*J I I Columbia (MnitJersiftp intijeCttpofi^fttigork College of ^fjp^iciansf anb ^urgeonsf Xibrarp *' PRE SENTED IN MEMORY OF -^WILLIAMHENRYDRAPER 1830-1901-P.A.ND S.-1855 AND HIS SON WILLL\M KINNKUTT DMPER 1S63-1926-P.AND S.-1888 f Digitized by the Internet Archive in 2010 with funding from Open Knowledge Commons http://www.archive.org/details/textbookofmentalOOIewi JUST READY GORDINIER. The Gross and Minute Anatomy of the Central Nervous System. By H. C. Gordinier, a.m., m.d., Professor of Physiology and of the Anatomy of the Nervous System in the Albany Medical College. With many full-page Plates and other Illustrations, a number of which are printed in colors and the majority of which are original. Large Octavo. Handsome Cloth, ^6.00; Sheep, ^7.00; Half Russia, ^8.00, net. SYNOPSIS OF CONTENTS. I. The Histologic Elements of the Nervous System. II. The Spinal Cord. III. The Medulla Oblongata. IV. The Cerebellum. V. The Mid-Brain. VI. The Region of the Third Ventricle. VII The Membranes of the Brain. VIII. The Cerebrum (Forebrain or Pros- encephalon). IX. The Histology of the Cerebral Cortex, Cerebrum Ovale and as- sociation, Commissural and Pro- jection Systems of Fibers. X. The General Anatomy of Interior of the Cerebral Hemispheres. XI. The Circulation of the Brain. XII. Localization. XIII. The Embryology of the Nervous System. This is a systematic, practical treatise upon a most important subject that has been in great part neglected by English writers on neurology. Dr. Gordinier has devoted many years to the special study of the nervous system, and has produced a most useful work that will mark an epoch in the literature of medicine. The illustrations, of which there are a large number, are chiefly from the author's own preparations. They have been reproduced in the very best manner, the publishers' aim being to give results that are scientifically correct and at the same time pleasing to the eye. In order that certain pictures may be more faithfully shown, they will be printed in colors ; this will bring out the details perfectly, and enable the student to quickly recognize their relative value. Those illustrations borrowed from others have generally been remade, so that they will harmonize with the general style adopted for the work. In some cases these have been improved upon in details that the originals failed to make clear. The typography and paper used in printing are of the best, and the publishers have spared no cost to make their part correspond in perfectness with the scientific worth of the text. P. BLAKISTON'S SON & CO., Publishers, Phila. A TEXT-BOOK MENTAL DISEASES WITH SPECIAL REFERENCE TO THE PATHOLOGICAL ASPECTS OF INSANITY » BY W. BEVAN LEWIS L.R.C.P. (LoND.), M.R.C.S. (Eng.) MEDICAL DIRECTOR, WEST RIDING ASYLUM, WAKEFIELD; LECTURER ON MENTAL DISEASES AT THE YORKSHIRE COLLEGE ; EXAMINER IN MENTAL DISEASES TO THE VICTORIA UNIVERSITY Secont) BMtion THOROUGHLY REVISED, ENLARGED, AND IN PART RE-WRITTEN WITH ILLUSTRATIONS IN THE TEXT, CHARTS, AND TWENTY-SIX LITHOGRAPHED PLATES PHILADELPHIA P. BLAKISTON'S SON & CO IOI2 WALNUT STREET 1899 I: SIR JAMES CRICHTON-BROWNE, -M.D., LL.D., F.R.SS. (LOND. AND EDIN.), lORD chancellor's VISITOR IN LUNACY, LATE MEDICAL DIRECTOR OF THE WEST RIDING ASYLUM AT WAKEFIELD. 3^11 BDmicatton of THE VIGOROUS INTELLECT, COMiMANDING ELOQUENCE, AND UNTIRING ENERGY BROUGHT TO BEAR ON THE SCIENTIFIC ASPECTS OF PSYCHOLOGICAL MEDICINE DURING HIS DIRECTORATE OF THE WEST RIDING ASYLUM ; Mnb in Tkeen Appreciation ot HIS WIDE-SPREAD SYMPATHIES AND G E >J E R O U S IMPULSES, ^bis IKIlorf^ is De^icateJ) BY THE AUTHOR. PEEFACE TO THE SECOND EDITION. Since the appearance of the First Edition of this Treatise great advances have been made in our knowledge of the intimate structure of the Nervous System. The new methods of Golgi and his School have thrown light on many obscure points in neuro-histology, and the doctrine of the neuron and neuron chains has had a paramount influence upon problems in nerve physiology and pathology. It is hoped that the revision of the Anatomical Section faithfully reflects the change which has come over our con- ception of the nerve-cell, and that this section correctly and concisely expresses the more essential facts which recent re- search has established. The large number of additional plates and illustrations in the text will, it is believed, be welcomed in this section : most of them are from original drawings by the Author. In the Clinical Section will be found, as additional mattei' a chapter on Progressive Systematised Insanity ; an account of certain forms of Impulsive Insanity ; a description of the reaction-time instrument ; estimation of muscular sense dis- crimination in general paralysis, alcoholism, &c. ; and a separate chapter on the Treatment of Insanity. Much matter considered less essential has been eliminated. In the Pathological Section, the Author has maintained his views of the importance of the Scavenger-cell as a morbid factor in the changes found in the brain of the insane, nor does he find in recent criticism of this view any solid reason for a modification of his opinion as to the role played by it in insanity. Finally, he would express his grateful appreciation of the flattering reception accorded to the First Edition, and his great indebtedness to his Publishers, who have spared neither expense nor care in the production of the Second Edition. West Riding Asylum, Wakefield, December, 1898. ' • PPtEFACE. Ix wj'iting a new Treatise on Mental Diseases, I have not been blind to the wealth of available literature in this department, nor to the claims upon the Student's attention of such works as the classical Manual of Bucknill & Tuke, the English translation of Griesinger's Treatise, and the admirable Lectures of Dr. Blandford, nor again to the more recent additions to Dr. Bristowe's Clinical Medicine, and the works of Drs. Sankey, Clouston, and Savage. It has, however, been my special object to present a risum4 of our knowledge of the structure and connections of the cerebro-spinal nervous system, of the architecture of the cerebral hemispheres, and more especially of the cortical envelope as the essential organ — the material substratum — of Mind ; and to afford a concise account of the morbid changes found in the brain of the insane, as viewed in the light of recent research. It appears to me that a disproportionate amount of attention has been paid in former text-books to the clinical aspects of Insanity, and it is hoped that this attempt to deal more fully with the organisation of the material substratum of mind, and with the evidences of morbid change to which it is prone, will not prove unwelcome to the Student of Mental Disease. In the Anatomical Section, I have endeavoured to comprise such information as shall prove of utility to a more thorough conception of the ground-plan and superstructure of the nervous system ; and it will be at once evident that special emphasis has been advisedly bestowed upon the cortical envelope — tlie structure, nature, and autonomy of the nerve-cell. X PREFACE. The Clinical Section comprises statistics based upon an analysis of 4,000 cases of Insanity in both sexes, treated at the West Riding Asylum. In the Pathological Section, I have endeavoured to do justice to certain morbid processes, which appear to me to be of paramount importance in the history of Insanity ; and more particularly would I here allude to the functions of the Lymph- connective system of the Brain, and the life-history of the " Scavenger-cell." To my Publishers I would desire to express my acknowledg- ments for the consideration uniformly received at their hands, despite the delay which has unavoidably occurred — for the liberal supply of illustrations, so essential to the success of a work of this description, and for the special care taken in their production. I can add my testimony to the admirable faithfulness with which my drawings have been reproduced by Mr. Danielsson. I have also to express my obligations to my colleague, Mr. St. John Bullen, for reference to a compilation of Statistics from the Pathological Records of this Asylum, and for material assistance in the revision of the proof-sheets. West Riding Asylum, Wakefield, November, 1889. GENERAL CONTENTS. PART I.— ANATOMICAL AND HISTOLOGICAL SECTION. The Spinal Cord — The Medulla Oblongata — The Mesencephalon — The Prosen- cephalon — The Encephalon as a whole — The Cerebral Cortex — Cortical Lamination, .......... Pages 1-141 THE SPINAL CORD. The Cerebro-Spinal Axis — The Central Grey Matter — The White MeduUated Columns — Transverse Section of Cord — Substantia Gelatinosa and Vesiculai- Columns — Caput Cornu — Sacral Nucleus of Stilling — Anterior Cornu — Inter- medio-lateral Tract of Clarke — Formatio Reticularis — Conducting and Com- missural Tracts — Direct Pyramidal Tract — Crossed or Latei-al Pj'ramidal Tract — Posterior Median Column — Postei'O-lateral Columns — Direct Cerebellar Tract — Gowers' Antero-lateral Ascending Tract — Anteiior Radicular Zone and Lateral Limiting Layer, ....... Pages 1-8 THE MEDULLA OBLONGATA. Region of the Calamus Scriptorius — Clavate and Cuneate Nuclei — Solitary Fasci- culus — Nucleus of Fasciculus Teres — Floor of Fourth Ventricle (lower half) — Hypoglossal and Vago-accessory Nuclei — Ascending Root of Fifth Nerve — Motor, Mixed and Sensory Systems — Restiform Tract — Dentate Nucleus — Inner and Outer Divisions of Cerebellar Peduncles — Roof Nuclei of Stilling — Arcuate Fasciculi — Fillet — Nucleus of Lateral Column — Inferior Olivary Bod^- — Internal and Extei-nal Accessory Olives — Corpus Trapezoides — Origin of Hypoglossal Nerve — Mixed Lateral System — Spinal Accessory, Vagus and Glosso-Pharyngeal — Upper Half of Medulla — Abducens, Facial and Acoustic Nuclei — Superior and Inferior Olivarj^ — Inferior Facial Nucleus — Nuclei of Acoustic Nerve — Sound-rod of Bergmann — Acoustic Striie — Facial Genu — Ascending Trigeminal Root — Abducens Facialis — Superior Olivar}- Body — Lemniscus or Fillet — Upper Angle of Fourtli Ventricle — Posterior Longitudinal Fasciculus — Nuclei and Root-fibi'es of Trigeminal — Locu.s Cceruleus — Nuclei of Oculo-Motor and Tiocldcaris — Root-tibrcs of Fourtli Nerve, Pages 8-28 THE MESENCEPHALON. Advance in Complexity — Tegmental and Crustal Tracts — Internal Caj)sulc — Corpora Quadrigemina and Tiialami — Locus Niger — Knee of Capside — Posterior Perforated S])aee — Basal Aspect of Mcsenceplialon — Tiunia Pontis — Corpora Albieantia — Infundibuiuni and Pituitary Body — Conducting Tracts xii CONTENTS. of Crusta— Fundamental, Mixed and Accessory Systems — Cortical Termination of Tracts— Relationships to Motor Cells of Cortex— Termini of Sensory Columns of Cord — Radiations of <4ratiolet — Sensory Peduncular Tract — Constitution of Internal Capsule —Ansa Peduncularis - Substantia Nigra — - Fillet or Lemniscus — Pineal Body — Posterior Commissure — Aqueduct — Tegmental Structures— Superior Bigeminal Body or Nates— Its Brachia and Stratum Lemnisci — Red Nucleus — Upper Cerebellar Peduncle — Posterior Longitudinal Fasciculus —Substantia Nigra — Relationships and Connections of Tegment and Crusta, Pages 28-41 THE THALAMENCEPHALON. "~ External Conformation— Inner and Outer Face of Optic Thalami— Sub-thalamic Body— Thalamic Peduncles— Fimbria of Fornix— Pillars of Fornix— Cortical Connections of Optic Thalami — Lamina MeduUaris of Burdach — Centre ^Median of Luys — Stratum Zonale — Anterior Tubercle of Thalamus — Pineal Body and its Connections— Fasciculus Retroflexus — Posterior Commissure — Corpora Geniculata, Pages 41-49 THE PROSENCEPHALON. Configm-ation of Fore-brain — Lenticular and Caudate Nuclei — Lenticular Axis of Revolution — Relationships of the Lenticular and Caudate Nuclei— Head of Caudate Nucleus — Tail of Caudate Nucleus — Amygdaloid Nucleus — The Surcingle — Loop-like Disposition of Ganglia — Stria Terminalis or Taenia Semicircularis— Olfactory Area — External Conformation of the Lenticiilar Nucleus— The Claustmm and Insula— Globus Pallidus— Lamina Medullares of Lenticular Nucleus, Pages 49-54 THE ENCEPHALON AS A WHOLE. Comparative and Embryological — The Neural Tube in Amphioxus and in the Lamprey — The Brain in Fishes and Amphibia— The Five Vesicles of the Neural Tube — The Cerebral Hemispheres in Fishes— Hypoaria— Predomin- ance of Optic Lobes in Insects— The Reptilian Brain— The Brain in Birds— Development of Neiu-al Canal — Vesicles of the Fore-brain — Olfactory Lobe — Foramen of Monro— Ganglia of Fore-brain— Pitiiitary Bodj^ — The Neuro- enteric Canal— Vesicle of Mid-brain— Formation of Quadrigeminal Bodies and Crura— The Iter— Vesicle of Hind-brain— The Cranial Flexures— Forma- tion of Fissures, Pages 55-60 THE CEREBRAL CORTEX. Methods of Enquiry— The Grey Matter the Tissue of INIind— Histological Elements of the Cortex — Nerve-cells— Minute Structure — The Cj-toplasm — Nucleus — Dendrites and Thorns— Chemical Constitution— Physiological and Patho- logical —Chromatolysis— Fatigue — Lesion — Conligiu-ation and Nomenclature —The Neuron— Angular Cells— Granule Cell— Pyramidal Cell— Motor Cell- Pyramids of Comu— Cells of Purkinje— Cells with Ascending Axons— Cells of Peripheral Zone— Mitral Cells of Olfactory Bulb— Inflated or Globose Cell — Spindle Cells— Nerve-fibres of Cortex— The Primitive Fibril— Non-MeduUated Fibres— Axis-cylinder Process — Mediillated Nerve-fibre — Myelin Sheath — Keratoid Sheath — Fromann's Lines — Lantermanu's Dissepiments — Centric CONTENTS. XI 11 and Pei-iplieric Fibres— Staining of Axis-cylinder — Arteries of the Cortex — Intima — Tunica Media — Tiuiica Adventitia — Perivascular Channel of His — The Capillaries of the Cortex — Stigmata and Stomata — The Veins of the Cortex — The Connective Matrix or Neuroglia — Local Varieties — Cellular Elements of Neuroglia — Lj'mphatic System of the Brain — The Perivascular Channels and Lymph Channels Proper— Epicerebral Space — Pericellular Sacs — Cells of Adventitial Tunic — A Description of the Lymph System — The Lympli-connective Elements or Scavenger-cells — Vascular Process of Scaven- ger-cell — iid/e of Spider- or Scavenger-cells, .... Pages 60-101 CORTICAL LAMINATION. Laminated Structure of Cortex — Cerebral Hemisphere in the Rodent — Eight Types of Cortex — Transition Regions in Man — Distribution of the several Types — Upper Limbic Type — Modified Upper Limbic Type — Outer Olfactory Type — Inner Olfactory Type — Modified Lower Limbic Type — Extra-limbic Type — Type of Cornu Ammonis — Peripheral Zone — -Ascending Axons — Lacunar Layer — Striate Layer — Pyramidal Layer — Polymorphic Cells — Alveus — Fascia Dentata — Stratum Granulosum — Type of Olfactory Bulb — Stratum Glomerulosum — Mitre Cells - Granule and Medullated Layer — The Retina — Rods — Cones Plexiform Layers — Granule Layers — Spongioblasts — Ganglionic Cells — Cerebellar Cortex — Peripheral Layer — Basket Cells — Cells of Piirkinje — Connective Cells — Granule Layer — Moss Fibres — Diversities of Cortical Lamination — Regional Distribution of Ganglionic Cells in Cortex — The Clustered and Solitary Arrangements — Distribution in the Pig, Sheep, Cat, Ape, &c. — Significance of Fissures and Sulci — Fissures defining Distinct Cortical Types — Contrasts between Brain of Man and Lower Mammals — Lamination of Motor Area in Man— Five-laminated Cortex — Histological Structure of the Several Layers of Motor Cortex— Distribution of Motor-cell Groups — Transition-realms of Cortex — Specialised Areas — Acquirement of Structural Variations — Significance of Cell-groupings — Comparative size of Brain-Cells — ^The Nucleus of the Nerve-cell and its Role, — Electrical Excita- bility of Cortex —Latent Period of Stimulation and the Summation of Stimuli — Conditions AiTecting P^xcitabilitj' — Functional Equivalence of Cortex — Faradic Stimulation of Cortex — Extra-polar Conduction — Conduction to Lower Centres — Proximity of Psycho-motor Centres, . . Pages 101-141 PART II.— CLINICAL SECTION. States of Depression — States of Exaltation — Fulminating Psychoses — States of Mental Enfeeblement — Recurrent Insanity — Epileptic Insanity — -General Paralysis of the Insane — Alcoholic Insanit}' — Insanity at the Periods of Puberty and Adolescence — At the Puerperal Period — At the Climacteric Epoch — Senile Insanity — Treatment of Insanitj', . . . Pages 142-485 STATES OF DEPRESSION. Mental Depression Defined — Decline of Object-consciousness — Rise of Subject- consciousness — Muscular Element of Thought — Failure in the Relational xiv COXTENTS. Element of Mind — Sense of Environmental Resistance — Reductions to Automatic Levels — Sense of Effort — Restricted Volition — Enfeebled Repre- sentativeness — Transformations of Identity — Tlie Physiological Aspect — Defective Circulation — Xutritional Impairment — Explosive Xeuroses — Hunger of the Brain-cell — Painful and Pleasurable Mental States — Re- action-time in Melancholia — Degi-ees of Mental Depression — Clinical Varieties of Melancholia — Simple Melancholia — Delusional Melancholia — Hj-pochondriacal Melancholia — Melancholia Agitans — States of Mental Stupor — Stupor and H}-pnotism — Acute Dementia, . . Pages 142-178 STATES OF STUPOR. Stupor and Dementia — Etiolog}' of Stuporose States — Stupor and Hypnotism — Stuporose Melancholia — Acute Primary Dementia, . . Pages 179-189 STATES OF EXALTATION. Mania<;al Reductions — Failure of Attention — Enfeebled Synthesis — Transient Delusive States — Exalted Sense of Freedom — Impulsive Conduct — Xoctumal Crises — Seclusion Fosters Hallucination — Sexual Illusions — Stadium Melan- cholicum — Enfeebled Imagination — Bodilj- Symptoms — Periodicity of Maniacal Phenomena — Acute Delirious Mania, . . . Pages 190-204 FULMINATING PSYCHOSES. Uniform and Partial Denudations— Defective Control — The Xeurotie and Criminal Subject — Xatiu-e of Impulsive Insanity — Insane Homicidal Impulse — Exist- ence of Aura — Epigastric Aura — L'ncovering of the Brute Instincts — Relief of Mental Tension — Illustrative Cases — Suicide in Homicidal Subjects — Etiology — Effect of Physiological Cycles — Epilepsy — Masked Epilepsy — Alcohol and Impulsive Insanity— The Mimetic Tendency — Suicidal Impulse — Other Forms of Morbid Impulse — Kleptomania — Dipsomania — Erotomania — Obsessions — Imperative or Dominant Ideas — Insanity of Doiiljt — Aboulia, . Pages 205-220 STATES OF MENTAL ENFEEBLEMENT. 31ental Deprivation in Contradistinction to Developmental Ai'rest — Persistent Enfeeblement — Chronic Residue of Asylum Communities — Recoverability of Maniacal and Melancholic Forms — Consecutive Dementia —Delusional In- sanity — Genesis of Monomaniacal States — Environmental Resistance — Trans- formation Completed — !Mystic Symbolism — Illustrative Cases of Delusional Insanity — Monomania of Pride {J.O., E.T.) — Religious Monomania (J.B.) — Monomania of Persecution [E.C.) — • Progi-essive Systematised Insanity ( Paranoia ) — Primary Implication — Systematisat ion — Xeuro-pathic Basis — Secondary Systematised States — Tj-pical Psycho-neurotic Form — Folie a deux, Pages 220-235 RECURRENT INSANITY. Definition — Establishment of Labile Equilibrium — Prevalence at Sexual Deca- dence — Heredity — Influence of Neurotic Heritage and of Ancestral Intem- perance — Ata\'ism — Recurrence in the Congenitally Defective Subject — Morbid Excitement and the Moral Imbecile — Alternations of Excitement and Stupor — Hysteria and Menstrual IiTegularity — Eroticism (A . S. , Jf.A . JI.) — Recurrence CONTENTS. XV in Adolescence {M. C. W.) — Recurrence at the Climacteric (H.O.) — at the Senile Epoch (./. ,S'. ) — in Puerperal Subjects (M . 5. )— in Traumatic Insanity (B. L.) — Morbid Impulsiveness — Hallucination and Delusion (/. B.) — Prog- nosis — Treatment, Pages 235-255 EPILEPTIC INSANITY. Definition — Epileptic Neurosis — Immediate and Remote Results of Epileptic Discharge — Diffusion-currents — Nascent Nerve-tracts — Discharge from Sen- sory Areas — The Aura in Sensory Epilepsies — Epileptic Amaurosis, Hemian- opsia and Hemiantesthesia — Champing Movements — Pre-parox3'smal Stage — Pi'emonitory Stage — Special Sense Aur;e — Vasomotor and Visceral Auras — The Epileptic Paroxysm — Grand and Petit Mai — Post-paroxysmal Period — Post-epileptic Automatism — Case of E. C. — Status Epilepticus — Inter- paroxysmal Stage — P]pileptic Hypochondriasis, Automatism and Impulsiveness — Medico-legal Relationships — Impulse — Delusion — Malingering — Reg. v. Taylor — Treatment of Epileptic Insanity, .... Pages 255-284 GENERAL PARALYSIS OF THE INSANE. Prodromata — Egoism — Eaily Moral Perversion— Faihire of Re-representative States — Enfeebled Attention — Transient Amnesia — Vasomotor Derangements — Early Paresis — Second Stage — Delusions of the Paralytic and Monomaniac — Vanity and Decorative Propensities — Sexual Perversions — Facial Expression — .\rticulatory Impairment — Cerebral Seizures — Syncope — Epilepsy (/. F.) — Unilateral Twitching (/. S.) — Epileptiform Attacks — Conjugate Deviation — Case of H. P. — Apoplectiform Seizures — Monoplegise — Hemiplegia; — Muscular Sense Discrimination — Apparatus for Testing Appreciation of Weight: — Re- action-time — Oculo-motor Symptoms — Spastic and Paralytic Myosis — Mydriasis and Amaurosis — Reflex and Associative Iridoplegia — Cycloplegic Iridoplegia or Ophthalmoplegia Interna — Size of Pupils — Statistical Tables — Consensual Movements — Reflex Dilatation — Associated Irido-motor States — Significance of Pupillary Anomalies— Cycloplegic Forms — Spinal Symptoms — Deep Reflexes — Tabetic (jrait — Incontinence and Retention — Atrophy of Vesical Muscle — The Blood in (xcneral Paralysis — Clinical Croupings of (reneral Paralysis, ......... Pages 285-327 ALCOHOLIC INSANITY. Alcoholism and Age— Susceptibility at Certain Developmental Phases — Adolescent Period {F. S.) — Prevalence of Impulse — Influence of Sex, Heredity, Epilepsy, Cranial Injury — Ancestral Intemperance — Anomalies of Systemic and Visceral Sensation — Aural Hallucinations (J. J.^) — Delusions of Suspicion — Optimistic Delusions — Clinical Forms of Alcoholism — Mania a Potu — Amblyopia — Cutaneous Anajsthesia — Relapses — Case of W. W. — Homicidal Impulse (t/. S.) — Chronic Alcoholism — Physiological effects of Alcohol— Evolutionary Period — Mental, Sensorial, aiid Motoi'ial Sj'mptoms (./. J.^) — Anniesic Forms (,/. F.) Conditions of Mental Revivability (M. It. L.) — Delusional Forms (T. S.) — Instances of "Environmental Resistance" — Visceral Illusions — The Epi- gastric Voice — Various Illusory States {E, A. F.) — Evolution of Psychical Phenomena — The Nervous Discharge— Hallucinations as Determining Morbid Ideation — Augmented Sjjecific Resistance — Sensory Anomalies — Motor XVI CONTENTS. Enfeeblement (./. i?.) — Twitchings, Tremors, Stolidity — Reaction-time in Alcoholism — Muscular Spasms and Cramps — Oculo-motor Immunity — Nystagmus — Epileptiform Attacks — Hemiplegiaa ( T. P. and /. C. ) — Classifi- cation, Pages 327-370 INSANITY AT THE PERIODS OF PUBERTY AND ADOLESCENCE. Evolution of Puberty and Adolescence — Pubescence as Distinguished from Adol- escence — Antagonism of Growth and Development — Excessive Metabolism of Infancy — Acquisitiveness and Mimetic Characters of Childhood — Initiative Tendencies of Adolescence — Pubescent Insanity in the Female — Delusions and Hallucinations — Relapses at Menstrual Periods— Hysteric Type of Mania — Stupor Coincident with Menstrual Derangement — Case of F. W. — The Blood in Stuporose States — Case of M. A. H. — Etiology — Ancestral Influence — Periods of Susceptibility — Statistics of Hereditary Factors — Ovarian Derangements and Pubescent Insanity (^1. H.) — Amenorrhceal and Anjemic States— Influence of the Environmental Factors — Percentage of Haemoglobin in Cases of Stiipor — Pubescent Insanity in the Male — Sexual Divergence — Symptoms of Piibescent Insanity — Modified Forms {J. M. ) — Masturbatic and Uncomplicated Form of Pubescent Insanity — Etiology — The Moral Imbecile, Pages 370-398 INSANITY AT THE PUERPERAL PERIOD. Symptoms — Predominance of Mania — Intensity of the Morbid Process — Obtrusive Sexual Element — Hallucinations — Delusions of Suspicion — Prevalence of Suicidal Feelings — Etiology — Susceptibility of the Puerperal Period — Illegitimacy and Puerperal Insanity — Frequency in Primiparje — Condition of the Blood — Diminution of Htemoglobin — Prognosis — Treatment — Insanity of Pregnancy — Relatively Infrequent — Primiparte Show no Special Liability — Symptoms — Recoveries, ........ Pages 398-409 INSANITY AT THE PERIOD OF LACTATION. Risks attendant upon Lactation — Period of Uterine Involution — Period of Mammary Excitation — Symptoms — Depressing Delusions — Impulsive Nature (3/. W.) — Suicide(i/. Z>.) — Caseof ^. E. C. — Intensity of Maniacal Excitement — Sexual Perversions — Hallucinations — Etiology — Exhaustion and the Sequelae of Labour — Protracted Uterine Involution — Lactation during Profound Ansemia — Hyperlactation — Qualifications of the Nursing Mother — Period for Weaning — Prognosis — Treatment, .... Pages 409-424 INSANITY AT THE CLIMACTERIC EPOCH. Symptoms — A Subacute Delusional Melancholia — Suicidal Tendency [S. H.) — Nymphomania {A. A.) — Etiology — Incidence of Insanity at dilferent ages in 4085 cases — Influence of the Climacteric— Tlie Psychological Transformations of this Epoch — Instinctive Actions— The " Time.-eltment" in Prognosis — Alcoholism and tlie Climacteric — Treatment, . . . ' . Pages 424-437 SENILE INSANITY. Mental Derangements Incident to Senility — Senile Mania — Senile Melancholia — Chronic Cerebral Atrophy — Senile Convulsions — Senile Epilepsy — Senile CONTENTS. XVII Dementia — Inheritance as a Factor in Senile Insanities — Exhaustive Brain- work — Alcohol and Senility — Case of T. G. — Onset and Prodromata — Character of the Senile Reductions — Senile Hypochondriasis (/. .4.) — Senile Atrophy and Thrombosis (/. i?.)- Acute Senile Melancholia and .'>yiicopal Attacks {H. D. ) — Partial Exaltation in Senile Insanity — Delusional Per- versions of the Monomaniac and Senile Subject Contrasted — Senile Amnesia — Cases of Senile Insanity [M. H. and M. M.) — Elimination of Urea in Chronic Cerebral Atrophj- and Premature Senility — A Local Manifestation of Chronic Bright's Disease, ......... Pages 437-4(30 THE TREATMENT OF INSANITY. Physiological Element in Treatment — The Moral Element — The Mental Nurse — Individualised Treatment — Hospitals for the Acute Insane — Modern Revolu- tion in Treatment — Rest and Exercise — Treatment of Delusion — Destructive Habits — Suicidal Tendencies — Therapeutics — Rdle of the more important Sedatives Employed : — Chloral — Chloralamide — Paraldehyd — Sulphonal — Trional — Tetronal — Hyoscyamine — Duboisine — Opium — Cannabis Indica — Conium Pages 4G0-4S5 PART III.— PATHOLOGICAL SECTION. Morbid Condition of Cranial Bones — Investing Membranes — Brain-Substance — Histological Elements of Cortex — Forms of Tissue Degradation — Patho- logical Anatomy of General Paralysis, of Epilepsy, and of Chronic Alco- holism, ........... Pages 486-594 GENERAL PATHOLOGY AND MORBID ANATOMY. The Cranium-— Dura Mater — Pia-arachnoid — Arachnoid H;emorrhage — Adherent Pia — Vascular Apparatus — Congestion — Inflammatio7i — Softening — Atroplijr — Miliary Sclerosis — Colloid Degeneration— Granular Disintegration of Nerve- cells — Pigmentary or Fuscous Degeneration— Developmental Arrest of Nerve- cells — Vacuolation of Cell-j)iotoplasm — Vacuolation of Nucleus — Destruction of Intra-cortical Nerve-fibre plexus — Tissue Degradation from Over-strain — Tissue Degradation from Active Morbid Processes — Tissue Degradation from Disuse — General Summary, ....... Pages 486-547 PATHOLOGICAL ANATOMY OF GENERAL PARALYSIS. Tilt Brain and ils il/emhranes: — Early Implication of Vascular Tissues — Vital and Mechanical Efiects — Effects on Lymph-connective System — Intracellular Digestion — Jidle of Phagocytes, or Scavenger-cells — Character of Scavenger- element — Its Vascular Process — Fuscous Degeneration of Nerve-cells — Three Stages of Morbid Evolution : — InHainmator}' Engorgement — Implication of Pia-arachnoid— Nuclear Proliferation of Adventitia — Paralj'sis of Arterial tunics — -Diapedesis^ — Exudalitui — Htemorrhagic Transudations — Arachnoid b xvill CONTENTS. HEemorrhage — Second Stage : — Hypertrophy of Lymph Connective System — Fuscous Change and Removal of Nerve-cells — Nature of the Destructive Process — Early Implication of Apex Process — Third Stage : — Fibrillation and Atrophy. The Spinal Cord: — Spinal Cases in Four Groups — Evolution of Pseudo-tabetic and Spastic Paraplegic Forms — Angio-Neuroses — Pathogenesis of Transient Tabetic Forms — Change ^ in Vascular, Connective, and Nervous Elements — System-implication of Lateral Columns — Secondarj' to Cortical Lesions — Respects Sj'stematic Barrier — Chronic Parenchymatous Myelitis — Dependent on Gradual Degeneration of Cortical Cells — Amyotrophic Form — Degeneration of Cornual Elements in Cervical Associated with Descending Lateral Sclerosis in Dorsi-lumbar Regions — Combined System Implication of Columns — Pseudo-tabetic Forms — Ataxic Tabes — Loss of Knee-jerk— Anorexia — Flashing Pains and Sensory Symptoms — Genuine Tabetic Form in General Paralysis, Pages 548-675 PATHOLOGY OF EPILEPSY. Modern View of its Nature — An Impalpable Trophic Change — Objections to Methods of Examination — Change in Elements of the Second Cortical Layer — Fatty Change in Nuclei of Nerve-cell — Common also to Alcoholic Insanity — Vacuolation of Nucleus — Ultimate Break-down of Nerve-cell— Implication of Motor-cells— Absence of Vascular Implication— Functional Endowments of Nucleus— Resistance of Cell to Discharge — Nutritional Rhythm — Significance of Size of Cell and Nucleus — Primitive Type of Nerve- cell — Degraded Type of Nerve-cell — Cell- conformation as indicative of a Convulsive Constitution, Pages 575-581 PATHOLOGY OF CHRONIC ALCOHOLISM. Morbid Change in Cerebral Vessels— Scavenger-cells in Outer Zone of Cortex — Sclerosis of Outer Zone — Amyloid Bodies beneath Pia — Implication of M otor and Spindle-cells — Significance of these Changes — Deepest Layers more generally Involved — -Early Vascular Implication — Aneurysmal Bulgings — Atheromatous and Fatty Change — Pigmentary Degeneration of Motor-cells — Scavenger-elements in Spindle-layer — Degeneration of Medullated Nerve- fibre — Spinal Lesions — Vascularity — Hypertrophy of Tunica Muscularis an Inconstant Feature — Relationships to Chronic Bright's Disease — Sclerosis of White Columns of Cord — Spinal Degenerations in Typical Case — Implication of Clarke's Column — Immunity from Multiple Neuritis — Neiu'otic Heritage — Chronic Endarteritis — Fatty and Sclerous Tendency — The Brain of the Criminal Class — Exceptional Resemblance to General Paralysis — Coincidence of Grandiose State and Delusions of Persecution — Inconstant Vertical Impli- cation of Cord — Constitutional State that of Chi'onic Bright's Disease — Exceptional Transition to General Paralysis — Significance of Arterial Changes — Aflection of the Visceral System, Pages 581-594 DESCRIPTION OF PLATES. PLATE I. Illustrative of the "Motor Type" of Cortex. Page 62. The section taken from the extra-limbic area of rabbit's brain, near the frontal pole of lienaisphere. x 200. The three small figures represent the mesial, basal, and coronal aspect re- spectively of the rabbit's brain, showing distribution of the various types of cortex. A. Sub-frontal and sub-parietal segments of the limbic fissure. B. Gyrus hippocampi or lower limbic arc. C. Limbic fissure. D. Occipital pole. E. Frontal pole. F. Olfactory bulbs. G. Optic tract. L Olfactory root. J. Corpus callosum. K. Parietal sulcus. S. Sylvian depression. T, Limbic fissure. The types of cortex ai'e indicated bj' the siibjacent scheme. PLATE n. Page 70. Fk;. 1. Cerebellar Cortex of Young Rat. Section in direction of lamina, showing the arrangement of cells of Purkinje, and the T-shaped fibres arising fi'om the underlying graniiles. Sublimate pre- paration. X 110. Fig. 2. Section across lamina showing arborisation of dendrites of cells of Purkinje and their descending axons. Sublimate preparation. x 110. PLATE in. Page 74. Fig. 1. Cerebellar Cortex of Mouse. Showing cells of Purkinje, granules, and moss-fibres. Sublimate preparation, X no. Fig. 2. Cerebellar Cortex of Max. Showing stellate cells in neighbourliood of cells of Purkinje, distributing branches upwards to the peripheral zone; section taken across lamina. Sublimate pre- paration. X 110. PLATE IV. Page 82. Fig. 1. Cerebellar Cortex (Human). Section in direction of lamina sliows large stellate nerve-cell near cells of Purkinje; the latter are truncated; a few terminal dendrites occu])y outer half of peripheral zone, tlie liorizontal fibres being derived from the granules below. Sublimate preparation. ■- 110. XX DESCRIPTION OF PLATES. Fig. 2. Olfactory Bttlb of Rat (Semi-diageam). Arrangement of mitre-cells in relationship to their centric and peripheric dis- tribution. PLATE V. Page 92.. Fig. 1. Illustrative of the "Motor Type" of Cortex. Taken from the left hemisphere of the brain of the pig to exhibit its five- laminated type with the nests or clustered ganglionic cells. x 76. Fig. 2. "Motor Cortex" of Pig. Nerve-elements of second, third, and fourth layers respectively. x 306. PLATE VI. Page 100. Fig. 1. Illustrative of the "Sensory Type" of Cortex. Taken from the first annectant gyrus of human brain. x 65. Fig 2. ' ' Sensory Type " of Cortex. Nerve-elements of the third, fourth, fifth, and sixth layers of first annectant gyi'us of human brain. x 157. PLATE VII. Page 104. Fig. 1. Cerebral Cortex : Nerve-cells of Second Layer (Rat). Showing characteristic groupings of cells : the dendrites are notably hirsute^ and the terminal dendi'ites of a large nerve-cell from the subjacent layers pass, upwards betwixt the two groups. Sublimate preparation, x 110. Fig 2. Cornu Ammonis of Young Rat. Shows pyramidal nerve-cells of Cornu and granules of fascia dentata. Sublimate preparation. x 110. PLATE VIII. Page 108. Fig. 1. Cerebellar Cortex of Mouse. Shows granular layer between cells of Purkinje, and tlie T-shaped fibres of peripheral zone. Sublimate preparation. x 110. Fig. 2. Cerebral Cortex of Rat. Arrangement of stellate connective element beneath pia coA-ering peripheral zone ; transitional elements are also shown interblending with the terminal arborisations from nerve-cells of underlying strata. Silver-chi'ome pieparation. X 110. PLATE IX. Page 114. Fig. 1. Cerebellar Cortex (Mouse). . Shows basket-work around cells of Purkinje : axons from intrinsic cells of pei'ipheral zone are seen contributing bj' numerous descending collaterals to this ■DESCRIPTION OF PLATES. XXl basket-work, which in the specimen is much obscured by the dense deposit of sublimate. Sublimate prepaiation. x 360. Fig. 2. Scheme of Retinal Layers ix Mammals. As indicated by the more recent researches of Ramon-y-Cajal. PLATE X. Page 116. Fig. 1. " Sensory Cortex." Taken from the "modified upper limbic" ty^^e in the brain of the rabbit. This area is represented in the small figures on the i-abbit's brain on this plate by the dotted area covei'ing the inner and mesial aspect posteriorly, internal to the parietal sulcus, K. Fk;. 2. "Modified Olfactory Type." Taken from the posterior extremity of the lower limbic arc of the rabbit's brain. The area is represented in the first of the three figures, and is lettered T, The large swollen cell of the second layer is a notable feature of this cortical area. X 210. PLATE XL Page 124. Fig. 1. Cerebral Cortex (Rat). Arrangement of nerve-cells of cortex beneath "outer olfactory root." Silver- chrome preparation. x 1 10. Fig. 2. Cerebral Cortex of Rat. Nerve-cells showing extremely hirsute dendrites and descending axons. Sub- limate preparation. x 110. PLATE XIL Page 132. Section through "Motok Cortex" of Brain of Cat. Specially prepared to show relationships existing between the nerve-cells and the lymphatic channels and saccules of coi'tex. The connection of the pericellular sacs with the blood-vessels is clearly indicated, as is also the arching of the nutrient vessel around the nerve-cell. The perivascular nuclei in some cases alone indicate the position of the Ij'mphatic sheath. PLATE XIII. Page 136. Fig. 1. Spinal (iAXGLrox-CELL ok the Frcx; (Von Lenhossek). The nucleus shows a deep stained nucleolus, a faint linin network beset with chromatin granules. The filar element is faintly represented in the cytoplasm, but tlie chromophilic granules of Nissl are a notable feature, together with tlie finer granular vortex licncatli the nucleus, representing the so-called coitrosonie. Two small connective tissue cells appear to the left. This figure is l)orr()wed from Wilson's The (Jell in DfreJopmciil and Iiiheritaiire. Fjc. 2. CoR'I'KX OK ClORKBRUM (PiG, TWO 1)A>'S OM)). Sublimate preparation taken from near mesial aspect of iicmisplicrc. SubUmate prepai-ation. x 110. Xxii DESCRIPTIOX OF PLATES. PLATE XIV. Page 512, Fig. 1. DEGE>-EBATioy of Meduulated Eibees rs" Lateeal Coltthxs of Spts'Ai. Coed foemikg so-calxed "Miliae.t Sclerosis," as seex ttn^der a Low- power Objective. The pale nodular stmctures are the " miliary" patches into which varicose and moniliforni fibres are seen to pass ; most of such patches are multilocular, and are snrroimded by deep-stained sclerosed tissue. Fig. 2. -'Colloid" Patches resultixg from Degexeratiox of Medfll-^ted Fibres of Spix-^l Cord — more Highly Magnified. The multilocular constitution of the patch is mdicated by delicate outlines ; axis-cylinders devoid of medulla are seen passing into the degenerated focus ; and scavenger-ceU is seen thrusting its ramifying processes into the substance of the colloid patch. Fig. 3. "Colloid" Patch still more Highly ^vIagnified to show the Oi:TLT>"E of ^IrLTTLOCrLAE iL\TERLA^L WITH A FiXE StEOMA OF ELASTIC Fibres Arol-^'d. x 350. PLATE XV. Page 516. Degener-A-TIOX of K'erve-fibres of Lateral Columns of Spinal Cord ix so-called '-'Colloid Degeneration" of these Tracts. The axis-cylinders are seen stripped of their meduUated sheath, or surrounded by moniliform medulla undergoing granular degeneration, each with a super- imposed nucleus, which reaUy represents a young seavenger-ceU. At other parts of the field the medulla is seen in process of segmentation, and attacked bj- scavenger-cells crowded -with granular contents. PLATE XVI. Page 520. " Colloid Degeneration." Portion of inferior olivary and accessory olivary bodies in a case of glosso-labio- larjTigeal paralysis {T. W., p. 520), showing spheroidal products of degenerated medullated fibre, and the complete immunity from morbid change presented by the grey matter. x 22. PLATE XVII. Page 5-26. Fig. 1. "Colloid Degeneration." Showino- degeneration of the medullated arciform fibres of the first layer of the cortex in a case of chronic alcoholic insanity. A deep-stained sclerous belt bounds the outer zone, crowded -with degenerated products of medullated fibre forming "colloid" bodies arranged in linear series. Active scavenger-cells are seen scattered profusely throughout the morbid patch. x 350. Fig. 2. "Colloid" Transformation of Medullated Fibre foeming the Aecifoem Steipe of the Peeipheeal Zone of the Coetex in a Case of Advanced Senile Atrophy of the Beain. Numerous scavenger-cells are scattered amongst the degenerate fibres, and oil- globules crowd upon the vessels in their vicinity. x 350. DESCRIPTION OF PLATES. xxiii PLATE XVIII. Pageo8(J. To Illustrate Sclerosis of Cerebellar Cortex in a Case ok Ei'iLEPTic Imbecility. A. Healthy leaflet closeh' adjoining diseased tract, showing the cells of Puikinje iininvolved. B. The sclerosed leaflets united lirmly together, the cells of Purkinje absent, and the normal structiu'e completely altered. PLATE XIX. Page 544. To Illustrate Sclerosis of Cornu Ammonis in Epileptic Insanity. A. Peripheral zone in gyius hippocampi. B. Vacuolated cells beneath the above, PLATE XX. Page 5.V2. Vacuolation of Nuclei of Nerve-Cells. To illustrate extreme degrees of this change in the elements of the second and third layers of the cortex — human brain. The nucleus, which in the normal state should stain much deeper than the cell, remains colourless or is swollen into one large vacuole. At times sucli a vacxiole appears to have burst through the cell-protoplasm, or the latter contains numerous small vacuoles, the unaffected protoplasm still staining deeply. PLATE XXI. Page 5.56. Extreme Degrees of Vacuolation in the Multipolar Ganglionic Cells of Spinal Cord. The coincidence of granular degeneration, and vacuolation is here seen. The cells are swollen with bright translucent contents, or indurated and devoid of branches. Tlie retracted protoplasm and displaced nucleus are evident features. X 350. PLATE XXIL Page 560. Fuscous Degeneration of Large Ganglionic Cells from the Motor Cortex of Human Brain. The pale patches repiesent the areas of pigmentary degeneration, the mass being coarsely granular and of bright yellow tint. The unaffected pi'otoplasm is seen retracted and stained by the aniline dye, and the apex of one of these cells is pigmented and stunted. x 350. PLATE XXIII. Page 568. Fji:. 1. Fatty Degeneration and Atrophic Shrinking of the Pkkii'Heral Zone (1st Layer) of Cortex in a Case of Advanced Senile Atrophy of the Brain, taken from a Section of the Ascending Frontai- Con- volution. Scavenger-elements are seen profusely scattered through the upper layers, invading the vascular-tracts and surrounding the nerve-cells. The latter arc diminished in numbers, and a notable sclerous sluiiiking of tliesc superficial layers of the cortex is evident. x 130. XXIV DESCRIPTION OF PLATES. Fia. 2. Patty Disixtegkation of Cortex i>" a Case of Advanced Senile Atrophy of the Braix. The peripheral zone (first layer) is here delineated, the vasevdar walls as well as the scavenger-cells are surrounded by collections of oil-globules and fatty dtbris. X 350. Pk;. 3. Xerve-cells from the Deeper Layers of the Cortex undergoixg DlSIXTEGBATIOX AND REMOVAL BY THE AgEXCY OF SCAVENGER-CELLS WHICH st'rrouxd them. The uerve-cells are seen in difi'erent stages of degeneration — swollen, irregular, and deformed, devoid of branching processes, or reduced to a formless heap of granular debris still enclosing the nucleus. The scavenger-cells contain numerous coarse granules in their interior, deeply stained and similar to the products of nerve-disintegiation around them. x 350. PLATE XXIY. Page 584. Fig. 1. Granular DegenepvAtion of Ner^-e-cells. Taken from the fifth layer of motor cortex in a case of chronic alcoholic insanitj'. Proliferation of the perivascular nuclei as well as the pericellular elements is evident, and a coarse granular degeneration of the protoplasm of the nerve-cell has occurred. x 210. Fig, 2. Invasion of Deepest or Spindle-cell L.\yer of the Cortex by" Scavenger-cells in a Case of Chronic Alcoholic Insanity. The blood-vessels which are undergoing fatty degeneration are crowded by perivascular nuclei and surrounded by numerous scavenger-elements. x 180. Fig. 3. AxErRVSMAL Dilatation of Perivascular Sac ix' a Case of General Paraly'Sis. x 210. PLATE XXV. Page 588. Scavenger-elements in fihst layer of Cortex of Human Brain, Illus- trating TILEIR MODE OF PROLIFERATION AXD PERMEATION OF THE C0RTIC.\L Structure by' their Fibrillation. The coarse vascular processes may readily be distinguished from the finer fibrils given off bj' these organisms. x 240. PLATE XXVI. Page 592. Degexeratiox of Nerve-cells ix Humax" Cortex. Taken from a section of the motor region to illu.strate the mode of connection and relationships existing between the scavenger-elements of the lymph-connective system and the cortical blood-vessels. The swollen granidar degenerated nerve- cells are seen attacked on all hands by the scavenger-elements, x 210. DESCRIPTION OF WOODCUTS. xxv DESCRIPTION OF CHARTS. Chart A. Chart of recoveries in insanity of the climacteric. Page •2')2. ,, ,, recurrent insanity. Chart B. Chart of recoveries in insanity of pubert}- and adolescence in both sexes. Page 396. Ch.\rt C. Chart of recoveries in insanity of the puerperal and lactational j^eriods. lacre 422. DESCRIPTION OF WOODCUTS. FIG. PAGE 1. Section across transitional region of medulla, showing decussation of pyramidal tract aiid nuclei of posterior columns, . {ScJura/he), 6 2. Cross-section of medulla oblongata at the upper decussation of the pyramidal tract, ....... {Schira/haj, . 9 3. Section of medvilla oblongata through the inferior olivary bodies ( ,, ) , 11 4. Medulla oblongata and pons with neighbouring structures seen from behind ; schematic representation of the nuclei of origin of the several cranial nerves, .... {Landois ci- Stirliwj), 18 5. Section through pons on a level with the origin of the great root of the trigeminus, ........( Wernic/ce), 22 6. Scheme of segmentation of third nerve nuclei, showing presiuned anatomical and physiological relationships, ..... 27 7. Horizontal section through hemispheres, the right at a deeper level than the left, ...... {Landois d Stir/imj), 30 8. Section thi'ough hemispheres (vertical transverse), passing through plane of middle commissure, ..... {Gegenhaiir), 50 9. Cerebral cortex ; nerve-cells of second layer; higher ampliiication (rat), . 72 10. Pyramidal nerve-cell of motor cerebral cortex (hunian), ... 77 11. Cerebral cortex ; nerve-cell from deeper zone of cortex (human), . . 79 12. Cerebral cortex ; nerve-cells of second layer wth extensive lumificatiun of axons (rat), ........... 81 13. Cerebral cortex ; spindle-cells of deepest laj'er (human), ... 81 14. Cerebi-al cortex ; granules of fascia dentata — cornu Annnonis (ral)bit). 111 15. Cerebral cortex ; nerve-cells of second layer ; descending axons, . . 127 16. Left ascending frontal and parietal convolutions seen from the side, with tlie attached frontal gyri included in scheme of examination of detached cell groups, . . . . . . . . . .129 17. Left ascending frontal and parietal gyri, with the attached frontal convolutions as seen at vertex, to illustrate site of detached cell- groups, ............ l.SO 18. Reaction-time apparatus, ......... 162 19. Scheme of segmentation of third nerve nuclei, siiowing presumed anatomical and physiological relationships, ..... 308 20. Scheme illustrative of pupillary reactions, 318 XXVI DESCRIPTION OF WOODCUTS. FIG. PAGE 21. Mesial aspect of human brain, illustrative of the more frequent site of localised softenings in order of precedence, ..... 506 22. Lateral aspect of human brain (right hemisphere), illustrative of the more frequent site of localised softenings in order of precedence, . 506 23. Section across hemispheres and basal ganglia, illustrative of the site of localised softenings, in order of precedence, ..... 507 24. Lateral aspect of left hemisphere of human brain, illustrative of the sites of election of atrophy, in order of precedence, . . . .511 The Author would wish to acknowledge his indebtedness to the Cambridge Scientific Instrument Company, the Editors of the British Medical Journal, and Messrs. Macmillan for permission to utilise Figs. 18 and 20, and Plate xiii. , Fig. 1. A TREATISE MENTAL DISEASES, PART I.— ANATOMICAL AND HISTOLOGICAL SECTION. Contents.— The Spinal Cord— The Medulla Oblongata— The Mesencephalon— The Thalamencephalon — The Prosencephalon— The Encephalon as a whole — The Cerebral Cortex— Cortical Lamination. THE SPINAL CORD. The cerebro-spinal axis consists of a series of longitudinally disposed columns of white medullated nerve fibre, arranged around a central axis of grey ganglionic substance, which in its turn surrounds a central cavity or cavities. Both white and grey columns constitute symmetri- cal and bilaterally disposed halves, reminding us of the double gangli- onic cord in the invertebrata, and are connected across the middle line by a system of commissural fibres, and by certain tracts of the white columns which decussate from the one half into the other at diff"ei'ent levels of the system. We have certain points to allude to, both as regards the idea of "medullated columns" and the central "grey axis," as v/ell as the "bilateral symmetry" referred to, ere we describe in detail the structures themselves. In the first place, the central grey axis surrounding the central cavity should not be considered altogether in the light of a uniform column of grey matter, but rather as a series of ganglionic masses, which, fused together along the whole length of the spinal cord, become dissevered into separate masses in the medulla Oblong'ata, and into the much larger and more important ganglionic masses at the base of the cepebPUm. Although fused in the spinal cord, an indica- tion of the primitive ganglionic type of the invertebrata can still be traced in the enlargement of the grey substance at the level of each spinal nerve, mapping off, as it were, each spinal segment from its neighbour above and below it. In the next place, the columns of white medullated fibre are not continuoiis tracts throughout the whole cerebro-spinal axis; they consti- 1 2 THE SPINAL CORD. tute rather a multiplicity of smaller columns, each of which varies in. its destination, and consequently in its longitudinal extent. We may correctly presume that — taking as the longest course pursued by these medullated tracts that of fibres arising from the grey cortex of the cerebrum, and passing down the whole length of the cord to terminate in the motor cells for the lower extremities in the lumbar region — we have between these and the shortest every intermediate length of medullated fibre, interrupted by the ganglionic masses to which it is destined. The shortest fibres will probably be a series of fibres running as longitudinal commissures between the neighbouring spinal segments alluded to : these occur in the anterior and posterior columns of the cord. We have spoken of the fibres as being " interrupted " by the grey matter, by which we must understand the important fact, that at such points a functional connection is established between the nerve fibres and the nerve cells which abound in the grey matter at these points of interruption, and from which cells a fresh start of fibres is made into other realms. With respect to the bilateral symmetry of these two halves of the cerebro-spinal axis, it must be also stated that although at a first glance the various parts constituting the brain, medulla, and spinal cord, would appear to exactly reproduce such symmetry of arrangement, yet in the former, a lateral asymmetry is detected by a more careful consideration of its cortical envelope, the convolutionary surface of which varies very considerably in either hemisphere as regards arrangement and complexity of gyri, and superficial area of grey matter. This bilateral asymmetry apparently conforms to the extreme differentiation in structure, accompanying the more independent func- tional activity of the cerebral hemispheres, and histological research teaches us still more forcibly how infinite become the possibilities for this hemispheric differentiation. We must likewise attend to the reversal of conditions in the case of the white and grey matter constituting the large cranial ganglionic structures and the spinal cord respectively. In the latter — the spinal cord — the grey matter is central, and is invested externally by the white nerve fibre ; in the former, as the cerebral hemispheres and cere- bellum, the white medulla is central, and invested externally by an envelope of grey cortex. We need only state here that the last is the type assumed by those ganglionic levels which form the starting- point of fibres for centric destinations : the first is the type assumed for the reception of such centric diffusions. Wherever centric fibres terminate in this radiate manner, there the grey cortex assumes a sort of outer capsular investment and the medullated fasciculi its centric core. We shall find this appertain to the radiating fibres received by SUBSTANTIA GELATINOSA AND VESICULAR COLUMNS. 3 the cerebFum, the cerebellum, the thalamus, quadrig-eminal bodies, the geniculate, and the inferior olivary, in particuUir ; and, in almost all alike, we shall find even to the cerebral hemi- spheres, a zonular layer of medullated fibres bounding the grey capsular investment. To revert to the lower or subordinate levels, or the spinal axis, we learn to familiarise ourselves with the disposition and longitudinal direction of its various columns and ganglionic centres, by studying a series of sections taken in different planes and at various levels. For our present purpose, however, it is but necessary to acquaint ourselves with the appearance of its parts as seen in transverse section. In such a section carried through the lumbar enlargement of the cord, we see the irregularly crescentic masses of grey matter disposed on either side and connected across the median line by the anterior and posterior commissures, between which lies the minute orifice of the central canal. The anterior cornu or horn is at this site thick, broad, and bulbous; the posterior horn, as in other regions, is longer and narrower, directed towards the groove on the outer surface of the cord, which separates the lateral from the posterior columns, and where it receives the lateral section of the fibres of the posterior roots. Somewhat expanded at its extremity, the posterior cornu is obliquely truncated from within outwards, and capped at this site by a translucent sub- stance, the substantia gelatinosa of Rolando ; the expanded part so capped, and forming the greater part of the posterior horn, being called the caput. The connection between it and the anterior horn and median grey is called the cervix or nCCk of the horn. Whilst the lateral segment of the posterior roots passes into the caput cornu, the median-lying fasciculi arch inwards around the gelatinous substance and ascend in the outermost zones of the posterior column to enter the horn at a higher level, whilst others bend downwards into the grey tract. Just anterior to these arched fibres, where they enter the cornu, and on either inner side of the neck of the horn, is found in the upper lumbar region an insignificant cluster of nerve cells, which at higher levels become an important feature, the vesicular COlumn of Lockhart Clarke. In transverse sections the cells of this column look inflated and spherical ; they are really fusiform, as seen in longitudinal sections.* Commencing above the third lumbar nerve, this formation extends up to the ninth dorsal, and in still higher regions are found, occasionally distinctly clustered, similar cells which appear as the representatives of the same formation. Dr. Ross traces this formation as reaj)pearing in the lower end of the medulla oblongata, where he considers it to be * These cells have been spoken of as " bipolar," but this is an error, since few of tlie elements fail to show several processes. ^ THE SPINAL CORD. represented by the nucleus common to the origin of the spinal acces- sory, vagus, and glosso-pharyngeal nerves. A similar formation appears in the sacral region (origin of the second and third sacral nerves) as the sacral nucleuS Of Stilling" ; so that we have throughout the length of the spinal cord and lower end of medulla an interrupted column appearing at the two extreme ends and in the thoracic division, of which the latter is far the more conspicuous, and lies exactly along the plane of emergence of the visceral nerves (Gaskell). In the posterior horn, we find sparsely scattered cells of fusiform contour and of small size (15/x), which are regarded as sensory elements probably in connection with the posterior roots. The anterior horn presents in cervical and lumbar regions most conspicuous groups of multipolar cells, which vary much in their dis- tribution with changes in the form of the grey matter ; these groupings are fewer and far less conspicuous in the narrow anterior cornu of the dorsal region. The more important groups to be distinguished are five viz., an inner or median, an anterior, an antero-lateral, a postero- lateral, and a central. Of these clusters the first and last (median and central) are the least constant ; and, in the lumbar region in particular, do we note the absence of the inner or median groups, although even in this region minute elements tend to appear occasionally in a somewhat clustered arrangement along the mesial border of the grey matter ; at all times the inner is one of the least prominent clusters in the horn. Of the antero-lateral and postero-lateral groups, the latter is the more conspicuous, both as regards size of cluster, dimensions, and number of cells. These two groups occupy the outer margin of the grey horn, the one lying in front of the other, and usually occasioning a well-defined anterior and posterior angular projection of the horn. Internal to these, between them and the inner, and behind the anterior, lies in certain regions a central cluster, also a well-defined group. In the upper dorsal and lower cervical region a prominent lateral projection from the outer side of the grey matter betwixt anterior and posterior horn has long been known as the intepmedio-lateral tract of Clarke : it contains a cluster of cells which higher up fuse with the postero-lateral group already alluded to. The outer margin of grey is behind this site blended in a coarse meshwork with the neighbouring white medullated strands, consti- tuting the so-called formatio reticularis. The white medullated substance of the cord investing this central crrey mass is roughly distinguished into an anterior, lateral, and PYRAMIDAL TRACTS. 5 posterior column — the former extending to the outermost roots of the motor nerves : the second from this point back to the attachment of the sensory roots : and the last to the posterior median fissure. These columns are each of them further mapped out into separate tracts, indicated anatomically by a distinct groove on the surface and by a difference in the dimension of their fibres ; or by the results of emhryological research indicating their inechillated development at distinct periods of life ; or again, by the facts of the AVallerian degeneration resulting from physiological experimentation or the processes of disease. By one or other of these means we ascertain that at least eight physiological tracts may be distinguished in the white substance of the spinal cord. These may be classed as follows : — Anteriorly- — ^1. Anterior or direct pyramidal tract (also termed the column of Turcl:). . 2. — Anterior radicular (or root) zone (also termed anterior ground fibres of Flechsuj). Laterally — 3. Lateral or crossed pyramidal tract. ■4. Direct cerebellar tract. 5. Antero-lateral ascending tract of Gowers (also extending forwards). 6. Lateral limiting layer. Posteriorly — 7. Postero-extemal (or postero-lateral) column [column of Burdach : posterior radicidar zone : posterior ground fibres of Flechsig). 8. Postero-internal (or postero-median) column (column of Goll). Direct PyPamidal Tracts. — A certain proportion of the fibres of the anterior pyramids which escape decussation in the medulla, descend direct on the same side of the cord, forming the tract which more or less completely bounds the anterior median fissure. This anterior pyramidal tract progressively diminishes in size from above down- wards, as it becomes distributed to the anterior cornu of the opposite side by a continuous decussation of its fibres throughout its course along the anterior commissure. It usually extends to the mid-dorsal region, but occasionally passes down as far as the lumbar cord, and is found to vary very considerably in size for the same levels of the ■cord in different individuals, according to the more or less complete pyramidal decussation at the lower end of the medulla. There are strong reasons for regarding the fibres of this tract as chiefly destined for the upper extremities. Crossed or Lateral Pyramidal Tract.— Constituted by the larger proportion of fibres from the anterior pyramids •'' which undergo decus- sation at this high level (see fig. 1, py), the lateral tract passes down in the posterior section of the lateral columns, becoming like the anterior direct tract, progressively diminished in size to the lowest level of the * In rare exceptions even less than half the pyramidal fibres decussate at this level (Flechsig). 6 THE SPIXAL CORD. cord. The fibres of this tract pass into the grey substance of the anterior cornu between the two horns, to become connected with its motor nerve cells through the intervention of their terminal arboriza- tions. The direct and crossed pyramidal tracts represent a continuous connection with the cortex of the motor area of the brain, passing un- interruptedly in this course through the CrUSta of the cePebral peduncle and the internal capsule. r.jj.aL Fig. 1. — Section across transition region of medulla, showing decussation of pyramidal tract and nuclei of posterior columns. s.l.p. Posterior longitudinal sulcus. n.g, Xucleus funiculi gracilis. n.c, Xucleus funiculi cuueati. 5"!, Funiculus gracilis. H'^, Funiculus cuneati. c.e, Xeck of posterior liorn. g, Head of posterior horn. r.p.C.L, Posterior I'oots of first cervical nerve. X, Cell group in base of posterior cornu. cc. Central canal. S, Lateral column. a and 6, Cell clusters in anterior cornu. C.a, Anterior cornu. pii. Pyramidal tract from lateral columns decussating at d as they ascend. f-l.a, Anterior longitudinal tissure. /. r, Formatio reticularis. Posterior Median Column. — This wedge-shaped column of fine nerve fibres lying on each side of the posterior median fissure, scarcely recognisable below the dorsal region, extends from this site upwards to its termination in the clavate nucleUS in the medulla. It increases steadily in size from below upwards, and undoubtedly receives fibres in part from the sensory nerve-roots which pass into this column by way of the postero-external column as well as the posterior commissure. LATERAL COLUMNS AND TRACTS. 7 PostePO-Lateral Columns cannot, like the postero-median, be regarded as largely a continuo^is tract throughout the spinal cord. A great part of their bulk is constituted by the inner division of the posterior nerve-roots which, curving round the caput cornu, run obliquely upwards or downwards, or directly outwards, to enter the grey matter of the horn ; and also, in part, by a system of short, vertical, commissional fibres passing betwixt different levels of the grey matter throughout the whole extent of the cord. Higher up in the medulla we shall find that this column terminates in the CUneate nucleus. Direct CePebellar Tract. — A somewhat flattened fasciculus form- ing the marginal zone of the lateral columns, from the end of the dorsal region upwards, lies upon the outer side of the lateral or crossed pyramidal tract; its fibres arise from the posterior vesicular column of Clarke, which, as we have seen above, com- mences at the level of the third lumbar nerve. It gradually augments in size, and eventually terminates in the cerebellum, passing up to it along the restiform tract of the inferior peduncle. At its origin, and high in the cervical region also, the lateral pyramidal tract becomes superficial behind, so as to separate it from the posterior cornu. Antero-lateral Ascending* Tract (Gowers).— A column of fibres extending up through the whole length of the cord (occasionally the seat of ascending degenerative changes) has been described by Dr. Gowers as situated in fi'ont of the crossed pyramidal and direct cere- bellar tracts. The tract is regarded as a sensory tract originating from root fibres of the sensory nerves decussating across the posterior commissure.* Arising in the lumbar region where it lies across the lateral columns on a level with the posterior commissure, it becomes placed, higher up in the cord, more superficial. Here, ia transverse section, it forms a comma-shaped tract, its head lying betwixt the crossed pyramidal and direct cerebellar tracts, whilst its attenuated tail extends along the outer border of the cord, almost as far as the anterior median fissure. It can be traced up into the medulla in front of the direct cerebellar tract, but where the latter unites with the restiform body, its further course is not definitely ascertained. Some authorities {BecJderevj, Bruce, and Tooth) suggest its termination in the lateral nuclei ; others {Iladden and Sherrington) trace it into the restiform tract. Anterior Radicular Zone and Lateral Limiting: Layer.— These may be considered together as constituting, likf many of the fibres of the posterior radicular zone, a series of short commissural fibres uniting the grey matter at different levels. In the case of the * Diayno.-iis of Di'^nw^ti of the Spinal Cord, First Kd., 1879 ; and Diseases of the Nervoiis System, vol. i., page 12-2 ((iowers). ■ Q THE MEDULLA OBLONGATA. anterior root zone a certain portion of the fibres decussate at the anterior commissure, and thus " a connection may be established between the two anterior cornua at different levels." (Gotcers^'). We might summarise in the following short scheme the probable relationships of these tracts, as taught us by the Wallerian degenera- tions following upon disease or physiological experiment : — Short commissural vertical tracts^ (1) Anterior root zones. (2) Lateral limiting layer. (3) Burdacli's columns (in part). Descending motor from cortex cerebri — (4) Tiirck's columns. (5) Crossed pyramidal tract. Ascending sensory tracts — (6) Goll's columns from posterior roots. (7) Direct cerebellar from visceral tract. (8) Anterodateral ascending from crossed sensory roots. THE MEDULLA OBLONGATA. A transverse section taken just below the calamUS SCriptoriuS so as to reveal the central canal intact, ere it opens out on the free surface of the fourth ventricle, shows us the central grey matter thrust back to the posterior margin of the section — ^yet encroached upon laterally by the mass of the clavate and CUneate nuclei on either side. The central grey substance situated in the middle line is symmetrically disposed around the central canal, which here forms a mere elongated slit. Most prominent in front, it presents an eminence on each side of the median raphe, with a rich nucleus of large nerve cells, really disposed in double clusters — the nuclei of origin of the hypoglossal nerve, the fibres of which conspicuously run forwards towards the olivary reg'ion. On each side of these median pro- minences, a lateral projection of grey matter also occurs in front of and partially surrounding a conspicuous column of medullated fibres seen in transverse section encircled by medullated loops — the Solitary fasciculus, fasciculus rotundus or respiratory fascicle. From this point the central grey matter inclines backwards to the middle line, behind the central canal, and at an acute angle to the former. This sudden inclination backwards is necessitated by the prominence of the clavate nuclei, which, lying behind the central grey substance and to its outer side, approach each other near the mesial line ; along the backward inclination of this and the lateral prominence, nuclei for the origin of the accessory nerve and vago-accessory system are found. On each side of the central canal a column of fibres enclosing an * Loc. cit., i^. 123. eminently: teretes. . 9 elongated nucleus of nex-ve corpuscles, measuring 23 /x x 11 /x, extends forwards towards the middle line ; and here, still enclosed in the central gx'ey area, is a mesial compact cluster of small cells. The former represents the nucleus of the emineniia or fasciculus teres, which at higher levels becomes a prominent feature on the floor of the ventricle. Root fibres of the accessory nerve will at this level be traced from the lateral angle or eminence of the grey substance to their site of emergence behind the olivary body. Ip ^ ' /'S J^' n.c. Fig. 2. — Cross-section of medulla oblougata at the upper decussation of the pyramidal tract. s.l.p. Posterior longitudinal sulcus. H^, Funiculus gracilis. a.g. Nucleus of funiculus gracilis. H-, Funiculus cuoeatus. n.c. Nucleus of funiculus cuneatus. n.c^, External nucleus of funiculus cu- neatus. a.V, Ascending root of trigeminus, f/, Substantia gelatinosa. f.a, f.a^, f.a-, External arciform fibres. F.r, Formatio reticularis. n.l, Nucleus of lateral column. u^, Accessory olive. 0, Inferior olivary body. n.ar. Nucleus arciformis. P>/, Pyramid. f./.a, Anterior longitudinal fissure. d.a, Anterior or upper decussation of pyramids. n.XII, Nucleus and root fibres of hypo- glossal nerve. c.c. Central canal. n.XI, Nucleus of siiinal accessory. Above the calamus scriptorius, the opening up of the central canal on the floor of the fourth ventricle, is necessarily attended by a recession of the clavate nuclei ; and the eminentise tCPCtes, which, as we stated, formed the antero-lateral V)Oundaries of the central canal, become now exposed on the surface, as the innermost column seen in this lower half of the ventricle, on either side of the median lO THE MEDULLA OBLONGATA. raphe. Formerly covered by the ala CinePea, which represents the nucleus of the vagus, the fasciculi teretes pass upwards as white columns, strongly contrasting with the grey of the vagus nucleus ; and, as they take this course, they lie superjacent to the hypoglossal nuclei. These eminences, therefore, map out the course of the hypo- glossal nuclei, but must not be identified with that nerve, as they belong to a wholly distinct system. Whilst the white columns of the eminentife teretes become wider and more pronounced upwards, the ala cinerea disappears between them and a more external eminence — the acoustic tubercle — so that transverse sections exhibit on either side of the median raphe from within outwards the eminenti3& teretes, the tuberculum acousticum, and lastly the restiform, columns. At this plane the central grey matter is consequently unfolded outwards — a strongly marked concavity directed backwards, still characterises this region of the ventricle; but this process of unfold- ing proceeds at higher levels, until on a plane with the strlSB acOUStiCSe, the floor of the ventricle is almost flattened out, pre- senting only a gentle depression at the middle line. As this process of unfolding of the central grey matter proceeds, the restiform tract diverges more and more from the mesial line ; whilst simultaneously the central grey substance, notably in the region of the hypoglossal nucleus, becomes shallower, and the nucleus itself nearer the surface. Superficially viewed in the fresh medulla one readily sees between the diverging restiform columns a large central V, divided midway by the vertical raphe and separated from the plump acoustic tubercles on either side by a well-marked depression, into which the upper wedge-shaped apex of the ala cinerea plunges and loses itself. The anterior border of the grey substance now loses its abrupt pro- minences, and assumes a gentle sinuous course across the medulla from one solitary fasciculus to the other — the several wave-like summits representing the site of the hypog'lOSSal and the vag^O- aecessory nuclei (fig. 3). Having so far followed the disposition of the central grey matter,, from just below the opening-up of the central canal to the level of the strise medullares of the acoustic nerve, the student should now direct his attention to a cross-section of meduUated fibres of a notable crescentic configuration, and encircling on its outer side a coarsely reticulated region largely made up of deep stained connective tissue. These conspicuous structures lie laterally disposed near the margin,, on either side, in all sections of the medulla up to the emergence of the trifacial nerve : the dark-stained reticulum is the representative of the substantia g'elatinosa of the posterior cornu. The medul- lated crescent is the ascending root of the fifth nerve, representing ANTERIOR AND LATERAL COLUMNS. II the ascent of the remaining portion of the posterior root zone of tlie spinal cord. Drawing an imaginary line from the solitary fasciculus outwards to this crescent — to its anterior border in the lower levels, and to its posterior border in the higher level near the acoustic — we map off a region which corresponds to the posterior columns of the spinal cord, and their continuation as the inferior peduncles of the cerebellum : this region lies behind the imaginary line so drawn. In like manner, a line drawn from the mesial eminence of the central grey matter obliquely outwards to the root of the inferior olivary „ nX' , nX' iiJUl f/ /J J' 3 t I __.?!>^-^*^' n c oa.TTi fla ,,„,. p Section'of medulla oblougata tlirough the inferior olivary bodies. o.a.l. Outer accessory olive. Fig. 3 ■n.t. Nucleus of fasciculus teres. n.XII, Nucleus of hypoglossal nerve. n.X and n.X^^ Nuclei of vagus. f.s. Funiculus solitarius. n.g, Nucleus of funiculus gracilis. n.c, Nucleus of funiculus cuueatus. n.am. Nucleus ambiguus. C.i\ Restiform tract. g. Substantia gelatinosa. a. F, Ascending root of trigeminus (fifth nerve), i^'.r, Reticular formation of lateral column a.X, Root fibres of vagus. n.l, Nucleus of lateral columns. f.a.e, External arciform fibres, o, Inferior olivary body. XII, Emergent root fibres of hypoglossal. J), Anterior pyramid. n.ar. Nucleus arciformis. f.l.a. Anterior longitudinal fissure. p.ol. Olivary peduncle. o.a.ni, Inner accessory olive. V, Anterior column. r, Median raphi.'. body, maps off the remaining portion of medulla into two divisions, an inner, between it and the median raphe ; and an outer, between it and the former line drawn to the crescentic root of the fifth nerve. The inner of the two divisions corresponds to the antePiOP COlumil of the cord; the outer division, to the lateral COlumns. I'he two imaginary lines, so drawn, correspond to the direction taken by a 12 THE MEDULLA OBLONGATA. motor system and a mixed motor and sensory system of cranial nerves — the former line corresponding to the spinal accessory, pneumo- gastric and glosso-pharyngeal' nerves, the latter to the hypoglossal. On the other hand, the purely sensory acoustic nerve arises from a position further back than the mixed lateral system ; and above the plane of the latter nerve we find the origin of other cranial nerves disposed in like manner — viz., an antePiOP or motOP, a latePal or mixed system, and a postepiop or sensopy system. Reverting now to the posterior of these three divisions of the medulla, which, as we have stated, is bounded in front by the emergent root fibres of the mixed lateral system of nerves — we note, first, that in the lower plane (below the calamus) the greater mass of this region is constituted by the derivatives of the posterior columns of the cord and their nuclei. The columns of Goll with their clavate nuclei, and the columns of Burdach (or posterior root zones) with their cuneate nuclei, encroach by their mass upon the posterior aspect of the central grey matter, nearly meeting at the middle line ; and concealing, in this way, the deeper seated nuclei of origin of the vagus and hypoglossal. In front of the clavate and cuneate nuclei in the same tei'ritory, lies the solitary fasciculus, and the ascending root fibres of the fifth nerve around the gelatinous substance of Rolando. The crescentic root-area of the fifth nerve is covered externally by the fibres of the direct cerebellar tract from the cord ; whilst behind this tract, but still forming the outer margin of the medulla, is a narrow zonular layer, representing the commencing PestifOPm tPact of the cerebellum. At this level it will be apparent that large numbers of delicate arched fasciculi are thrown ofi* from both clavate and cuneate nuclei ; and these, passing forwards through the lateral columns of the medulla, terminate in the inferior olivary body of their own side partially, from whence fresh fasciculi start to reach the opposite resti- form tract, whilst the more posterior fasciculi cross the raphe, and traverse the opposite olivary body on their way to join the restiform tract on this side. Hence the clavate and cuneate nucleus of each side discharges itself by an extensive series of arcuate fibres into the opposite restiform tract, through the intermediation of the olivary body partially of its own side, and partially through that of the other side. As a direct result of this projection, we tind in our sections above the calamus, the rapid attenuation of these nuclei of the posterior columns, with a corresponding enlargement of the restiform tract for the cere- bellum. At these higher levels the unfolding of the central grey matter is permitted by the lateral recession of these structures, partly induced by the attenuation and disappearance of the two nucleated masses, and partly by the divergent course assumed by the resultant RESTIFORM AND INNER DIVISION OF CEREBELLAR PEDUNCLE. I 3 restiform tract to reach the cortex of the cerebellum and its dentate nucleus. Near the lower angle of the fourth ventricle, our transverse sections show us the restiform tract as a very conspicuous, somewhat pyriform area, and pale-stained in contrast to the parts behind it, from the close approximation of the meduUated fibres seen in cross-section, no7ie being arranged in fasciculi. Immediately behind this tract, however, is a much deeper-stained area, lying between it and the central grey matter and solitary fasciculus ; it is notable for the great number of small round or oval bundles by which it is constituted, measuring usually 90 /a, x 22 /a, pale-stained, and enclosed in grey matter with a meshwork of deeply-stained tissue. In this area, which covers an irregularly quadrilateral space, appear many large multipolar nerve corpuscles with large nuclei ; these corpuscles attain the dimensions of 32 /x X 20 /x. The medullated formation so constituted is an important division of the medulla to recognise. It has been long known as the inneP division of the inferior cerebellar peduncle— the restiform tract forming the OUter division of the same structure. Its connections above are with two nuclei, situated one on either side of the median line, beneath the superior vermiform process of the cerebellum, and are called, since their discovery by Stilling, the roof nuclei. Its connections below have been variously given. Stilling believed them to be the origin of the clavate and cuneate columns ; Meynert shows that this view is incorrect, and we have already seen that the latter columns are in complete connection with the restiform tract. It would appear to us that these internal divisions of the cerebellar peduncle break up into arcuate fibres, which partly pass behind, but partly traverse the hilus of the olivary body of the same side, and thence, crossing the median raphe, terminate in the grey matter of the opposite olive. Nearer the calamus we find that the clavate and cuneate nuclei, not as yet completely resolved into arcuate fasciculi insinuate themselves between these two divisions of the inferior peduncle — a little higher, where the nuclei have disappeared, these cerebellar columns are, as we have intimated, in juxtaposition. Passing now to the region lying in front of the emergent root fibres of the lateral system of nerves, between them and the motor svstem (hypoglossal), we find the greater part of this area occupied by the cross-section of ascending fibres, broken up into numerous minute groups by the intertwining of complex arcuate fibres, as they curve forwards and inwards to the raphe and the olivary district. To this fasciculated meshwork the term reticular formation has been applied, the ascending fibres being the continuation of the outermost part of the anterior root zone. Two well-defined nuclei characterise 14 THE MEDULLA OBLONGATA. this lateral column of the medulla: one, which is the richer in cells, is elongated and directed from without inwards, approaching the margin of the lateral column, lying parallel with the roots of the lateral mixed nerves, between the substantia gelatinosa behind, and the olivary and its fillet in front. Further inwards in this lateral column, and carried backwards parallel with the lateral mixed roots, is a second smaller group of cells more closely clustered than the former ; not traversed, as these are, by dense fasciculi of arcuate fibres passing to the olivary body. The former called the nucleuS of the lateral eolumn by Stilling and Clarke, might be more conveniently termed the external, and the second cluster the internal, nucleus of the lateral column •. or following Dr. Ross, the anterior and posterior nucleus. They almost certainly represent motor cell-groups of the anterior cornua of the spinal cord, severed from the rest of the central grey matter by the decussation of the pyramids across to the lateral columns, and the interposition of the mass of the inferior olivary body. By Dr. Ross they are regarded as detached from the antero- lateral and postero-lateral group of cells in the anterior cornu by the cleavage effected by the arcuate fasciculi of the medulla, whilst those main groups are still found as the motor nuclei in the central grey matter of the medulla. Intercalated between the lateral columns of the medulla and the anterior or median, is the inferior olivary hody, extending throughout the region we have been studying, but terminating at the level of the lowermost fibres of the jootzs. Prominent on the superficial aspect of the medulla, between the pyramids and the lateral and restiform tracts, it looks like a small almond-shaped body, which upon trans- verse section reveals a grey nucleus, imbedded in a medullated invest- ment of longitudinal fibres — the so-called fillet or ollvary fasClCUluS. The grey nucleus is in the form of a plicated capsule of many folds, constituted of numerous cells imbedded in grey matter, and open towards its inner side. The investing medulla of longitudinal fibres passes inwards and forms a central core for this grey capsule, the fibres of which then spread out into its various convolutionary plica- tions to terminate in the cells here distributed : the remaining fibres, which do not so turn inwards to the grey capsule, pass downwards into the lower regions of the medulla and cord. We have already alluded to the dense intertwining and connections of the cerebellar arcuate fasciculi within these olivary bodies. In the lower planes below the calamus, our sections exhibit the olivary capsule open in front at its hilus, the anterior line of plica- tions being shorter than the posterior : and hei*e, bordering upon this opening in the capsule, is an elongated belt of grey matter, containing ACCESSORY OLIVES— MIXED LATERAL SYSTEM. i 5 cells similar to those in the olivary body — this is the internal accessory olive. Sections taken midway through the inferior olive show us two such bodies ; the one, as before, situated in the anterior column, separated from the olivary body by the root fibres of the hypoglossal, and greatly segmented by the passage of a rich system of arcuate fibres to the raphe : the other, in the lateral column just behind the hilus, like a concave lens with its concavity towards the olivary capsule — this latter is the external aCCeSSOPy Olive.* In histological structure both resemble the larger olivary body, and by Meynert they are regarded as continuous with its capsule. The most external arcuate fasciculi entering the restiform tract come to it by way of the anterior pyramid and olivary body, partly in front of and partly behind (and so encircling) the former, and forming a thick stratum of fibres over the external aspect of the olivary body — its stratum ZOnale : and lastly, covering in like manner, the ascend- ing root of the fifth nerve to end in the restiform tract. A similar investment of the upper olivary body exists, as we shall see later ou ; this in the lower mammals is uncovered by the fibres of the pons, which do not conceal them, as in man, and constitute the so-called corpus trapezoides. The zonular layer passing over the anterior aspect of the anterior pyramids is aptly referred to by Meynert as a small anterior pons. This inferior half of the medulla, with which we have for the present concerned ourselves, contains the nucleus of origin and emergent root fibres of but one purely motor nerve — the hypoglossal — but of four of the mixed lateral system, viz., the spinal-accessory, vagus, glosso- pjliaryngeal, and (the ascending root of) the trifacial or fifth. At the anterior mesial prominence of grey matter in these planes, we find the nuclei of the hypoglossal, which, prior to the opening up of the central canal, are arranged in a double cluster usually termed the internal and external convolute of the hypoglossal, owing to the loop-like arrangement of the centi ic and peripheric fibres connected with them. The external also lies at a posterior plane to the internal. The cells are large and multicaudate, forming the most conspicuous cell-groupings in the whole sectional area of the medulla : they measure 60 /x x 20 /x. The centric connections of these nuclei consist of certain straight fibres of the median raphe, which run back- wards as far as the central gi"ey matter, and then arching outwards, form spirals around the front and outer border of each nucleus, and are connected with its large cells : thence, similarly curving around the inner Ijorder to pass obliquely outwards, are the peripheric Juscicrdi — the root-Jibres of the hypoglossal. These emerge from between the * These bodies ai'e also known as tlie "external and internal paiolivary bodies." I 6 THE MEDULLA OBLOXGATA. yjyramid and the olivary body, some fasciculi traversing the latter in their course. In vertical extent this centre of origin stretches from just above the level of the decussation of the pyramids to the strice medullares of the acoustic nerve; but, as we distance the calamus, the groups become less definite and merged into a less characteristic form, far less rich in, cells. Throughout the whole of this extent, the vertical column of cells gives origin to emergent radicles, which issue anteriorly. External to the hypoglossal nuclei lie the lateral projections of the central grey matter, in the angle of which we find the sensory nuclei of origin of the mixed lateral system of nerves, so named from their possessing both motor and sensory filaments. Some seven roots of origin are enumerated by Meynert for this system of nerves ; and it is probable that the three nerves of this system in the lower half of the medulla arise in a very similar, if not identical, manner from closely associated nuclei, some of which are common to two nerves. The two more important nuclei of origin for this system are — the motor nucleus of the mixed nerves, and the sensory already alluded to. The viotor nuclei are found in advance of the central grey substance, disposed in the lateral columns of the medulla. A somewhat elongated cluster of large nerve cells, from which motor fasciculi emerge and run hachwards parallel to the emergent root fibres of this system of nerves, is the more important of this mode of origin ; but, fibres running in the same direction can also be traced further outwards, to the nucleus of the lateral column, between the ascending root of the fifth and the inferior olivary body. Much discrepancy appears with respect to the descriptions given to these anterior roots of orio-in of the mixed lateral system ; some authorities speak of an anterior and posterior nucleus of the lateral column ; others describe these fibres as being doubtfully roots of the system ; whilst others with Meynert refer to one nuclear column of origin distinct from the nuclei of tlie lateral column. In fact, Meynert traces this motor nucleus as a nucleus of the spinal accessory downwards to the lateral process of the anterior horn, and finds its analogue on higher levels in the inferior facial and motor nucleus of the trigeminal.* Our own view of the case would be in accordance with that of Meynert ; in addition to which, however, we would assign to the external nucleus of the lateral column a partial site of origin for these motor rootlets. The important fact for the student to bear in mind is that these motor nuclei are, in accordance with the spinal cornual scheme, in advance of the sensory division, and reach the main roots by recurrent fasciculi curving round the vagus nucleus from the inner to the outer side ; and this type is repeated for the motor roots- of the facial and trigeminal nerves. * Psychiatry, translated by Sachs, part i., p. 124. 1 SENSORY NUCLEUS. ^7 The sensory nucleus or posterior nucleus for these three mixed nerves is a somewhat compact formation of nerve cells, clustered within the lateral angle of the central grey substance at the lower planes of this region, where it forms the vago-accessory nucleus. A little difficulty may be experienced in distinguishing between the nucleus of origin for the three nerves, if we do not attend to the fact that so long as arcuate fibres are seen distributed to the solitary funiculus from the raphe, we are in the region of origin of the spinal accessory and belovj the, vagal nucleus ; the latter centre can also be differentiated into two groups, an external and internal nucleus, de- sci'ibed by Lockhart Clarke, a similar arrangement prevailing for the glosso-pharyngeal centres. From the apex of this grey prominence, which represents a sensory column of origin for these nerves, pass outwards the main root fibres — the spinal accessory between the olivary bodies and gelatinous substance ; the vagus and glosso-pharyn- geal througli the latter, and traversing in their course the ascending roots of the fifth nerve ere they emerge at the surface. Immediately outside the origin of this root, at the posterior or sensory nuclei, is the conspicuous cross-section of the solitary faSClCUlus, which really represents an ascending" FOOt for the same mixed lateral system. We have seen that a dense arcuate system passes into it from the median raphe {centric fibres) below the origin of the vagus; we may also juSt as readily trace fibres issuing from this ascending root to join the emergent roots of the accessory, vagus, and glosso-pharyngeal nerves. The posterior sensory nucleus is regarded by Dr. Hoss as the representative of the VCSiculaP COlumnS Of Clarke in the spinal cord from their relative position, connections, character of the cells, and their distribution. A fourth root easily traced in the region of the vagus, is one which, emerging from the raphe, traverses the front of the hypoglossal nucleus, and, following the curve of the grey substance anteriorly, enters the vagus nerve. In traversing the gelatinous substance also, the vagus and glosso-pharyngeus both derive fibres from the former ere they issue fi'om the medulla ; this is the fifth root of origin for these nerves. Another small fnsciculus has been described by Clarke as passing from the faSCiCUlUS tCrCS into the vag'US. The Upper Half. — Passing now to the upper half oi the medulla, which upon its ventricular aspect is, like the lower half, triangular in outline, its base being mapped out by the acoustic strijB and its lateral boundaries formed by the superior cerebellar peduncles, con- verging to the quadrigeminal bodies, we meet first witii two motor nerves closely associated in their origin, and arising, as do the motor cranial nerves generally, from an anterior or median position on either side of the raphe ; and one purely sensory nerve, which takes THE MEDULLA OBLONGATA. its origin, in accordance with the same morphological principle alluded to, from a lateral and posterior plane. The two motor nerves are the sixth and seventh })air, or the abducens and facial ; the sensory nerve is the eighth, the aCOUStiC or auditOPy. The nuclei of origin for these three nerves do not occupy the same vertical plane ; that for the sixth is the highest, next below it comes the facial nucleus, M'xidle cerebellar tieduncle. Conariuni or pineal gland. Hrachiani coxtjunctiviuu anticiuik. Uracil ium conjunctivum posticum. PedimcaluB cerebri. ad corpora quadri-^ ^emina, or -auperior cerebellar peduncle. id medullam oblon- eratam, or inferior cerebellar peduncle. Aecflssorias nucleus Obex. Clava Funiculus cuneatus (Part of restiform bodj ). Funiculus gracilis (Posterior pyramid) Fig. 4. — Medulla oblongata and pons with neighbouring structures seen from behind : schematic representation of the nuclei of origin of the several cranial nerves. and lowest of all the acoustic nuclei — yet they each successively overlap the other, the internal acoustic nucleus, as we have already seen, descending also below the base of the arbitrary triangular space drawn by the striae medullares (see fig. 4, 6, 7, 8). As in the loiver triangular area of the grey floor below the striae, we found the nuclei ot a motor (XII) and three mixed nerves (IX, X, XI) associated OLIVARY BODIES— FACIAL NUCLEUS. IQ through a great part of their extent with the accessory body — the inferior olivary ; so in this upper triangular division we find a very similarly constituted structure — the SUperiOF Olivapy body — occupying a vertical plane corresponding very nearly to that of the two motor nerves — the sixth and seventh. Transverse sections across the levels of emergence of these latter nerves exhibit a notable change in the distribution of the various ■structures in front ; the inferior olivary bodies have disappeared ; the pyramids still maintaining their integrity as independent, comjiact ■columns, are now concealed beneath the most anterior fibres of the middle cerebellar peduncle (pons), which enclose them between their transversely disposed fasciculi, as a more superficial and a deep series of fibres; whilst laterally the toPachia of the pons diverge to the ■cerebellum, and farther back the restiform and internal divisions of the inferior cerebellar peduncles in like manner pass to their dis- tribution. On a level with the striie medullares an intermediate transitional stage is apparent ; and, as we pass to higher planes, the inferior olivaiy body loses its outward inclination, its long axis becoming disposed autero-posteriorly and immediately behind the two pyramids. Thus a lateral constriction occurs which gives the niedulla here from before backwards an apparent but not absolute increased depth. This antero-posterior depth appears still more •exaggerated by the lower loops of the pons capping the pyramids in front, which have just been caught at this plane and divided. In such sections the nucleus of the lateral column is still well seen between the diminished olivary body and the ascending root of the Jifth nerve, whilst immediately posterior to the inferior olives is a group of large fusiform and multicaudate cells, the former in connec- tion with the arcuate system here, the latter in apparent connection with fasciculi which pass backwards to the median or motOP column of grey mattep. Still somewhat higher, the inferior olive ceases, or may present its upper extremity as a single minute plication ; and, in this region, we find the nucleus of the lateral column compressed into a long narrow tract by the interposition between it and the ascending root of the Jifth nerve of a very notable lai'ge nucleus of almost spherical outline, and, by the disposition of its enclosing fibres, severed appar- ently into a series of convolutes of large multicaudate cells. This is the antepiop or infepiop nucleus of the facial nepve, and from it a somewhat wide belt of sparsely scattered fasciculi pass back to ascend, as we shall see later on, as the g^enu of the facial nerve, whilst the compressed nucleus of the lateral columns sends in- distinctly marked fibres towards the median grey. The superior olivary does not as yet present itself; in this plane we may study 20 THE MEDULLA OBLONGATA. the various nuclei of origin of the auditory nerve. Following the grey matter of the floor of the ventricle outwards from the median promi- nence (which here is remarkably shallow), we find it progressively increases in depth to its extreme lateral limits, where the lateral or sensory projection is a notable feature, and the large intemaL auditory nucleus is seen. Immediately outside this sensory nucleus is the tessellated area characterising the inner division of the inferior cerebellar peduncle ; followed still further outwards by the transverse- section of the crescent-like rentiform tract. To the inner side of the restiform tract the conspicuous ascending root of the fifth nerve is applied. The whole of the structures above noted — the grey floor with its lateral prominence, the inner peduncular tract, and tl.e restiform column — are embraced superficially by a zonular investment of fibres issuing from the region of the raphe; in fact, the strice medullar es,. which, reinforced further on by others emerging from the restiform. tract, constitute the posterior root of the auditory nerve. On the other hand, these same structures above enumerated, are- embraced from within by the anterior auditory root, which runs chiefly between the restiform tract and the ascending root of the fifth, although many of its fasciculi traverse the structure of the latter. The student should remark here that the fifth ascending- root serves always to distinguish to him the emergent roots of the facial from those of the aiiditory ; the facial lying to the inner, and the auditory to the outer side of this root. We should, therefore, regard the auditory nerve as possessing two roots of origin — (1) anterior, also called the internal, deep or vesti- bular root, whose peripheral destination is the semi-circular canals ;, and (2) the posterior, also called the external, superficial or cochlear root, which in its turn ends in the cochlea. POSteriOP Root.— -The fibres from this source enter the anterior or accessory auditory nucleus, which forms so prominent a feature as it lies upon the deep root of the auditory nerve, and, as suggested by Bruce, might much more justly be termed the auditOPy g'ang'lion. From this centre they form connections with the following- structures : — (a) Through the striae acousticEe with the fopmatio reticularis. (6) With external and internal acoustic nuclei of -same side, (c) With the flocculus. {d) With superior olive of the same and opposite sides by the medium of the trapezoid body. - Anterior Root. — The vestibular root, continuous with the so- called ascending, root of the auditory nerve, arises in part from the cuneate nucleus, in part from the large cells of the external acoustic ANTERIOR ROOT. 21 or Deiter's nucleus ; whilst a further division passes into the internal or chief nucleus of the auditory ; and some fibres are traced into Bechterew's nucleus. From the internal and external acoustic nuclei the following connections are traced : — (a) With the sixth nerve nucleus of the same side, (h) With the inferior olivary of the same side. ((•) With the fioceuhis of the same side. (d) With the ojDposite formatio reticularis. (e) With the opposite roof nuclei of the cerebellum. (/) W^ith the opposite posterior longitudinal fasciculus and thus possibly with the third nerve nucleus. Lastly, the cuneate nucleus may bring the vestibular root into ■connection with the fillet and with both restiform bodies (Bruce). At higher planes of the medulla wherein the superior olivary body appears, we reach the radicular zones of the facial and the abducens. The motor area of the grey floor oi the ventricle at these levels presents in transverse sections, two strongly defined eminences separated by the median groove and raphe — these are the eminences over the facial genu and the nucleus common to both facial and abducens nerve. The sensory area, of the grey floor flanks these eminences on either side like walls, diverging from them at a somewhat obtuse angle, the enclosed space being bridged over by the cerebellum. On either side of the median line at the extreme posterior end of the raphe, is an oval cross-section of medulla 1^ mm. by a ^- mm. in size, sharply defined and lying between the grey matter of the floor and the hindmost series of arcuate fasciculi given off' fx'om the raphe ; it represents the root of the facial nerve in cross-section at its curva- ture upwards, otherwise called the facial g'enu. From its neigh- bourhood medullated fasciculi sweep in a wide curve, following the inner margin of the grey matter as far as the sensory area, when they pass forwai'ds and outwards to their emergence from the medulla, forming in this latter course the boundary between sensory and motor divisions. In the sensory division outside this root lies, as we before indicated, the ascending" trigeminal root. The sweep of the facial in its course beneath the grey floor encloses a large and important nucleus, measuring 3 mm. in widest diameter, very rich in cells which are multicaudate, and are disposed in an almost circular area ; from the outer side of this nucleus emerge root fasciculi, which are distinctly seen to join the facial as it sweeps forwards round the nucleus to its point of emergence. On the other hand, from the posterior and inner margin of this nucleus other fibres emerge, which upon the inner side strike forwards, becoming gradually more divergent from the raphe in sharply defined fasciculi, to leave at the lowest border of the pons as the abdUCens 22 THE MEDULLA OBLONGATA. nerve or sixth pair. The nucleus itself is the abducens-facialiSy also termed the posterior or superior facial nucleus, or again the nucleus of the sixth nerve. It is placed in communication with the cerebrum by means of the fibrse rectse of the raphe, which can be readily traced as the most posterior of the arcuate fibres curving first around the facial genu in front, and then passing round the lower hemisphere of the nucleus. Considerably in front of the abducens- facialis nucleus, and in the motor division of this region, lies the inferior or anterior facial nucleus, almost parallel with the trigeminal root, but separated from it by the facial emergent fasciculi ; its fibres pass backwards, as we have already seen, at lower levels to G.f. Fig. 5. -Section through pons on a level with the origin of the great root of the trigeminus. G.j, Gem; of facial nerve. VII, Root fibres of facial. Vm, Motor nucleus of trigemiuus. Va, Ascending root of trigeminus. i?, Restiform tract. M.P, Fibres of pons varolii. •p.l.h. Posterior longitudinal fasciculus. F.B, Reticular formation, s.o, Superior olivary body. F, Lemniscus or fillet. Fy, Pyramids. s.t.f,d.t.f. Superficial and deep transverse fibres of the pons. arch beneath the abducens-facialis to its inner and posterior aspect,. and thence running upwards as the gemi of the facial, again bend around its upper border in the graceful sweep of the emergent roots. The Superior Olivary Body. — We have seen that this body extends from the lowest border of the 2yons through the whole tract of origin of the facial nerve, being well-exposed in cross-sections, lying between the inferior facial nucleus and the emergent root fibres of the sixth nerve. The transversely disposed fasciculi lying upon its anterior surface, extending from the decussation at the raphe to ascend in the inferior cerebellar peduncle, form the so-called COrpus trapC— ZOideS which becomes exposed superficially in animals where the- UPPER OLIVARY BODY. 23 dimensions of the pons are greatly reduced with the diminished supply of fibres i-eaching the medulla from the crusta. The lemniscus or fillet lies in these planes to the inner side of the superior olive/ forming the pale-stained area of truncated triangular outline next the raphe, the base traversed by the most posterior fasciculi of the pons and trapezoid formation. Into the fillet at higher planes, fibres of the upper olive pass to be connected with the central grey of the lower quadrigeminal body, the testes ; functionally these fibres should be regarded as centrifugal, since they have been found by Flechsig to degenerate doivnwards to the superior olive. A cerebellar connection is established between these bodies and the roof nuclei (nuclei tecti) of the middle lobe of the cerebellum ; whilst other fibres pass back from them also to the nucleus of the sixth and of the auditory nerve, as well as to the lateral columns of the spinal cord. Motor impulses, therefore, emanate from this body to the sixth nerve nucleus, which, being connected by decussating fibres with the nucleus of the opposite motOP OCUli, subserve the conjugate deviating movements of the eyeballs. In like manner, motor impulses to the lateral columns of the cord explain the associated movements of the head to the same side. The quadrigeminal bodies, on the other hand, which are connected with the optic tracts, transmit stimuli thence emanating, to the superior olivary bodies through the medium of the fillet, and so to the oculo-motor apparatus of the sixth and third nerves. We have already seen that the posterior columns of the cord resolve themselves through the intermediation of their clavate and cuneate nuclei into tlie restiform tract of the cerebellar peduncle. They also by the anterior sensory decussation of a portion of their arcuate fasciculi pass upwards on either side of the median raphe as the fillet, and thence to the quadrigeminal bodies. This portion of the fillet, it will be observed, is a centripetal or sensory tract, so that the fillet really contains systems of ascending and descending fibres, as is indicated also by the results of lesions aS'ecting the tract. At the level of origin of the sixth and seventh cranial nerves, the central grey forming the floor of the ventricle is, as we have seen, extended laterally, shallow from before backwards, and bounded on either side by the restiform tracts : as we ascend to a higher plane we find the superior cerebellar peduncle on each side, which, in their descent, restrict the lateral extension of the ventricle and its investing grey substance. This occurs in such sections as are carried through the emergent roots of the fifth nerve. If we now follow the ventricle towards its upper angle, we find with the convergence of the superior peduncles towards the quadrigeminal bodies, the following changes in the disposition of the central grey : — First, the ventricle becomes 24 THE MEDULLA OBLONGATA. narrower ; the prominent lips of the grey matter become more pronounced, from the increase in the thickness of this formation; and at the same time the ventricle is roofed over posteriorly by the antePiOP medullary velum. The nuclei of the fifth pair alone of all the cranial nerves characterise this plane. Higher still, we come upon the root fibres of the fourth nerve which decussate across the aqueduct posteriorly. The central grey here forms two notable protruding lips on either side of the mesial line, converting the aqueduct into a Y-shaped figure. Progres- sive thickening of the central grey substance occurs as we carry our sections through the posterior and anterior quadrigeminal bodies ; the aqueduct restricted in size is completely surrounded thereby, and suffers minor alterations in its outline until it opens up into the central cavity of the third ventricle. From the level of the crossing of the fourth nerve upwards, the Y-shaped grey exhibits the heak of the Y interposed between the two notable bundles of the posterior long-itudinal fasciculus, behind which we may con- tinuously follow an anterior or motor column of grey matter, containing nerve cells, and externally a lateral ot sensory column, such as charac- terised the cranial nerve origins in the lower half of the medulla. Betwixt these planes and the iipper roots of the facial nerve lie the emergent fasciculi of the fifth nerve, which, in accordance with its mixed motor and sensorial function, also assumes a lateral site of origin. We will now take these upper cranial nerves seriatim. The fifth or trigeminal has the nucleus of oi-igin for its motor root within the motor area of the pons somewhat similar in position to the nuclei of the latei'al division, which at lower levels sent fasciculi to the mixed system of nerves — the so-called nucleus ambig'UUS. It, however, lies considerably behind this nucleus of the facial — the anterior facial nucleus- — and cannot be mistaken for it, since it does not present the same convoluted structure ; and, moreover, is not in the mid planes of section of the upper olive, although the up[)er end of this structure is still seen. Its centric fibres pass to it from the median raphe where they decussate. The origin of the sensory root of this nerve is far more extensive. We have thi'oughout the whole of the medulla followed up in our sections the aSCCnding" root of the trig'eminal, noting how in the lower planes of the medulla the vagus and glosso-pharyngeal, and higher up, the facial, traversed its cross-section near their points of emergence, and now find it lying between the motor nucleus and the restiform tract in the sensory area of the pons, throwing forwards its root fibres to emerge between the transverse fasciculi of the pons. Below, this ascending root appears to end in the tubercle of Rolando, and so would seem to have a close connection with the caput COrnu NUCLEI OF THE OCULO-MOTOR AND TR0CHLEARI8. -0 pOSteriOPiS. A median root is described as originating from a nucleus ;it the level of emergence of the sensory root, in contiguity with tiie ascending root and the motor nucleus. On the lateral margin of the central grev around the aqueduct, as high as the anterior quadrigeminal body, or nateS, we find the cross-section of the descending root of the fifth nerve with very characteristic spherical or vesicular cells lying in the central grey upon the inner side of the root fasciculi. Both cells and descending lasciculi become more conspicuous at lower levels, and the latter extend to the level of the exit of the sensory root, where they join it to emerge from the pons. Internal to this descending root is a series of deeply pigmented nerve cells, iorming the substantia ferruginea, which is seen through the grey floor of the ventricle at the site named the loCUS COBPUleuS. From these cells, according to Meynert, pass root fibres to the opposite trigeminal root (sensory trunk), which in their course surround and traverse the posterior longitudinal faSCiCUlUS, decussate at the posterior extremity of the raphe, and thence, following out the anterior margin of the central grey, arch into the opposite sensory root. Associated with these latter fibres are described others which issue from the median raphe posteriorly, and after decussation terminate in the sensory root likewise. Lastly, a Cerebellar rOOt is described by some authorities. In the motor column of the central grey, lying immediately behind the posterior longitudinal fasciculus, on either side of the median line and beneath the nates, is a well-defined grouping of cells, which, however, usually presents an apparent segmentation into distinct clusters. These nerve cells, commencing as high up as the posterior commissure, are at first somewhat scattered, but assume a more compact form as they run backwards towards the upper half of the testes, in which region they appear lodged in a hollow of the ])Osterior wall of the longitudinal fasciculus. This nuclear column represents the origin common to the OCUlO-motor (third) and the trOChleariS (fourth) nerves; the upper, scattered, segmented portion is the nucleus of the third more especially ; the lower compact segment lying at the junction of the two quadrigeminal bodies is the nucleus of the /orirth nerve. Both are believed to receive their centric fibres through tli'e median raphe, those of the third nerve decussating ere they reach their nucleus. Tlie segmentation of the nucleus of the motor oculi nerve has been variously described by diff'erent authorities : we follow here more strictly the account given by Bruce,* which certainly accords most *"()ii the Segmentation of the Nucleus of the Third Cranial Nerve," Alex. Bruce, Pror. Roy. Soc. Ediii., 18S9. Ill n.st rat ions of I In- X! %J On the other hand, the outer division of the striate body — the lenticular nucleus — lias its base directed forwards and outwards parallel with the insula, whilst its wedge-shaped apex is directed backwards and inwards towards the base — its upper surface, convex from before backwards, being adapted to the concavity of the internal capsule. The general relationships of the internal capsule are well seen in horizontal sections carried across the hemisphere at succes- sively higher levels. Near the base it forms a quadrilateral section directed from within outwards and separating the tegmentum from the lenticular nucleus, having immediately behind it the loCUS nig'eP, and parallel with it in front the antePiOP COmmiSSUre as it passes outwards through the lenticular nucleus. It will be recalled that the crust, ere it disappeared into the depths of the hemisphere at the base, was bounded by the broad optic tracts : these are destined to terminate in the external g'eniculate body, and at a slightly higher plane these bodies appear immediately behind the extremity of the internal capsule as seen in transverse section. At a higher level, however, above the anterior, and through the ])lane of the middle commissure, the antei-ior portion is bent outwards, forming an obtuse angle or " knee," so that a horizontal section through both hemispheres at this level represents both internal capsules as an X-shaped figure with two anterior and two posterior segments (fig. 7). In the lateral angle of this X the wedge of the lenticular nucleus insinuates itself; between the anterior segments of the X the nuclei caudati appear ; whilst the posterior segments include the optic thalami. At the central junction of these limbs, we find the foi-nix cut across. Lastly, turning our attention again to the base of the brain, we see that the divergence of the peduncles, as they ascend, leaves between them in the middle line, first, the posterior perforated Space — a bridge-like extension of grey substance at the angle between both peduncleSj perforated by numerous vessels which enter the base of the thalamus at this point ; and from which white fibres emerge and course round the upper margin of the pons — the taenia pontis. In front of the perforated space are two rounded bodies — the corpora albicantia — round which the pillars of the fornix turn ; and from these bodies a thin grey lamella stretches forwards as far as the optic commissure, forming the floor of the third ventricle : from its floor a hollow tube — the infundibulum — descends, to which is attached the pituitary body. Having thus far defined the limits of crusta and internal capsule, we can the more clearly appreciate the course and distribution of their fibres. Since the crusta represents the continuation upwards of the pyramidal tract of the cord, it must receive a considerable rein- 32 THE MESENCEPHALON. forcement of fibres within the pons to account for the much larger size which the crusta bears to the corresponding anterior pyramid of the medulLi. An examination of the pyramidal tracts and crusta in a- nine months' embryo giv^es us most valuable information respecting the origin of the fibres found therein. In the pyi'amids the outer portion of the tract consists of distinctly medullated fibres ; the inner and anterior of non-medullated fibres ; and an intervening portion contains a mixture of both. In the pons a similar distribution of these fibres is seen : whilst higher up still in the crusta, the non- medullated fibres occupy the inner third ; the medullated fibres, the middle third ; whilst the admixture of both is found behind and between the two former. This embryological dissection maps out three- distinct systems of fibres which have received the respective names of {a) the fundamental or medullated ; {b) the mixed system ; (c) the accessory or non-medullated system. Still further outwards, occupying the outer fourth of the crusta,. just beyond the fundamental system, are found the fibres which represent the continuation upwards to the hemispheres of the sensory columns of the cord. If we now trace these tracts upwards into the internal capsule, we find that the accessory or innermost passes up along the anterior segment of the capsule ; the mixed occupies in its ascent the anterior third of the posterior segment, the fundamental the middle third of the same segment ; whilst the posterior third or outermost part of the capsule is occupied by the sensory tract of the peduncles: or tabulated also with reference to their distribution, thus : — Tracts. Accessory Tract, Position in CiirsTA. Inner third, Ix Internal Capsule. Anterior segment, Cortical Termini. Mixed Tract, Fundamental Tract, Sensory Tract, Third frontal gjrus posteriorly. As- cending frontal and parietal gj-ri at their lower end. Behind and between Posterior segment. Second frontal and Accessory and ,.,„ Anterior third, middle portion of central gj'ri. Posterior segment, First frontal pos- Middle third, teriorl3\ Upper end of centi-al gyri ; paracentral andi^arietal g^Ti. Do., do., Temporo - occipital Posterior third. regions. The aboA'c table reads as follows : — The accessor}'- tract occupies the inner third of the crusta, passes up the anterior segment of internal capsule, and terminates in the cortex of the third frontal and lower end of ascending frontal and parietal gyri — and so for the remaining tracts. Fundamental, Middle third, Outer fourth, CONDUCTING TRACTS OF THE CRUSTA. 33 The student must bear in mind that the crust of the peduncle contains a very large system of fibres, and that the tracts above described by no means represent the whole series. Thus each crusta includes, not alone tracts of the pyramidal fibres, but the sensory tracts of the cords — fibres to the thalamus and caudate nucleus, ifec. The tracts we have now traced have 7io connection with the basal ganglia : they pass between the lenticular nucleus, thalamus, and caudate nucleus uninterruptedly to their cortical termini. Thus, as we have seen, the pyramidal tract extends uninterruptedly between the cortex of the central g'yPi and their immediate neighbourhood, and the motor cells of the anterior eornua of the cord at different levels throughout its course : and this pertains both to the motor or pyramidal and to the sensory tracts. It is obvious that the body and lower extremities have the least specialised movements, just as the arm and hand and the muscles of articulate speech are excessively specialised ; and, that whereas the accessory group of fibres which supply the latter chiefly have to deal with very complex and specialised co-ordinations, the muscles being very numerous, but relatively very small in size : the fundamental group which supply the large muscles of the limbs, and especially of the lower extremity, have to deal with simple massive movements, the musculature being correspondingly bulky and less specialised. We see, therefore, that the accessory group, arising from the cortex at the lower end of the central gyri, pass chiefly to the spinal levels of the face, mouth, and hands, whilst the greater bulk of the fundamental group arising at the upper end of the central gyri extend as far as the lumbar enlargement for the musculature of the legs. Hence the former constitute comparatively short loops, and supply many though small muscles ; the latter constitute very lengthy loops, and supply the largest muscles of the frame. The latter originate in the largest nerve cells discoverable, the former or accessory in cells of greatly reduced dimensions. The fibres of the sensory columns of the cord, according to Meynert, undergo a decussa- tion upon a level with the decussation of the crossed pyramidal fibres by arching forwards around the central grey column, and after decus- sation passing up on the outer side of the anterior pyramid to the hinder third of the internal capsule, when they suddenly turn back, and are distributed to the OCCipital and temporO-Sphenoidal lobes. As before stated, they have no connection with the thalamus or lenticular nucleus. So far for the direct sensory and motor fasciculi of the crusta and internal capsule. The internal capsule, however, includes several other systems of fibres, of which the following are the more readily followed out : — (a) Fibres to the cortex from the outer surface of the optic thalamus. 3 34 I'HE MESENCEPHALON. (b) Fibres to the cortex from the outer surface of the caudate nucleus. (c) Fibres to the cortex from the upper and inner surface of lenticular nucleus. (Denied by Wernicke.) (a) Thalamic radial fibres given off from the whole length of the outer surface of the thalamus radiate forwards, outwards, and back- wards ; the anterior radiations towards the frontal lobe ; the median, outwards to the parietal ; and the posterior, arching backwards and upwards to the occipital lobe. The latter or posterior is a very important formation, and is separately distinguished as the radia- tions of Gratiolet : it serves as the means whereby the several roots of the optic nerve, which terminate in the quadrigeminal, geniculate bodies, and thalamus are connected with the OCCipital cortex. These optic radiations pass through the posterior third of the internal capsule, and are consequently brought in close relation- ship to the sensory peduncular tract already described as occupying this position. (6) In like manner, fibres radiate from the outer surface of the caudate nucleus to reach the cortex in planes internal to and above those of the thalamus. (c) Fibres arise from the upper and inner surface of the lenticular nucleus to interlace with the fibres of the internal capsule. (d) A system of fibres has been assumed by Flechsig to pass con- tinuously through the red nucleus of the tegmentum and thalamus into the internal capsule, to be distributed along with the fibres of the pyramidal tract to the region of the RolandiC fiSSUre. By this means a crossed connection would be established between the central g-yri and the cerebellum. (e) Fibres from the Olfactory bulb, after decussation in the anterior commissure, are also believed by Meynert to join the optic radiations of Gratiolet, to be distributed to the occipital and temporo- sphenoidal cortex. We have seen how those peduncular fibres forming the crusta diverge and enter the hemisphere as the internal capsule, supporting upon its upper surface the caudate nucleus and thalamus, and receiv- ing, into its concavity below, the wedge-shaped mass of the lenticular nucleus, whose apex contributes a mass of fibres to the formation of the crusta. If we now imagine a lamella of fibres spread over the broad base or outer aspect of this wedge to pass downwards beneath the lenticular nucleus towards its apex, and there meeting the crusta, whose fibres it crosses transversely to its inner side, we have an arrangement which, from the insula and temporal regions of the brain, passes to the base, as a sort of sling-like loop supporting these ganglionic masses, and binding them and the peduncles together. ANSA PEDUNCULARIS. 35 Such an arrangement is represented by the external capsule and its continuation at the base, where it is termed the ansa pedunculapis •of Gratiolet or substantia innominata of Reil ; and a portion of ■which crossing the crusta to its inner side is named the COUaP, fillet, or loop of the CPUS. This important formation of the ansa peduncularis consists of four systems of fibres, according to the statement of Meynert. They 4irise from the under surface of the lenticular nucleus, from a ganglion lying in this position, and from the cortex of the sylvian fissure, upper and temporal margin of the insula. Tlie capsular portion of the ansa has no organic connection with the base of the lenticular wedge over which it spreads, so that it can be readily separated, .and the latter enucleated by the handle of the scalpel. Its fibres, which arise from the cortex of the insula and upper margin of the sylvian fissure, necessarily pass to it through the claustpal formation ; and then, forming the compact lamella of the external •capsule, converge to the fasciculi at the base of the lenticular nucleus, which we referred to as the ansa. At this point it is crossed superficially by the anterior commissure, which has to be removed to •expose it in its entirety. The deepest layer of the ansa peduncularis takes its origin from fibres issuing at the base of the lenticular nucleus — from those concentric lamellse, the laminSB medullaPes. The fasciculus completely crosses the crusta pai'allel to, and immediately in front of, the optic tracts, and passing to the inner or median border of the crusta, forms the innermost series of fibres, here destined to pass back along the paphe to the central grey substance, where they terminate after decussation in the nucleus Of OPigfin of the oculo-motor and trochlear nerves within the nates. The second layer originating in fibres from the cortex of the upper margin of the sylvian fissure, the temporal lobe and the cortex of the insula, is interrupted in an elongated ganglion at the base of the lenticular nucleus. From this ganglion fresh fibres proceed at first parallel to the course of the rest of the ansa, but then suddenly bending back- wards and upwards, pursue their further course just within the grey substance of the third ventricle, forming the well-marked bundle of medullated fibres known as the pOStePiOP long-itudinal fasciculus, which may possibly be traced down into the spinal cord as the most posterior of the fibres of the anterior column in front of the grey commissure of the cord. The third layer of the ansa arises from the sylvian foSSa :nid the cortex of the temporal lobe, runs parallel to the above-mentioned fasciculi at the base of the lenticular nucleus ; and then turning upwards into the thalamus, forms a brush-like radiation of its fibres far back into the interior, constituting the so-called infePiOP peduncle 3^6 THE MESENCEPHALON. of the thalamus. A fourth layer of the ansa which overlies the latter joins the Stratum ZOnale, or capsular investment of the thalamus. This substantial belt of the ansa firmly binds together the region of the Operculum and island to the central structures at the base of the brain, forming a complete sling around the lenticular ■wedge, consisting of a series of loops, the deepest of which connects- the base of the lenticular body to the motor nuclei in front of the aqueduct ; the others having a more lengthened course from the cortex to the anterior column of the spinal cord, or blending in- timately with the structures of the optic thalamus and its capsular investment. Dissection. — Place the brain with the base uppermost, ha-\"ing carefuU}' removed the membranes and the large vessels. Remove ■with the blade a shallo'w horizontal slice from one of the temporo-sphenoidal lobes so as to pass throiigh the hook of the uncinate gyrus ; this lays bare the section of the pes hippo- eampi, and the amygdaloid nucleus. Pass the handle of the scalpel vertically through the medullary strands outside the pes — it enters the extremity of the descending horn of the lateral ventricle. Carry an incision along the floor in a somewhat curved direction, backwards and outwards, along the whole length of the occipito-temporal g}Ti. This exposes the descending comu throughout its extent, and the worm-like COPnu ammonis is seen descending from above to terminate at the pes. The tails of the caudate and lenticulaP nuclei are now seen along the roof of the cornu, and a white, glistening, narrow band of fibres curves do^svnwards towards the amygdala— the taenia semicipculapis. Upon raising up the inner border of the uncinate gjTus, we expose the optic tracts- arching backwards over the crusta, and terminating in the two elevations of the corpora geniculata, externa and interna, beyond which the pulvinar of the thalamus projects. In front of the optic tracts the crusta has dis- appeared in the depths of the hemisphere, and we see here the triangular floor of the anterior perforated space bounded by the delicate white roots of the OlfaetOPy nerves. Dra^sving the optic ner^^es backwards we expose the deUcate, almost translucent grey lamina forming the floor of the fifth ventricle, bounded by the two white peduncles of the COPpuS CallOSUm, which at this juncture probably tears across, revealing a broad white fasciculus, the antePiOP COmmiS- SUPe, crossing from one to the other hemisphere. Upon gently drawing apart the divergent crura cerebri in fronts a dark line is apparent, running the whole length of its mesial aspect at a greater depth than the innermost fasciculi of the crusta. This, upon dividing the crura, is seen to be the inner margin of a dark pigmented body, of lenticular section, the substantia nigra of Soemmering, which forms a distinct boundary wall betwixt the crusta in front and tlie tegmentum behind : it extends from the level of the corpora albicantia as far as the upper border of the pons. We shall refer more minutely to its relationships when studying transverse sections of this region. Immediately behind lie the tegmentum and the large ganglia of the mid-brain or quadri- geminal bodies. LEMNISCUS OR FILLET AND AQUEDUCT. 37 We can best study this posterior division of the mesencephalon by first examining the external configuration and connections of these ganglia, and subsequently following the course of their fibres, and next by studying the relative position of the fasciculi as seen in transverse section at different levels. If now we so separate the occipital lobes of the hemispheres as to expose these bodies to view from above, they appear in a mid position between the two -divergent cushions of the thalamus, overriding the cerebral peduncles in tlie form of two pairs of tubercles — ^an upper and a lower, each oval in form, the upper pair the larger and darker in colour, both having their long axis disposed transversely. Both pairs of tubercles extend upwards and forwards an arm, which is really a connection for the cerebral cortex, and spreads upwards as COPOnal radiations : each pair in like manner sends downwards a meduUated lamella on either side of the crus cerebri — that from the nates being deeper-placed and over-laid by that from the testes ; this downward extension is called the fillet of the quadrigerainal bodies, or the lemniscus. We shall trace the arms or brachia and the lemniscus at a subsequent stage of our examination. Between the nates, at the upper end of their median raphe, lies the COnarium or pineal body (fig. 4), with its long delicate brachia extending forwards, and bounding the white ventricular surface of the thalamus above, from the median grey walls of the third ventricle. Beneath and in front of the pineal body is the posterior commissure, under which we observe the opening of the aqueductUS Sylvii into the third ventricle. Hence the <^uadrigeminal bodies lie immediately over the continuation of the central gi'ey substance of the ventricles surrounding the aqueduct, which would be, therefore, laid open by a vertical incision carried through the median raphe of the quadrigeminal bodies. Moreover, the descending ribands called the upper and lower lemniscus are, in their descent, closely approximated to the outer wall of the central grey substance, which is, therefore, as it were, very largely embraced by the quadrigeminal system. Just external to the central grey substance are tlie various structures entering into the formation of the tegmentum, extending as far forwards as the substantia nigra. If the peduncular fibres of the crusta exposed on one side of the pons, upon removal of the superficial layers of the middle peduncle, be divided, raised, and tlie deei)er transverse fibres be dealt with in like manner, we come down uiion the most anterior layers of the tegmentum, and these, wlien traced, are found to be the fibres of the lemniscus, which winds round from behind, and insinuating itself beneath the overlapping crusta, passes in a compact belt extending even as far as the median 35 THE MESENCEPHALON. raphe of the pons (fig. 5, F). In its further course it is traced downwards as the most external zone of the lateral columns of the spinal cord in front of the direct cerebellap column. Whilst the lemniscus in this part of its course gradually encloses the several other tegmental structures, including the red nucleus, the fibres given off from the latter after decussating at the median raphe pass to the outer side of the tegment in its further course, and eventually escape from the embrace of the lemniscus about the level of the upper trans- verse fibres of the pons, and, becoming superficial, pass downwards to the cerebellum as its superior peduncles. We must now follow the fasciculi of the mesencephalon in transverse sections taken across the upper and lower pairs of quadrigeminal bodies. In such sections passing through the region of the nateS, five structures which demand examinution are exposed to view ; these are in order from behind forwards — (1) The nates; (2) the central grey substance ; (3) the tegmentum ; (4) the substantia nigra ; (5) the crusta. (1) The Nates (Superior Bigeminal Body). — The ganglionic struc- ture presents four distinct strata — {a) Outer grey or cortical layer of Forel. {h) Longitudinal fasciculi from the external geniculate body, (c) Bundles of fasciculi passing outwards on all sides from the central grey substance in delicate radiations into the substance of the ganglion. {d) The stratum of the lemniscus, embi*acing the grey substance, and through which the radiating fibres last named have to pass outwards. To appreciate correctly the structure of these ganglia, we must imagine a coronal fan of fibres from the cei'ebral cortex converging in the direction of the external geniculate bodies into the substance of the cortical layer of the nates ; passing back to the median raphe behind the aqueduct, and decussating here with a similar fasciculus from the opposite hemisphere ; thence arching around the central grey as the lemniscus already described. The coronal fasciculus constitutes tlie superior brachium of the nates ; and, j)'>"''Or to the decussation, it terminates in the ganglionic cells of the nates. From these cells arise the fibres which decussate, and pass as the lemniscus downwards into the spinal cord. We have, therefore, the two ganglia of the nates,, so to speak, enclosed between the upper and lower arms of a decus- sating medulla with an internode of ganglionic cells on the centric side of the decussation ; and the lower arms or Stratum lemnisci enclosing in like manner the central grey, and supporting as a girdle the mass of tegmentum lying in front of the latter. The upper arm or brachium passes in part direct to the cortex beneath the pulvinar of the thalamus, and thence through the posterior division of the internal capsule ; and in part, passes into the external geniculate body behind and covering the former. This decussating system of RED NUCLEUS OF TEGMENTUM. 39 medulla extends downwards, presenting a similar formation for the testes as for the upper bigeminal body (nates). Here also we have an inferior brachium of the testes, and an inferior lemniscus given off to the medulla and cord. As we shall see later on, an almost exact counterpart of this medullary system carried forwards explaiiis the formation of the posterior commissure of the third ventricle. Both geniculate bodies receive coronal radiations, both transmit fibres through the brachia to the corresponding quadrigeminal body, and the external geniculate is intimately connected with the optic tract as the latter passes to the nates by the medium of the upper brachium ; whilst the inferior brachium and hence the testes have likewise a connection with the inner side of the tract. Hence the nates and testes are brought into relationship with the cortex of the occipital and temporal lobes, as well as with the retina. The radiating fibres spreading from the central grey substance in the nates through the lemniscus, are probably direct connections between tlie nerve cells of the nates and the nuclei of the oculo-motor nerves within the central grey area. In the region of the nates also we observe in cross-sections numerous fasciculi lying between the antero-lateral margin of the grey substance and the lemniscus ; they pass inwards between the red nucleus and the posterior longitudinal fasciculus and decussate at the raphe — their destination being obscure. By Meynert they are supposed to be connected with the nucleus of the descending root of the fifth nerve. Red Nucleus of Tegmentum and Upper Cerebellar Ped- uncle. — When describing the structure of the thalamus we shall find, as an important constituent, a rounded nuclear mass named the red nucleus, which, upon the one hand, receives coronal radiations, and, on the other hand, gives off medullary fasciculi extending downwards through the quadrigeminal region, where they decussate at the median raphe to terminate as the superior peduncles of the cerebellum. The nucleus itself, consisting of much grey matter enclosing large and small nerve cells, is continued into the region of the nates, below which its grey matter disappears, and white medullary fasciculi with interspersed nerve cells are alone continued downwards to the points of decussation. In the upper sectional planes (transverse) of the mesencephalon, we see this red nucleus of almost rounded contour embraced between the substantia nigra and commencing lemniscus (in front and laterally) and the other fasciculi of the tegment behind. At this level also the arched roots of origin of the oculo-motor nerve lie on its inner side, and partly travex'se its structure. In the lower planes of the mid-brain, through the testes, the medullary fasciculi derived from the red nucleus, now called the 40 THE MESENCEPHALON. SUperiOF eePebellaP peduncles, approach the median raphe and decussate completely with the fasciculi of the opposite side. Through- out this decussation the fasciculi are embraced between the loop of the lemniscus in front, and the posterior longitudinal fasciculi behind. From the line of decussation the fasciculi now arch outwards and backwards, and, still covered by the lowest fibres of the lemniscus derived from the testes, emerge opposite the greatest convexity of the pons to' enter the cerebellum u|)on the same side. If we adopt Meynert's view of the projection System, the nucleus ruber forms an internode or point of interruption between the coronal fibres of the cerebral hemisphere and the superior peduncular fibres of the opposite cerebellar hemisphere ; and, as with the first link of the projection system generally, the internode occurs on the same side as the hemispheres supplying the coronal attachments. Posterior Longitudinal Fasciculus.— A compact column of lai-ge nerve fibres, oval, somewhat pyriform or lenticular, according to the plane of section, presents itself immediately in front of the central grey area, and, therefore, behind the red nucleus or its decus- sating medulla — the cerebellar peduncles. This very obvious column of fibres is seen as one of the most striking features of the tegmentum throughout the mesencephalon and down the whole of the medulla oblongata. We have already seen that this fasciculus originates in a compressed ganglionic mass forming the second stratum of the ansa peduncularis, and that its coronal origin is from the cortex of the operculum, insula, and temporal lobs ; it is traced into the posterior fibres of the anterior columns of the spinal cord. Substantia Nigra of Soemmering.— Another formation seen in these transverse sections is the grey matter of Soemmering. It begins near the posterior plane of the corpora albicantia, and stretches down- wards to the lowest limits of the mesencephalon, terminating, there- fore, where the transverse fibres of the pons appear. For the greater part of its course it stretches completely across the mesencephalon in an oblique direction forwards, a line which, if continued, would meet that of its fellow at an acute angle. It owes its dark colour to an abundance of large pigment cells. We shall, when referring to the thalamus, find that in transverse vertical sections through the hindmost part of the third ventricle, this grey matter lies between the crusta and the red nucleus, and that a fan of coronal fibres is here seen passing outwards beneath the thalamus to the cortex (Mei/nert). Fibres pass downwai'ds and forwards from this grey belt into the middle and inner divisions of the crusta, and hence this substance forms a ganglion of origin for certain portions of the crusta of the cerebral peduncles, and although resting close upon the tegment behind, has no organic connection therewith. CONNECTIONS OF TEUMENT AND CRUSTA. 4 1 We have thus traced the several ganglionic structures and medullary fasciculi, entering into the formation of the mesencephalon, and it remains but to summarise the results of the inquiry. The mid-brain consists of two pairs of ganglia, the quadrigeminal bodies seated upon the brain stem or pedunCUlus CePebri, where it diverges as two branches or crura, uncovered by the transverse layers of the pons, and up to the point of its entrance into the base of the brain. The cerebral peduncle consists of CPUSta and tegment severed by the intervening substantia nigra. The nates and testes have intimate connections through their hrachia with the cerebral cortex and retina ; and below tlirough the girdle-like lemniscus with the olivary bodies, and, accord- ing to Meynert, the lateral columns of the cord ; both ganglia are also connected by their radiating central fibres with the oculo-motor nuclei in the central grey substance continued from the ventricle. In front of the ganglia and central grey substance lies the structure of the tegment, viz. : — The posterior longitudinal fasciculus ; the superior cerebellar peduncular fibres and its red nucleus of origin ; certain fasciculi crossing the median raphe from the quadrigeminal bodies ; and lastly, the layer of the lemniscus. Anterior and external to the tegment, is the crusta with the substantia nigra lying behind it. In the crusta we recognise the pyramidal tract as occupying the inner, middle third, and the portion behind and between these areas, representing respectively the accessory, fundamental, and mixed tracts ; whilst in the outer fourth pass the fasciculi of the sensory tract. In high planes and upon the innermost fibres, the deepest layer of the ansa peduncularis passes backwards to the nucleus of the oculo-motor. Lastly, the substantia nigra, peculiar to this region, represents a ganglion, from which the crusta in part arises, and which in itself is but an internode for coronal radiations. THE THALAMENCEPHALON. The region of the thalamencephalon is best exposed within the lateral ventricles, for the study of its superficial parts and their relations ; and, for this purpose, a dissection, such as shown in fig. 7, should be made, in which the relative position of the mesencephalon and epencephalon are equally exposed. We here see the tA/o great ganglia, the optic thalami, the pineal gland with its peduncle, the central grey substance (and the commissure) of the third ventricle passing downwards towards the infundibulum, and the two corpora geniculata beneath the hinder extremity of the thalamus indicating the termination of the optic tracts. These, tlien, form the chief structures constituting the "tween-brain " or thalamencephalon. To appreciate their mutual relationships — their centric and peripheric connections — a 42 TBE THALAMEXCEPHALOX. careful study of sections carried througii this region in three different planes is requisite — viz., horizontal, longitudinal, and vertical-trans- verse; but a preliminary study of their more obvious external conformation is necessary ere a more minute inquiry is instituted. The reader should refer here to the ilh;stration (fig. 7) given on p. 30. The optic thalami are somewhat oval, -wedge-shaped bodies, broadest behind, where they diverge from each other so as to expose the quadrigeminal bodies ; and narrowed anteriorly where they approach the middle line. They are limited externally by the StPia teFminaliS (cornea), which is also the upper and outer boundary of the thalamen- cephalon — immediately external to which is the caudate nucleus and its attenuated tail. In front, the thalamus presents a notable prominence, the anterior tubercle ; behind, it projects back as the pulvinar, and forms in the descending horn of the lateral ventricle the anterior wall or roof of the cornu. Mesially, the thalami are bounded by the peduncles of the pineal body ; and the vertical median grey walls of the third ventricle do not, as might be conjectured, represent the median aspect of the thalami, but must be carefully distinguished therefrom. In fact, the mesial aspect of these grey masses is here completely concealed beneath the grey matter of the third ventricle, which is identical and continuous with the central grey substance of the cerebro-spinal tube throughout its length. To the thalami, however, belong the middle and posterior commissux'es which cross the ventricle, and which are really deCUS- sating" medullated tracts of the thalami. AVhilst the inner face of the thalami covered by the central grey substance is perpendicular, the outer presents a kind of obliquely sloping roof resting upon the fasciculi of the internal capsule : and hence the vertical transverse section of the thalamus is likewise somewhat wedge-shaped in configuration. Then again, it must be re- membered that the thalami in lower vertebrates— birds and reptiles — are very evident projections on the upper surface of the peduncles not included within the hemisphere at all ; and that in man, although they appear thus to project within the ventricles and to be included within the more extended hemisphere, they, in reality, are outside the hemisphere of which the fornix constitutes the median boundary. We have spoken of the thalamus as a somewhat wedge-shaped mass, as seen in transverse vertical sections. In similar sections through both thalami, they conjointly appear like the transverse-section of a boat, keel downwards, in whicli the arched side rests as on a couch in the concavity of the internal capsule, whilst in the hollow of the keel the thalamus is separated from the capsule by a region known as the sub-thalamic region, in which a sharply-defined, biconvex, lens-shaped body is situated, to which we shall refer later on as the THE THALAMIC PEDUNCLES. 43 SUb-thalamic body (lenticular body of Meynert, or Luys' body of Forel). The anterior end of the thalamus, therefore, is placed at a considerable distance from the base of the brain and the sub-thalamic region — the mass of the cerebral peduncles and the intervening substantia nigra being immediately beneath it ; whilst the whole extent of the central cavities of the third ventricle and its grey walls continuous below with the infundibulum and posterior perforated space, must be excluded from the true thalamic structure. The interior of the thalamus consists of a large mass of grey matter, split into layers in various directions by the medullated strands passing into its structure. The grey matter encloses numerous nerve cells, which are the thalamic termini for coronal radiations connecting the most diverse regions of the cortex with this body, and the centres of origin for fresh strands which pass down into the te^mentiim of the crus. The arrangement of medullated and grey elements is peculiar. The cortical fasciculi, as they enter the thalamus, diverge within its structure in brush-like fashion, forming concentric lamellfe, between which are intercalated the layers of grey matter with their nerve cells. Since medullated fasciculi enter the thalamus from very distant regions of the cortex — from the frontal, occipital, temporo-parietal gyri and gyrus fornicatus — they necessarily meet at varied angles, and cross each other in their course within the ganglion : thus it is that this body becomes moulded by its medullary cones into apparently distinct segments — not, however, true centres or nuclei in the usual acceptation of the term, since their grey substance freely intermingles with that of neighbouring .structures. The cortical contribution to the medullated system of the thalamus approaches that body in part at its anterior extremity by three so-called peduncleS — the anterior, superior, and internal (or infer- ior). These thalamic peduncles connect its structure with the frontal lobe, the sylvian fossa, temporal lobe, and gyrus fornicatus res})ec- tively. Such cortical fasciculi have necessarily a lengthened course to pursue, and none more so than that from the g'ypus fomi- CatUS, which reaches its destination after a peculiarly complex spiral course. The anterior peduncle approaches the thalamus from the frontal cortex througli the strands of the anterior segment of the internal capsule between lenticular and caudate nuclei, interlacing here with its tibres, and eventually passing into the front end of the thalamus, expands brush-like in its interior, its fibres arching backwards, crossing the fibres of the inferior peduncle, and passing chiefly to the outside of the latter. A portion decussates at the middle and posterior commissure, whilst the rest continues directly down the tegmentum. Part of its fibres help in the formation of the capsule of the thalamus or so-called stratum ZOnale. The inferior 44 THE THALAMENCEPHALON. peduncle, already alluded to as the third layer of the ansa pedun- ciilaris, connected with the cortex of the temporal lobe, passes from beneath the lenticular nucleus up into the thalamus, expanding also in brush-like manner chiefl}^ along its internal portion, and forming the inner boundary of the thalamus. It also deciissates at the middle and posterior commissure to pass down as tegmental fasciculi of the crus. Both these peduncular expansions are interrupted by the nerve cells in the grey intercalated layers of the thalamus ere they decussate at the commissures. The SUpePiOP peduncle takes a still more com- plicated course ; its centric connection is with the cortex of the gyrus fornicatus — appearing first in the two fimbpice or posterior pillars of the fornix arising from the cornu Ammonis ; and ascending as the body of the fornix connected by the transverse fibres of the lyra upon the thalamus, it arches forwards at the front end of this body, and thence passes downwards as the two descending" pillars of the fornix. These latter pass back to the corpora albicantia, around which they form a distinct loop, and again turn upwards as the ascending" pillars or bundles of Vicq D'Azyr to terminate within and around the anterior tubercle of the thalamus. Whilst forming this loop around the COrpuS albicans, a portion of its fibres is interrupted by nerve cells within this body, and a fasciculus starts from this site and passes directly backwards into the tegmentum. Whilst the frontal, insular, and median cortex is thus connected by the thalamic peduncles to the anterior end of the ganglia, the posterior or hinder half of the thalamus receives along its outer mai'gin coronal radiations from the occipital and mid-regions of the hemisphere. These fasciculi radiate from the upper and outer border of the thala- mus to corresponding regions of the brain opposite them ; the middle section spreading towards the mid-regions ; and the posterior arching backwards towards the occipital pole. These latter, as they pass outwards and backwards to the occiput, are associated with similar radiations proceeding in like direction from the geniculate bodies and the brachia of the nates and testes. This system of fibres arches around the outer wall of the posterior cornu of the ventricle, and has long been known as the optic radiations of Gratiolet. In their course they are brought into close association with the sensory fibres of the cord destined for the occipital and temporal lobes ; and, as we have previously seen (fig. 7, S), occupying the outer fourth of the crusta. This peduncular sensory tract, it must be remembered, has no con- nection with the optic thalamus, but runs directly into the occipital and temporal regions of the cortex. The coronal radiations which enter this outer border of the thalamus, pass through its structure as arcAeo? /asciCM^i towards the median line — /.e., across the long axis CORTICAL CONNECTIONS OF OPTIC THALAMUS. 45 of the thalamus ; the meduHated tracts being intercalated by the grey matter common to the whole ganglion. Upon a lower level than the entrance of these cortical radiations, other medulltited fasciculi pass into its substance in an identical manner from the middle root of the optic tract, and this double origin partially severs this hinder region of the thalamus into an upper and a lower segment. In botli systems of fibres, hemispheric and retinal (through the optic tract),, union of the fibres is effected with the cells of the grey intercalated layers. It has been shown that the peduncles — anterior and inferior — entering the anterior pole of the thalamus, run backwards through its structure as brush-like formations to terminate in cells of the grey matter ere they decussate at the commissures ; and that a larger pro- portion of these fresh fasciculi do not decussate, but pass directly down- wards into the tegmentum. The latter direct fascicidi, in passing inta the hinder half of the tegmentum, run immediately across the axis of the optic and COPtical Padiations just described ; and necessarily form apparent concentric dissepiments in these regions. These laminated dissepiments form the new meduUated tracts for the tegmentum arising within the grey matter of the thalamus. The anterior peduncle especially, passing backwards through the thalamus, is not crossed by these transverse radiations, and its region is bounded on the outer side by a strongly-marked meduUated belt, the innermost of the concentric dissepiments alluded to, and known as Burdach's lamina medullaris. This well-marked boundary and absence of transverse radiating fibres, maps out a kind of nucleus in this region of the anterior peduncle, which is known as the centre median of Luys. On examining the thalamus from above, after opening-up the lateral ventricles, it is found that the grey matter forming the tail of the caudate nucleus may, by gentle pressure with a brush, be raised away,, together with the stria cornea, from the subjacent parts ; and, imme- diately beneath it, radiating fibres in coarse fasciculi are seen passing from the whole extent of the upper margin of the thalamus, either directly outwards towards the parietal lobe, or arching back towards the occipital region. These fasciculi consequently form the outer wall of the lateral ventricle in their course towards the parietal lobe. If the scalpel divide these fibres across parallel to the direction of the stria cornea, the blade passes directly into the internal capsule, and it becomes evident that the outer obliquely-placed surface of the thalamus rests upon the internal capsule as upon a couch, and gives off from the whole of its outer aspect meduUated fibres which enter into the constitution of this capsule, and then spread as coronal radiations to the various districts of the cortex of the parietal and 46 THE THALAMENCEPHALON. temporo-sphenoidal lobes. The greater bulk of these pass dea'ply into the thalamus, and, as before said (p. 43), are crossed by the brush- like fasciculi of the thalamic peduncles. The more superficial layer iirst revealed upon raising the tail of the caudate, enters into the con- stitution of the white capsular investment of the thalamus (stratum ^o?^a/e), which gives to this ganglion its peculiar white hue within the ventricle, as contrasted with the greyish aspect of the caudate nucleus. The capsule of the thalamus spreads inwards as far as the peduncle of the pineal gland ere it turns downwards to form part of the inner investment of the thalamus ; and at this line it disappears from view, and the grey matter of the third ventricle becomes apparent. This capsule or stratum zonale is itself of complex formation : it receives also fibres from the optic tracts, the uppermost of those which join the thalamus ; so also fasciculi from the frontal lobe enter it by the anterior })eduncle of the thalamus, and in like manner the most superficial stratum of the ansa lenticularis ; lastly, the gyrus fornicatus sends its contribution by means of the ascending pillar of the fornix, which in this course embraces a nodular segmented portion of the thalamus at its anterior extremity, termed the anterior tUbercle. Hence the zonular layer or thalamic capsule receives fibres from almost every region of the brain — the frontal, parietal, temporo-sphenoidal, and occipital lobes, and the mesial aspect or gyrus fornicatus, as well as the retina. This very extensive retinal and hemispheric connection of the thalamus may be tlius tabulated : — Fasciculi from Frontal lobe, .... Through anterior peduncle of thalamus. Temporo-sphenoidal lobe, . Coronal radiations and superficial layer of ansa lenticularis. Parietal and occipital lobes, . Coronal radiations along its whole outer surface. Gyrus foiiiicatus, . . . Through pillars of fornix. Retina, ..... Through uppermost thalamic connections of optic tract. The Pineal Body and its Connections.— Surrounding the upper pair of the quadrigeminal bodies, immediately beneath the posterior extremity of the callosal commissure and in the middle line between the mesencephalon and diencephalon, lies a small, reddish, somewhat conical structure — the pineal body. It is closely attached to the velum interpositum, so that it is frequently torn away with the membranes investing it. It is hollowed into several small sacculi, which contain the gritty, earthy, and amylaceous material termed acervulus cerebri : and the structure is peculiarly vascular. In microscopic structure we find it consists, like other ganglionic struc- THE PINEAL BODY AND FASCICULUS RETROFLEXUS. 47 tures in the brain, of closely aggregated cells, varying considerably in size from 5 /x to 18 ,a. Its connection with the rest of the cerebrum is effected by means of two processeSj which are directed forwards along the inner border •of the thalami optici, forming a boundary between the latter and tlie grey matter of the third ventricle ; and descending in front in conjunc- tion with the taenia semicircularis and the pillar of the fornix : these are the two peduncles of the pineal body or habenula. These peduncles are distinctly ganglionic in structure, and together with the pineal body are probably to be regarded, as Meynert believes, as ganglia of origin for the tegtnentum,. The connections by meduUated tracts are twofold — centric and peripheric. The former, as a connection with the cerebral hemi- spheres, takes place through the medium of the stratum zonale, already described as investing the optic thalamus. The latter or peripheric connection is effected by a large and important fasciculus, which passes down vertically from the habenula or peduncle, covered by the grey matter of the third ventricle, and towards the region (at the base) of emergence of the motor oculi nerve on the inner side of the converging crura. In this course it describes a sigmoid bend, and near the base of the mesencephalon it lies between the posterior longitudinal fasciculus, on the median aspect, and the red nucleus of the tegmentum, external to it. Some of its fibres radiate into the nucleus ruber (Meynert) ; but the larger proportion bend at this point immediately backwards at right angles to their former course, and appear to pass into the tegmental areas of the pons and medulla, in conjunction with thQ posterior longitudinal fasciculus. This rectan- gular bend has gained for it the appellation of the fasciculus retFO- ilexUS — it is often termed the Style of the peduncle of the pineal body, where it passes vertically towards the red nucleus of the tegmentum. The style or fasciculus retroflexus may be best exposed by trans- verse vertical sections carried through the ganglion of the peduncle just in front of the quadrigeminal bodies, but it may also be traced in longitudinal vertical sections near the mesial plane of the ^Hween and mid brain." In these sectional planes, however, owing to its sigmoid flexure, a part only of its course can be usually seen. Thus in a vertical longitudinal section of the brain of the dog, near the mesial plane, we find the lower end of this fasciculus about to bend backwards at right angles, and on this plane it is seen to descend in fi-ont of meduUated fasciculi passing downwards from tlie i)Osterior commissure and the emergent roots of the third nerve. In a section carried still nearer the mesial plane, we see its course about]complete, whilst a portion ■of both ascending and descending pillarsof the fornix is revealed likewise. 48 THE THALAMENCEPHALON. Posterior Commissure, — We have already traced the anterior and inferior peduncles of the thalamus as far as their decussation in the posterior commissure, and it would seem extremely probable that the fasciculus retrqflexus undergoes partial decussation through the medium of this commissure also. Near the mesial line, we can readily trace these decussating fibres of the posterior commissure in their further course passing downwards into the tegmentum, where they bend backwards to pass into the medulla and spinal cord ; whilst prolonged from the posterior commissure backwards is also seen the cross-section of the medullated fibres of the corpora quadrigemina. In these vertical longitudinal sections taken near the mesial plane, we therefore see three systems of decussating fasciculi crossing at the middle line, and forming peripheric extensions from a series of gang- lionic bodies, viz. : — The fasciculus retroflexus, the mass of the posterior commissure, and the quadrigeminal fasciculi called the- lemnisci or fillets of the nates and testes. Corpora GeniCUlata. — Beneath the pulvinar of the thalamus in man we see a small club-shaped body about the size of a coffee-bean,, directly continuous with the optic ti-act anteriorly, and by a notable border separating mesencephalon from thalam,encephalon, connected with the upper quadrigeminal body or nates. This small ganglion, for ganglionic it is in nature, is the OUter g'eniculate body, and lies in the course of the arm of the nates or superior hrachium,, with which it is intimately connected, as it proceeds to the cortex of the occipital lobe. Upon vertical longitudinal section it is found to- possess a peculiar plicated arrangement of a medullated and a grey lamina, exhibiting alternating layers of grey and white substance. Internal to this body, that is, nearer the mesial plane, lies another small structure of spindle-shaped outline, immediately beneath the upper and between it and the lower brachium ; it is directed towards the nates by one of its pointed extremities. This structure is the inner g'eniculate body. Both geniculate bodies are connected with the corpora quadrigemina on the one hand, and with the cortex of the occipital lobe along with the other centric fasciculi of the brachia. In a vertical section we find an extensive portion of the optic tract directly continuous with this plicated outer geniculate ganglion, and hence also with the nates. An inner segment of the optic tract, but much more limited in extent, passes into the internal geniculate and thence to the nates also ; no fibres from the optic nerve are believed to pass by this tract to the testes. The remaining connections of the optic tract are the optic thalamus (to the stratum zonale and radiating fasciculi previously described) ;. and the basal optic ganglion, a small body of grey matter lying beside the tuber cinereum immediately covered by the optic commissure. CONFIGURATION OF CEREBRAL HEMISPHERES. 49 The ganglia of origin of tlie optic nerves, therefore, are the upper quadrigeininal, the outer and inner geniculate bodies, the optic thalamus, and the basal optic ganglion : the centric or coronal extensions arising in these ganglia pass by means of the posterior section of the internal capsule as the optic radiations of Gratiolet to the cortex of the occipital and (?) temporo-sphenoidal lobes. PROSENCEPHALON OR FORE-BRAIN. Conflg'UPation. — We have already seen that divergence of the brain-stem in the crura cerebri to reach either hemisphere, entails also the more and more complete severance of the various ganglionic masses at the base with which it is brought into connection : and that from the bilateral fusion of the mesencephalon, we pass forward to the divergent masses of the thalami (diencephalon), and thence to the still further severed corpora striata, constituting the ganglia of the fore-brain (prosencephalon). We have seen how these more divergent masses are braced together by sling-like loops of medulla, such as the ansa lenticularis, and united mesially by the anterior and other commissural tracts. The ganglia of the prosencephalon form the most anterior mass of grey matter surrounding the peduncular extensions, and are so disposed as to constitute two incompletely- severed masses of grey substance, whose configuration shadows forth the form assumed by the hemispheric envelope moulded around them. The flexure of the fore-brain, whereby this hemispheric arc reproduces the contour of these ganglionic structures, has its site at the fissure of Sylvius ; and, in foetal brains, ere the further differentiation of the cortex into its varied longitudinal and transverse fissures has proceeded, we see readily how the hemispheres are, so to speak, moulded after the form of their subjacent ganglia. The axis of this flexure is constituted by the most external of these ganglionic masses, the so-called lentiCUlaP nucleus, wedge-shaped in form, its base directed forwards and out- wards, covered by the cortex of the insula — its apex downwards and inwards towards the crus cerebri. Around this wedge-shaped axis, the ganglionic and hemispheric arcs are severally formed — the ganglionic, in the form of the caudate nucleus : the hemispheric, beginning at the orbital aspect of the frontal, sweeps round the fronto-parietal to the tip of the temporo-sphenoidal lobe. The more flattened aspect of the region of the insula, therefore, bears the impress of the base of this lenticular axis of revolution, whilst the more spheroidal contour of the hemisphere conforms to the curvature of the caudate body. Upon this constructive principle largely depends the divergence observed in the primitive contour of the cerebrum in various animals and in man. Although identical in the nature of their histological 4 50 THE PROSENCEPHALON. constituents, these two ganglionic masses differ widely, not alone in their rough contour, but in their quantitative relationships : in certain brains, the caudate nucleus assumes a mass far out of all proportion to the lenticular ; whilst in man, the former is dwarfed, and the latter assumes a relatively important role. The greater magnitude assumed by this lenticular axis of revolution, the greater the scope for the unfolding of the hemispheric arc, and the more important the develop- mental features assumed by the regions of the insula and sylvian Fig. 8. — Section through hemispheres (vertical transverse) through plane of middle commissure. N.c, Tail of caudate nucleus. Cx^ Corpus callosum. F, Fornix and choroid plexus. a. Internal capsule. T.o, Optic thalamus. C.?;i, Middle commissure. C.e, External capsule. CI, Claustrum. 1, 2, 3, Three segments of lenticular nucleus. fossa. On the other hand, the smaller lenticular body, and the larger proportionate development of the caudate bespeak a brain of simple configuration, more spherical, less complex in convolutionary arrange- ment, and of more uniform symmetry throughout. In thus indicating their impress in the configuration of the cerebral vault, these striate ganglia differ widely from the diencephalic ganglia previously considered, the thalami Optici ; in fact, these latter bodies, so far from having any portion of the cerebral hemi- sphere moulded to their form, are themselves wholly outside the cerebral envelope in their mesial position. Thus, the adult brain CAUDATE NUCLEUS. ^I "witnesses to the genetic relationship of the cerebral hemispheres, and the related striate ganglia ; the whole mass in front constituting the fore-brain or prosencephalon in advance of the thalamence- phalon. We have spoken of these basal ganglia as incompletely severed masses of grey matter, a statement at once verified by vertical sections taken in anterior planes through these bodies. In such -iinterior planes, the medullated interval elsewhere separating these bodies is bridged by numerous broad bands of grey substance which are but extensions from one to the other ganglion ; whilst, at the base, •complete fusion occurs between the two, the head of the caudate nucleus merging into the frontal extension of the lenticular, becom- ing so superficial at the base as to be merely covered over by the ■orbital medulla and the grey matter of the anterior perforated space with which it becomes continuous. Caudate Nucleus. — The innermost or intraventricular nucleus of the ganglion of the fore-brain, and the only portion superficially seen within the lateral ventricles, is of pyriform shape, with a long attenuated tail-like process extending into the temporal lobe. In this course, as before explained, it is bent upon itself, its axis of ■revolution being the lenticular body. The head of the ganglion fusing at the base with the lenticular, arches forwards and inwards towards the septum lucldum, and, lying on the inner aspect of the internal capsule, embraces, in this first part of its course, an important medullated fasciculus, which connects the cortex of the frontal lobe with the anterior extremity of the optic thalamus, the so-called anterior thalamic peduncle. In its further course, it ascends .above the level of the lenticular, and lies upon the internal capsule ; its tail-like extension resting, opposite the thalamus, upon the hemi- spheric fibres which pass beneath it, to form the capsular investment of the thalamus {stratum zonale). Still further back, the tail arches downwards into the descending hoi-n of the ventricle, and can be ti'aced upon the roof of the latter as far forwards as its anterior extremity, where it terminates in a somewhat bulbous end, having immediately in front of it a mass of grey matter, termed the amyg'dala. It will be seen from this description that the bulbous extremity of the tail extends almost as far forwards as the head of the caudate nucleus, and thus describes an almost complete loop around the internal capsule and thalamus, hence termed the " SUrcingle." The whole course of this loop can be well demonstrated by vertical longitudinal sections of the hemisphere ; whilst vertical transverse sections anywhere between the amygdala and posterior end of thalamus reveal the upper and lower segments of the surcingle, as isolated grey masses above and below the thalamus. Each of these prosencephalic 52 THE PROSENCEPHALON. ganglia has a surface perfectly free, that is, devoid of medullated attachments — and other aspects, which present the termini of centric- and peripheric strands. Thus, the ventricular aspect of the caudate, together with the base of the lenticular wedge (insular aspect), are alike smooth and devoid of medullated connections ; whilst the opposed surfaces, separated by the intervening capsular fibres, as well as the basal or inferior aspect of the lenticular nucleus, are the surfaces for the termination and departure of the numerous medul- lated connections of this with distant regions. Since the lenticular body lies beneath the internal capsule, its temporal extremity is separated for some distance by that formation from the temporal extremity or cauda of the intraventricular nucleus. Posteriorly, however, they approach each other, and bridges of grey matter connect them, separated by medulla. They are also separated here by the centric extension (brachium) from the external geniculate body ; and, finally, along the roof of the descending cornu these two temporal extremities fuse together, forming the lower segment of the surcingle. The constitution of the surcingle, therefore, is different in its upper and lower arc, being pvirely an extension of the innermost nucleus above ; but formed out of the fused tem])oral extremities of both prosencephalic nuclei below. It cannot fail to impress the student that the ganglionic structures and their extensions, so far described, encircle in a series of loop-like formations the medullated core which passes from the spinal cord and medulla upwards as peduncles and capsule to the cerebral hemispheres. First, there is the mesencephalon, the quadrigeminal bodies, each throwing downward its loop-like fillet or lemniscus ; and throwing upwards its centric arm in the form of the brachia. At a higher level, the thalamencephalon shows us the optic thalamus astride the posterior edge of the internal capsule, arching backwards around it to form the roof of the descending horn of the lateral ventricle ; whilst its centric extensions pass upwards to the cortex from its outer surface beneath the tail of the caudate. Then still higher we get the arc of the caudate body astride the anterior edge of the internal capsule with its long tail-like loop also passing down the roof of the descending cornu in conjunction with that of the lenticular : whilst still further outwards is the mass of the internal capsule becoming free as coronal radiations to the various parts of the hemisphere. For descriptive purposes it is convenient to distinguish between upper or ventricular, and lower, cornual or temporal arc of the surcingle : the caudal extremity, the body of the caudate nucleus, and its caput directed towards the base : whilst we also speak of its ventricular and capsular aspects. In like manner the lenticular nucleus has its frontal, its temporal. STRIA TERMINALIS -OLFACTORY AREA. 53 And peduncular or crustal extremity ; its insular aspect (or base of wedge), its inferior aspect, its capsular aspect. So also the capsular constituents may also bear the convenient terminology — COPtiCO- Striate and COrticO-lentiCUlar fasciculi for the centric bundles : pedunculo-striate and pedunculo-ventricular for the peripheric bundles : direct pedunCUlaP for those uninterrupted by the prosen- •cephalic ganglia. Stria Terminalis. — A glistening white band of fibres, strongly contrasting with the adjacent grey cauda, varying from one to two millimetres in diameter, lies along the inner border of the tail of the •caudate body throughout its whole length, extending from the tip of the temporo-sphenoidal lobe along the roof of the descending cornu, ^nd along the upper arc of the surcingle betwixt it and the thalamus, as far as the anterior end of the latter. Inferiorly it is ' 3 o. YihA. M-.--. or- Cortex of Pi^ -left KemispKere '<>i'bilm«' its five-UiTun?«U-d type ^ ith tlvej>lests or clusterf: r-ic cel\s. y /o Bale &Darvielsson,Lt<3 , Sculp THE XEUR0C4LIA MATRIX. 93 different regions of the cerebro-spinal system. Tliis, we find, accords ■with actual fact ; for, as a supporting, as well as embedding and protective material, the requirements demanded will differ widely in the white medullated structures from those of the grey centres ; whilst individual sections of these territories will also differ in the special ' qualities of this matrix requisite. Thus, the large closely-approximated medullated fibres of the Spinal Cord will be found to possess a strong binding material in the form of large-sized nucleated cells, with numerous lengthened ramifying processes, together with a plexus of fine fibrils (probably elastic tihre — Gerlach) ; whilst a structureless or very finely granular material is found here but sparingly. Still nearer the periphery of the cord, this supporting structure becomes a veritable fibrillar connective sheath of great strength, with trabeculse of like constitution passing inwards to the cord. In the central grey matter of the cord, however, the finely granular or molecular basis-substance predominates, as most essential for the protection of the extremely delicate nerve fibres present in this region. Farther up in the medulla of the brain, as in the neighbourhood bordering upon the grey cortex, the large bundles of medullated fibre again demand a predominance of the connective fibre element, so that here we meet with numerous though delicate ramifying cells. Wherever the medul- lated fibre reappears, there we find the association of these branching cells, and thus they are seen along the outermost or peripheral layer of the cortex as a normal element. In the grey matter of the cortex, however, the delicate nerve-cell and fibre network appears largely to dispense with this modification of the connective tissue, and we find a structureless matrix vastly preponderating over the cell and nuclear elements of the neuroglia. A still further modification of the neuroglia element is found on the free surfaces of the cortex immediately beneath the pia, where the branching cell before described fulfils the function of a flattened epithelial investment, whilst the surfaces not exposed to pressure, as the central canal of the cord, show us the element as a columnar epithelium. Thus, generally, we may affirm that, when dealing with nerve cells and their delicate extensions, the supporting material will be chiefly the structureless or finely molecular basis-substance ; whilst as we approach the medullated tracts, we shall find that the COnnectiV© cell and fibre networks increase at the expense of the former. The elements of the neuroglia are usually described as nucleated cells and free nuclei imbedded in a structureless, or, according to some, finely fibrillated matrix, and to this view tlie appearance of chrome-hardened preparations certainly lends support. The less we subject our sections to reagents, and the more recent the section examined, however, the 94 THE CEREBRAL CORTEX. more evident it becomes that the supposed free nuclei are invested by- protoplasm, and, in fact, are likewise nucleated ceils. These two cell elements differ much as regards their relationships and also their ■dimensions. (1) The smaller of the two kinds of cell vary from 6 /x to 9 ,'x in diameter ; have a spheroidal nucleus, surrounded by an extremely ■delicate protoplasmic investment, which, as before intimated, is shrunken, often beyond recognition, in hardened specimens. The nucleus is, proportionately to the cell itself, very large, and invariably ■Stains of an intense depth of colour with aniline blue-black. These elements appear disposed in three definite situations — (1) irregularly in the neuroglia framework ; (2) in regular series around the nerve cells; (3) in more or less regular succession along the course of the Mood-vessels (capillary and arteriole). (2) The larger cellular elements of the nexiroglia are usually 13 /^ in •diameter, and supplied with a relatively larger mass of protoplasm as compared with the nucleus. They are distinguished from the former not alone by this greater size and the preponderance of cell over nucleus ; but also by their frequent flask-like configuration, as seen in situ, and the presence of a very faintly stained nucleus, or even sometimes two or three nuclei, observed within them. If these elements are teazed out from the surrounding matrix, they are seen to possess numerous extremely delicate radiating processes ; not only the nucleus, but the cell and its extensions are likewise tinted by the .aniline dye ; not uniformly, however, for the nucleus is always of a slightly deeper tint, but neither cell nucleus nor processes betray anything like the vigour of staining shown by the former element -described. The nerve cell, its processes, and the enclosed nucleus had, as we said, a special afl&nity for this staining reagent, a fact, which indicates very conclusively the non-nervoiis character of these larger elements of the neuroglia. In healthy brain, at least in the human subject, we find these elements chiefly in the outermost layer -of the cortex and the central cone of the medulla, but their delicacy, itenuity of branches, very faint staining, and poor differentiation are not favoui'able to their immediate detection. In certain morbid conditions -of the cortex, as we shall see later on, these elements become a most notable and important feature, undergoing excessive proliferation, and betraying their morbid activity by the intensity of colouring which they acquire. If now we appeal to the silver-chrome methods of preparation, the appearances are very dissimilar. Two forms of cell obtrusively present themselves in both the white and grey matter of the brain ; the one with a poorly defined cell body, obscured by much deposit of silver, throwing off on all sides short, shaggy, protoplasmic processes, THE NEUR0C4LIA MATRIX. 95 dendritic or dichotomously dividing ; the other, a cell element, also badly defined, forming a centre from which radiate outwards exceed- ingly fine fibrils, often of gi-eat length, sharply defined contour, not ■dividing dichotomously, nor presenting the thickened hispid aspect of the processes of the first described element. These exceedingly fine fibrils are characterised not only by their length, tenuity, and clean contour, but also by their tendency to exhibit many sharp angular bends along their course. When the former element is more closely examined, we find the invariable presence of one or more lengthened thick processes by which the cell is attached to a neighbouring blood-vessel ; such pro- cesses end in a sort of conical or flattened sucker-like thickening on the vessel's walls. We have here, in fact, modified by the silver- chrome method, the same element described as the large cellular element of the neuroglia, and which, later on, we shall refer to as the spider cell. The second form, or stellate cell, has by some been regarded as a distinct element ; it is found more particularly in the white matter of the brain, and under the pia covering the cortex. A point of great significance, however, is the fact that they lie invariably in close contiguity to a blood-vessel, and that in favourable prepara- tions, when the dense deposits of silver are dissolving ofi", we can identify them as spider or Deiter's cells, the vascular attachments being clearly apparent. Between these two forms, superficially so different, there exists a further transitional element readily distin- guished ' in most regions of the brain and medulla, but especially beneath the intima pia, and which at once indicates to us the essential identity of the two former kinds of cell. The transitional forms alluded to are strung beneath the pia, attached by a vascular process to a vessel of the latter, whilst from the central end of the cell, which is usually ovoid in form, a large number of extremely long delicate fibrils extend into the first and subjacent layers of the cortex, resembling in all respects those of the stellate cell just described. These fibres, however, arise from short stunted protoplasmic processes which are truly dendritic and hispid, and take their origin immediately from the cell body, whilst some of these fine fibrils present numerous moniliform enlargements along their course. To summarise, therefore, we find three stages of development presented by these elements, viz. : — (a) Cells with short, thickened, moss-like protoplasmic processes, dendritic or branching dichotomously, and possessing also thick vascular attachments ; (b) Long and exceedingly fine unbranclied fibrils radiating from an obscurely marked central cell, also (under cei'tain conditions) showing vascular processes ; g6 THE CEREBRAL CORTEX. (c) Transitional forms with vascular processes, short dendritic branches, from which are given oft' long delicate fibrils like those of the stellate cell, and often distinctly moniliform. As stated elsewhere, we regard all spider-cells as destined to pass through the three stages — the embryonic moniliform, the fully developed spider-cell or "lymph-connective" stage, and, lastly, the stellate cell; whilst in normal devolution the lymph-connective or spider-cell passes back into the fibre state of the stellate cell, losing its active functional manifestations, and assuming, therefore, the purely mechanical rtU of a support for the blood-vessels and meduUated nerve fibres in its vicinity.* This study of the constituent histological elements of the cortex prepares us for the consideration of the lymphatic system of the brain, and the ultimate relationships of ISTerve cell to the Blood and Lymph channels. {d) Lymphatic System of the Brain.— To Obersteiner is due the credit of first definitely indicating the existence and relationships of these lymph channels.t Their existence since then has been re- peatedly denied, but the evidence hitherto brought forward against Obersteiner's views is most inconclusive in all respects, and in most cases apparently based upon incomplete methods of examination. This is not the place to enter on debateable ground ; but we are compelled, owing to the'supreme importance of the subject as aflFecting the physiology and pathology of the brain, to state the results of our own investigations, which were made the subject of a special memoir in 1877. + All hardened sections of brain exhibit along the course of their blood-vessels a distinct and more or less wide interval between the vascular walls and the brain-substance; in fact, the brain cortex is channelled throughout, in sucli a manner, that the vessels when con- tracted are enclosed within a channel of much greater calibre. The disparity betwixt the diameter of vessel and brain-channel will be affected undoubtedly by corrugating reagents ; and hence, we never fail to find these channels disproportionately large in brain which has been subject to extremes of hardening by chromic acid, &c, ; but recession of the brain-substance may occur from many other causes actinc^ during life — notably extreme atrophic degeneration ; and then, in like manner, such channels will appear inordinately large, however skilfully the brain be prepared. These channels are known * "Structure of the First or Outermost Layer of the Cerebral Cortex," Edhu Med. Joiirn., June, 1897. + "t)ber einige Lymphraiime im Gehirne" [Sitzh. d. K. Akad. d. Wis-seu-sch., Jan. Heft, 1870). + " The Relationships of the Nerve Cells of the Cortex to the Lymphatic System of the Brain,"' Proc. Boy. Soc, No. 182, 1877. LYMPHATIC SYSTEM OF CORTEX. 97 by the name of the perivascular channels of the brain — the peri- vascular channels of His : these are not the lymph channels proper, as several writers seem to have supposed, but are simple channels in the brain-substance, devoid of an endothelial lining, and communicating freely with the space between the investing pia mater and surface of the cortex, the eplcerebral space. The adventitial sheath of the blood-vessels becomes closely appressed to this limiting channel, and its (adventitial) nuclei often thus give it the appearance of being lined by endothelial cells. This, however, is not the case, as re- peated investigations by silver staining have shown. The student cannot too persistently bear in mind the fact that in these channels he deals purely with what seems equivalent to an involution of the naked surface of the brain, and yet the epithelial elements of the epl- cerebral surface are not continuous along this tubular channel. In the next place we find, under precisely similar conditions to those above enumerated, a wide space around the larger nerve cells ; the brain-substance, as it were, seems to have receded from the cell, so that it is enclosed within a circular, oval, or pyriform space. These spaces we will designate the pericellular SaCS. Genuine sacs, and not mere artificial gaps in the brain-substance, they undoubtedly are, as is abundantly proved by careful examination. To exhibit the true relationships of these perivascular channels and pericellular sacs, let us revert to the smaller cellular element described in the neuroglia (p. 94). It was stated that beyond the scattered elements in the basis substance of neuroglia, these cells were arranged in two other direc- tions. Let us particularise : — (1) The nucleated cells along the arterioles belong to the adven- titial tunic, and map out its course very accurately ; occasionally closely applied to the perivascular channel, as before stated, or separated as irregular ampullae from the vessel itself, this investment more frequently lies directly upon the media, and affords one (but an equivocal) evidence of the existence of a lymph channel surrounding the vessel. That a complete tubular membrane exists for a certain distance along the smaller arterioles is demonstrable ; that it is con- tinuous, as a membrane, further on to the arterio-capillary plexuses, is more than dubious. It is certain, however, that its representative cells are to be found surrounding these minute channels to their ultimate ramifications ; and thus, the perivascular lymph space of the adveutitia becomes continuous in these districts with the general perivascular channels and sacs around the nerve cells. (2) The nucleated cells found in connection with the nerve cell in certain states not only accumulate upon the nerve cell itself, but follow closely the outline of the cavity, or, properly speaking, the sac in which the nerve cell lies. Many pericellular sacs will show a 7 98 THE CEREBRAL CORTEX. complete series of such nucleated cells around it, still more frequently will they follow out a segment only of its circular outline ; occasionally none may be seen — an exception due probably to displacement during section-cutting or further manipulation. Upon closer observation, however, it becomes apparent that in the immediate neighbourhood of every large nerve cell there is a minute arteriole or capillary, not indicated so often by a well-diflFerentiated contour (for these minute vessels are usually most difficult to follow) as by the direction of its nucleated cells. Thus, the fusiform nuclei of the intima, alternately placed on opposite sides of the capillary, will lead to the discovery of the outline of the vessel faintly indicated in a graceful curve or spiral in close approximation to the nerve cell ; but the presence of the deep- stained nuclei of the adventitial cells taking the same course, plainly indicates the direction of these ultimate nutrient channels. It is these adventitial elements which give us the clue to tracing the obscurely marked capillary, and when this is followed out, the eye becomes accustomed to trace without any difficulty the vascular loop around the nerve cell. Around a segment of the pericellular sac, mapped out by adven- titial elements, we then see a delicate tubular loop, evidently con- tinuous with the neighbouring arteriole, and to the sides of which the pericellular sac appears to be attached, the nerve cell itself being, as it were, suspended within the latter. It would appear as if the general perivascular channels at their ultimate ramifica- tions around the arterio-capillary plexuses were enlarged here aud there laterally along the vessel by the growth of an element included within it which becomes the nerve cell, and which does not come in contact with the neuroglia matrix except through the medium of its processes, which, passing through the pericellular sac, permeate the neuroglia in every direction. It would appear also from examina- tion of specially prepared sections, that the adventitial elements are not entirely limited to the vascular loop, but may line the interior of these sacs — not as a regularly applied endothelial layer, but as loosely distributed and branching cells. In like manner, similar cells may be found free within the cavity of the sac between its wall and the nerve cell, resembling in all particulars lymph corpuscles. Beyond the system of perivascular channels, adventitial lymph space, and pericellular sac, we have a lymph-COnnective system which plays an important role in the pathology of the brain. This system is constituted by the larger connective element referred to above — the delicate branching masses of protoplasm supplied usually with one, sometimes with two, or even three large nuclei. These elements, when more closely examined, ai'e found, as already indi- cated, to have a definite and constant relationship to the cortical LYMPH-CONNECTIVE SYSTEM. 99 blood-vessels ; and are always discovered in larger numbers in their immediate neighbourhood, external to the perivascular channels. The latter present, where they are well seen and the adventitial sheath is appressed to the vessel's side, a series of delicate processes, which, traversing the channel, look like fibres extending from the adventitia into the brain-substance. What are these fibrous prolongations 1 Careful examination of one of the large neuroglia elements reveals the fact that they throw off two sets of processes — (1) an enormous number of extremely delicate fibres, which spread into the intervascular area around, and (2) a much thicker, coarser process, which, often after a tortuous course, ends in the adventitial sheath of the blood-vessel. In crossing the perivascular sac, these processes give rise to the fibres just described as extending between adventitia and brain-substance. It is in certain morbid developments of these cells that we can the more readily distinguish their real relationships. We find that the stouter process, which we may provisionally term the VaSCUlaP, 'terminates in a nucleated mass of protoplasm on the sheath itself. In morbid states, as we shall see, this terminal protoplasm of the vascular process becomes spider-like, in its turn throwing ofi" numerous branches, which embrace the vessel's wall. In the healthy state, it is most difficult to trace the vascular branch ; but that this can be done by proper methods, we liave frequently satisfied ourselves. The branched cells which we have now described have often been jrecognised in their morbid modifications, and variously interpreted. Their representatives in healthy brain were first described by Deiters,* .and subsequently by Ball and Golgi ; but we do not think their true significance has been recognised either as normal or pathological •elements of the central nervous system. We incline to regard these ele- ments as comprising the distal extension of a lymphatic system, in fact as a lymph-COnnective system permeating the neuroglia in the intervascular area. The individual elements are excessively delicate and pellucid, their protoplasm appearing almost of fluid con- sistence, and the vascular process invariably establishing its connection with the lymph sheath of a blood-vessel. In whatever manner these spider cells effect the reabsorption and distribution of the eflfete material and surplus plasma — whether by direct assimilation into their -own structure, and its removal by currents within the protoplasm of the cell and its processes, or by means of a true canalicular system terminating in the lymph sheath — it is an undoubted fact that any arrest to the escape of perivascular lymph from the cortex is imme- diately followed by a morbid development and hypertrophic condition * Hence they are often named after liira — Deiters' cells. — Untersuchimyta liber •Gehirii und Riickenmark der Menschen und der Smiyethiere, 1865. lOO THE CEREBRAL CORTEX. of this system of spider cells, as vre shall for the future call these elements of the "lymph-connective system." ■^•' !Meynert long since drew attention to their frequent presence as associated ■with congestion and degeneration of the lyn:phatic glands of the head and neck, and we have assured ourselves of the frequent association of this morbid development in tuberculosis, and in several affections of the cortex and its membranes which lead to obstruction of the perivascular lymph channels.! The morbid changes undergone by this lymph-connective system and the effects of its morbid activity will be more fully dealt with when treating of the pathology of the cortex. 1 For the present we shall summarise the above statements as follows : — The lymphatic system of the brain consists — (1) In the first place, of a distensible lymphatic sheath, loosely applied around the arterioles and venules, containing numerous nucleated cells in its texture — the adventitial lymph sJieath, the whole being included within a non-distensible channel of the brain-substance, devoid of endothelial lining— perivascular channel of His. (2) In the second place, of a continuation of the cellular elements of this sheath, loosely applied to the arterio-capillary plexuses, still contained within a perivascular channel, which now exhibit along the capillary loop sac-like dilatations — the -pericellular sacs, within which the nerve cell lies, surrounded by plasma. * It mav prove of interest to give here references to some few of the articles bearing directly upon phagocjirosis : — " Poisonous and Defensive Albumoses," by Dr. Hankin in Brit. Jled. Journ., May, 1890, p. 126. "Lectures on Phagocj"tosis and Imnittnity,'" by Sims Woodhead, Lancet. Jan. and Feb., 1892. " Discussion on Phagoc}i;osis and Immunity," Brit. Med. Journ., Feb. and Mar., 1892. " Ptomaines and Animal Alkaloids," Brit. Med. Journ., Xov., 1892. " The Spider (so-called Scavenger) Cell of the Brain," by Edwin GoodaU in Journcd of Pathology, Feb., 1894. "The Spider or Phagocyte Cells of Senile Insanity, General Paralysis, and Alcoholic Insanity," by Alfred Campbell, Journ. Mental Sc, Oct., 1894. t We have elsewhere alkided to the comparative significance of these elements as follows : — " In man they appear ia scanty numbers ; in the Barbary ape, they become more frequent ; in the cat and ocelot, they are still more abundant ; in the pig and sheep so profusely scattered are they that they form a most characteristic stratum immediately below the pia mater, and the meshwork formed by their fibres is dense and coarse, binding the blood-vessels to the cortex and rendering the pia mater strongly adherent. We find these corpuscles in human brain which has undergone senile degeneration — in other diseases attended by reduction in fimctional activity, and in vascular affections resulting in retrogressive changes and a reversion to a low type of structure." — " Comparative Structure of the Cortex Cerebri." Trans. Royal Soc, part i., 1880. t See in this connection a suggestive article by Dr. Shaw on Aprosexia in Children — a condition where adenoid growths (post pharj-ngeal) obstruct the flow of lymph from the frontal lobe, which naturally escapes through the lymphatics of the ethmoid plate. — Practitioner, July, 1S90. r'UneVJ. . i/> ,ix ■ I »^ -A i^ r ,'■ * . : ' 1 •Ny'jr^v• •*>\; ; '' ^ s L/-~-*f ^^ ,R ;■>>- Zarge Pyramids of lov,\ , strata of Third lajc Cranizle elements in Fourfh layer. Ganglionic cells of Fifth layer. Spindle eLeinenis of Sixth layer K|-^ : : 6.1 "S e ns ory typ e " „ , ^ „ l^erve elements of 3- .^'QJ, 5 - &.6*^ layers t)ensor>^ \vpe of First AriTiectaiit Gyrus of uuor. OvroMsn rirst Anrvectant Human Brain. Tvn;? of hinr,;^:, Cortex. x65. xl57. Bale iDanielsson. Ltd Sculp. RETICULATED WHITE SUBSTANCE. lOI (3) Lastly, of a system of plasmatic cells with numei'ous prolon- gations, which are always in intimate connection with the adventitial lymph sheath, and which drain the areas between the vascular branches : these we have termed the lymph-connective elements. If we take a comprehensive view of the whole system — the channelled vascular tracts, the saccular ampullae along the capillary tube, the canalicular-like formation of the lymph-connective elements, all em- bedded in a homogeneous matrix of neuroglia — we cannot but be struck by the sponge-like arrangement of the cortex, and the facilities so afforded for the free circulation of plasma throughout its most intimate regions. CORTICAL LAMINATION. Having familiarised himself with the individual histological elements of the cortex — the nerve cells, blood-vascular and lymph-vascular systems, and the neuroglia framework — it becomes the student's duty to examine their general arrangements and the local deviations to be observed. A vertical section of fresh cortex of human brain reveals to the naked eye a distinctly laminated aspect, the various laminae of which are more or less clearly marked out by difference in colour, the outer being usually of a pale translucent grey, and the deeper of alternating pale and dai'k grey layers, more opaque in aspect, and in certain regions exhibiting a sharply-detined white streak. The outer trans- lucent layer has superimposed on it a delicate white stratum, scarcely appreciable on the convexity of the hemisphere, but well-marked in the convolutions bordering upon the corpus callosum, and the convolution of the hippocampus, at the base, where its peculiar aspect has gained for it the name of the reticulated white SUb- Stance. As we shall see later on, this is a superficial layer of white medullated fibre running parallel to the surface of the convolution ; whilst the paler intersecting streaks deeper down in the cortex are similar systems of arciform intracortical fibres intervening between layers of grey substance. The deeper layers owe their opacity to the relatively large proportion of medullated fibres passing through them ; the upper layers are translucent from the preponderance of the neuroglia element and fine protoplasmic processes of the nerve cells ; the warmer grey tints are due not only to large numbers of pigmented nerve cells, but chiefly to the amount of blood in the vessels of the layer. As might be supposed from the above, the distinctness of lamination not only varies with the local peculiarities of structure, but with morbid states of the cortex and with the full or empty state of its vessels. Probably the best introduction the student can have to the I02 LAMINATION OF CEREBRAL CORTEX. study of the human cortex is to commence first with the brain of one of the lower mammals, choosing one of the smooth non-con- voluted brains, as of the rat or rabbit, ere he attempts the more complicated brain of those animals which exhibit a convoluted surface. He thereby learns to appreciate the great diversity of lamination which may exist in so small an organ as the brain of the rodent, as also the abrupt transition from one type of cortex to that of another wholly difierent from it, and lastly he becomes familiar with types of lamination which are strictly reproduced in higher forms up to the brain of man. Figs. 1-3 in Plate i. represent the brain of the rabbit seen from its upper, lower, and median aspect, of somewhat pyriform contour below at the base, and triangular above ; its frontal pole is much attenuated, and rests upon the olfactory lobe. On its inner aspect we see two very delicate furrows (fig. 1, A) which represent the sub-frontal and sub-parietal segments of the limbic fissure, which is strongly marked in the brains of the pig and of the sheep ; this rudimentary fissure limits the upper limbic arc (between A and J) from the extra- limbiC or parietal mass of the hemisphere (fig. 3, Z, Y). If we follow this upper limbic arc from before backwards, we find that its anterior extremity is deep, and that it gradually becomes more shallow towards the sub-parietal furrow ; beyond this it is hollowed out by the prominence of the mesencephalon and overhung by the occipital pole (fig. 1, D), and curving downwards behind the corpus callosum, it bends forward as the gyrus hippocampi or lower limbic arc (figs. 1, 2, B). Looked at from the base, we see the lower limbic arc separated from the extra-limbic mass by a well-defined fissure — the limbic fissure, which here separates the lower limbic arc from the extra-limbic mass, the latter being still prominent and not concealed from this aspect, as in the rat, where the lower limbic arc extends farther outwards. Extending back from the frontal pole are the olfactory lobes, the outer roots of which (or superficial olfactory medulla) terminate neai- the extremity of the gyrus hippocampi. These two external olfactory roots enclose between them two pyriform grey areas, one on each side, separated by the middle line, bounded behind by the optic commis- sure — the optic nerves lying superficial to them. This grey area is the olfactory field of Gratiolet. Between the olfactory area and the lower limbic arc, a very slight depression indicates the site of a rudimentary Sylvian fissure. Looked at from above, we find the surface of what Broca would call the extra-limbic portion, perfectly smooth, and showing no indica- tions of rudimentary furrowing beyond a very delicate, shallow, linear depression, mapping off" the sagittal region of the brain from the parietal LAMINAR TYPES OF CORTEX. 103 or extra-limbic portion ia the posterior half of the hemispheres. This is the representative of the primary parietal sulcus, which in the Pig, Sheep, and other Gyrencephala, separates the sagittal from the sylvian gyri of the parietal lobe. In the rat no such linear depression exists ; but, this region bordering on the sagittal margin posteriorly, is clearly mapped out by its distinctly pcde aspect as compared with the cortex external to it. The difierent regions which we have now indicated are all dis- tinguished by a type of cortex peculiar to each ; and thus the upper limbic arc, the lower limbic arc, the olfactory area, the extra-limbic or parietal portion — areas obviously differentiated roughly from one an- other by sulci or faint indications of furrowing — all exhibit absolutely distinct types of cortex. But this differentiation does not stop here ; the pale strip of cortex bordering upon the sagittal margin in the rat, although not mapped off by a distinct furrow, has also its own peculiar type of cortex ; and in the rabbit, as we have seen, this region is further differentiated by a linear furrowing. Then, again, the lower limbic arc, if traced backwards, presents us beneath the occipital pole with a further modification, which can only be regarded as a distinct type of cortex. If we add to the above the formation of the cornu Ammonis and of the olfactory bulb, we have presented to us eigh' distinct • types of cortex, not mere fanciful distinctions based upon trivial peculiarities ; but, in all cases, abrujjt transitions from one kind of cortex to another. This divergence in laminar type is peculiarly abrupt in these lower forms of life, the demarcation usually being sharply drawn at the furrows intervening between these regions. In higher animals, and especially in man, no such abrupt demarcation occurs ; distinct transition reg'ions He between either territory, so that the gradual passage from one form of cortex to another is a dis- tinctive element in the evolution of the higher brains * {Brain, vol. i., page 84). The eight laminar types of cortex which are thus distinguishable in these small mammalian brains, we have named as follows : — (1) Type of the upper limbic arc. (5) Modified olfactory type. (2) Modified upper limbic type. (6) Extra-limbic type. (3) Outer olfactory type. (7) Type of cornu Ammonis. (4) Inner olfactory type. (8) Type of olfactory bulb. * In his earlier memoir, published in Strieker's Human and Comparative Histology, as well as in his later views expressed in Psychiatry, Meynert defines but five types of cortical lamination as distinctive of the brain in mammals. We find our- selves unable to agree with Meynert, not only as regards his enumeration of types of lamination, Init in some cases as regards his description of the specific characters of individual types of cortex. lOA LAMINATION OF CEREBRAL CORTEX. On the other hand, we find that Meynert enumerates but five types as follows : — (1) Common type. (3) Sylvian type. (2) Occipital type. (4) Type of cornu Ammonis. (5) Type of olfactory bulb. In addition to these types of cortical lamination we have also to consider the cerebellar cortex, and what is really an ofi'shoot of the cerebral cortex — viz., the retina. If we turn to our outline scheme of the rabbit's brain (Plate i.), we shall find these diverse forms of cortex distributed in the following regions : — (1) The first, or the type of the upper limbic arc, occupies the median cortex of the hemisphere from the frontal pole to the end of the sub-parietal furrow (figs. 1-3, + ); it moreover spreads beyond the sagittal margin, and embraces the pointed frontal extremity of the extra-limbic region at the vertex. (2) The second, or modified upper limbic type, prevails also on the median cortex behind the above type, extending to the occipital pole, but also spreading outwards over the sagittal border to the upper aspect of the hemisphere, where it terminates abruptly at the parietal furrow (dotted area). (3) The third, or OUter olfactory type, characterises the cortex of the (greater segment of the lower limbic arc to its extremity — the gyrus hippocampi (figs. 1, 2, B). (4) The fourth, or inner olfactory type, covers the grey pyriform areas enclosed within the last mentioned and the outer root of the olfactory bulb (fig. 2, dark area). (5) The fifth, or modified olfactory type, occupies the posterior segment of the lower limbic arc, where it sweeps round posteriorly to meet the upper limbic arc. This form of cortex, unique of its kind, is also abruptly limited externally by the great limbic fissure. (6) The sixth, or extra-limbic type, is peculiar to the whole of the extra-limbic or parietal portion of the hemisphere, except the regions already described as presenting a peculiar lamination. Thus it occupies the whole of the vertex except the portion internal to the parietal furrow, and the pointed end of the hemisphere in front, whilst elsewhere it is strictly demarcated from other regions by the great limbic fissure. (7) The seventh, or type of the COrnu Ammonis, characterising the involuted free margin of the cortical envelope, is, of course, con- cealed from view in these aspects of the hemisphere. (8) The eighth, or type Of the olfactory bulb, has its distribu- tion sufficiently indicated by its name (figs. 1, 2, F). Plate VII. Fig. I. X 110 Fk;. 2. CORNU CELLS AMMCJNIS OF YOUNC; RAT. SHOWING PYRAMIDAL OF CORNU, AND GRANULFS OF FASCIA DENTATA. Bale, Sons <& Daniehson, Ltd.,LUh. UPPER LIMBIC TYPES. 105 A brief description of the peculiarities of these cortical belts of nerve cells will be all that is needful for our present purpose. (1) Upper Limbic Type. — The cortical lamination here referred to is illustrated in Plate i. The area it covers is represented in figs. 1-3^ + . It is essentially a four-laminated type ; its first or superficial layer being a light grey belt of delicate neuroglia matrix, with connective elements and their fine prolongations supporting the extremely de- licate subdivisions of the apical processes of nerve cells in the subjacent layers. This layer we term the " peripheral cortical zone" (Plate i.). Next to this succeeds a layer of small pyramidal cells, which, down to the confines of the third layer, remain equable in size throughout; in all respects these elements bear close re- semblance to the upper half of the third layer in higher animals. They diff'er from the human cortex (1) in not, as in the latter, rapidly increasing in size with their depth, and (2) in following immediately upon the peripheral cortical zone with no intervening belt of small oval and angular cells, ^uch as characterises the second layer in man. A few bifurcate cells in sparse detached clumps occur on the outermost confines of this layer, probably rudimentary elements of the second layer of man. Beyond the layer of small pyramidal cells is a pale belt containing the largest cells of the cortex — a pale poorly- celled zone demarcating them from the superimposed layer of pyra- midal cells. These elements are, however, distinguished from the latter not alone by their great size, but by their distribution into confluent groups or clusters, which, as we shall see later on, is a special character of the large nerve cells of the motor cortex. Their apex pro- cess extends right through the pyramidal series into the peripheral zone. We cannot now stop to inquire into their many striking features. Beneath these large cells is a series of fusiform elements similar in all respects to those found in higher mammals. This type of cortex, thepefore, is constituted by (1) A peripheral cortical zone. (3) Ganglionic layer. (2) Small pyramidal layer. (4) Spindle cell layer. (2) Modified Upper Limbic Type.— This form of cortex, like the last is also a four-laminated type. Near the posterior extremity of the corpus callosum (Plate x., fig. 1), the upper limbic arc exhibits the in- tercalation of a series of g'ranule CellS between the small pyramidal and the large ganglionic cells ; but, as we proceed farther back, this belt of granule cells deepens, and, approaching the surface, eventually entirely displaces the small pyramids, and becomes in their place the second layer in this region. The granule-like aspect is due to the relatively large nucleus, as compared with the investing protoplasm : they form a belt of densely crowded elements. The cortex, therefore, I06 LAMIXATIOX OF CEREBRAL CORTEX. of the area represented in Plate x. (figs. 3 and 5, dotted area) is constituted of (1) Peripheral cortical zone. (3) Ganglionic belt. (2) Deep belt of granule-like cells. (-4) Spindle cell layer. (3) Outer Olfactory Type. — Passing now to the lower limbic arc at the base, we tind that the area marked Plate i., B. has a much simpler form of cortex than those hitherto described — two belts of nerve cells only are found in this region subjacent to its outer or peripheral zone. This peripheral zone is specially characterised by the distribution throughout its greater extent of fibres derived from the superficial olfactory fasciculus, which lies embedded in this iirst layer of its cortex ; fibres which ramify at all depths in this layer to unite with the meshwork derived from the apex processes of the cells beneath. Xext to this succeeds a shallow belt of irregular cells, pyramidal, oval, or fusiform, small in size, each with a bifurcate apex process, which immediately undergo rapid subdivision. They are arranged in peculiarly appressed clumps. Then amongst them appear a few large cells of pvramidal contour, which deeper down increase in num- ber and form a distinct belt, in which a few rather large elements are seen. Traced outwards, beyond the limits of the great limbic fissure, these larger elements appear to pass into the ganglionic series. whilst the small clumps of irregular cells pass into the small pyramidal cells of the extra limbic region. This cortex, therefore, comprises (1) A peripheral cortical zone. (2) Dense appressed clusters of small cells. (3) Scanty large pyramidal cells. (4) Inner Olfactory Type. — Covering Gratiolet's -olfactory area'" is a three-laminated cortex, comprising (1) A peripheral zone. (2 > A granule cell layer. (3) Layer of spindle cells. The second layer is formed of cells measuring 9 ,a x 6 /x, with a large spheroidal nucleus, 6 a in diameter; with these are associated numerous minute granules only 5 a in diameter, like the granule cells of the modified upper limbic region. This layer is duplicated in numerous folds, in which the outer layer does not participate. The layer of spindle cells is notable for the large size of these elements ; they are reclinate — i.e., their long axis lies parallel with the surface of the cortex. (5) Modified Lower Limbic Type.— This unique formation, occu- pying the small triangular area, shown in the figure (Plate i., T), is a five-laminated type, the chief feature of which is presented by the peculiar second layer of cells. These nervous elements are more than double TYPE OF CORNU AMMONIS. 107 the size of those occurring in the second layer of the cortex elsewhere ; they are large, swollen, globose, inflated-looking cells, which almost invariably branch from the apex by a bitid or bieorned process. This belt of inflated cells is superimposed on a series of small pyramidal bodies, which succeeds them (Plate x., fig. 2). A pale belt, devoid of nerve cells, follows the latter, and is in turn succeeded by a seiies of spindle cells. To recapitulate, we have here (1) Peripheral cortical zone. (3) Small pyramidal cells. (2) Layer of globose inflated cells. (4) Pale belt devoid of nerve cells. (5) Spindle cell layer. (6) Extra-LimbiC Type differs from that of the upper limbic cortex solely in the intercalation of a belt of granule or angular cells between the small pyramidal and ganglionic series. This form of cor- tex exhibits a very gradual transition to the upper limljic type, and, therefore, presents an exception to the rule of abrupt demarcation shown by other varieties of cortex. The gradual passage of one into another form we shall have reason to refer to later on ; for the present, it will sufl&ce to enumerate the relative layers of this formation. (1) Peripheral cortical zone. (3) Belt of granule or angular cells. (2) Small pyramidal layer. (4) Ganglionic series. (5) Spindle cell series. (7) Type of the Cornu Ammonis. — The cortex of the cornu presents several features common to other regions of the hemispheres : we here have reproduced a peripheral zone to which run the radiate apex processes of underlying cells : then a dense belt of ganglionic cells : beneath which again we trace a spindle-form series of elements. The distribution, however, of these several nervous constituents is so far different as to stamp this type of cortex with features peculiarly its own. Peripheral Zone [Limiting Zone or Molecular Layer).- — This super- ficial layer receives the terminal dendrites ascending from the sub- jacent pyramidal cells of the fourth layer, and the terminal ramifications of axons also derived from subjacent cells — ascending" axons. Apart from these, however, we find, according to Cajal, certain nerve cells •proper to this layer — cells of fusiform or more or less triangular form, with several dendritic branchings, whilst a tine axon ramifies very extensively through the whole thickness of the stratum. Meynert recognised spindle-formed cells in this his " Nuclear layer." The cells whose axons enter into the constitution of this peripheral zone are the nerve cells horizontally disposed within the second layer — the so-called "lacunar layer" of the cornu ; as well as the polymorphic cells of the fifth layer, lying just beneath the belt of pyramidal cells. I08 LAMINATION OF CEREBRAL CORTEX. Lacunar Layer. — This, the second layer of the cornu Ammonis, is formed by (1) Horizontally disposed meduUated fibres, the origin, of which has been traced to the large size collaterals ascending from the axons of the giant pyramids and ramifying across the primary dendrites of the small pyi-amids, which run vertically at this level ; (2) Terminal arborisations from collaterals ascending from the white matter or alveus ; (3) Terminal arborisations of the ascending axons of subjacent cells ; (4) The intrinsic cells of this layer, usually triangular in form, possessed of several dendrites, and an axon which terminates in an arborisation horizontally distributed along this layer, losing itself, as before stated, in the lower region of the peripheral zone. Striate Layer. — This is the part characterised by its radiate appear- ance due to the ascent of the dendrons of the pyramidal cells. Several species of nerve cell are found in this layer, distinguished by the dis- tribution of their axons. All have dendritic expansions, which are distributed to the two upper layers on the one hand, and to the layer of polymorphic cells on the other. All such cells give origin to axons which ramify extensively ; some upwards into the two superficial layers ; others horizontally within the striate layer ; and, lastly, others descending to end in free arborisations around the pyramidal cells and the subjacent polymorphic cells (Cajal). Pyramidal Layer.— These nerve cells, so characteristic of the cortex of the cornu Ammonis, vary much in form in different animals ; they are closely appressed and often appear in several tiers one over the other. More frequently oval or spindle-shaped, they, at times, assume a more spherical contour; but, in all cases, whilst several protoplasmic processes descend from their base, an apical dendron ascends towards the peripheral zone. This latter throws ofi" several collaterals on its ascent through the striate layer, and upon its arrival at the lacunar layer, its primary splitting up into dendrites occurs, and a rich terminal plume results, whose branches extend up to the pial surface of the cortex. Like the corresponding pyramids of the cerebral cortex these dendrites are covered by rough thorny projections. The axon of these pyramids (long known to be continuous with the medullated fibres of the alveus) descends to the white stratum below, throwing off several collaterals in their course, and then, bending at right angles, becomes continuous with the medulla of the alveus — many bifurcating into two branches which run in opposite directions. A very important distinction must be made between the inferior or giant pyramids near the fimbria, and the superior or small-sized pyra- mids beneath the alveus as regards their conformation, relationships, and functional significance. The former are not only larger in size, Plate VIII. X 110 Fig. I. X 110 Fig. 2. Bale, Sons & Daniehson, Ltd.,LUh. TYPE OF CORNU AMMONIS. 109 have thicker dendrons with stunted dendritic arborisations, but also their axons are continuous with the fibres of the fimbria, not with those of the alveus as is the case with the smaller pyramids ; at the same time these axons originate the recurrent collaterals alrt^ady alluded to, which pass through the lacunar layer to ramify over the primary dendrites of the smaller pyramids. A further distinction is established by the fact that the axons of the granules of the fascia dentata (moss fibres) are brought into close velationshii)S with the dendrons of these giant cells alone and not with those of the smaller cells. Folymorjihic Cells. — The irregular and fusiform cells beneath the pyramidal layer constitute the "polymorphic layer "of Cajal — the old " stratum raoleculare " of Kupffer. Formerly classed as simple spindle- shaped cells recent authorities describe certain notable features as presented by them. They may be divided into three groups — all the cells of which are dendritic — these dendrites ramifying amongst the protoplasmic processes and the collaterals from the axons of the pyramids. These groups are characterised as having — (1) Ascending and recurrent axons ; (2) Horizontal and plexiform axons ; (3) Deep spindle cells with ramifying axons. In the first group of ascending and recurrent axons this process ascends through the radiate stratum, throwing off occasional collaterals in this course, some of which pass up into the lacunar and peripheral layers, whilst the main axou curves downwards again and, passing into the pyramidal layer, forms a rich terminal plexus of fibres around these cells. In the second group, the axon passes off' horizontally and divides into numerous branches after the manner of Golgi's sensory cells. From these originate collaterals, which ascend to the pyramidal layer and surround its cells with a plexus of branches mingling with the similar plexus from the recurrent axons. In the third group, the cells are spindle shaped; they have long been recognised in the neighbourhood of the alveus, but only recent research has revealed the fact that they possess axons which ramify extensively in this stratum and ascend upwards to a higher level. Besides these three, Cajal describes a cell which gives off" an ascending axon terminating by ramification in the lacunar and peripheral layers, but having no recurrent branch. We need scarcely include here certain cells, as Cajal does, which, together with others in the stratum radiatuni, are admittedly similar in type to the pyra- midal cell, and are really dislocated cells not special to these layers, any more than we should speak of those cells found in the lowermost stratum of the peripheral layer which are really elements dislocated from the cells of the second layer of the cerebral cortex, and which are also seen in the region of the fascia dentata. 1 I o LAMINATION OF CEREBRAL CORTEX. Medullated Layer or Alveus. — This deepest layer, bounded centrally by the ependyma of the ventricles, is largely constituted by the axons of the pyramidal cells, many of which, as before stated, bifurcate into a thick and a slender process running in an opposite direction. These medullated fibres throw upwards collaterals which ramify in the upper layers of the cornu. The Fascia Dentat'i. — In this region we have to distinguish three layers, viz. : — (1) Peripheral or molecular ; (2) Granule layer or small pyramids ; (3) Polymorphic cells. Molecular Layer. — This, like the corresponding layer of the cortex generally, and that of the cornu Ammonis formation proper, receives the terminal plume of dendrites given off from the subjacent cells, and which are here brought into relationships with the axons of certain nerve cells peculiar to this layer. These cells are distributed in a double series, superficial and deep (Cajal) ; they are ovoid, stellate, or spindle-shaped — the deeper cells being the larger and more freely supplied with protoplasmic processes. Both have an axon thicker in the deeper cells, and ramifying to great distances in a horizontal direction, but much finer in the smaller cells and far more restricted in range. Stratum Granulosum. — The elements of this layer are very small, densely appressed, forming a deep stratum of minute ovoid, pyramidal cells, throwing off numerous protoplasmic processes towards the peripheral zone : whilst its axon, descending through the subjacent layer of cells (polymorphous), gives origin here to numerous delicate collaterals, which ramify amongst the polymorphic cells. Continuing its course, it bends as a knotty, non-medullated fibre along the length of the cornu, upon the body and dendrons of the giant pyramids. These axons from the granules appear similar in all respects to the so-called "moss fibres" of the cerebellum. Cajal affirms that such moss fibres never trespass on the region of the alveus or the lacunar layer of the cornu proper ; but are invariably confined to the giant pyramids, their moss-like bunches resting in close contact with these cells. It will be observed that these axons of the granules form a terminal arborisation, bringing tliem into relationship with the giant pyi'amids and their dendrons, just as the collaterals ascending from the axons of the latter ramify over the dendrites of the smaller pyramids of the cornu. Pyramidal cells of notable character occupy the upper regions of this granule zone. They have an apical process (protoplasmic), which ascends to be distributed in branches to the molecular layer ; several TYPE OF CORNU AMMONIS. Ill protoplasmic branches given off from the base of the cell, and an axon "which running horizontally immediately above the granule layer sends downwards numerous collaterals to envelope the granules in a rich plexus of branches. Fig. 14. — Cerebral cortex : granules of fascia dentata — Cornu Ammonis (rabbit). Stratum of Polymorphic Cells. — Irregularly shaped cells are seen beneath the granule layer, throwing off protoplasmic processes in several directions, and with an ascending axon passing upwards through the granule into the molecular layer, there to bifurcate and ramify in extended horizontal planes. Some of these cells are stated, however, to give off an axon which runs horizontally along the summit of the granule zone, taking part with the pyramidal cells in the formation of the intergranular plexus already described. Cajal, moreover, describes in this layer nerve cells similar to the sensitive cells of Golgi ; and others with a descending axon continued into the alveus. Lastly, immediately above the molecular layer of the cornu proper, where it is concealed by the fascia dentata, are large fusiform cells, whose axon can be followed into the alveus. The Olfactory Bulb. — The cortex of the olfactory bulb consists of a superficial and a deep fibre-tract ; the former non-medullated, connecting the nervous apparatus of the Schneiderian membrane with the olfactory bulb ; the latter, medullated, and connecting the bulb I I 2 LAMINATION OF CEREBRAL CORTEX. tlirough the olfactory tract with the centric termini in the cerebral cortex. These two tracts of nerve fibre are brought into relationship through the medium of the intervening layers, three in number, of nerve cells and fibre systems. We have thus, from without inwards, five distinct strata : — (1) Flexus of peripheral nerve fibres. (2) Layer of olfactory glomeruli. (3) Molecular layer with small nerve cells. (4) Layer of so-called " mitre cells." (5) Layer of granules and medullated nerve fibres. The superficial layer of the bulb is formed by non-medullated fibres which, arising from bipolar nerve cells of the Schneiderian membrane, pass through the cribriform plate to form a densely woven plexus, on the surface of the bulb ; the fibres eventually terminate by a free arborisation in the interior of the glomeruli which form the character- istic feature of the second layer of the bulb. These spheroidal bodies were first observed by Leydig in the fish, and constitute Meynert's "stratum glomerulosum.' Each glomerulus consists almost entirely of the interlacements of the olfactory fibres derived from the super- ficial layer on the one hand, and, upon the other, of those derived from the cells of the fourth layer: whilst a few nuclear-like cells are found within and around the glomerulus. The silver-chrome method shows that the non-medullated fibres on entering the glomerulus split up into arborisations of excessively flexuous, thickened and varicose fibrils (Cajal), which never again emerge from the glomerulus. Here thev intertwine, but do not inosculate, with a rich arborisation of dendrites derived from a process of the cells of the fourth layer — the so-called "mitre cell." Immediately beneath the glomerular layer is a stratum of finely grani^lar material, in which are imbedded small fusi- form nerve cells which throw off a coai'se protoplasmic process (dendron) towards the glomerulus, in the interior of which it loses itself as a dendritic arborisation. An extremely fine axon passes from the cell deeply towards the granule layer of the bulb and, bending at right angles, passes backwards amongst the medullated fibres of the tract. Layer of Mitre C'eZ^i\ —Between the preceding and the deepest layer of the bulb lies a stratum of cells, remarkable for their size, configura- tion, and relationships. These elements are usually of large size, of mitre-like form, throw off lateral processes ramifying through the neiofhbouring molectilar layer, whilst, from their lower aspect descends a coarse dendron, which, entering a glomerulus, breaks into a terminal ramification of dendrites interlacing with the olfactory nerve fibres, as already noted. From the deep aspect of the mitre cells a stout axon passes upwards to the granule layer, and here, bending backwards, THE RETINA. I I 3 becomes continuous with a medullated fibre of the olfactory tract, for distribution to the cerebral cortex. In this course through the granule layer, the axon throws downwards several vertical collaterals, which ramify in the molecular layer ( Van Gehichten and Martin). Layer of Granules and Medullated Fibres. — This, the deepest layer of the grey matter of the bulb, is of considerable thickness, and is formed by granules similar in form to those of the cerebellar granule layer, but separated into compact groups by the passage of fasciculi of medullated fibres. According to Cajal and others they diS'er morpho- logically from the granules of the rust-coloured layer of the cerebellum ; these possess a distinct axis-cylinder, while those of the olfactory bulb are devoid of such.* These spherical or angular cells throw off two processes ; one, downwards to the mitre layer, where it ends in a terminal plume in close contiguity to the lateral dendrons of the mitral cells ; the other, far less conspicuous, passes inwards to ramify amongst the granule groups at a deeper site. The terminal dendrites of the peripheral plume are thickly beset with minute spiny projections along their course, like the similar formations along the terminal plumules of the pyramidal cells of the cerebral cortex. The pePiphePal dendron is always present ; the central dendron may be insignificant or altogether wanting {Pedro Ramon). Golgi has, moreover, described large sized stellate cells sparsely scattered in the granule layer, which, besides protoplasmic processes, possess an axon which (according to Cajal) always ends in a rich arborisation within the molecular layer of the bulb. The medullated fibres found in this deepest layer of the bulb consist very largely of the axons derived from the mitre and small fusiform cells ; but also embrace medullated fibres which pass from the brain to the bulb (centrifugal), and end in free ramifications in the granule layer. The Retina. — ^The nerve elements of the retina consist of a super- ficial layer — the well-known rods and cones ; of an outer and inner granule layer, separated by outer and inner plexiform relational fields ; and of an innermost layer of ganglionic cells and optic nerve fibres. The rods and cones of the superficial layer are morphologically continuous with the elements of the outer granule layer, but are usually, for convenience, described separately. The inner granule layer differs essentially from the outer not alone in the conformation of its constituent elements — the granule cells, but in the presence of two other nerve elements — the horizontal cells and retinal spongio- blasts : whilst the outer granule layer is a comparatively simple * This opinion, which has given rise to the doctrine of "amacrine cells," or nerve cells without axis-cylinders, appears to us to be satisfactorily confuted by Dr. Hill, who has discovered the axon and figured it. See "Notes on Granules," by Alex Hill, Brain, vol. xx., p. 125. 8 I 1 4 LA^illNATION OF CEREBRAL CORTEX. structure — the inner is much more complex and may be subdivided into three layers : — (a) Superficial horizontal cells ; (b) Bipolar granule cell ; (c) Spongioblasts of the deepest stratum. Lastly, the inner plexiform or relational field is one of very great complexity, embracing, as it does, the ramifying branches of the bipolar granule cells, the dendrites of the ganglionic series, and the branches of the retinal spongioblasts. RefAnal Rods. — The rods are cylindrical bodies, fine in mammals and nocturnal bii'ds (in man 50 /^ to 60 'i, long by 2 ,a thick, according to Max Schultze) ; but of large size in Batrachia, diurnal birds and fishes (Cajal), consisting of an inner segment or body, and an outer segment perfectly cylindrical and unpointed. Unlike the cones they are not attached directly to the body of the granule cell of the next layer, but through the medium of a peripheral process extending from the latter ; so that the granule cells of the rods resemble a bipolar ganglion cell, ■of which the finer or centric process extends to the outer plexiform layer. Here this process ends in a small spherical swelling embraced by the dendritic peripheral branch of the cells of the deeper granule layer. The rod fibres are always finer than the cone fibres. Retinal Cones. — The cone, always shorter than the rod, consists of a flask-shaped body or inner segment, about 6 /i in thickness, terminated in a pointed conical extremity ; and an outer segment, which is more strongly refractile than the inner segment. Schultze has described a longitudinal striation of the outer segment of both rods and cones in fishes, amphibia, and mammals, including man ; and likewise a similar striation of the inner segments in man and mammals. The body is distinctly continuous with the granule cell of the cone — an ovoid body with large nucleus, from which a centric process descends to the outer reticulated or plexiform layer, where it ends in a conical swelling with a. few lateral free fibres. In reptiles the rods are absent, the cones alone being found. It is interesting to note that Krause has shown that the rods and cones both persist after section of the optic nerves, their appearance suggesting no degenerative change. The external limiting membrane forms a natural boundary between the rods and cones and their subjacent granule cells. Outer Plexiform Layer. — In this layer the protoplasmic branchings from the peripheral ends of the bipolar cells take part — some inter- lacing with the termini of the centric branch of the cone cell ; others receiving between them the spherical globules of the centric branch of the rods. Bamifying fibres extend also into this layer from the horizontal cells of the subjacent stratum. Plate IX X 360. Fig. I. Layer of Rods and Cones. External Granule Layer. Outer Plexiform Layer. \ Internal Granule Layer. Internal Plexiform Layer. Ganglion Cell Layer. ..Fibres of Rods. -Fibres of Cones. -Interlacement of inner Granules ivith termini of Rod and Cone Fibres. -Bipolar Cell connected with Rods. -Bipolar Cell connected 7uitk Cones. SpongiobUists. Field of Interlacement of Ganglion Cells with inner Granule Elements, and Spongioblasts. Ganglion Cells of Retina. jOptic Nerve Fibres. V\r,. 2. SCHEME OF RETINAL LAYERS IN MAMMALS. Bale, Sons £ DanUlsson, Lld.,Ltlh. THE RETINA. II5 Inner Granule Layer. — Three subdivisions are described by Cajal — {a) horizontal cells ; (h) bipolar granules ; (c) retinal spongioblasts. The former consist of small and large horizontally disposed nerve cells, of which the large-sized are the most internal ; both throw off numerous protoplasmic processes and a long axon ; the axon from the smaller and more superficial cell is very fine, giving origin to several short ramifying collaterals. In the deeper cells this axon is much larger, and terminates, after a lengthened course, in a rich arborisation in connection with the spherules of the rods {Cajal). These ramifica- tions add, therefore, very materially to the plexiform meshwork of the third layer. The bipolar cells of this layer are fusiform in contour, give off from their peripheral pole a tuft of protoplasmic branches, which spread laterally into the outer plexiform layer ; whilst the fine centric branch descends to different levels of the inner plexiform layer, ending in a plume of free branches in close relationship to the ascend- ing dendrites of the ganglionic cells. Cajal specially distinguishes these bipolar cells into such as have a peripheral ascending plume of branches, and those with a flattened plume : the former being dis- tributed to the spherules of the rods ; the latter to the branches of the cones. The terminal arborisations of the centric or descending branch of these, bipolar cells meet the dendrites of the ganglionic cells at different levels of the inner plexiform layer, thus dividing it into five or more strata. Retinal Spongioblasts. — These cells are remarkable in the fact that they possess no axon. They are disposed in the lowermost stratum of the inner granule layer, in immediate contact with the inner plexiform layer and all their processes are directed centrally. Schafer doubts their nervous nature.* Inner Plexiform Layer. — We have, therefore, immediately beneath the inner granule layer, a very rich relational field, in which three distinct retinal elements are brought into immediate apposition, viz. : — (a) Bipolar granule elements ; (6) Retinal spongioblasts ; (c) Ganglionic cells. This elaborate system of fibres is known as the inner plexiform layer. Ganglionic Cells. — These cells, like those of other parts of the nervous centres, vary much in size, from 15 /a to 30 /x in diameter in the fresh state ; they are flask-shaped or ovoid in contour, usually devoid of pigment, and possess a large nucleus with a prominent nucleolus. They give origin to a peripheral and centric process — the former directed into the depths of the inner plexiform layer, as a dendritic arborisation, extending laterally in the horizontal plane as far as one or other of the five stratifications of this layer, where they * "The Nerve Cell considered as the Basis of Neurology," Brain, 1893, p. 138. = Il6 LAMINATION OF CEREBRAL CORTEX. blend with the corresponding terminal fibres of the retinal spongio- blasts and the bipolar granule elements. The centric process is really its axon, '-which becomes continuous, as an optic nerve fibre, with the deepest layer as it courses towards the optic nerve. Oajal divides these ganglionic cells into three series : those whose dendrites are restricted to one stratification only ; those distributed to two or more; and those which observe no such law of stratification,. but are equally distributed throughout the whole depth of this plexi- form layer ("cellules unistratifiees, multistratifiees, et diffuses"). So far, therefore, from the optic nerve fibres being in direct continuity with the special visual elements of the retina — the rods and cones — we find, according to the researches of Cajal, Van Gehuchten, and others, that there are two breaks in this course : the first, betwixt the dendrites of the ganglionic cells and the bipolar elements ; the next^ betwixt the peripheral ofishoots of the latter and the termini of the centric branches of the rods and cones respectively. This is in accord with the results of physiological and pathological teaching, for we know that section of the optic nerve fibres entails fatty degeneration of the ganglion cells of the retina {Erause), whilst the rods and cones- escape intact ; moreover, in blindness from optic nerve atrophy and in glaucoma from intraocular pressure the ganglion cells suflPer in like manner. The Cerebellar Cortex — Lamination. — The cerebellar cortex may be regarded as two-laminated, a superficial or external, commonly termed "the molecular layer," and an internal or granule layer; whilst upon the confines of both is a series of voluminous cells peculiar to the cerebellar structure, termed the "cells of Purkinje." The leaflets or foliola of the cerebellum are mostly disposed trans- versely to the antero-posterior axis of the brain, and sections may be taken either in the direction of the plane of these lamellae— z.c.^ frontal sections, or across their plane antero-posteriorly — i.e., sagittal sections. The microscopic appearance is very diflerent in these two cases. In the antero-posterior or sagittal section, the dendritic expansions of the cells of Purkinje attain their maximum development and are displayed to the greatest advantage, whilst the nerve cells peculiar to the superficial layer, with their neuraxons, are also seen following the same plane. In frontal sections, on the other hand, the cells of Purkinje are seen as it were in projile, their dendritic branchings, insignificant and flattened laterally, would indicate a somewhat fan-shaped conformation in the direction of the sagittal plane. At the same time, we lose the characteristic branchings of the nerve cells peculiar to the molecular layer, and, in lieu thereof, we see a horizontal striation of this layer, due to bifurcation of the axons of . the granule cells of the deeper layer, and which, being cut across in ■)^ A Plate X. Fii.2. '^V A '<^ • ^ "MoSifiea oifactoi^- type" r/ '^^- ■• f Iiom posterior extremity of I^wex limbic j^-«y- " Brain of Rattil xilO Fii.l. "SensOTy Cortex' Prom tlie "Modified upper liml)ic' type m Brain of Rab^iil. x 214- CZ3 i ; Brain of H^fb t Ivlesij.l ^Tpe^t > asal aspect Distritixtion of ^^a^lo.J;s Tiated types cf Cort. e iDaliieUson.ltd , Sculp THE CEREBELLAR CORTEX LAYERS. I I 7 sagittal sections, appear simply as innumerable dots scattered through- out the field at this level. Superficial Layer of Cortex. — The external layer immediately beneath the pia has, in the embryo, a structure quite peculiar to itself, of which, however, all trace is lost in later life ; it forms a peripheral zone (not as yet encroached upon by the dendrons of the cells of Purkinje) and consists, according to Cajal, of — (1) A superficial series of granule-like cells placed vertically to the surface ; (2) A deeper layer of horizontally disposed bipolar cells lying next to the molecular layer. In the fully developed cortex, however, the elements demanding attention are : — (a) The intrinsic nerve cells or elements peculiar to this layer; {b) the dendrons and collaterals of the cells of Purkinje, together with the nerve fibres extending from undeidying structures ; (c) non-nervous or connective elements. (a) Nerve Cells of the Superficial Layer. — These are irregular, stellate, or polyhedral cells, most numerous in the deeper realm of this layer, having their long axis disposed antero-posteriorly — i.e., in the plane of the dendrons of the cells of Purkinje. Numerous long protoplasmic processes are given off" by these cells ; but, in particular, a fine and greatly elongated axon, which also courses along the plane of these same dendrons. Along this course, wdiich is more or less parallel to the surface of the cortex, vertical ofifshoots or collaterals descend to the cells of Purkinje, and, becoming coarser near these cells, split up into a terminal ramification, which forms a basket-work* embracing the body of the latter as far as the commencement of its axon. Contracted over the lower pole of the cell, this basket-work extends brush-like a short distance along the neuraxon, where it is still devoid of a medullary sheath. It is remarkable how this terminal basket-work escapes staining in aniline preparations, where the cells of Purkinje are yet admirably displayed. The cell with enclosing basket-work is named by KoUiker a " basket cell " [Korhzellen). Close to the pial surface of the cortex a few stellate cells, much smaller, are found ; their protoplasmic processes are richly developed, and the axon, which runs antero-posteriorly, ramifies extensively, but its destination is imknown. (6) Cells of Purkinje. — These flask -shaped cells, so characteristic of the cerebellar cortex, throw ofi^ from their upjier or outer pole a single or double dendron (in the latter case, giving the cell a somewhat horned appearance), from which arises a luxuriant arborisation by frequent dichotomous division as far as the surface of the cortex. These dendrites neither anastomose with each other nor with the * Endkoerheiiy Faserkorhe, Fa-'icrpiimtl of Kolliker. I 1 8 LAJMIXATIOX OF CEREBRAL CORTEX. neighbouring cells ; they are strictly terminal and often turn back upon themselves. The dendritic system is spread out along the antero-posterior plane, and the branches are, moreover, thickly studded with minute thorny processes like the corresponding projections (epines) upon nerve cells in other regions {Van Gehuchten, Cajal, Betzius). From the lower or internal pole of the cell descends the neuraxon, which, passing through the granule layer of the cerebellum (becoming medullated in this course), enters the central medullated core of the lamina, probably passing out of the cerebellum to distant centres. Shortly after its origin the axon throws off some two or three delicate collaterals, which, passing upwards into the peripheral layer, end by a terminal ramification in contact with the lower dendrites of the neighbouring cells of Purkinje ; thus, according to Cajal, they ensure a certain degree of functional solidarity. Besides the structures just described the molecular layer displays in frontal sections [i.e., sections taken along the course of the lamincB) a distinct striation due to fibres running at all depths parallel to the surface, which, on close examination, are seen to arise in all cases from the T-shaped bifurcation of numerous vertical fibres passing upwards into this layer from the stratum of granules below. These delicate fibres, which are really the axons of the granules of the cerebellum, give off no collaterals, pass horizontally through the dendritic expansions of the cells of Purkinje, and thus bring the granule layer into intimate connection with those arborisations. They are stated by Cajal to end after a lengthened course in a free and varicose thickening near the white matter of the lamina. If we refer to frontal sections as the " plane of lamination," and to the sagittal sections across a lamina as the "plane of arborisation," i.e., of the dendrites of Purkinje, then it may be stated that the cells of Purkinje are brought into intimate functional relationship along the plane of arborisation by the axons of the cells of the peripheral layer, and the basket-work resulting from their collaterals ; whilst they are bound in functional relationship along the laminar plane by the parallel T-shaped axons arising from the granules of the second layer ; and at the same time their functional co-operation may be established by the recurrent collaterals arising from their axons, and ramifying in the proximity of the lower dendrites of the neighbouring cells of Purkinje. A further relationship, however, is established between the cells of Purkinje and certain coarse fibres ("fibres grimpantes " of Cajal) which ascend from the central medulla through the granule layer, and form a dense plexus upon the primary and secondary dendrites of these cells, just as the nerve cells of the peripheral layer form the basket- work enveloping the body of the cell. The peripheral origin of these fibres is unknown. THE CEREBELLAR CORTEX LAYERS. I I 9 (c) Non-nervous or Connective Elements. — These are of two kinds : — (1) A large irregularly shaped dendritic cell, lying in close proximity to the cells of Purkinje, which throws off, in the granule layer, a number of thick short processes and several long vertical branches. The latter pass upwards into the peripheral layer, each fibre ending immediately beneath the pia in a small triangular or conical thicken- ing ; these fibres have long been known as the fibres of Bergmann. (2) Deeper in the granule layer and extending into the medullated centres are numerous stellate cells in no wise differing from the stellate glia cells common to the nervous centres at large. Granule Layer of the Cortex. — -Beneath the peripheral layer is a compact stratum of small, nearly spherical nerve cells, possessing very little protoplasm, and throwing off on all sides some few short pro- cesses, which end in miniature arborisations around the bodies of the neighbouring granules. From the cell body or one of its processes arises an extremely delicate axon, which passes vertically upwards, and, at varying levels in the outer zone, bifurcates into the T-shaped fibres which have already been alluded to in this stratum of the cortex. Golgi and Cajal also describe large stellate cells as occurring sparsely in the granule layer, with extensive protoplasmic processes spreading in all directions — often far into the peripheral layer, and with an axon which, passing downwards into the granule layer, ramifies to an extraordinary extent amongst these granules, ending, according to Cajal, by free varicose extremities. Moss Fibres of the Cerebellum. — In the granule layer certain notable fibres, first described by Cajal as moss fibres, appear ; they ascend from the medulla as coarse medullated fibres which, on entering the granule layer, split up and ramify extensively among the granules, presenting here and there along their course peculiar rough knotty thickenings or moss-like growth, and do not extend beyond the granule layer. These fibres are presumed to bring the granules of the cere- bellum into relationship with distant nerve centres, and are suggested as the possible central nerve termini of the cerebellar tract {Cajal). Our review of the foregoing types of cortical lamination in the mammalian brain prepares the way for certain deductions which have an important bearing upon the physiology and pathology of the cerebrum. In the first place, let us note that the simpler formS Of cortex are confined to the lower margin of the cortical envelope, where it folds round the cerebral peduncle at the base — the COmu Ammonis, the lower limbic lobe ("outer olfactory type"), and also the olfactory area of Gratiolet. The more complex form of cortex, however, spreads over the upper limbic arc and the whole of the extra limbic region of I20 LAMINATION OF CEREBRAL CORTEX. the hemisphere. It is these more complex forms of cortex which concern us chiefly ; they comprise in man the extensive areas at the vertex and the whole convoluted surface of the hemispheres, as seen from above. Now, in studying the small brain of the rodent and higher animals, we find structural modifications in the cortex of this region, which appear to foreshadow the divergences observed in man. Thus, if we examine successively the cortex at different points from within outwards in a vertical section through the hemisphere, passing through the Sylvian depression, we find that — (a) The first layer of the cortex is deepest at the sagittal border, and steadily diminishes in depth as we proceed outwards towards the limbic fissure ; (6) The second layer of small pyramidal cells increases rajndly in depth and in loealth of cell-structure in a reverse direction — i.e., from within outwards ; (c) The ganglionic series of cells (which assume thick clustered nests in the upper limbic arc and over the vertex bordering on the sagittal fissure), gradually loses its confluent tendency and becomes spread out in isolated units (" solitary type ") as we approach the limbic fissure externally. On the other hand, if we examine similarly a vertical section taken through the posterior moiety of the upper limbic arc (Plate vi.) we find that — {d) The intercalated series of granule cells increases in richness of elements and depth of formation as we proceed outwards to the lateral aspects of the hemispheres, and backwards to the occipital pole ; and reaching the limbic fissure terminates abruptly, whilst the other layers pass on uninterruptedly. If we now examine vertical sections of the hemisphere in the antero-posterior plane, we find that — («) The outer layer (peripheral zone) progressively diminishes in depth from the frontal to the occipital pole ; (/) The small pyramidal cells of the second layer diminish in size in the same direction ; (g) The granule or angular cells intercalated in the five-laminated cortex increase in richness conspicuously towards the occipital pole ; {h) Lastly, the ganglionic series, which near the frontal pole forms a deep layer rich in cell elements, thins out considerably backwards into a laminar or " solitary " formation ; but, at the extreme occipital pole, these cells again form a somewhat deep belt with granule cells superimposed. The obvious deductions to be made from the foregoing are that certain elements preponderate in certain fixed areas of the cortex, and that the development of certain layers appears to exclude that of another series. Thus the frontal pole and frontal extremity of the upper limbic arc are especially characterised by the preponderance of DISTRIBUTION OF THE GANGLIONIC CELL. 121 the g^ang'lioilic series, which accumulates here in rich clustered groups ; towai'ds the Sylvian border this element is insignificant, and it is the small pyramidal layer which here prevails. Towards the occipital pole mesially (" modified upper limbic type ") the granule cell att-ains like importance from its notable wealth of elements and its more or less complete exclusion of the small pyramidal series; whilst outside this formation, in the extra limbic cortex, the intercalated granule belt is a notable feature, accompanying a corresponding impoverishment of the small pyramidal and ganglionic series. A certain relationship also would seem to exist between the depth of the first layer or peripheral zone and the ganglionic series of cells ; since it notably diminishes in depth as these elements thin out into the solitary type of arrangement, and this despite the marked increase in the small pyramidal series above. This mutual dependence seems to us explained by the fact that the apical processes of these large elements pass up into, and terminate in, this peripheral zone, so that any regional difference in the depth of the outer layer will be dependent on the greater or less development of these ganglionic cells. It must be borne in mind, however, that the average depth of the first layer increases in lower mammals and becomes shallower as we rise to the more highly organised brains — a fact which does not militate, as might at first appear, against the preceding conclusion. In the lower mammals, the absolute and relative increase in the depth of this outer layer probably means a large preponderance of the connective over the nervous element.* Regional Distribution of the Ganglionic Cell.— Attention was first directed to the peculiarly clustered arrangement of these cells in the cortex of man and the higher apes by Professor Betz,t who denominated them "giant pyramids," and suggested their probable motor signification from their form, arrangement, and connections. Subsequent research appears fully to confirm the conclusion arrived at by Betz,J and it becomes, therefore, important to indicate the regional distribution of these elements. We find that this series of cells in man * See upon this point, Meynert, " Brain of Mammals," Syd. Soc, p. 383 ; also, Brain, vol. i., p. 358. t " Anatomischer Nachweis zweier Gehirncentra," Prof. Betz, Ctntralhlatf f. d. Med. Wisseiisch. , Aug., 1884. t It is true that Meynert would dispose of the assumed significance of these cells on the ground that their large size depends on the distance which their apical process has to traverse in reaching the outer laj'er, and their gradual increase iu dimensions being, as he states, proportionate to this distance. The "gradual increase in size" alluded to proves to us tliat Meynert lias failed to identify the elements I'eferred to — probably mistaking for them the larger pyramids ; and finally his argument falls to the ground when it is seen that the second layer of tlie "modified lower limbic type" contains larger elements tlian any of its subjacent layers. 122 LAMINATION OF CEREBRAL CORTEX. and the higher mammals (Pig, Sheep, Dog, Cat, Ape, and Man) assumes- in separate regions of the cortex a different arrangement, which we have termed the clustered or nested and the laminar or solitary arrangement* — the former showing these large cells aggregated into- distinct oval clusters stationed at intervals apart — the latter ap- proaching the arrangement of these cells universally met with at the base of a sulcus, viz. : — solitary cells, stationed like sentinels wide apart, showing no tendency to grouping beyond two or three at most in certain exceptional areas. In lower mammals (Rabbit and Rat), these discrete or distant clusters do not appear ; but what we take to be the homologue of this series forms COnfluent grOUps — the nested ar- rangement being scarcely indicated, and a deep and dense formation replacing the latter. As already observed, however, these confluent groups thin out, in certain regions, into linear file, assuming the laminar or solitary arrangement. The cells of this series in these lowly-organised brains are peculiar in their extremely elongate pyra- midal or fusiform contour, and approach in this respect the form of the larger pyramids in the human cortex rather than the configuration of the motor cell. As we pass from the confluent groups of elongated elements in the Rodent to the more specialised areas of higher mammals, we find that — (1) The cells become less elongate, more swollen, and irregular in contour ; (2) Their groupings become more and more discrete ; (3) The individual groups grow larger in size ; (4) The clustered arrangement occupies a wider range of cortex. In Plates i. and v. this series of cells is richly represented; they are densely congregated towards the margin of the hemisphere, and thence^ continued to the limbic fissure, occupy the whole area embraced by Nos. 7 and 9 in Terrier's work.f Further back, however, this layer diminishes in depth and in wealth of cells, except at the exposed margin of the hemisphere, where it still remains a rich formation ; beyond the margin and over the extra-limbic region, as far as the limbic fissure, the cells rapidly thin out into a simple linear series, and the five-laminated cortex appears. Still further back the series, in like manner, thins out into a mere insignificant formation — yet always most richly developed along the sagittal margin of the hemisphere. Plate v., fig. 1, represents the arrangement of the ganglionic series in the pig, the regional distribution of which is almost identical in formation with that of the sheep. For both these animals, it may be stated that a five-laminated cortex, with clustered cell- groups, spreads over the anterior half of the upper limbic arc (which • "Comparative Structure of Cortex Cerebri," Trans. Roy. Soc, part i., 1880. Functioiis of the Brain, second edition, p. 259, fig. 78. DISTRIBUTIOX OF THE GANGLIONIC CELL. 123 in these animals becomes superficial on the upper aspect of the hemi- sphere) over the frontal pole and along the first (or Sylvian) and second parietal convolution. Between these tracts is embraced the area of the third and fourth parietal convolutions, which have a six-laminaied cortex and a distinctly solitary arrangement of these cells. If we examine the regional distribution in the cat, the anterior portion of the upper limbic arc in front of and above the crucial sulcus; the frontal lobe; the first parietal, or Sylvian; and the anterior extremity of the fourth parietal or sagittal convolution, will all be found to exhibit the laminated cortex and nested cells ; yet the formation, excessively rich in the sigmoid gyri around the crucial sulcus, becomes much poorer in other regions. The six-laminated type extends over the whole extent of the upper limbic arc, behind the crucial sulcus, as far back as the retro-limbic annectant. The distribution of these nested groups of ganglionic cells in the ocelot, reproduces, in fact, very nearly the arrangement met with in the cat. The distribution of the same formation in the Barbary ape fore- shadows the arrangement which pertains to the more highly-developed cortex of man. It will be observed from the foregoing remarks that the CPUCial sulcus in all these animals forms a distinct limit to two types of lamination — peculiar to the vertex — the five- and the siX-laminated types, and that this distinction is continued upon the mesial aspect of the hemisphere into which this sulcus extends ; that, similarly, at the frontal pole of the hemisphere, the vertical sulcus, regarded by Broca as the representative of the fissure of Rolando, also separates an inner or five-laminated from an outer or six-laminated cortex ; whilst the first parietal or Sylvian convolution in the pig and sheep partakes, in front of the Sylvian fissure, of the five-laminated type. In 1882,* after a minute enquiry into the cortical envelope of the brain in mammals, the author had reason to express himself as follows : — " The more fully I investigate the minute structure of the cortex and its deep connections, the more forcibly am I impressed with the belief that the various fissures and sulci are not mere accidental productions,! but have a deep significance of their own, dividing off the cortical superficies into morpliologicaUy, if not physiologically, distioict organs. Hitherto the fissures and sulci which I have found to be boundary lines of distinct cortical realms are the followinjr : — * 0]}. cit., p. 724. t That is, the result of pressure merely during the development of the cranial arch. 124 LAMINATION OF CEREBRAL CORTEX. "(1) The limbic fissure. (4) The superior parietal sulcus. (2) The infra-parietal sulcus. (5) The inter-parietal sulcus. (3) The crucial sulcus. (6) The olfactory sulcus. (7) The fissure of Rolando." Contrasts between the Brain of Man and of Lower Mam- mals. — When we contrast the cortex of the human brain and of the ape with that of the mammalian series below these types, certain strongly-marked resemblances in intimate structure, as well as equally notable divergences, present themselves. With respect first to the resemblances, it is to be noted that the various types of cortical lamination described in the lower mammals are reproduced in the brain of the ape and man ; and that the several layers maintain the same relative position throughout their depth, except where in certain cases a layer is wanting, or a new layer is interposed. Again, the individual elements constituting these layers — the granule cell, the angular cell, the spindle, the pyramidal element — although differing somewhat in dimensions and general contour, are yet sufficiently alike for their identification apart from their mere position in the cortex. In the next place, the lower limbic margin of the cortical envelope always presents the simpler forms of cortex ; while, towards the vertex and mesially both towards frontal and occipital poles, the more complex forms of cortex prevail. Another striking resemblance occurs in the distribution of these laminar types — that characterised by the g'ranule cell predominating towards the OCClpltal polC ; that of the five- laminated type being especially developed towards the frontal pole : with this there is associated finally the gradual diminution in size of the one element towards the temporal and occipital lobe, and the increased dimensions and richness of formation of the other element in the same direction. These are some of the more striking resemblances presented between the cortex cerebri of man and that of the lower mammals. As to the divergences presented by these structures, we are early struck by the fact that the abruptness Of transition from one to another type of cortex, seen, e.g., in the rodent, is not a feature in the human brain ; in fact, transition-realms invariably intervene betwixt different types of lamination. The one fades into the other form so gradually that a line of demarcation can rarely be drawn. Thus, the five-laminated cortex characterising the " motor area " of the human brain affords no abrupt transition into the six-laminated cortex lying external and posterior to it ; a mixed type intervenes, to which we have applied the term of " transition-realm." In the second place, the cells which we have ventured to term '■'■ motor " in the fourth layer of the human cortex, differ from what we Plate XL Fig. I. Connective Tissue Cell.^ •.'-Dendrites thickly clothed with Geniinules. Axon. Jrotn cells o/ louver layers with tcfDiinal arl>orizatio}i. Axons with Collaterals. X 110 Fu;. 2. BRAIN OF YOUNG RAT: SECOND LAYER. OF CORTEX NERVE-CELLS, SHOWINC EXTREMELY HIRSUTE DENDRITES AND DESCENDING AXONS. lidU, Sons £ Danielsson, Ltd. Lith. LAMINATION OF THE MOTOR AREA. 125 have regarded as the homologous series in lower mammals, in beino- restricted as a typical formation to a comparatively limited area of the cortex — that of the rodent, e.g., being spread over a far wider propor- tionate area of the hemisphere. This concentration of these cell- groups is best seen in carnivora, where, as already shown, they crowd around the crucial sulcus, especially at the angle of the sigmoid gyrus. They exhibit the tendency in a less marked degree in the higher apes, whilst in man they are concentrated in three or four districts occupy- ing, as before stated, but a comparatively limited area. A still more notable distinction between the higher and lower forms of brain presented by this formation, is the nested arrangement observed by Betz in the human brain. This segregation is complete, the groups being large and far apart. As we descend the scale, however, the more do we observe the tendency for such groups to become confluent,, and the series to be disposed as an equable stratum. Lamination of the Motor Area in Man.— That region of the cortex which has been shown in animals to be electrically excitable, and which upon stimulation calls forth responsive movements, has been termed the "motor area." It is, as we have just seen, chatac- terised by a highly specialised structural arrangement. It is all the more essential that its structure in man should be clearly defined here, since it has been the subject of dispute between such writers as Meynert, Betz, Baillarger, Mierzejewski, and others, some authorities speaking of it as a five-laminated and others as a six-laminated type. At the outset, therefore, it is well to define our own view of the case, which is briefly as follows : — The cortex typical of motor areas is a five-laminated formation, and the more absolutely the granule- cell formation (which, when intercalated, gives us the six-laminated type) is excluded, the more highly specialised become those groups of enormous nerve cells which go by the name of the " nests " of Betz. Where, therefore, these cell-clusters are best represented, there we find a five-laminated, not a six-laminated, cortex ; in other words, at these sites the granule-cell layer no longer exists. Such a specialised cortex is not spread uniformly over a large convolutionary surface at the vertex — any such notion would be very far from correct ; but it occupies very irregular, limited, and unequal areas along the course of the ascending frontal and the junctions between it and the frontal gyri, as well as the " paracentral lobule." These positions we shall more clearly define later on. Such irregularly-disposed areas are severed from each other by a transitional form of lamination, whereby these districts gradually merge into the six-laminated cortex surrounding them. This highly- specialised cortical formation is constituted as follows : — First Layer. — An extremely delicate pale zone limits tlie cortex 126 LAMIXATIOX OF CEREBRAL CORTEX. ■externally ; it presents all the features already described as peculiar to the cortical neuroglia (see p. 94). The outer surface, upon which the intima pia rests, presents numerous flattened cells, from which excessively delicate processes pass downwards into this layer. These •cellular elements are often found, detached from fresh sections, float- ing in the medium around : they form, in fact, a kind of epithelial limiting layer, extremely delicate and translucent. This first layer, ■or peripheral zone, exhibits a pellucid homogeneous matrix (becoming finely molecular with reagents) and three structural constituents — (a) non-nervous, (b) nervous, (c) vascular. (a) The non-nervous constitiients are not numerous, are widely dis- persed, and belong to the two categories of the perivascular or -adventitial elements and the elements of the lymph-connective system already refer-red to (p. 94). The former measure 6 ,tt to 9 /z in diameter, possess a spheroidal nucleus, stain well, and are seen disposed along the course of the blood-vessels. The latter often measure 13 n, in diameter, possess one and occasionally two or three nuclei, are spher- oidal, flask-shaped, or irregular in contour, stain uniformly and very faintly, and throw off numerous excessively delicate processes, which in healthy fresh cortex can only be distinguished with difficulty. (6) The nervous constituents embrace a series of medullated, non- medullated nerve fibres, and nerve cells peculiar to this layer. The medullated nerve fibres course along the outer division of this zonular layer, in a horizontal direction, lying parallel to the pial surface of the cortex ; many of these are observed to pass downwards into the deeper layers of the grey matter. In certain regions, this medullated tract or tangential belt, so characteristic of the first cortical layer, lies .&t a somewhat deeper level near the second layer of cells. In most small mammals these fibres take also an antero-posterior direction. The non-medullated nerve fibres are of two kinds, viz., protoplasmic processes or dendrons from the underlying cells (p. 128): and terminal distribution of axons from the cells peculiar to this layer, and also from those of subjacent tracts. The nerve cells, very sparse in number, are chiefly limited to the lower two-thirds of the peripheral zone. Three kinds of nerve cells are described by Cajal : — the polygonal or stellate, with numerous protoplasmic processes, but with one axon only, which, arising from the body of the cell, ramifies very extensively in all directions in this zone, terminating in fine, varicose and free fibrils ; long fusiform, bipolar cells disposed antero-posteriorly, and, from either pole, giving origin to an exceedingly long protoplasmic process, which along its course throws off" from time to time collateral branches upwards to the pial surface. The peculiar feature, however, in these cells is the plurality of their axons ; each main protoplasmic branch appears con- LAMINATION OF THE MOTOR AREA. 127 tinuous with, or gives off at some distance from the cell, an axon which, coursing horizontally, throws off numerous ascending collaterals, ■ending in extremely delicate branches throughout a very extensive region of this layer. Such axons are pi'obably medullated (Cajal). Supernumerary axons arise also from the secondary protoplasmic branches, usually ascending in their course. These cells, therefore, exhibit the unusual feature of protoplasmic processes ending in axons. Lastly, there are triangular elements which usually throw off three main protoplasmic processes ; two of which run horizontally to the surface, or course somewhat obliquely upwards ; whilst the third passes downwards to bifurcate into arcuate processes, from which arise two, three, or more axons probably medullated, and of extensive distribution. A smaller and more rounded element is also found here, which not only originates axons from its protoplasmic processes, but sends off an axon from the cell body, from which numerous collaterals arise and ascend to the surface. In all these nerve cells of the peripheral zone, besides the plurality of axons, their extensive rami- fication, and their origin from protoplasmic processes, the characteristic features are the great paucity of dendritic branchings, the great length of the dendrons, and the absence of varicosities and collateral spines (Cajal). (c) The vascular elements pass as long straight vessels for deep distribution, and as short branched and smaller vessels through its structure ; they call for no special re- mark here. Second Layer.— A nar- row belt of very closely agcjref'ated nerve cells of irre- gular marginal contour, oval, pyramidal, or angular, with a proportionately large nu- cleus, forms this stratum. The cells vary much in size, and, as we have previously re- marked, are much more richly developed in some than in other regions of the brain. They measure (fig. 15) from 11 /x to 23 /«, in length, G /A to 9 ,'Jb in breadth, the nucleus being often 6 /ti in diameter. Fig. 15. — Cerebral cortex : nerve cells of second layer : descending axons. 128 LAMINATION OF CEREBRAL CORTEX. They exhibit numerous delicate processes, radiating from the base and sides ; but a distinct apical process or frequently a bi-corned apex passes up radially to the surface of the cortex and undergoes rapid subdivision. In the cortex at the vertex in a rat and rabbit this second layer is practically absent ; although it may be traced as small, scattered, appressed groupings of cells on the lateral aspects of the brain. To see these cells to advantage in such brains we must examine the lower arc of the limbic lobe. The same may be said for the sheep ; but, in the pig, this layer is a fairly notable one throughout, reaching a thickness of 138 {Jj. In the cat, dog, and man the depth attained may be 279 /a. Third Layer. — Subjacent to the above lies a deep belt of nerve cells, the elements of which are characterised by their more or less elongated or pyramidal contour, and by the tendency to gradual increase in their size as they lie deeper in the cortex. The summit of these cells is elongated into a long delicate apex process, which passes radially upwards towards the peripheral zone. The opposite pole of the cell is irregularly dentated by the extension of numerous delicate processes, which are thrown off from the cell in all directions around : none of these processes turn upwards and pursue the course of the apex process. The dimensions of these cells in the outermost zone average 12 /a x 8 /a; those of the deeper regions of this layer 22 ih up to even 41 /a in length, and 23 /a in shorter diameter. Each cell possesses a large nucleus and a distinct nucleolus. Small pyramidal cells, however, no larger than those at the commencement of this layer, occur even at the deepest part, side by side with the largest. Fourth Layer. — This layer presents us with the highly charac- teristic nerve element which we have already dealt with under the name of "motor cell." These great elements are found modified in different cortical realms as follows : — (a) In the highest regions of the motor area (summit of central gyri and paracentral lobule) they are not only of gigantic size, as compared with other nerve cells around, but they form here the large clusters recognised by Betz. (b) In the lowest regions of the motor area (lower end of central and junction with third frontal gyrus) they become small in size, even less than the superjacent elements of the third layer, but still retain their clustered disposition. (c) Towards every sulcus these cells, be they large or small, lose their groupings, and at the base of the sulcus they always assume the drawn-out single file, spoken of as the " solitary " type of arrangement. (d) Lastly, as this laminar type passes into that of the sensory realms, these cells have superimposed on them a layer of granule cells, but still retain a somewhat clustered disposition so characterising MOTOR-CELL GROUPINGS. 129 the transitional cortex; and they ultimately assume the solitary arrange- ment always seen in a sulcus, throughout the convolution at all heights, becoming, in fact, the six-laminated cortex typical of sensory areas. Such are the modifications undergone by these elements at different localities in the cortex. Fifth Layer. — This layer is represented by the series of spindle cells, which, beneath the summit of a convolution, are disposed radially to the surface in regular columns, separated by bundles of medullated fasciculi, ascending from the central medullated core of the gyrus. Towards a sulcus they lose this radial disposition, and at the bottom of the sulcus are disposed in a narrow belt, their long axes horizontal to the surface, a position aptly termed reclinate by Dr. Major. These cells measure from 25 /a to 32 /a in length, by 9 /a to 13 /.o in breadth, and exhibit a large oval nucleus. Distribution of the Motor-cell Groups.— The specialised five- p.— Fig. 16.— Left ascendiug frontal and parietal convolutions seen from the side, with the attached frontal gyri included in scheme of examination. H-K, Third group of ganglionic cells. N-0, Fourth group of ganglionic cells. M-N, Barren area. R, Region of large elongate cells. P, Fifth group of ganglionic cells. laminated cortex, with the cell clusters above referred to, has been stated to occupy certain areas of the ascending frontal, the three frontal gyri and the "paracentral" lobule; it remains for us to indicate more particularly the exact site occupied by this type. In the scheme now presented, the results of an investigation into the localisation of these areas in eight human brains, made in 1878 by the author in conjunction 9 I30 LAMINATION OF CEREBRAL CORTEX. with Dr. Henry Clark, are given.* The arrangement and distribution were strangely uniform in all these cases (see figs. 16 and 17). Variations in the extent of these areas, of course, presented themselves, but not to such an extent as to vitiate the general result arrived at, viz., that such cell-clusters were grouped into several distinct areas, very clearly and definitely interrupted by the transitional type of cortex. The variations in the extent of such areas are no more than might be anticipated from the developmental variations indicated by the form of the central and neighbouring gyri. The upper end of the ascending frontal and its junction with the upper frontal gyrus are, as is well known, very variable in form and complexity, and such variations are, in our opinion, closely related to the more or less rich development of the specialised cortex under consideration. Reference to the scheme shows us that the ascending frontal gyrus may, in general, for con- venience of description be considered as consisting of two segments — an upper, comprising two-thirds its length, into which run the superior and middle frontal; and a lower, comprising the remaining third, continuous with the inferior frontal in front, and with the ascending parietal behind. Taking first the upper two-thirds, we find that the upper end has R-->. Fig. 17.— Left ascending frontal and parietal gyri, with the attached frontal convolutions, as seen at the vertex. A-D, First group of ganglionic cells. E-G, Second „ H-K, Third „ M-N, Barren area. N-O, Fourth group of ganghonic cells. R, Region of large elongate cells. a somewhat broad attachment to the upper frontal. The lower end receives the middle frontal usually as a narrower-folded convolution, whilst between either junction a sinuous knee-like bend of the convo- lution exists. The broad upper extremity continuous with the upper frontal is the site of two important clustered groups (A-D and E-G) ; the plump lobule intervening between both upper frontals is the site * " The Cortical Lamination of the Motor Area of the Brain," by Bevan Lewis and Henry Clarke, Proc. Roy. Soc, No. 185, 1878. TRANSITION-REALMS. I^I of two other similar groups (H-K) ; lastly, the extreme posterior end of the middle frontal gyrus shows similar cell groupings (N-0), the areas of which extend into those of the ascending frontal at their lines of attachment. The upper group (A-D) presents by far the larger cells and the more perfect and dense clusters. Such clusters occupy especially the parietal aspect of the convolution, which is adjacent to the ascending parietal convolution. They appear, therefore, in the cortex forming the wall ■of the Rolandic fissure, and creep up towards the summit, where they rapidly thin out and disappear. The second group (E-G), connected with the lower attachment of the Tipper frontal, is entirely restricted to the frontal aspect of this gyrus, -and does not overleap the confines of the vertex and spread into the Rolandic fissure, except at its most inferior part. The third group (H-K) forms a large area, covering the parietal or Rolandic wall of its knee-like lobule (upper two-thirds), and spreads over the summit of the convolution at this site. Between it and the fourth group occurs a narrow territory wholly devoid of this formation; transitional cortex extending until we reach the latter group. At the junction of the middle frontal (N-0).— This group, as before stated, becomes continuous with that of the middle frontal ; it also begins with the fissure of Rolando and sweeps over the vertex. The fifth and sixth groups (P) are indicated approximately on the scheme, but appear subject to considerable variations in extent. To the foregoing groups must be added a further area, occupying the posterior two-thirds of the lobule on the inner or mesial aspect of the central gyri, lying in front of the fissure of Rolando and above the gyrus fornicatus, usually termed the paracentral lobule. Some enor- mous cells are found in the groupings of this area. Tpansition-Realms of Motor Cortex.— It will be observed that, in the above enumeration of specialised areas, we have by no means covered the ground assigned to the motor area by Prof Terrier : the lower end of the ascending frontal, the whole of the ascending parietal, as well as the postero-parietal lobule have been omitted. In fact, these latter regions do not exhibit the specialised cortex referred to, but are covered by cortex transitional in its character between the former and what Ave find existing in sensory realms. If, for instance, the upper extremity of the ascending parietal be subjected to examina- tion, we find that its interior aspect, dipping down into the Rolandic fissure, also possesses large ganglionic cells similar to those in the motor area in advance of this site. The nests or clusters, however, are not only thinly scattered, but contain few cells, and the latter diminish rapidly in size at lower levels along this convolution ; it is only at the upper extremity of the gyrus that large cells are found. 132 LAMINATION OF CEREBRAL CORTEX, Throughout by far the greater extent of this convolution, the cells of this layer are exact representatives of those found in the ascending frontal, hut are greatly diminished in size, and although often arranged in clustered groups, the groups are poor in elements and sparse. The major distinction between the transitional and specialised motor cortex is in the presence of a gradually increasing belt of small pyramidal or angular cells, which are almost identical with those of the second layer, and which here insinuate themselves between the largest cells of the third layer and the sparse nests of the ganglionic cells. Thus, with the fading-off of this rich clustered formation, we- get the intercalation of an entirely new layer of elements, which grows in importance as we approach sensory realms. Now the whole ascend- ing parietal, postero-parietal, and lower end of the ascending frontal divisions, partake of this six-laminated type of cortex; and, moreover, as we approach the margin of the brain-mantle — i.e., the lower end of the central gyri — the "motor" cells become smaller and yet smaller,, forming eventually insignificant clusters of minute elements. Roughly stating the case, we may say that the fissure of Rolando in the upper two-thirds of its extent, separates the typical motor cortex from the transitional cortex ; whilst, in like manner, the interpai-ietal fissure is the boundary between the transitional and the typical sensory cortex below and posterior to it. It will be apparent from the foregoing chapter on the histological structure of the cortex cerebri that its many varieties of type depend, for the most part, upon the operation of one or more of the following^ circumstances. There may be — (1) Inverse development of superimposed layers — such, for in- stance, as was noted in the rodent's brain, where the third layer of cells invariably became shallower with increasing richness of the second layer of angular elements and vice versd. As the one formation, tends to die out, the other tends to increase in thickness and density. (2) Substitutional stratification may occur— i.e., a layer of cells may have other elements mixed with it, and gradually pre- ponderating to the exclusion of its own cells, and then a change in type may occur ; e.g., the granule cells may gradually intermingle with the angular elements, and excluding them entirely, form a deep belt in their place, or vice versd. (3) Intercalation of new layers, as in the appearance of a six- laminated type, where the angular elements gradually insinuate themselves between the third layer of pyramidal cells and the sub- jacent ganglionic series. Or, again, an altered type of cortex may proceed from — (4) An unusual development of the elements of a certain layer, as- Vhen the angular element of the second layer develops into the large- PlaleJd], s %e- tweeri the Nerve -cell?; R ^be Lymphatic •AMttiel^ 'ex. Bale 8tX)awdssoii Ltd: Sculp SIGNIFICANCE OF SIZE OF CELLS. ^33 globose cell of this layer in the modified loweF limbic type of the rat, rabbit, mole, ,, (lower third), . 41 M X 24 m ... We find this law fully borne out by the results of an examination of the bulbar and spinal cell-groups in different regions — the g"reater musculature being presided over by the groups of larg-est Cells. We, therefore, see reason for regarding the dimensions of these cells in the cortex as influenced by — (1) Range of discharging distance. (2) Size of musculature innervated. (3) Age of nerve cell. (4) Resulting multiplicity of cell connections. It will be seen from these conclusions that tlie deepest elements are not necessarily the oldest, for some of the lowest of a series are very small and very simple in their connections. The reason for this was shrewdly given by Dr. Ross from observations on the develop- 136 LAMINATION OF CEREBRAL CORTEX. ment of the motor cell-groups in the anterior cornu of the spinal cord.* His statement is to the effect that the younger cells are in close contiguity to the blood-vessels : that as growth proceeds, they are thrust further aside, so that the larger and older cells lie midway between parallel vessels. No one familiar with the structure and disposition of the cortical elements of the brain will fail to see the force of this sugges- tion. These small pyramidal elements which we meet with constantly side by side with the older cells, are found often with very few lateral branchings, and the apex-process thins out rapidly and is lost to view at a short distance from the cell, notably contrasting in this respect with the older elements, whose apex-process can be traced up into the first layer or peripheral zone. It is important to note this fact — new elements are being continually formed, which for some time have no connection with the grey meshwork of the outer zone of the cortex. These extensions from the apex or centric pole of these young cells continue to thrust themselves further outwards, and are brought into apposition by delicate lateral offshoots, with nerve-fibre plexuses around. Can we suggest the significance borne by the nucleus in the autonomy of the nerve cell ? The results of physiological experimen- tation by Ferrier, Hitzig, Horsley, and Beevor, and clinical investi- gations, especially those of Hughlings-Jackson, appear conclusively to indicate the anterior or fronto-parietal realms of the cerebrum as especially motor; and the occipital and temporo-sphenoidal lobes as especially sensory, in their endowments; and it is, to say the least, highly suggestive that the large pyramidal and ganglionic cells peculiarly characterise the former, just as the smaller elements and densely aggregated granule cells characterise the latter — that, in fact, as we pass from motor to sensory realms, so we find the nerve cells progressively diminishing in bulk and the granule cell progressively preponderating in number. Dr. Hughlings-Jackson long since sug- gested the representation of small muscles by small cells, requiring as they would, in their almost ceaseless lively activity, rapid and frequent, though short, discharges of energy; in fact, he regards such small elements as necessarily of unstable equilibrium. His words are as follows : — " I have suggested that the size and shape of cells, as well as their nearness to the tumour, or other source of irritation, will have to do with their becoming unstable ; other things equal, the same quantity of matter in many small cells will present a vastly greater surface to the contact of nutrient material than the same quantity in one large cell. I have also suggested that small muscles, or, more properly, movements which require little energy for the displacements they have to effect (those of the face and of the hands in touch, for example), are represented by small cells. Such movements are rapidly changing during many of the * Diseases of the Nervous System, vol. ii., p. 26, 1881. Plate XIII. :^<^ff'^^^ 'J, i: .{.-•^V-. JVV \ / Fig. I. Fk;, 2. X 110. CORTEX OF PIG'S BRAIN, TWO DAYS OLD, SUBLIMATE PREPARATION. Bale, Sons & Datiielsson, Ltd.JAth. THE NUCLEUS OF THE NERVE CELL. I 37 ■operations they serve in — writing, for example — and require repetitions of short liberations of energy, and necessitate quick recuperation of the cells concerned. Movements of the upper arm are, in comparison, little changing, and require persistent steady liberation of energy." * When, however, we consider the assumed sensory element of the cortex — the minute angular and granular cells — we must not lose sight of a remarkable distinction between them and the assumed motor unit, and that is, the great proportionate preponderance of the nucleus to the cell itself in the former. That the nucleus does exert some mysterious influence over the nutPitive and functional activity of the cell has long been surmised ; and the results of our histological inquiry indicate that nuclear degeneration within the nerve cell is peculiarly associated with certain states of mental and motorial insta- bility. We have long been accustomed to regard it as related more definitely to the functional activity of the cell, and less directly related to the nutPitiVC activity of the cell. In other words, the cell is subject to a constant supply of nutritive plasma — it gradually assumes a state of nutritive instability, and will necessarily discharge its accumulated energy in accordance with the simple law of nutri- tive rhythm — the resulting stable equilibrium is succeeded by a measurable period ere the potential energising of the cell has once more brought it up to its former state of instability. Were this all that occurs, the process of storage and liberation of energy would be a simpler rhythmic process than the more compounded rhythm which actually pertains to mental operations. If, however, we regard the nucleus as affecting the functional activity of the cell, as in fact, restraining or inhibiting ItS discharge, as a kind of imperium in imperio exercising a controlling influence upon the perturbations which reach the cell from sensoi'y surfaces : then the presence of a healthy nucleus would become an all-important feature in the cell-life — a feature of the utmost significance to us in our patho- logical enquiries. What really does occur when these nuclei are especially affected by morbid processes, we shall refer to more particu- larly in our chapter on the epileptic neuroses. The view we have here taken of the significance of the nucleus would lead to the conclusion that when, from its degeneration or morbid state, it fails to inhibit the cell, these nerve elements would be subject to a rapid running-dotvn on trivial excitation, and in servile obedience to the law of nutritional rhythm ; in fact, we should here find an explanation of morbid insta- bility such as, e.g., in motor realms results in convulsive states, and in the substrata of mental operations in varied psychical states and reductions in consciousness. * "On Temporary Paralysis after Epileptiform and Epileptic Seizures," Brain, vol. iii., footnote to p. 43G. 138 LAMINATION OF CEREBRAL CORTEX. It is these considerations which induce us to regard the dispropor- tionately large nucleus of these small angular elements of the second layer of the cortex as being of some significance. Subject as such minute cells are to a rapid accumulation of energy, we might presume that some restraint must be established to prevent their reckless liberation of energy, and, hence, we believe such restraining capacity to be atforded by the very lai'ge nucleus. In the next place, we have every reason for believing that this superficial belt of angular cells is in direct functional connection with the subjacent cells of large size, and that their morbid instability would, therefore, affect these larger units, which, from the small size of their nucleus, would be more subject to the law of nutritional rhythm in their discharge of energy. As indicated by Dr. Ross, and also in the preceding note by Dr, Hughlings- Jackson, the large cell would present a far smaller area in contact with nutrient material than the same amount of protoplasm broken up into numerous minute elements ; and hence, such large cells would labour under nutritive disadvantages — would be reservoirs for the slotu accumulation and storage of energy, which, when liberated, would again result in a tardy re-instatement of nutritive instability. Electrical Excitability of the Cortex.— Fritsch and Hitzig were the first to demonstrate, in the year 1870, the excitability of the cortex in animals to the galvanic current ; and three years later Prof. Ferrier prosecuted with the faradaic current his first investigations into the functions of the cerebral hemispheres. The method of stimulation employed by Ferrier was, to use his own description, " The application of the electrodes of the secondary spiral of Du Bois-Reymond's induc- tion coil, connected with a cell of the mean electro-motive power of one Daniell. The resistance in the primary coil was such as to give a maximum current of 1'9 absolute unit, as estimated for me by my colleague. Professor Adams. The induced current generated in the secondary coil at 8 cm. distance from the primary spiral was of a strength sufficient to cause a pungent, but quite bearable, sensation when the electrodes were placed on the tip of the tongue." * We can but briefly summarise here some of the more important facts elicited by these experimental methods respecting the reaction of the cortex to electric stimuli. Latent Period of Stimulation and Summation of Stimuli. — It is from these phenomena we infer that the cortical areas found to be excitable are really centres, in the proper acceptation of the term. Tt must be remembered that a ganglionic centrum is an elaborative structure, and that stimuli applied to it meet with delay ere the result- ino- response be elicited. The excitation of a centre is therefore accom- panied by the time element seen in nerve stimulation in a marked * Functions offJie Brain, 2nd edition, p. 223. LATENT PERIOD OF STIMULATION. 139 degree, and this is very appreciable in the stimulation of the so-called psychO-motOP centres of the cortex. This is well brought out on contrasting the effects of a carefully-regulated current applied to the cortex of this realm, with the effects of the same current as applied to the medullated strands immediately beneath, by first excising the overlying cortex. In tlie first place, we find (after, of course, abstracting the time required for transmission down spinal cox'd and motor nerves and the latent period of the muscle) that the retardation is 0'045 of a second, and in the latter place, 0'03 of a second (^Franck and Pitres). So, also, if very feeble stimuli be applied to the cortical centres their summation occurs, so that no contraction takes place until several stimuli have been delivered. Of the many interesting facts revealed by the researches of Schafer and Horsley, Franck and Pitres, relative to the effect of electric stimuli on motor centres, the more important may be stated as follows : — (1) In the same animal the numbeP Of Stimuli per second requisite to produce a continuous contraction is always the same for cortex, motor nerve, and muscle. (2) A continuous contraction does not occur on stimulating a motor centre, until the rate of stimuli reaches 46 per second ; below this, single contraction occurs for each shock or thereabouts. (3) The contractile rhythm of muscle, whether it be cortex, corona radiata, or spinal cord that is stimulated, has been shown to follow this rule : — Rhythm of stimulus below 10 per second = muscular rhythm identical. Rliythm of stimulus at above 10 per second = muscular rhythm constant and independent (Schafer and Horsley). Rhythm of stimulus about 46 per second = continuous muscular contraction [Franck and Pitres). (4) The muscular curve of cortical stimulation is less sudden in its rise and more sustained than the curve shown in subcortical stimula- tion, and all voluntary muscular contractions show a similar rate of undulation in the muscular curve. Modifying" Circumstances. — The excitability of the cortical areas is subject to great variation. Thus, different animals vary in the intensity of stimulus required to produce the adaptive movement ; and the same animal will vary from time to time as regards this susceptibility, according to the conditions in which it is placed. Severe hSBmorrhag^e greatly reduces or even abolishes, whilst moderate loss of blood exalts, the excitability of these parts (Munk, llitzig). Prolonged expOSUre and stimulation rapidly exhaust, whilst apnoea, and the deep narCOSiS of chloroform, ether, chloral, and morphia abolish it [ScJiiff), so that all animals completely ansesthetised fail to reveal such excitability. So if the T40 LAMINATION OF CEREBRAL CORTEX. cortex be in a state of inflammatory iPPitation, its excitability can be readily aroused by even mechanical stimuli, which in health have no such effect. In new-born puppies, Soltmann obtained early response to stimulus, of the corona radiata, whilst it was not until the tenth day that he was able to obtain such response by stimulation of the motor cortex. Functional Equivalence. — Some authorities have inclined to the belief that a process of functional compensation occurs when injury, disease, or experiment has removed a motor centre : that either the opposite sound hemisphere, or even some other portion of the same hemisphere, may assume the functions of the area destroyed. It is undoubtedly true that centres bilaterally associated and least inde- pendent recover soonest from a lesion of one centre, and are least affected in the issue, as is indicated in the history of all cases of ordinary hemiplegia ; but this can scarcely explain what we meet with in experiments on dogs. Here it has been shown that if the motor centres of one hemisphere be destroyed, the resulting hemiplegia is soon recovered from, and if this were due to the substitutional activity of the other hemisphere, ablation of the centres in the latter would presumably paralyse both sides. This, however, is not the case ; for, as Carville and Duret clearly proved, the reinstated power of the limb first paralysed is not affected by the second operation. The explanation is, therefore, not one of functional substitution by another region, but is really due to the more automatic ChaPactCP of the movements in these animals ; in other words, these movements are far more dependent upon the activity of lower centres and are less represented in psycho-motor or cortical realms. In man and the monkey such movements are brought more under the control of the volitional centres — they are removed, as it were, to a higher plane of activity, are less automatic, more independent, and their removal by disease or injury is followed by absolute paralysis of the opposite members. Phenomena of Electpic Stimulation of Coptex.— Professor Ferrier gives preference to the faPadiC Stimulation of the cortex, rather than the galvanic, since the first requisite is a stimulus of a certain dUPation, and not the momentary effect of the opening and closing of a galvanic circuit ; the latter also has the further objection of inducing electrolytic decomposition of the brain-surface if its action be long sustained. If the intensity of current be greater than necessary, diffused stimulation occurs, so that neighbouring areas are aroused into consentaneous activity. ExtPa polaP conduction has also been proved to occur by Carville and Duret, as seen in contractions of a frog's gastrocnemius, the sciatic nerve of which rested on the occiput of a brain, the motor area of PROXIMITY OF PSYCHO-MOTOR CENTRES. 141 which was stimulated. This fact, however, does not vitiate the result* of a minimum current applied to the motor cortex. Conduction to lower centres, as the basal ganglia, has been by some assumed to be explanatory of the results of stimulation of this motor area. This argument is, however, wholly disposed of by the fact that (1) direct stimulation of these ganglia (corpora striata) results in entirely different movements, not the adaptive, purposive movements which the psycho- motor centres elicit ; and that (2) when we bring the electrodes upon their immediate superficial aspect, at the insula, no response whatever occurs. As might have been surmised, the radiations of the coronal medulla, entering into connection with the motor cortex,, are in like manner functionally differentiated ; and, as shown by Burdon-Sanderson, when the cortex is removed and they are stimulated, similar purposive movements can be called forth. Proximity of Psycho-Motor Centres. — It has been seen that the so-called motor cortex, distinguished by the nested cell-g'roups of the fourth layer, is so distributed as to occupy distinct areas, separated only by narrow intervals from each other. This fully accords with the fact that the phenomena of electric stimulation of the cortex demonstrate the clOSe proximity of wholly distinct centres, as Professor Terrier remarks — " Areas in close proximity to each other, separated by a few millimetres or less, react to the electric current in a totally different manner." * * Loc. ciL, p. 229. 142 PART II.— CLINICAL SECTION. €eneral Contents. — States of Depression — States of Exaltation— Fulminating Psychoses — States of Mental Enfeebleraent — Recurrent Insanity — Ejjileptic Insanity — General Paralysis of the Insane— Alcoholic Insanity — Insanity at the Periods of Puberty and Adolescence— At the Puerperal Period — At the Climacteric Epoch— Senile Insanity. STATES OF DEPRESSION. Contents. — Mental Depression Defined — Decline of Object-Consciousness— Rise of Subject-Consciousness— Muscular Element of Thought— Failure in the Rela- tional Element of Mind — Sense of Environmental Resistance— Reductions to Automatic Levels— Sense of Effort— Restricted Volition- -Enfeebled Repre- sentativeness—Transformations of Identity— The Physiological Aspect— Defec- tive Circulation — Nutritional Impairment— Explosive Neuroses— Hunger of the Brain-Cell— Painful and Pleasurable Mental States— Reaction-Time in Melan- cholia—Degrees of Mental Depression— Clinical Varieties of Melancholia- Simple Melancholia— Delusional Melancholia— Hypochondriacal Melancholia- Melancholia Agitans— States of Mental Stupor— Stupor and Hypnotism— Acute Dementia. Painful mental states are of course normal under certain conditions in health and sanity. As in the intellectual sphere it is but human to err, so in the emotional sphere it is but human to suffer, and to feel acutely : hence it is not the intensity of mental pain (although this is often far greater than in healthy states similarly aroused) that characterises this phase of disease, for if the anguish be the outcome of commensurately painful circumstances, we regard it as but a natural reaction. It is in the fact that the emotional storm is out of all proportion to any exciting cause, that we recognise the departure from the standard of health. It is essential, therefore, that we carefully in- .quire into the antecedent circumstances of our patient's disorder, so as to determine whether there are adequate causes to account for the distress apparent — if so, there is but normal physiological reaction, and cerebral function cannot be regarded as deranged. If, however, the mental pain is the result of trivial exciting agencies, if moral or physical agencies arouse emotional states out of all propoi'tion to what would occur in the liealthy mind, then we infer that the grey cortex of the brain is so far disordered as to functionate abnormally, and we speak of the result as pathological depression. It is clear, therefore, that our chief diiEculty in distinguishing normal from abnormal states of depression depends on our correct estimate of the correspondence of emotional states and their excitants, due allowance being made for special peculiarities of temperament. We cannot apply the same rule to a callous, unemotional nature as to one refined and sensitive. MENTAL DEPRESSION. h: In our search for adequate causes we do not confine our attention to the patient's environment ; we must look for possible moral agencies, sucli as shock, disappointment, domestic affliction, together with physical agencies, such as injury, disease, privation, or, again, overstrain of mind, or vicious habits of life — in all alike, the real causes are centric, and consist in a disordered function — the incapacity of reacting commensurately in the conditions in which the organism is placed — in physiological terms it is a "disproportionately excessive" reaction. ^'The melancholia which precedes insanity is distinguished from the mental pain experienced by healthy persons by its excessive degree, by its more than ordinary protraction, by its becoming more and more independent of external influence, and by the other accessory affections which accompany it " (Griesinger):-'' By one thoughtful writer it has been suggested that melancholia might be spoken of as a homologous, wl)ile mania and monomania might be termed heterologous affections.! This, of course, would imply a quantitative and qualitative distinction; but, since" emotional and intellectual states may be disordered quali- tatively as well as quantitatively, the parallel is scarcely applicable. Emotional disturbances as the result of disease differ from the normal reactions of health, not only in volume but also in nature : as Herbert •Spencer indicates, the correspondence may vary in two directions, quantitatively and qualitatively, in degree as well as in kind. With respect to the non-relational feelings — the appetites, pains, ^c. — Herbert Spencer says : — " Their great indefiniteness of limitation and accompanying want of cohesion forbid unions of them, either simultaneous or successive. Obviously, the emotions are characterised by a like want of combining power. A confused and changing chaos is produced by any of them which coexist." | This very want of relativity, this dissociability and absence of a tendency to form strong coherent groups, at once account for the comparative difliculty of estimating the degree of mental alienation in melancholia, as contrasted with states of delusion, where we are dealing with definitely measurable factors. Simple pathological depression is ushered in l)y that failure in object-consciousness which invariably inaugurates a corresj^onding rise in subject-consciousness ; and which, we have reason to infer, implies a diminished functional activity in those realms of the cerebrum correlated thereto. The patient exhibits a growing indifference to his former pursuits and pleasures : the ordinary duties of life and business become irksome and devoid of interest : especially do all forms of mental exertion cause ennui and distaste — the attention cannot as * Mental Diseases, p. 210. t Pfiychological Medicine, Bucknill & Tuke, 3rd edit., p. 440. Principles of Psychology, vol. i., p. 177. 144 STATES OF DEPRESSION. formerly be directed without undue effort, and so reading becomes laborious and thought sluggish and monotonous. The environment fails to call up pleasurable associations — a dreariness and gloom pervade the outside world, since it is interpreted in terms of the predominant feeling. All aspects of object-consciousness alike indi- cate the negative state. There is a want of vigour in the representation of the environment, and feelings aroused thereby are at a low ebb. Corresponding to this there is a pise in SUbject-COnsCiOUSneSS, shown in the prevalence of painful mental states — the predominance of gloomy emotions. This is the positive aspect of the patient's mental state, and this aspect is the one which chiefly obtrudes itself upon our notice. It is characterised especially by an all-prevailing gloom, varying in degree from mild depression up to acutely painful mental states. The subject may complain of vague anxiety — a feeling of some impending evil — an indefinite prevision of coming sorrow, which gives its own colouring to objective existences : he retires from social converse, which but adds to his irritation and mental distress, gives himself up to introspective states, in which he dwells upon the present contents of his mind, broods over his morbid feelings, and falls into long reveries, the subject-matter of which partakes of the same gloomy colouring. He is hyper-sensitive over trifles, irritable and impatient, or his querulous humour may alternate with sullen silence and obstinacy. Even in this reticence and retirement from social responsibilities, this growing apathy to all around or feeling- amounting to dislike or direct hostility, we recognise the origin of that subjectivity, that egoistic state which, in more advanced affections of the mind, conjures up delusions of encroachment and persecution. In every case of mental depression we have this duplex state to study — the negative afi"ection of object-consciousness, and the positive affection of subject-consciousness. Griesinger also asserts that forms of mental depression are due to states of cerebral irritation and mental excitation; but he apparently fails to recognise the duplex nature of the phenomena in neglecting the distinction between the two realms which comprise the totality of consciousness. Thus he says : — " In employing the term, ' states of mental depression,' we do not wish to b& understood as implying that the nature of these states or conditions consists in inaction and weakness, or in the sup2Jression of the mental or cerebral phenomena which accompany them. We have much more cause to assume that verj' violent states oj irritation of the brain and excitation in the mental processes are here very often the cause ; but the general result of these (mental and cerebral) processes is depression, or a painful state of mind. It is sufficient to recall the analogy to physical pain ; and to those who imagine they make things better by substituting 'cerebral torpor' and 'cerebral irritation' for 'depression' and 'exaltation,' DECLINE IN OBJECT-CONSCIOUSNESS. 145 it may fairly enough be objected that in melancholia there is also a state of irritation." * Had he asserted that both conditions co-existed, a state of cerebral torpor in the physical substrata of object-COnSCiOUSneSS, and a state of cerebral ippitation in the substrata of SUbject-COnsCiOUSnesS, he would, we think, have faithfully recorded the morbid phenomena. The normal variations in these antithetic halves of consciousness, with which reverie and dreamy states render us familiar, have been thus lucidly expressed by Herbert Spencer, when in reference to the vivid and faint aggregates of consciousness he says : — " Though entire unconsciousness of things around us is rarely if ever reached, yet the consciousness of them may become very imperfect ; and this imperfect consciousness, observe, results from the independence of the faint series becoming for the time so marked that very little of it clings to the vivid series." t Decline in Objeet-COnseiOUSnesS. — The various states of con- sciousness and the changes from one to the other constitute collectively the sole elements of mind ; and our considerations, therefore, apply to feelings and the relations between feelings. First, let us note that the variations from the normal state embrace a quantitative and a quali- tative change. Feelings may succeed each other in rapid order, or in slow, monotonous file ; they may arise in serial order, or numbers of disconnected states may simultaneously thrust themselves into the field of consciousness, producing turmoil and indefinite vague emotion and thought. On the other hand, mental phenomena may exhibit a qualitative alteration, such as, e.g., is shown in degrees of intensity of feeling, or again, of definiteness as due to the more or less relational character of the product. The decline in object-consciousness which occurs in states of pathological depression, such as we are now dealing with, presents us with the following features : — (a) Enfeebled representativeness : (b) a lessened seriality of thought (weakened attention) : (c) diminution or failure in the muscular element of thought. The last appears to us so important a factor in these morbid states, as to demand here somewhat careful and detailed consideration. Failure in the Muscular Element of Thought.— The constant accompaniment of depressed mental states is a diminished range of perception ; and, since every perception is a complex phenomenon of composite states of consciousness — if one or other of the e.ssential elements of an idea or of a presented object be wanting — the definite realisation of such object or idea is defective. The loss may be in the more sensuous element of the perception — in those qualities, in fact, of body which are categorised as dynamic ("primordial"), e.g., colour, * " Mental Diseases," Syd. Soc, p. 210. •J: Principles of Psychology, vol. ii., p. 459. 10 146 STATES OF DEPRESSION. odour, taste, or the pure sensations appreciated by the specialised senses of sight, hearing, taste, or smell ; again, the loss may pertain chiefly to the statical or primary attributes of the perception— i;hose of size, position, form. A vigorous perception of these primary or space attributes of body is dependent largely upon our " sixth " or muscular sense. If, therefore, this sense undergo any diminution, so will the space attributes of body become less vividly conceived — the cognition is hut feebly produced. The sense of sight is pre-eminently interwoven with the muscular mechanism involved in our perception of objects : and, since the retinal field can only receive the impress of these dynamic attributes of body by means of a musculature, which rotates the eyeball and so disposes the visual axis suitably, the know- ledge of such movements, comprising figure, bulk, and position in space, becomes inextricably blended with these dynamic attributes. There is little doubt that the retinal impressions are, in states of melancholic depression, but feebly produced ; but whether the muscular ■element of perception is first or simultaneously afiected, is an enquiry of special interest. And here we must distinguish between that portion of the muscular element which enters into our higher intellectual con- cepts, and that grosser factor of the large musculature of the limbs, &c., which subserves the purpose of locomotion and coarse movements. The sense of muscular contractions which forms the basis of the prim- ordial ideas of form, size, position, lapses eventually in consciousness as a pure sense of muscular contraction. With the larger musculature this is not so : it is essential that the movements of the limbs, their contraction, and tension should be exquisitely registered centrally, as thereby alone can we gain an idea of their position in space apart from the sense of sight, and appreciate the relative weight of objects and the resistance ofi"ered by them. The unrestrained action of these muscles signalises to our minds the absence of external resist- ance, and the rise in the muscular sense which accompanies any resistance opposed is the direct measure of such resistance. Similarly, with the *' muscularity of thought," which in the normal state is of free and easy play, the rise into consciousness of its primordial muscular element means efibrt, and at once suggests to the mind the same notion of resistance in the environment. It is obvious, we -think, that the muscular element is the first to decline : for cases of intense grief, as from a sudden mental shock, are associated with a notable contraction of this sphere, and space dimensions are altered and contracted. This feature is one of importance, since it clearly points to the decline of the more relational elements of the perceptive process. The" relations of bulk, configuration, and position are recognisable only by the intellectual operations of the mind, and it is this intellectual element which is SENSE OF ENVIRONMENTAL RESISTANCE. 1 47 earliest enfeebled. This follows, therefore, the inverse order of the evolution of psychical powers. Muscular sense, which appears much later in the evolution of the nervous system than do the general or the specialised sensations of sight, hearing, &c. , is in morbid states the first to succumb. The infant learns to appreciate the colour of an object long before he has received the visual percep- tion of its form, bulk, and position : he learns to recognise sounds ere the direction whence they proceed establishes the organised series of reflected changes in certain nuclei of the medulla, which enables him to turn the head and localise the source of such sounds. Just as in the infant we trace the sensuous element of mind as preceding in evolution the relational element, so, in dissolutions of the nervous system in the insane, the inverse order is followed, and the relational decline before the sensuous or "primordial" sensations : and, since a relation can best be defined as a state of consciousness " holding together other states of consciousness " {Herbert Spencer), so individual conscious states become dissociated or unrelated. The loss of such relational element implies a certain degree of intellectual torpor ; but, as we shall have reason to see, the sense of volitional freedom, which is probably an abstract product of the muscular sense, must in like manner decline. Our vigorous perception of the outside world depends largely upon vivid states of consciousness : our realisa- tion of such related states by muscular sense and its derivatives may be compared to a mental gPasp of the environment: and, in direct proportion to the vigour of such grasp, does our power over the envi- ronment predominate, and the resistance of the latter diminish. In states attended by decline of the muscular or relation element of mind, therefore, external resistance must be pari passu intensified, and the apparent energy and freedom of the will restricted. ■ Let us analyse this component of ideation more thoroughly, and we shall find that not only is every perception evolved from a series of com- plex related states of consciousness, but that every concrete perception or idea is attended by certain vivid primary states of consciousness and other secondary component impressions which fail to rise into consciousness, or are more or less revivable or representative. Now such unconscious components of an idea which we take, so to speak, for granted — these lapsed states of consciousness, although they form an integral com- ponent of the perception or ideal representation, are chiefly of TCVWS- CUlar origin. If, in every conception of a sphere, the roll of the eyeball on its axis were induced, the objective origin of the perception of its form would be evident : but, although such actual muscular movements do not occur, yet the musculatures productive of such movements have their centres innervated by each such perception. Still, such innervation as a direct muscular state or sense of muscular tension nnd movement fails, in health, to rise into consciousness — an automatic play calls up vivid representation of form and figure without 148 STATES OF DEPRESSION. any consciousness of muscular action or strain. As before stated, the frequent repetitions of the muscular act essential to the knowledge of figure, position, &c., have eventually resulted in a lapse of the same muscular action in consciousness. If, however, delay occur in the production of such relational states, the statical attributes of body will be perceived only after COnseiOUS effort ; even actual muscular movement and the tension so brought about for the realisation of more vivid conception of form, configur- ation, and bulk, will give that sense of strange efibrt which metes out to us the resistance of the environment. Do actual muscular move- ments occur in the deranged states with which we are now concerned, and does eonSCiOUS effOFt thus arise upon planes which are normally devoid of such feelings 1 The melancholic exhibits to a notable degree the effort which it causes him to think, reflect, or attend to what is said, or to what he reads. It appears to us that the true explanation is due to mental operations being reduced in level so far as to establish conscious effort in lieu of the usual unconscious operations, or lapsed states of consciousness which characterise all intellectual processes. The restless movements of the intellectual eye (in the artist, poet, &c.), as well as those of the state of maniacal excitement, bespeak in the former case the exalted muscular element of thought, and in the latter a highly reflex excitability ; but in the melancholic these muscles of relational life are usually at rest, the eye is fixed, dull, heavy, sluggish in its movements and painful in effort, the eyelids are drooped, the limbs motionless. The only muscles in a state of tension are those which subserve emotional, and not relational life, viz., the small muscles of expression. Hence, the failing vigour of representative states aroused in simple perception or ideation, issues in the sentiment of objective resistance. The environment encroaches pari passu with the failure of that faculty whereby the mind projects out of itself, so to speak, an environment, or revives in idea impressions received from the environment. It is the motor element of mind which is here at fault — the relational element of thought, since it is the space attributes of bodies which are involved. Now, since in the appreciation of these attributes of body (form, bulk, &c.) the subject is active and the object passive, it results that the motor constructive element of the idea is the one which suffers. In other words, failure in the muscular element of thought has as its results on the subjective side, enfeebled ideation and the sense of objective resistance. With respect to the sense of resistance from the environment, it is of interest to note its artificial production in the reductions of consciousness by the agency of anaesthetics. To any one who has been anaesthetised, and who recalls his experiences, say, with nitrous RESTRICTED VOLITION. 1 49 oxide, it mu3t be obvious how the environment crowds in upon one more and more, and how the ego, or personality enslaven by its power, finally feels that thought itself is succumbing to its resistless advance. Restricted Volition. — As in the sphere of perception, so when taking into account consciousness in its totality, we likewise find the same failure in those complex muscular centres, which, in their adjust- ment to the environment, issue in what we term conduct. All volitional acts categorised under this head are the resultants of many factors, or rather the result of the struggle between many contending forces. A certain line of conduct or a certain action being determined upon, presupposes the representation in consciousness of the several possible lines of action. This, in other Avords, is equivalent to saying, that various feeble motor excitations are represented in consciousness, a^nd that the stronger the aggregate of excitations in any special direc- tion, the more does it tend to issue in action. Volitional actions are hence preceded by naSCent motOP excitations. Such excitations are the basis of the act represented to the mind in ideas "which more or less vividly precede the act as realised. In this conflict volition may be enfeebled as the result of failure of those initiative emotions, desires, and sentiments which are in abey- ance in states of depression ; or it may be convulsively restricted as the result of two opposing antagonistic forces, as when such groups of motor excitations divide the attention between them, and the mind sways from one to the other in hesitation and doubt ; or, again, such motor excitation as forms the impulse to action cannot be definitely and strongly represented, and this enfeeblement of muscular represen- tativeness issues in apathy and inaction. A clearer conception of the resultant phenomena may be gleaned by contrasting the voluntary and the involuntary or automatic acts. In the latter, the ideal movements have lapsed in consciousness — the stimulus, whatever it be, is followed so rapidly by the appropriate reaction that the nascent motor excitations do not rise into consciousness. The start of surprise, the suddenly assumed attitude of self-defence, the mechanical movements employed in conveying food to the mouth, and the masticatory actions following thereupon, as well as other complex though automatic acts, have no initial motor antecedent represented in our consciousness ; yet all these movements are exquisitely co- ordinated and rapidly executed. In the enfeeblement of motor representations preceding volitional acts during states of depression, the actions themselves, if performed, are sluggish, mechanical, and devoid of normal energy ; and herein lies the distinction between healthy automatism and these abnormal states. The distinction is more important than at first sight may be apparent, for upon it hinges the explanation of the automatic 150 STATES OF DEPRESSION. freedom of maniacal states, which implies, as we shall see further on, a grave and more serious reduction. The apathy and sluggish reaction of melancholia appear in part due to this want of vigorous motor representation ; the true characteristic of a normal and vigorous mind is the vivid Pealisation in consciousness of the action or line of -conduct to be pursued — the ideal recognition of all alternative lines of conduct by the contrasting faculty, together with the repre- sentations of similar actions previously performed, with the result as affecting the organism. In normal states, each group of the feelings which we class as desires and sentiments rapidly tends to swell the aggregate of its own motor excitations : so rapidly does this natural attraction of " like to like " go on, that the contrasting faculty whereby the resiilt is obtained appears often to act with incalculable rapidity by a process which Spencer calls *' automatic segregation." This process is impaired in states of depression, and becomes sluggish, feeble, and hesitating. Those faint summations of ideal movements which are aroused as the incitants to volitional acts may mutually antagonise each other ; and their very want of vigour will of itself neutralise that distinctive quality which enables the one group to preponderate and overcome the other in action. To employ a figurative illustration — thus do we witness in the surging tide advancing upon a rocky shore, two waves diverging at an angle ; the one, receiving fresh impulses from minor wavelets which take the same course, swells into a rising crest ; the other, receiving no additions, subsides exhausted. Or, two such waves of different size advancing the one upon the other, the higher, representing the aggregate swing of numerous undulations, overcomes and carries with it the surging elements of the weaker. Or, again, to illustrate the feeble representations alluded to, let us picture the uniform ripplets advancing by thousands on the surface ; from want of co-operation, each maintains its own distance from the other, no great con- trasting aggregate of movement is formed collecting to itself stray pulses of force, and hence all alike come to the shore with similar insignificant results. Want of vigorous representation, enfeebled contrasting faculty of thought, antagonistic tendencies, or, lastly, recession or restriction of those feelings which normally excite to voluntary reactions may one or all take part in that restriction of the ego which we speak of as a restrained volition. Here, again, we have suggested to the mind that resistance of the environment which inevitably results where subject-consciousness has a diminished range. It may at first sight appear contradictory to speak of a/aU in object-consciousness and a rise in subject-conscious- ness as issuing in a sense of resistance from the object-world, and a state of enfeebled subjectivity : this is, however, the case, since the less definitely the mind conceives of external realities, the less vivid their representations — the wider the margin for doubt, suspicion, and ideas FAILURE OF PERSONAL IDENTITY. 151 of encroachment from without. We fail to grasp the environment : we do not know it, in the sense of measuring our strength against it — aoid hence lue fear it. So again, the enfeeblement of subject-consciousness pertains only to that "faint aggregate of conscious states which the vivid aggregate tends to draw after them into being " (^Spencer), viz., the ideas con- nected with the outside world, and the representation of our reactions upon the same — hence the faculties of ideation and volition are im- paired. Far otherwise is it, however, with the more sentient element of the self-consciousness — that mass of bodily sensations, visceral, muscular, articular, cutaneous, and the feelings and emotions and sentiments which in the aggregate constitute the sentient or pasSivC e^O — it assumes a concentrated and exaggerated intensity, and this is what we refer to as the rise of subject-consciousness as distinguished from the decline of object-consciousness : a truly self-analytic state. Failure of Personal Identity. — If, now, we attempt to trace further the decadence of mind, in progressive forms of mental disease, we arrive at a very notable stage, and one of profound import, when the failure of object-consciousness is so far advanced as to lead to alterations in the patient's notions of his relationship to the outer world, and to a confusion in his own identity.* A considerable differ- ence is observable in these cases of confused identity, but the more important distinction appears to exist between — (a) Oases of transformed identity associated with general feelings of regard or good-will to the outer world, and a universal sense of well- being, or, at all events, a complete indifference to the environment; and — (6) Cases where, with the transformation of the ego, the environ- ment or non-ego is also transformed in the patient's mind into a formidable, encroaching, and persecuting foe ; whilst all its manifesta- tions usually tend to call up a sense of repugnance and hostility. The ego may exist as a double personality, each independent of the other, or the one swayed by the other, and utterly dissentient in their nature. We need not here deal with these minor differences, but rather consider the development of the latter class, where the identity is transformed and the non-ego is estimated in terms of the malign. It is well, perhaps, at once to state, that these latter forms appear to us to arise out of the various melajicholic types of alienation, whilst the former are educts of the more purely rnaniacal affections. How does this mysterious transformation arise 1 The ego is consti- tuted by the vast aggregate of sensations derived immediately from the body, which are a complexus of all grades of sensory manifestation, * See on this point especially Ribot, p. 107-110; also Griesinger's Mental Diseases, p. 51 ; cf. Spencer, Sully. 152 STATES OF DEPRESSION. from visual and other special senses to tactual and general sense, as well as the far less definite organic or visceral sensations. All those ingoing currents which arouse, more or less definitely, our knowledge of the existence of a body, its limbs, musculature, and viscera, conjointly aid in the elaboration of the ego or personal identity. But the ego is far more than this. We must associate therewith those representations of the same, and moreover the '■'■ faint aggregate" as Spencer terms it, of states aroused by presentative cognitions of the outer kosmos. Our sentiments, ideas, emotions, as well as our memory of presenta- tive states, all, alike go to form that complex elaboration — personal identity, which is severed sharply from the " vivid aggregate " known as the non-ego — the physical in contradistinction to the physiological environment. Now, since in all normal states, the internal order bears a definite relationship to the "outer order" of things, when either of these is profoundly disturbed, the identity tends to suffer con- siderably, as indicated by Sully. We are all acquainted with transient confusion of identity, in those waking states when we fail to realise the impressions suddenly received from the environment ; and were the latter compl«?tely and suddenly transformed, we should fail to restore immediately the balance necessary to re-establish our own identity. So, when the internal mechanism is deranged, and the orderly relationship of inner to outer kosmos is confused, personal identity is apt correspondingly to suffer. We have already seen how this may occur in the progressive failure of object consciousness. The failure to ap])reciate external relationships, again, is associated with that gathering gloom, that sense of outward resistance, fear, and insecurity of the non-ego already alluded to. Impressions from the outer world fail to arouse the normal representative states of cogni- tion, but aid in the welling-up of the emotional life of the subject, and it is from this latter source that falsifications Of Sense arise. As subject-consciousness becomes more and more pronounced with failure of object-consciousness, all impressions alike, received from the non-ego, become the pabulum for the growth of an all-pervading eg'Oism. The subject broods over his multiform and novel feelings — morbid introspection and egoistic musings replace the healthy altruistic feelings and sentiments : and, since the emotional life is itself in part the origin of representative cognitions of the outer kosmos, so out of this source there now arise falsifications of the environment. The pervading gloom, the sense of objective restriction, and the emotional states so aroused, attract to themselves like groupings of ideas—" attempts at explanation," as Griesinger has it ; and this state progressing, tends eventually to the establishment of a neW nexus of ideas correlated to impressions received from without, in TRANSFORMATIONS OF IDENTITY. I 53 lieu of the old and normal relationships pre-existing. It would be a fallacy to assume that the falsifications of the environment precede the emotional disturbance, or that delusions of persecution beget gloomy and malignant passions — this would really invert the actual sequence of phenomena. A gloomy emotional background begets a gloomy interpretation of the non-ego, and all delusions of persecution are begot in like manner out of disordered emotional states. Such translations, if we may so speak, from emotional realms to the realms of thought are, even in normal states of mental life, of frequent occurrence : they peculiarly characterise the poetic faculty, and distinguish the purely emotional and imaginative from the intellectual tj^e of mind ; but, where such emotional incitants to thought are in themselves the product of morbid action, the intellectual result of such operations is liable to be delusive and false. The more immediate concepts, as we may term those which are the result of pure intellectual operations, unassisted by, or only associated with, emotional states, are more subordinate to accurate laws of logic : the more mediate concepts, emotionally derived, are less susceptible to such exactitude of classifying and grouping. Such concepts, in the morbid states now under consideration, are utterly illogical, unclassifiable, frag- mentary, and betray but the disjecta m,emhra of a once rational mind. It appears to us that such distinctions between the immediate and the mediate knowledge, so acquired in the case of the insane, are all-important in our conception of the genesis of these morbid conditions of the ego. We have been tracing in these mental operations the transformation of the environment to the alien's mind : out of the old tissue, by a species of re-arrangement and reconstruction, is woven a fabric repre- senting to him the reality of external things, and which to him is the only reality, but, to his former state of sanity, is an utter falsification. Since this morbid concept is projected out as the actual kosmos, and since internal order must correspond to the external, so a transfor- mation of the ego itself responds to this altered state — the former identity is lost and replaced by the new. And here we have an explication of that newly-acquiPGd freedom which, at this juncture, appears to dawn upon the mind of the mono- maniac. No longer are phenomena in the outer world laboriously investigated and subordinated to rigid laws of logic and of science — they pass, as through a magic crucible, the morbid tissue of his brain, and are transformed in accordance with no objective laws, but take their colour wholly from the morbid emotional states present. Self- creations arise with wondrous celerity and of protean form ; and the morbid imagination conjures up fantastic groupings utterly devoid of coherence and objective reality, A feeling of new freedom replaces the old one of restriction and aggression by the environment, and the ego is consequently endowed with new faculties, new powers — becomes a mighty potentate or a god. Still, the environment is indelibly stamped with the malign character which the former emotional state fostered, and it is only in late stages of the malady that such realisa- 154 STATES OF DEPRESSION. tion of a new-got freedom entirely effaces the enmity of the non-ego from the mind. Like all sudden and extensive transformations of miud, the change thus delineated must be accompanied, as Griesinger has indicated, by great emotional disturbance, "as the results of the conflict between the old and the new." He says, referring to the new sensations and instincts which become generated : — " At first these stand opposed to the old /in the character of a foreign thou, often exciting amazement and fear. Frequently their forcible entrance into the whole sphere of the perception is felt as if it were the possession of the old I by an obscure and irresistible power, and the fact of such forcible possession is expressed by fantastic images. But this duplicity, this conflict of the old I against the new inadequate groups of ideas, is always accompanied by painful opposing sensations, by emotional states, and by Adolent emotions. " * It will be seen that we differ from the above statement, in regarding the emotional perturbation not as the outcome of the "conflict between the old and the new ego," since it appears more in accord with the sequence of the phenomena to regard the morbid emotional storm of this period as being the direct origin of the newly-generated identity. Reductions such as ensue from nervous dissolutions alone, can scarcely explain the phenomena with which we meet: we must, in addition, suppose a process of Pe -integration to ensue. The level to which the mental life is reduced is still one of active, nascent, mental life,t and, like all such nascent life, is accompanied by much emotional disturbance. Even in these morbid minds there is no reason to suppose that the same process does not proceed which we assume to occur in profound sleep, where the re-energising in lower planes, while the individual is for the time unconscious, still proceeds, and so mental potentialities are unconsciously acquired. So also in the monomaniac, though the activities be those of lower planes, still they indicate developmental activities, and those groups of sensa- tions and ideas are conserved which are the fittest to survive : irrational as may be the beliefs, inconsistent the new concepts with the actual truth, still, as Hughlings-Jackson indicates, they are the best possible in the patient's state of reduction. 1 * "Mental Diseases," Syd. Soc, p. 50. t Dr. Hughlings-Jackson has repeatedly insisted upon the negative and positive results of epileptic seizures. + We are prone, by the loose phraseology of common life, to regard the subjec- tive as a permanent possession — to speak of our mind as a something bej'ond the simple active contents of the moment and as the accumulated psychical acti^^ty of our total existence ; as if thoughts could be bottled-up permanently and unchangeably. It is the material substratum of thought^the organised nervous plexuses — which represents the permanent and the potential revivabilities of former experiences, as Herbert Spencer says : — " Just as the external nexus is that which continues to exist amid transitory appearances, so the internal nexus is that which continues to exist amid transitory ideas." — Principles of Psychology, p. 485. THE PHYSIOLOGICAL ASPECT. 155 As the tide of intellectual life retires, so does it well-up into emotional states ; but such emotional wave must have its rebound, and this is expressed in the re-integration which pervades the mental organism with fresh ideas and concepts ; and when such groups acquire a certain definite cohesion amongst themselves, we have the g"enesis of a new identity. It is only at an advanced stage of dissolution that this transfor- mation of the ego can occur — we may safely assert that extensive connections between distant nervous mechanisms must be deranged or dissolved, ere that failure of association of ideas could occur which always precedes this morbid change. Fresh connections probably arise, through the newly-forced channels of the emotional wave, and new centres of internal cohesion are begot and evolve the fresh association of ideas of the transformed ego. And here we might note what we shall later on deal with more fully — viz., those transformations of the personality which characterise certain critical or climacteric periods of life — notably that of puberty. It can readily be conceived how powerfully the mental life is affected by the re-integration of the new encroaching sensations into fresh instincts, desires, impulses ; or, as at the menopause, by the ablation, so to speak, of one of the strongest instincts of the nervous constitution, the sexual. Can it be a matter of wonder that, at these critical periods, the risk to the mental integrity should be great or that, in many subjects, permanent damage should ensue? So interwoven are these instincts with the whole fabric of mind, that a complete transformation of the sentiments and feelings follows, as the result of such incorporation. Obscure longings and yearnings, imperfect, in- definite perceptions, emotional surgings which have no obvious origin or purpose, characterise a period of perturbation of the mental life, which may readily lead to misdirected ejQForts or morbid impulse and disease.* The Physiological Aspect. — In dealing with states of mental depression, did we attempt anything like an artificial division of this class of the vesanise, it would appear to us more important to lay emphasis upon the morbid processes to which they are traced, when- ever such processes can with justice be assumed. It is clear that the symptomatic indications of the so-called varieties of melancholia point not so much to a fundamental distinction in their essential natuie as to one in their mode of origin : they indicate quantitative as well as qualitative variations in tlie nutritive functions of the nervous centres, and, hence, are roughly divisible into groups, comprising those which arise from direct disturbance of the blood-current, and those which are induced in the nervous tissues primarily. *See on this subject the section on the "Insanities of the Period of Pubes- cence." 156 STATES OF DEPRESSION. Two groups stand strongly contrasted here : the one, in which a defective CerebPal circulation is the more prominent feature ; the other, in which an acute nutritional anomaly of the nerve-centres expresses itself in still more unmistakable symptoms. A further group may be constituted by the various qualitative variations of the blood-plasma — tOXSemia, &c. — a group conveniently placed between the two former. D'Abundo has recently affirmed that the toxic and bactericidal action of the defibrinated blood-serum is much increased in all forms of insanity, except in mental depression, in which, on the contrary, it is lessened. His experiments demonstrated the fact that some 10 c.c. of serum to the kilogramme of blood kills rabbits by acute intoxica- tion. It by no means follows, however, that mental symptoms due to the products of auto-intoxication should bear any direct relationship to the toxic effect on lower animals of the blood-serum injected. The effect of certain poisons administered to rabbits upon the nerve cell and its histological constituents has recently been carefully studied by Nissl, very definite changes being induced in the nerve cells of the spinal cord and brain by such agencies as alcohol, morphia, strychnine, lead, phosphorus, and arsenic* It will at once be evident that this is a very arbitrary grouping, the one condition being often associated with the other — nay, evolved out of the other. Thus, defective circulation leads eventually to grave nutritional anomalies, so that the symptoms of the first group may pass into those of the third, although the usual result is not its passage into the acute but into chronic forms of nutritional impairment. Again, quantitative and qualitative variations of the blood, affecting centric nutrition, may co-exist, whilst such nutritional disturbance of the nerve-centres reacts again upon their blood supply. Yet this inter-dependence of functionally related systems, although it renders any sharp demarcation into separate groups impossible, does not impair the practical value of a division into the three groups, since it always holds good that we may clearly distinguish those affections in which the prominent indication is that of simple depressed circulation, from a state in which the vitiated quality of the blood chiefly appeals to us, and, lastly, from those grave affections in which acute and chronic nutritional anomalies are the chief factors con- cerned. States of defective circulation will comprise all the simpler forms of melancholia characterised by lowered cerebral activity. Excitations from the environment do not arouse the normal reaction ; they are sluggishly transmitted, slowly elaborated, and wholly fail to react with due vigour or purposive result. The registry of all impressions * Rivista Sper. di Freniatria, vol. xviii. PATHOLOGICAL DEPRESSION. 157 is faint or imperfect, the latent period prolonged, the reaction-time delayed. The very earliest signs preceding genuine pathological depression are really the symptoms of cerebral anaemia and nervous exhaustion. The cerebral functions are torpid, there is diminished activity both of the impressive and of the expressive realms of the cortex, as above described, and negative states predominate throughout. The subject is heavy, languid, sleepy; frequent yawning occurs — not the insomnia of a more advanced stage ; intellectual efforts are oppressive, and thought becomes dreary, monotonous, and painful. If the warnings thus afforded be disregarded, there arises the frequent recurrence of a painful idea, occasional sensory hallucination, sleeplessness, all indicative of a commencing pathological change — of impaired centric nutrition. In the earlier stage, where warning is not taken, and where, despite such clear evidence of cerebral exhaustion, the brain is still made to do its daily round of duty, in a state utterly inadequate for such exer- tion, unless absolute rest be here enjoined, the next step will certainly issue in pathologfical depression. The morbid nature of this change is sufficiently evident in the fact, that the diurnal cycles of nutritional rhythm are frequently inverted, or at least gravely disturbed. Viewed from the mental aspect, the highest psychical operations are first enfeebled : abstract thought becomes oppressive or impossible ; attention impaired or restricted ; sensations are less vivid, and per- ception is incomplete or wanting in detail or imaginative vigour — the representative faculty especially being enfeebled. Apathy and indif- ference to the surroundings, associated with painful gloom, pervade the mind, betraying the decline of object-consciousness and the rise of subject-consciousness. * In these states, the reaction on the outer world may be characterised by fitful irritability, impatient conduct, sluggish, mechanical actions, or by entire suppression of volitional initiative. Both sensorial and motorial functions are sluggish or in abeyance, and the functions of organic life are all depressed. The vitality of the organism as a whole, being largely dependent upon the activity of the nervous centres, must necessarily suft'er when this important regulative system is deranged : the condition is truly one of devitalisation — life is carried on at a lower level. Should the nutrition of the nerve-centres suffer materially, a fresh series of symptoms is aroused : illusory states and hallucinations distract the attention — the mental pain and disquiet is intensified thereby : apathy and indifference may be replaced by timidity, fright, * So the converse of an over-active circulation reveals itself in increased cerebral activity — often in extraordinarily vivid memories. This we see in fevers, also after the use of certain drugs, as opium, hashish, &c. — (Ribot, Op. cit., p. 198). 158 STATES OF DEPRESSION. or terror ; and the reaction becomes expressive of such emotional states — restless movement and agitated demeanour replacing the former negative condition. All this indicates impaired nutrition of the nerve- -centres, owing to the defective supply of blood : the nerve cells, im- poverished, exchange their normal functional irritability for an exag- gerated abnormal explOSiveneSS, and fitful irregular discharges replace the rhythmic outflow of the nervous discharges which regulate the subordinate centres and relational apparatus of animal life. These nutritional anomalies reach their climax in the third group to -v^hich I have referred, the explosive or fulminating" psychoses. These affections are characterised by the suddenness and explosive nature of the nervous discharge, which relieves the pent-up and accumulating energy of highly unstable centres. In lieu of the •equable rhythm of discharge and repair, corresponding to the wants of the organism, and adapting it to its environment, there is dispro- portionate accumulation of energy ; the centres are brought up to a •degree of high nutritional instability, and the least excitant, however trivial, may, like the spark to the fulminate, issue in an explosion of serious intensity. The nePVe pulse is irregular, fitful, intermittent. This group comprises certain varieties of so-called impulsive insanity — the homicidal and suicidal — the subjects of epileptic neuroses, and affections arising at the climacteric cycles. The cortical expanse of the cerebral hemispheres is certainly the site of the highly representative and re-representative opera- tions : a defective circulation here results in a genuine Starvation of the nePVe-elementS. How does this starvation betray itself 1 In replying to this enquiry let us briefly refer to the physiological appetites for food, &c., and parallelise them with the case in point : we shall then find that all animal appetites are dependent upon two essential factors : — (a) The reception of peripheral excitations by a centric register. (6) The supply of blood to this centre. Thus, the sense of hunger is the indication of a want of this due excitation of the peripheral nerves of the gastric mucous membrane : and for its alleviation the centres must receive impressions so created. But excitation of the peripheral ends of the vagus, produced by any mechanical contact other than by the ingestion of aliment, temporarily suffices to restore the nutritive equilibrium of the nerve- ceutres. The rhythmic pulse of excitations thus transmitted to the centrum calls up the increased vascular flux associated with all brain functionising — and thus, these two agencies combine to raise the nerve-elements into their normal physiological condition of satiety. The reinstatement of molecular equilibrium in the centric nerve cells depends not alone upon the transmission of the physiological stimuli. MENTAL DEPRESSION. 159 but also upon the collateral flow of blood to the part. So — as regards the special senses — the abolition of the usual afferent impressions begets a condition which is truly a pathological hunger. Strikingly is this the case with the sense o? hearing : depression of spirits is a well- marked phenomenon in suddenly-induced deafness, partial or com- plete. The depression so induced we regard as a genuine instance of sensorial hunger — as the expression of starvation of the nerve cells, thus deprived of the normal ingoing currents. Sameness and monotony of sensory impressions produce identical states : and the want of " a change " is nothing more than the ex- pression of this physiological hunger of the nerve-centres. On the other hand, the more highly representative the special sense faculty is in its evolution — the less dependent is it in this respect upon ■presentative excitations ; and thus the sense of sight, when similarly affected, fails to indicate in the same notable degree a corresponding depression of the emotions — idealising or centric initiation so com- pletely supplementing the loss that the results are far different : yet, if the sphere of such operations is in itself implicated, if the nervous mec-hanisms initiating representative processes are starved out by deficiency of blood, then there is begotten a corresponding hunger of the brain-Cell. For, cerebral activity in these realms being restricted, as shown in the poverty of active ideation and thought, there is an arrest of diffusion-currents provocative of the pleasurable emotional states which always accompany healthy energising of these centi-es. Corre- sponding, therefore, to the dreariness of thought in cognitive realms, we have in the region of feeling such painful mental depression as accords with what we should term the hung'eP Of the brain Cell. Again, those ganglionic structures which are the regulative centres for the organs of vegetative life, subserve the wants of the system through the agency of an inscrutable law of nutritional rhythm, differing for each organ concerned : yet, whether we consider the ganglia connected with the visceral sensations, or those which receive epi-peripheral excitatitms, as those of the special senses — i.e., whether the physiological stimuli are continuously received, or have intervals of some duration, or, as in the case of the heart, are equable and periodic — in all cases alike, the excitation of such centres depends much in its degree upon a due supply of blood to the part ; unless this be the case, the centre, exhausted by discharge and not renovated by due nutritional flux, must lose in its excitability. Further, this exhaustion means a weakening 0/ that associative affinity which arouses correlative centres, and Avhich is the physical basis of ideal association. In like manner, the directive agency of such ex- hausted centres must be enfeebled, and the blending of impressions and associated states into the "serial line of thought" (Spencer) l6o STATES OF DEPRESSION. must be correspondingly enfeebled. As we shall see further on, this weaving of the crude material into forms of thought becomes a greater and yet greater effort : " gang'lionic fpiction," as Eomanes aptly terms it, becomes arrestive of the higher processes of thought, and this resistance in the intellectual sphere is associated with a diffusion towards the more purely emotional sphere. Painful and Pleasurable Mental States. — Since these states form so important an element in conditions of mental depression and exaltation, it will repay us here to summarise briefly our views as to their nature. Mental pain has been defined as the result of under-action or over- action — its antithesis, pleasure, finding a place midway between these extremes ; as though, we might say figuratively, an ocean of sluggish waters and of stormy billows lay on each side respectively, with a mid- region of rippling sun-lit wavelets. We think this definition fails — under-action certainly leads to apathy and torpor — over-action to all the various grades of painful mental states ; yet, the essence of this mental pain is surely not over-action, but pent-up activity. Mental pain varies in degree from mild indefinite gloom up to extremes of anguish and despair, in which restricted volition is replaced by agitated and frenzied movement. Now, immediately the sphere of object-conscious- ness declines in functional activity, the minus quantity of the one sphere becomes the plus quantity of the other ; which, in physiological terms, implies that ingoing nervous currents which normally would arouse appropriate reactions in the intellectual and motor realms, become diffused in the realms of feeling and emotion : what is lost for per- ception is gained in feeling. The restricted accumulation of energy is surely at the basis of states of mental pain. If we allude to such states as the result of under-action, the under-action is distinctly that of the higher planes, whilst there is a corresponding surplus of activity aroused in the subordinate planes of feeling and emotion. The so- called states of over-action, again, are similar conditions more definitely expressed — the over-action being that of the recipient, afferent, or im- pressive sphere, with a corresponding under-action of the efferent, intellectual, and expressive sphere — in fact, all grades of mental pain are dependent upon OVCF-action on the impreSSiVC and restricted activity on the expressive plane. In normal states, ingoing currents, or impressions centrally initiated, are translated into realms of motor activity or high intellectual phases; in states of mental pain, such translation is restricted, and such activities expend their energy in those diffused spreading discharges which are the correlatives of emotional conditions. We find pleasurable states invariably associated with the translation oi feeling into thought and action: we likewise find painful mental THE REACTION-TIME IN MELANCHOLIA. i6l •states associated with the surging tide of feeling vainly struggling to bui'st the barriers, in order that it may appear under the varied forms of intellectual or muscular activities: yet, we find all degrees of the latter — from that of high-strung emotional potentiality, down to those minor states where feeling is expressed in terms of general gloom or irritable impatience and fretfulness — passing, in fact, towards states whei'e feel- ing as a higher emotional state seems well-nigh abolished, and passive indifference and apathy indicate a purely negative state of mind. These latter cannot be comprised under the head of states of mental pain, however consistently they may be classed as states of mental depression or anorexia. The Reaction-Time in Melancholia. — Any estimate of the reaction-time in health or disease must take account of many possible sources of error; and such fallacies are but intensified when dealing with the insane mind. A large proportion of the insane do not, of course, admit of such methods of examination : and even amongst such as cheerfully respond to experimentation, a certain proportion are likely to falsify results from individual peculiarities, and the unpredicable vagaries of the insane : delusional cases are in this connection the most doubtful subjects. Where the reductions involve much impairment of memory, or profound mental torpor, the test of 'reaction-time, however taken, is perfectly futile and unreliable : and it is only in those instances of incipient mental derangement, where the intellectual ojjerations are not grossly involved, that a failure in the energy of cerebral reflex can be regarded as of important signifi- cance. In applying the test, the patient should, so far as possible, be made to take an interest in the experiment, and this can frequently be done with great success, by a little tact, even in serious cases of melancholic depression, or acute maniacal outbursts. Each subject should be repeatedly tested short of actual fatigue ; and no average struck of the rapidity of reaction from less than tiventy trials. In our own experi- ments with the insane we have restricted ourselves to the estima- tion of the total time required for reaction to the stimulus of sound or light; and have not attempted to investigate the more complex reaction-time of a more involved process. Reaction-Time Instrument. — We have employed an instrument made by the Cambridge Scientific Instrument Company and designed by Mr. Galton, and have found it admirably adapted for our purpose in testing the reactions in the insane. A short description of this apparatus (fig. 18) may not be out of place here.* * See description of this apparatus by the author in Hack Tnkc's Dictionary of Paychological Medicine, vol. ii., Art. " Psycho-pliysical Methods," No. 3, p. 1022. 11 l62 STATES OF DEPRESSION. Fig. 18. — Reaction-Time Apparatus. f;^The above represents the apparatus as subsequently modified for obtaining a prolonged reaction-time. It will be noted that four graduated rods take the place of a single rod ; that each rod is suspended by an electro-magnet to secure silent release, and that the six Bunsen cells (quart) supply the current for working the electric signals, the upper series of magnets, and the electro-magnet for the clamp. The simpler form of apparatus — devoid of the extra rods, battery, and foot-board — was the instrument mainly used for the results given throughout this work. REACTION-TIME INSTRUMENT. 163 A square standard of pitch pine, 5 feet 10 inches in height, is fixed into a solid, unyielding tripod, in which levelling screws ensure its exactly vertical position. Half way down this standard a i-ectangular piece of mahogany or teak is screwed at right angles to its long axis ; this supports a horizontal table upon whicli rests the hand of the party to be tested. To the same rectangular piece a small electro-magnet is fixed, which holds in position (as an armature) a spring stirrup so long as the electric circuit remains unbroken. This latter circuit passes over the table to a contact breaker, so that the finger of the operator, by depressing a button here, breaks the circuit, releases the stirrup, which, being in its turn drawn back by a powerful spiral spring, clamps the registry rod in its fall. The steel base of the stirrup is fitted inside with rubber, thus forming a more effective brake. From the summit of the standard a box- or lancewood rod is sus- pended, three feet in length, and accurately graduated along its edge in hundredths of a second, up to thirty divisions, the limit of its complete registry being therefore three-tenths of a second. Astride the summit of the registry rod rides a heavy brass plate which falls a short distance with the rod, being then arrested by a diaphragm, its impact causing the SOUnd signal ; or by the make and break of an electric current here it starts an electric bell as the sound signal. The registry rod as it hangs suspended is concealed from the subject's view by a narrow projecting ledge of pine wood fitted to the standard betwixt the rod and the person to be tested. In this ledge, at a con- venient height for the eye, is a small vertical slit or window, and a corresponding slit exists in the rod through which the light is seen ; but, on the release of the rod, the window is closed by its fall, and a Si^ht signal is thus afforded. This apparatus should be as simple as possible, solid and absolutely steady ; the rod should hang perfectly vertical and should not come in contact witii any surface in its fall ; by arranging the levelling screws in the tripod the clamping of the rod should secure it from any sliding, and in its descent the rebound should be reduced to a minimum at the foot, and as little clatter as possible allowed. The release of the rod should be effected with absolute silence, and this is best secured by suspending it to a straight bar electro-magnet by a short cylinder of soft iron. The fall of the rod should be on a cushion or i)ad filled with sand to deaden the resulting thud and check the rebound. The rod is released by pressing a button conveniently placed behind the standard out of observation, which breaks the circuit of the electro-magnet suspending the rod. The Test. — The subject sits supporting his right hand on the table, his forefinger on the interrupting button of the clainp-magnet. The operator silently releases the rod which gives the sound signal, and the 164 STATES OF DEPRESSION. forefinger is instantly depressed, releasing the stirrup and clamping the rod. The figure on the front of the rod where clamped gives the reaction-time for the SOUnd signal. In the next place the brass weight is removed and the subject, sitting as before, is directed to keep his eye on the light seen through the slit in the rod. The rod is now released, the light disappears, and the subject, as before, clamps the rod as rapidly as possible. The time taken in the fall, as read ofi" on the edge of the graduated rod, gives the total reaction-time for a sig'ht signal. The general results, so far obtained, would indicate a decided prolon- gation of the reaction-time in many forms of insanity. Simple afiective forms — as in melancholic depression or maniacal excitement of a simple nature — as well as insanity, the outcome of alcoholism, or of epilepsy, and associated with general paralysis — were made the subjects of enquiry. In none of these were the results more strikingly uniform than in alcoholic forms of insanity, where, after eliminating every probable source of fallacy, the reaction to an optic stimulus was almost invariably delayed, and, in most instances, the reaction to the acoustic stimulus was likewise involved. K one of the patients tested suffered from any serious degree of dementia, such as would have prevented their fully entering into the interest of the trial. ^'' In general paralysis, also, the same delay in reaction occurred, but for such cases we must refer the reader to the series of experiments as given in the section treating of these forms of derangement. Here we more particularly desire to record the results obtained from the subjects labouring under melancholic depression, simple or otherwise. In the following table we have contrasted the results obtained from a series of individuals presumed to be healthy, and from- the subjects of more or less acute melancholic depression : — Reaction-Time ix Health and Disease. Acoustic Stimulus. Sec. •13 •15 •16 Self, . R. H., T. H., R. L., -13 D. A., •le R. W., Simijh Melancholia, . . "29 M. L., „ „ . . . •22 S. W., Climacteric Melancholia, . 29 J. W., Hypochondriacal „ . . "23 G. A., Delusional ,, . . '20 C. K., „ „ . . •U * See also " Reaction-time in certain Forms of Insanity," in Tuke's Dictionary of Psychological Medicine, vol. ii. ; also "Reaction-time" (in same), by Prof. Jastrow, with Bibliography. Optic Stimulus. Sec. •16 •17 •18 •21 •21 •30 •25 •29 •24 •26 •24 REACTION-TIME IN HEALTH AND DISEASE. 165 Acoustic Stimulus. Optic Stimulus Sec. Sec. 0-149 0-200 0-151 0-2-25 0-180 0-188 0-182 0-194 0-128 0-175 0-136 0-150 0-1-22 0-191 0-120 0-193 With the above we also contrast the results given in a table by von Kries and Auerbach, embracing the investigations of several observers * : — Observer. Hirsch, Hankel, Donders, Von Wittich, Wundt, Exner, Auerbach, . Von Kries, . It will be apparent from the observations on healthy subjects, that whereas from ^jf^ to -^-^jj of a second formed the limit of variability for acoustic stimuli, and y^^ to ^Yij ^^^ visual stimuli — in the insane, the former is only exceptionally below -f^jj, and the latter rises from tVo ^^ TijV °^ ^ second. In healthy states the reaction to visual stimuli is slower than to acoustic impressions : — There seems good reason to suppose that the reaction-time of sight is necessarily longer than that of hearing or touch, on account of the photo-chemical nature of its more immediate stimulus. One observer (von Wittich) has even gone so far as to conjecture that the speed of conduction in the optic nerve is less than that of the other nerves of sense ; it is rather to be concluded, however, that the latent time of the sensory end-apparatus, and of the cerebral processes by which sensory impulses pass over into motor impulses is different (Ladd). t The prolongation of the reaction-time in cases of insanity generally, would indicate a special impairment in the visual as contrasted with the auditory sphere : both are often involved ; but the former often suifers to the exclusion of the latter, it being frequently observed that a subject who responds readily and normally to an acoustic stimulus, exhibits notable delay in the response to a visual stimulus. There are many reasons for agreement with Professor Ladd that the distinction in reaction-time for these two kinds of stimuli is due, not to a different rate of conduction, but to the different atent period of end- and centric-org'ans : and we may assume, with nearly as much certainty, that in the deranged states met with in the insane, the protraction of reaction-time found is due, either to impli- cation of the sensory end-organ, or to the intra-central link whereby the sensory is transformed into the motor impulse ; the former is probably illustrated by certain subjects of chronic alcoholism \ — the latter in ordinary forms of aftective insanity. Deg'reeS of Mental Depression. — Of the innumerable combina- tions of mental symptoms embraced under states of mental depression, * Archiv. f. Anat. u. Physiol., Physiolog. Abth., 1877. t Elements of Physiological Psychology, p. 477. X Vide infra, under Alcoholic Insanity. 1 66 STATES OF DEPRESSION. certain forms present themselves, having many features in common of sufficient distinctive value to constitute them arbitrary varieties for the purposes of systematic study : such so-called varieties, however, it must be understood, are by no means other than purely artificial or arbitrary divisions, which are nevertheless essential for the orderly grouping before the mind's eye of what otherwise would form but a chaotic and confusing assemblage of facts. It is thus we hear of a purely affective Melancholia, in which the emotional or affective sphere is chiefly at fault ; and of a delusional melancholia, in which the intellectual or ideational sphere suffers. Whilst fully recognising the utility of such grouping — whereby we keep in view the more notable affective implication in the one case, and the more prominent intellectual perversion in the other — we must insist that the student is here likely to fall into the serious error of regarding such arbitrary divisions as the negation of a principle which we regard as one of the greatest importance in our studies of insanity — viz., the universality of implication which characterises mental disease. By tliis universality of implication we do not mean that all mental faculties suffer alike in extent or degree — this would be obviously absurd : but, that however prominent and obtrusive may be the implication of any special faculty — however limited at first sight may appear the derangement, further investigation shows that the mind in its totality has suffered. The psychological aspect of mental depression presented to the student in the foregoing remarks, will have prepared him for the recognition of this fact of the universal implication of the mental sphere in cases of morbid depression — as subject-consciousness rises in intensity, so he has learnt to appreciate the wane of object- consciousness. It matters not how mild the form of pathological depression — how slight the degree of mental pain — object-conscious- ness invariably presents this corresponding enfeeblement : but this latter feature has to be carefully looked for, whilst the former is the obtrusive and prominent indication of the derangement.* We do not here allude to delusional perversions, but simply to those minor grades of failing representativeness which we have already traced in the sluggishness and poverty of ideation, its lessened vigour, and the dubiety of mind respecting objective existences, which, later on, culminates in delusions of suspicion. When we consider how, in transient functional disturbances falling far short of pathological depression, we find a gloomy emotional tone associated so frequently with a morbid suspicion, bordering at times upon actual delusive states — inconsistent, irrational misjudgments of our fellow-men and * These considerations sufficiently indicate the fallacies of implication to the student's mind of the term partial as contrasted with generalised insanity. SIMPLE MELANCHOLIA. 167 universal distrust, we may be fully prepared in states of genuine melancholia, however mild in type, to recognise in the sphere of the intellectual operations a corresponding wane. Whilst minor degrees of pathological reduction result in a welling-up of feeling as the more obtrusive feature (simple melancholia) — deeper reductions, resulting in more serious implication of representative operations, issue in delusive perversions (delusional melancholia) as the more striking feature, whilst the emotional gloom, in its place, aids in the creation of further delusional notions as " attempts at explanation," to use Griesinger's phi'aseology. It will at once be apparent how this view differs from that which enunciates, as its leading doctrine, an affective origin for insanity : our own view being that the relational and the sentient elements of mind must be conjointly implicated, and that the priority of implication pertains to the relational. Of the clinical groups arbitrarily constituted, from amongst the sub- jects of mental depression, we may cite as the more important — (a) Simple melancholia. (6) Melancholia with delusions, including the hypochondriacal form. (c) Melancholia agitans. (d) Melancholia atonita, or melancholy with stupor. To these separate groups we must now devote some attention : in the first place, it is necessary to indicate, that the varied states which these terms connote are the outcome of the same morbid process in the cerebral cortex, and represent but different depths of dissohUion — serial stages in the same disease. A still lower stage of reduction is that of maniacal excitement : and we mention this fact here, since it is so frequently implied that mania and melancholia are distinct diseases, rather than different stages of the same morbid process. (a) Simple Melancholia. — Under the term of simple melancholia are embraced forms of a purely emotional or affective insanity, where there is mental pain or emotional distress apart from obvious intel- lectual disturbance — if such mental pain be abnormal in its intensity and disproportionate to any exciting cause, we liave a species of simple melancholia. Here, at the outset, we must qualify the phrase, "a purelj' emotional or affective insanity:" for it requires but little insight into the operations of the sound mind, to lead us to the conclusion that so interblended are all the mental faculties in their mutual co-operation, that no such division can be drawn, in a strictly scientific sense, between the purely emotional and the intellectual states. When we speak of emotional states, we must ever bear in mind that the term connotes more or less of the intellectual element of mind — that every mental operation presupposes in its very simplest form — feeling, memory, reason, volition ; or rather tliat these are but different aspects of the same state. It is, therefore, onlj' in tlie greater pre- 1 68 STATES OF DEPRESSION. ponderance of the one or the other factor that we distinguish between abnormal mental states. Simple melancholia, therefore, really embraces those states of morbid depression in which the painful emotional element of mind preponder- ates to the exclusion of disorder of the more relational element ; or, to- be more exact, where the disordered feelings by their intensity and obtrusiveness overshadow any slight intellectual disorder which may be present. Definite delusional states, therefore, are evidently excluded from our definition ; reason still asserts herself; there is no enfeeble- ment of memory ; volitional control is not withdrawn. The cerebral dissolutions which such states of melancholia imply tend certainly towards a lower level, towards more complete dissolutions — and the psychical expressions then vary with it to those of disordered reason, memory, and will : yet, for purposes of clinical study, it is convenient, although this tendency be obvious throughout the attack of insanity, to fix the mind's eye upon the affective disorder. The subject may long have struggled against the gradually increasing depression, and may have concealed his actual state from the notice of relatives or associates — any undue reticence, absence of natural buoyancy, or change in demeanour being usually explained away upon any other grounds than those of mental implication : and thus the barrier between simple functional disturbance and a genuine pathological process is passed without notice. A universal gloom pervades his mind, and a distaste for all previous avocations and interests declares itself: exercise and all forms of recreation no longer appeal to him, and a dull uniform level of indifference is engendered towards the outside world. Life has lost its freshness — Nature pre- sents him "with no delights, and whatever there be of beauty or happiness or gaiety around, but serves to emphasise his gloom as he feels their want of kinship to his nature. With still greater emphasis can he say with Nature's poet : " But yet I know, where'er I go, That there hath pass'd away a glory from the earth." Retiring into the solitude of his own self-consciousness, he broods abstractedly over his alien state — fully cognisant of the nature of his malady ; often dreading to reveal his condition to those most interested in his welfare. But though the object-world has lost for him its pleasurable aspects, and thought and feeling with regard thereto are laboured, restricted, and wanting in vigour — yet subjective states of introspec- tion, of self-analytic activity are keenly dominant, and this self- inflicted torture grows apace as sleep is lost, as defective appetite and sedentary habits of life still further retard the processes of nutrition and repair, and sap the foundations of his mental vigour. SIMPLE MELANCHOLIA. 169 It is at this period that SUicidal pPOmpting'S often come to the front ; but, here we see Reason asserting herself — the patient recognises his moral obligations clearly — often shrinks with horror from tlie suggestion — or may be driven to implore protection such as may be afforded by asylum supervision. Many, however, in the gentle forms of depression, are equally conscious of a degree of self-control which enables them to meet any such suggestion with perfect confidence : they may utter the usual formula of being wearied of life, but, with the utmost self-assurance, deny that they could ever be induced to lay violent hands upon themselves. As we shall see later on, it is in much more serious nutritional anomalies, that the helplessness of the victim and the dread of impulsive acts prevail, as in the fulminating psychoses. Every degree of mental pain may prevail in the subjects of simple melancholia, from such as do not materially interfere with their pursuits — home or business duties, to such as result in utter paralysis of volitional energy : and, in these cases, their daily wants have to be strictly attended to— as they would starve rather than exert themselves to eat. It is by no means unusual at this stage of depression for homicidal actions to replace the suicidal deed. From motives of intense love for her offspring — to save them from the terrible calamity which appears- to await herself — the mother will occasionally sacrifice her children or anyone nearest and dearest to her (^Nicholson). '■•' If the patient does not improve, a further stage is reached in which we observe a still greater wane in object-consciousness: the jaundiced view of the environment is no longer correctly interpreted as due to the subject's own indisposition, but doubts arise — distrust prevails — and a SUSpiciOUS bearing" towards those around inaugurates this -further state of dissolution. The patient, not actually deluded, begins to misinterpret all interference, however kindly meant — looks suspi- ciously at his nurse — struggles violently at the most trivial attention paid him ; whilst the preparations for feeding him or other necessary procedures may be met with every sign of terror. And yet, on questioning him, he admits no definite deluded state — is readily re- assured — only the next moment to relapse into his state of all- prevailing fear that something may happen — he knows not what. The volitional restriction here is serious — self-confidence is greatly enfeebled, and suicide is not unusual : the stage is one of transition to the more definitely deluded or acutely melancholic forms. The follow- ing is a typical illustration of simple melancholic depression : — M. A. W., aged sixty-tiiree, a married woman with a family of two children, was admitted suffering from an attack of depression, which liad commenced about * See Dr. Nicholson's remarks at the discussion of Dr. Semalaigne's Art. on the "Insanity of rersecution," Journ. oj Mental Science, vol. xl,, p. 516. 170 STATES OF DEPRESSION. three weeks previously. She had an earthy complexion — the cheeks mottled with dilated capillaries : her bodily condition approached the obese. No pronounced cardio-vascular change was apparent, although the heart's action was somewhat feeble : the genito-ixrinary system appeared healthy. It was stated that she had just attempted to drown herself in a water-butt. For three weeks past she had slept but little ; had become more depressed from day to day. From her friends' statement, it appeared tliat she had led a perfectly steady, temperate life ; had never before exhibited any mental disturbance or eccentricity, and was not known to have had insane or neurotic ancestors. She was extremely depressed, wept constantly, was reluctant to enter into particulars about her mental state. She admitted that she had been failing in health for some months prior to her attack, and that she did not know the cause of her bodily or mental ailment — could not explain why she was distressed, but was constantly the subject of vague fear, and frequently asked what was to become of her. No delusions were apparent : she had suffered from no hallucinations. This condition continued for some three weeks : she always presented a most melancholic expression, but slept well at night without sedatives, and only on one occasion required forcible administration of food. Through the day she sat rocking herself to and fro, sobbing aloud, and at times became greatly agitated. The more acute symptoms then subsided, and she turned her attention to household work. She became more reticent, and when pressed with questions grew miserable and wept bitterly : could still give no explanation for her fretting. In less than a month she fully realised her own improvement in health — grew more hopeful, less reticent, but now troubled herself much at having attempted suicide. She was now given small doses of opium and ■ether (15 and 10 minims respectively) twice daily, and in a few weeks later was cheerful, active, industrious, longing to return to her friends and home, and left the asylum some ten weeks after her admission. (b) Delusional Melancholia.— This form, as before stated, we regard as presenting us with a deeper stage of reduction than that ■of the simple form of affective insanity (simple melancholia). Gloomy- apprehension and suspicion have here 'passed into definite and per- sistent delusional states ; and, intense as may be the emotional implication, the intellectual derangement now appeals more forcibly to us : and, being constantly insisted upon by the patient, is apt to be regarded by the friends as the real cause of the malady. It would be ■quite apart from our purpose here, even if it were practicable, to illustrate the various features assumed by these cases of delusional melancholia — they will receive sufficient notice in the several clinical forms of insanity which we shall deal with later on. The perversions of the intellect may apply to any one of the whole range of things outside the subject, or may be entirely restricted to the bodily and organic sensations : or again, to his relationships to anotlier state of existence — to his moral being. An infinity of delusive notions, there- fore, necessarily presents itself, often in such strange and contrasting combinations as to be utterly unclassifiable. Prominent, however, amongst such delusive notions are those which •deal with the subject's corporeal frame — the head, the body, limbs, or viscera — often of a grim, and as often of a grotesque character, and DELUSIONAL MELANCHOLIA. I 71 which, if the cattention be riveted thereupon, constitute the so-called hypochondriacal melancholia. Then again, we meet with de- lusions relative to the moral being — the victim has committed the unpardonable sin, or for some, perhaps, insignificant action, his soul is entirely lost — or passages in Scripture constantly recur to him of a gloomy denunciatory nature as applicable to his own state, forming one of the class of so-called relig'iOUS melancholia. Or again, the encroachment of the environment is the more perceptible feature — and the mind conjures up those malign agencies therein which are expressed in the multitudinous ideas Of persecution, tyranny, treachery. And yet again, a well-marked class of patients infer demoniacal possession, witchcraft or other unseen agency as accounting for their states of mental pertui'bation. In these con- ditions of delusional melancholia, hallucinations are not only frequent, but often form the chief material out of which such delusional states are framed. Aural hallucinations more frequently occur than visual, and both far more generally than affections of smell or taste. Halluci- nations of smell are of ominous import — they are frequently associations of irreparable alcoholic brain disease — of epileptic states — of traumatic forms of brain disease, &c. In the following case, however, we find such hallucinations of smell in acute insanity induced by alcohol, but rapidly recovered from : — M. A. S. , a married woman, aged forty-eight, suffering from her first attack of insanity, stated to have been of ten days' duration. She had lived an immoral life for twelve years past, and lately had been of intemperate habits, drinking heavily up to the onset of her attack of insanity. She is not known to have inherited insanity. She was of corpulent proportions, her complexion dusky, and expression dissipated : her bodily health had not very materially suffered. On admission she was greatly agitated and terrified, shaking her limbs violently in bed, or trying to rush from the room. She slept for a few hours after taking 30 minims of paralde- hyde. Much melancholic agitation continued next day : she struggled violently to open the doors, declaring "there was a charm to open them," that there was "a woman after her to burn her," that "she saw the flames." The following night six hours' sleep ensued upon the administration of 25 grains of chloral : she awoke calm and ordei'ly. At this period she was composed, attentive, and coherent, but betrayed the presence of numerous delusions and recent aural liallucinations : thought slie was in an asylum : came here for protection from the noise in her own house. There was such shouting and calling ; if she did not aihsiuer, the.y got louder and louder. Voices kept calling, "Mrs. Birkett, Mrs. Smith, Mrs. Birkett, Mrs. Smith, come and help me, come and help me out !" when she asked, "Who has put you there?" they shouted " Rustan, Rustan, Rustan ;" that had been the cry in her house for the last two years. (She had gone to the police for protection against a man she called Rustan, and had stated that she then saw liis body blowni up by dynamite.) She had "smelt an earthy ■smell like that of a dead body in lier house for the past few weeks." When she went to tlie police for protection, "hundreds of blackguards followed shouting after her." She went the same day to get water from a tap in 172 STATES OF DEPRESSION. the yard. " She Avas sure it was drugged, it dried up her mouth, which began burning and swelling ; she was confident somebody wanted to stab her." As usual in such cases, she absolutely repudiated anj- suggestion of intemperate habits. Her calmness of demeanour continued, but she required a sedative each night to secure any sleep. A week later she affirmed that at home she constantly heard a voice from beneath a stone table calling out, "0 Mrs. Rustan, Mrs. Rustan, Mrs. Birkett, Am y, come down here ; Im down here under the stone." She went and searched under the stone, and saw what "would have blown her up if she had remained in the house." Xone of these voices have been heard since coming to the asylum. A fortnight after her admission she exhibited but verj- gentle depression, and for the first time began to question the real or imaginary nature of the voices heard. Xo relapse occurred, and in less than two months from admission she left, perfectly recovered. (c) Hypochondriacal Melancholia.— In this form of delusional insanity, the morbid interest of the patient is concentrated upon his bodily organism and its functions. In healthy states of activity the ingoing currents arouse, as we have seen (p. 160), none but the massive feeling oi pleasurable well-being, and it is only when the bodily functions are deranged that we become directly conscious of the existence of our organs. So interblended, so inextricably interwoven is the web of sensuous feeling produced by such activities, that out of it arises the central core of the personality — the ego ; around the latter there crowd the impressions received from objective existences — the 2}hysical in contradistinction to the phj/siological environment; yet, these two halves are dissevered, and although they help to form the aggregate mind of the individual, the characteristic stamp of healthful mental operations consists in the continuous and vivid realisation of this distinction between the subject and the object-world. In the lower forms of life, we conceive of the subjective element as forming by far the larger factor of mental states — a vast series of impressions received from the phj'sical en\'ironment are not referred thereto ; and, although the appropriate reaction maj'' occur, this by no means proves that such sensations are not referred to some part of the organic or physiological environment. The higher we rise, the more definite becomes the reference of its O'wn series of excitations to the object-world ; and this, in certain special lines, we see to a remarkable degree in certain insects, such as bees and ants. In man, of course, we attain the complete severance between these antithetic halves which renders possible his knowledge of nature as embraced in the various sciences. "We have seen how the failure of the one half (object-consciousness) may proceed to a serious extent without implicating the groundwork of our being — the personality ; and states of advanced dementia realise this still more fully : we may equally well conceive how the other half (subject-consciousness) may suffer disruption, if this " sensUOUS core " of the personality be implicated either by peripheral or centric derangements. That the former is possible, we have confirmed by numerous instances of hypochondriasis with mental derangement. HYPOCHONDRIACAL MELANCHOLIA. 17, arising from disease of the abdominal and thoracic viscera ; that the hitter occurs, is sufficiently obvious in the excitation of similar states by menial agencies — the perusal of morbid and sensational books, obscene pamphlets, and the association with similar ca,ses of hypo- chondriasis. The anxieties and delusions of the hypochondriacal patient may have reference to any part of his bodily organisation : amongst the insane, howevei", prominence is given to the tract innervated by the pneumogastric nerve, and thus the regions of the throat, thorax, and abdomen — the respiratory, circulatory, and gastro-intestinal organs — are peculiarly the subjects of the patient's anxious attention and complaint. The hypochondriacal subjects of epileptic insanity almost invariably refer their ailments to the stomach and bowels — obscure feelings, pains or imaginary diseases, torpidity or obstruction, are incessantly dwelt upon by them ; and in most instances, if not all, have some basis in actual derangement ; but it is in the constant brooding over these states, and the exaggerated colouring of their ailments, that the hypochondriacal condition is revealed. In the alcoholic subject, on the other hand, hypochondriacal notions have reference often to the peripheral ends of the nerves of common sensation : thus, they continuously examine their limbs, complain of pricklings and other strange sensations in tlie skin; assert that they are poisoned, so that the skin is black, diseased, or "corrupted" (see Alcoholic Insanity). To take the more frequent ailments complained of in hypochon- driacal melancholia, there is the idea so frequently leading to obstinate refusal of food, that "the throat is made up," or that the gullet is wanting, an idea which persists in spite of the frequent passage of the feeding-tube. In such cases, a spasmodic stricture of the oesophagus is not infrequently met with as an obstruction in feeding — the spasm is always high up : it is a reflex spastic state intensified at ouce by the introduction of the cesophageal tube. Although met with in men, it is of more frequent occurrence in women, and then often associated with functional uterine disturbances, as in the cesophagismus of hysterical suVjjects. Organic stricture we have very rarely met with in such subjects ; but direct compression from enlarged thyroid we have frequently found, associated with such delusive conceptions of the absence or total occlusion of the gullet. Such patients are intensely dejected, often seen with the head bent forward, the hands grasping the throat, and fully persuaded that they are dying of inanition, whilst fed artificially with ample meals. They will point to their limbs (often well nourished) as evidence of their advanced emaciation; and they will often induce vomiting, by irritating the fauces after 174 STATES OF DEPRESSION. feeding, declaring that the food so introduced can do them no- good. In like manner, other subjects declare that they have no stomachy and transform various dyspeptic symptoms into indications of grave disease : they may on these grounds resent any attempt at feeding, and struggle violently to thwart one's efforts. Others may take food heartily, yet declare it does not nourish them, and that they are slowly undergoing starvation. Obstruction of the bowels is a most frequent idea, aperient medicine is asked for repeatedly, and despite the daily action of the bowels, the insane patient reiterates his belief that no stool has been passed for days or weeks. Such patients are pictures of misery, importunate- about their treatment, querulous, irritable, wholly absorbed in their own feelings, and can be induced to talk upon no subject without at once reverting to their miserable pliglit. In other cases, the genital organs are the source of anxiety — the- subject believes himself to be impotent or the subject of syphilis, and no possible argument can be used to assure him that his whole system is not permeated by the virus. One patient, at the West Riding Asylum, believed his generative organs had been displaced : another that his sexual organs were diseased and mortified. Numb- ness of the epigastrium was a sore grievance to another patient, who,, moreover, believed that his stomach contained pins and needles. In the case of a middle-aged man who died of tubercular phthisis, an accident (from which he had really suffered some years since, and in which he fractured an arm and two ribs) was made the basis of extravagant delusional notions. He insisted that his skull was nearly hollow — "half his brains having been scattered about at the time;" that he also lost "two gallons of blood ;" and that a screw placed in his bowels by some unknown agency caused him continual and terrible a