the lower end of the tuberculum cinereum Rolando in the oblongata. The trigeminal root encircles its gray nucleus with medullary substance, which becomes attenuated as it proceeds downwards, and is finally lost in the spinal caput cornu posterioris (Stilling). This root is of very considerable size; (5) Cerebellar fasciculi also join the large trigeminal root. These fas- ciculi lie either immediately upon the outer edge of the processus cerebelli ad cerebrum, or actually invade the latter; (6) the large trigeminal root is joined beyond a question by cerebellar bundles, which are closely moulded to the outside of the superior peduncle (Stilling), and possibly invade the latter. Below and indeed within the planes at which the fifth nerve emerges, begins the central origin of the sixth and seventh nerves. At the highest level we notice decussating bundles leaving the raphe to enter the root of the facial nerve (Fig. 45, right side). Dorsad of the bundles from the raphe lies a circular spindle-shaped section, the genu facialis, the bundles of which arise from the Facial and Abducejis lYerves. 111 anterior facial nucleus, which is situated in an anterior inferior portion of the pons (Fig. 45 ncl. 7). The knee of the facial nerve is made up of root bundles, which curve around the common nucleus of the sixth and seventh nerves, and, in so doing, pre- sent a convex surface facing dorsad. For this reason, the knee seems separated from the emerging root (Fig. 45, left, g) on sections which are laid below this connecting arch, between the (From the same series as Fig. 43.) Cross-Section through the Pons at the Level of the Exit of large V. roots. P. Chi., nucl. Superior peduncles of cerebullum. V. 4. Fourth ventricle, r. dest. Descending root of V. nerve. Sf. 5. Trigeminal root from sutstantia ferru- ginosa. Sg. Substantia gelatinosa. 5, Large V. root. Ol. s. Superior olive. Br. p. Brachium pontis. Tr. prf. Deep transverse bundles of brachium pontis. F. p. l.a. Anterior longitudinal fasciculus. P.tr.spf. Superficial transverse bundles. Fibr. ecfl. Circumflex fibres of pons. Lms. Lemniscal layer and stratum inter- medium. 6 VI. roots. Th. Thalamic area of posterior pons segment. L. Posterior longitudinal fasciculus. 7. Facial roots. knee and the emerging portions of the root. The genu, descend- in"- vertically, appears cut on cross-sections. The common nucleus of the sixth and seventh nerves lies between the two portions of the root. The large and radiating cells of this nucleus are similar to those in the nuclei of the third, fourth, and in the outer nucleus of the fifth nerve. The roots of the sixth nerve bend outward, and undoubtedly connect with the cells of this common nucleus 112 Psychiatry. (Deiters). It is equally certain that the root of the seventh nerve also connects with a lesser portion of this nucleus, the lateral por- tions being taken up entirely with the emerging fibres of the facial, which on their course from the nucleus forward constitute a much narrower bundle. Like all nerve-nuclei, this nucleus is in reality nothing more than the more compact portion of a formation of cells scattered all around it (Deiters). From this nucleus fibrae arcuatae pass into the raphe, thus connecting this nucleus with Stained with Potassium and Gold Chloride. Cross-Section at the Level of Emergence of Seventh Nerve. (The Right Side Represents a Higher Level than the Left.) Cbl. cerebellum, nc. Dentate nucleus of cerebellum, ncl. 6 + 7. Joint nucleus of sixth and seventh nerves, g. Central knee of facial nerve. L. Region of posterior longitudinal fasciculus. Th. Thalamic area of posterior pons segment. 8. Fibres of auditory nerves. 7. Root of facial nerve. 6. Root of sixth nerve. Py. Bundles of the pyramid. R. Raphe, fa. Anterior column. Ols. Superior olive. ncl. 7. Anterior facial nucleus. Lms. Lemniscal layer and stratum intermedium. B. p. Brachium pontis. 5. Ascending trigeminal root. S. gl. Gelatinous substance in the ascending trigeminal root. V. IV. Fourth ventricle. topographically higher centres. There appear to be commis- sures also uniting both nuclei. The anterior facial nucleus is distinct in higher facial planes, increases in size as we proceed downward, and is traversed by the root-fibres coming from the genu (Fig. 45. ncl. 7; Fig. 46, ncl. 7, and on the right to the inner side of 7). This nucleus lies close to the trunk of the facial, which passes between it and the ascending trigeminal root. The Nervus Facialis. 113 knee gathers from the anterior nucleus a number of fine parallel bundles, and at its upper level of origin does not yet present a compact mass to the front of the tubular gray (Fig. 46 ; compare right and left). The nerve fibres from the lower nucleus do not narrow down at once towards the genu, but the external fibres form a loose capsule around the ventral surface of the posterior nucleus. It is possible also that root-fibres pass directly and ob- liquely outward and forward from the lower nucleus to the emerging portion of the facial root. Analogous to the descending, vertical, and ascending roots of the trigeminal nerve, we find that those facial bundles which issue Fig. 46. Stained with Potassium and Gold Chloride. Cross-Section through the Highest Level at which the Auditory Nerve Emerges- The Right Half of Specimen Represents a Higher Level than the Left. ncl f. Nucleus tecti (Stilling). V IV. Fourth ventricle, g Knee of facial nerve 'pdi. Internal division of the cerebellar peduncle, ncl. 7. Lower nucleus of seventh nerve, which receives the bundles from the knee of the facial, ol. 1. Inferior olive P Pons. Py. Pyramid. 01. s. Superior olive. 7. Iacial root. 5. Ascend- ing root "of trigeminus. 8. Auditory root not designated on left side. Br. p. Brachium pontis. (Medipedunculus.— Wilder). from the raphe, are in reality descending fasciculi, arising possibly from the internal capsule, but certainly from the nucleus lenticu- laris. The planes of origin and emergence are identical in the case of these roots which enter the posterior nucleus ; but the root derived from the inferior nucleus of the facial, the genu n. facialis, is an ascending, root (as is evident from fig. 46). The bundles emerging from the lower nucleus pass toward the gray ii4 Psychiatry. substance, anteriorly from this, and move upward as the ge7iu n. facialis; the knee is continued into the emerging root. The genu has the shape of a horse-shoe; its lower branch travels from the inferior nucleus to the gray substance, its upper branch from the middle of the gray substance forward. The knee of the facial nerve constitutes the middle piece and connecting link be- tween these two branches of the horse-shoe formation, lying one over the other. This formation is placed diagonally in each half of the posterior division of the pons. The connecting arches converge in a median and posterior direction. The facial does not run parallel to the anterior roots of the spinal cord, as do the third, the sixth, and the twelfth nerves, which form the ex- Preparalion Stained with Potass. Gold Chloride. From the Newborn. Cross-Section through the Lower Level of the Exit of the Auditory Nerve—The Right Half Represents a Higher Level than the Left. NC. Dentate nucleus of cerebellum. Bp. Brachium pontis. R. Rst. Corpus restiforme. e. Fibrae arcuatas from the corpus restiforme. 5. Ascending trigeminal root. oli. Inferior olivary body. R. Raphe beginning between the pyramids. Z. Stratum zonale. nclSa. Anterior acoustic nucleus, v. IV. Fourth ventricle. 8'. External auditory nucleus. 8. Inner acoustic nucleus. T. Eminentia teres. flf8. Bundles of the flocculus and the external acoustic root which are not separated from each other. ternal boundary of the region of the anterior columns; it lies to the outside of the lateral columns, next to the formation analo- gous to the posterior horn—i. e., the substantia gelatinosa of the asc. trigeminal root. On sections through the level of the origin of the facial nerve the gray substance exhibits two prominences. The inner prominence, the eminentia teres, is due to the common Cerebellum. "5 nucleus of the sixth and seventh nerves ; the external prominence is to be placed to the account of the auditory nucleus (Fig. 45). The origin (?) of the eighth nerve, however, can only be studied in connection with the region surrounding it, including the cere- bellum, THE CEREBELLUM AND THE N. ACUSTICUS. It is not my intention to give an exhaustive account of the minute anatomy of the cerebellum ; for many gaps in our present knowledge of the subject will undoubtedly be filled up by the great monograph which Stilling is now preparing.1 It would be a gratuitous and unnecessary task to attempt to anticipate the work of such a great master. We have already described cerebellar bundles in the internal capsule in the form of radiations from the cerebrum into the nucleus ruber of the tegmentum. We have observed the processus cerebelli ad cerebrum emerging from the red nucleus of the opposite side ; we noticed, furthermore, the unfolding of the frenulum into the vermis superior, after passing through the ante- rior velum medullare, and in the vellum medullare we found the cerebellar portion of the lemniscus. The bundles of the brachium pontis (processus cerebelli ad pontem) consist (1) of a superficial layer, (2) of the interwoven layer which gives rise to secondary bundles in the pes pedunculi, and (3) of deep, transverse fasciculi. These bundles lie to the front of the stratum intermedium, which is joined by the inner bundles of the crus arising from the ansa lenticularis. Gray substance with its ganglion cells, which invade the entire anterior division of the pons, reduce the volume of the crus proper. Through the mediation of these cells, and a gradual reduction of its volume, the bundles of the crus are transformed into a cross- section of the pyramidal tracts, and then enter the brachium pontis. For this reason alone the bundles of the brachium pontis are not commissural fibres, but the continuations of recurrent fibres of the crus. Their commissural nature is disproved furthermore by the fact that a large number of bundles of the brachium pontis pass on the outside around groups of peduncular fasciculi (Figs. 43, 44). Both ends of this circular formation turn toward the opposite brachium pontis, and belong to different levels of the pons, passing, as they do, either from the superficial layer to the interwoven layer, or 1 This work was completed in 1S78.—S. n6 Psychiatry. from this layer into the layer of deep, transverse fibres. Each arch of the fibrce circumflexoe pontis exhibits two branches, one of which enters the crus after taking a superficial course from the brachium pontis of the opposite side ; while the other leaves the pedunculus again, mingling with gray substance and emerging through the opposite brachium pontis. Its processus ad pontem forms a considerable mass on the outer side of the meditullium of the cerebellum. The cerebellum, like the cerebrum, is surrounded by a mantle of convoluted gray substance, the cerebellar cortex, which serves as the point of origin of its medullary substance. It is developed from the pos- terior wall of the posterior (fcetal) cerebral vesicle. Between the ends of the large horizontal fissure lies the hylus, which permits the entrance of the four medullary processes into the gray mantle of the cerebellar cortex. Owing to the medullary substance of the velum medullare (Marksegel) and of the processus cerebelli ad cerebrum entering through the hylus, the vermis superior presents a white anterior surface (Fig. 55). Up to the date of Stilling's great investigations of the cere- bellum, the only gray cerebellar masses known were the cortex and the nucleus dentatus. We are indebted to him for our knowl- edge of the ganglia tecti, which unite in the median line, and of other nuclei, to which he refers in a preliminary notice published in the Centralblatt fur die Medicinischen Wissenschaften. The cortex of the cerebellum consists of three layers : an ex- ternal gray layer proper and an inner grayish-red layer, which are distinguishable by the naked eye. On microscopical examination of the cortex, we discover between these two layers another layer con- taining cells of Purkinje. The basis substance of the cerebral cor- tex, of the ganglia, and of the cerebellar cortex remains the same throughout. According to Obersteiner the cerebellar cortex in the child is covered by a layer of formative cells, which are trans- formed into spindle-shaped fibrils, thus constituting an innermost stratum of the pia mater. The gray layer proper exhibits small nerve-cells, the destructible protoplasm of which often makes it difficult to distinguish between these and non-ganglionic elements. In addition to these generally triangular granules, there are, near the grayish-red granular layer, transverse, spindle-shaped bodies, which run parallel to the convolutions around the fissure between two convolutions, but, unlike the cerebral, spindle-shaped cells, lie to the outside instead of to the inside of the granular layer. I Cortex Ccrebelli. 117 H> agree with Stilling in regarding the cells of the granular layer as multipolar ganglion-cells ; but, instead of looking upon the widely separated groups of cells as the breeding-place of regenerating cells, I would prefer to assume a connection between them and the ramifications of the inner processes of the cells of Purkinje. In doing this, I must demur, however, to the view of Koschewni- koff, who believes that one solitary, non-ramifying process issues from the inner side of Purkinje's cells. The large cells of Pur- kinje, which occur at con- siderable distances apart, and form but a single row, are connected with the in- ternal and external layers l a of the cerebellar cortex. The external powerful pro- cesses of these cells are formed by a gradual atten- uation of the latter's pro- toplasm. They branch dichotomously at first around the sulcus between two convolutions, at an angle of nearly 180 de- grees. From these main branches many others are sent off at right angles; these undergo secondary ramifications like the an- tlers of a deer, and are con- tinued into, and give the appearance of parallel stria- , ,1 ,1 r I a and I b. Gray layer proper; (i b with spindle- tion tO tne OUter layers OI cellsand transverse fibres.) 2. Cells of Purkinje, 3. the cortex. They seem to Granular layer, m. Medullary substance. ramify in those planes of the cortex which belong to one of the delicate, separable laminae of the medullary substance. Accord- ing to Hadlich these processes take a recurrent course when near the cortical surface. The appearance of recurrrent processes may be due to the terminal ramifications turning back in order to unite with the triangular cells of the external layer. The inner halves of the cells of Purkinje are vesicular in shape. Purkinje compared these large cells to field-flasks—ampullae,—from the neck Fig. 48. Transparent Section of the Cerebellar Cortex of Man. n8 Psychiatry. of which the strong process just described would issue. Those processes which issue from the bottom of the flask are much more delicate, and though they may give rise to more than one process (contrary to Purkinje's opinion) they ramify but little. They split up soon into a network of fibres connected with the granular layer. The medullary fibres of the cerebellum have, in all likeli- hood, a twofold origin in the cerebellar cortex: (i) from recur- rent, ramified branches of the external process of the cells of Purkinje; and (2) from the network of the granular layer into which the inner processes of these cells of Purkinje enter. The nucleus dentatus of the cerebellum is traversed by radiat- ing fibres, which probably connect the bundles of the processus ad cerebrum with the cells of this body. At the level of the facial and at the upper levels of the auditory nerve, the nucleus denta- tus is separated as yet from the processus ad cerebrum, and lies very close to the lateral wall of the oblongata, after it has become distinct from the trunk formations in the oblongata. At the level of the central origin of the auditory nerve, the processus ad cerebrum is crowded away from the lateral wall of the floor of the iv. ventricle: (1) by bundles which decussate in passing through the ganglia tecti; and (2) by another group of bundles which covers the nucleus dentatus (Fig. 46, left, to the outside of pdi.); (3} by still another, which at higher levels was situated between the nucleus dentatus and the processus ad cerebrum (Fig. 45, Cbl.). These groups, constituting the corpus restiforme lie to the outer side of the cross-sections of the ascending trigeminal root. At the level of the ganglion tecti (Fig. 46), the first group of bundles passes either, without crossing, to the outer side of this ganglion, or, after decussating through the ganglion tecti on the inner side, downward to the oblongata. These bundles appear truncated on the outer side of the auditory nucleus in the gray substance. They intersect with bundles of the auditory root which, entering In a reverse direction from the anterior surface of the trunk, turn inward on the outer side of the ascending trigeminal root, and after mingling with the former are cut across also. At the junction of these con- verging bundles from the cerebellum and from the auditory root, an angle is formed opening outward. This angle is completely filled by the corpus restiforme, which descends from the region of the nucleus dentatus cerebelli, and edges its way into this angle. These bundles, which have been first described as entering the gray auditory nucleus to the inner side of the corpus restiforme, occupy the area of Clarke's external auditory nucleus. Corpus Restiforme. 11 q In his publications on the lingula and the lobulus centralis, Stil- ling has shown that the neighboring cerebellar convolutions are united to one another by wreath-shaped bundles, and that on sagit- tal sections of the vermis we perceive arciform fibres connecting various regions of the cortex; these arciform fibres varying in length and running in different directions. These arciform bundles lie nearer to the cortical surface than a certain system of fibres which Stilling has described as transverse and decus- sating fibres. The cortex of the cerebellum possesses single and general systems of association bundles, similar to those of the cerebral cortex. The corpus restiforme must be divided into two parts : one part belonging to the pons, and the other to the oblongata. That portion of the posterior division of the pons which is not crowded out by arciform bundles emanating from the nerve- nuclei, and taking the place of the gray substance in the raphe, is occupied by transverse fibrce arcuatce starting from the corpus restiforme. Above the auditory root they emerge distinctly from that portion of the corpus restiforme which is situated between the processus ad cerebrum and the nucleus dentatus, which passes through the trigeminal region at the level of the superior olivary body, and sends forth that superficial stratum of bundles which answers to the corpus trapezoides in animals (Fig. 45, Ols., Lms., 5). These bundles appear at first to course to the front of the superior olivary body, and after crossing each other at an acute angle, to enter the latter body. It is not as evident in man as it is in animals that pedunculated bundles issue from the superior olivary body on the inner side of the ascending trigeminal root; and that these bundles course backward between the nucleus and root of the eighth nerve. This relation bears a striking resem- blance to that existing between the corpus, restiforme and the lower olivary body. Longitudinal bundles from the region of the funiculus lateralis are also connected with the superior olivary body; as is exhibited on longitudinal sections of the brain-trunk, these longitudinal bundles lie behind the layer of the lemniscus in which the lower olivary body is lodged. The lower portion of the corpus restiforme—that part belonging to the oblongata—is separated in the cerebellum, by the nucleus dentatus, from the other portion belonging to the pons. This lower restiform body, which is a prominent feature on the surface of the medulla, together with its stratum zonale, we described above as it edges its way be- 120 PsycJiiatry. tween the cerebellar and the auditory bundles into the section of the external auditory nucleus. The innermost cerebellar bundles, which pass down to the oblongata, descend far down in the com- pany of the anterior roots of the auditory nerve, through the area of the external auditory nucleus. They are situated to the inner side of the corpus restiforme (Fig. 49, Rsl., 8', R.). These bundles are evidently related to the auditory root. Examination reveals that these cerebellar bundles in the audi- • tory nucleus are part also of the auditory nerve-tract. On its course between its cerebellar origin and its central root, the audi- tory tract receives an addition in the form of the fibrae arcuatae. The fibrae arcuatae cross the median line, and undergo decus- sation between a large portion of auditory roots and the cere- bellum. Auditory bundles may indeed be divided into such which decussate in the oblongata and such which do not decussate in that organ. Crossed auditory bundles. a. External auditory bundles, striae medullares. They pass either directly to the point of decussation—the raphe,—in order to enter Clarke's nucleus (Fig. 49, 8') through the intervention of fibrae arcuatae, or they pass from the outer side into the cross-sec- tion of Clarke's nucleus (Figs. 50, 52, left), which, connecting with fibrae propriae, conduct the former to Clarke's nucleus of the opposite side (Fig. 52, right). The external auditory bundles surround the inner limiting bundles of Clarke's nucleus, and then divide up into posterior bundles, travelling through a part or the whole of the inner auditory nucleus, and as far as the raphe, and into anterior fibrae arcuatae which pass far front in the raphe (Fig. 49, right and left). In the upper levels of the oblongata almost all fibrae arcuatae constitute part of the auditory tract. These lend a medullary ap- pearance to the whole of the raphe, through which they pass, and the stratum zonale even seems to be derived from bundles coming from Clarke's nucleus (Fig. 49, se.). b. The inner crossed auditory bundles form in conjunction with cerebellar bundles cross-sections in the area of Clarke's nucleus. They descend to within half the height of the hypo- glossal triangle. Throughout this area these bundles are con- nected with fibrae arcuatae, y. The acoustic bundles, as well as the cerebellar bundles, after they have descended so far, may form either simple commissures with the ends of the fibrae Nervus Acusticus. 121 arcuatae, or they may in passing through the raphe connect the cerebellar bundles of one side with the acoustic bundles of the opposite side. Of these two possibilities, the latter is the more plausible. The existence of commissural fibres in the raphe has not been proved at all. In descending, the inner crossed acoustic tract extends from the region of origin of the facial, through that of the tenth, to the central origin of the twelfth nerve. The raphe, as well as the area of the acoustic arched fibres y, contains Fig. 49. Stained with Potassium and Gold Chloride. Section through the Level of Emergence of Auditory Nerve. Brain of a New-born Infant. (The Right Half Represents a Higher Level than the Left.) Bp. Brachium pontis. NC. Dentate nucleus of cerebellum surrounding the superior peduncles of the cerebellum. To the inner side of the radiation of the su- perior cerebellar peduncles into this gray lamina there lies a dentate ganglion, which I have termed the accessory nucleus ol the nucleus dentatus. v. IV. Fourth ventricle. 8. Inner auditory nucleus 8 ' .Externalauditory nucleus (Clarke's), fl + 8. Medullary substance of the flocculus and an external auditory root. On the left side the pedunculus flocculi is distinctly separated from the external auditory root. Rsl. Left corpus restiforme, with the inner acoustic root resting upon the median mar- gin. 5. Ascending trigeminal root. nclSa. Anterior auditory nucleus (Stilling). oli. Inferior olivary body. 12. Hypoglossal root. R. Raphe. £. Most anterior fibrae arcuatas. T. Eminentia; teres. ganglion cells which are connected with the network of fibres issuing from the above-mentioned motor-nuclei, and which Deiters supposes to be scattered ganglion cells belonging to the nuclei of the nerve-roots. The uncrossed acoustic bundles are the following: The inner acoustic bundles ' are connected with the cerebellum 1 I prefer to speak of acoustic bundles rather than of acoustic nerve-roots, for their value as nerve-roots is, as Kolliker has shown, as questionable as in the case of the optic nerve and the so-called olfactory nerves. 122 Psychiatry. directly by means of fibres which ascend toward the ganglion tecti, invading and covering the processus ad cerebrum (Fig. 46, right). There is a partial interruption of fibres through nerve cells on the external barren portion of Clarke's nucleus (Fig. 49 between 8' and Rs.l., left). Acoustic bundles bend over also directly into the corpus restiforme, and particularly so, near the lower edge of this body. Finally there is the anterior acoustic nucleus of Stilling lying between the inner acoustic bundles, the corpus restiforme and the Stained with Potassium Gold Chloride. Section through Oblongata of New-born. (Left Side Represents a Higher Level.) A. Internal auditory nucleus. 8'8. External auditory nucleus ; external and internal auditory root on left side. R, C. R. Corpus restiforme. v. Ascending trigeminal root, to the outer side of the gelatinous substance. The trigeminal section is traversed by glosso-pharyngeal roots, oi., oi. Olive and accessory olivary body. Hypo- glossal roots emanating from nuclei in the eminentia teres disappear in the olivary body. a. Fibras arcuatae from the nuclei of the tenth nerves. (These nuclei consti- tute the prominence between the eminentia teres (hypoglossal nucleus) and the inner auditory nucleus, and this prominence is very slight on the left.) (5. Fibne arcuatse from the ascending vago-glosso-pharyngeal root, y. Fibrse arcuata? from the external audi- tory nuclei. 6\ Fibrae arcuatoe from the post, column. Hst. Posterior column. flocculus (or the medullary substance of the cerebellum, Fig. 49, ncl8a). This nucleus receives those parts of these bundles which take an outward course, and then again it is unquestiona- bly connected with the medullary substance of the cerebellum. EXIT OF THE 9, IO, II, AND 12 PAIRS OF CEREBRAL NERVES. With the disappearance of the fasciculi of the pons the pyra- midal tracts lie immediately to the front of the anterior columns, from which they are not distinctly separable. The gray substance of this region exhibits three prominences: (1) the eminentia teres; Nerve Roots of Oblongata. 12 3 (2) the posterior column of origin of the ninth, tenth, and eleventh cerebral nerves—the ala cinerea—pneumogastric nucleus ; (3) the internal acoustic nucleus (Figs. 50 and 51). The eminentia teres contains heaps of nuclei from which circu- lar bundles emanate, which encircle their own cells of origin, and ultimately give rise to roots of the hypoglossal nerve (Figs. 50 and 51); but the superficial margin of the eminentia teres also con- tains smaller cells, through which definite portions of the root of Stained with Potass, and Gold Chloride. New-born. Transparent Section from the Oblongata. (Left Half Represents a Higher Level.) XII. Hypoglossal nuclei. 12. Hypoglossal roots. The lateral 12 exhibits fibres from the nucleus XII., which are scattered in among the gray substance of the funicu- lus lateralis (\,}). 8'8'. External auditory nuclei. Pst., list. Posterior columns. A. (ought to be further to the outside) Ala cinerea (nucleus of vagus). 8. (ought to be far- ther to the outside) Internal auditory nucleus. CR., CR. Corpora restiformia. g. Substantia gelatinosa surrounded by the ascending trigeminal root. On the left to the inner and outer side of g, the vagus roots. O., oi., oe. Olive, internal and external accessory olives, a. Fibrae arcuatae from the nucleus of vagus. /?. Fibrae arcuatae from the ascending vago-glosso-pharyngeal roots, y. From the external auditory nu- cleus. 8. From the posterior column, e. From the corp. restiforme. Z. Stratum zonale. the tenth nerve pass on their way to the raphe. These fibres pass around the hypoglossal cells in a circle presenting a posterior convexity. The joint nucleus of the ninth and tenth nerves, which disappears toward the inner side of the auditory nucleus in the plane of the rhomboid fossa, does not approach at all near the surface on sections at levels immediately below the striae transversae. On such sections it is merely a mass of gray sub- stance, which takes a brighter carmine staining than the neighbor- ing gray substance (" eminentia teres and auditory nucleus "), and gives rise, first of all, to roots of the glosso-pharyngeal nerve. At 124 Psychiatry. lower levels (Fig. 50, left) a small nucleus of the tenth nerve is perceptible. A very large part of this nerve at its origin seems on carmine preparations to form a groove about a number of small cells (Fig. 50, glosso-pharyngeal nucleus on the left). This section of medullary fibres represents not only an ascending root of the glosso-pharyngeal nerve, but also of the tenth nerve, from which delicate fibrils are unrolled throughout its entire length ; these delicate fibrils, as they are unrolled, mingling with small cells. This ascending root lies on the gray floor close to the trigeminal root (Fig. 50, right; Figs. 51, 52). The ascending trigeminal root has been justly compared to a posterior root of the spinal cord, and the hypoglossal nerve to an anterior spinal-cord root. The region of the vagus is accordingly situated between these two nerves, which are analogous to roots of the spinal cord. In- asmuch as it sends its fibres through the funiculus lateralis, it deserves to be termed the lateral system. In the funiculus lateralis we find a small cluster of cells, which, as it is analogous in position, should be considered the prolongation of the lower facial nucleus, and of the motor trigeminal nucleus ; it contains also large multipolar cells. This is the motor nucleus of the tenth nerve, or, as it has been termed, the anterior column of ori- gin of the lateral mixed system. The expression " lateral mixed system " was suggested by Deiters, and might be appropriately applied to the central nuclei of the 9th, 10th, and nth nerves. In reality the anterior central nucleus of the vagus is a nucleus of the accessory nerve, which passes downward into the gray reticu- lar lateral process of the anterior horn, from which the lowest roots of the accessory nerve emanate. Those bundles of this sys- tem which are analogous to the posterior roots of the spinal cord traverse the gelatinous substance of the trigeminal nerve, and are derived from the posterior column of origin of this system, from the ala cinerea (Fig. 51, 8). The ascending root arises on deeper levels, from fibrae arcuatae issuing from the raphe (Figs. 52, left fS). Roots of the vagus from the raphe course behind the hypoglossal nucleus as bun- dles of the eminentia teres, while bundles from the ala cinerea pass in front of the hypoglossal nuclei, past the posterior columns of origin, and directly into the root of the tenth nerve (a). Bundles from the anterior column of origin curve knee-shaped into the vagus, and from this nucleus roots appear to arise, which pass directly to the front (radial fibres, Lenhossek). (Fig. 51, left, in front of the fifth nerve). The region in which the motor nucleus of the tenth nerve and the hypoglossal nucleus are situated, though it appears to be a Cross-section of Oblongata. 125 centre of aggregated cells, has no distinct boundaries; and re- sembles the region of the facial nucleus (higher up) in having uniformly shaped cells scattered all around it. These cells, which contribute to the formation of a network of gray fibres, are traversed by arciform fibres, which can be traced into the bundles of the external auditory nucleus (Figs. 50, 51, 52—y). Fig. 52. Stained with Potassium and Gold Chloride. Transparent Section through the Medulla Oblongata of New-born Infant. 12 Hypoglossal nuclei. IO. Vagus nuclei. A., A. External auditory nuclei with auditory roots CR., CR. Restiform body. fp. Posterior column v. Trigeminal root g. Gelatinous substance. fl. Lateral columns. L. External bundles of lateral columns, on the inner side of which deep arciform fibres of the restiform body are situated. oe. External accessory olive. o. Olive. oi. Internal accessory o ive XII Twelfth nerve. P. Pyramid, fa. Funiculus anterior. CH. (left). Roots ot vagus, n.a. Region of the anterior nucleus of the vagus. Xr. Recurrent roots of vaaus. X. Roots of tenth nerve : a fibrae arcuatae from the vagus nuclei ; 0 from the joint ascending root ; y from the acoustic nuclei ; 8 from the posterior column ; and e from the restiform body. CROSS-SECTION OF THE OBLONGATA AND ITS TRANSITION INTO THE SPINAL CORD. On the outer side of the median-anterior columns and the pyramidal area of the oblongata, the lateral columns extend as far as the cross-section of the trigeminal nerve. They are dis- tinguished from the very dense and coarse fasciculi of the anterior 126 Psychiatry. columns by the lesser density and the lesser thickness of their fibres. The lateral column fills out the segment of the cross-section of the oblongata between the roots of the hypoglossal and trigeminal nerves. The funiculus lateralis encloses the lower olive, and is joined to it by horizontal bundles (Fig. 57, o). Immediately be- hind the lateral column lies the trigeminal root, and behind this, to the outside, lies the scythe-shaped section of the corp. restiforme (Figs. 45 and 50, R., Cr.), and in a median direction from the resti- form body lies the external auditory nucleus. This nucleus, under the name of Clarke's nucleus, forms the lateral boundary of the gray substance. The groove-shaped section of the trigeminal root surrounds a gray column, the substantia gelatinosa, which, as in tne spinal cord, forms an indentation or wave visible on distinct preparations. This formation is traversed by tranverse arciform fibres, which issue in part from the nerve-nuclei and in part from the corpus restiforme. 1. The most posterior (dorsal) of the fibrae arcuatae connect the nuclei of the vagus with the raphe (Figs. 50, 51, 52—a), and, in so doing, separate the hypoglossal root from its nucleus. 2. Then there follow fibres which have been unravelled from the ascending root of the ninth and tenth nerves (/?). 3. Fibrae arcuatae arise from bundles of the eighth nerve, which extend far down into the oblongata, and do not disappear until just above the closure of the central canal (y). The region of the arciform bundles issuing from the nerve origins is distinctly separate from the more anteriorly situated systems of fibrae arcuatae ; nor do the former traverse either the olives or inner olivary bodies. The corpus restiforme soon forfeits a number of its bundles, which go to the surface of the oblongata, there forming the stratum zonale, covering and invading the olivary body. Inspec- tion of the surface of the oblongata reveals the fact that the stratum zonale does not lie altogether in transverse planes, but that it descends obliquely. These bundles are situated on the outer side upon the trigeminal root. Deeper layers from the corpus restiforme send coarse bundles through the section of the fifth nerve, but invariably to the outside of the substantia gela- tinosa. Superficial bundles of the restiform body, which lie upon the fifth nerve, often continue the stratum zonale'iar enough to cover the pyramids. If such bundles unite on the cross-section of the pyramids with islands of gray substance, then we consider Origin of Spinal Cord. 127 them bundles of the pons which cross the corpus restiforme. In the lowest strata of the pons, we find bundles taking such a course, into the corpus restiforme and not into the brachium pontis, the two types of bundles separating distinctly in the cerebellum. On the other hand, the stratum zonale allows an external layer of the funiculus lateralis to touch the surface, surrounding the latter on its inner aspect (Fig. 52, left L.). Such bundles protruding behind the olivary bodies give rise to an external limiting fasiculus and in the same way longitudinal bundles on the inner side of the olivary bodies may be exposed and form the inner limiting fasicu- lus; (Fig. 15), owing to a deeper course of the stratum zonale. As a rule the pyramids remain free. The superficial layer of the corpus restiforme, the stratum zonale, not only covers, but actually crosses the olivary body of the same side. The deep layers of the restiform body take the same course through the olives and inner accessory olives (Figs. 50, 51, 52—s). The bundles of the restiform body cross in a transverse direc- tion through the raphe from one side to the other, emerging from the hilus of one olive and passing into the hilus of the other. These fibres cannot represent a true commissure of the olives, for the fibres have dissimilar (ungleichnamige) endings. From the opposite olivary body restiform bundles pass into a dense, co- lumnar1 mass of fibrae arcuatae, which are found to the inner side of the gelatinous substance, and instead of entering the corpus restiforme pass into the posterior column. On sections, which pass between the point of emergence of the ninth nerve and the region of the nucleus of the tenth nerve, a new formation is seen to edge in between the corpus restiforme and the region of the auditory root. This formation is the funicu- lus posterior of the oblongata, which passes into the posterior columns of the spinal column, through the funiculi graciles and cuneati (Figs. 50, 51, 52—Hst., Pst., fp.). The funiculus pos- terior has a net-like structure, the fibres of that net enclosing clusters of middle-sized ganglion cells. This formation stands in striking contrast to the external auditory nucleus, where distinct sections of bundles are imbedded in a floor of gray substance. In the one instance we find bundles imbedded in gray substance ; in the other, gray substance in a network of bundles. These posterior columns gather fibres from that set of fibrae arcuatae (Figs. 50, 51, 52—6) which are situated between those derived from This refers to the appearance on cross-sections. 128 Psychiatry. Clarke's auditory nucleus and those from the restiform body. The fibrae arcuatae in question also pass through the olivary body, and, as is worth mentioning, through the external secondary olive also. In the arciform course of those bundles which connect the pos- terior columns and the corpus restiforme, no cells are seen scattered in between these fibres, from which the posterior columns might be derived. It would be difficult, were it not for a single circum- stance, to decipher the origin of the bundles which go to make up the posterior column ; this fact is, that in proportion as the corpus restiforme diminishes in volume by yielding up fibrae arcuatae (Figs. 50, 51, 52—zone e), so the posterior columns develop by the ad- dition of the posterior fibrce arcuata? (Figs. 50, 51, 52—zone S). There can be but little doubt, therefore, that the corpora resti- formia, after passing the olivary bodies, pass into the posterior columns of the opposite side. In the olivary bodies the restiform bodies end, and the posterior columns begin. It is known of the olive (Deiters), that very coarse fibres of the corpus restiforme pass through it, and that very delicate bundles are distributed within it. The coarse restiform bundles simply pass through the oli- vary body of the same side, but terminate in the olive of the opposite side. Proof of this may be found in the fact that if one half of the cerebellum and the restiform body atrophy, the olivary body of the opposite side will also degenerate. The posterior columns once formed, the transition from the cross-section of the oblongata to that of the medulla spinalis is easily effected ; for the floor of the IV. ventricle changes into the gray substance surrounding the central canal. At such levels at which the corpus restiforme lies next to the beginnings of the posterior column, it is situated to the outer side of the latter. The posterior column forms on its inner side (Figs. 50-52), the substance of the restiform body diminishing on the outer side, and the posterior columns increasing on the inner side. In conse- quence of this mode of growth the region of the corpus restiforme and the funiculus posterior is pushed toward the median line, in the direction in which new growth takes place. The result of this is that in the adjoining halves of the gray floor upon which the posterior column encroaches after the disappearance of the acoustic nucleus, the nuclei of the tenth nerve occupy a more median position. The hypoglossal nucleus, the representative of the anterior spinal-cord nucleus, does not change its position, for the posterior column turns inward back of this nucleus. Cer- Origin of Spvial Cord. 129 tain roots of the tenth nerve intersect the gelatinous substance in the trigeminal nucleus, just as its continuation in the spinal cord is intersected by posterior roots. The nucleus and roots of the vagus are prototypes of the posterior horn of the spinal cord. The substantia gelatinosa represents the caput of the posterior horn; the vagus roots are its cervix, and the vagus nucleus, which is connected with the anterior horn (the hypoglossal nucleus) and turns toward the cervix the apex of a triangle, corresponds to the trigonum ccrvicale of Goll, the triangular section of the union of the Stained with Potassium and Gold Chloride. Cross-Section through the Oblongata of the New-born. c. Central canal. P. Pyramids. X. Decussating bundles, o, oi. Region of olives, g. Gelatinous substance, v. Trigeminal bundles, c. p. Caput of posterior horn. f. b. Fibrae arcuatae. c. a. Anterior horn. n. Nucleus of fasciculus cuneatus. fr. Fasciculus cuneatus. f. gr. Fasciculus gracilis. N. B.— The commissure back of c. does not exist in reality. posterior horn with the entire gray nucleus. This posterior horn of the oblongata lies, as it is, between the lateral and posterior columns, and there is a difference only in its being covered by trigeminal roots, and in the angle formed by the two posterior horns. The roots of the vagus, if prolonged, would intersect behind the oblongata, while the posterior roots of the spinal cord would intersect anteriorly. The former diverge anteriorly, and the latter posteriorly. The nucleus of the tenth nerve turns inward, approaches the median line, unites with its fellow of 130 Psychiatry. the opposite side, and thus closes the canalis centralis. The fossa rhomboidea has disappeared. The posterior columns crowd toward the median line, behind the united nuclei of the tenth nerve, leaving naught but the posterior fissure between them. But the entire posterior horn has followed the inward twisting of the vagus nuclei, so that the posterior horns, from which the I. and 2. cervical nerve-roots below the vagus arise, are no longer convergent posteriorly, but are juxtaposed at an angle of i8o° ; as soon as the fibrae arcuatae disappear, the posterior horns with the nerve-nuclei in the funiculi graciles and cuneati are separated less and less until they approach one an- other and converge anteri- orly. In consequence of the disappearance of the fibrae arcuatae and of the inner olive, the anterior columns become more compact; and this is true also of the lateral columns, which, instead of presenting a convexity, ex- hibit, as soon as the olivary bodies disappear, a flattened surface behind the pyramidal tracts (Fig. 53). Behind the lateral column, the trigeminal fibres which are no longer covered by fibrae arcuatae are replaced by the convex region of the substantia Rolandi (Fig. 54, V., SR.). Back of this the gray substance scattered in among the posterior columns, gives rise to two well-defined convex formations—the funiculus cuneatus, and funiculus gracilis (Fig. 53, fr., f. gr.), the latter lying immediately adjoining the posterior fissure. Before the total disap- pearance of the inner accessory olive, the lower boundary of which Stilling thought coincided with the lower limit of the hypoglos- sal roots, the posterior column receives additions from a decussa- tion of fibres between the anterior columns ; these fibres are Fig. 54. Carmine Preparation. Decussation of the Pyramids in the Adult. Hst. Funiculus posterior. C. Central Canal. V, V, SR. Caput cornu posterioris, substantia Rolandi, on the left side with vestiges of the ascending trigeminal root. LS. Lateral column. VS. Anterior column. P. P. Pyramids. Cross-Section of Spinal Cord. 131 stouter than those of the fibrae arcuatae which decussate higher up. Clarke and I thought they were derived from the external bundles of the pyramidal tracts. There is every reason to doubt the correctness of this view, for the inner accessory olive adjoins those bundles which take a transverse direction from the outside into the decussation. If they pass through this ganglion it would be natural to suppose that they were derived from the corpus restiforme. This view would not be contradicted by an inspec- tion of gold preparations (53, oi., X., f. gr.). Assuming this to be the case, they would be distinguished from the rest of the forma- tion of the posterior column in this, that the decussation would not consist of restiform bundles decussating at the hither side of the enclosed olive, but that it would consist of fibres which enter the funiculus gracilis, decussating at the further side of the enclosed, inner secondary olive. The entire pyramidal tracts are lost from the anterior fissure on through the great decussation of the pyramids ; they crowd the an- terior columns far from the raphe which they fill out and expand ; they separate the anterior horns from the gray substance, but how- ever much they may push the posterior horns backward, these pyramidal tracts still pass entirely and continuously into an area of the lateral column which lies next to the posterior horn, but does not extend to the external surface of the most posterior part of the lateral column. The position which the pyramidal bundles occupy after decussation as longitudinal fibres of the lateral columns, passing gradually into the anterior roots, they retain throughout the entire length of the spinal cord (Fig. 54, P. P.). CROSS-SECTION OF THE SPINAL CORD.1 We remarked above that the cross-section of the brain-trunk, at the lowest division of the mesencephalon exhibits three distinct masses of fibres, bearing downwards, and superimposed one upon the other. These were the tegmentum, the stratum intermedium, and the pes pedunculi. The cross-section thus constituted re- 1 Prof. Meynert's interpretation of the cerebral origin of the spinal-cord tracts dif- fers widely from that given by other authors. The reader who is especially interested .in this part of the subject is referred to the following publications : Flechsig : Leit- ungsbahnen ini Gehini u. Ruckenmarke ; Leipsic, 1876. p. 263, etc. Flechsig: Plan des Mensdilichen Gehirnes; Leipsic, 1S83. Alby : Schema des Faserverlaufes im Menschlichen Gehirn und Ruckenmark ; Bern, 18S3. Roller: Die Schleife. Arch, f mikroskopische Anatomie ; Bd. xix , p. 240, 138 r. Wernicke: Lehrbuch der Grhim- krankheilen; Kassel, 1881. Starr: Sensory Tract, JournalofNervous and Mental Dis- ease, July, 1SS4, Spiszka: On the Lemniscus, etc., New York Med. Record, 1884. Edin- ger: Zekn Vorlesungen uber den Bau der nervbsen Lentratorgane ; Leipsic, 1885.—S. 132 Psychiatry. ceived an increase in the oblongata, in the form of an addition from the cerebellum, which was joined to the other behind the continuation of the tegmentum. At this point there were four layers: corpus restiforme, or funiculus posterior; tegmentum, stratum intermedium, and pyramidal tracts. The formation of the spinal cord is the result of the union of bundles coming from the different centres of the cerebrum and cerebellum. Whether their wealth of fibres.increases or not, there Transparent Sagittal Section through the Brain of the Monkey. (More Lateral than Figs. 56 and 57.) Fr. Frontal end. Occ. Occipital extremity, c.d. Corpus callosum. n.l. Lenticular nucleus. nc. Caudate nucleus. Lp.f. (o) Lamina perforata anterior, ged.dept. pelt. Region of the pedunculus septi pellucidi. II. Optic tract., above it the optic commissure, ci. Internal capsule. Th. Optic thalamus. L.lf. Discus lentiformis. The dark (shaded) bundles, forming an < around its anterior end, constitute the radia- tion of the posterior longitudinal fasciculus. Between this formation and the thala- mus is the radiation of the nucleus ruber. Cp. Posterior commissure. Q. Corpus quadrigeminum in which the brachium corp. big. superius (bearing downward and back- ward), and the fillets (Lms 2, Lms 3, bearing downward and forward) are seen. The lowest white bundle is the cerebellar lemniscus (Lms 3) from the valvula cerebri. Behind Lms 3, the trigeminal root. ss. Soemmering's substance with its medullary sub- stance. P. Pes pedunculi. v. Pons Varolii. C.tr. Oblongata (corp. trapez.). R. Corp. restiforme. Cbl. Cerebellum, nd. Dentate nucleus of cerebellum. is no reason to suppose that these bundles would occupy any other position relatively to each other than they hold in the crus cerebri and in the oblongata. Inasmuch as continuity of the bundles can be studied best on sagittal sections, Figs. 55-57 are added as supplementary to the previous descriptions of the brain. Cross-Section of Spinal Cord. 133 The tegmentum is composed of four distinct sets of fasciculi of origin (Ursprungsbiindeln). 1. The fasciculus longitudinalis posterior does not seem to end altogether at the lower margin of the pons and in the oblongata. It might be traced to those posterior bundles of the anterior columns (on the spinal-cord section) which are situated in part immediately in front of the gray spinal commissure—the pro- longation of the gray floor of the brain,—and in part are crossed by the white bundles of the anterior commissure (Fig. 58). The radiation of the posterior longitudinal bundle from the cerebral cortex contributes a small number of fibres to the cross-section of the crus cerebri, and there is no doubt that in the brain-trunk its bundles are interrupted by masses of gray substance, each cerebral nerve-nucleus constituting an internodium. And yet the fasciculus longitudinalis posterior forms the most direct connection between the gray substance of the spinal cord and the cerebral cortex, for the gray internodia high up in the brain, such as the ganglion optician basalc, are equivalent to the central gray substance of the spinal cord (Figs. 18, 21, 31, 37, 40-44, 55>and S6- Cf. p. 96, etc.). 2. The bundles of the anterior pedicle of the thalamus, which, as laminae medullares, emerge like the rays of a fan from the thala- mus,and can be traced in the tegmentum to bundles situated to the outer side of the red nucleus (Figs. 29, 34-36, 38, 55. Cf. pp. 89, 91, etc.). 3. The fasciculus retrofiexus might be considered the continu- ation of that portion of the stratum zonale which participates in the formation of the ansa peduncularis ; it passes through the peduncle of the pineal gland into the posterior commissure. An ependymal connection with the posterior surface of the fornix might lead one to believe that there is a connection of the pineal peduncle with the posterior surface of the fornix. These bundles twist about in such a way that in the tegmentum they constitute the innermost anterior bundles, and in contrast to the posterior long, bundles participate in the formation of the anterior column of the tegmentum (Figs. 34, 35, 38, and a frag- ment in Fig. 56. Cf. p. 90). 4. The bundles from the posterior commissure, which enter the thalamus along its inner margin, after passing through the ansa peduncularis, and constitute its stilus anterior. These bundles and the fasciculus retrofiexus represent the crossed thalamic origin of the spinal cord (Figs. 17, 18, 31, and 57). The 134 Psychiatry. former bundles are well exhibited on Fig. 57, where they are seen entering the tegmentum ; according to Fig. 34 they lie behind the origin of the fasciculus retrofiexus, and by comparison with the brains of carnivora seem to me to constitute an opaquely bounded column lying in front of the gray substance and to the outside of the posterior longitudinal bundle (in the mesencephalon). This column is intersected by the fasciculi marginalcs aquaeducti Sylvii. On the supposition that it descends directly to the spinal cord we should have to look for this formation in the funiculus lateralis, next to the external margin of the anterior horn (Fig. Fig. 56. Transparent Longitudinal Section through the Brain of a Monkey. Fr. Frontal end. OCC. Occipital end. Rd. Cortex cerebri. M. Medullary substance of the fore-brain. Nc. Caudate nucleus. L. Lenticular nucleus. C. Anterior commissura. N. Globus pallidus. A. Amygdala. II. Optic tract. CL Internal capsule. Th. Thalamus. Br. Brachium corporis quadrigemini. Lb. Discus lenti- formis (by mistake of the engraver united to the stratum intermedium. 1 he dark- pointed triangular mass in front of it is the radiation of the posterior longitudinal fasciculus. Underneath Br. The radiation of the nucleus ruber). L, L, , LT]T. Lemnis- cus of the superior and inferior corpus bigeminum, and of the valvula cerebelli. P.P. Pes pedunculi. P. Pons Varolii. Rh. Corpus rhomboideum. O. Inferior olive. Cbl. Cerebellum. Pr. Processus cerebelli ad cerebrum. R. Corpus restiforme. Fp. Funiculus posterior. 5. The quadrigeminal origin of the spinal cord is embodied in the lemniscus, which can be pursued quite to the medulla spinalis on Figs. 55 and 56; particularly in the former. The divi- sions of the lemniscus constitute the most anterior layer of the funiculus lateralis, are crossed in or below the pons (according to the structure of the animal's brain) by the transverse bundles Origin of Spinal Cord 135 of the corpus rhomboideum, and in the oblongata enclose the lower olive, in the cells of which possibly some of their bundles termi- nate, unless they should alter their position and pass to the back of the olivary bodies. In the spinal cord the layer of the lemniscus would correspond to the external stratum of the lateral column (Fig. 58), and possibly to the most anterior layer of the anterior columns. The analogue of the quadrigeminal layer of the lemniscus would, in the spinal cord, not extend as far as the caput of the posterior horn (Figs. 17, 22, 40-45, 55, and 56. Cf. pp. 34, 102, 103 . Fig. 57- Sagittal Section through the Brain of a Monkey. The Most Median Section of Figs. 55, 56, or 57. Ca. Anterior commissure. II. Optic tract. Th. Thalamus. st. i. Internal pedicle of thalamus covering the ascending crus fornicis (white). cp. Posterior commissure. III. Third nerve. Between cp. and III. the tegmentum, on the back of which lie the roots of the fourth nerve. Pr. Decussation of the superior cerebellar peduncles. Between pons and third nerve innermost bundles of the crus, which run from the pes pedunculi into the posterior division of the pons, and are cut off on their course toward the median line. L. Stratum lemnisci. Rh. Fossa rhomboidea, in front of it the roots of the sixth nerve. 12. Hypoglossal roots, v. Pons Varolii; be- hind the transverse basilar bundles, the pyramidal tract, al. Behind the pyramidal tract we find the lemniscus passing through the olivary body, for lower down the sec- tion inclines strongly to the outside ; this lemniscal layer was traversed above by the roots of the sixth nerve and the transverse fibres of the corpus rhomboideum. Cbl. Cerebellum with the valvula ad. corp. quadrigemina. Fg. Funiculus gracilis. The gap left in the spinal cord by the quadrigeminal lemniscal layer is filled in by the cerebellar or lower lemniscus, which is derived from the vermis superior of the cerebellum, passes through 136 Psychiatry. the valvula cerebri, and joins the quadrigeminal lemniscus. From the manner in which the lemniscal layer advances, we can under- stand why this last addition should represent the most posterior (or, in the pons, the most external) bundles of the lemniscus; for in the cross-section of the lemniscus the quadrigeminal fibres are so arranged, that those originating highest are pushed farthest toward the median line, and the lower bundles join the lemniscus externally in the order of their descent. The lowest addition to the lemniscus would, therefore, form its most external bundle. But in the spinal cord the lemniscus cannot spread as readily in a transverse direction as it could in the crus cerebri and in the pons. Its bundles will necessarily lie behind one another, and the outermost bundle will become the hindermost (Fig. 58). Genetically, this portion of the external layer of the lateral column would have a distinct anterior boundary, for the spinal- cord mantle develops around the anterior and posterior horn in the form of an antero-lateral and a postero-.lateral column. The latter adjoins the posterior horn. The hindermost lemniscal layer undoubtedly forms in the spinal cord that portion of the postero- lateral column which constitutes the anterior boundary of the caput cornu posterioris (Figs. 17, 55, and 56. Cf. p. 34). 6. The stratum intermedium is the representative of the nu- cleus lenticularis in the spinal cord, though some of the crural bundles of the latter may have entered the pyramidal tracts. On sections through the crus cerebri the strata intermedia lay to the outside of the outermost bundles of the pes pedunculi, and were confluent in the median line. The stratum intermedium is composed of the ansa lenticularis, the innermost bundles of the pes pedunculi, and of other fibres of origin of the lenticular nucleus, which, after interweaving with the pes pedunculi, or the capsula interna, respectively, enter the lateral portions of Soemmering's substance. In the oblongata, this stratum lies in the same trans- verse position immediately behind the pyramids. Upon diminu- tion in the size of the stratum intermedium, followed a diminution of the angle between the two continued halves of this stratum (originally amounting to 180°), for these two halves, forming the posterior boundary of diminished pyramidal tracts, had to approach each other more closely. As the decussation increases, the angle grows more acute, and as soon as the last traces of the pyramidal tracts have disappeared, the two halves of the stratum intermedium lie immediately adjoining the anterior Cross-Section of Spinal Cord. 137 fissure of the spinal cord, there forming the innermost bundles of the anterior columns (Figs. 22, 23, 31, 34, 36, 37, and 58. Cf. pp. 45, 51, 54, 55, 80, 106, 107). 7. The pes pedunculi has a twofold origin: as the fasciculus of Arnold of the internal capsule from the cortex bf the fronto- parietal lobe, and as the fasciculus of Gratiolet, from the occipital and temporal lobes. Its cross-section area in the mesencephalon is so constituted that its innermost bundles belong to the stratum intermedium (ansa lenticularis), that its outermost bundles are those described by Gratiolet and myself, and that the largest mass between the parts referred to contains the remaining pyramidal bundles, and the medipeduncular1 bundles of the crus cerebri. In point of calibre, the lenticular portion is the small- est, the bundles of Gratiolet of far greater dimensions, while the last-named of the three divisions is the most powerful of all. In the spinal cord, the pyramida1 bundles are lodged to the front of the posterior horn, but do not extend either to the outer sur- face of the lateral columns corresponding to the fillet, or to the anterior portions of the posterior horn (Stilling, Flechsig). That portion of the spinal-cord section which immediately surrounds the gray substance from the anterior horn to the caput of the posterior horn, contains all the thalamic fibres of the spinal cord ; though the origin of some of these bundles may still be hypothetical. (Figs. 15, 17,22,23, 30, 31, 33, 34, 36, 37, 40,41 ; regarding bundles of Gratiolet, Fig. 33. Cf. pp. 27, 50, 81, 84, 85.) 8. The outermost area of the brain-trunk, the corpus resti- forme, is added to the oblongata from the cerebellum; it is derived from the laminated medullary substance immediately sur- rounding the nucleus dentatus, and passes through the stratum zonale, and through the mediation of the two most anterior sets of fibrce arcuatce into the posterior column. The term "funiculus cuneatus of Goll" may easily lead to confusion in spinal-cord terminology. This strand, which takes on a bright red color when stained with carmine, and in the foetal spinal cord is distinctly separate from the lateral portions of the funiculi cuneati in the oblongata, is the spinal continuation of the funiculus gracilis and not of the funiculus cuneatus. That the bundles of Gratiolet in the pyramidal tracts may participate in the formation of the posterior columns, or their inner portion at least, is not at all improbable (Figs. 15, 17, 22, 23, 45- 47, 52, and 53, Cf. pp. 28, 36, 119, 126). In Wilder's sense. ANATOMICAL COROLLARIES AND PHYSI- OLOGY OF CEREBRAL ARCHITECTURE. In the preceding section of this work we discussed the structure of the brain. We have now to consider a number of anatomical corollaries which will explain certain functions of this mechanism upon which physiological experiments have shed no light hitherto. Our understanding of those functions will be advanced, too, by close attention to the pathological anatomy of the organ under consideration. A mechanism may operate before us without our recognizing the exact relation between its function and its architecture. But, on the other hand, if we are acquainted with the principles upon which this mechanism operates, we may infer its function from its structure, regarding the former as the natural outcome of the latter. This method of reasoning would be applicable to the brain, even though the principles involved in its activity were entirely un- known. Moreover, matters are simplified very much in the case of the brain, for, without committing serious error, we may regard it as made up of a large number of entirely similar structural elements. In order to establish anatomical corollaries, wre need postulate but a single principle, abundantly proven by physiological experi- ments. This is Bell's law of the conduction of nerve-force in a centripetal direction through the posterior, and in a centrifugal direction through the anterior, spinal roots. We need not, how- ever, accept Joh. Miiller's opinion, that, at the very outset, differ- ent parts of the brain display different functional energies. A single functional energy only, though as inexplicable as all physi- ological forces, is inherent in the brain-cell, and that is Sensitive- ness. Actual sensation is developed by the evolution of equally unknown external forces, which we must suppose differ very materially from one another. These differences imply a difference in anatomical structures, not of the brain, but of the terminal organs of peripheral nerves. T38 Relation of Cortex to Outer World. 139 This is well illustrated in the case of the anterior roots, which are motor in function, simply because the terminal organs of nerves derived from the anterior roots are connected with muscles. The latter alone are motor elements. Nerves and nerve-cells possess no motor power. Indeed, there is nothing more certain about the functions of the cerebral organism than that the centripetal sensory nerves are the keys which wind up the mechanism con- nected with the muscles, and excite the latter to action. A varying functional energy of brain-cells, according to the special organ of sense with which they may be connected, is quite indemonstrable, since we are acquainted with the physiological conditions favorable to the action of external forces, and can prove easily enough that it devolves upon the terminal organs of the nerves to meet these conditions. Even if the auditory nerves possessed a specific visual energy, the media between the auditory nerve and the waves of light would be totally unfit to transmit visual rays. Specific energies therefore depend altogether upon the peculi- arities of the end-organs, and sensitiveness is the only specific property of brain-cells. Within the fore-brain sensitiveness is converted into actual sensation. The relation of the fore-brain to the other parts of the cerebral mechanism is easily understood. To this end we may recall the structure of the retina, which constitutes a hollow into which the visual rays from the external world are, as it were, entrapped. And, in the same way, we may look upon each half of the cortex of the fore-brain as a concave organ, duplicated in parts, enveloping all the nerve tracts, which conduct to it the impressions from the outer world. In this organ these impressions are converted into the phenomena of sensation. In assimilating totally unknown physi- cal impulses, the cerebral cortex, a complicated protoplasmic structure, resembles the protoplasm of the primitive amoeba, which can transform itself into a hollow mass, and can thus encircle any hody which it wishes to assimilate. Just as the mollusca possess tentacles which they protrude toward the outer world, and claws by means of which they take possession of their booty, so this complicated protoplasmic organism, the prosen- cephalic cortex, possesses centripetally-conducting processes,— the sensory fibres of the nervous system—which we may con- sider its tentacles, and motor fibres, which are its claws. The 140 Psychiatry. remainder of the body, with its sensitive surfaces, its muscles, and the skeleton to which these muscles are attached, serves to sus- tain these tentacles and claws, which enable the fore-brain to re- ceive the images of the external world, and to react upon the latter. Comparing the disposition of the gray substance in the spinal cord with its arrangement in the cortex, we observe that the gray substance in the spinal cord is very much crowded by the aggregation of medullary strands on its periphery. Nerve-ele- ments corresponding to tentacles and claws are thus closely united, and the functional result of this union is exhibited in the reflexes which the spinai cord necessarily develops, as soon as those higher cerebral structures which have an inhibiting influ- ence upon its vital activities have been removed. In the fore- brain, on the other hand, the gray substance is not lodged in the white, but the white substance in the gray, and the former pushes the cortex asunder. Knowing the difficulties of nerve-conduction through the gray network of fibres, we may infer that this en- larged surface will be able to perform a number of totally inde- pendent functions; that a sensory perception, for instance, need not give rise immediately to a motor act. Every spinal-cord segment embraces the whole of the gray substance, whereas sections through the cortex contain but a small portion of the cortical gray. This distribution of gray substance will naturally pre- vent the entire cortex from acting to one single end, while it favors the isolated action of various cortical regions. Irradiation of functions is facilitated in the gray substance of the spinal cord, and rendered difficult in that of the cortex. We learned, further- more, that the cortical structure was not the same throughout; that the arrangement of the cortical elements varied in different divisions of the cortex. Purely morphological data arid a single pathological anatomical fact will enable us to determine which re- gions of the cortex, in the probable division of labor, take upon themselves centrifugal functions in the sense implied in Bell's law. The pathological fact referred to is the hemiplegia resulting from destruction of the prosencephalic ganglia, and particularly from destruction of. the nucleus lenticularis. These ganglionic masses divide into a club-shaped body (nucleus caudatus), and a wedge- shaped body (nucleus lenticularis), both presenting their broadest surfaces anteriorly (cephalad). Upon the number of cortical cells, depends the number of radiating fibres which the ganglia receive ; accordingly the anterior portions of the prosencephalic ganglion Proof of Cortical Localization. 141 must contain by far the largest number of cells for the reception of these radiating fibres. We may therefore establish this morpho- logical corollary, that the cells of these ganglia will be able to re- ceive more fibres from the anterior cortical regions than from the posterior ventral regions of the hemispheres, for these ganglionic masses, tapering as they do toward this latter region, will naturally have a lesser number of cells to put at the disposal of fibres coming from that direction. Besides, Gratiolet and I observed that the oc- cipital and temporal cortex received fibres from those ganglia with which the optic tract is connected. And into these latter regions of the cortex those bundles of the internal capsule enter, which, when implicated in a lesion, give rise to hemianaesthesia (Tiirck). It was well known also that the anterior commissure, which was sup- posed to be an olfactory chiasm, was not connected with the cortex of the frontal lobe, but according to Arnold with those of the temporal lobe ; Burdach and myself insisting on a further con- nection with the cortex of the occipital lobe. This separation of sensory and motor areas in the cortex was later on substantiated by the results of physiological experiments. Before discussing these physiological experiments, I wish to call attention to three anatomical facts which render a functional differentiation of the various cortical regions highly probable. The first of these facts is taken from comparative anatomy, and refers to the enormous difference in development of the olfactory lobe in different animals. Animals that are accustomed to run their noses close to the ground, and to obtain their food by following the scents they perceive, are characterized by a highly developed olfactory lobe, as is shown in Figs. 7 and 10. In man, whose erect posture has lessened such sense-impressions, as well as in the monkey and in all climbing animals, the olfactory lobe has deteriorated very much. Among water mammals, the sea- lion, which spends part of its life on land, has an olfactory lobe about equal in dimensions to that of man, while cetaceans have no olfactory lobe at all. The second fact refers to the difference in the relation between the median and the convex surfaces of the cortex, in animals with strongly developed olfactory lobe, and in man. In the latter, the convolutions of the convex surface everywhere overtop the gyrus fornicatus ; whereas, in the former category of animals, this same gyrus, which is connected with the olfactory lobe, is enormously developed, forming, together with the external olfactory convolu- 142 Psychiatry. tion, the gyrus uncinatus of the convexity which continues the formation of the cornu ammonis to the middle of the corpus callosum ; while in man this formation but barely grazes the pos- terior surface of the corp. callosum. Thirdly. Among the convoluted regions of the convexity of the human brain, the walls of the Sylvian fissure are most highly developed. This includes its floor, the island of Reil, as well as the operculum covering the island from above, and the first tem- poral convolution covering it from below, and also the transitional convolution, together with the posterior margin of the orbital surface closing, in upon the anterior border of the island. In keeping with these peculiarities of structure, the human brain exhibits the most extensive claustrum. But I have shown that the disturbances of psychical speech, which are classed under aphasia and its allied conditions, depend upon lesions in the claus- trum—in general terms, upon lesions in the walls of the Sylvian fossa. Man as far excels in the development of these psychical regions of speech, and in the number of convolutions belonging to these regions, as animals with highly developed olfactory lobes excel in regard to the size of these lobes. From this we may unquestionably infer that evolution of certain psychical functions will go hand in hand with a proportionate development of certain regions of the cortex. We may as well add at once that there are quantitative dif- ferences in the brain-trunk, both in man and in animals, dependent upon quantitative variations in the different parts of the fore- brain. An inspection of the basilar surface of the cerebrum confirms this view. The ideas suggested by such considerations as these were referred to on page 82, where it was shown that, in keeping with the quantitative development of the fore-brain, those structures, which are connected with its centrifugal tracts, such as the crus cerebri (excluding the tegmentum), the pons, and the pyramidal tracts of the medulla oblongata,' are the most powerful formations in the human brain ; there seems, therefore, to be a harmonious dependence between the form of the brain- trunk-structures and the quantitative development of the fore- brain. These well-ascertained, though general and incomplete ana- tomical facts, would justify us, from a purely morphological point of view, in affirming the localization of cerebral functions. Having obtained this firm and safe anatomical foothold, it will Critical Review of JMotor "Areas." 143 be especially interesting to examine those from among the many contradictory physiological experiments which confirm the view of a localization of functions in the cortex of the brain. This question has been studied in all its details by different physiologists and in different ways. The older brain-physiologists asserted that direct irritation of the cortex produced no effect; but Hitzig,1 by means of electrical irritation, and Nothnagel,2 by the use of mechanical irritants (principally pricking with needles), suc- ceeded in proving that irritation of certain regions of the convexity produced movements in the opposite side of the body. Ferrier,3 too, has furnished many details. It would be a difficult task to give a critical review of all these experiments, which agree with each other in principle only, and not in the dimensions of the psycho-mo- tor, functional areas governing special groups of muscles. Nothnagel operated on the rabbit, and could not make out as large a number of centres as Plitzig determined, though Nothnagel's centre for the anterior extremity occupies the same position as the centre to which Hitzig has ascribed similar functions. On the other hand, Nothnagel has described another possibly motor (leap) centre, which Hitzig cannot corroborate ; and the difficulties of a critical review are increased still more by the fact that Hitzig, experimenting on monkeys, wishes to limit all centres to the gyrus /nrcentralis, in flat contradiction of his description of the motor areas in dogs, in which the facial centres, as well as those for the straight ocular muscles, are situated posteriorly to the fissura centralis. No one will be disposed to agree with Hitzig's complicated interpretation of the gyrus praecentralis in the dog, and yet, in spite of the artificial expansion of the gyrus praecentralis over several other gyri, we may locate the centre for the muscles of the neck and trunk in front of the gyrus praecentralis, as Hitzig defines this convolution, and as was explained on page 18 of the previous section. No mention is made of this particular motor area in the brain of the monkey. Ferrier's " centres " cover the largest area, extending from the middle frontal con- volution over and beyond both central convolutions as far as the occipital fissure, and including the first temporal convolution. By way of anticipation, I will remark that Munk also looks upon a line drawn from the posterior end of the Sylvian fissure to the margin of the hemisphere as the posterior boundary of the motor area of the cortex. The radiating fibres which we observe entering the nucleus lenticularis and capsula interna, on longitudinal sections of the brain (Fig. 30), would seem to be radiations from that portion of the cortex which lies anterior to and including the region of the pos- terior central convolution, into these centrifugal conducting tracts. Hitzig's centres are located principally in the frontal lobe, although he has made an entirely gratuitous change, as was shown in the previous chapter, in making the gyrus pracentralis part of the parietal region. 1 Fritsch u. Hitzig: " Ueber die elektrische Erregbarkeit des Grosshirns, DuBois-Reymond's Archiv, 1870. Hitzig : Untersuchungen zur Physiologie des Gehirns, ibid., 1873. " Untersuchungen des Gehirns," Berlin, 1874. 3 Virchow''s Archiv, vols. 57 and 58. 9 " Experimental Researches in Cerebral Physiology," 1873 ; cf. also "Functions of the Brain," Am. Ed., 1876. 144 Psychiatry. Hitzig has mapped out the centres in the praecentral convo- lution in the following order of succession from above downward : (i) Centre for the posterior extremity ; (2) for the anterior ex- tremity; (3) for the facial nerve; (4) for the movements of mastication. The disagreement among authors in regard to the topographical distribution of psycho-motor centres in the cortex makes it incumbent upon us to examine into the nature of those motor disturbances upon which all are. agreed. In parenthesis be it said, that Ferrier's " centres " have met with opposition from all other authors. Hitzig believes that the motor disturbance in the anterior extremities, consequent upon an excision of cortical substance, is to be explained onthe ground that the animal has lost the muscu- lar sense which informed it of the position of its extremities. An animal, thus operated, will allow its foot to be placed in the most uncomfortable position, with the dorsal surface to the ground, and so on, and yet is able to use the anterior extremity well enough in response to volitional impulses. Nothnagel is inclined to the opinion that ataxic movements result from the lesion of the so-called psycho-motor areas of the cortex, agreeing with Leyden, who attributes ataxia (i. e., the condition consequent upon the disease of the posterior gray col- umns of the spinal cord, which effects irregularities in lifting, and in the forward and lateral movements of the foot) to an inter- ruption in the nerve-tracts issuing from the brain. The present author believes he was the first one (in the pamphlet entitled " On the Twofold Cerebral Origin of the Spinal Cord," 1869) to insist that the processes of innervation from the hemispheres, which constitute what we term volitional acts, are nothing more than the perception or memory of the sensations of innervation; for such a sense of innervation accompanies each reflex act, and is registered in the cortex ; there this same sense of innervation serves as a fundamental basis for similar secondary movements produced by excitation of the fore- brain. By association, these memories {Erinnerungsbilder) acquire suffi- cient intensity to excite secondary movements starting from the fore-brain and passing along centrifugal nerve-tracts. I shall refer to this subject again later on. Munk ' has given a very lucid account, based upon experi- 1 Ueber die Functionen der Grosshirnrinde. Berlin, 1SS1. Sensory Character of 'Motor Centres. 145 ments, of the relation of the cortex to the movements of the body. Munk subscribes to my views, believing that in order to explain conscious movements—volition, in psychological phraseology,—it is sufficient to postulate sensations of innervation. Munk dis- tributes sensory areas over quite the whole of the cortical surface, and therefore terms the region from the occipital lobe to very near the frontal margin (in the monkey) the "sensory sphere " of the brain. The character of these sensations he described as tactile sensations, sensation of pressure (the knowledge of which is not enhanced by experiment), and as sensations of innervation. The tactile and pressure sensations are thought to be a means of regulating the excitation of sensations of innervation. Spiess and Lotze entertained the same opinion long ago, but were too one- sided in supposing that the " muscular sense " depended simply upon the perceptions of touch and pressure by any part of the skin and the joints. Thus we see that an experimental investigator reaches the conclusion, that sensitiveness is the only specific energy common to ganglion cells ; and we are led to infer a localization of sense- impressions only, which, in keeping with the above views, would be synonymous with the localization of so-called motor areas of the cortex. In corroboration of the conclusions of Hitzig, who observed that blindness ensued upon the removal of a part of the occipi- tal cortex, Munk finds that removing a piece (15 mm. broad and 2 mm. thick) of the same region gives rise to what he calls "psychical blindness." Removing another piece of about the same dimensions, from the temporal cortex, caused symptoms to which Munk gives the name of " pyschical deafness." The dog which has undergone the former of the two opera- tions, will show the following peculiarities, provided the healthy eye 011 the side of the lesion be properly bandaged. The dog, so conditioned, " no longer scents for his food as he used to do in the accustomed corners of the room, and if both the mess-pot and the pail of water be placed in his way he will go round about them again and again without taking notice of them. He takes no notice of his food until he smells it ; neither a finger nor a flame brought close to his eye moves him ; and the sight of the whip, which used to drive him into the corner, does not intimidate him in the least. He had been trained when the hand was passed by his eye to give his paw; now he does not respond to 146 Psychiatry. that movement of the hand until he hears "paw" ; and other like observations are to be made. But such a dog can again learn to see as in his earlier days, and to regulate his action according to his visual images; he is, therefore, mentally blind—not organi- cally blind; he has lost the visual images registered in his cortex previous to the operation. This phenomenon, the revival of pre- viously registered images, Munk explains by the fact that the visual area is larger than the part extirpated, and that up to the time of excision the entire visual area was not filled with visual images, but that after the removal of the part formerly entrusted with the care of the visual images, another part, connected also with the retina, assumes the duty of harboring new visual images. If the entire occipital cortex be removed, cortical blindness will ensue ; the animal becomes totally blind, although the subcorti- cal visual centres are intact. We are, therefore concerned with the loss of cortical functions only. More recently Munk's " visual sphere" has been extended considerably, so as to include in the monkey the whole of the well-marked occipital lobe, including its median surface. Munk states, that after psychical blindness has disappeared, in the course of four to six weeks, the dog does not pounce directly upon meat which is placed before him, that he cannot snatch it until it has become distinctly visible by appropriate movements of the head. He draws the inference that after the extirpation of the cortical centre, a new blind spot is created on the retina, for a definite portion of the retinal elements is no longer connected with the cortical cells upon which retinal impressions were registered ; and he contends, furthermore, that the arrangement of the cortical elements repeats the exact distribution of the retinal elements around the spot of clearest vision. The psychical deafness resulting from the removal of a por- tion of the temporal lobe, brings about the following changes in the behavior of the dog : " The dog has retained the power of hearing, every unusual noise exciting an equal pricking of both ears, but he cannot in- terpret what he hears. The meaning of ' pet,' ' come,' ' up,' 'there now,' ' paw,' and all calls to which he was accustomed to respond, he does not understand ; so that he no longer per- forms movements which at one time had the value of reflex acts." Gradually the dog learns to hear again, and after four or five weeks behaves about as he did before the operation. In this Distribution of Different Areas over Cortex. 147 instance also Munk assumes the existence of a peripheral auditory area around the part extirpated ; this peripheral area now receiv- ing and storing up the newly acquired impressions. Removing this peripheral area and leaving the concentric inner sphere of the auditory area, produces no effect whatever; for no auditory images had as yet been stored in the former (peripheral) area, as was the case also with regard to the visual centre. This theory would explain not only the re-acquisition of facts which had once been familiar, but also every increase of knowledge effected by the mediation of cerebral cells which up to that time had not been functionally employed. To this literal quotation of Munk's views, I wish to add that they give a plausible explanation of mental (psychical) deafness only. The introduction of projections of the macula lutea into an inner area of the " visual sphere " must necessarily lead to in- calculable confusion. This re-acquisition of facts is simply the result of newly-registered images which are lodged in the environs of the parts removed. According to Munk's theory we should have to believe that these later impressions were indistinct, because they did not happen to be registered within the cortical (projec- tion) area of the macula lutea. Munk thinks perception and memory are identical. We, hope to prove later on that a cortical image has no material background (sinnliche Klarheit). Even darkness leaves its image upon the cortex. The term " psychical blindness " would suffice, and from this, cortical blindness would differ simply in being an incurable state. There is no reason compelling us to assume that tactile and pressure-sensations, because of their relation to the muscu- lar sense, are registered in the same parts of the,cortex as are the sensations of innervation: first, because the external fasciculi (of Tiirck) of the crus cerebri, which carry tactile impresssions, are not united to one another throughout their entire course; and secondly, because association-fibres effect the co-operation of widely separated cortical districts. Referring once more to that portion of the sensory sphere, (Fiihlsphare) which was called the psycho-motor centre until Munk called it the centre of sensations of innervation, Ave find that this author ascribed limits to this sphere in the brain of a dog, which, if followed in the brain of a bear (Fig. 7), would,leave a functionally useless region occupying the convex surface between the two parietal convolutions, and a portion of the marginal convolution 148 Psychiatry above. The sensory sphere would be situated cephalad, the auditory and visual areas caudad, of this functionless area. Between this boundary and the olfactory lobe there lie, one above the other, the centres for the hind limb, for the fore limb, and for the head. In a later publication (December, 1878), Munk has added to the number of sensory areas in which the sen- sations of innervation are registered. These newly added areas are an eye-region on the summit of the second parietal arch of the dog, governing the protecting and motor apparatus of the eye ; and an ear-region on the summit of the first parietal arch, removal of which disturbs or annuls the movements and sensibility of the concha auris. The head-region, which he subdivides into centres for the tongue, facial nerve, etc., Munk distributes over the whole operculum as far as the anterior branch of the parietal arch. The centres for the fore- and hind-limbs lie in front of the sulcus centralis, and together do not occupy an area larger than the " head-region." The centre for the hind-limb lies nearer to the margin of the hemisphere. The extremities being repre- sented in the frontal lobe, Munk locates the neck-region still farther to the front, as Hitzig did also, in that part of the frontal lobe lying in front of Leuret's transverse fissure; while he relegates the trunk-centre to the most anterior portion of the frontal lobe, resting upon the lobus olfactorius. With the excep- tion of the gyrus fornicatus and the olfactory lobe, a correspond- ing portion of the median surface, which begins in man in front of the occipital fissure, is appropriated to the domain of the sensory sphere, including the various muscular areas. (" Ver- handlungen der physiologischen Gesellschaft," Berlin, 1878.) In the monkey (Fig. 8) the occipital lobe constitutes the visual sphere ; the temporal lobe presides over auditory functions, as has 'been repeated often enough. Within the sensory sphere we have the eye-centre on the posterior (superior) parietal arch, and the ear- centre on the anterior (inferior) parietal arch. The head-region ex- tends from the margin of the operculum upward as far as beyond the lower portion of the sulcus centralis, and anteriorly it extends as far as the sulcus praecentralis, in the concavity of which the neck- region is lodged. Above it and partly behind it lies the centre for the hind-limb, along the posterior margin as far as the sulcus occ. ext. The sensory sphere of the trunk is situated in the frontal lobe, in front of the sulcus praecentralis and in the posterior por- tion of the orbital surface. The Fore-Brain the Seat of Intelligence. 149 In keeping with his visual and auditory areas Munk estab- lishes consistently enough a psychical paralysis after the removal of the so-called motor centres. This psychical paralysis, which is curable, is the result of the destruction of a small amount of cqr- tical motor substance ; but if the peripheral as well as central area of a motor centre be entirely destroyed, then permanent cortical' paralysis will be established. Munk's sensory centres do not respect definite, prescribed boun- daries, such as the convolutions would present; but there is this in favor of his views, that he rriakes the entire frontal lobe do serviec as a sensory sphere. Other authors, and among them Hitzig, who has misinterpreted the results of his own experiments, have pro- nounced the frontal region to be the exclusive seat of intelligence. On this head Munk justly observes: " Intelligence is located everywhere in the cerebral cortex and nowhere in particular." I wish to add in corroboration of this view, that no author of the present day would be likely to insist on one special seat of memory, for memory is the common property of all cortical cells and fibres which are able to receive and conduct external stimuli of all sorts. Consciousness and intelligence also, which is evolved in the fore- brain, depend upon a mechanism, the minute details of which enable us to understand the restriction of intelligence to the fore-brain. This discussion, which we now take up, shall be guided by the fundamental experiments of physiologists, and by certain morphological views which I published years ago (Leidesdorf, " Lehrbuch der Psychiatrie," p. 45, et seq.—1865). There may be a difference of opinion among physiologists regarding the localization of definite impressions and functions in the cortex of the fore-brain, and their distribution among the med- ullary fibres of the brain ; but there can be no question about this, that the intelligence of an animal is seriously impaired (obliterated) by the extirpation of its fore-brain. Those who removed the fore-brain with the knife, who destroyed it by freezing, or who, like Goltz, trephined the skull and washed out the fore-brain with a stream of water, all are agreed on this one point. Goltz (Ueber die Verrichtungen des Grosshirns, in Pflugers Archiv, vol. xiii. and xiv.) presents the largest number of striking and critical observations, although he removed less of the fore-brain than 'The German text has Rindenldhmung. I have translated it cortical paralysis. The term is misleading, however: for the " psychical " paralysis is also based upon changes in the cortex. It would be better to speak in the one instance of " psychical " paralysis, and in the other of " absolute " or " complete " paralysis.—S. 150 Psychiatry. others, for he could wash out' only about as much of the cortical substance as is visible after removing the roof of the skull. But on this very account Goltz has supplied facts which support the view of the localization of sensory areas on the cerebral surface. He observed a diminution of cutaneous sensibility, of the sensa- tions of innervation due to ataxia and impairment of sight; but having left intact the base, and the median surface containing the olfactory area, and the basilar portions of the temporal lobe con- taining Munk's and Ferrier's auditory areas, the senses of smell and hearing were in no way affected. The mere loss of a few grammes of cerebral substance suffices, according to Goltz, an extraordi- narily keen observer, to produce a degree of idiocy which he recognizes from the appearance and gaze of the mutilated dog. Idiocy develops in these dogs to such an extent, that they stumble into their meat-pots, bite their own legs, and, not remem- bering this experience, repeat the very same thing from day to day. Not knowing their whereabouts in space, they wander about the whole room in answer to their master's call; nor are they able to' find with their snouts any portion of their skin which has been teased by serres fines, but simply run about restlessly and helplessly in con- sequence of the pain ; they are not bright enough to discover or to lift up their whining pups, though they be very near them. Apart from the difference in the arrangement of its nervous cortical elements (vid. p. 56, et seq.), the white substance of the hemi- spheres has everywhere the same structure ; and we have shown that each hemisphere consists : (1) of projection-systems, which, with the aid of gray internodia interspersed among the medullary fibres, connect the cortex with sensitive surfaces and motor or- gans ; (2) it (the white substance) consists of association-systems— i. e., of arciform nerve-bundles. The wealth of such fibres, and their variation in length, connecting as they do near and remote parts of the cortex, will suffice, without formulating an anatomi- cal hypothesis, to unite any one part of the cortex to any other. The phenomenon of mental blindness is exhibited in the case of the animal that no longer associates the sight of the whip with the idea of punishment, or in that other animal (of Goltz), from which the right hemisphere had been removed, and which, when its right, healthy eye was covered, took no notice of a servant wearing a mask and decorated with red rags; but as soon as its 1 Goltz has long since abandoned this method of removing cortical substance. (For his present method see Pflilgers Archiv, Sept. 17, 1881.) S. Memory—A Cortical Function. 151 healthy eye (connected with the left hemisphere) was uncovered, pounced upon this fantastic figure, as every normal dog would do. If an animal has become psychically blind, it will recognize the sound of the whip but not the visual image. We may note, therefore, that there is a localization of certain intellectual activities dependent upon, and coinciding with the localization of definite sensory areas. Longet was among the first to contend that a dog which had forfeited its hemispheres could be made to swallow colocynth, show displeasure, and per- form all the retching movements which a disagreeable or bitter taste naturally excites ; we must therefore assume the existence of sensory perception entirely independent of the fore-brain. Schiff, who has given further details regarding these experi- ments- of painting the tongue with colocynth, calls attention to the fact that he can again and again open the mouth of the ani- mal, show it the instrument of torture (the brush), without elicit- ing the slightest repulsive movement; whereas young kittens give indubitable facial expression of the disagreeable gustatory impression they have received. Sensory impressions are there- fore not wanting; but the recollection of previous pain, the recognition of movements preparatory to the infliction of pain, so-called volitional impulses which would induce the animal to escape, all these are wanting. In order not to be dealing with intelligence in the abstract, let us determine by the simplest scientific analysis the contents of or, at least, a demonstrable factor of intelligence; and with this let us compare the brain-mechanism. We shall thus be able to con- clude whether the cerebral mechanism can, or can not, account for the manifestations of intelligence which existed prior to the impairment of the mechanism. First of all, animals deprived of certain cortical areas (Munk) or of the entire fore-brain, show no recognition of former impres- sions; the dog does not recognize the call of his master, nor the cat the preparations which are being made to insult her gusta- tory nerves. This non-recognition constitutes loss cf memory, and loss of that special memory which is based upon the juxta- position of successively received impressions. This is memory of the fore-brain, and must be regarded as a cortical function. It is not part of our present purpose to dilate upon other phases of memory, such, for instance, as affect the entire or- ganism, and are independent of the functions of the fore-brain, 15 2 Psychiatry. though related to other parts of the nervous system. Among these would be classed those phases of memory which are mani- fested by the spinal cord in repulsive movements, the exact nature of which is varied, knowingly as it were, according to the character of a painful, exciting stimulus. Lotze regards this phenomenon as an after-effect of oft- repeated innervations of motor spinal nerves, responding to definite stimuli reaching the hemispheres. Later on we shall refer in detail to this view, to whick Brticke also subcribes. The spinal cord of the new-born infant would not exhibit such phe- nomena. As long ago as 1867 I endeavored, in an essay entitled " Ueber den Bau der Rinde und seiner ortlichen Verschieden- heiten," to demonstrate the fitness of the cerebral cortex to be the seat of memory, by showing that it contained more than a milliard nerve-cells which could serve as the functional posts of succes- sively received impressions; while in the retina, in which after- images occur as the result of protracted normal stimuli, these images are not permanently lodged ; for every succeeding series of images occupies the retina in its totality, and finally annuls the after-effects of former images. Munk's experiments have verified this assumption in regard to the nature of cortical mem- ory. They have demonstrated that animals possess a function- ally unoccupied region in the vicinity of the visual and auditory spheres, the removal of which (region) produces no effect in regard to psychical blindness, and psychical deafness; but if the central functional portions of these spheres be destroyed, these peripheral areas will be the recipients of new visual and auditory impressions. From this it follows that in the normal (physiological) course of events, more and more of the cortex is called into requisition to receive new impressions, and that upon the increase in the number of registered images will, depend the enlargement of the child's mental sphere. It is very probable that the number of cortical cells fixes the limits of memory—the foundation of all intellectual activity. The recollection of former experiences, of which the animal that has forfeited its fore-brain is incapable, implies the association of one phenomenon with some other. The dog, for instance, would associate the sound of the whip with the bodily pains it inflicted. In examining the structure of the hemispheres, and remembering that different, distinctly limited and functionally separated por- tions of the cortex receive impressions from the various senses,' The Induction-AIechanism in the Brain. 153 we may naturally infer that the association-bundles, the fibrae proprice of the cortex, which form anatomical connections between the different cortical regions, effect the physiological associations of the images which are stored in these various parts; these images implying excitation of the cells of these parts. Since these medullary bundles are beyond a doubt conducting nerve-tracts, this interpretation is not a mere hypothesis. But recollection constitutes a process of induction (Schluss- process). Wundt was justified in calling an induction the fundamental logical function (vide his lectures " Ueber Menschen und Thier- seele "). I was the first one to demonstrate the association and in- duction mechanism of the fore-brain, in my pamphlet on the " An- atomie der Hirnrinde alsTragerdes Vorstellungslebensund deren Verbindungsbahnen, 1865," which was reprinted verbatim in Leidesdorf's " Manual." Both ends of the association-fibres are connected centrally with cortical cells ; the projection-bundles (vide pp. 42, 43), consisting chiefly of fibres of the corona radiata, spreading into the medullary substance of the fore-brain, conduct to the cortex the excitations from the external world, and distrib- ute them over its different sensory spheres. All objects which, as soon as perceived, engage two different sensory spheres, may well serve to prove the existence of an induction mechanism, present everywhere in the brain, and anatomically dependent upon the association-, and projection-systems. John Stuart Mill, who cer- tainly possessed no anatomical knowledge of this sort, furnishes a simple instance of this process of induction. A person who would happen to find a watch on an uninhabited island would infer not only that this island possessed a fauna and a flora, but that man must have been on this island ; for the idea of a clock or watch is inseparable from the idea of a human being. To this I will add the example I employed in Leidesdorf's " Manual." Let us imagine the cortex to be a tabula rasa, and let us present to it a phenomenon which, perceived by two different sensory surfaces, would, through the mediation of the corona radiata, stimulate two distinct areas of the cerebral cortex. Let the phenomenon in question be a lamb, and let us suppose that it emits a bleating sound. The sight of the lamb will stimulate cells of the visual area, and the bleating sound will arouse cells in the auditory sphere. The lamb disappears and the two kinds of images which it caused to be registered will grow fainter. 154 Psychiatry. If, in the course of time, one of these registered images be revived, through the bleating, say, of the lamb hidden in a stable, then not only the auditory, but the visual image also of the lamb will be reproduced. In the cortex an inference is made from the sound to the body that gave forth the sound. We can readily understand this process of induction, if we will but assume that the original excitation of both sensory areas included the excitation of those arciform bundles which united the cells of the visual and auditory areas of the cortex, which areas had in turn received stimulating impulses through the pro- jection-bundles. In this way both registered images are associ- ated, and whenever the one of them is re-excited, the excitation will extend along the association-fibres to the other cells, which on a previous occasion had been taken out of a condition of repose simultaneously with the cells harboring the former image. The association-bundles may be compared to a connecting thread,' which enables one image to lift the other, as it were, over the threshold of consciousness. Inferring one attribute of a phenomenon from the presenta- tion of another attribute, constitutes an induction; it is a recognition in the direction of causality, for the bleating sound is taken to be the result of the presence of the lamb. And here it will be well to add that, though the seats of these registered images are connected with many other cortical regions, there is no need of supposing that all these association-bundles will be simultaneously excited. In the first hypothetical case we supposed that the perception, the recognition of these images, or the induc- tion regarding them, were effected by a brain in which no other images had as yet been registered in any part of its cortex. In that event, of course, but one association could be estab- lished. A second, not nearly so hypothetical a case, will show us that not all the existing association-bundles connecting certain groups of cells with other cortical regions need be called into activity when any one member of the associated groups receives an exter- nal stimulus. In spite of the existence of numerous functionally perfect connections between such cortical groups and others, recollection is due to the excitation of accessory impressions (simultaneous sensations), which were registered simultaneously . with the impression just reproduced, and which were established with especial reference to the revivified image. In the example Spontaneous Movements : Subcortical Centres. 15 5 cited above, the visual image of the lamb was inferred from the sound ; this can be explained by assuming that the visual image was an accessory impression, a sensation, received simultaneously with the perception of the bleat. Hence we argue that in real cases of numerous and complicated associations in various cortical regions, recognition depends upon the re-presentation of accessory sensa- tions which had entered into a union with certain registered im- ages. Among its numerous associations each registered image may be considered to be a special group of simultaneously perceived sensa- tions. We shall see that this is analogous to the recognition of single retinal areas through accessory sensations, through local signs, the contents of which comprise sensations of innervation, by means of which the perception of space is effected in the cortex. A person who met with some stirring experience on a large, monotonous meadow, will recall, in passing over this meadow, the very spot at which this occurred ; but all other parts of the meadow he will not be able to identify in this way, With this one spot impressions and secondary presentations have been asso- ciated, which will serve as local signs for recognition of this one spot in the midst of this uneventful barren meadow. It is safe to suppose in regard to all inductive processes, that certain obstacles which impede the excitation of cells in full repose are very much lessened after a single, and particularly after repeated, identical excitations of association-bundles uniting the cells of two distinct areas of the cortex ; while the transmission of such stimuli to association-tracts, which have been called upon to unite other, previously established, groups of associations, becomes wellnigh impossible. We may add that Wundt,1 in his great psychological work, has adopted the term, " association-bundles." Another characteristic of animals which have been deprived of the cortex is the lack of so-called spontaneous movements ; move- ments not of a reflex nature, but resulting from cortical impulses which seem to bear the stamp of volition (freedom), and are not caused by undoubted momentary impulses. They are the result of innumerable registered images, the residual-effects of past motor stimuli. We are told also, on the authority of Goltz, that animals which have had the medullary substance of their hemispheres removed (by a stream of water) were able to walk; in these cases subcortical motor apparatuses must have remained functionally intact. x " Grundziige der physiologischen Psychologie," 1873. 156 Psychiatry. Longet, Schiff, and Goltz report that animals without hemispheres were able to fly and swim, and Goltz found that the frog could move forward well enough, as long as the mesencephalon and cerebellum were undisturbed. Plainly then co-ordinated move- ments may be initiated by subcortical centres ; and these move- ments responding to momentary stimuli, among which we must class the retinal image, will come under the heading of the primary mode of movement. It was stated above that all excitations of the fore-brain are secondary. By reason of the anatomical connections, at what- ever level, between the cortex and subcortical centres, memories of innervation are deposited in the former, which record the active processes of the latter. Every registered image, in fact, depends upon impressions received primarily in the subcortical centres. Let us illustrate this by analyzing the act of conscious closure of the eyelids. Let a sharp instrument touch the conjunctiva of the eye (O., Fig. 59); the impression of this will be conveyed by the tract Va. 5 to the trigeminal centre in the pons. A subcortical centre of the seventh nerve (7) is connected with the former, which will transmit this stimulus to a facial branch in the tract Vila., in fact to the facial branch ending in the sphincter palpebrarum (Sph.). Three impressions will also be recorded in the cortex. 1. The image of the sharp instrument conveyed by the tract II. aa EII. 2. The sensation conveyed by the trigeminal branch of the conjunctiva and carried by the route a5 EV. 3. Sensations of innervation, transmitted by the closure of the lids to the cortex through the tract 7 a J7. All of these simultaneously excited cortical centres will at once be united by association-bundles, and in the following order : The visual centre which received the image of the sharp needle will be joined to the trigeminal centre of the stimulated conjunc- tival branch ; then, again, the visual image of the needle will be joined to the innervation-centre of the facial branch supplying the sphincter palpebrarum ; and, lastly, the above-mentioned trigeminal centre will be united to the said centre of facial innervation (J7). In this illustration we have proceeded on the supposition that this is the first impression of conjunctival irritation which the child received, and that those images, which were conveyed by the tracts described above, were aroused and associated with one another in the cortex. Explanation of Conscious Movements. 157 If hereafter a needle should happen to be brought close to, without injuring, the conjunctiva of the child, the image of the needle will revive the sensation of pain in the conjunctiva, and with the image of the needle there will be associated also the sen- sation of innervation of the facial branch ; thus the mere sight of the needle will suffice to excite the act of innervation within the fore-brain. The association-bundle EII. ]y. conveys the optic stimuli to the centre of innervation, and this may be re-inforced by stimuli conveyed along the tract connecting the trigeminal centre with J7. The recognition of the needle, and an inference fig- 59- Diagram Explaining the Mechanism of Conscious Closure of the Eyelids. 0. Eye. Sph. Sphincter palpebrarum. L. Levator palpebrse superioris. nuc. Laud. Nucleus caudatus. nuc. b. Nucleus lenticularis. Th. Thalamus. Q. Corpus quadrigeminum. Lbl. Cerebellum. MC. Medulla spinalis. 3, 5, 7. Central nuclei of the III., V., and VII. nerves. II. Optic nerve. III. Third nerve (n. oculomo- torius). V. Trigeminal. VII. Facial. Jdv. Individuality. J3, [7. Cortical centre of innervation for III. and VII. nerves. EV. Cortical centre of Trigeminus. EII. Cortical centre of the optic nerve. Lines marked a denote centripetally conducting projection-fibres; b., c. centrifugally conducting proj.-fibres. Red lines denote as- sociation-fibres (systems). as to the pain it inflicts, will, without any further injury, produce a closure of the eyelids through the centrifugal tract, ]j. by., by means of a volitional impulse starting from the cortex. The cortex and subcortical centres may, therefore, be said to possess a mechanism which, intelligently unites two motor acts occurring at different times. Without such a mechanism con- scious movement would remain inexplicable. The cells of the cortex are endowed merely with the power of perceiving the origin of sensations of innervation ; but the cortex could not Fig. 60. Diagram Explaining the Mechanism of a Conscious Movement of the Arm. F. Frontal cortex. ccO. Occipital cortex. CN. Nucl. caud. LN. Nucl. lenticu- laris. hT. Thalamus opticus. D. Mesencephalon. L. Pons Varolii. O. Med. oblongata, characterized by the olivary body. M. Medulla spinalis, terminating with a cross-section of the cervical spinal cord. cd. Cerebellum. Blue lines indicate centrip- etal ; red lines, centrifugal tracts ; the red and blue circles in the spinal cord and fore-brain denote the central gray nuclei ; black lines mark the association-fibres. 1 ai. Sensory tract of the arm. B. Part of the cortical centre for cutaneous sensation. 2. Tract for effecting the movement of the arm. 3A. Conducting tract of the optic nerve. A. Part of the visual centre. 4C. Tract conducting sensation of innerva- tion, interrupted in the thalamus. C. A centre in the cortex for sensations of inner- vation. 5. Centrifugally-conducting tract, originating in the cortical area C. 158 Secondary Form of Movements. 159 originate movements, if the innervation-sensation of definite forms of movement were not conveyed to it from subcortical centres by the primary form of movements. In consequence of its connec- tions with subcortical centres, and as a result of the sensations of innervation deposited in its substance, the cortex becomes a spec- tator of the reflex acts evolved in the subcortical centres. And thus we find,that, in the normal brain, no reflex actions can be performed without exciting to action the second portion of this mechanism— namely, secondary volitional movement which no longer requires the stimulating influences of a reflex action. The influence of innervation-sensations on centrifugal nerve- tracts is explained by assuming the former to be images regis- tered in the cortex ; and that the assumption is a correct one is rendered probable by the fact that intense volitional efforts may be directed to paralyzed muscles, without producing the slightest contractions in these muscles, thus proving that the innervation- sensations in question do not arise from a perception of muscular contractions, but from sensations which the reflex centres, in their capacity as subcortical sensory centres for motor-sensations, transmit to the cortex of the brain. The sensation of innervation which reaches the cortex, and from the cortex innervates centrifugal nerve-routes, does not present the fundamental motor-images; for the registered im- ages, and the sense-perceptions are altogether incommensurable. Furthermore, if for the term " registered images " we should substitute the expression " presentation," we would find that in these instances of secondary stimulation of the fore-brain, the presentation of sensory impressions, be they ever so intense, does not in the least reproduce the image of the original (primary) sense-perception. This is true also of the sensory images of the innervation-sensations, which are distributed over various cortical regions as motor presentations (Bewegungs vorstcllungen). We must look for the indirect causation of even the remotest effects of cortical movements in the centripetally-conducting sensory nerves. Using a former simile, and comparing the gan- glionic cells of the central nervous system to living organisms, we may say that stimuli acting upon their tentacles invariably produce excitation of the claws, and movements of their muscular apparatus. The light of the sun produces contracture of a muscle of the iris, the constrictor pupillae ; and if it be excessive, it will bring about closure of the eyelids. Sensory fibres of the optic nerve convey i6o Psychiatry. this stimulus through nerve cells and motor fibres connected with these muscles, the optic stimulus thus extending beyond optic- nerve cells. The optic nerves supplied the primary motor im- pulses, and gave rise in the first instance to the cortical sensations of innervation. Conscious closure of the eyelids is, therefore, an after-effect of the intense sunlight which stimulated the optic nerve. Whence it follows that the motor effects of our conscious- ness reacting upon the outer world are not the result of forces innate in the brain. The brain, like a fixed star, does not radiate its own heat; it obtains the energy underlying all cerebral phenomena from the world beyond it. The secondary order of movements, hitherto described, com- prised mere repetitions of movements which were primarily part of reflex actions. The reflexes had been transplanted to the fore- brain, and naught but the means of exciting such movements from the fore-brain had been changed. The initiatory impulse was con- veyed by the association-bundles. But in the fore-brain any order of movements can form an indefinite number of associations, and so the associations engendered by reflex movements become part of the whole mechanism of association. The result is, that movements which were originally reflex in character can be set in action by any one of a larger number of associated impulses. In order properly to understand the relation between primary and secondary movements, it will be wise to give another illustra- tion ; for in the former illustration we failed to explain the anatomical substratum of the functional activity of subcortical centres with regard to the sensations of innervation and the motor impulses. Let us suppose a flame to have injured the hand of a child, and that the latter withdrew the hand from the flame. This movement (the withdrawal of the hand) will be effected, without the intervention of conscious impulses, by an impulse conveyed from the injured part by the centripetal tract I (Fig. 60), and transmitted through a spinal-cord centre along the path 2, which withdraws the hand from the flame. This movement and the altered position of the arm are indicated by dotted lines. The following records of this act will be transmitted to the cortex through the agency of the projection-systems : First, the visual image of the flame from the eye along the tract 3A.; secondly, a painful sensation from the injured part along the tract 4B. This nerve-tract passes uninterruptedly through the cerebral ganglia Optic Thalamus and the Upper Extremity. 161 and the capsula interna to the occipital lobe, as is evident from a study of the course of the outermost and posterior bundles of the internal capsule (p. 4, Fig. 5, POM.; p. 85 ; Fig. 22, Tm.). Thirdly, the sensation of innervation resulting from the reflex movement. The interruption of this tract in the thalamus is in accord with the physiological fact, that the movements of the upper extremities are influenced by this ganglion, but that destruc- tion of the thalamus is not (necessarily) followed by an interruption of the tract conveying cortical impulses to the muscles of the arm. The innervation-sensation C. acts through the centrifugally- conducting tract 5 upon the central origin of those anterior roots which, through reflex excitation, protected, the arm against the flame. This central origin of the anterior roots is controlled first by primary motor impulses carried to it by the posterior spi- nal roots, and secondly by secondary motor impulses emanating from the cerebral cortex. Since the centre C. is connected with the association-bundles CB., CA., and the latter with one another by the bundle BA., the child need not actually burn its hand again before guarding against the flame ; but the memory of the flame and of its effect (through association with the centre in which the painful sensation has been stored), will suffice, through the one or the other of these associations, to initiate a movement which will put the arm beyond the reach of the flame. We have taken our illustration from the movements of the upper extremity, because in so doing we can refer to a ganglion which we know positively to be connected with this extremity, and to be intercalated in the path of its sensations of innervation. This ganglion is the thalamus opticus. The optic thalamus is not a ganglion of the fore-brain. All pa- thologico-anatomical lesions prove that the lenticular nucleus is the ganglion of the fore-brain, which conducts cortical impulses from the one hemisphere to the anterior roots of the opposite side of the body. Nothnagel has given the most accurate proof of this by observing the condition of rabbits after he had de- stroyed their lenticular nuclei by means of chromic-acid injections. He reports that the animals acted as though their hemispheres had been rernoved. They exhibited not a sign of spontaneous movements, and yet irritation of the skin caused them to leap for- ward. Nothnagel proved also that complete destruction of the thalamus did not in the least impair the movements of the ani- mal, and that consequently centrifugal nerve-tracts between the l62 Psychiatry. cortex and the anterior roots did not pass through the thalamus. In a case of right hemiplegia I found symmetrically situated cysts (of the size of a hazel-nut) in each thalamus. These symmetrical lesions of the thalami could not account for the hemiplegia, but Basal Surface of a Brain-Axis with Tubercular Tumor of Right Side. I. Island. T. The cut temporal end of the corona radiata. F. The cut frontal end of same. II. Optic nerve and tract. P. Pedunculus cerebri, broader on right side, and passing without a definite boundary line into the region of the lamina per- forata posterior ; crowding at the same time the right half of the pons backward. V. Pons Varolii. O. Oblongata. Cb. and Cbt. Cerebellum. The left half of the cerebel- lum is larger in volume in consequence of a tumor enclosed within. I found in addition to these a unilateral lesion in the left nucleus lenticularis. This observation shows of what little moment the thalamic lesions were, and what great importance was to be Lesion of Thalamus. 163 attached to the unilateral destruction of the nucleus lenticularis. Yet, on the other hand, Nothnagel was able to infer from his ex- periments that extirpation of the thalami, like the removal of the innervation-centres of the cortex, seemed to produce a condi- tion in which the animals remained ignorant of the position of their extremities, from which it follows that the optic thalami Fig. 62. Dorsal Aspect of Same Brain-Axis (vid. Fig. 61). I. Island. Nc. Nucleus caudat. Th. thalmus. III. Third ventricle. Q. Corp. quadrig. V. Valvula cerebri. P. Region of lemniscus and processus cerebelli ad cerebrum. The right corp. quadrig. is flattened ; the left is crowded away from the median line ; furthermore the broadened thalamus was pushed to the left. The third ventricle is not mesial. Two incisions, one into the corp. quadrig., the other into the optic mass, enabled us to inspect the convex surface of the tumor, which had attained to the dimensions of a pigeon's egg. 164 Psychiatry. are a signal-station (Wundt) for sensations of innervation, akin to those contained in certain cortical centres. According to the experiments of Longet, Schiff, and Nothnagel, incomplete and unilateral lesions of the thalamus influence the position of the head and the upper extremities. The muscles of the spinal column turn these in such a manner that the head looks toward the healthy thalamus; while on the side of the healthy thalamus the flexors of the upper extremity, and on the side of the lesion the extensors of the upper extremity, appear contracted. From the description given in the previous section of the thala- mic origin of the tegmentum (p. 90, et scq. and p. 132), we learned that decussating and non-decussating fibres passed from the thala- mus to the spinal cord ; the decussating fibres through the poste- rior commissure, the non-decussating through the laminae medul- lares, which originate, in radiating fashion, from the thalamus. As this twofold thalamic origin of the spinal cord holds good for its relation to the upper extremities, we can readily understand that both upper extremities should be represented in one and the same thalamus. The deviation of the upper extremities and the turning of the head toward one side account for the manege- movements of operated animals. This circular movement is lost in a very few days, and after that the animals which have been thus operated upon show no sort of paralysis. In the year 1872, I had occasion to observe a five-year-old boy, who was affected with a large tuberculous tumor (figs. 61 and 62)—of the mesencephalon and the thalamus. The diagnosis was easily made, for the boy showed marked paralysis of the right third nerve, and slight paralysis of the left facial nerve, and of the left extremities. The architecture of the mesen- cephalon (fig. 41), and particularly the origin of the third pair of nerves, lead us to infer the dependence of such a grouping of crossed paralyses upon lesions affecting the right crus cerebri and the third nerve. In addition to the above symptoms, it was noticed that the head of this boy was turned toward the left side; and that, while at rest, there was flexion of the left upper ex- tremity, and extension of the right upper extremity; a rel- atively great resistance was required to counteract this flexion or extension. From this circumstance it was fair to suppose that the tumor involved the right thalamus opticus. Figs. 61 and 62 represent the post-mortem appearances. Symptoms from Thalamus Lesion. 165 Whenever this boy was otherwise occupied, this compulsory position of the head and upper extremities disappeared; and, in spite of the paresis of the left facial nerve and of the extremities, he played quite naturally with the objects which engaged his at- tention for the time being. A lesion of the thalamus does not produce paralysis, and for this reason we cannot agree with Schiff, who supposes that rota- tion of the head to the right results from paralysis of the left rotatory muscles of the spinal column, and that the flexion of the right arm was due to a paralysis of the extensor muscles of the same arm ; and extension of the left extremity, to a paralysis of the left flexor muscles. But this typical pathological position following upon a thalamic lesion will admit of a different explana- tion. We may suppose that the boy, lacking the innervation- sensations of the left rotatory muscles of the spinal column, was induced to excite the missing sensation of innervation by a volitional rotation, to produce the lacking sensation of innervation of the left-arm flexors, by forcing a flexion of this arm, and under similar circumstances to compensate for the disturbed sensation of innervation of the right extensor muscles by exciting a forcible extension of the right arm. The lack of innervation-sensations created delusions with re- gard to the position of his extremities, which in turn gave rise to volitional movements, in consequence of which he assumed the typical pathological position, the so-called forced position. If he had been a quadruped this position would have been fol- lowed by circus movements. Making use of our knowledge of the twofold, asymetrical origin of the spinal-cord bundles in the thalamus, we conclude that the rotatory muscles of the spinal col- umn and the flexors of the upper extremity are represented by bundles which decussate below the thalamus, whereas the extensor- muscles of the upper extremity are represented by direct bundles from the spinal cord to the thalamus. In my pamphlet published in 1869, " On the Twofold Origin of the Spinal Cord in the Brain," I showed that the origins of the tegmentum and the ganglia of the di- and mes-encephalon were not connected with the centrifugal nerve-tracts of the fore-brain, but that they served to initiate the primary forms of movements, as we conceived of them above; that on the other hand the nucleus lenticularis was connected with no centripetal tracts which could possibly convey reflex stimuli to it, but that in accordance with all pathological data as 166 Psychiatry. well, it carried the motor impulses from the cortex ; and increas- ing with the dimensions of the hemispheres, attained the greatest magnitude in man. As a corroboration of this view, I have found the area of the tegmentum which contains the spinal-cord origins from the di- and mes-encephalon, relatively and abso- lutely more developed in animals than in man. The secondary movements which receive their innervation from the fore-brain are effected, therefore, by nerve-tracts, which pass through the nucleus lenticularis (Vide Wiener med. Jahrbiichcr, 1872). Soltmann ' has discovered a physiological fact which postulates the necessity of finding the origin of conscious movements. He has found that those regions of the cortex which if stimulated in the adult brain produce muscular movements, are in the new- born animals unexcitable and " not yet motor " in function. But as the reflex movements are perfect in these very young animals from the time of birth, we are doubly justified in believing that the reflex movements constitute the primary form of movements, and that centres for the innervation of the secondary forms are established later, and are derived from the sensations of innervation connected with the former class of movements. That centripe- tal tracts connecting the cortex with the subcortical centres for sensations of innervation, such as the thalamus, constitute the ana- tomical link in the chain producing secondary movements, is my own well-established conclusion, and one that is supported by Soltmann's discovery. In its relation to the upper extremity the thalamus was shown to be a motor mechanism, through the agency of which decussat- ing and non-decussating representatives of certain muscles effected certain definite forms of motion. This was demonstrated by the nerve-tracts of the case pictured in Fig. 60. The nucleus lenticu- laris, which, with its second division of the projection-system (p. 27), winds around the inner side of the pes pedunculi as the ansa peduncularis (constituting the innermost and most posterior of its bundles), and in part traverses this same pedunculus, in order to reach the stratum intermedium and the substantia Soem- meringi behind the pes pedunculi ; this nucleus lenticularis, so connected, effects movements only on the opposite side of the body. The point at which this impulse crosses to the opposite side is not to be looked for in the decussation of the pyramids, : Jahrbuch f. Kinderkrankheiten, N. F., IX. Experimentelle Studien fiber die Functionen des Grosshirnes des Neugebornen. Sensation of Innervation. 167 but since I proved anatomically (p. 136), in unison with Tiirck's processes of degeneration after destruction of the lenticular nucleus, that the stratum intermedium is transformed in the spinal cord into the innermost bundles of the anterior columns, it must be looked for in the anterior commissure lying next to these. There is still a third grade or order of innervation from move- ments of the body which must engage our attention. The move- ments of this order, like those of the second order, are initiated through the mediation of the association-mechanism from the cortical centres of innervation. They are distinguished from movements of the second order by the fact that the form of move- ment is not a mere repetition of reflex movements, and that amid circumstances, and after stimuli, which would produce a definite form of reflex movements, or which would excite secondary cortical movements, copying the form of. movement prescribed by the reflex,—under such circumstances entirely different forms of movement would be initiated in the cortex. Thus a cool and collected person, who submits to an operation on his eye, will, in- stead of allowing the sphincter palpebrarum to act as soon as the knife approaches the eye, be able to keep the eye wide open by in- nervating the levator palpebral superioris. Mucius Scaevola, of legen- dary fame, was able to retain his arm in the sacred flame, instead of withdrawing it in obedience to the ordinary reflex excitation. The first of these two instances is explained in Fig. 59. From the cortical centre for the innervation of the (musculus) levator palpebroe superioris (J3), the roots of the third nerve may be innervated through the tract J3 C3 in response to irritation of the conjunctiva; whereas irritation of the conjunctiva ordinarily produces a reflex movement of the sphincter palpebrarum ; this primary form of movement is followed by conscious closure of the eyelids—the secondary form. Consequently we may say that the innervation centre of the third nerve is controlled also by stimu- lating tracts, giving rise to movements which have no prototype in the reflex mechanism. The innervation centre for the third nerve is anatomically connected with a number of mutually asso- ciated centres arbitrarily drawn in Fig. 59 which should be repre- sented as distributed over the entire cortical area. These centres, connected with one another by every variety and length of asso- ciation-bundles, are nothing special, their " memories " are defined by the peculiarities of localization referred to above. The sum of these " centres" constitutes the " individuality," the 168 Psychiatry. " ego " of abstract-psychologists. I attach some importance to the word " individuality," because it is founded upon the ana- tomical structure of the cortex, and the simple physiological process which enters into our present discussion. Individuality implies the sum of firmest associations, which under ordinary circumstances are wellnigh inseparable ; the aggregate of " memo- ries " forming a solid phalanx, the relation of which to conscious movements can be defined apparently with mathematical precision. This unequal activity of the fore-brain, constituting individuality, varies as regards contents and degree with each person ; it is designated also as the character of the individual. It has been justly observed, if the character (individuality) of a person were entirely known we would be able to predict the thoughts and deeds of such an individual, however complicated they might be. The observation of Munk quoted above, and the conclusions of experimental physiology, strengthen the opinion that intelligence is not limited to definite cortical areas, but that, being based upon perceptions including the sensations of innervation, it results from the activity of the entire fore-brain. Anatomically speaking, the exercise of intellectual activity by every part of the cortex depends upon the uniform structure of all parts of the fore-brain, which makes of each part a centre for inductive processes, and supplies to each part nerve-elements capable of perceiving and re- taining "images." This is true also of individuality, the motives of which are to be sought in the most frequently repeated im- pressions of every sort. These impressions hold a causal relation to one another, and by means of nerve-arches establish a connec- tion of causality between residual images' as motives for move- ments, and certain definite motor-acts. In order to be clearly understood, I will begin by explaning the simplest relations. Kussmaul has shrewdly remarked that certian perceptions and movements are common to the foetus in utero ; that the foetus feeds itself, as it were, by swallowing amniotic fluid, and that there may be special motives, such as the more-stimulating taste of the fluid after the occasional depletion into it of the allantois, which induce swallowing. Here, then, there are two al- ternatives, the foetus swallows or does not swallow. We are already presupposing fcetal consciousness, which would exist, however, un- der circumstances which give but little opportunity for perception. 1 The reader will by this time have noticed that I have given several translations for the German "Frinnerungsbild"; no one English term appears to me to be entirely satisfactory.—S. Primary Individuality : Instinct. 169 The new-born infant at once discovers signs by which it distinguishes between one set of perceptions and another. One set of perceptions helps it to define the circumference of its own body, another set belongs to the world beyond it. However obtuse this perception may be, and though the child may at first not even be able to discriminate between the various impressions of space, it is certain that the perception of its own body-circumference is established very early in childhood. Among other signs by which it learns to distinguish between impressions received from its body and from the outer world are these: Con- tact of a strange finger with its own skin excites but one tactile sensation ; contact between two parts of its own body excites two tactile sensations—one from the touching, and the other from the touched part (Wundt). Furthermore, a number of strange audi- tory sensations strikes the ears of a child, but only the sound of its own voice is accompanied by muscular sensations ; and so the attendant muscular sensations help the child to discriminate be- tween movements of its own body and any other forrn of move- ment which it may have occasion to see. The number of such examples could be increased with ease. And, besides, it is im- portant to remember, that the continued presence of impressions which its body yields—as, for example, the sight of its own hand, and the ever-increasing number of muscular sensations, due to movements of the hand, will soon become fixed in its con- sciousness, and that, by repetition, the intensity of such percep- tions will soon outweigh the interrupted occasional impressions received from the outer world. In the work of registering images in the cortex, the memories conveyed by those nerve fibres, which enter into definitely asso- ciated groups, and constitute tracts along which a motor impulse is most easily conducted, lend a helping hand ; it is fair, therefore, to expect that frequent and simultaneous sensations, which emanate from the body itself, will enter into firm connections, which they will never again be able to dissolve, and which will thus form the first primary conception of the ego, based upon the perceptions of the body's circumference. And here it will be proper to remark, that there is no order of movements which, under the cover of instinct, can be pushed in between conscious and reflex movements. The first instinct of a child would be its instinct for food ; but the origin of that has been alluded to. There is absolutely nothing in the sensation of 170 Psychiatry. hunger which would acquaint the child with the means of remedy- ing this pain, of initiating movements suited to this end. It ob- tains naught but the concept of pain. In the general restlessness it displays, and in the convulsions ultimately resulting from anaemia, there is nothing which could be likened to an instinct for food. If the child is not left to depend upon its own resources, but has a nipple put into its mouth, then the sensation thus excited starts the reflex mechanism of sucking. The child has thus acquired the concept that the sensation of satiation is connected with the act of sucking, and these two sensory memories are associated with the sensations of innerva- tion aroused by the sucking-act, probably, also, with the scent of its mother's breast. That the child should suck at every finger, we may attribute to a reflex mechanism ; but that the child should suck in its dreams, proves that the act of sucking has produced images which have been registered in the cortex. If a child looks for the nipple by turning its head on its mother's breast, it is not moved by a reflex mechanism, and there is no logical reason to suppose that this is not a conscious movement, which is based upon the association of painful sensations and sensations of innervation, both implanted upon the fore-brain by the reflex mechanism, which gave relief to the pain. This would be analogous to the conscious closure of the lids and the movements of the arms, instanced above.. There is no gap between conscious and reflex movements to be filled in by instinct. The factors of this primary, abstract ego are not definitely defined. No child is likely to lack the sensation of hunger; but memories collected from certain sensory spheres may be wanting in consequence of deafness, blindness, or some other defective perception ; and, in proportion to the lack of perceptive powers, there will be a corresponding defect in the sensory fac- ulties of the ego; and yet through the more intense applica- tion (education) of the other senses (as has been shown in the case of deaf mutes who were blind also), an extraordinary wealth of images may be amassed, which can lead to an excellent knowl- edge of the external world. Munk's views may help us to under- stand the refinement and amazing development of any one sense which may do service, possibly, for other defective senses. Just as single sensory spheres do not occupy the entire cortical area connected with their special sensory nerves, but have a peripheral The Cortex and Special Sense-Development. i 71 zone which is left open for later occupation, so we can imagine, as long as the cortical surface is practically one, through the media- tion of the gray network of fibres, that a greater area is left at the disposal and for the development of any one sense ; for a larger area, generally occupied by other sense-perceptions, is left func- tionally vacant by the occlusion of its sources of perception, and through association-fibres this area may be joined to the only sensory spheres in active operation. Thus we are told by the blind, that from slight changes of atmospheric pressure, the reflexes excited from the skin of the head and extremities by currents of air give rise to innumerable im- pressions and inferences sufficient for them to perceive from these sensations the presence of a large quiescent body in their vicinity, and still better the presence of moving objects. The first case only is to the point; no doubt moving objects produce auditory sensations as well. A similarly exalted degree of sensory perceptions, which play a subordinate role in the life of normal man, is revealed by the bat. In Auteuriet/is Archives experiments are recorded which have been performed on bats; their eyes were destroyed, their nostrils stopped up with molten sealing-wax, and their ears were also effectually occluded. But though blind, deaf, and minus the sense of smell, these bats contrived to fly through a regularly-constructed thread network without touching any of the threads. In accordance with the opinions then prevalent, it was thought necessary in order to explain this phenomenon to call up a sixth sense, akin to that sense which formed the basis of som- nambulistic (?) perception. If we remember, however, that the bat possesses large wings, and very delicate, sensitive webs, which present an enormous cutaneous surface, we need not be surprised to find that, as in the blind, this large sensitive surface enables the bat to perceive changes in atmospheric pressure and atmos- pheric waves, created by bodies moving about in a direction opposed to the current of air. From such changes the bat infers the extent and direction of its movements. The nature of the ego does not depend upon any definite order of memories; it is determined simply by the most firmly fixed memories. As soon as we apply the test of more com- plicated relations to the formation of the ego, we must bear in mind that the ego can be influenced only by images of permanent intensity, which are associated at the same time with other and 172 Psychiatry. as firmly fixed images. The idea of individuality is an artificial one, though valuable from a practical point of view, for the degree of intensity by which these images and their connections adhere to this conception will not admit of accurate measurement; and it is plainly impossible to say that at a certain intensity a presentation becomes a factor of the ego, and not yet at another. There is but one safe stand to take on this question, and that is to attribute to the ill-defined conception of individuality only those presentations which, as soon as the " character" of an individual is known, will enable us to predict his deeds; whence it follows that the deeds of the individual obey certain laws. On the other hand, these same conscious deeds, as they are not mere copies of acts suggested by re- flex movements, remain incomprehensible as long as the character of the individual is unknown. This very mystery attending the deeds of others is another and highest expression for the freedom of the will. There can be little doubt but that the primary order of move- ments is effected by the diencephalon, the mesencephalon, and other centres lying still farther from the hemispheres; the first- named ganglion contributing largely to the formation of the spinal- cord columns, through the continuation of their bundles through the tegmentum of the crus. But now the question arises, whether there are distinctly separated cortical centres, or nerve-tracts, which mediate in the execution of secondary movements—copies of reflex movements—and of tertiary movements, which bear the stamp of individuality ; both kinds of movements being distinctly cortical in origin. Charcot contended that paralyses, resulting from (lesser) lesions of the lenticular nucleus, were distinguished by their shorter duration from those resulting from lesions of the crus, or of such parts that spring directly from the hemispheres through the internal capsule; and that (complete) destruction of the crus entails permanent paralysis. But from the study of the ana- tomical structure of these parts, we learned that a considerable portion of the internal capsule and of the pes pedunculi is inter- woven with -peduncular portions of the lenticular nucleus, with the stratum intermedium. It is evident, therefore, that destruction of the pes pedunculi will also destroy tracts from the lenticular nucleus. It was absolutely necessary for the lenticular fasciculi, lying in front of the crus, to interweave with the peduncular fibres, in order to reach the substance of Soemmering, lying behind the crus cerebri. Conduction to and from Cerebellum. i y\> Charcot's statement becomes intelligible only by supposing that the gray substance of the spinal cord conducts motor im- pulses after the interruption of the connection between the white anterior columns, inasmuch as all white bundles pass into the gray substance, and establish conducting tracts in the gray network of fibres, though the speed at which impulses are conducted is very much reduced. This might possibly hold good for a moderate- sized lesion affecting the gray substance of the lenticular nucleus, and amends might thus be made for imperfect conduction ; but lack of gray substance precludes the possibility of such patch- work in that part of the crus in which lenticular and cortical bun- dles of the crus are interwoven. Lesion of the crus is, therefore, necessarily followed by impairment (annihilation) of lenticular conduction. There is one difference, to be sure, between the course of the stratum intermedium and the pyramidal tracts, and that is that the cortical bundles of the latter develop into the anterior longitudinal fasciculi of the pons, and as such, connect with the cerebellum. The possibility of a similar connection between the cerebellum and the stratum intermedium cannot be absolutely denied, for the latter, instead of being crossed by the brachium pontis, might possibly interlace with the superior division of the corpus restiforme. And yet the intimate union between peduncular and cerebellar fibres is the only one that can be regarded as established beyond controversy. The difference in regard to the function of the pedunculus and the bundles of the stratum intermedium might become intelligible by assuming that the peduncular fibres co-operate with the brachia pontis in effecting complicated (more co-ordinated) movements, while the lenticular nucleus constitutes a co-ordinating centre per se. Ac- cordingly the co-ordinations of movements conducted by the stratum intermedium would be effected in the gray masses of the fore-brain. But I am not willing as yet to draw any positive con- clusions as to the importance of one or the other of these tracts engaged in the transmission of motor-messages from the cortex. The occurrence of sling-shaped bundles in the pons can be utilized to illustrate the manner in which the brachium pontis endows the cerebellum with a co-ordinating influence over con- scious motor-innervations (p. 115, et scq.; Figs. 43 and 44). These circular (sling-shaped) bundles, which unite in the pons with fibres from the crus, enter and terminate in the brachium pontis of the same side. I can conceive that the one arc of this sling 174 Psychiatry. constitutes a tract along which cortical motor impulses, passing through the pes pedunculi, reach the cerebellum, and by which the cerebellum is notified of the cortical sensations of innervation, the work of association ; that, on the other hand, the other arc, taking a recurrent course through the gray substance of the pons, conducts to the pedunculus cerebri the influence of the cere- bellum. On page 28 it was demonstrated that the dimension of the pes pedunculi, the nerve-tract along which cortical motor im- pulses travel, varied with the dimensions of the fore-brain in general, and that for this reason all structures directly connected with the pes pedunculi, such as the pons and the pyramids, were particularly well developed in man. We showed also that the alternating visibleness of the olivary and trapezoid bodies in man and animals was determined by the fundamental factor in brain-trunk architecture—the relative development of the fore- brain. Were any animal able to increase the size of its fore-brain, the basilar aspect of its trunk-structures would approach more closely to the human " type." The stratum intermedium forms an integral part of the section of the anterior tract of the brain-trunk, which far exceeds, in dimensions, the cross-section of the tegmentum. But the stratum intermedium is derived from the lenticular nucleus, which attains its greatest development in man, and from the pes pedunculi, with which it interlaces. Consequently the development of this stra- tum will keep step with the development of the fore-brain, at least as far as conducting nerve-tracts are concerned. If appearances deceive, it is due to the presence in animals of larger quantities of scattered gray connective tissue possessing no functional value. The exact function of the radiation of the posterior longi- tudinal fasciculus is still a matter of conjecture: its connections with the nuclei of cerebral nerves, exhibited above with regard to the third nucleus, give undoubted evidence that it is partly motor in function ; but, on the other hand, it is connected also with sensory tracts, those of the fifth and eighth nerve ; and possibly also with the retina through the mediation of the basal ganglion of the optic tract. Lesion of the lenticular nucleus produces hemiplegia ; and yet there is no connec- tion between the fibres of the anterior columns (descended from this ganglion) and the coordinating centre in the cerebellum save low down in the central gray substance, where the bundles just mentioned unite with the pyramidal tracts. There is no conscious stimulus starting single complicated acts of cerebellar coordination. And then, too, mere ataxia resulting from loss of cerebellar influence does not affect the conscious in- The "Ego " and the Individuality. i 75 nervation of uncomplicated movements ; while clinical observation goes to show that there need be but slight consciousness and hampered memory attending the distinctly coordinated movements in the hypnotic state and in cases of chorea magna. We may argue, therefore, that during normal cerebral activity coordinated and conscious move- ments are tolerably distinct phenomena. Since coordinated movements are no criterion for the activity of the lenticular nucleus, and since the initiation, direction, and inhibition of a movement, which constitute its conscious features, remain intact as long as the lenticular nucleus is preserved, and in spite of the presence of ataxia result- ing from injury to the cerebellum, we must infer the only condition necessarily preceding a consciously excited movement to be this, that t!:c lenticular nucleus be stimulate sim.- ultaneously with the cerebral cortex. Moreover this very ganglion, forming together with ihe nucleus caudatus, part of the gray substance of the fore-brain, stands in closer genetic relation to the cortical substance than do the other (strictly speaking) sub-cortical ganglia. I return now to the discussion of the phenomena of the ego evolved by the cortex cerebri and its medullary mass; but I do not intend to treat this exhaustively at present, for we shall have occasion to study this subject more fully in the clinical portion of this work. Let us suppose, in order to give a concrete meaning to the term, that there be a primary ego, a nucleus of the individuality, defined by the limits of the infant's body. And, as a rule, the image of the primary ego will coincide with the mental concept of the body; there will be, as it were, a nucleus of the individuality. But the most fre- quently repeated perceptions of the outer world, as well as the most frequently revived memories, and particularly those joined to the emotions, will enter into firm associations, and will consti- tute the nuclei of a secondary individuality. Such memories will be more readily reproduced, and will exert a greater psycho-motor influence than transient impressions and less intense perceptions will do. And yet the substance of this secondary individuality lies beyond the circumference of the body itself. The entire indi- viduality becomes decentralized and is made to include much of the external world. The primary ego expands through permanent and intense secondary perceptions joined to it by association; so that in- timately related persons, property, skill obtained by constant practice in any art, science, a fondly cherished aim in life, convic- tions, patriotism, and honor become part of the ego. That from among these component factors of the ego, the primary ego should be consciously endangered by surrendering one's own body, can be explained by assuming that component factors of the secondary ego have attained such a psycho-motor intensity in the play of association, as to become more effectual motives 176 Psychiatry. than the original powerful motive of self-preservation. In fact, the very person who sacrifices his life believes, in so doing, that he preserves his own individuality, which now includes so much that lies beyond his physical self. Whatever explanation one may choose to give of this phenomenon, the simplest will be this : In all the actions of man, be they ever so complicated, problematic, or incomprehensible, the avoidance of greater pain is the deter- mining motive. The satiation of hunger is a powerful presentation in the infant's mind. But if we should attempt to put any thing which had been steeped in acetic acid into the mouth of a starving child, the perception of a momentary still greater pain would be joined to the sensation of hunger, and a reflex mechanism would be set in action which would prevent the swallowing or retention of this noxious substance. Whenever, later on, a substance that merely smells of acetic acid is brought near it, the infant's mind, having already learned to utilize its registered images and their associa- tions, as well as to start simple processes of thought which deter- mine the initiation of this or the other movement, the child will spit out, retch, cry, kick; it will consciously repeat repulsive move- ments which were originally reflex in character. These movements of repulsion are the direct opposite of those aggressive movements exemplified in the act of sucking, and in clinging to the mother's breast. The child may associate the pleasure.of satiation with the swallowing of food, and yet this so-called pleasurable sensa- tion will not be the determining motive for the child's action; but as long as it is exposed to two painful sensations, that of hunger and of the pain produced by a caustic substance—the choice of the lesser pain will be the determining motive. This discussion of the individuality has led us far astray from the consideration of such facts in regard to the architecture of the fore-brain as are supplied by a study of its anatomy, and such as are supplied by physiological experiments. A more definite observation of experimental physiologists, though clad in vague language, is this : They maintain that loss of fore-brain entails loss of intelligence in animals. In order to give tangible shape to this term " intelligence," I would say that under the guidance of this symbolic faculty, only those associa- tions prevail (from among the anatomically possible associa- tions of registered images and concepts) which represent the ordinary regular connections of things. All parts of this organ Methods of Thought, Associations. 177 of intelligence are joined to each other by an evident anatomical and functionally perfect association-tract. The anatomy of the cerebral structure, and the proof of the presence everywhere in the brain of an induction apparatus, render it highly probable that all perceptions received simultaneously or in continuous suc- cession become correlated with one another. Such connections explain the relations of successively and separately received im- pressions to one another. Our methods of thought and of speech have designated this relation as one of causality; but this is a purely cerebral function, for there need be no bond in the outer world corresponding to these cerebral relations of causality; nor does this relation constitute intelligence. The so-called logical sequence in the evolution of association, which yields the factors of intelligence is effected in various ways and to a varying degree of perfection in different brains. First the intensity of established associations, dependent upon their conscious and frequent re-excitation, is of greatest import- ance. An accidental succession of impressions is seldom repeated, and relations thus established vanish quickly in the brain. But, as soon as the subjective bond of causality represents an actual union of things, the re-occurrence of external stimuli will estab- lish a permanent association within the brain. Thus by the renewal of perception, such associations are turned into the elements of inductive logical thought. Deduction we need not consider now for it is the simple outcome of Induction. Secondly, a number of approved associations are transmitted through the medium of language, through conversation and instruction from one individual to another. According to the degree of culture, the intentions in transmission, and the logic of those who communicate them, the mass of such transmissible asso- ciations will be more or less remote from intelligence. The imprint- ing of prevalent associations by mere repetition upon the brain is demonstrated also in the negative phase of cerebral activity, in the dissolution 1 of conscious associations. It is characteristic of senile loss of memory, that the earliest memories are preserved best, for these had been revived innumerable times in the processes of thought during more than half a century, while the recent associations, acquired shortly before the death of the aged, can have been recalled but seldom to memory. The loss of foreign languages, following upon pathological processes in the brain, does 1 Used in Jackson's sense.—S. i 78 Psychiatry. not entail ignorance of the sounds common to the foreign and the mother tongue, but includes only recently acquired knowledge of languages. The conclusion is forced upon us, that the indi- viduality also, apart from its relation to emotions—which in- clude the most frequently repeated associations,—is influenced largely as regards the union of its component factors, and the diffi- culty of disassociation by the more powerful bonds established between the earliest factors of the individuality. If intelligence and individuality be not evolved from per- sonal associations of ideas, and if the motives for the direction of thought, and actions connected therewith, be simple copies of psychical associations started in another's mind, then the growth of intelligence, as well as the expansion and composition of the individuality, will be seriously impeded. As correlated sensations of sensory perceptions, motor im- ages play a very important role even in the simplest processes of the fore-brain; in ordinary perceptions, such as those of space, for instance. The perception of space is by no means dependent upon the visual sense alone, for a blind person has as full a knowledge of the surface of his body as one with normal vision ' has. Such knowledge is enhanced by sensations accompanying tactile impressions received from dif- ferent parts of the skin. Thus Lotze has shown how markedly the tactile sensation, dispatched from the skin over the ster- num, differs from that forwarded from the skin of the thigh; and going still further into details, he mentions the different tactile impressions received from the finger joints, and from the more elastic and poorly supported skin between the fingers ; from the movability of the skin over the tendons, and its compressibility between the tendons on the dorsum of the hand. Lotze insists, furthermore, on the great difference between the sensations accom- panying compression of the volar surface of a finger against the nail—a perception influenced by the form of resistance presented at both ends of the nail margin—and the touch of the nail itself. He refers also to Spiess' remark, that the local signs of correlated sensations undergo further change through motion of the parts, through the varying pressure acting upon joint surfaces, the tension and relaxation of skin during flexion and extension, and through the contact between different skin areas. Correlated sensations play a very important role, in the endeavor to establish, through the sense of touch alone, a skilful Space-Image the Result of Induction. i 79 and rapid recognition of all surface impressions from the body itself. To this category of sensations belong the sensations of innervation in varying intensity, produced by contact of near and distant parts of the body's surface, and the sensations dependent upon the difference of the muscles thus set in action. A cor- related sensation of this sort has evidently the value of a secondary (allied) presentation, for if any part of the body be touched by a foreign body, we can, without putting our own hand to the part touched, be sure of the location of this point, simply from the local signs which we know from former experience to be associated with certain sensations of innervation. Projection and association are the two fore-brain principles at the bottom of such concerted action, which constitutes an induction. Funke ("Physiology," ii., p. 177) has given a lucid exposition of our judgments regarding the origin of sounds and noises in space. Animals form a conclusion as to the origin of a sound by utilizing the sensations due to the reflex movements of the concha, and other sensations produced by the reflection of sound-waves from the convex or concave surface of the concha. Helmholtz and his pupil, Wundt, have given an explanation of the retinal space-image, by showing the genetic development of space-perceptions, instead of adhering to the nativistic standpoint of other authors; with the latter we are not concerned just now, The manner in which the retina evolves a space-image, which is apperceived by the cortex, seems, at first, easy enough to explain ; as a matter of fact, this process is quite incomprehen- sible. It requires but little reflection to discover that a uniform surface, which we are looking at, very soon evolves itself into a mosaic of smaller areas, or, at all events, appears divisible into smaller spaces; and if, on the other hand, the most variegated areas of any surface be presented to our eyes, we should never- theless conclude that they were parts of a continuous surface. Granted that every retinal fibre be endowed with a special per- ceptive power for definite colors, and even for definite shades of such colors, this would not suffice to produce a space-image, for one would be compelled to suppose that these distinct local peculiarities of perception are so distributed that the disposition of variegated or uncolored portions of any surface would corre- spond to the exact juxtaposition of nerve-fibres of varying and special sensibility—a view which no one will consider plausible. Supposing the various points of a visible surface to be pro- i8o PsycJiiatry. jccted upon different retinal cells, it would be possible to account for the division of this surface into smaller areas, but it would be difficult to form a concept of these smaller areas as a continuous surface. If, on the other hand, we assume the various parts of the retina to be connected with one another, and that their re- spective cells join in common action, we can understand then how conception of a continuous surface is engendered, but will be puzzled to explain the correlated concept of its divisibility into smaller areas. We may, however, conclude that each single spot of the retina receives its local signs in a different way, and we are justified in assuming that the development of a retinal space- perception depends upon the association of innervation-sensations of the ocular muscles and retinal impressions, just as in the develop- ment of space-perception through the sense of touch sensations of muscular innervation acted in unison with tactile sensations. I wish here to dwell upon Lotze's conception of the nature of local signs, and I do this not with the intention of combating the views of others, but because by reason of their very simplicity, Lotze's . views commend themselves to our purpose. It goes without saying that the remarks previously made re- garding the inadequateness of the retina to develop a space-percep- tion, apply equally to the inadequateness of the cerebral cortex, if the latter, in taking cognizance of space-perceptions, is to confine itself solely to the adoption of projected retinal images. The retina is composed of two zones: a smaller zone of distinct vision—the macula lutea; and a larger one of indistinct division —the horopter circle. Distinct vision is accompanied by move- ments of the bulbus, by means of which definite points on the horizon are conveyed to the region of distinct vision. Thus the visual act is attended by sensations of innervation, which are projected upon the cortex, and result from the play of the ocular muscles. The projection-areas of the retina, and of the sensations of innervation, are united in the cortex cerebri by association- systems. In the annexed drawing, the projection of the retina extends from the bidbus through the optic nerve to a segment of cortical surface, corresponding to the region marked I, 2, 3, on the outside of the skull (Fig. 63). Three bright points, I, 2, 3, lying in front of the bulbus, fall upon different spots of the retina, lying one above the other (1, 2, 3. The points are erroneously numbered, but this need not interfere with our understanding of the diagram). If these spots Space-Ids ion. 181 should lie above the macula lutea, it would be necessary, in order to produce distinct vision, to move the macula lutea so far upward that the ray of light from each bright point would, in due rotation, fall upon the macula.. As an effect of the varying intensity of its contractions, the inferior rectus muscle (J) will be able to bring the macula lutea into the line of incidence of each one of these bright points. A slight contraction of the inferior rectus will cause the macula lutea to change position with that Fig. 63. Space-Vision. part which has been irritated by the bright point 3; after a stronger contraction, the retinal point 2 and the macula will have exchanged places, and a still more powerful contraction will ad- vance the macula lutea to the portion of the retinal area, upon which the point I had been projected. Through certain fibres of the tractus opticus, the points I, 2, 3 are represented somewhere in the cortex, and in ancrther cortical region there is a representation of that branch of the third nerve which innervates the inferior rectus muscle. In the diagram one fibre is connected 18 2 Psych ia try. with three cells, Jx J2 J3, and these three cells are joined to one another in the net of gray fibres. The^ contents of the sense- perceptions stored in 3, 2, 1, in the cortex, are visual impressions; of those lodged in J1 J3 J3, are sensations of muscular innervation. There are certain resistances opposed to the conduction of nerve- force in this gray network of fibres, which in the case of the nerve-tract leading to the celta ] 1 J2 J3, as long as the intensity of innervation is very slight, will excite but the one cell J T ; if the intensity be increased, there will be sufficient irradiation to involve the cell J2 ; and if the intensity of innervation be increased still more, the irradiation will extend to all three cells—from which we may infer that the signs J x J 3 J 3 correspond to different intensities of innervation. The least intensity of variation results from the effort to advance the macula lutea to the position occupied by the retinal spot 3 ; a greater, from its attainment of the point 2 ; and the greatest intensity will result when the macula lutea has usurped the position occupied by 1. In a succeeding portion of this book we shall have occasion to refer to the fact, that the in- tensity of innervation attending emotions (Affectc), for instance, is revived with the memory of this state. The varying distances between retinal areas and the macula lutea on every meridian or parallel circle of the retinal hemisphere, are designated by these local signs of graded sensations of innervation, which are derived from different muscles, according to the change in the position of the macula lutea. The sensation of innervation, which is joined to the light-impression falling upon any spot on the retina, varies according to the muscles employed and the innumerable gradations of intensity. The perceptions of these sensations of innervation and of the optic areas become associated in the cerebral cortex ; and by an in- ference as to the retinal spot, from the nature of the correlated sensation inseparably connected with it, every retinal area has its proper place in the perception of space. In this way, the topographical relation of retinal areas is registered in the cor- responding cortical region ; and from this relation we complete the image of space. If I suppose the three bright points of the preceding figure to be stars, it may be said that the retina per- ceives space between the three stars. But we cannot suppose the dark ether to be a retinal irritant; as far as perception is concerned, it is nil. That this nil between the stars is filled in by space, proves that there is no direct perception of space, but that the Aggressive and Repulsive Movements. 183 latter constitutes a subjective inference. It is not the vacant ether which is actually measured, but the inference made, aid- ing in generating the idea of space, is based upon a measurement of the distances between the three spots (stars) on the retina. To the extent that the visual concept of space is utilized by the sense of touch, we shall find sensations of innervation derived from the ocular muscles associated with similar sensations of varying in- tensity from the upper extremity. The movement of stretching out the hand to seize objects lying at the side of us is accompanied by sensations of innervation from the sixth nerve, which directs the eye outward. If paresis of this nerve provokes a stronger sensation of innervation, then by association more forcible movements of the hand will result, so that the hand is extended beyond the object it had intended to seize upon. This subjective power of the brain to evolve the idea of space from nothing, expands under pathological conditions to the power of filling out space with images which are the creations of the brain—the outcome of cerebral irritation. To this account of the widespread activity of the fore-brain, not only as the recipient, but also as the creator, of sensations, I wish to add that it is the boldest hypothesis, shared alike by the ordinary mind and by scientific realism, to assume that the world is such as it appears to the brain to be ; that the latter can be likened to a mirror which simply reflects the forms of the outer world ; that the world as it appears to the brain exists inde- pendently of the presence or absence of mind. Indeed, it seems to me to be a crucial test of an individual's power of thought, to determine whether he can conceive or not of the unreality of the world clad in forms which our minds have bestowed upon it. It should be reiterated that the idealistic conception of the world is supported by physiological facts, and still more positively by the facts of cerebral architecture before alluded to. Residual images would not furnish adequate motives for our movements and deeds if the phenomena of feeling were not inher- ent in them. To the reflex of sucking we applied the term reflex of aggression ; to that of vomiting,, reflex of rejection, of repul- sion.^ The conscious movements based on these reflexes were termed respectively movements of aggression and repulsion. These resulted from association with the ideas of satiation and injury. Such concepts were not mere passive reproductions of former images ; but their reproduction was attended by a degree of 184 Psychiatry. intensity which we term emotion. In both cases the emotion varied, producing pleasure in the one case and pain in the other. A pleasurable emotion gives rise to movements of aggression, a painful emotion to movements of repulsion. Possibly we may be better able to fathom the different degrees of emotion or feel- ings by following a more complicated though physiological train of thought and omitting psychological definitions altogether. Our understanding may be furthered by a series of facts including more than mere bodily pain and the reflex of repulsion ; for we have to consider a movement accompanying the latter and affect- ing the arteries of the central organ, and the connections of this correlated movement with the change in chemical relations—phe- nomena which are at the foundation of all nerve-phenomena. It will answer our present purpose to designate these chemical rela- tions by the short expression : respiration of nerve-cells. The solution of this problem would be reached by continuing the line of thought to which I have hitherto adhered, and by proving that the conjoined processes above referred to are al- ready associated in the cortex as secondary processes with others involved in the mechanism of reviving associated images (or con. cepts). Even though a decapitated frog respond to pinching of his skin with a kicking, repulsive movement, there is no need of sup- posing that a sensation of pain preceded this act. Nor are we compelled to regard the conduction of the cutaneous irritant to the muscles as the only process here set up, to which, if the brain had not been removed, sensation would have been added. We must remember rather that every sensory act produces a number of secondary effects acting upon the central nervous organ ; and that such secondary effects, in default of fore-brain consciousness, are nothing more than the consequence of an irritation which would be adequate to the excitation of pain. First.—The experiments of Schiff and others have taught us that in animals in which the conduction through gray substance has been impaired, by experiment or otherwise, cutaneous irrita- tion produces simply tactile or thermal sensations ; in spite of the adequate nature of the irritation (burning) and the presence of the fore-brain, there is no sensation of pain. To this changed condition of irritability we apply the term Analgesia. We know, furthermore, that nerve-conduction through the network of gray substance meets with a certain resistance which can be. Irradiation. —Inhibition. 185 gauged by the time which elapses in the transmission of a peripheral impulse. With an increase in the number of muscular groups excited to action by a reflex irritant, an increase in the resistance to nerve-conduction goes hand in hand. The irradiation of any irritation so as to involve a large number of muscular groups, say those of the side opposite the cutaneous irritation, will depend upon the duration and intensity of the latter, and will have an influence upon the character of the pain-sensation resulting from said irritation. The sensation of pain, therefore, is attended both by a reflex movement and by an inhibition of nerve-conduction in the gray substance of the spinal cord. The strength of the resistance to be overcome, in the case of motor irradiation, in- creases, even in the ca-se of the unconscious, decapitated frog, with the duration and intensity of the irritating cause. It is evident, then, that inhibition, resulting from resistances introduced in nerve-tracts, accompanies the simplest reflex processes, while consciousness in the main recognizes this inhibition as pain. In regard to inhibition, I wish to call attention to the retardation in the conduction of reflex nervous impulses. A nervous impulse takes, according to Helmholtz, about twelve times as long to travel through the gray substance as it does to be transmitted through the peripheral nerves. Transverse conduction largely increases the time required for the reflex act, as when a stimulus applied to the one side is to excite the muscles of the other side (Rosenthal, of Berlin). According to Exner, the stronger the stimuli, the less the amount of time required to effect a reflex action. Secondly.—Irritation of a sensory nerve excites in decapi- tated animals not only a repulsive movement of the skeleton muscles, but exerts an influence also over the circular muscular apparatus of blood-vessels, the centres for which are located in the spinal gray matter. The ex-and in-tensity of this influence vary directly with the amount of irradiation of the original stimulant impulse. The dilatation of the blood-vessels in the web of a frog's foot continues steadily to increase as layer after layer of the spinal cord is removed (Lister). Goltz has demonstrated the functional activity of the spinal vaso-motor centres in mammals, and W. Schlesinger (Strieker, Med. Jahrbiicher, 1879) has proved their presence by experiments with strychnia. We are justified in inferring that repulsive movement is attended by a reflex con- traction of the spinal-cord arteries, similar to the one which takes 186 Psychiatry. place in normal animals, and which is observed to result from an increase of blood-pressure in the carotid after the application of strong sensory stimuli. The rise of blood-pressure in the carotid following sensory stimulation was measured by Owsjannikow and Ditmar with the use of the manometer. This increase of pres- sure depends upon reflex contraction of the arteries (vid. p.---). In this way physical pain produces swooning, and explains why in former centuries when a confession was extorted in court by tort- ure, the person incriminated would fall asleep while undergoing the pangs of the rack. Ditmar believes that there is no better proof of the presence of sensory processes in animals than the rise of the blood column in the manometer, following upon ex- ternal stimulation, and that this increase of manometrical pressure is caused by the reflex vaso-motor contraction set up by the exci- tation of sensory nerves. Thirdly.—The constriction of the arteries, by impeding the respiration of the nervous elements, must necessarily engender a dyspncetic phase of nutrition; it will modify the chemical changes going on in these elements, and as a further result of this we shall find the sensory stimulus associated with the irritation resulting from a certain degree of dyspnoetic intoxication. Dysp- noea of tissues is alone sufficient to excite repulsive movements. Inspiration is the most ordinary form of repulsive movement evoked by a dyspncetic stimulus, which in extreme cases may, by irradiation, involve a number of muscles. But this dyspncetic stimulus results not only from defective breathing, but also from increased arterial pressure, as in convulsions, or from arterial con- traction, as in anaemia. The convulsions occurring in persons bleeding to death are an instance of the latter kind. The respira- tory centres extend, according to Procop Rokitansky, into the cervical portion of the spinal cord. From what has just been said we may assert that even in the spinal cord of decapitated animals movements of repulsion are connected with sensory excitation, inhibition of nerve-conduction, increased arterial pressure, and with dyspncetic stimulation of nerve-cells. Movements of aggression, also, can be demonstrated in animals that have forfeited the pros- and di-encephalon, and indeed in animals which have retained the spinal cord only. In his "Contributions to the Study of the Functions of the Nerve-Centres of the Frog," Goltz remarks : " If on a fine summer's Goltz s Croak-Experiment. —Aggression: 187 evening, we hear the croaking of the frogs, we conclude correctly enough that these inhabitants of the marshes are thoroughly happy in the enjoyment of the tepid waters." We infer the same cheerful spirit, when we observe the playful aggressive movements of a cat engaged in pushing a ball or a mouse before it, and always endeavoring to recapture it; when we see animals frolicking and tumbling about incessantly on the grass and in the open air, or birds that give expression to their mirthful restlessness by their song, and direct their aggressive movements upon crumbs, seeds, or worms. All these convey the same impression to our minds. As soon as a stone is thrown into the pool the frogs cease their croaking, in view of the danger to which they are exposed ; an enemy suddenly appearing before animals that have been tumbling about in the full enjoyment of their liberty will cause them to hide themselves, or to start movements of repulsion by taking to flight. The unhampered, purposeless movements of a lively animal, the song of a bird, the merry bark of a dog, are in reality movements of aggression. Goltz has excited similar movements of aggression in decapitated animals, such movements indicat- ing a certain self-consciousness, attended by undoubted pleas- urable sensations. He was able to elicit the croaking of frogs and the aggressive movement of clenching the female as in the sexual act. The stimuli which Goltz had to apply in this instance differed materially in character from those which elicit repulsive movements. The effective stimuli were of a gentle sensory and non-painful character. Gentle friction between the shoulder blades sufficed to provoke a croak in frogs deprived of their fore-brains; and (slight) pressure brought to bear on, or fric- tion of, the breast and flexor surfaces of the arms was followed by an embrace of any thing which had been placed within reach of the arms. The transmission of such gentle stimuli through reflex gray substance could not take place in the face of any great resistance, and, being permitted, it compels us to assume the existence of a distinct centre which is reached at once by external stimuli. The " centre," excited in the croak-experiment, lies in the mesencephalon, while the reflex centre for the sexual embrace in male frogs must be located in the cervical portion of the spinal cord. The term "embracing spasm" (Umarmungs- krampf) will do justice to the powerful muscular action, resulting probably from a hyperaesthesia of the frog's spinal cord, produced during heat by the nerves of the testicles, but existing for i88 Psychiatry. some time after the removal of the testicles. The exciting cutaneous stimulus need be merely of the slight (unpainful) intensity mentioned above. Hence we may infer that even in the case of frogs that are brainless as far as consciousness is concerned, the correlated effects (in the central organ) of aggressive movements and the accompanying circumstances differ from those attending repulsive movements, with which consciousness associates the sensation of pain. These two orders of correlated (secondary) effects are on the whole diametrically opposed to one another. I.—The stimuli are gentle and not painful; they must be con- veyed to certain centres direct without irradiating and without overcoming any resistance or inhibition. In the sexual spasm of the frog, the conduction of nerve-force was assisted, and not inhibited, by a periodic increase in the excitability of the centre. Just as the consciousness of a painful impression is based upon, or is attended by, inhibition of nervous impulses, so the con- sciousness of a pleasurable impression is attended by the free transmission of nerve-force. II.—In the one case, the irradiation of a stimulus adapted to the excitation of pain, sets up activity in the centre for the vaso- constrictors followed by increased arterial pressure (active arterial anaemia), as concomitants of repulsive movements; and in the other case, there is no irradiation produced by external stimuli, no inhibition, and no increase in arterial pressure accompanying aggressive movements. Later on I shall be able to show that the movements of aggression are associated with a diminution of blood-pressure, a dilatation of the arteries entailing the so-called functional hyperaemia. Ditmar's reflection, that increased arterial pressure keeps step with the process of sensation, does evidently not apply to the generation of aggressive movements. III.—The functional arterial dilatation necessarily produces an apncetic phase, in consequence of the increased tissue-breathino- of the nervous elements, and will bring about a chemical change also, differing from the one effected by a painful stimulus, which called forth a dyspncetic phase of nutrition in these elements. These antitheses are not artificial and far-fetched ; they are sug- gested by the fact that in the reflex centres aggressive and repul- sive movements inhibit one another, because of the different pro- cesses set up by the one order of movements or the other. Goltz has shown that the ordinary croak cannot be elicited, if painful Inhibition of Repulsive Movements. 189 stimuli be simultaneously applied; and that the sexual spasm will be inhibited if any part of the frog's body be at the same time touched with acetic acid. And, on the other hand, the great ex- citability of the sexual spinal-cord centre under the influence of the testicle-nerves may, during the period of heat, inhibit the repulsive movements which would follow painful stimuli, were they not inflicted during the sexual act. But have we any facts which will enable us to say whether the same accompaniment of vaso-motor innervation, of differences in the chemical changes of nutrition, attends the secondary mechanism of conscious movements, incited and set into action by subcortical reflex acts, which (movements) are based upon the revival of cortical images, and are effected through the mediation of association-tracts; and who will say whether among the im- pulses of the Ego, and within the bounds of free will, we shall be able to discover and to discriminate between a series of repulsive movements which had to overcome a resistance to nerve-conduc- tion, and aggressive movements which have had free pass through the nerve-tracts engaged in their excitation or transmission ? Although it has been shown (p. 141) that the great expanse of cortical surface favors the restriction of certain functions to certain »localities, it is certain that apart from the association through the arciform bundles, and on the supposition that its gray substance represents a network of gray fibres, the condition of its structure permits the irradiation of stronger stimuli, as is the case in the spinal cord and the remainder of the central gray substance. Association and Irradiation are two very different processes. We shall show that the process of irradiation in the cortex inter- feres seriously with that of association. As in the spinal cord so in the cortex, vaso-constrictor nerves will be excited by an irradi- ating sensory stimulus. Among German authors, Eulenburg and Landois,1 as well as Hitzig, have shown that upon stimulation of the cortex the extremities of the opposite side grow cooler (through arterial constriction), and upon removal of the cortex the tem- perature of these parts increases (Hitzig) so markedly (through arterial dilatation) that it can be discerned by the hand placed upon the extremity of a dog so operated upon, and without the corroborative evidence of a thermometer.2 1 Centralblatt f. d. med. Wiss., 1S76. 8 For full literature vid. Exner : " Grosshirnrinde " ; Hermann : " Handbuch der Physiologie," Bd. II., p. 318. 190 Psychiatry. Let us take up the example discussed above (p. 184), in which we supposed a brainless frog to be exposed to an intense sensory impression (pinching with forceps) and to display an inhibition sufficient to overcome the stimulus. We then discussed the in- fluence of the stimulus upon the vaso-motor nerves, and the repulsive movements executed by motor nerves ; but let us vary the example by supposing an animal to be in full possession of its brain. A person who has been teazed with forceps, or who has under- gone the pangs of an operation, will recognize this intense irri- tation as an impossible (unbearable) act of perception. Evidently irradiation meets with severe and widespread resistances in the gray substance of the cortex as well as in the central gray sub- stance. In this respect the pain answers to a sensation of inhibi- tion. The inhibition, obstructing the path of this irradiating stimulus, calls for an exhausting amount of excitation, and this effects inhibition of other cortical functions, and sets a limit to the activity of the cortex ; it inhibits attention, thought, and the association of ideas as well. Secondary conscious movements of repulsion are incited (p. 184 et seq.) to ward off the hands or in- struments that have inflicted the torture. As concomitant con- ditions, we get paleness, nausea, loss of consciousness, together with slowness of pulse, with or without convulsions, or uncon- sciousness together with rapid pulse and universal convulsions. Loss of consciousness, attended by nausea and slowness of pulse, may be ascribed to irradiation affecting the vagus—the cerebral nerve of the heart,—and to anaemia of the brain, due to the suppression of the systole. Though restricting myself to the above-mentioned facts of experimental physiology as evidence of the vaso-constrictor influence of the stimulated cortex, and reserv- ing other striking proofs of the fact (Kussmaul, Nothnagel) for the clinical chapters of this book, I have said enough to infer from the importance of the cortex as a vaso-motor centre, and from the above-mentioned relation between strong sensory impressions and an increase of blood-pressure, that the conduction of strong sensory stimuli into consciousness (into the cortical " gray ") increases the arterial pressure and arterial constriction, thus setting up active anaemia. Whether this leads to actual un- consciousness or not, we may say that the sensation of pain is associated with increased narrowing of the arteries. But con- traction of the arteries implies a chemical change, the dyspncetic Actual Irritation not Necessary to Produce Pain. 191 phase of prosencephalic nutrition, and, united to the latter, the disagreeable sensation of pain. This is the less hypothetical, inasmuch as external chemical changes, such as accompany diffi- culty of respiration in impure air, are attended by a sensation of discomfort and restlessness, leading to repulsive movements, loss of consciousness, even to swooning with convulsions. It has been shown, therefore, that strong sensory stimuli excite in a reflex way conscious movements of repulsion, and in originating sensa- tions of pain introduce inhibition, arterial contraction, and dyspnoea of the elements of the fore-brain. It is not alone the perception of actual pain which incites movements of repulsion, but the mere sight or touch of, or a sound from, objects which are associated in the fore-brain with the idea of pain, danger, or death, excite, as the pain itself, move- ments of repulsion, and create in the cerebral cortex all the con- ditions of subjective torture which we have learned to regard as concomitants of real pain. If the mere sight of the sharp end of a knife, of loaded fire-arms, of a wild animal let loose, of a fire, a corpse, an operation on others, or the sight of blood, suffices to produce loss of consciousness or swooning attended by the sensa- tion of inhibition of thought, or impels the witness to take to flight, we must suppose that our ideas of pain are intimately con- nected with the revival of reminiscences of these objects, and that this subjective pain is of sufficient intensity to arouse all the correlated physiological sensations accompanying genuine object- ive pain: namely, inhibition, increased arterial pressure, dyspncetic phase of nutrition, and repulsion. Referring again to the former example of the consequences of an intense sensory irritation upon the spinal cord, we observe that the entire complicated primary form of the reflex mechanism is transmitted secondarily to the fore-brain. The instances cited immediately above referred to associations started by the actual perception of objects calcu- lated to produce pain ; but the momentary painful stimulus was lacking. Physiologically speaking, the stimulus bears a special characteristic sign, resulting from the excitation of peripheral sen- sory nerves, and, more particularly from excitation of their terminal apparatus ; among such signs may be classed cutaneous irritants, dazzling light, unusually strong waves of sound, etc. Pain is classified with the Feelings; but it is distinguished from sensory perceptions by its widespread irradiation which interferes with localization. 192 Psychiatry. Feelings without physical pain are termed emotions (Ergriffen- heit). We are here concerned with painful emotion, psychical pain. That painful emotions depend solely upon associations, upon in- ferences pointing to the perception of pain, can be proven by a sim- ple analysis of the objects which excite pain. The retinal images of a tame and of a wild animal, of an indifferent person or of one whom we fear, are projected upon the same retinal area, possess the same color and the same intensity of light. An indifferent red fluid and blood leave the same impression upon the retina. Mere perception itself cannot excite emotion, but the associa- tions united to the former can. It is, therefore, the mechanism of the hemispheres only—the process of thought—which excites psychical pain and movements of repulsion, as well as the arterial contraction which may terminate in swooning, and in engendering fear and a feeling of incapacity for action (Unmoglichkeitsgefuhl). But sensory impressions are not needed to arouse the associa- ions connected with emotions ; recollection of painful impres- sions may revive such associations. Boerhave relates a story in point : He says that he passed a spot where years ago the smell from the cadaver of a horse made him vomit, and that the mere recollection of this occurrence produced nausea anew. Without wishing to dilate upon such corrob- orative evidence, we may add that reflexes attending painful emotions, such as crying, may be excited by reminiscences, as is the case too with the centre for the contraction of the blood-vessels. Further details about the connection between reflexes and emo- tion may be called from Domrich : " Die psychischen Ziistande " Jena, 1849. Explana- tions will be given later on. Equivalent in value to cortial images are the symbols of language associated with these. The news of the death of a person whose image would frequently be revived in our brain by the most manifold associations, and which when presented to the brain would arouse all sorts of secondary presentations and pleas- urable emotions, who was bound up with a good portion of our thoughts—such news, I repeat, would cause inhibition of all these associations, and the place of easily excited associations (through repetition) will be usurped by others that are not yet easily transmitted. Marked inhibition of nervous impulses from the fore- brain excites,likethe inhibited conduction of painful sensory stimuli, or the suggestion of torture, a concept of the impossibility of counteracting this inhibition which may ultimately lead to suicide. A physiological process occupying much time, and consisting in the dissolution of now purposeless associations and the formation of Processes of 7 bought and Their Relation to our Moods. 193 new ones, precedes the introduction of this death-news into the web of associations. Inhibition is attended by emotion and psychical pain. With inhibition and psychical pain there is connected in- crease of arterial pressure, which during an emotion may, by mere presentation to the mind, produce swooning. Inhibition or resistances on the lines of nerve-conduction through the gray sub- stance, as well as increased arterial pressure in consequence of strong sensory impressions, are physiological facts. Lastly, that the impoverishment of the brain substance in oxygen (Hermann) has the effect of a chemical irritant which excites dyspncetic respiration in the oblongata, and produces epi- leptic changes in the so-called convulsion-centre, is a well-established physiological fact. Since increased arterial pressure even in the cortex produces an impoverishment in oxygen, a dyspncetic phase of nutrition will be set up in the cortex as soon as the conditions of painful sensations exist. Sensation itself is the subjective form of perception of all these physiological processes ; it is, as it were, the expression of a special sense concerned with the nutritive phases of the cortex. The stimuli and their effects, which interest us in this connec- tion, may be of every conceivable degree of strength. The higher intensities of Feeling may be termed Passion, Emotion ; the lower intensities, Moods, Temperament. By calling the latter hampered and unhampered moods, we establish an evident analogy with the conditions which lead ulti- mately either to repulsive or aggressive movements. If we are actuated by appropriate motives, the pursuance of a complicated process of thought produces in us a rapid change of mood, according as this thought is hampered or furthered. If a beginner wishes to determine the rank of a plant in the natural system, he must perceive and associate in mind all those character- istics which have been united in such a way in his brain as residual images, as to enable him to properly classify this object in the botanical system. If he discovers a single characteristic which is poorly developed, this cirumstance will inhibit the association ; if he has forgotten the proper term for another characteristic, the flow of association will be checked also ; if still another quality of this plant does not harmonize with the others which he thought would suffice to determine its rank, further work in this direction is hampered; and if he cannot recall all the peculiarities of the family to which he supposes the plant to belong, then the very 194 Psychiatry. last step in the long process of association is rendered impossible —no conclusion can be reached. Confusion results, but confusion indicates an inhibited association of ideas. Displeasure is united with this inhibition of thought; the conception of a repulsive movement to give up the plant or to throw it away will probably follow very soon after. The hampered mood will grow more in- tense if the original motive was intense, say an examination in botany on the morrow. If the associations had not been checked, they would have formed a complete circle, beginning with the in- spection of the plant, taking in all possible associations on the way, and ending by identifying the real plant with the one constructed in the brain. During the development of this process of thought; the student would have experienced a pleasurable sensation, and a certain degree of happiness would have accompanied the attain- ment of the conclusion reached through this chain of associations. Every scientific investigator who endeavors to work out the answer to certain problems, whether it be his aim to determine the correspondence of two cerebral acts, say between the perception of a natural phenomenon and the conclusion reached by a long line of association, in the course of which he endeavors to evolve the conditions of this phenomenon in such a way that they appear as the conclusion reached by his own mental activity ; or any one who has successfully reached his Q. E. D. experiences a psychical happiness even while his thoughts are in full blast. This sensation (of happiness) is no doubt to be ascribed to a determination of arterial blood to the busied fore-brain, to a functional hyperaemia. Functional hyperaemia is the physical concomitant of thought, and in the succeeding chapter we shall endeavor to examine in detail the relation between these two processes. And the manner in which thought in general produces functional hyperaemia, I will endeavor to explain upon a physiological basis, and not merely by drawing an analogy between the fore-brain and other brain-organs when in a state of activity. Goltz explains the inhibition of the croak-reflex in the frog through painful constriction of one leg, by supposing the reflex centres to form a single complicated apparatus ; and having made the assumption, he contends that every mechanism will operate the more easily, the less the tasks it has simultaneously to perform. The croak-reflex is inhibited by the irradiation of another stimulus into the confluent gray reflex centres. Relation of Brain Nutrition to Mental Processes. 195 The cerebral cortex has two distinct tasks to perform: I. The innervation of processes of thought and of movements con- nected with these ; 2, the innervation of arterial vaso-constrictor muscles. The more inactive the brain is in the former respect, the more intense will be the constriction of the arteries. This latter process will be inhibited as soon as the first order of cortical activity is called into play; which means that arterial constriction diminishes during the innervation of mental processes. Hence a condition of functional hyperaemia is set up. Although different sensory stimuli reach special cortical areas, it was shown above that a revived memory, constituting the act of recognition, must be regarded as a complex phenomenon associated with secondary concepts, derived from different areas of the cortex. The majority of mental processes are based upon reminiscences or memories which have engaged the functional activity of widely separated regions of the cortex. Whence it follows that the functional hyperaemia will invariably affect circumscribed areas at some distance from one another; and this is equivalent to saying that a condition approaching a universal hyperaemia of the fore-brain will be established. A psychical condition, which engenders innumerable aggressive concepts, and puts no obstacle in the way of the expansion of associations, is attended by a feeling of happiness. Winning the first prize in a lottery will make the ordinary mortal happy; this feeling of happiness is the result of an unrestricted flow of asso- ciations. The wealth he has suddenly acquired puts a number of desirable objects within his reach ; with a number of objects his personality can now form associations which could formerly not be established from the lack of wealth ; a multitude of possible aggressive acts present themselves before his mind ; his brain enters into a condition of great though easy and untiring ac- tivity, attended by an apncetic phase of nutrition, thus standing in great contrast to the check put upon the flow of associations through the loss of persons or possessions. We have seen that the concept of motion, the revival of sen- sations of innervation linked to a long chain of associations, can arise in the mind unaccompanied by movements or acts of any sort. We are, therefore, impelled to the conclusion that the execu- tion of movements demands more powerful stimulation and stronger nutritive stimuli. The intensity of such motives for action vary with the feelings. It would be difficult to conceive of the 196 Psychiatry. existence of an animal not actuated by feeling, and, indeed, to conceive of one that is not influenced by the two sorts of feeling, of which the one provides motives for movements of repulsion, and the other for movements of aggression. An animal endowed only with processes of aggression, and lacking the conception of re- pulsion, or the power of performing acts of repulsion, would succumb to the inimical influences of sensitive and insensitive beings; while an animal that possessed no conception of aggres- sion, nor the power to perform aggressive acts, could not avail itself of the conditions of subsistence offered by nature. The phenomenon of (personal) freedom depending upon the variety of possible deeds, is best exemplified by the impulses which intensity of feeling imparts to our secondary individuality. Within the realm of the primary ego, the destruction of one's own body—death—constitutes the extreme concept of repulsion ;. yet through the intensity of feelings, joined to the secondary ego, infliction of death may simply be an act of repulsion by which it is intended to ward off destruction from other portions of our individuality, as in sacrificing our own lives to save the life of a person so dear to us that he has formed part of our own in- dividuality. Duty and Honor have become integral parts of the individuality, exceeding in the intensity of repulsive and aggressive acts which they arouse, the factors of the primary ego. Com- plications of the individuality may reach such an extreme that the reflection of the individuality in the brain of others becomes the leading motive ; and the primary ego is sacrificed in order to preserve an image in accord with our secondary individuality in the brains of the survivors. This constitutes the desire for so- called immortality. Let us turn now to the consideration of the anatomical corollaries and the results of the experimental investigation regarding the subcortical ganglia, which terminate cephalad in the diencephalon. The prosencephalic ganglion may be looked upon as a nodal mass among the centrifugal nerve-tracts of the cortex ; its conditions of excitation keep step with those of the cortex. The relation between the trunk ganglia, beginning with the thalamus, and the fore-brain is such that, although excitation of the former produces sensory apperception in the cortex, the greater the cortical excitation following upon the independent revival of cortical memories and of associations, and upon the exercise of thought, the more the influence of the subcortical centres Cortical Inhibition of Subcortical Ganglia. 197 will be diminished. This is cortical inhibition. I wish at this juncture to call particular attention to the incommensurability of cortical reminiscences with sensory perceptions—a fact of great importance in regard to all future discussions. Sensory perceptions are invariably referred to the external world, even in those cases in which, through blindness or amputation, the peripheral sensory surfaces have been removed. This is due to reasoning by analogy. By experience we have learned that , excitation of subcortical nerve-tracts and ganglia, which trans- mit sensory perceptions to the cortex, is due, in the first instance, to a peripheral stimulus ; and, therefore, excitation of these subcortical tracts and ganglia is invariably referred, by the laws of causality, to the external world. It never occurs to us, however, to suppose that the cortex is stimulated directly from the external world ; whence it follows that cortical images, which are due chiefly to excitation of the cortex, can never be invested with the qualities of true sensory impressions. That the peculiar formation of the prosencephalic ganglion de- pended upon its important connections with the anterior portions of the cortex was insisted upon on pp. 141 and 142 ; but although the nucleus caudatus and the nucleus lenticularis form one mass, their respective proportions are in striking contrast to one another. It is safe to say that the intraventricular expansion of the nucleus caudatus does not vary much in its relation to the brain-trunk throughout the mammalian series. In man its basi- lar portion, lying above the lamina cribrosa anterior, is flat, while in animals with highly developed olfactory lobes it is convex, and exceeds in size this formation in the human brain. Excess of development in this one particular is evidently de- pendent upon the presence in the basilar portion of the caudate nucleus of nerve-tracts coming from the olfactory lobes. Magen- die was the first to recognize the corpus striatum as an organ connected with locomotion, and Nothnagel discovered in the cor- pus striatum of the rabbit a nodus cursorius, which, when excited, compelled the animal to run forward. The movements of ani- mals possessed of highly developed olfactory lobes are influenced chiefly by olfactory impressions, for which reason we can readily understand the connection between these two physiological fac- tors and their relation to the structure of the nucleus caudatus. The lenticular nucleus is sparingly developed in mammals, in great contrast to the development of this ganglion in man and 198 Psychiatry. monkeys. It is well developed also in the brains of the mole and bat—whose anterior extremities subserve special forms of motion. In cases of hemiplegia, due to destruction of the lenticular nucleus, paralysis of the upper extremity exceeds in intensity that of the lower. Climbing animals, including the monkey and man, have from the very first adapted their anterior extremities to the performance of complicated movements, and in man the upper extremity has learned to do the bidding of psychical im- pulses ; but man and monkey are the two forms which are char- acterized by the possession of the most highly developed nuclei lenticular cs. Quite recently three cases of encephalitis affecting the right island of Reil have been reported, which were followed by monoplegia affecting the left upper extremity (Brodeur, Ray- mond). Severe paralysis cannot, as a rule, be attributed to lesions of the cortex, and according to Munk the cortical area governing the movements of the upper extremity is situated at a distance from the island; this monoplegia must undoubtedly, therefore, be referred to the lenticular nucleus which stands in close con- tiguity with the island of P.eil. Whence we conclude that in addition to its well-known relations to the hypoglossal and facial nerves, the lenticular nucleus has special relations to the move- ments of the upper extremity. The remaining masses of gray substance do not form part of the prosencephalon, but lie within the walls of the primary medul- lary tube, which extends originally only as far as the anterior cerebral vesicle. The relations of the powerful di-encephalic ganglia, the thalami optici, have already been discussed (p. 162), also their prominent connections with the optic tract. Due attention was bestowed upon the relations proved by physiological experiment to exist between the thalamus and the upper extremities, and thus the influence of one thalamus—whose mechanism is set into activ- ity by retinal impressions—upon different muscular groups in both arms was explained anatomically by the origin in the thalamus of decussating and non-decussating spinal-cord tracts. I wish to add that the greater caudal expansion of the human thalamus, exceeding as it does in this respect the thalamus of all other mammalian forms, may possibly bear a definite relation to the greater use of the upper extremities. The anterior ganglion of the thalamus is not more highly developed in man than in other mammals, and naturally enough, Relation of Thalamus to Visual and Muscular Senses. 199 -too ; for there is an evident connection between this part and the gyrus fornicatus, the cortical portion of which Munk connects with the sense of smell. This conclusion is borne out by its connection with the olfactory lobes, and the relatively great development of its substance. From an anatomical point of view, this anterior protuberance may also be considered a nucleus cauda- tus. The size of the human thalamus depends largely upon the development of the pulvinar, which arches over and beyond the corpus geniculatum externum, while in all mammals, with the exception of the monkeys, the corpus geniculatum externum tends upward toward the thalamus. But on the other hand the corpora quadrigemina are less developed in man. No ganglion has as broad connections on all sides with the cortex as the thalamus op- ticus ; on superficial examination it appears to be connected with the entire corona radiata. And since this broad expanse of cortex joined to the thalamus is connected with almost every cortical function, we must infer that the thalamus has an important bear- ing upon functions of widely different character. The functional activity of an animal which has not been de- prived of its thalamus, is in no wise impaired, save that it lacks the customary centrifugal impulses dependent upon cortical reminiscences. There is good reason, therefore, to believe that all forms of sensibility are represented in the thalamus and cor- pora quadrigemina. In corroboration of this belief we might cite the very great influence sensory impressions exert upon move- ments as long as the thalamus remains intact. First and fore- most we must recognize the projection of the retina upon the optic thalamus; and from Nothnagel's pathological experi- ments we learn that visual disorders result from lesion of the posterior segment of the thalamus—a conclusion which is in perfect accord with anatomical facts. As long as the thalamus is uninjured, and only then, animals are able to avoid obstacles thrust in their way, and birds that are thrown up in the air show by their behavior that they can measure with their eyes the distance to, and the direction of, the ground upon which they are bound to land. The posses- sion of a thalamus enables an animal to creep through a narrow opening between objects placed before it—a fact known even in Magendie's day ; or, as Goltz observed in frogs, enables an animal to jump past an object placed in its way, provided there be a sufficient stimulus for escape. Facial reflex movements seem also 200 Psychiatry. to be dependent upon the thalamus. On the one hand patho- logical cases are on record in which, in spite of an existing facial paralysis, the reflex movements involved in laughing, crying, in giving expression to pain, and in the protective closure of the eyelids, were well executed (Nothnagel, " Topische Diagn. der Gehirnkrankheiten ") ; but in all such cases it has been remarked that the lesion did not affect either the thalamus or its medullary radiations. And on the other hand Longet and Schiff have called attention to the grimaces of cats (which have been deprived of their hemispheres) following the application of Tincture of colo- cynth to the tongue. I do not mean to defend the term " psychi- cal reflex," nor to insinuate that the projection which receives its innervation from the hemispheres (from consciousness) suffers an interruption in the thalamus ; for of all negative physiological truths relating to the thalamus, there is none more certain than that so-called volitional paralyses are entirely independent of thalamic lesions ; and yet, if we do not wish to give too simple an interpretation to facts of great subtlety, we must keep in mind that the thalamus possesses a reflex mechanism of such a high order, of such marked psychical characteristics, that Goltz, who is not satisfied with the term " reflex," would designate the complex functions of these higher physiological mechanisms of the brain as " the power of adaptation." And then, too, the views I have expressed regarding the genesis of volitional paralyses, which were implied in the assertion that volitional movements are based upon the primary motor images derived from reflex movements, must not be accepted as a negation of the fact that the co-ordinating power of the fore-brain may enable it to exert an educational influence upon reflex actions. Lotze was no doubt justified in giving the following explana- tion of Pfliiger's so-called psychical spinal-cord-functions: He contends that, in the young animal, movements of expediency, ■ like those of cortical motor acts, are not yet to be ascribed to the spinal cord, but that definite cerebral co-ordinations, which are accustomed to utilize certain spinal nerve-tracts, leave an im- pression of fore-brain activity of such a nature that, after the removal of the brain, reflex innervation is directed to the groups of nerve-tracts most frequently engaged in the transmission of cerebral impulses. Anatomically, this can be proved more readily for the spinal cord than for the thalamus, for the former is con- nected with centrifugal cortical nerve-tracts, while the thalamus is Injluc7icc of Conscious-Movements upon Reflex Actions. 201 not so connected. The thalamus itself is the centre for and origin of the centrifugal tracts in the tegmentum of the crus ; their in- nervation is dependent upon stimulation of the thalamus. On the strength of the discussion at the beginning of this chapter con- cerning compulsory positions of the anterior extremities following lesions of the thalamus, we conclude that the fibres radiating from the cortex into the thalamus constitute centripetal nerve-tracts which conduct to the cortex the sensations of innervation created by motor processes started in the thalamus. We must necessarily, however, premise the existence of centrifugal, cortical tracts in the prosencephalic ganglion and in the internal capsule, initiating the same forms of movement, and emanating from the same areas of the cortex, to which the thalamic radiations conveyed the innervation-sensations of such movements. Though the co- ordinating fibres belonging to the cortex proper may refine upon and vary these forms of movements, they could not exert any in- fluence over the mechanism of the thalamus unless we assumed the twofold conducting power of cerebral nerve-fibres—a power which Dubois-Reymond proved that the excised nerve possessed. The association-tracts of the fore-brain possess beyond a doubt such a twofold power of conduction ; for of two associated corti- cal reminiscences, either one, when revived, will recall the other. I can find no grounds for denying a twofold nerve-conduction in the thalamic projection-systems conveying sensations of innerva- tion to the cortex ; and through these systems, movements which have been modified by cortical co-ordinations may react. But more of this later on. It was in regard to physiognomical ex- pression, which is not confined solely to the facial nerve, but is a result of the activity of the muscular mechanism of our entire body, that Darwin asserted the apparently paradox principle that the form of volitional movements in the ascendant are changed to reflex movements in the descendant. The inheritance of physiognomi- cal expression I do not consider proved. At all events Darwin could have made his views plausible only by showing that con- scious movements originated in reflex actions. In that event a reciprocal effect might be conceived ; reflexes might be proved to be the primary roots of the simpler manifestations of the con- scious motor mechanism ; and later on the cortical co-ordination might generate more complicated forms of movement which, from the presence of a twofold nerve-conduction, would react upon the motor functions of subcortical centres. 202 Psychiatry. The multiplicity of relations of the thalamus as a centre of automatic movements, and as such, its relations to the sensations of innervation, its bearing upon the sense of smell (restricted to its anterior protuberance), and its relations to sight are in keep- ing with the anatomy of this ganglion as described in the first section of this book (pp. 31, et seq., 48, 92, 101, 141, and 142). The dependence of physiognomical expression upon cutaneous sensibility leads us to look for the anatomical substrata of such expression in the nerve-tracts of the thalamus. At this juncture it is incumbent upon me to fill in a gap in the anatomical de- scription given above. I wish to add that the fibres of the lem- niscus, the fillet, derived from the trunk ganglia, do not issue merely from the corpora quadrigemina, but that some of its bundles, situated to the front (cephalad) of the corpus bigeminum superius (Figs. 55, 56, behind ss), and inseparable from the inner- most bundles of the brachium corp. bigem. inferius, originate undoubtedly from the thalamus. In searching for the anatomical substratum of the possibly sensory functions of the lemniscus, we must necessarily keep in mind the quadrigeminal origin of the fillet from the brachium corp. bigem. sup. Bundles of the lem- niscus constitute the anterior portion of the brach. sup. (p. 102), while the posterior margin of the latter contains fibres connecting the corpus geniculatum externum with the corpus quadrigeminum. These bundles of the lemniscus stand in close contiguity with the latter, centripetally conducting fasciculi of a sensory organ, and issue from the same regions of the cortex in which are contained optic radiations, as well as the sensory bundles of Tiirck which pass through the internal capsule and the pes pedunculi. There is no strict anatomical proof for the assumption that the occipital lobe is the chief central organ of sight ; but other radiating fibres connected with this division of the cortex furnish ample proof, as was stated at the beginning of this section (p. 142), for the belief that it is connected with the special organ of touch (the skin). Following the track of the lemniscus into the spinal cord, we find that the fillet is continued into the outer portion of the funiculus lateralis (p. 134), which Mischer has conclusively shown to con- tain sensory tracts. The lemniscus, therefore, connects the thalamus with the corp. quadrigemina through tracts conveying reflex influences of tactile stimuli. The anatomical connection of the lemniscus with the oli- vary bodies-—from which the posterior columns emanate (p. 134)— Thalamus and Corp. Quadrig. are Subcort. Centres. 203 is also worthy of remark. According to Renzi, detonation pro- vokes movements of the eyes in frogs possessing normal thalami; we must, therefore, suppose connections to exist with the sense of hearing. Remembering the entrance of central auditory tracts from the cerebellum into the superior peduncles, and on the other hand the connection proved by Wernicke to exist between the thalamus and the nucleus ruber of the sup. peduncle,—re- membering these facts, we may say, although I wish to speak reservedly regarding the importance of these peduncles as an auditory chiasm, that a connection of the thalamus with the laby- rinth is not beyond the bounds of anatomical possibilities. In the oblongata the voice- and auditory-centres crowd upon each other. Possibly the motor function which Bechterew ascribes to the thalamus in locating a centre for screaming in that ganglion, and the " croak centre " which Goltz locates in the region of the mid-brain, may argue in favor of the existence of anatomical connections between the VIII. nerves and the mechanism of articulation. Since voice manifestations in animals represent the acoustic factor in their physiognomy, we might consider a phy- siological connection established between this function of the thalamus and the above-mentioned function of reflex mimical ex- pression. Possibly, also, the difference of opinion between Goltz and Bechterew regarding the voice-centre might be explained by supposing that functionally at least the thalamus and mid-brain are not strictly separable. Caudad of the mesencephalon (mid-brain) begins the region of the oblongata, which cannot be separated easily from the pons, and which, properly speaking, does not contain centres for loco- motion, but for restricted movements only. Goltz mentioned as characteristic of animals deprived of the mid-brain and cerebel- lum, in contradistinction to. those possessing a spinal cord only, that the former could change from the dorsal to the abdominal position. Undoubtedly, however, the pronounced influence of sensations upon bodily movements, such as leads us to presup- pose a centre (for the maintenance) of equilibrium, is to be ascribed to the region of the'thalamus and mesencephalon. The possession of this centre enables a frog, deprived of its fore-brain, which had been made to squat on the palm of the hand, to change its position step by step when the hand is tilted down- ward, until it is landed on the dorsal surface. To the di- and mes-encephalon we must, therefore, look for the subcortical sen- 204 Psychiatry. sory centres, which help to perfect all possible movements of the highest order, with the aid of impressions they receive, say, from the retinal image and the surface of the skin. In the modifica- tions here imposed upon the forms of movement by retinal im- pressions, we may recognize the first beginnings of the visual impressions of space ; though our fore-brain consciousness does not take cognizance of them until they have been handled by the mechanism of association. The architecture of the corp. quadrig. (Figs. 38 and 39) showed that tracts leading to it from the main external terminal mass of the tractus opticus—the corp. genie, externum—through the posterior margin of the brach. corp. quadr. superius, take a longi- tudinal course in the mid-brain, thus avoiding a union with the formation of the lemniscus. But these longitudinal bundles are united to the gray substance of the aquaeductus Sylvii in conse- quence of a radiating course which they enter upon, after taking up quadrigeminal cells, and then traversing the layer of the lemnis- cus. In this gray substance the nuclei governing the movements of ocular muscles are imbedded. This allows us to conclude that ocular movements are stimulated by retinal impressions, and that this mechanism constitutes a simple automatic apparatus. Since both the gray substance of the corpora quadrigemina and the nerves supplying the ocular muscles are projected upon the cere- bral cortex, a union there takes place between the sensations of innervation derived from the ocular muscles and the impressions of different retinal areas, as was explained by Fig. 63, p. 181. The sensations of innervation of the ocular muscles, together with the projection of individual retinal areas, established in the cortex the local signs which help us to determine our whereabouts in space. Anatomical facts would go to prove the truth of the old view of Wundt, now, however, abandoned by him, that the mechanism producing the psychical conception of space is started into activity by reflex processes. The union of retinal impressions with muscular innervations in the corp. quadrigemina constitute, in my opinion, the primary reflex process assisting in establishing the conception of space. The nature of this reflex process would be similar to the one involved in the closure of the eyelids fol- lowing upon conjunctival irritation (p. 156, and Fig. 59). When analyzed into its primary factors, this conception of space will be seen to consist of sensory perception and sensations of innervation which have been secondarily transmitted to the cortex. To the Centres for Inter7ial and External Ocular Muscles. 205 entire di- and mes-encephalon Eckhardt's remark is applicable : that, " experience renders it highly probable that visual percep- tions attain a certain degree of perfection in the thalamus." But we have further evidence to show that in addition, to its relation to the centre of equilibrium governing locomotion—a centre which may possibly be connected with the functional mechanism of the corp. quadrigemina,—the thalamus exercises an important influence over the movements of the upper extremities, as was demonstrated by the pathological case exhibited in Figs. 61 and 62 ; and this will be corroborated by the observation of Noth- nagel, that after destruction of the thalamus, the fore-limbs, which had been extended, cannot be retracted. This is due to the loss of the focus for the sensations of innervation engendered by the movements of these limbs. The gray substance at the posterior margin of the third ventricle and around the aquaeductus Sylvii was investigated by Adamiick and again by Hensen and Volckers. The latter contend that the cen- tres for the intra- and extra-ocular muscles succeed each other in the following order : In the posterior portion of the gray substance of the third ventricle lies the centre of accommodation, to the front (ventrad) of the centre for the sphincter pupillae, both centres being connected with the most anterior III. root. At the boundary between the III. ventricle and the aquaeduct. we come upon the centre for the rectus internus, and to the outer side from this the VI. centre. At the cephalic end of the floor of IV. ventricle immediately below the corpora quadrigemina, the centre for the vaso-motor fibres of the iris is located, which, in the opinion of some, are solely responsible for the dilatation of the pupil by influencing the volume of the pupillary tissue, while others feel called upon to assume the presence of a special dila- tator pupillae. However this may be, this region (containing the above centre) lies immediately adjacent to the general vaso- motor centre of Owsjannikow, and, assuming irradiation to take place, we may be able to explain the simultaneous occurrence of dilatation of the iris and of constriction of the arteries. Immediately below the gray floor we are confronted by the descending arm of the formatio fasciculi long, posterioris. Wernicke had ample reason to trace one origin of this latter system to the lenticular nucleus; the projection-system of this nu- cleus would thus take a shorter route to the central motor roots and nuclei of the cranial nerves scattered throughout the central 2 06 Psych ia try. gray substance. The variable thickness of the posterior longi- tudinal fasciculus, its connections with the trigeminal and acoustic tracts, preclude the thought that it is principally the projection- system for the motor nuclei of the brain ; its purpose seems rather to be functionally to unite different levels of the central gray substance and to effect a co-ordination of movements. It may contain nerve-tracts uniting the abducens with the topo- graphically higher centres of co-ordination superintending the play of the ocular muscles. But the nerve-nuclei of the mesenceph- alon, of the pons and of the oblongata are joined by decussating fibrae rectce to the cortical bundles of the pes pedunculi. As was exhibited in Fig. 43, et seq., these fibrae rectae, after traversing the deep transverse fibres of the pons, extend to the anterior longitudi- nal bundles and in the oblongata to the pyramids. The oblongata, which but for its cerebellar relations, need not be considered apart from the pons, impresses us with its great importance as a vaso-motor centre. Owsjannikow has studied the oblongata most carefully from this point of view. This author has demonstrated that removing layer upon layer of the brain- trunk causes a fall of manometrical pressure, as measured in the carotid, even before the pons has been reached, which signifies that that portion of brain substance has been destroyed which, when normal, maintains the blood-vessels in a state of definite contraction. Blood-pressure does not depend altogether, as Bezold would have us believe, upon the heart and an increase of cardiac innervation, but in an opposite direction, chiefly upon the resistance which the contraction of the fine capillary vessels op- pose to the forward movement of the blood. Strieker very properly remarks that we are here concerned not only with the contractility of the finer arteries, but also, as has lately been shown, with the smaller capillaries endowed with nerves, and even with the more delicate veins. The blood-pressure ceases to fall as soon as the sections have been laid within 3-4 mm. above the apex of the calamus scriptorius. If the destruction by layers has not transcended the level of the origin of the facial nerve, the dimin- ished blood-pressure can be restored by irritating the sciatic, the trigeminal, or the auricularis magnus. But from this point on, the possibility of restoring the blood-pressure by stimulation of sensory nerves sinks until we reach the before-mentioned lower limit of the centre for the vaso-constrictors. Owsjannikow con- sequently distinguished between two vaso-motor centres: an The Question of Vaso-Dilator Centres. 207 automatic centre extending farther cephalad, and a reflex centre extending farther caudad, but for the most part both centres occupy the same level. He regarded the relations of these two kinds of centres to the innervation of blood-vessels very much as I did that existing be- tween primary and secondary movements of skeletal muscles. He insists that the automatic centre is subject to psychical influ- ences, as is shown by the influence of the emotions on the dilata- tion of blood-vessels ; and that the other is a reflex centre for the vaso-constrictors. From an anatomical point of view we may make the following distinction: the one centre innervated from the cortex belongs to the anterior division of the brain-trunk (pes pedunculi and stratum intermedium), say, possibly to the re- gion of the crus cerebri, and at all events to the anterior division of the pons ; while the reflex centre is included in the various groups of cells which are interwoven with the posterior (tegmen- tal) segment of the pons and oblongata. Ditmar thought the reflex centre might possibly be located in the superior olivary body of the oblongata. The extension farther forward of the centre governing psychical influences can be accounted for by the fact that the tegmentum begins to develop lower down than do the structures of the pes pedunculi ; and the termination of this cortico-vascular centre at a higher level would be sufficiently ex- plained by the fact that the pyramidal tracts get rid of their gray substance before reaching this level. Upon the mooted question of vaso-dilator centres I will not now enter. It would appear as though they simply regulated the antagonistic influences of the circular muscles, and exerted a secondary influence upon the width of the vessels ; furthermore, that a genuine dilatation resulted only from a lessened stimulation of the vaso-constrictors. It is, to say the least, very significant that if they (the vaso-dilators) are to be forced into action by stimulation of the sciatic, the latter must have been cut across several days previously, which compels us to assume that an ante- cedent condition of exhaustion of the vaso-constrictors paves the way for the subsequent preponderance of a vaso-dilator influence. The known phenomena of dilation proper have reference to the periphery—to cutaneous irritants. Inasmuch as these phenomena depend, according to Vulpian, upon peripheral ganglia, we can make no application of them to the central processes now under discussion. We take no notice for the present of an active pro- 208 Psychiatry. cess of vaso-dilation which has been most carefully investigated by Goltz and Strieker. The acoustic region occupies about the middle portion of the oblongata (Fig. 17, p. 32). Here our morphological attain- ments will stand us in good stead ; for, as Burdach puts it, the structure and significance of cerebral organs are united by a secret bond. The auditory nuclei are spread along the entire line of those central motor masses from which the nerve-roots emanate, and which are engaged in the production of sound, including the facial, hypoglossal, and vago-accessory (laryngeal) nerves ; also the respiratory and particularly the expiration-centres, which play an important role in this mechanism. Thus we find the conditions exceedingly favorable to the reflex origin of auditory communi- cations ; we have given a focus which functionally contains all the elements of speech, since speech is based upon the repetition of heard sounds. But this is not intended to be a teleological conception of our organization. Since sensory stimuli irradiate in the gray substance we must expect stimulation even of the lowest lumbar and sacral nerves to irradiate into other areas, including the sound-producing centre. This is shown in the re- flex cry following in earliest childhood even, upon the traditional educational handling of the skin over the glutei. According to Pfltiger's laws, particularly favorable conditions exist for the translation of sensory stimuli into reflex movements of a special character, if the gray substance in which a sensory nerve terminates, and that from which certain motor nerves originate, lie in adjacent levels of the cerebro-spinal axis. Stimu- lation of the auditory nerve will, according to Pfliiger's laws of reflexes, excite most readily movements of the sound-producing mechanism. The primary fundamental reflexes of speech are based upon a type of structure as simple as that of the ordinary spinal-cord section ; upon centripetally and centrifugally conducting roots of gray substance. A sound constitutes an auditory stimulus, and a sound results from the excitation of this sound-producing mechanism. It is the function of the motor nerve-tracts engaged in this process to effect a form of movement which will reproduce the contents of the original stimulus which was a sound. This is a purely imitative process. Just as the reflex-centre, mediating between a conjunctival nerve and the sphincter palpebrarum, transmitted to the cortex through the cortical projection-fibres a sensory stimulus and sensations of in- Imitative Mechanism of Sound. 209 nervation, so, in the same way, the stimulating sound, the sensa- tions of innervation derived from the muscles of this sound-pro- ducing mechanism, and the produced sound, are associated with one another in the cortex. Through association, an auditory stimulus must consequently lead to the imitative mechanism in- volved in the reproduction of sound. For the secondary motor mechanism of speech which goes on developing by an endless number of co-ordinating acts in the fore-brain, and for the methodi- cal imitation of heard syllables which is at the very root of the acquisition of language, there is a good anatomical basis. This consists in the presence of a simple reflex-centre which effects and permits a joint action of the adjacent nuclei of VII., VIII., X., and XII. cerebral nerves with the nerves of expiration. In the preceding section (p. 120 et seq.) I dwelt upon the con- nection existing between the auditory fasciculi and the cerebellum. Two physiological facts argue in favor of such a connection : the importance of the cerebellum as an organ of motor co-ordi- nation is a firmly established fact; and equally well founded is the influence which auditory impressions exert upon the rhythmical evolution of co-ordinated movements in walking, dancing, as well as in the formation of notes in the rhythm of a song. But there is still another fact to be mentioned as corroborative of the above anatomical connection. According to all experiments, the nervus vestibuli is not auditory in function ; but, as Goltz, Mach, and Breuer, who repeated Flourens' sections of the membranous semi- circular canals of the labyrinth, have been able to determine, it is responsible for the cerebellar maintenance of equilibrium. De- struction of these membranous canals entails the forced move- ments attending static vertigo. Together with Briicke, I regard these movements as actions resulting from delusions which pro- duce abnormal sensations of innervation, as I explained on p. 164 to be the case with regard to the forced movements resulting from lesions of the tegmentum and the optic thalamus. Flourens holds that similar forced movements are produced whether the semicircular canals of the labyrinth or certain portions of the cerebellum be cut across. He says : " If the horizontal semicir- cular canal be cut, the animal will turn on its own axis ; if the anterior, the animal will perform a number of somersaults for- ward ; and if the posterior canal be cut, the somersaults will be made backward. Similar to these movements are the rolling movements observed after section of the pons, and the running 210 Psychiatry. backward and forward following upon section, in the same direc- tions as above, of the anterior and posterior pedunculi ccrebelli." The direction of the movement would invariably be parallel to the course of the fibres; thus the rolling movements subsequent to lesion of the pons take place from one side to the other. Huschke, quoting Hamilton, communicates a fact which has an important bearing upon cerebellar co-ordination. It is this : The cerebellum is far more developed in animals which are able to help themselves immediately after birth (such as the chick, pheasant, partridge, goat, colt) than in animals which are born blind, are helpless, and find great difficulty in learning to walk. In man the cerebellum is surprisingly undeveloped. If the brain- trunk be dissected out from the rest of the brain, as in the manner indicated in Figs. 15-17, and if the cerebellum be removed also, as shown in Fig. 17, then we shall be able to determine the pro- portional weights of three portions of the brain : (1) the brain- mantle, (2) the brain-nucleus from the island to the beginning of the spinal cord, and (3) the cerebellum. In the adult these parts hold about the following relation, 79 : 10.5 : 10.5 ; whereas in the brain of the new-born the proportions are, 83:11:5. Magendie and Demoulins found long ago that injury to the cere- bellum produces backward movements. No one at the present day objects to the view that the cerebellum is a way-station for the muscular sense ; ataxic phenomena may, but actual paralyses never do, result from pathological disturbance of this organ. The muscular sense is based upon sensations which, when modi- fied by disease of the cerebellum, give rise to numerous delusions, such as falling into a pit, and so on. We must represent the matter to our minds somewhat as follows : Those movements which assume definite forms under the guiding influence of the cerebellum have their single acts associated with one another in the fore-brain from childhood up; and the sensations of innerva- tions grouped together in the cerebral cortex are transmitted to the cerebellum through centrifugal tracts—say, the pontine fibres of the crus cerebri. The posterior columns originating in the cerebellum are enabled to convey to that organ sensations regu- lating the co-ordination of movements, making the intervention of the cerebral cortex in the execution of movements quite super- fluous. Here the muscular sense comes into play in the manner described by Spiess. The varying sensations perceived through the posterior roots, changing with each phase of a motor act, with Co-ordination and Sensation. 2 11 the tension and relaxation of the skin on the extensor and flexor surfaces of the extremities, with the change in pressure due to contact of different portions of the skin, are transmitted to the cerebellum through the mediation of the posterior columns; and in the cerebellum, through the action of its association-system, are utilized as sensations regulating the form of movements. Taken in this sense, the gymnastic brain-organ is also an organ of feeling. Considering that the movements of the eyes and the extremities are accurately co-ordinated, as in taking aim with a gun, we can readily understand why cerebellar lesions should be followed by deviations of the ocular muscles, which would come under the head of forced movements in the sense in which the latter have been before referred to. Even the simplest forms of movement (such as changing from the dorsal to the abdominal position, dis- entangling the interlocked legs in the frog—Goltz) are performed fairly well after removal of the mid-brain, as long as the medulla remains; but they are executed with greater accuracy when the cerebellum is uninjured, in which case sensation plays an impor- tant role. Some authors have proceeded still further in making the cerebellum an organ of sensation. Eckhardt mentions, in a critical review which he gives of the subject, that Foville observed a general diminution of sensibility following upon disease of the cerebellum, that Renzi found that destruction of the cerebellum entailed impairment of the visual and auditory senses, while Lusanna noted visual destruction only following a similar lesion. Among the remaining centres of the oblongata the so-called diabetes-centre is probably identical with the vasa-motor centre for the arteries of the liver, and the chemical results of the diabetic puncture are dependent altogether upon the paralysis of their circular muscles. In corroboration of this view, we may cite the fact that if after paralysis of the splanchnic nerve, which exercises an all-powerful influence over the blood-pressure in the abdominal cavity, a general hyperaemia is set up, there is not a sufficient hyperaemia of the liver to produce mellituria in spite of Bernard's diabetic puncture. Moreover, the oblongata is the centre for a number of forms of movement, partly or wholly inde- pendent of the will, which come within the domain of the nerve- roots originating in the oblongata. Closure of the lids, lachrymal secretion, deglutition, constrictions of the pharynx, of the larynx, and of the oesophagus, all are dependent upon the oblongata. Inasmuch as the lower portion of the calamus scriptorius in- 212 Psychiatry. eludes the centres for in- and ex-piration, it contains also the centres for all the modified forms of respiration, under the influence of nerves starting from the oblongata such as'laughing, crying, sighing, gaping, and sneezing. Of greater importance is the existence in the medulla of a centre for the contractions of the uterus and the vagina, which is repeated, however, in the lumbar enlargement of the spinal cord. Goltz found that mammals whose spinal cord had been cut above the lumbar enlargement could still bear young. The influence of the oblongata on the salivary secretion, through the n. lingualis and the chorda tympani, as well as upon the lachrymal secretion, depends upon its significance as a vaso- motor centre. According to the demands of an existing dysp- noea an irradiation takes place, from the mere innervation of the diaphragm such as is practised in ordinary breathing, to an innervation of all respiratory muscles ; to the extremities for the purpose of fixing the thorax, to the muscles of mastication for the purpose of " snatching " air.1 If there be a lack of oxygen the oblongata becomes a general convulsive centre; and this centre is situated, according to Nothnagel, between the lower por- tion of the pons and upper portion of the oblongata in man. The solitary bundle situated externally from the X. nuclei was termed respiratory bundle by Kraus. Beyond a doubt the oblongata also contains in the tract of the vagus a centre regulating the cardiac nerves. Finally, we are told that a lesion of the oblongata along the auditory protuberance produces derangement of hearing, and Renzi speaks of a spatium opticum extending from the corpora quadrigemina to above the auditory protuberance, injury to which causes amblyopia. In regard to movements, most authors are agreed not to attribute to the oblongata the power of co-ordinating locomotion. 1 " Luftschnappen." THE NUTRITION OF THE BRAIN. In the preceding section (pp. 138, 145) we contended that in order to understand the functional activity of the brain, it is necessary to attribute but a single transcendental quality, i. c, sensitiveness, to the ganglion-cell and its protoplasmic annexes. From this faculty of sensitiveness actual sensation and its con- comitant or resultant states, representing the functional activity of the hemispheres, can be evolved, provided two conditions be met: there must be adequate nutrition of the ganglion-cell, and peripheral stimuli. As regards the latter, the external causes of perception—the stimuli—are influenced by the peripheral end- organs of the nerves. The central nerve-tracts, which effect the transformation of external stimulation into sensation, were deter- mined by anatomical and physiological methods in the preceding chapters of this book. On morphological grounds we reached the conclusion that the brain was composed of two grand divisions : the cortex with its medullary substance—the brain-mantle ; and the brain-nucleus, en- veloped by the brain-mantle, and consisting of subcortical centres and their nerve-tracts. We remarked also that association was favored by the broad expansion of the cortex, while the possibili- ties of irradiation were increased by the accumulation of the sub- cortical form of gray substance in the ventricular cavities, and in the walls of the medullary tube. As regards the nutrition of the brain, the ramification of the arteries meets the requirements of " association " and "irradiation." In my monograph on the structure of the cerebral cortex (published in 1868), I observed that the broad expansion of pia, answering to the extensive sur- face of the cortex, provided the latter with the largest possible number of arterioles, all of about equal diameter, entering adja- cent portions of brain-tissue, and of which each one, represented to a certain degree, an independent circulatory area, and that in a mass of tissues supplied by a smaller number of larger arterial branches, it would be quite impossible for differences of arterial blood-supply to exist simultaneously in adjacent portions of that 2'3 214 Psychiatry. tissue. Whence it is to be inferred that the broad expansion of cortical surface, and the large number of arteries descending ver- tically from the pia into the cortex, are well calculated to permit partial, functional hyperaemia of separate cortical areas. This peculiarity in the distribution of nutritive supply would allow us to suppose that the so-called cortical "centres" could be in a state of functional hyperaemia, at a time when the other cortical regions were enjoying a functional rest. But we must beware against ingenious exaggerations of this theory of local- ization. I have shown, when treating of the nature of re- sidual images (p. 154), that in a single process of thought widely separated areas of the cortex are thrown into a state of functional hyperaemia. A detailed analysis of the complexity of residual images (special memories) with their manifold cind widely distributed areas of functional hyperaemia, will lead us to accept an opinion of Fechner, which I referred to in my monograph on the cortex. The excitation and nutrition of the brain are subject to a change between two phases: one of sleep, and the other of waking. Sleep implies a universal diminu- tion of activity, but waking by no means implies increased activity of every part of the cortex. Fechner looks upon the partially increased functional activity of the hemispheres in the condition of wakefulness—that is upon the phenomenon of re- stricted attention— as a limited wakefulness, and accordingly he supposes that during the state of cerebral activity extensive areas of the cortex remain in a state comparable to that of func- tional repose. Association as effected by the hemispheres does not therefore depend upon the possibilities of a single limited area of functional hyperaemia, but upon the hyperaemia of numer- ous cortical areas, united for combined simultaneous action by the aid of association-fibres. This end is furthered by the pecu- liar distribution of arteries derived from the pia. The multi- plexity of cerebral functions cannot be explained by comparing the brain with any other organ whose physiological function is di- vided equally among all its parts, and of which every portion re- ceives an equal supply of blood. We can explain why every portion of the spleen, of the liver, and of the lungs should receive an equal amount of blood, but we can understand also, why, in regard to the brain, all parts of which are never simultaneously active, the case should be different. The blood-supply in the brain is determined by the functional hyperaemia of the areas called into activity; Local Hyper a mias in the Brain—Brain Movements. 215 while in the case of other organs, blood-pressure and blood- supply are limited only by the resistance of their membranae propriae, or their trabeculae of connective tissue. In the physio- logical condition the skull appears to constitute a rigid wall limiting the quantity of blood in cerebral vessels, and controlling any general change that may take place in this respect; by reason of which it becomes an important factor in the nutrition of the brain. A closer examination into the details of this sub- ject will convince us that the skull regulates the pressure of the fluid within its cavity. If the brain were surrounded merely by rigid cranial walls, a partial change in the distribution of arterial blood would be conceivable. Functional hyperaemia re- sulting from arterial dilatation, set up and regulated by the brain itself, might be effected by a mechanism such as I have described on p. 195, and would be explained by the fact that the cortex itself is a vaso-motor centre. But within the cranium with its rigid walls a functional increase would be possible only upon one of two conditions : there would have to be either collateral arterial diminution (olygaemia), for which it would be difficult to suggest an appropriate mechanism, or a transfer of venous blood in the direction of the sinuses, thus leaving more room for the play of the arteries. But this sort of venous transfer would be alto- gether too slow; besides there could be no continuous action, for the repulsion of the venous current dependent upon the respiratory movements, would give rise to a frequently interrupted flow of venous blood in the brain. A different explanation must be attempted for the manifold rapid changes of functional hyper- aemias, upon which depend cerebral excitation and the effective- ness of the mechanism of association. We shall see, however, that the brain does not completely fill the cranial cavity, but that the latter contains a number of spaces, filled with so- called lymphatic fluid. The skull is therfore not per se the cavity in which the brain rests, for on its basal side it contains a number of lymph-cisterns (Key, Retzius, Schwalbe). This explains why the forms of the convolutions should be imprinted upon the concave walls of the roof, and not upon the basilary (sphenoid) bone. The base of the brain would by its own weight rest upon this bone, and would leave impressions of its form but for the intervention of these water-cushions. Nor is the brain quiescent within the cranial cavity; it passes through three different phases of motion, as Burkhardt was able to demonstrate (in his 2l6 Psychiatry. monograph on the movements of the brain) on four patients with defective skulls. The movements of the brain in these four cases were registered upon a rotating cylinder, and gave tracings: (i) of the systole and diastole of the pulse, varying between sixty and eighty in the minute ; (2) of the expiratory rise and the inspiratory fall, numbering from fifteen to twenty in the minute (described before him by Ecker); (3) of the so-called vascular wave. This last is a peristaltic arterial movement, regulated by the vaso-motor centre, and occuring from two to six times per minute. These forces, affecting the intercranial pressure and the movements of the brain, are equally active in a normal closed cranial cavity, but do not produce any compression of the cerebral mass, for the lymphatic fluid, the more movable contents of the brain, recedes from or flows into cranial spaces as the compress- ing forces may demand. It will be necessary, first of all, to consider the anatomical and physiological conditions affecting the quantity of, and changes in, cerebral blood-supply, and after that to discuss certain independent cerebral influences, resulting from the activity of the brain itself, in their relation to the distribution of blood ; these influences unite with the former conditions in spite of variations of pressure due to peripheral causes,' to perfect the nutrition and function of the brain. We shall find that these cisterns, which are of variable volumes modify the cavity in which the brain is lodged, and that there is another mechanism by which an increase or decrease in the vol- ume of the brain itself can be compensated. This consists of the venous spaces, described by Cruveilhier, adjoining the sinus longi- tudinalis, which, like the sinuses themselves, do not possess a wall proper, but merely a simple limiting endothelium. These venous spaces represent, according to Langer, cavernous spaces situated between the separated trabeculae of the tissue of the dura. Vari- cosities of these venous spaces, by wearing away the vitrea, produce, as Langer and Trollard have shown, the foveae glandulares, in the aged, in drunkards, and in subjects of cardiac disease. These foveae glandulares, be it said, are not direct impressions of the Pacchio- nian dilatations of subarachnoidal spaces, to which we shall have occasion to refer later on. These cavernous spaces of Langer, Ludwig Meyer regards as a compensatory mechanism designed to secure at all times the repletion of the cranial cavity ; they dilate as soon as anaemia of the brain sets in, and collapse with the The Subdural Lymph-Space and its Connections. '217 return of a full current of blood. According to Schwalbe, the arterial system of the skull and brain divides into two distinct halves, according as the arteries are branches of the meningeal or cerebral arteries. The ramifications lying in the sulci arteriosi1 of the flat basilar bones are chiefly concerned with the nutrition of the bone and the dura mater, or at least with that part of the latter which acts as the periosteum of the cranial bones, and carries the blood-supply for the cranial walls. Besides investi- gating the cavernous tissues adjoining the sinuses, Langer has made a careful study of the dural arteries, and divides these into an outer and inner network. The outer network empties into the capillary net of the dura, and, which is quite remarkable, empties at once into very large veins, which grow narrower as they approach the finest arteries, while the inner network lying nearer to the surface of the cranial cavity, and forming a derivative net in contrast to the external nutritive one, does not divide up into small capillaries, but has its arteries empty directly into veins, thus warding off to a certain extent the effects of hyperaemia from the nutritive vessels of the dura. This observation of Lan- ger is of a kind with the views of Schroder van der Kolk, who, as will be explicity stated later on, maintained that there was a direct communication between the larger arteries and veins of the pia on the surface of the brain, having the effect, as he puts it, of allowing a strong current of arterial blood to pass over the cere- bral cortex directly into the veins, and permitting this vascular storm, as it were, to spend its force on the surface. Heubner's investigations have lent additional interest to this view. It will be shown that the increase and diminution of the volume of the brain in its cavity, as Burckhardt demonstrated in his four cases, bear significantly upon the nutrition of the brain, by transferring the waste products to the lymph-vessels, for which reason it will be necessary to consider the relations between the network of blood-vessels and the lymph-spaces of the brain. The exfernal (subdural) lymph-space has very different dimensions in the spinal and cranial cavities, for the spinal dura is widely separated from the arachnoid membrane, while in the case of the brain a capil- lary space is all that remains between the dura and arachnoid ; and this capillary space is lined by endothelium, communicates with the lymphatic glands of the neck, furthermore with subdural spaces which do not immediately surround the nerve-roots, but do 1 Erroneously so-called (Langer), for each artery is accompanied by two veins. 2l8 Psychiatry. so in common with the arachnoid and are connected with the i lymphatic spaces of peripheral nerves. The most important of these lymph-spaces, are those surrounding the auditory and optic nerves. As regards the former, it is found that the peri-lym- .phatic fluid of the labyrinth communicates with the subdural space. It is equally plain that the venous spaces of the sinuses and their surroundings communicate by a process of transuda- tion with the subdural spaces in the Pacchionian granulations. The subdural space is connected also with the lymph-spaces in the tissue of the dura. The arachnoid membrane, both surfaces of which are covered with an endothelium, shuts off the subdural space. This membrane envelops the brain, without forming any duplicatures upon it, but is intimately connected by means of a network of threads and trabeculae of connective tissue, and on its basilar aspect by means of perforated membranes, with the pia which covers every fold of the cerebral surface. Henle speaks of this network as physiological, hydropic connective tis- sue, which permits a complete interchange between all subarach- noidal spaces. These spaces are traversed at the summit of the convolutions by tense threads, and are narrower than over the sulci. On the basilar surface in particular, the subarachnoidal spaces are dilated, and in part devoid of trabeculae. This ex- plains the formation of cisterns (Key, Retzius Schwalbe). The following cisterns belong to the surface of the cortex : The space of the fossa Sylvii, which is merely spanned by the arachnoid, and a space which separates it from the dorsal surface of the corpus callosum, which space extends on the basilar sur- face as far as the tinea terminalis (of the central gray substance) situated beneath the corpus callosum. More caudad on the basi- lar surface we come upon the cysterna chiasmatis and the cystcrna intercruralis, the latter dividing again into a superficial and deep reservoir. From the cysterna intercruralis and to the outer side wide subarachnoidal spaces extend across the crus cerebri to the corpora quadrigemina, i. e., from the basilar surface to the dorsal surface of the trunk—the cysterna ambiens. Short tra- beculae unite the subarachoidal space just over the corpora quadrigemina to the surface of the latter. The most extensive subarachnoidal space on the dorsal side is the cysterna magna cerebello-medularis, extending from the dorsal surface of the oblon- gata to the cerebellum, on the superior surface of which exactly the same relations obtain as over the convolutions of the cerebrum. Subarachnoidal Spaces—Pacchionian Granulations. 219 The pia mater forms the inner wall of the subarachnoidal spaces, of which the tche chorioideae, in the superior cerebral ven- tricles, and in the fovea rhomboidea, constitute a part. This connection, and the penetration of the pia into the ventricles, are easily understood if we keep the fcetal brain (Fig. 1) in mind, and remember that the pia covers every part of the brain-surface. At the time when the hemispherical vesicle was situated to the front of the anterior cerebral vesicle, the pia mater passed from the former over the latter to the mesencephalic vesicle. But at a later period, when the hemispherical vesicles, enlarging caudad, had enveloped the anterior cerebral vesicle, the pia mater of the posterior surface of the vesicles of the hemispheres was, by mere flexion, made to lie upon the pia which covers the thalamic region (derived from the anterior cerebral vesicle). Just as the trabeculae of subarachnoidal spaces pass between the dupli- catures of the pia in the sulci between two convolutions, so the two laminre of the pia •which accompany the invagination of the velum chorioideum, are fastened to one another by subarachnoidal trabeculae. Behind the corpora quadrigemina, the arachnoid of the cysterna ambiens ascends to the upper wall of the cysterna corporis callosi. The flexion of the cerebellum over the oblongata produces, furthermore, a fold in the pia on its way from the cerebellum to the oblongata, the two laminae of this fold giving rise to the tela chorioidca of the IV. ventricle. The foramen Magendie leads through the pia from this ventricle into the sub- arachnoidal space of the spinal canal. In regard to the III. ven- tricle, it is to be remarked that its membranous tela does not correspond to the superior wall of the primary cerebral vesicle, but that the only vestiges of this which remain are the epithelial cells of the plexus choroideus, at the lateral margin and on the inferior surface of the velum. The larger branches of the arteries on the surface of the brain do not lie within the pia, but in the subarachnoidal spaces; the smaller branches only, enter the pia. Before describing these, we must stop a moment to note those prolongations of the sub- arachnoidal spaces, which are formed by the Pacchionian granula- tions. The latter occur in the course of all sinuses, particularly, however, alongside of the sinus longitudinalis. The sinuses and the veins adjacent to them are situated in the substance of the dura, and, according to Langer, are distributed in such a way that the veins of the anterior portions of the hemispheres meet with 2 2 o Psych iatry \ the veins of the posterior lobes in the wall of the sinus longitudi- nalis, as though the former (veins) stood in the relation of vasa vasorum to the latter. a The posterior cerebral veins take a similar longitudinal course forward, between the layers of the dura, so that the cerebral veins empty into the sinus for a distance of only 2 cm., and about below the middle of the parietal vertex. The Pacchionian formations push forward into the cerebral veins as diverticula of the subarachnoidal spaces. The veins lie intra- dural, and the subarachnoidal spaces are shut off from the subdural space. A definite brain-pressure, which we shall con- sider hereafter, forces the subarachnoidal serous fluid into the subdural space, whence, by a process of filtration, it empties into the veins and the sinuses. The subarachnoidal spaces communi- cate, moreover, with the lymph-channels of peripheral nerves, which encircle the roots, as does the dura also. From these sub- arachnoidal spaces we can throw injecting fluid into the lymph-space surrounding the optic nerve, into the perilymphatic space of the labyrinth, and the lymphatic vessels of the nasal mucous membranes. The pia cerebralis receives nerves from the plexus around the circle of Willis; these divide into small branches, which accompany arteries of only I mm. in diameter into the brain-substance. Bochaldek contends that branches from the III., V., IX., and XI. cerebral nerves join these vaso- motor nerves. According to Kolliker, the plexuses do not possess nerves. The sensitiveness of the dura in all experiments proves that it has sensory nerves ; they were demonstrated by Luschka and Riidinger. On the other hand, Nothnagel's experi- ments, confirmed by Krauspe, show the relation existing between the nerves of the pia and arterial contraction, which is generally reflex in character. The arteries of the fore-brain are derived from the carotid and vertebral arteries. The former supplies the art. cerebri ant. seu corporis callosi (the anterior cerebral), and the art. cerebri media seu fossa Sylvii (the middle cerebral). I. The anterior cerebral artery ramifies on the orbital surface over the convolutions which surround the sulcus rectus and the olfactory lobe; on the external surface, over a wedge-shaped region, which includes the second and third longitudinal convo- lutions (Fig. 9, L2, L3, cm., S.occ.) together with the uppermost portion of the central region as far as the occipital lobe ; while on the median surface, the corpus callosum and the entire region Arterial Blood-supply : Heubner s and Duref s Views. 221 from the frontal apex to the sulcus occipitalis receive their blood from the anterior, median, and posterior internal arteries, all of which are branches of the art. corp. callosi. II. The arteria fossae Sylvii embraces, with its ramifications, the operculum and the superior temporal convolution ; and extends to the convexity of the cortex, after sending off several secondary branches for the island of Reil. Charcot has designated the principal branch-arteries as follows: 1, the external frontal artery supplying the inferior frontal convolution; 2, the ascend- ing frontal artery for the region of the anterior central convolu- tion ; 3, the ascending parietal artery for the posterior central convolution and the superior parietal lobule ; 4, a parietal artery for the region of the parietal convolutions; and, lastly, temporal arteries ramifying over the first and second temporal con- volutions. III. The terminal branch (end-artery) of the vertebral artery —the arteria profunda—supplies the cuneus, the gyrus lingualis, the gyrus fusiformis, and the third temporal convolution, to- gether with the uncus and the gyrus uncinatus. Both Heubner and Duret deserve great credit for their methods of studying (by injections) the details pf cere- bral arterial distribution ; but there is a great difference of opinion between these two authors regarding the significance of the derivative arterial network of the cerebral surface, which unites with veins of an equal calibre,' as opposed to the nu- trient network of arteries, which connects with capillaries, and from which the end-arteries of the cortex and medullary sub- stance of the fore-brain are derived. Duret's views are corroborated by Charcot. Heubner does not consider the circle of Willis the final channel for effecting a collateral compensation of the cortical blood-current. After passing the subarachnoidal spaces into the pia, the cortical vessels issuing from the circle of Willis go on ramifying until the minute branches possess a diameter of but 1 mm. These branches then break up into a net of arteries, commu- nicating one with the other, extending over the entire pia, each part of which can be supplied by one of the chief arteries. The resistance among these derivative arteries is said to be so slight, that it would be easier to inject the entire pia of one half of the fore-brain through one of the six large arteries, than to make a 1 As was known to Schroder van der Kolk. 222 Psychiatry. successful capillary injection of a circumscribed region of the brain. From this common reservoir issue the more delicate arterial nets of the pia, and, at right angles, the cortical arteries. Even arteries of the white substance, larger than the latter end- arteries, which traverse the cortex without dividing, but are at a greater distance from the injection-tube, can be injected more easily than the vessels of the cortical network. According to Heubner, therefore, the existence of this anastomosing net forbids our saying that an artery supplies this or that region or convolution, except inasmuch as more distant arterial nets in the pia and cortex can- not be as easily injected from one vessel as from one another. But these are differences which it would be difficult accurately to define. In opposition to this view, Duret held that the super- ficial anastomosing vessels have a diameter greater than I mm., and that the nutrient cortical vessels are in reality capillaries, as was maintained by Robin also. In proof of this Charcot and Duret refer to the destruction of small cortical areas, due to embolism. Whence it follows that the arteries supply definite nutritive areas, and that influences of the derivative network of arteries is not as powerful as Heubner would have it. Heubner, however, quotes in support of his views, cases in which embolism of the pia-arteries is not followed by softening of any sort. Charcot grants that there are such cases, but he thinks them very rare. The long course relatively stout arterial branches take before dipping even into the wall of the longitudinal sinus, as Langer has observed, would argue in favor of Heubner's views. Though the latter's opinion regarding localized nutritive areas may be more unfavorable to the cortex than that of Duret and Charcot, yet it is chiefly a question of difference in the number of cases of cortical softening due to embolism, which the various authors have had occasion to observe. On another point, however, both authors are completely at one—namely, as regards the difference in the mode of arterial blood-supply for the cortex, and the subcortical masses which for the nonce include the prosencephalic ganglion. None of the cere- bral arteries which unite with others to form the circle of Willis gives rise to larger arteries carrying blood to the subcortical organs, and diminishing in size after undergoing dichotomous division ; but these subcortical masses are supplied with blood by a large number of arteries, which issue from the dorsal surface of the circle of Willis, differ but slightly in calibre from one another, and The Terminal Arteries of Subcortical Ganglia. 223 resemble, in their arrangement and number, the so-called lamina cribrosoe on the basilar surface of the brain. This sudden diminution in calibre of the art. corp. call., of the art. communic, of the art. foss. Sylv., of the art. prof., to a series of vessels but \-\\ mm. in diameter, departing from the main arteries at right angles, and penetrating at once into the base of the cerebrum, affects about 2 cm. of the length of the three large cerebral arteries outside of the circle of Willis and the whole length of the rami communicant cs. There are no anastomoses existing between the larger branches of this system of trunk-arteries ; and so these branches represent what Cohnheim termed end-arteries, none of which furnishes anastomosing branches before dividing up into capillary ramifi- cations. Inasmuch as there is no derivative network beyond the circle of Willis, and as these arteries, because of their shortness, are under the more immediate influence of cardiac action, they are more liable to hemorrhagic rupture than the cortical arteries —a fact of great importance in cerebral pathology. Following up the details of the distribution of arterial blood-vessels to the brain- trunk, we learn, according to Heubner, that the head of the corpus striatum (the nucleus caudatus), together with the anterior wall of the infundibulum and the anterior portion of the chiasma, is supplied ] by arteries coming directly from the art. corp. callosi.; that the other portions of the caudate nucleus, together with the entire nucleus lenticularis and the anterior portion of the internal capsule, receive arteries from the trunk of the art. foss. Sylv., which have passed through the foramina of the lamina perforata anterior ; while the ramus communicans post, is destined for the anterior tubercle of the optic thalamus, for the posterior portions of the caudate nucleus, the posterior wall of the infundibulum, the posterior portion of the chiasma, the medullary bodies, the ant. tubercle, and the gray substance of the third ventricle with its com- missure. The area supplied by the arter. chorioidea would include the environs of the cornu infer., whereas its choroid plexus would be distributed to the posterior division of the internal capsule and the outer half of the anterior portion of the thalamus. The art. profunda (2 cm. in length) supplies the posterior half of the thala- mus, the pes and tegmentum of the crus cerebri, the corpora quadrigemina, and the choroid plexus of the posterior horn and of the third ventricle. The pons and the medulla receive small branches from the adjacent basilar, vertebral and anterior spinal arteries. Heubner established these details of arterial distribution by showing that the (blue) injection-fluid thrown into the organ from definite portions of the basilar arteries would invariably extend to and through the same well-defined areas of the brain. The veins of the brain run parallel to the brain arteries on the surface of the brain, and empty into the sinus falciformis. The veins coming from the vermis su- perior, and the vena magna Galeni empty into the sinus rectus. Each one of the latter receives blood from an inferior vein of the corp. callos.; from veins of the cau- date nucleus, of the choroid plexus, of the optic thalamus, and of a basilar and pos- 1 Not regularly. 224 Psychiatry. terior cerebral vein. Rudinger found the sinus transversi and the jugular veins wide on the right and narrow on the left side in 71 cases ; in 27 out of 100 cases the reverse "was true, and in only 2 cases were they equally wide on either side. The broad sinus transversus receives its blood only from the cerebral surface through the longitudinal sinus ; through the sinus rectus the blood from the surfaces of the ventricles is carried into the narrow sinus transversus. No author has been as happy as Burckhardt in explaining the general relations between the membranes and blood-vessels of the brain, and their bearing upon the nutritive mechanism and the vital activity of' that organ. Burckhardt's conclusions, based upon observation of patients with defective skulls, were published in' his " Experimental Investigations of Brain Movements." The pulse-wave, the respiratory and vascular waves were familiar enough, as they are common to blood-vessels everywhere in the body. The respiratory waves, indicating a process of venous aspiration, point to one of the chief factors which determine the flow of venous blood to the heart. The movements of the arteries dependent upon the vaso-motor centre, and indicated by the so-called vascular wave of Mosso and Burckhardt, had been previously demonstrated on the ear by Schiff, on the web of the frog's foot, on the curarized tongue of the frog, and on the saphe- nous artery in the skin by Riegel, who gave a detailed account of these movements in a monograph on the influence of the nervous system upon the circulation of the blood. Riegel insists upon the intensity of these peristaltic move- ments of the arteries under the influence of the vaso-motor centre causing not only capillaries, but quite broad arteries, to contract to such an extent as only to permit the passage of a single column of blood-corpuscles, or indeed temporarily to oc- clude the arteries altogether. The dependence of this form of movement upon a vaso-motor centre is proved for peripheral arteries by the effect upon them of section of the cervical spinal cord. Plering and others think it very probable that this form of peristaltic movement is the result of the respiration of the vaso- motor centre itself, but that the stimuli causing such contractions have not accumulated sufficiently to exert an influence with every respiratory act; and that special influences from other sources, acting upon the vaso-motor centre through stimulation of sensory nerves, disturb the rhythmic peristaltic movements of the arteries. But in the case of the brain, the fact that it is surrounded by- rigid cranial walls, by the subarachnoidal spaces, and that it is placed under considerable pressure, modifies the effects of this Trier otic and Tricuspid Pulse-wave. 225 general vascular movement. Since the membranes surrounding the brain enclose spaces filled with lymph and cerebro-spinal fluid, these vascular movements constitute an important factor in the nutrition of the brain and in the exchange of metabolic products. The brain is subjected to considerable pressure. This is proved by the fact that from other organs tracings of the pulse- waves can be obtained only by placing the sphygmograph upon the artery itself; but, owing to the pressure it is under, the whole mass of the brain is moved with such precision by the pulse- waves, that accurate and distinct tracings of a tricrotic or tri- cuspid pulse-wave are readily obtained by placing the pulsometer upon the mass of the brain which happens to protrude from an opening in the skull. The truthfulness of these graphic tracings is brought home to us by noting that tracings from the carotid are exactly like those representing brain movements; though the resemblance between the latter and the tracings from more distant arteries (say the radial) is not quite as striking. The details of the pulse-wave can be exhibited more clearly with Marey's apparatus than with the rotating drum. As regards the radial pulse, Landois and Wolff considered the tricrotic "s^^^/VA^JvN/ the normal form ; in this the descending line of the wave-like tracing; due to systolic contraction, exhibits a top-wave and two others traced nearer to the period of diastolic diminution. Mosso, on the other hand, contended that the tri- cuspid VVVYVYx was the normal form, showing in its systolic (ascending) lines a point in front of the apex which corresponds to the one on the diastolic side of the pulse-wave. That the pulse- waves are not represented by well-rounded elevations and depres- sions, must be attributed to the sudden influence of arterial con- traction at the beginning of the diastole, causing an acute angle to be formed between the ascending and descending lines; and the two elevations following the apex of the wave are explained by the closure of the semilunar valves, and by the bounding back of the blood-column against the valves (Mendel). Both act antagonistically to the depression which is due to the tonus of the vaso-motor centre, and which causes the rapid fall of the pulse-wave. If we increase the contraction of a radial artery by immersing the elbow in cold water, we observe a distinct tricrotic pulse; if the other elbow be immersed in warm water, the arterial walls 226 Psychiatry. will relax, and the radial artery of this side will give a distinct tricuspid pulse. In this case, the arterial contraction is so weak that the first leap of the pulse-wave does not prove to be the highest of the three, but the closure of the valves and the re- bounding of the blood-current caiise the second elevation to rise higher than the first, and the succeeding lower, pointed wave on the diastolic side of the pulse-wave dilates the artery to such an extent that the line reaches the level from which it was inade- quately depressed through the influence of the vaso-motor centre. Burckhardt divides the cerebral vessels into subtentorial vessels in the posterior cranial fossa, and into basal and cortical vessels which are situated above the tentorium. The former resemble, as regards distribution, the blood-vessels of other vascular areas of the body, and supply cerebellum, pons, and medulla oblongata. The region provided by the basilar and cortical vessels includes the ventricles. In order to understand the influence of the systole upon the pulsatory movements of the brain, Burckhardt shows that the ar- teries ascend for a long distance from the circle of Willis, and that the venous and arterial currents take the same direction. The systolic enlargement of the brain, in consequence of the engorge- ment of the arterial network of vessels extending as far as the capillaries and veins, begins at the base of the brain and in- creases in the direction toward the vertex. This enlargement of the brain increases gradatim in the direction of the vascular rami- fications, so that at any one time all vessels at an equal distance from the circle of Willis are in the same phase of pulsation. The roof of the skull and the dural processes offer a direct resistance to the swelling convolutions with their shallow sub- arachnoidal spaces; for which reason the brain can only en- large concentrically toward the ventricles. In consequence of the fact that the basal arterial trunks and the long arterial branches do not simultaneously fill with blood, the tumefaction of the basal walls of the ventricles diminishes at a time when the swelling, brought about by the engorgement of the vascular net- work which is supplied by the higher arterial channels, is begin- ning, under the opposing pressure of the cranial roof, to constrict the ventricles, the force being exerted in the direction from the vertex. Since the basal and the dorsal portions of the ventricles are not simultaneously constricted, the pressure is partially neutral- Pulsatory and Respiratory Brain J lavements. 227 ized through the displacement of the cerebral fluid within the cerebral cavities. The concentric swelling of the brain is very nearly universal, since both the cortical and basal vascular ramifi- cations tend toward the surface of the ventricles. But since the intraventricular dislodgment of cerebral fluid does not altogether neutralize the arterial pressure, fluid from the ventricles passes outward through the foramen Magendie, while the concentric pressure continues to exert its influence upon the contents of the ventricle, and, favored by the thinness of the gray substance of its floor, spends its force on the subjacent cystern. chiasm, and intcrped., and, since the cisterns are confluent, upon all the cis- terns in general. Through the pressure of the liquor cerebri the pulsation is transmitted to the membrana obturatoria atlantis. Still another portion of the systolic pressure which is responsible for the increase in the volume of the brain is neutralized, through the circumstance that the engorged parenchymatous arteries effect the exudation of lymphatic fluid from those perivascular spaces which lie between the blood-vessels and the adventitia, the latter being prolongations of the pia mater which accompany these vessels on their course through the cortex and white substance. The respiratory act produces a fall of the pulse curve during inspiration, and a rise of this curve during expiration (depression and elevation of the pulse-wave). This curve depends upon the fluctuation of venous pressure. The expiratory curve-eleva- tion is due to the blood-stasis in the jugular vein, owing to the insufficient aspiration of blood from the thorax, and is increased by the arterial blood-current, since the pressure in the aorta increases with the expiratory pressure prevailing in the thorax. The influence of the expiratory wave is noticeable in the cere- bral curve, even during quiet respiration ; but it is most marked during violent expiratory acts, such as coughing and screaming. The retrogressive stasis acts directly upon the rigid walls of the sinuses, and dislodges at once the venous blood column. The sinus rectus is the shortest sinus. Relatively to the length of the sinus, the veins of the choroid plexus which empty into the vena magna, are the longest. The stasis, which starts from the torcular, and is felt first in the sinus longi- tudinalis, and the sinus rectus, with the exception of the veins of the subtentorial cerebral parts, will tell sooner upon the veins of the cortex than upon the long, flexible veins of the ventricular 228 Psychiatry. plexus. The stasis will also aid in producing the concentric swelling of the hemispheres, for its influence is felt last of all within the ventricle, and is similar to the effects of the pulsa- tory wave, with this difference, that it is felt from four to six times less frequently, and during quiet respiration is less in degree, than the influence of the pulsatory wave. This concentric tumefaction of the brain again causes compression of the liquor ventriculi against the basal cisterns, and its escape through the foramen of Magendie, whence it follows that the respiratory curve also will be noticeable at the membrana atlantis. The retrogressive stasis in the veins must necessarily interfere with the forward movement of the lymphatic fluid, and thus prevent its transfer by endosmosis into the blood-vessels. The brain, which has become enlarged by reason of this stasis, presses against the cranial walls, and exerts a similar influence upon the pressure in the direction of the ventricles, as in the case of the pulsatory wave ; but there is this distinction : The above pressure acts from the vertex downward, instead of from the base upward. The vascular wave is the most powerful, but the least fre- quent ; it gives from 2-6 tracings per minute. In height it may exceed the pulse-wave, ranging from several millimetres to 1.5 cm. The vascular wave causes a hemispherical protrusion of the cerebral mass, followed by a bowl-shaped retraction. Height and length of this wave are not equal. The wave flattens in a cool bath, and is raised in a warm bath. Its lowest point (wave-depres- sion) corresponds to the contraction, and its elevation (wave- summit) corresponds to the relaxation of the arteries. It is most distinct and regular during sleep; during the hours of waking its regularity is interfered with. Moderately warm baths of 77-91° F. lessen the number of waves, but make each wave longer ; warm baths increase the number and shorten the single waves. Es- march's bandage diminishes, galvanization increases, their number. According to Mosso the vascular wave is independent of the pulse and respiratory waves. But the vascular wave exerts a decided influence on the respiratory and pulse waves. During vascular contraction (wave-depression) the pulsatory waves are lower, for the artery is already contracted, but during relaxation they are higher, more rounded, tricuspid in shape. During the stage of vascular depression the respiratory wave, too, seems weak. All stimuli acting upon the sensorium create vascular move- The Vascular Wave—Arterial Systole of Base. 229 ments and disturb the periodic changes in the condition of the vessels. Elevation is brought about chiefly by psychical influences, and less by purely intellectual processes than by emotions. Burck- hardt had opportunity to observe in his patients the influence of pain due to panaritium ; the variations were marked, depression was persistent, the wave-length occupied one minute. Sudden fright, produced by an unexpected noise, caused a rapid rise in the curve, which was followed as quickly by a fall. While one of his patients was busy playing at chess, low but long extended waves with a few larger protuberances were noted. One of them was quietly reading (to himself) a humorous tale ; while thus engaged the curve showed many irregular variations. The pronounced vascular movement in the ear of the rabbit is not noticeable unless the animal be frightened. WJiile doing arith- metical work—a purely intellectual function—elevations were noted at the beginning and at the end, in between depressions were more frequent. The vascular wave, an arterial systole and diastole, advances in peristaltic fashion. Burckhardt appreciated the importance of this wave, constituting, as it does with the aid of the rigid cranial walls, a motor-mechanism designed to carry off waste products through the lymphatic fluids by establishing currents within the brain fluid. To prove his views he refers to Quinke's cinnabar injections into the spinal subarachnoidal space. The greater portion of the cinnabar penetrated as far as the glandulae Pacchi- onicae, and also into the dura ; a smaller portion passed into the sheaths of the cerebral nerves and into the cervical lymph-glands, but the injected substance did not reach the ventricles or the perivascular spaces. The arterial systole, as was the case with the pulse-wave, be- gins at the circle of Willis, constricts the base of the brain, pushes it away from the floor of the cranium, and impels the blood for- ward toward the cranial concavity, into the superior arterial branches, which dilate because they have not yet experienced sys- tolic contraction. The basilar constriction of the brain crowds the brain all the more against the cranial roof, and shuts off superiorly the advancing basilar liquor cerebri. The systolic contraction of the base compels a portion of the ventricular fluid to escape through the foramen of Magendie. The basal systole and the simultaneous collateral arterial diastole in the upper portions of the hemispheres, oppose the injection of cinnabar from the ven- 230 Psychiatry. tricles, for the roof of the ventricles is compressed by the con- centric swelling of the upper hemispheres, which are pressed tightly against the skull. The pressure under which the ventricu- lar fluid stands in this first phase of the vascular systole, which starts from the base of the brain, causes another portion of it (not that which escapes by the for. Magendie), through a current of absorption, to flow into the veins of the choroid plexus. Thereupon follows the second systolic phase—namely, the systole of the superior cerebral arteries which course along the concavity of the skull. The escaped ventricular fluid does not now return from the subarachnoidal space into the ventricle, for the basal collateral arterial diastole has set in simultaneously. In consequence of the swelling of the basal portion the diastole pushes the liquor past the upper cerebral parts (which have been removed to a distance from the skull by the arterial systole) into the Pacchionian bodies and the sinuses, and then into the basilar nerve-sheaths and into the cervical glands. The foramen of Ma, gendie lies within the region of the basal vascular diastole, where the swelling then present prevents the entrance of the liquor cere- bri spinalis, so that a current moving in the direction of the spinal canal keeps the injected cinnabar out of the ventricles. During the period of vascular systole in the hemisphere the ventricle itself is wide, for in the collapsed state the hemispheres do not crowd against the concavity of the skull, and do not present a concentric swelling toward the ventricle. The choroid arteries are in a state of diastole simultaneously with the basal arterial zone, and their dilatation produces secretion of ventricular fluid, the flow of which resists the return of the fluid that was expelled during the preceding stage of vascular systole. It was stated above that Quinke's cinnabar injections did not pass into the perivascular spaces between the pia and media of the arteries. Burckhardt gives the following explanation for this phe- nomenon : If an artery, lying in the midst of a perivascular space which communicates with the subarachnoidal spaces, contract, lymphatic fluid will pass from the parenchyma into the peri- vascular space (in a direction opposed to the course of the injec- tion from the cerebral surface into the perivascular space), for the passage to the subarachnoidal spaces on the convexity of the brain is now unobstructed ; but if this artery in the perivascular space be dilated, it obstructs this passage by filling out the above space, and no,cinnabar will be allowed to enter the subarachnoidal spaces. I'ascular U\zve in Sleeping and Waking. 231 During this stage the lymph-current is impelled toward the veins, as by the pulse-wave, which, with less success during cardiac systole, enables parenchymatous lymph fluid to be absorbed by the veins, and during cardiac diastole opens up the passage into the subarachnoidal spaces. The lymph-current is opposed only during the stage of venous stasis in the respiratory wave, which stage is accompanied by a condition of cerebral tumefaction. This stage implies a constric- tion of the perivascular lymph-spaces in the parenchyma, and the advance of venous blood pressing in the opposite direction. The influence of this retrogressive stasis, however, is not to be rated too high, for, according to Burckhardt, the influence of the vas- cular wave far exceeds that of the respiratory wave, and produces greater variations in the quantity of blood. Knowing that the vascular movements of the brain are respon- sible for the circulation of lymphatic fluid in the brain, we may infer from the regularity of these movements during sleep, that the refreshing influence of the latter is due not only to diminished consumption, but to a greater extent to the removal of waste products. On the other hand, the irregularities of vascular move- ments during waking indicate, as Burckhardt puts it, that the process described above can or must possess a certain independence in definite provinces of the brain, as is the case with regard to the (localized) reflex arterial constrictions on the surface of the body. Later on I will dwell upon the corollaries to be drawn from the mode of vascular nutrition, both as regards the emotions, and in particular as regards the independence of functional hyperaemias, as far as the general vascular movements of the fore-brain are concerned. The study of the modes of cerebral nutrition should be followed by an examination into the chemical nature of the sub- stances and products of nutrition. But on this head our knowl- edge is far more fragmentary than it is in regard to the mechanism of functional tracts and the channels of nutrition. The reactions showing the chemical composition of the brain can be carried on to a limited extent even in microscopical in- vestigation, thus enabling us to discriminate between chemical substances belonging to the elements of the gray and the white substance respectively. Other facts, however, can be obtained only by an examination of the entire masses of the brain ; and it is difficult, therefore, to make an absolute distinction between the gray and the white substance. 232 Psychiatry. In the human brain the white substance constitutes the main mass of the fore-brain, while the gray substance is the chief ingredient of the brain-trunk. In order to judge of the distribution of the chemical bodies among the gray and the white substance, the simplest procedure would be to determine the largest number of elements in each, and to ascribe the form which preponderates in each case to the gray or the white substance. But such studies have never been carried on with the aid of my method of dissecting out the brain-trunk and cerebellum from the hemispheres—a method peculiarly adapted to such investigations (cf. Figs. 1.6 and 17). Danilewski is the only one who attempted to estimate the elements of the gray and the white substance, basing the estimate upon a comparison of the differences in specific gravity. He found that the sp. gr. of the gray substance varied between 1.029 and 1.038, and that of the white substance between 1.039 anc^ x-°43- He investigated also the relative proportions of both substances in the brain of man, and found that it possessed 37.7 to 39 per cent. of gray substance, and 61 to 62.3 per cent, of white substance - while in the dog the gray and the white substances are present in the proportion of 50: 50. It was shown above that the preponderance of gray substance in animals as compared with the gray substance in man, depended by no means upon the greater number of nerve-elements, but upon the excess of amorphous connective tissue in animals. As regards the latter, we know that it is not, like other connective tissue, to be placed in the category of glutin-substances, but that this non-nervous substance is also albuminous in character. Boll, nevertheless, considers it allied to connective tissue, for, as he claims, all connective tissue contains remnants of albumin, derived from formative cells, and this non-nervous gray substance is simply distinguished from other connective tissue by the possession of a greater quantity of albumin. A very small quan- tity of amorphous connective tissue is, by the way, to be found, too, in the medullary substance of the fore-brain. And as for the nerve-cells, of which the axis-cylinders of the entire nervous system are a uniform part, Kuhne has already shown that both the nerve cells and the axis-cylinders give the reaction for albumin. This is based upon the reaction of the axis-cylinder with acetic and very dilute nitric acid, concentrated and diluted alkalies, in which the axis-cylinder swells and partly dissolves ■ and upon the contraction and yellowish discoloration of the axis-cylin- Nerve-Cells and Axis-Cylinders. 233 der in hot nitric acid ; and, furthermore, according to Rumpf, upon the characteristic reaction of the axis-cylinders of peripheral nerves, which are stained red by Millon's fluid. Among the various kinds of albumin, it was formerly supposed to consist of myosin, and then again to be identical with the contents in muscular fibres ; but that is disproved by its insolubility in a ten-per-cent. solution of sodium chloride. From the investi- gations of Kundt, and the digestion experiments of Kuhne, and Ewald, who used trypsin—the pancreas ferment,—we now know that the axis-cylinder is surrounded by a sheath, containing indi- gestible keratoid substances. These keratoid substances are contained in a lime-yielding substance of the axis-cylinder sheath, similar to Schwann's sheath, which surrounds the peripheral nerves. Accordingly, the medullary sheaths of peripheral nerves, as well as the sheaths of the axis-cylinders in nerves and in central medullary fibres, which lack Schwann's sheath, are horny forma- tions, and those parts only which remain after the whole has been subjected to the digestion process deserve to be called keratoid sheaths. Rumpf demonstrated the axis-cylinder sheath by ex- tracting the medullary substance of the nerves with alcohol and ether, and removing it with chloroform ; or else by adding distilled water, which expels the medullary substance, causes it to foam and to divide up into globules of myelin. The expulsion of nerve-marrow by water can be explained on mechanical principles by the synchronous swelling of the axis-cylinder, which dissolves slowly in water, and pushes its horny sheath against the external sheath of Schwann, thus crowding out the white substance be- tween the two. The chemical nature of the nuclei of the ganglion-cells was revealed through Jaksch, who proved the presence of Mischer's nuclein in the gray substance of the brain ; while Geoghegan showed that it was present in the proportion of 1.4 to every 1,000 parts of the entire cerebral mass. Although Jaksch did not thoroughly isolate the gray substance, he was able to prove the excess of nuclein in the gray substance relatively to the white. The presence of albumin and nuclein in the gray substance im- plies the presence of phosphorus in the ganglion cells and the axis- cylinders. Meyer and Cornwinder proved that in plants, the quantity of phosphorus increased in direct proportion to the quantity of nitrogen, and Bischoff calculated that the urine of starv- ing animals contained phosphoric acid in definite proportion to the 234 Psychiatry. quantity of nitrogen—say, I : 6.4 ; while if an animal be properly fed, the quantity of phosphorus and nitrogen taken in with the food is greater than that in the excretions. From these facts Voigt infers that the albuminates and phosphates unite, so that we must classify the fundamental connective tissue, as well as the nerve- cells in the gray substance of the brain, with those substances which contain phosphorus. Direct chemical proof has been proffered of the high percentage of phosphorus in nuclein; and the quantity of nuclein varies in all the organs of the body with the number of cells and nuclei which they contain. Kossel showed that the liver and spleen yield more nuclein than muscle, which contains a lesser number of nuclei; that leucaemic blood with its wealth of cell-nuclei contains more nuclein than the blood corpuscles, .which are devoid of nuclei. But, in the second in- stance, the quantity of nuclein depends upon the reproductive power of cells in which the division of the nuclei plays so im- portant a role ; so it happens that the percentage of phosphorus is higher in fcetal muscles and other foetal organs than in the fully developed muscles ; and the same obtains in plants 'with regard to the momentary foci of growth. The phosphorus of nuclein is attached to an albuminoid body which can be readily isolated by chemical means, because of its power of resisting digestive agents. The preceding remarks apply to the phosphorous elements of the gray substance, which constitute an important factor in the chemical composition of the entire brain. Relying upon ash- analyses and the investigations which Schlossberger, Bibra, Pollak, and Jarisch have made on the various tissues of the body, a fresh brain contains 0.49 per cent., phosphoric acid in its gray sub- stance, and 0.89 per cent, in its white substance, while the calf's muscles contain 0.48 per cent., woman's milk 0.38 per cent., human blood 0.10 per cent. The egg-yolk alone exceeds all, con- taining 1.15 per cent, phosphoric acid. In the brain of the dog, which, by the way, contains less white substance than the human brain, Forster found 0.83 per cent, phosphoric acid; in the muscles, on the other hand, only 0.48 per cent., in the blood 0.13 per cent., which is in thorough accord with the results of the different inves- tigators mentioned above. We are not warranted, however, in concluding that the nerv- ous system contains an absolutely larger quantity of phosphorus. The quantity of phosphorus in the nervous system cannot be gauged by the amount of phosphorus in the excretions, for, as Aqueous Extract—Lactic Acid- —Hypoxanthin. 235 Voit has determined, the entire nervous system of man contains but 12 gr. of phosphoric acid as compared with 130 gr. in the muscles, and 1,800 gr. in the bones; and, besides, we know, ever since Chossat's starvation experiments were published, that during starvation the nervous system shows no appreciable loss of weight. Petrowski has determined the percentage of water in the brain, and has found it to amount to 81.6 per cent, of the gray substance, and 68.35 Per cent, of the white substance. Quantitative examination alone is able to furnish positive evidence of the nature of cerebral elements, for, as Drechsel has shown in Hermann's " Handbook of Physiology," substances vary- ing in quality cannot be obtained in a pure state from one and the same brain ; and the terms lecithin, cholesterin, and cerebrin designate mixtures only: of which lecithin applies to the sub- stance composed of phosphorus which has been dissolved by ether and alcohol ; cholesterin is the term given to the ethereal extract which remains after removing the lecithin ; and cerebrin desig- nates the substances which form crystals in hot alcohol, but are insoluble in cold alcohol. The aqueous extract of the brain was examined by W. Mtiller, and was found to yield large quantities of inosite, 8 per mille ; lactic acid, 5 per mille ; kreatin, 4 per mille ; in lesser quantities, uric acid, xanthin, hypoxanthiu, urea, and leucin. It is worthy of notice that the acid reaction of the brain, which becomes more acid as the brain dies away, is attributed by Gscheidlein to a fermenting lactic acid, which can be obtained in the form of a lactate of lime (Hoppe). The reaction of the white substance, however, is not acid, but neutral, turning alkaline with the onset of death. Consequently, the brain does not, as Funke supposed, give a purely acid reaction. Kossel derives the hy- poxanthin from nuclein, which is obtained by weak reagents from the latter without setting up a general decomposition. He con- siders it a temporary product in the development of urea from the decomposition of more highly organized nitrogenous sub- stances. Hoppe-Seyler found that caustic potash acting upon nuclein at a temperature of 2000 Celsius, liberatedprussic acid and ammonia ; and Kossel remarks that the conditions for the develop- ment of the cyanides, which are intermediate products of the meta- bolic processes, are to be found in substances obtained from cell- nuclei. This chemical fact is interesting enough physiologically 236 Psych iatry. as proof of the correctness of Pfliiger's views, who held that certain phenomena due to irritation, such as epileptic convulsions, which are believed by Kussmaul and others to be caused by the withdrawal of arterial blood, could be explained by the reten- tion of metabolic products exerting an influence similar to that of the cyanides. Cholesterin is one of the chief products to be extracted by ether from the brain-substance. Of this body Hoppe-Seyler says that it is common to all living vegetable and animal cells, but that it plays no important part in the development of the cells; that it is in all probability merely suspended, and not dissolved, in proto- plasm ; that it is a product of decomposition resulting from the organic changes during cell-life. One of the chief constituents of the brain is protagon, which was first obtained by Liebreich. It contains phosphorus, and that too in far greater quantities than nuclein, which was described above. Protagon and nuclein are the main depositories of brain- phosphorus. For this reason, if for no other, we might incline to the view, that other chemical products of the brain- substance, such as lecithin, first prepared from the brain by Diakonow, and Mtiller's cerebrin, are ingredients of Liebreich's protagon. The method of obtaining cerebrin was refined upon by Parcus, who divided cerebrin still further into homocerebrin and encephalin. This view was expounded first by Kuhne, and later on Blankenhorn and Gamgee investigated the subject again, with the result of modifying the chemical formula of protagon. Very recently Drechsel expressed the conviction that this view was most probably the correct one, but that the atomic weights of lecithin and cerebrin did not suffice to make up a mixture or union of the nature of protagon, and that still a third sub- stance, containing more nitrogen and less carbon, would have to be shown to exist, if Diakonow's and Hoppe-Seyler's views1 are to be credited. It has been urged that the percentage of phosphorus in pro- tagon is exceedingly variable in quantity, and Diakonow con- tended that it contained no phosphorus at all, and in this respect resembled cerebrin. But Diakonow himself was unable to prove this, and the former objection is answered by the investi- gations of Blankendorf and Gamgee, who found that though protagon had been re-crystallized four or five times, the percent- 1 That protagon is merely a mixture. Protagon. 237 age of phosphorus remained the same; and this view is upheld by Drechsel. The formula of protagon, as determined by Blanken- horn and Gamgee isC160 H308 N5 P035. That it is present in the very earliest stages of brain development maybe inferred from its importance as one of the main constituents of the brain, and from the fact that it is easily precipitated by weak chemical reagents as a crystalline precipitate. After extracting the cholesterin with ether, it is separated out by alcohol (85 per cent.) at a tempera- ture of 450 C, after the dehydration of the residual brain-sub- stance, amounting to 31.65 per cent, of the entire brain-substance; by gradual cooling it forms into needle-shaped or star-like groups. To the arrangement of these crystals in nerve-marrow, in a direc- tion radial to its axis, Kuhne attributed the double refraction of this substance. Ebner, in his studies on the anisotropia of organic substances, takes issue with Kuhne, for he finds that treatment with cold ether destroys the double refraction, whereas protagon is entirely insoluble in ether. Lecithin and cerebrin are pre- pared by much more complicated chemical methods. To obtain pure crystals of cerebrin some thirty odd re-crystallizations must be performed, while lecithin (according to Diakonow) does not crystallize at all, or is precipitated with great difficulty only by ether at a temperature of O0 C. The presence of a body like protagon in the brain is rendered plausible by the physical properties of the latter, which, as Kuhne puts it, do well in participating in the composition of so peculiar a substance as the medullary sheath. Among these peculiar qualities is the power the medullary substance of the brain possesses, of reducing osmic acid, and turning a black color ; but these pecu- liar qualities are common also to the myelin forms of protagon resulting from prolonged contact of protagon with water. Like nerves themselves, protagon develops forms of myelin in the initial stages of decomposition. Before protagon is thoroughly dehy- drated by drying over sulphuric acid, it gains a waxy appear- ance ; it swells in water, changing to a transparent, starch-like mass ; when dissolved it invariably becomes opalescent. Here- after we shall see that pathological conditions of nerves give rise to similar appearances. Though lecithin and cerebrin (the latter a substance without phosphorus) exhibit the starch-like properties of solutions and ;nyelin-like forms, there is not sufficient ground to doubt the o;-i'/i:i of these substances from the protagon of the brain, but 20u Psychiatry. their marked hygroscopical properties stand in broad contrast to the lack of such qualities in protagon. This was first mentioned by Diakonow and Miiller, and has been insisted on since by Blank- enhorn and Gamgee. If protagon were a mere mixture of cerebrin and lecithin, it would be difficult to conceive how a non-hygroscop- ical body could result from the union of two hygroscopical bodies. It would be more natural to suppose that the hygro- scopical properties were the result of the more elaborate methods by which cerebrin and lecithin are recognized as secondary brain constituents, while protagon, a primary brain-substance, is obtained in advance of these. The chemical composition of cerebrin is indicated by the for- mula CG9.08 H11-47 N2.13 (Parcus), and of lecithin by the formula C44 FI90 NP09 (Diakonow). The formulae do not contradict the premise of Liebreich and others, that protagon divides primarily into cerebrin and lecithin. Hoppe-Seyler holds that lecithin, which is a phosphorous body, can be found in all cells capable of development, in rapidly proliferating pathological tumors, in sperm, and in the yolk of eggs, from which it has derived its name. Both lecithin and pro- tagon split up at a temperature of less than ioo° C. into glycerin, phosphoric acids, fatty acids, and the base, neurin, which lacks phosphorus, and is precipitated in the form of needle-shaped crys- tals. Protagon yields stearic acid, while the various forms of leci- thin divide up into palmitin and oleic acids. Considering the small amount of nuclein, the phosphoric acid of the brain is certainly not confined solely to the two phosphorous bodies, for Geoghe- gan found, after removing the substances classed under lecithin, that the brain-ash still contained 23 per cent, of phosphoric acid. Petrowski has made quantitative analyses of the gray and the white substance. Drechsel's critical observations, regarding the inaccurate preparations of lecithin and cerebrin must be taken into account. PETROWSKl's ANALYSIS : Water .... Solid residue contained therein Albumin and glutin . Lecithin .... Cholesterin and fats . Cerebrin .... Substances insoluble in dehydr; Salts..... Gray. White. 81.60 68-35 18.40 31-65 55-37 24-73 17.24 9.90 18.68 5I-91 °-53 9-55 tted ether, 6.71 3-34 1.45 °-57 Neurokeratin. 239 According to Bernhardt, the percentage of water varies very much in the different portions of the brain. The cortex contains 86 per cent. ; the medullary substance of the hemispheres, 70 per cent. ; the oblongata, 74 per cent. ; while the cervical spinal cord contains 73 per cent, of water, the lumbar cord y6 per cent., and the sympathetic 64 per cent. Ktihne's and Ewald's neurokeratin is soluble only in a hot • concentrated solution of caustic potash and sulphuric acid, and amounts to but 15 or 20 per cent, of the dried residue of the alco- holic or ethereal extract of the brain. Petrowski insists that lecithin and cholesterin found in the gray substance do not originate from the white substance mixed with it, but from the cells of the former. That the phosphorous substance in the medullary substance of the brain is not lecithin, but a body peculiar to the brain substance—say, protagon,—would seem to be proved by the fact that the biological impor- tance which Hoppe-Seyler attributes to lecithin would presup- pose an increase in the number of tissue elements. But for some time previous to birth there is no such increase of the elements of the normal brain-tissue. There is one occurrence, however, which could be held responsible for an increase of the tissue-elements, and that is the formation of fatty granular cells, preceding the development of medullary sheaths, and taking place about the time in which the original gray cerebral substance is transformed into white substance, and before the medullary substance has been developed. But the process is not re-enacted during normal life; whence Hoppe-Seyler, following Kuhne, infers that the white substance of the brain is intended for isolated conduction, and on the other hand points to the fact that rapid perception and movement are effected by medullated fibres, while the unstriated muscles (non-volitional) are supplied by non-medul- lated nerve-fibres. Nowhere in the system, though widely scattered, is lecithin called upon to effect such conduction as falls within the province of the cerebral medullary substance. There is better reason, there- fore, to presuppose the existence in the above tissue of a uniform but more complicated chemical substance. Pathological increase of nuclei, and possibly also of nerve-cells, certainly does occur; and as it is always preceded by division of the nuclei, we may look for a second phosphorous body well adapted to effect such changes of form, which could be naturally associated in our minds with an increase of such elements. 240 Psychiatry. As regards the relation of protagon to the rapidity of conduc- tion, we have two reasons for supposing the medullary substance of the nerve-fibres to be functionally connected rather with the nutrition of the fibres than with conduction. Rapidity of conduction might be held to depend upon favorable conditions of nutrition, furthered by the composition of the medullary sub- stance. Conduction is effected by non-medullated fibres also. It is initiated at the periphery, where the terminal branches of the nerves, and probably of the axis-cylinders also, are non-medullated, and at the other end conduction is originated in the gray centres in which the axis-cylinder passes into the non-medullated pro- cesses of the ganglion-cell. From the laws of Ritter we know that motor nerves, after sec- tion, begin to die at their central end, whence they have received their stimuli, and that cut sensory nerves begin to atrophy at their peripheral ends, for it is at the periphery that sensory stimu- lation is started. On the one hand, nutrition depends upon the non-medullated tissue of the axis-cylinder, which receives central and peripheral stimuli; and furthermore the stimulus itself supplies the incentive for nutrition. This would signify, according to Vir- chow, that the axis-cylinder exercised a powerful attraction upon the nutritive plasma, which is rendered most effective through the mediation of stimulation. We must look upon the medullated sub- stance as a tissue favoring rapidity of conduction, and aiding the nutrition of the axis-cylinder. If the venous character of the blood which leaves the brain necessitates our considering a process of oxydation in connection with nutrition, then we may assume this medullated substance to be concerned in this, for this substance is able to reduce osmic acid by withdrawing oxygen, a chemical process in which its large percentage of phosphorus plays an im- portant role. Kuhne calls attention, from the chemical stand- point, to the slowness of nerve-conduction, first proved by Helmholtz. This slowness does not argue in favor of the conduc- tion of a physical force such as electricity, but suggests rather the progress of chemical changes from section to section of a nerve, while it is acting as a conductor. The axis-cylinder alone, because of its continuity, can be engaged in nerve-conduction. The medullary sheath is not continuous; it is interrupted (only) in the region of Ranvier's constrictions; here the medullary sub- stance appears interrupted, but not so the primitive sheath. Rumpf has shown, that if the medullary substance is forced out Keratoid Sheaths Regulate Nutrition. 241 by allowing the axis-cylinder to swell in water, that the white substance is checked momentarily in its course at the constriction- rings, but the current pushes on within the primitive sheaths through and beyond the constriction rings. Secondly, the septa of Lanterman divide the white substance into a number of segments, which seem as though they had slipped into one another like the two divisions of a box. The digestion experiments of Kuhne and Ewald, and the investigations of Rumpf, have shown that the septa contain horny substance, which intervenes between the sheath of Schwann and the keratoid sheath of the axis-cylin- der, and furthermore that a framework of horny substance traverses the white substance between the segments. Those complicated structures, which Stilling was the first to describe as a system of tubes and trabeculae are probably synonymous with this keratoid network. If the expression keratoid substance could be applied properly to the sheaths of the white substance and of the axis-cylinder, then the axis-cylinder could be considered isolated from the white sub- stance by a body far less permeable than either the white substance or the axis-cylinder. The correct expression, horn-bearing sheaths, points to the fact that the keratoid substance is interrupted by substances favorable to a nutritive endosmosis. Since the axis- cylinder is better adapted during stimulation to attract chemical substances from the white substance, and in keeping with the greater rapidity of conduction in medullated fibres to increase the chemical processes involved in conduction, its sheath, consisting of horny and glutinous substances, may be held to represent a sieve which allows the nutritive plasma, as much at least as is attracted from the white substance, to fall upon the axis-cylinder, not with the intensity of a full current, but with the more delicate force of a rain ; and we must regard the partial endosmotic per- meability of the neurokeratin sheath as an apparatus regulating the physiological needs of the axis-cylinder. But the white sub- stance of peripheral nerves also is surrounded by a similar horny sheath, and interrupted in its interior. The aforesaid regulation of the nutritive plasma applies to the sheath of Schwann as well. In this way the nutrition of the white substance is made indepen- dent of the plasma exuded by the blood-vessels, and a uniformity in regard to its chemical changes is established. This is sug- gested also by the peculiar conditions of nutrition, since the broad meshes of the vascular network of this comparatively 242 Psychiatry. anaemic white substance are to be found only on the surface of complicated bundles, whereas the single nerve-fibres have no sort of contact with the blood-vessels from which the plasma that is exuded permeates the sheath of Schwann, and mingles with the white substance drop by drop, instead of pouring in upon the latter with its entire mass at once. The numerous interruptions through horny substance in the white substance would impede any simple mode of nerve-conduction, but would favor an endosmotic pro- cess, advancing slowly from segment to segment. The electrical nerve-current is probably to be regarded merely as a secondary phenomenon accompanying chemical changes. This secondary phenomenon has been specially studied, how- ever, by nerve-physicists. Dubois' (Reymond) nerve-current of rest, as is well known, is not connected with any process of stimu- lation. It suffers disturbance through negative variation. The chemical nature of nutritive processes could be studied by careful examinations of daily secretions; and many chemists have sought to determine the quantity of phosphorus contained in the excretions of the body, starting out on the supposition that the nutrition of the nervous system is to be held to account for a consid- erable portion of the phosphorus in the urine and faeces. On this head, Mendel's observations on the percentage of phosphorus in urine are specially worthy of notice, as well as some other obser- vations to which Mendel refers in this same treatise. Mendel inquired into the variations in the quantity of phosphorus present in the urine during waking and sleeping. He found the nocturnal urine richer in phosphoric acid than the urine passed during the day; whence we infer that the waste products of the brain are increased during sleep. This fact is in perfect accord with Burckhardt's observations regarding the movements of the brain. Burckhardt established a direct mechanical relation between the vascular wave and the removal of the lymph-current; and main- tained, moreover, that while the brain was awake and active— e. g., while the brain was in a state of excitation—the effects of the vascular wave were interrupted and irregular ; but that regular peristaltic vascular movements were peculiar to sleep, and that the complete chemical restitution of the brain during sleep must be ascribed to the increased deportation of its waste pro- ducts. Mendel refers in his article to the observation of Wood, that mental activity slightly increased the alkaline phosphates of the urine, but that it caused a greater and decided diminution Nerve-Current and Negative Variation. 243 of the earthy phosphates. He concludes, therefore, that nerve- tissue increases, like muscular tissues, during mental work, thereby entailing a diminution in quantity of the phosphates excreted. The chemical phase of excitation seems to him to involve a synthetical chemical process and a decrease of waste products. This view becomes attractive, if we remember that the phenomenon of memory is a positive action which has out- lived the process of excitation, and it is much more conceivable that a synthetical chemical process should leave lasting functional consequences, than that functional acts should arise from the decomposition of waste products soon to be discharged. The nerve-current of rest, together with the increased products of chemical decomposition and the mechanical methods for the removal of waste products, exists also during sleep. If the func- tional excitation of the axis-cylinder, whether central or periph- eral, results in the withdrawal of a phosphorous substance from the white substance through the horny sheath, and in utilizing this phosphorous substance in a synthetical chemical process, then the chemical process going on in the white substance and connected with the nerve-current of rest will be disturbed ; and, further- more, the electro-negative variation would correspond to a synthet- ical chemical process in the axis-cylinder and the nerve-cells as the very opposite of a metabolic process of division (Spaltungsvor- gang). Rumpf's investigations have shown that the nutrition of the axis-cylinder depends, apart from the possibility of isolated and rapid conduction through the medullary substance, upon stimuli, and consequently upon its connections with a sensory apparatus at the periphery, and with a central organ. Rumpf's observations were made on nerves which remained within the living body, but which were severed both at their peripheral and central ends. Under these circumstances the axial band disappeared and was dissolved, within twenty-four to seventy-five hours, even if the nerve had been kept in a proper menstruum. The influence of the medullated substance and the blood-vessels does not suffice for the nutrition of the axial fibre. This experiment proved at the same time that the fibrillary structure of the axis-cylinder, which Max Schultze inferred from the striated appearance of the axis-cylinder and of the cell-pro- cesses, after treatment with silver, did not in reality exist, but that this striation was due simply to a precipitate of silver in the 244 Psychiatry. folds of the keratoid sheath. Rumpf was able to demonstrate this same silver-striation in sheaths which we but just learned did not enclose axis-cylinders. The keratoid sheath of the axis-cylinder and white substance might possibly, by reason of the pressure it exerts, be of some mo- ment as regards the more delicate conditions of normal nutrition. This applies to the horny sheath of the axis-cylinders of the brain, but all brain fibres as well as all fibres of the optic nerve and of the spinal cord lack the sheath of Schwann. But in lieu of the pressure which the sheath of Schwann presumably exerts on the white sub- stance, another force comes into play in the cranial and spinal cavities, which, as far as the medulla is concerned, is indicated by the tension of the membr. atlantis. From the chemical standpoint, the greater wealth of water of the brain and spinal cord must be borne in mind ; this might be thought to correspond to the greater quantities of plasma exuded in consequence of the ab- sence of the sheath of Schwann. This indirectness and independence in the matter of the nutri- tion of the axis-cylinder would seem to be a safeguard to prevent the disturbance of its function by cerebral hyperaemias. Under normal conditions of stimulation and attraction of the axis- cylinder, as well as of the ganglion-cells, and with an abundant ramification of wide arteries, the brain appears by its normal structure to be protected against the possibilities of anaemia ; for in a body wasting from starvation the central nervous system, it is claimed, suffers less in weight than other organs of the body, and it is well known, furthermore, that cerebral activity is possible in spite of general anaemia of the body. In consequence of its larger percentage of water, the gray substance would influence nutrition in a very different way from the axis-cylinder, which, throughout its course in the central white substance, and in the peripheral nerves, has the function of a conducting organ. The variable nutritive conditions are depen- dent upon the gray substance to the extent that they are in- fluenced by the blood-supply. Since albuminoid substances aggregate to more than one half of the chemical constituents of the gray substance, we may infer a direct relation to exist be- tween the albuminoid bodies and the percentage of phosphorus, and the latter would be greater than in the case of other cellular tissues, for even the connective tissue of the gray substance contains more albumin than the connective tissue elsewhere does. Nutrition of Axis-Cylinder. 245 This then, is another factor accounting for the greater accumula- tion of brain-phosphorus. Because of its greater wealth of cells, the gray substance con- tains a larger quantity of nuclein ; but nuclein stands next to protagon in the series of cerebral bodies containing phosphorus. LTnder normal conditions, and in the fully developed brain, nu- clein does not seem to play an important role, as regards the increase of tissue, as it does in pus, in yeast, and wherever else it is found. After excluding this particular function, and in addi- tion to its adding merely to the percentage of brain-phosphorus, we may insist on the fact that the nucleus exercises a nutritive in- fluence over the albuminoid protoplasm. This nutritive function for the maintenance of the protoplasm is annulled in pathologi- cal processes of the brain as soon as the division of the ganglionic nuclei enables the nuclein to develop its faculties for regenerating tissue, while the protoplasm of the cell-body is diminished and finally destroyed. The nutrition of the axis-cylinder, which is a part of nerve- cells and of their prolongations, some of which become axis- cylinders, is effected in a similar manner. Jastrowitz considers the gray substance (pp. 58, 59) the medium of isolated conduc- tion, to the extent that such conduction is imposed upon the gray substance. If the striation of the axis-cylinder, and the striation which Remak observed on the ganglionic processes, and Schultze (p. 62) within the cell-protoplasm, indicate an identical process, (while, according to Rumpf, it is simply an expression of the folds in the keratoid sheaths,) then we should have to suppose such striation to exist also in the case of the nerve-cells and their protoplasm. It is quite conceivable also that a sheath surrounding ramified nerve-cells, which has as many points of attachment in the connective tissue as it has branches, should, if isolated and separated from these points of attachment, lose its normal tension, and be thrown into folds occupying all possible directions. The lines of Schultze, which Boll demonstrated, with the aid of osmic acid, in cortical cells also, both in those forms which extend from one process to the other and in the dark lines running concentrically around the nucleus, might be ascribed to simple folds without a minute fibrillary structure. The larger quantity of plasma in the cortex depends upon its denser network of blood-vessels. The nutrition of the nerve-cells is regulated specially by the sieve-like perforations of the keratoid 246 Psychiatry. sheaths, while the short distance which the naked nerve-fibre has to travel from its origin in the gray substance before it is sur- rounded by a medullary sheath, does not influence the rapidity of conduction. And on the other hand, the network of gray fibres does not become medullated either. But the very obstacles to irradiation, causing slowness of conduction in the gray sub- stance (p. 185), might be explained by the dropping off of the medullated substance from the anastomosing processes of nerve- cells, since the medullated substance is justly considered to favor rapidity of conduction. In order to comprehend the mode of nutrition during cerebral activity, we must, if we wish to avoid confusion, omit the consid- eration of irradiation in the gray substance, although there is no reason to suppose that any active cerebral process excites but a few nerve-cells only. For this reason we must presuppose an irradiation of moderate extent to take place whenever a number of cells unite for common action. The development of a single thought is effected by the functional activity of association- bundles, which unite in a very complicated way the com- ponent elements of a so-called residual image of the cortex. These groups of associated cells which harbor residual images, are the starting-point for the excitation of more comprehensive associations, constituting simple processes of induction (Schlussprocesse) (p. 153). It has been explained above that the entrance of numerous and comparatively indepen- dent arteries of the pia into the broad expanse of cortical surface favors the excitation of separate and well-defined groups of ganglion cells, and we must remember that every cortical image and every inference depend upon the union of special groups of cells. The projection-system alone stands under the influence of centres pf excitation, for as soon as the fore-brain comes into play the activity of the association-fibres precludes the possibility of localizing the cerebral excitation. We must admit, therefore, that the localization is but an aid to the grouping of stimuli differing from one another in their relations to time and space ; they are united for a common action by the process of association—a process which cannot be confined to any one portion of the cortex, as is amply demonstrated by the varying lengths of the association-bundles which unite the remotest parts of the cerebral lobes. It is worthy of remark, that every process of thought ema- Nutritive Attraction. 247 nates from a residual image of the cortex, and that this image is connected with a large number of distinct cerebral foci. The individual mental act is so constituted, however, that the initiatory residual image enters into a lesser number of connec- tions than the actual anatomical connections would warrant. In every mental act but a portion of the possible available associa- tion-tracts are employed. If the functional activity of the cortex implies a synthetical chemical process, then the active brain- cells and brain-tracts will require an increased quantity of plasma. We can understand the possibility of isolated conduction by attributing to the nerve elements a nutritive attraction along the course of the arciform bundles. If we accept Fechner's theory, that the cortical images and their connections may be stimulated to one of two variable degrees of intensity, and that in any particular mental act those images which are actively utilized stand above the threshold of consciousness while others remain below the level of consciousness, then accepting this theory, we may interpret it to mean that elements bearing processes standing above this level exhibit a greater nutritive attraction than those elements which are not then called into play. This is a distinc- tion of degree, not of kind, for certain stimuli can be perceived without being clearly recognized by consciousness, and, further- more, synchronous or successive stimuli can become associated. These associations may be formed unconsciously, i. c., below the threshold of consciousness, and yet they may rise above this level, and into consciousness as complete inductions (inferences), if any groups of elements in these regions will by association stim- ulate nutritive attraction to such a degree of intensity, that these groups can pass from the condition of partial sleep to that of partial wakefulness. From a consideration of the cerebral mechanism we learn that in addition to the influence of attraction, there exists a nutritive process which, apart from the influence of the heart, relies upon the effectiveness of the vaso-motor centre; whence it follows that functional hyperaemia cannot be put solely to the account of attrac- tion. Analyzing the independent manner in which the vaso-motor centre influences the brain, as is exhibited by the vascular wave, we find that we get the vaso-constrictor influence with the arterial systole, and the vaso-dilator influence with the arterial diastole. But the mental processes are not interrupted by each arterial systole, for which reason they must, to a certain degree, be inde- 248 Psychiatry. pendent of functional hyperaemia. This independence of mental acts may possibly be due to the fact that the cortex itself acts as a vaso-motor centre in its relations to subcortical centres. This cortical function is the very opposite of functional hyperaemia. If functional hyperaemia is caused by stimulation of the cortex, starting the mechanism of association, then the constriction of the blood-vessels is the result of cortical stimulation, and this dila- tion (p. 189) must be considered the result of inhibition of cortical function. We must look to another fact if we wish to understand the nature of the functional hyperaemia which helps and, for a certain length of time, enables the plasma to exhibit its phenom- ena of attraction. Fechner denies the spontaneity of motor acts, which, if they were spontaneous, would not obey the universal law of the conser- vation of energy. According to this law, energy cannot be generated, except at a loss of other energy. He proves the applicability of this law to cerebral activity by referring to the common experience of all that muscular and mental energy can- not be generated simultaneously by the brain. A person who is engaged in physical labor will, as soon as a mental process is set up, allow an arm that was raised to fall; and, conversely, severe physical exercise disturbs any process of thought. The inference is, that functions of the fore-brain inhibit one another, according as one or the other happens to predominate in the fore-brain. Goltz demonstrated the inhibition of the croak-reflex in the frog through other stimuli acting simultaneously upon the gray substance. But from the investigations of Burkhardt we learned that the influence of the vaso-motor centre upon the peristaltic vascular movements is modified by the activity of the hemispheres, and that when this activity is lowest, as it is in sleep, arterial systole and diastole succeed one another with the utmost regularity. It is evident from the influence of cerebral activity over the vaso- motor centre, that the vaso-motor nerves of the cortex do not reach the blood-vessels at once, but that they are interrupted in the subcortical vaso-motor centre ; and that for the vascular innervation of the cortex, the subcortical centres must constantly be called into requisition. I remarked above (p. 207) that the vaso-motor centres governing cortical influence must be referred to the oray sub_ stance of the anterior division of the brain-trunk, in which are Influence of Cortex upon Vaso-Motor Centre. 249 situated also the other motor tracts subject to centrifugally trans- mitted cortical innervation. If the cortex be excited in its capacity as a vaso-motor centre, the influence of the arterial systole upon the vaso-motor centre will be augmented, thus causing active anaemia of the brain, which, as a rule, remains entirely independent of the anaemia of the rest of the body. But since a functionally active cortex cannot impede the devel- opment of functional hyperaemia, we must assume that the physiological excitation of the cortex increases in a centrifugal direction the arterial diastole which forms part of a peristaltic move- ment. This would be quite comprehensible, not only if an active cortical process, such as a thought, would, as Fechner contends, interfere with the evolution of intense conscious movements, or vice versa, but also if the process of association would inhibit the second motor function of the cortex—viz., the constriction of the arteries. I made mention of this division of cortical energy in favor of functional hyperaemia, in speaking of the relation of this hyperaemia to an aggressive emotion (p. 195). On the other hand, again, deficient or diminished cortical activity, as expressed in conscious movements, or in thinking, would be attended by an augmented excitation of the vaso-motor nerves connected with this very portion of the cortex, and would deprive the attraction of the nervous elements through exosmosis of adequate material, just as reflex impulses from the periphery cause a constriction of tne cerebral arteries through the mediation of a subcortical vaso-motor centre, and thus influence the blood-supply as well as the chemical changes in the brain. Figures 64 and 65 are intended to illustrate in a diagrammatic way the manner in which excitation of the cortex and excitation of the vaso-motor centre inhibit one another. Each one of the three different cortical regions performs two different kinds of work : Through the U-shaped bundles JA., JA., processes of asso- ciation receive their innervation,—thought is rendered possible; while the projection bundles JV. and JN. transmit cortical stimuli to the vaso-motor centre CMU. and CVM.1 Figure 65 exhibits the association-bundles with the arith- metical sign > (larger) appended, and the vaso-motor bundles with the arithmetical sign < (smaller) ; in addition to this we 1 The wood-engraver is responsible for this difference on the two diagrams in the lettering of parts that are identical. 250 Psychiatry. note that the blood-vessels of the brain-substance are indicated, and are larger on Fig. 65 than on Fig. 64. This difference of in- nervation exists at all times, and the diagram is intended to show that increased functional activity of the cortex is followed by a diminution in cortical vaso-motor influences ; whence it follows Fig. 64. Diagram of Vascular Innervation during Increased Vascular Pressure. C, C, C. Three regions of the cerebral cortex, connected with association-bundles A, and with vascular nerves V. A. Association bundles. > J A. Inhibited associa- tion. < JV. and < JN. Increased vascular innervation, consequently the blood- vessels are drawn narrow, nucl. lent. Lenticular nucleus. Th. Thalamus. CMU. Vaso-motor centre. Diagram of Vascular Innervation during Functional Hyperaemia. C, C, C. Three regions of the brain which are cpnnected with association-tracts A, and vaso-motor nerves. < JA. Increased associations. JV. < Vascular inner- vation checked, consequently diagrammatic blood-vessels are dilated. Lent. N. Nu- cleus lenticularis. Th. Thalamus. CVM. Vaso-motor centre. Functional Hyperemia—Blushing. 251 that a cortical process of association by inhibiting vascular inner- vation will result in immediate functional hyperaemia. In the diagram, Fig. 64, the association bundles have received the mi- nority sign <, and the tracts for vascular innervation the plurality sign >. The three diagrammatic arteries are here represented narrower than in Fig. 65. The diagram signifies, furthermore, that if a slight functional activity of the association-tracts does not produce functional hyperaemia, the influence of the vaso-motor centre will not be checked. At the same time it is shown in both diagrams by means of arrows that the cortex imparts a centrifugal impulse to the vaso-motor centre, and that in some way or other this impulse is transmitted in a centripetal direction from the sub- cortical centre reacting upon the vascular system. A graphical statement of these facts would read thus : Process of association > = vascular innervation < and Process of association < = vascular innervation > || } Innervation of the arteries is invariably the result of the functional activity of a subcortical centre ; from which we infer that the whole condition of functional hyperaemia is based upon the cortical inhibition (p. 197) of a subcortical centre. The existence of general cerebral fluxion during and after mental work forbids the inference that functional hyperaemia depends altogether upon the attraction of stimulated nerve-elements. That this fluxion extends beyond the brain would seem to be proved by the physiognomical phenomenon of blushing. Blushing is often the concomitant condition of a multitude of associations. The wealth of simultaneously excited nerve-ele- ments interferes with the orderly development of thought; for in this instance, too, the simpler the character of mental work the more unrestrained will be the flow of association. A girl who recites a poem from memory will, by dint of cere- bral association, be able to repeat words and verses in the proper order of succession. The child will not blush if she happen to recite the poem before children only ; but the presence of the kin^ would make it blush and stammer forth its lines in mere confusion. The sight of the monarch is connected with a larger number of associations which have reference to his unusual personality; his presence will start a number of indistinct associa- 1 The author uses these signs rather oddly. The signs > and < denote "in- creasing " and " decreasing."—S. 252 Psych iatry. tions. The sum of the cortical elements thus excited will be followed by paralysis of the vaso-motor centre, and will cause much more widespread functional hyperaemia of the brain than the task of reciting the poem did. This degree of general hyper- aemia will, in consequence of the numerous and simultaneous inhibitory influences acting upon the vaso-motor centre from all sides, cause a paralysis extending beyond the domain of cerebral vessels, and will thus produce blushing ; secondly, we may argue that the many different ideas started in the child's brain will inhibit one another and confuse the acts of the child. Every one experiences a sensation of confusion at the be- ginning of a difficult mental task, for from the residual cortical images which are at the bottom of any special train of ideas, more association-tracts will be excited into action than can be utilized in the orderly evolution of these ideas. Excessive functional hyperaemia will be engendered, upon which will follow a state of mental confusion. Walking up and down, bodily exercise of any sort, the sight of green fields, will suffice to dis- tribute a general hyperaemia to other parts of the cortex, by means of which the initiatory conception, with all the associations connected therewith, will be more sharply defined, and simplicity as well as precision of thought will be secured. It is only when the disturbing influence of superfluous hyperaemia has been re- moved, that the process of attraction will insure the development of thought to a purpose. By injecting the vessels of the brain it can readily be demon- strated that the blood-vessels run a radial course parallel to the projection-systems, and an arch-shaped course along the associa- tion-bundles. The course the various systems of fibres take could be inferred from the distribution of the arteries as seen on corrosion-preparations. We can thus understand why functional hyperaemia should favor well-ordered attraction along the lines of association-tracts ; but we perceive, too, that a large number of associations connected with the image just then above the threshold of consciousness will be equally favored ; in which case many re- sidual images will be called into activity—i. c, will pass into a state of superior nutrition, but to no purpose whatever. Such secondary images can, however, be consciously utilized. There are easily aroused, though generally neglected, associations ex- isting between each word and its assonances. If thoughts are to be expressed in rhymes, these assonances rise into consciousness Assonances—Evolution of Thoughts. 253 and are interwoven with the ordinary train of ideas to the extent that other associations will allow. Every oratorical effort is likely to be marked by secondary associations which were sug- gested by the thought in words alone. If we speak of certain ideas as bright, we are easily led to think of things that are bright, and we may compare such ideas to the light of the sun, and so on. The physiological isolation of certain association-tracts, to the exclusion of the functional activity of secondary associations, may be explained somewhat as follows : A train of thought starts from a residual image in the cortex. All the associations con- nected with this image are, as it were, ready for action. But no sooner has the cortical image been revived than a second image is presented before our consciousness, which also exercises a definite attraction. The result is that the associations connecting these two images are under double attraction, and will conse- quently be more intensely excited than any others. The orderly evolution of any one thought implies a starting-point and a goal between which it runs its course. The two images are at either end of this course; and through a strict observance of this course a firm union is established between them. Just as a marksman, in spite of numberless objects around him, establishes a direct re- lation between his finger on the trigger and the bull's-eye which he is to hit, so a similar relation is established between the two terminal images, which controls the direction of the play of association. And at the same time other images from a parallel direction are perceived in the horopter, which, after the shot has been fired, may alter the circle of vision. The so-called unity of consciousness may be likened to the activity of the macula lutea, and the secondary association to that of the horopter. Below the threshold of consciousness parallel presentations may arise, due to attraction exercised along associa- tion-tracts not connecting the starting-point with the goal of the thought then before the mind. Such parallel presentations may, but do not necessarily, arise from previously well-established associations. They may have terminal images of their own before risinc to the level of consciousness. The forces of attraction which push them above this level find these images and associa- tions well organized and ready for use. I shall have more to say of these parallel images and associa- tions in a later chapter of this book in which I propose to treat of the mechanism of Expression. 254 Psych ia try. Both diagrams representing the inhibition of the vaso-motcr centre through cortical function, and the inhibition of cortical. function through the vaso-motor centre, apply, as was stated above (p. 195), to different stages of emotion. Burckhardt tells us, moreover, that the influence of emotion upon the vascular wave is far more intense than the influence of thought. This greater intensity is easily explained if we remember that an emotion involves our whole individuality, and the functional excitation of the widest possible range of associations. Whether the arterial systole will be checked or increased will depend in the one case upon the proportionate number of functionally liberated nerve-elements, and in the other upon the number of nerve-ele- ments already functionally engaged. This is true, above all things, of bodily pain. A finger dipped in water having a temperature of 500 Celsius will give rise to a sensation of warmth ; if the arm or the entire body be immersed into water of the same tem- perature, the sensation of pain will be engendered, for in the latter instance a larger number of nerve-tracts will have been excited. So, too, moderate illumination of the retina effects simple vision ; but the disc of the sun striking the eye produces actual pain, through the larger number of elements excited by irradiation from every optic-nerve fibre. The retinal images of a stranger, and of a person whom we love or fear, are entirely the same, yet the sight of the latter two may produce an emotion in us. For, in this instance, innumerable associations will be aroused which are connected with the images of these persons. These are associations of time and place when and where we met them ; there will be a play of vascular innervation dependent upon the excitation of fettered or unfettered moods; many association- fibres above and below the level of consciousness will be called into activity,— all of which, and a complicated process it is, will be due to the sight of these two persons. The sight of the one beloved will excite an unfettered mood which is based upon a functional hyperaemia, and which will inhibit the vaso-motor centre at the beginning of the systole of its vascular wave. The sight of one we fear will excite a fettered mood, giving rise to associations, with sensations of vascular innervation, which in- crease the arterial systole of the vascular wave. ■ The nutrition of the brain will be impaired by rapulsive emo- tions, for it is undoubtedly the arterial diastole which removes waste-products from the brain. We have learned to recognize Weight of Brain. 255 that the brain, by dint of the variable volume of the cranial cavity, with the aid of the muscular apparatus of its blood-vessels, con- tains within itself its metabolic mechanism ; it is a pressure- and suction-pump, forcing out ventricular fluid, absorbing lymphatic fluid from the various tissues into the perivascular spaces, and pumping it from these into subarachnoidal spaces. The state of the nutrition of the brain has an important bearing upon its weight which we must now consider without regard to its specific weight. Neglecting, for the present, the statements of older German authors, such as Huschke and Rudolf Wagner, we find, according to the valuable statistics of Bischoff, that the weight of the brain varies in man between 1,018 and 1,925 grammes; in woman, between 820 and 1,565 grammes. The average would be: For the male brain, 1,362 grammes ; and for the female, 1,219. Pfleger's investigations have shown that this statement must be modified if the calculation is based upon adult brains alone, between the ages of 19 and 58, by which means we get rid of a possible error due to senile atrophy. Weichselbaum calculated the average brain-weight of 390 Austrian soldiers, between the ages of 20 and 48 years, to be 1,373 grammes; while Pfleger, who took the brain of men up to the 59th year, proved this weight to be but 1,321 grammes. This discrepancy I would not ascribe altogether to the ten years of more advanced age, but rather to the more powerful organization of soldiers, whose exalted nutrition would tell in the case of the brain also. As regards sex, there is an undoubted difference of ten per cent, (according to Pfleger of thirteen per cent.), in favor of the male. Nor is this difference proportionate to the difference of stature. The average statures of man and woman are to each other as 100 : 93.2, while the male and female brain hold the rela- tion of 100 : 90.93. Furthermore, various authors, and among them Le Bon and Bischoff, are agreed to this, that there is a certain proportionate difference between the length of the body and the weight of the brain, but that the difference between the brain-weights of various individuals is far greater than could be accounted for by the rela- tive lengths of the body alone ; and that this difference must undoubtedly depend upon other conditions. It is worthy of note that short persons have a relatively larger brain than large persons. The largest brain relatively speaking, is that of the new-born 256 Psychiatry. infant. Its increase is so marked during the first period of extra- uterine life, that Tuczek, basing his statements upon the investi- gations of Huschke and Bischoff, contends that during the first year of extra-uterine life the daily increase in the size of the brain amounts to more than one cubic centimetre,—i. c., about the size of a bean. There is a direct connection between the growth of the brain, the size, and the quantity of blood of the new-born infant. According to Hermann, the blood of the new-born amounts to -^, of the adult to T\ only, of the entire weight of the body. There is a direct connection also with the greater width of the arterial system in the child, as was shown by Benecke ; and there is no doubt that the greater number of pulse contractions has an important influence upon metabolic changes in the body. Benecke attributes these to the smallness of the infantile heart, for the cardiac contractions are effected by shorter muscular fibres in a shorter space of time. In the female, the lesser size of the heart and the narrowness of the vascular system are important factors in determining the growth of her brain, and to these fac- tors must be added the higher percentage of water in her blood. Briicke quoting Valentin's tables on the sexual difference in the composition of blood, states that the percentages of water in the blood of man and woman are as 77.19: 79.11 ; of the solid resi- due as 22.1 : 20.89; ^he number of blood corpuscles as 14.1 : 12.79. No positive conclusions can be drawn from the weight of the brain ; the statistical average might be utilized, but the individual case certainly not. To take a single instance : The large size of the skull in taller persons depends more especially upon the gen- eral osseous system and the length of body (Pfleger). The brain filling this cavity need not necessarily contain a larger number of nerve-elements, but it is more likely that it will contain fibres of greater length, for their development will be hampered less, the greater the width of the skull. But that the greater length of nerve-fibres implies greater functional activity, as a larger num- ber of elements would do, has not been proven. I have given preference to the statistical results of Weich- selbaum and Pfleger over those of other authors, simply because the former have enabled us to draw some conclusions as to the proportionate weight of the different divisions of the brain. They have made use of my method of dividing the brain : Follow- ing my example, they do not join the cerebellum to the pons and Method of Weighiiig Brain—Statistics. 25 other parts of the brain ; and, furthermore, they regard the brain- trunk, including the prosencephalic ganglion, as the third impor- tant factor in calculating the weight of the brain. This method of separating the brain-trunk from its mantle, by cutting through the corona radiata, is illustrated in Figs. 16 and 17. The brain, together with its membranes, is weighed first, and its total weight is calculated from the sum of the weights of each division of the brain. The brain-trunk also is divided up in the manner I have indicated, into the lobus candicis, including the fore-brain gan- glion and the island of Reil; into the optic thalamus, the region of the corp. quadrig. and crura cerebri; and into the pons and oblongata. In the following table I propose to give (for the sake of comparison, and to bring out prominent and instructive differences in weight between the various divisions of the brain in man and animals) some statistics regarding the proportional weight of the different parts of the brain in various animals and in man. The animal brains here referred to were selected by chance rather than by intent, from a far larger (compara- tive) anatomical collection : Ba O C O 1-. 0 si 'o 0 '5-■5 3 u 3 O Cfl 3 m-E 3 c _o ■3 J3 a u a V c 0 Ph 420 230 350 58 19 6-5 12 394 256 349 350 180 470 40 30 10 IO 40 16.9 15.3 IO.7 300 255 444 328 379 290 34-5 20.4 16.9 10.7 25 12.5 19 15 Man (adult) '' (new-born) Monkey Elephant * Horse a " b . . . .Seal Bear .... Dog ... Pig . . . . Roe ... Cat .... New-born rabbit 780 830 708 630 604 698 ^73 644 728 615 622 614 611 105 057 085 239 190 105 148 146 090 120 122 105 112 20S 125 204 196 177 209 1S1 265 255 140, 245 063 324 4-5 10 16.9 17-3 19 In the preceding table, the proportional weights of the fore-brain constitute a pro- gressive series, beginning with the new-born infant, ascending thence to adult man, and in due succession to the dog, the monkey, the horse, the seal, the elephant, and the bear ; whereas one horse (a) follows after the pig, roe, cat, and the new-born rabbit. To make this series thoroughly instructive, we must note that the greater percentage of hemispheres in the new-born infant is due to the slight weight of its cerebellum ; the same is true of the new-born rabbit. We must note, furthermore, that the other division of the fore- 1 The total weight of the elephant's brain was 4,576 grammes ; the brain-mantle •weighed 2,906, the cerebellum 1,097, and the brain-trunk 575 grammes. 258 PsycJiiatry. brain, the lobus-caudicis stands highest in the series, amounting to 58 per cent., or more than half of the entire brain-trunk. From the point of view of comparative anatomy, we shall find that the larger the animal, the greater the weight of its cerebellum. The elephant excels in weight of cerebellum, both relatively and absolutely, doubling the weight of the human cere- bellum, and having a disproportionate advantage as regards cerebellar percentage over other animals : On the other hand, among more closely allied mammalian forms. we find that there is a difference of twenty per mille (as regards cerebellar weight) be- tween man and a young monkey, in favor of the former. The new-born infant and the new-born rabbit stand greatly in arrears, as compared to adult man, the infant showing a difference of nearly fifty per mille (057). The preponderance of the elephant as regards cerebellar weights sets his hemispheres far back in the series ; the seal, and even the bear, going ahead of him. The recognition of such broad differences as these seems to me to be of far greater value than the consideration of refined minutiEe. In the above table the frontal lobe is regarded as extending as far as the fissura Rolando ; the parietal lobe thence as far as the occipital fissure ; and the remainder is put down as occipito-temporal lobe. Comparing the development of these several divi- sions in man, the monkey, bear, and dog, we find that the frontal lobe is more highly developed in man than in the monkey ; that of the monkey stands higher in the series than the frontal lobe of the carnivora (dog and bear). The parietal lobe is also less developed in monkey than in man. His larger occipito-temporal lobe is due to the excessive development of the occipital portion. The bear, like all carnivora, dis- tinguishes himself from man by the superior development of the parietal region. In. the new-born infant, this region is proportionately larger than in the adult, a fact which can be explained by reference to the observation of Zuckerkandl, to be stated in full hereafter. Zuckerkandl states that the growth (lengthwise) of the median portion of the hemispheres is furthered by the two cranial sutures (the coronal and lambdoidal), which are responsible for the longitudinal growth of the skull itself ;. but that there are no such transverse sutures favoring the unchecked development of the occipital and frontal lobes. As regards the subcortical ganglionic masses from the thalamus on, we note that man is inferior to other animals as far as the thalamus, and the mesencephalon in particular, are concerned ; in the monkey the thalamic region stands higher than the mesencephalon. The greatest proportionate difference is found in the case of the medulla oblongata. Its relative size in the cat is four times that in man. This is due in part to the larger quantity of connective tissue in animals, and to the excessive de- velopment of the posterior division of the brain-trunk as compared with the pyramidal tract. Taking into account the difference quoted above between the brains of two horses, and remembering that in another series of similar investigations I found the hemispheres—the brain mantle—in dogs and horses to amount to 67 per cent, of the entire brain surface, we may draw the inference that the difference, as regards brain- weight between animals of the same genus, is no less than between various human subjects. Satisfactory results must, therefore, be based upon a very large statistical collection. The greater difference will probably be found between species differenti- ated by a process of selection. From Pflegers and Weichselbaum's statistics, based upon the weight of adult brains, we take the following table giving the absolute and proportional weights of the three grand and primary divisions of the brain : Weights of Brain-Divisions. 259 Weight (in grammes). Per mille. of the entire brain. u a a 5 E _3 ai 1 o S _3 c.a W-5 5 e J3 4> 0 •5 = - 3 "IS pa a 1) 0 Brai trun 1.373 1,092 148 133 795 108 97 From 390 soldiers of different nationalities, between the ages of 20 and 48 years. Average length of body = 171 ctm. = 5 ft. 6 in. 1,321 1,044 142 135 790 107.5 102 From 107 healthy (sane) men between the ages of 20 and 59 years. Average length of body, 166.5 ctm. = 5 ft. 4^ in. 1,189 936 131 122 787 110 103 From 148 healthy wo-men between the ages of 20 and 59 years. Average length of body, 156 ctm. = 5 ft. 1 in. 1,154 908 128 118 787 III 102 From 377 healthy (sane) women varying between 20 years and old age. Average length of body, i« crm. = 5 ft. 1 in. Pfleger insists on the relatively greater development of the hemispheres in man as compared with those of woman; the exact relation being as 795 to 787, on the scale of 1,000. Engel has shown that this assigns the larger cerebellum to woman during the prime of life. No inferences can be drawn from the tables furnished above as regards the composition of the hemispheres; neither does the literature of the subject give us any clue either to the propor- tional weight of the several lobes of the hemispheres, or to the relative weights of the various subdivisions of the brain-trunk. I propose, therefore, to supply this deficiency, and to amplify the physiological law of brain-weights by an analysis of a large num- ber of brains, which, though the brains of insane patients, were not characterized by any marked degree of atrophy. The sub- jects from whom these brains were taken were all classified in the official list of the Vienna Insane Asylum under the head of what Griesinger termed " Conditions of maniacal excitement." As Pathologist to that institution from the years 1866-71, I had occasion to weigh 733 brains, and from among these I have the notes of the brain-weights of forty-six male and seventy female subjects, who had been classed under the heading mentioned im- 260 Psychiatry. mediately above. This group of cases comprises the heaviest brains, both male and female. For the sake of comparison, the absolute and relative weights of these " heaviest " brains are tabu- lated below. Adopting Bischoff's method, I have also divided the cerebral hemispheres into frontal, parietal, occipital, and temporal lobes, and have ascertained the average weights of 400 frontal lobes, 235 parietal lobes, 135 occipital and 210 temporal lobes. A more rational division could, however, be made between the occipital and temporal lobes. Weight in Grammes. 4) • u S "5 '3 C i_ w s i | 2 ? 23 ~ "3 . 'SI Occipito-temporal lobe. in 2 0 H ■ e s 0 a, ,-. it -C 1=5 d O to C O Ph ci ■ — SO. •c J S'o Male Female . 1,383 1,221 1,085 954 148 135 148 132 450 400 251 2l6 383 338 81.36 74.84 26.40 23-74 9-03 7.6l I6.58 14 23 6.2s 5-50 Per Mille. 1 V £ i 4) ^ X2 •5 g "e3 5 a; remember something often raises his eyebrows, as if to see it.1 Engel calls attention to the fact that a person who suddenly finds his thoughts checked will walk slowly or come to a complete halt; he will move on again, as soon as his thoughts move. From the above we see that physiognomical expression is chiefly a matter of sec- ondary presentations, which are evolved, like dream presentations, from the condition of partial sleep ; and that expression is dependent altogether upon the simultaneous excitation of such secondary presentations as are associated with our emotions or our thoughts. I have a few words yet to add in regard to the emotions, of which I have spoken repeatedly. I discussed their mechanism in the preceding chapter, but for the sake of simplicity did not go into details. We had contrasted only the emotion of passive grief with that of joy and con- tentment. The emotion of passive grief is accompanied by ideas of repulsion in regard to the outer world, but not by active impulses of aggression. This emotion is expressed by a narrowing of the cavities of the face and by a dearth of movements—a failure to assert one's own personality among the objects of nature. The corrugators draw the skin under the brows inward, as if to form a roof to shut out the light ; the lips are pressed together, the nostrils are depressed, for respiration is weaker, and the features are immovable, in consequence of a monotony of the thoughts. The expres- sion may become complicated through the united action of the inner fasciculi of the frontalis and the corrugator bringing into prominence the so-called wrinkles of grief. The inner end of the eyebrow will be raised, and we then get an expression which, if dependent upon allied associations, may be interpreted to mean that the afflicted one is looking up at his fate or for help. Passive grief is marked by inattention ; the emotion of fear, by greater excitement and attention. In grief there is nothing like the defensive cry brought on by fear. In fear the eyes are kept open, and flight is the active expression of such fear. Move- ments of flight maybe started by allied associations, even in the absence of real danger, and in spite of the will of a person. Fear is probably the neurosis of a subcortical centre (of the oblongata), excitation of which defies cortical inhibition. Darwin relates the following experience : " I put my face close to the thick glass-plate in front 1 In this paragraph Meynert quotes almost verbatim from Darwin's " Expressions of the Emotions in Man and Animals." Vid. Am. cd., pp. 33 and 33. The translator has, therefore, rendered this passage largely in the words of Darwin.—S. Appendix. 277 of a puff-adder in the Zoological Gardens. Every idea of danger was entirely sec- ondary ; but as soon as the animal struck at me, my resolution went for nothing, and I jumped a yard or two backward with astonishing rapidity." ' Fury is a painful emotion, which throws, the entire cortex, including its sensa- tions of innervation, into a condition of hyperaesthesra ; and this hyperaesthesia is characterized by great tension of all the muscles, which are in readiness, as it were, to perform aggressive, destructive movements. The phenomena of expression, which we considered hitherto, depended upon irradiation or secondary associations (concepts). But the thoughts and impressions of the physiognomist himself bear the stamp of secondary (parallel) associations. Recognition of an oft-repeated impression is effected with astonishing ease, and as surprising is the mass of physiognomical material stored up in our memories. My own eyesight is very good, and I can recognize at a distance a person whom I never thoroughly looked at. I recognize him, even if he be turning a corner at the very moment I spy him, by the peculiarity of his gait, by a movement of the shoulders, by the way he carries his head, all of which peculiarities have uncon- sciously been imprinted upon my mind. Physiognomical associations are based, therefore, upon innumerable details, which have been registered in the brain. The special features of a person aid us none in judging of his conscious actions, and yet we cannot help inferring similarity of volitional actions in two persons from simi- larity in their cast of features. Such conclusions—secondary associations—are the result of impressions which we can scarcely define in words, of innumerable images deposited in the fore-brain. The features of the face, the development of permanent wrinkles, will depend much more upon nutritive changes, upon tension or relaxation of the skin, upon the loss of its elasticity in old age, than upon fixed innervations repre- senting actual forms of expression. Secondary associations (allied concepts) lead us to put an entirely erroneous interpretation upon movements of expression, which are indeed so variable that we are frequently unable to distinguish between the beginning* of crying or laughter. Hanns Virchow has succeeded admirably in suggesting a very fine physiognomical secondary association in regard to the interest attaching to the pupil of the eye. The pupil, he says, is the portal through which we look into the "innermost recess" of another person. A psychical secondary association is connected with the term " inner- most recess." We are reminded of the sensation of an " unfathomable depth, "which a child experiences on looking through the door of a cellar or down into a well. There can be no doubt, therefore, that our interpretation of expressions is simply a matter of secondary associations. Physiognomists—and all persons are such—are apt to be deceived as to what they really see. A person's voice and external appearance may remind us so vividly of another's that, at first sight, we recognize that she must be the sister of the second person. Placed side by side, these two sisters may present entirely different features ; the resemblance will lie altogether in certain movements of expres- sion, including those of speech, which one has copied from the other. But the physi- ognomist referred the resemblance to a similarity of features. I have referred to the condition of the observer—the physiognomist—in order to prove that his judgment is as largely dependent upon the secondary associations of partial psychical sleep as are the phenomena of expression which he has been observing. And to such a conclusion we can surely not ascribe the value of an unerring judgment. There is another order of secondary concepts, exerting their influence in every phase of consciousness, and dependent upon those sensations of muscular innervation which represent one's own features and carnage, as copied by one individual from. 1 Darwin, toe. cit., p. 38. 278 Psychiatry. another. This image of the personality is subject in most people to certain uncon- scious aesthetic concepts, which help the individual to govern the innervation of his own features and movements. If movements of expression indicate, to the extent mentioned above, all possible relations to the outer world, or the lack of such relations, then the features and carriage of a person may be said to define his own personality independently of all external considerations. These aesthetic considera- tions modify personal appearance according to the power of the fore-brain over all innervation, in proportion to the influence of imitation, of taste or lack of taste, which are important factors in the matter of personal appearance, and according to the culture and mental calibre of the individual. The control which a person possesses over the most movable body-opening—the mouth—may be taken as a measure of his mental strength. Uneducated persons lack this control altogether. The finest shades of expression, which cannot be aptly described in words, are indicated by changes in that depressed region of the face sur- rounding the lips, bounded above by the naso-labial and below by the .genio-labial fold, which region, according to Langer, is never disfigured by the accumulation of adipose tissue. By far more universal are those physiognomical symbols which the eyes offer and which vary with slight changes in pressure, with a change in the lustre, the position, and the shadow of the eyes. Every one will be able to distinguish whether certain forms of expression, as indi- cated by the features or the carriage of an individual, result from unconscious innerva- tion with the aid of secondary concepts, or from conscious innervation in a condition of partial wakefulness. Volitional mimical movements lack the character of spon- taneity, and become ridiculous as expressions of foppishness or excite contempt as expressions of falsehood. Intentional movements of expression lack that very charm which is inherent in the unconscious mechanism of expression. We shall gain a clearer insight into this subject from the study of the pathological changes of expression ; and such changes we shall have reason to consider pathog- nomonic of certain morbid states of the mind. INDEX A Acoustic nerve. (See nerve, auditory.) Adamiick; investigations on centres ocular muscles, 205 After-brain. (See medulla oblongata.) Aggression. (See movements.) Ala cinerea, 36 Alveus, 24 Amphioxus, 1 Amygdala, 46 Animals, speech of, 275 Ansa lenticularis, 50, 81 Ansa peduncularis, 45, 47, 50 Ape-fissure, 9 Apex-process, 65 Aquaeduct of Sylvius, 30 Arched fissures of carnivora, 17 Arndt, 59, 62 Arnold, 24, 36 ; fasciculus of, 85, 137 ; lattice-like layer of, 88 Arteries of brain, 220 Association-fibres, 39 ; functional im- portance of, 153 Association, logical aspect of, 177 ; rela- tion to induction, 179 Associations, secondary, physiognomical importance of, 271, 275 Auditory bundles, 120, 121 ; communica- tions, reflex origin of, 203 Axis-cylinder, composition of sheath, 233 ; development of, 64 ; fibrillary structure of, 243 ; nutrition of, 243, 245 B Baer, Carl v., 1 Base-process, 60 Bat, sensations of, 171 Bechterew, on voice-centre, 203 Bell, law of conduction, 138 ; on closure of the lids, 273 TO PART I. Bergmann, 36 Besser, 54, 62 Betz, 70 Bindearme. (See process, cerebell. ad. cerebrum.) v. Bischoff, 13-15 ; on weight of brain, 255 Blankenhorn and Gamgee, on protagon, 235 Blushing, 251 Bochaldek, on vaso-motor nerves, 220 Boll, 56-67 Brachia of the corp. quadrigemina, 101 Brachium-pontis, 28, 115 Brain, aqueous extract of, 235 ; cisterns of (see lymph-cisterns) ; convex sur- face, 3 ; median surface, 18 ; minute anatomy of, 56 ; movements of, 215, 227 ; nutrition of, 213 ; of bear, 6 ; of hamadryas, 7 ; of monkey, 23 ; of mus- tela, 9; weight of, 255 ; sexual differ- ences, 261 Brain-axis, dissection of, 25 ; develop- ment of, in man, 28 Brain-trunk, dependent upon fore-brain, 142. (See also above.) Breuer, on static vertigo, 209 Briicke, on forced movements, 209 Burckhardt, on movements of brain, 215, 217, 229 C Calcarine fissures, 23 Canals, semicircular, as organs of equilib- rium, 263 Capsula interna, general anatomy of, 82 ; relation to cerebral peduncle, 81 Cell-process, 58 Cells, cubic, 61 Central gray substance, 35 | Centres, subcortical, 156 279 28o Psychiatry. Cerebellar co-ordination, 210 Cerebellum, conduction to and from, 173 ; connection with auditory fasciculi, 209 ; cortex of, 116; description of, 115; development of, 1-33 ; nucleus dentatus of, 118 ; as organ of sensation, 211 Cerebrin, 237 Cervical flexure, 3 Charcot, arterial terminology of, 221 ; views on paralysis, 173 Cholesterin, 236 Chorda dorsalis, 3 Choroid plexus, 19 Cingulum, 24, 40 Clarke, 35, 54 ; auditory nucleus of, 118, 126 ; structure of cortex, according to, 75 Claustrum, formation of, 71 Commissura anterior, 45 ; atrophy of, fol- lowing extirpation of olfactory lobes, 266 ; medullary formation of, 72 ; rela- tion to olfactory lobes, 75 Commissura inferior, 95 ; media, 94 Commissura, posterior, bundles to thala- mus, 133 Commissure of Wernekinck, 53 Conarium, 31 Convolutions, homology of, 16 ; index of intelligence, 260 ; laws of development of, 14 ; mechanical laws governing de- velopment of, 262 Convulsion-centre, 212 Cornu ammonis, 3, 24 ; cortical structure of, 70 Corona radiata, bundles to optic thalamus, 76 ; relations to lenticular nucleus and crus, 78 Corpora quadrigemina, 31 ; connections with corp. genie, ext. et int., 100, 101 ; connections with cortex, 102 ; periph- eral connections of, 103 Corpus, callosum, 3, 20; candicans, 27 ; genie, externum, connections with optic tract, 100; restiforme, 28, 36, 126; origin of, 118 ; relation to olivary bodies, 128 ; striatum, development of, 3 (see also nucleus caudatus and nucleus lenticularis) ; trapezoides, 28 Cortex, cerebellar, 116 Cortex, cerebral, anatomy of, 37, 56, 59 ; arterial supply of, 214 ; functions of, 195 ; of occipital lobe, 68 ; relation to outer world, 139 ; separation of sensory and motor areas, 141 ; as vaso-motor centre, 249 Cortical inhibition, 197 Cortical localization, proof of, 141 Crus cerebri. (See pedunculus cerebri.) Crusta. (See pes pedunculi.) Cruveilhier, on venous spaces, 216 Crying, in children, 272 Cuneus, 15 ; thalami optici intergenicu- laris, 102 D Danilewski, on proportion of gray and white substance in brain, 232 Darwin, on inheritance of ideas and criti- cism of his views, 274 Deiter, 35 ; lateral mixed system of, 124 Demoulins, on injury to cerebellum, 210 Diakonow, on protagon, 236 Di-encephalon. (See inter-brain.) Discus lentiformis, 53, 85 Ditmar, on rise of blood-pressure after sensory stimulation, 1S6 Drechsel, views of, on protagon, 236 Duchenne, on movements of expression, 276 Dura mater, blood-supply of, 217 Duret, on arterial blood-supply of brain, 221 Dyspnoea of tissues in relation to move- ments of repulsion, 186 Ecker, convolutions of, 16 Ego, primary and secondary, 175 ; primary conception of, 169 Eminentia collateralis Meckelii, 23 Eminentia teres, 35, 123 Emotion, influence upon vascular wave, 253 ; in relation to aggression and re- pulsion, 184 ; the result of association, 192 Engel, 36 ; on relation between thoughts and movements, 276 Ep-encephalon. (See cerebellum.) Ependymal formation, 56 Eulenburg and Landois, experiments of, on relation of cortex to changes in tem- perature, 189 Index to Part I. 281 Expression, mechanism of, 271-278 ; movements of, in a child, 272 Eye, in expression, 277, 278 Eyelids, mechanism of closure of, 156 F Fascia dentata Tarini, 21, 24 Fasciculi bigemino-geniculares, 101 Fasciculi marginales aquaeducti, 104 Fasciculus, arcuatus, 42 ; basalis internus (Burdach), 41 Fasciculus longitudinalis posterior, 96, 133 ; function of, 174 ; formation of, 205 ; radiations of, 98 Fasciculus retrofiexus, go, 133 Fasciculus uncinatus, 42 Fear, expression of, 276 Fechner, views on activity of brain, 214 ; views on spontaneity of motor actions, 248 Feeling, divisions of, 193 Ferrier, motor areas of, 143 Fibrae arcuatae, different divisions of, 126 ; relation to auditory tract, 120 Fibrae propriae, 39 Fimbria, 18 Fissura calcarina. (See sulcus calc.) Fissura, interparietalis, 8 ; parallelis, 9 ; prae-occipitalis, 15 Fissure of Rolando. (See sulcus centralis.) Fissure of Sylvius, 4 Flechsig, 55, 64 ; investigations of, on de- velopment of medullary substance, 268- 270 Fleischl, 58 Flocculus, 29 Flourens on forced movements, 209 Foramen of Monro, 3 Fore-brain, anatomy of, 3 ; development of, 1 ; ganglia of, 25, 75 ; physiology of, 139, etc.; as seat of intelligence, 149 Forel, 53 Fornix, 3, 18, 21, 8g Fossa, ccerulea, 35 Fossa of Sylvius, 2 Foveae glandulares, 216 Foville on cerebellar lesions, 211 Freedom, phenomena of, 196 Frenulum, 102 Frontal flexure, 3 Frontal fissures, etc., 9 Frontal lobe, 9, 10 Funiculus, cuneatus, 36, 127; gracilis, 36, 127 ; lateralis, 28, 125 ; posterior of medulla, 127 Fury, 277 G Geniculate bodies, 32 Geoghegan, on nuclein, 233 Gerlach, 63, 65 Girdle-formations of ganglia, 95 Globus pallidus, 80 Glomerulus olfactorius, 70 Goll, column of, 37 Goltz, croak experiment of, 188 ; on localization of functions, 149 ; on spinal vaso-motor centres, 185 ; on static ver- tigo, 209 ; on voice centres, 203 Granules, 60 Gratiolet, divisions of corp. striatum ac- cording to, 76 ; fasciculus of, 137 ; on movements of expression, 276 Gray substance, of cortex, 56, 58 ; spe- cific gravity of, 232 Grief, emotion of, 276 v. Gudden, 27 ; experiments of, 266 Gyrus fornicatus, 3, 20; in relation to sense of smell, 141 ; fusiformis, 15; longitudinalis, 10 ; orbitalis, 10 ; transi- torius, 10; uncinatus, 22 ; cortical structure of, last, 70 H Habenula, 31 Helmholtz, on space-image, 179 Henle, 14, 39, 59, 62 ; mechanics of brain development, 262 Hensen and VSlckers, investigations on centres for ocular muscles, 205 Hering, on peristaltic movements of arte- ries, 224 Heubner, on blood-supply of brain, 217, 221 Hind-brain. (See cerebellum.) Hitzig, 18 ; motor centres of, 70, 143 Hollander, 94 Hoppe-Seyler, on chemical composition of brain, 235, etc. Huschke, 12, 22 ; on cerebellar co-ordi- nation, 210 2S2 Psychiatry. Hyperaemia, functional, of brain ; changes of, 215 ; how guarded against, 247 ; relation to emotion, 274 ; relation to thought, 194, 214 Individuality, discussion on, 168 Induction, mechanism of, in brain, 153 Infundibulum, 20, 27 Inhibition, 185 Inspiration, physiognomic value of, 272 Instinct, 169 Intelligence, seat of, 149 ; mechanism of, 151 Inter-brain (see also optic thalamus) ; de- velopment of, 2, 3 ; ganglia of, 25 Irradiation, 185 ; in movements of ex- pression, 275 Island of Reil, 2,4; development of, in man, and relation to speech, 142 Isolation, 64 J Jaksch, on nuclein, 233 Jastrowitz, 58, 59, 63, 64, 67 K Keratoid substance, 241 Key, on lymph-cisterns, 215 Kissing, physiognomical aspect of, 274 Koschewinkow, 65 Kossel, on chemical composition of brain, 235, etc. Kussmaul, on perceptions in utero, 168 Kuhne and Ewald, digestion, experiments on the axis-cylinder, 233 L Lamina perforata anterior, relation to corp. striat., 77 Laminae medullares, 80 ; connections with internal capsule, 83 Landois and Wolff, on pulse-waves of brain, 225 Langer, on venous spaces, 216 Language, symbols of, equivalent to cor- tical images, 192 Lateral column. (See funiculus lateralis.) Lateral mixed system (Deiter's), 124 Laughter, 273 Lecithin, 237 Lemniscus, 34, 52 ; origin of, 103 , con- nections of, 134, 135 ; relation to opt. thai, and spinal cord, 202 Lenhossek, 124 Leyden, views on ataxia, 144 Leveille, 34 Lids, closure of (see also eyelids); physi- ognomic aspect of, 273 Liebreich, on protagon, 236 Lingula, 33 Lobus candicis, 32 ; occipitalis, 14 ; parie- talis superior, 12 ; inferior, 13 ; quad- ratus, 13 Localization. (See cortical localization.) Loewe, 58 Longet, experiments of, on dogs, 151 Lotze, on psychical spinal cord functions,. 200; views on local signs, 180 ; views on muscular sense, 145 ; on repulsive movements, 152 ; on tactile sensation, 178 Lusanna, on cerebellar lesion, 211 Luys, 70 Lymph-cisterns within the skull, 214 Lymph-spaces, subdural, 217 M Mach, on static vertigo, 201 Magendie, on injury to cerebellum, 210 Malacarne, 34 Medipedunculus (Wilder). (See hrach- ium-pontis.) Medulla oblongata, development of, 1 ; centres in, 211 ; transition into spinal cord, 128 ; as vaso-motor centre, 206 Medullary sheaths, 64, 269 Membrane of Schwann, 60 Memory, seat of, 149 Mendel, phosphorus in urine, 242 Mes-encephalon. (See mid-brain.) Met-encephalon. (See after-brain.) Meyer, Ludwig, on cavernous spaces, 216^ on mechanism of brain development,262 Meyer & Cornwinder, phosphorus in plants, 233 Meynert, base-processes of, 65 ; fasciculus of, 90 Mid-brain, development of, 1 ; ganglia of, 25, 29 Middle cerebellar peduncle. (See bracfu ium-pontis.) Index to Part I. 283 Mischer, on spinal-cord connections of lemniscus, 202 Monoplegias, depending upon lesions of lenticular nucleus, 198 Mons, 33 Moods in relation to movements, 193 Motor areas, criticism of, 143 (see also cortical localization); distribution over cortex, 147 ; sensory character of, 145 Movements, aggressive, 176, 185 ; forced, 209 ; influence of, upon reflex actions, 201 ; inhibition of, 188 (see also under expression) ; primary and secondary, 156, 159 ; repulsive, 176, 185 ; spon- taneous, 155 Munk's views on localization, 145 N Nerve, as a conductor, 240 Nerve or nerves: I. (See olfactory bulb.) II. (See optic tract, etc.) III. (oculo-motor), nucleus of, 107 IV. (trochlearis), origin of, 108 V. (trigeminal) descending root of, 107 ; description of all roots, 110 VI. and VII., common nucleus of, 111 VII. (facial), anterior nucleus of, 112 ; knee of, no VIII. (auditory), anterior nucleus of Stilling, 122 ; external nucleus of, 120; relatio* to cerebellum, 115; relation to reflex impulses, 274 IX., X., XL, XII., nuclei of, 125, etc. Nerve-cells, chemical composition of, 232; respiration of, 184 Nerve-current, electrical, 242 Nerve of Lancisi, 21 Nerves of pia, etc., 220 Nothnagel, chromic-acid injections into lenticular nucleus, 161 ; convulsion centre of, 212 ; nodus cursorius of, 197 ; on motor areas, 143 Nuclein, 233, 245 Neuroglia, 56 Neurokeratin, 239 Nucleus caudatus, 2, 26, 76 ; proportion- ate development of, in man and ani- mals, 78 Nucleus, caudatus thalami, 89 Nucleus dentatus (of cerebellum), 53 Nucleus lenticularis, 2 ; description of, 79 Nucleus (tegmenti) ruber, 53 ; relations to internal capsule, 85 Nutrition, phases of, in cortex, 193 Nutritive attraction, principle of, applied, 247 0 Obersteiner, views of, on cerebellar cor- tex, 116 Obex, 36 Occipital lobe, convolutions of, 16; fissures of, 16 Ocular, movements, reflex mechanism of, 204; muscles, centres of, 205. (See also under Eye.) Olfactory bulb, 16 Olfactory lobe, 6, 18 Olivary bodies, 28, 127 ; relation to corp. restiforme, 128 Operculum, 5 Optic radiations, 48 Optic thalamus, description of, 86 ; con- nection (?) with auditory tract, 203 ; re- lation to upper extremity, 161 ; relation to various senses, 199 ; symptoms from lesion of, 165 ; parts of, 31, 47, 133 Optic tract, 95 ; connections with corp. genie, ext., 100 ; radiations from, 101 Orbital surface, convolutions of, 12 Owsjannikow, on oblongata, 206 ; on rise of blood-pressure after sensory stimula- tion, 186 P r Pacchionian granulations, in relation to subarachnoidal spaces, 219 Pain, in relation to movements of repul- sion, 191 Pansch, 14 Parietal arches, 13 Parietal flexure, 3 Parietal lobe, 12 ; fissures of, 12 Pedicles, of thalamus, 84, 89 Pedunculus (crus) cerebri, 27, etc. Pedunculus ganglii habenulae, 90 Pes pedunculi (crusta), 28 ; double origin of, 84, 137 Petrowski, analysis by, of gray and white substance 238 ; on percentage of water in brain, 235 284 Psychiatry. Pfleger, on brain-weights, 255, 259 Phosphoric acid, amount of, in body, 235 Phosphorus, in brain tissue, 234 ; in fcetal muscles, 234 ; in urine, 242 Praepedunculus (Wilder). (See processus cerebell. ad cerebrum.) Proc. cerebell. ad cerebrum, 34, 53, 106 Proc. cerebell. ad corp. quadrig., 34. (See frenulum.) Processus cerebell. ad med. oblong. (See corpus restiforme.) Process, cerebell. ad pontem. (See brachium-pontis.) Projection, relation to induction, 179 Projection-fibres, 39, 46 ; from occipital lobe, 86 Projection-system, divisions of, 79 ; func- tional importance of, 153 Prosencephalon. (See fore-brain.) Protagon, 236, 237 Psychical blindness, 145. (See also Munk's views.) Psychical deafness, 145 Purkinje, cells of, 117 Pulse-waves of brain, 225 Pulvinar, 31 Pyramidal tract, 50 Pyramids, 28 ; decussation of, 131 Q Quinke's cinnabar injections, 229 R Rage, expression of, 276 Ranvier's internodia, 60 Red nucleus. (See nucleus ruber.) Reinisch, 62 Renzi, on cerebellar lesions, 211 Repulsion. (See under movements.) Restiform body. (See corpus restiforme.) Retina, images of, how utilized, 180 Retzius, on lymph-cisterns, 215 Riegel, on peristaltic movements of ar- teries, 224 Rindfleisch, 59, 63 Ritter's laws of nerve nutrition, 240 Rohony, on amphioxus lanceolatus, 1 Rokitansky, 56 Rokitansky, Procop, on respiratory cen- tres, 186 Rudinger, on development of convolu- tions, 261 Rumpf, experiments on axis - cylinder sheath, 233 S Sander, 15 Schiff, experiments with colocynth, 151 ; on cutaneous irritation, 184 Schlesinger, on vaso-motor centres, 185 Schmidt, 22 Schnopfhagen's decussation, 95 Schroder, 35 Schultze, Max, 62, 64 ; on axis-cylinder, 243 Schwalbe on lymph cisterns, 215 Schwann, sheath of, 60 Screaming in children, 272 Sensations, correlated, 178 ; of innerva- tion, 144, 159, 182 ; in the blind, 171 Sense or senses, muscular in relation to cerebellum, 210 ; special, development of, 171 ; tactile, in relation to space, 178 Sensitiveness, of brain cells, 138 Sensory stimuli, in relation to arterial changes, 190 ; nutrition of nervous tissues, 269 Septum pellucidum, 20 Sighing, 273 Skull, measurements of, 263 ; relation to convolutions, 262 Soemmering, substance of, 51 Soltmann, on motor functions in children, 166 Sound, reproduction of, 208 Space-image, retinal, 179, 182 Space, perception of, 178 Space-vision, 181 ; in children, 274 Spiess, views on muscular sense, 145 ; on local signs of correlated sensations, 178 Spinal cord, cross-section of, 131 Stannius, 27 Stilling, 33, 35 ; anterior acoustic nucleus, of, 122 Stilling's wreaths, 51 Stilus of thalamus, 88, 93 Stratum intermedium caudicis, 55 Stratum intermed. pedunculi, 51, 54, 82, 136, 174 Stratum zonale, of medulla, 126, 127 ; of thalamus, 86 Index to Part I. 285 Stria cornea, 29 Striae medullares, of thalamus, 91 Substantia, ferruginosa, 109 ; connection with root of fifth nerve, no ; innom- inata, Reil, 94 ; nigra, 51 ; of Rolando, 130 Sucking, physiognomic aspect of, 272 Sulcus calcarinus, 23 ; structure of, 69 Sulcus, centralis (Rolando), 7 ; of bear, 17 Sulcus cruciatus, 11 ; magnus horizontalis, 29 ; praecentralis, 8 ; rectus, n Sylvian fossa, development of, 2 Tapetum Reilii, 44 Tegmentum, 55 ; origin of, 90 ; continua- tion of, in spinal cord, 133 Temporal lobe, fissures and convolutions of, 16 Thalam-encephalon. (See inter-brain.) Thalamus. (See optic thalamus.) Thought, methods of, 177 ; orderly evolu- tion of, 253 Tractus transversus pedunculi, 27 Trapezoid bodies. (See corpus trape- zoides.) Trigonum, cervicale (Goll), \i% Trigonum olfactorium, 20, 21 Trollard, 216 Tuberculum cinereum, (Rolando) 37 Tuczek, on increase of brain-weight, 256 Tiirck, 55 ; bundles of, 85 'Tween-brain. (See inter-brain.) U Uvula, 29 Vagus (X) see nerves. Vascular waves, of brain-movements, 228 ; during sleep and waking, 231 Vaso-dilators, 207 Vaso-motor centres, in cortex, 249 ; in medulla oblongata, 206 ; in spinal cord, 185 Veins of brain, 220 Velum medullare superius, 33 Vesicles of hemispheres,' of fore-brain, etc., 1, 2 Vieussens, 47 Virchow, 56 Virchow, Hanns, on expression of the eye, 277 Visual sphere, of Munck, 146 Volckers. (See Hensen.) 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