Part 14

_Means and Present Aims of Physiological Study of the Brain._--The aspects of cerebral activity are therefore twofold. There is the contribution which it makes to the behaviour of the animal as seen in the creature's doings. On the other hand there is its product in the psychical life of the animal. The former of these is subject matter for physiology; the latter is especially the province of psychology. Physiology does, however, concern itself with the psychical aspect of cerebral functions. Its scope, embracing the study of the bodily organs in regard to function, includes the psychic as well as the material, because as just shown the former inextricably interlace with the latter. But the relation between the psychic phenomena and the working of the brain in regard to any data of fundamental or intimate character connecting the two remains practically as unknown to us as to the Greek philosophers. What physiology has at present to be content with in this respect is the mere assigning of certain kinds of psychic events to certain local regions of the cerebrum. This primitive quest constitutes the greater part of the "neurology" of our day, and some advance has been made along its lines. Yet how meagre are really significant facts will be clear from the brief survey that follows. Before passing finally from these general considerations, we may note that it becomes more and more clear that the brain, although an organ than can be treated as a whole, is complex in the sense that separable functions belong in some measure to its several parts.

The means principally adopted in studying the functions of the brain--and it must be remembered that this study in its present phase is almost exclusively a mere search for localization--are four. These are the physiological, the clinico-pathological, the histological and the zoological. The first named proceeds by observing the effects of artificial excitation, chiefly electric, of various parts of the brain, and the defects produced by destruction or removal of circumscribed portions. The clinico-pathological proceeds by observing the disturbances of body and mind occurring in disease or injury, and ascertaining the extent of the disease or injury, for the most part _post mortem_. The histological method examines the microscopic structure of the various regions of the brain and the characters and arrangement of the nerve-cells composing it. The zoological follows and compares the general features of the brain, as represented in the various types of animal creation.

It is on the functions of the fore-brain that interest now mainly focuses, for the reasons mentioned above. And the interest in the fore-brain itself chiefly attaches to the functions of its cortex. This is due to several causes. In man and the animals nearest him the cortex forms by far the larger part of the whole cerebral hemisphere. More than any other part it constitutes the distinctively human feature. It lies accessible to various experimental observations, as also to traumatic lesions and to the surgeon's art. It is composed of a great unbroken sheet of grey matter; for that reason it is a structure wherein processes of peculiar interest for the investigation in view are likely to occur. To make this last inference more clear a reference to the histology of nervous tissue must be made. The whole physiological function of the nervous system may be summed up in the one word "conduction." This "conduction" may be defined as the transmission of states of excitement (nerve-impulses) along the neural arcs composing the system. The whole nervous system is built up of chains of nerve-cells (neurones) which are nervous conductors, the chains often being termed arcs. Each neurone is an elongated cell which transmits nerve-impulses from its one end to its other, without so far as is known modifying the impulses in transit, unless in that part of the nerve-cell where the nucleus lies. That part of the neurone or nerve-cell is called the perikaryon or cell-body, and from that part usually many branches of the cell (each branch being a nerve-fibre) ramify. There is no evidence that impulses are modified in transit along a branch of a nerve-cell, but there is clear evidence of manifold modification of nerve-impulses in transit along the nerve-arcs of the nervous system. These nerve-arcs are neurone-chains. In them one neurone continues the line of conduction where the immediately foregoing neurone left it. That is, the neurones are laid in conductive series, the far end of one apposed to the near end of its precursor. The place of juxtaposition of the end of one neurone against the beginning of another is called the _synapse_. At it the conduction which has so far been wholly intra-neuronic is replaced by an inter-neuronic process, in which the nerve impulse passes from one neurone to the next. The process there, it is natural to think, must be physiologically different from that conductive process that serves for transmission merely within the neurone itself. It may be that to this inter-neuronic conduction are due the differences between conduction in nerve-_arcs_ and nerve-_trunks_ (nerve-fibres) respectively. Significant of the former are changes in rhythm, intensity, excitability and modifications by summation and inhibition; in fact a number of the main features of nervous reaction. These characters impressed upon conduction in nerve arcs (neurone-chains) would therefore be traceable to the intercalation of perikarya and synapses, for both these structures are absent from nerve-trunks. It is therefore probably to perikarya and synapses that the greater part of the co-ordination, elaboration and differentiation of nervous reactions is due. Now, perikarya and synapses are not present in the _white_ matter of the central nervous organ, any more than they are in nerve-trunks. They are confined exclusively to those portions of the central organ which consist of _grey_ matter (so called from its naked-eye appearance). Hence it is to the great sheet of grey matter which enfolds the cerebrum that the physiologist turns, as to a field where he would expect to find evidences of the processes of cerebral co-ordination at work. It is therefore to items regarding the functions of the great sheet of cerebral cortex that we may now pass.

_The Cerebral Cortex and its Functions._--The main question which vexed the study of the physiology of the cerebral hemispheres in the 19th century was whether differences of function are detectible in the different regions of the hemisphere and especially in those of its cortex. One camp of experimenters and observers held that the cortex was identical in function throughout its extent. These authorities taught that the various faculties and senses suffer damage in proportion to the amount of cortex removed or injured, and that it is a matter of indifference what may be the particular region wherein the destruction takes place. Against this an opposed set of observers held that different regions perform different functions, and this latter "differential" view was raised in two wholly dissimilar forms in the first and last quarters of the 19th century respectively. In the first quarter of the century, a school, with which the name of Gall is prominently associated, held that each faculty of a set of particular so-called "faculties," which it assumed constituted intelligence, has in the brain a spatially separate organ proper to itself. Gall's doctrine had two fundamental propositions. The first was that intelligence resides exclusively in the brain: the second, that intelligence consists of twenty-seven "faculties," each with a separate local seat in the brain. The first proposition was not new. It is met with in Hippocrates, and it had been elaborated by Descartes and others. But Bichat in his _Anatomie generale_ had partly wandered from the gradually established truth and referred the emotions to the visceral organs, returning to a naive view popularly prevalent. Gall's first proposition was probably raised especially in reaction against Bichat. But Gall's proposition was retrograde from the true position of the science of his time. Flourens and others of his contemporaries had already shown not only that intelligence was resident exclusively in the brain, but that it was resident exclusively in that part of the brain which is the fore-brain. Now Gall placed certain of his twenty-seven intellectual faculties in the cerebellum, which is part of the hind-brain.

_Phrenology._--As to Gall's second proposition, the set of faculties into which he analysed intelligence shows his power of psychological analysis to have been so weak that it is matter of surprise his doctrine could obtain even the ephemeral vogue it actually did. Among his twenty-seven faculties are, for instance, "_l'amour de la progeniture, l'instinct carnassier, l'amitie, la ruse, la sagacite comparative, l'esprit metaphysique, le talent poetique, la mimique_," &c. Such crudity of speculation is remarkable in one who had undoubtedly considerable insight into human character. Each of the twenty-seven faculties had its seat in a part of the brain, and that part of the brain was called its "organ." The mere spatial juxtaposition or remoteness of these organs one from another in the brain had, according to Gall, an influence on the constitution of the mind. "_Comme l'organe des arts est place loin de l'organe du sens des couleurs, cette circonstance explique pourquoi les peintres d'histoire ont ete rarement coloristes_." All these "faculty-organs" were placed by Gall at the surface of the brain. "This explains the correspondence which exists between craniology and the doctrine of the functions of the brain (cerebral physiology), the single aim of my researches." Gall wrote that he found the bump of pride (_la bosse de l'orgueil_) as far down in the animal series as the goat. Broussais traced the "organ" of veneration as far down as the sheep. Gall found the bump of murder (_bosse du meurtre_) in the carnivora. Later it was traced also in herbivora. Broussais added apologetically that "the herbivora cause a real destruction of plants."

Gall's doctrine enjoyed enormous vogue. He himself had the gifts and the demerits of quackery. His doctrine possessed, apart from its falsity, certain other mischievous qualities. "_Que ces hommes si glorieux, qui font egorger les nations par millions, sachent qu'ils n'agissent point de leur propre chef, que c'est la nature qui a place dans leur coeur la rage de la destruction_." One of his scientific opponents rejoined, "Nay, it is not that which they should know. What they should know is that if providence has allowed to man the possibility of doing evil, it has also endowed him with the power to do good." The main cause of the success of phrenology (q.v.) has been no doubt the common desire of men to read the characters and hidden thoughts of others by external signs. Each bump or "bosse" on the cranium was supposed to indicate the existence and degree of development of one or other of the twenty-seven "faculties." One such "bosse" showed the development of the organ of "goodness," and another the development of the organ of "murder." Such an easy means to arrive at information so curious delighted many persons, and they were not willingly undeceived.

_Modern Localization Doctrines._--The crude localization of the phrenologists is therefore too clumsy to possess an interest it might otherwise have had as an early expression of belief in cerebral localization, a belief which other labours have subsequently justified, although on facts and lines quite different from these imagined by Gall and his followers. Patient scientific toil by the hands of E. Hitzig and D. Ferrier and their followers has slowly succeeded in obtaining certain facts about the _cortex cerebri_ which not only show that different regions of it are concerned with different functions, but, for some regions at least, outline to some extent the kind of function exercised. It is true that the greater part of the cortex remains still _terra incognita_ unless we are content with mere descriptive features concerning its coarse anatomy. For several scattered regions some knowledge of their function has been gained by physiological investigation. These scattered regions are the _visual_, the _auditory_, the _olfactory_ and the _precentral_.

The grey matter of the cerebral cortex is broadly characterized histologically by the perikarya (nerve-cells bodies) which lie in it possessing a special shape; they are pyramidal. The dendrite fibres of these cells--that is, their fibres which conduct _towards_ the perikarya--are branches from the apex and corners of the pyramid. From the base often near its middle arises one large fibre--the axone fibre, which conducts impulses away from the perikaryon. The general appearance and arrangement of the neurones in a particle of cortical grey matter are shown in fig. 15, above. The apices of the pyramidal perikarya are turned towards the free surface of the cortex. The figure as interpreted in terms of functional conduction means that the cortex is beset with conductors, each of which collects nerve-impulses, from a minute but relatively wide field by its branched dendrites, and that these nerve-impulses converge through its perikaryon, issue by its axone, and are carried whithersoever the axone runs. In some few cells the axone breaks up into branches in the immediate neighbourhood of its own perikaryon in the cortex. In most cases, however, the axone runs off into the subjacent white matter, leaving the cortex altogether. On reaching the subjacent white matter it mingles with other fibres and takes one of the following courses:--(1) to the grey matter of the cortex of the same hemisphere, (2) to the grey matter of the cortex of the opposite hemisphere, (3) to the grey matter of the pons, (4) to the grey matter of the bulb or spinal cord. It is noteworthy that the dendrite fibres of these cortical neurones do not transgress the limits of the grey cortex and the immediate neighbourhood of the perikaryon to which they belong; whereas the discharging or axone fibre does in the vast majority of cases transgress the limits of the grey matter wherein its perikaryon lies. The cortical neurone therefore collects impulses in the region of cortex just about its perikaryon and discharges them to other regions, some not cortical or even cerebral, but spinal, &c. One question which naturally arises is, do these cells spontaneously generate their impulses or are they stirred to activity by impulses which reach them from without? The tendency of physiology is to regard the actions of the cortex as reactions to impulses communicated to the cortical cells by nerve-channels reaching them from the sense organs. The neurone conductors in the cortex are in so far considered to resemble those of reflex centres, though their reactions are more variable and complex than in the use of the spinal. The chains of neurones passing through the cortex are more complex and connected with greater numbers of associate complex chains than are those of the spinal centres. But just as the reflex centres of the cord are each attached to afferent channels arriving from this or that receptive-organ, for instance, tactile-organs of the skin, or spindles of muscle-sense, &c., so the regions of cortex whose function is to-day with some certainty localized seem to be severally related each to some particular sense-organ. The localization, so far as ascertained, is a localization which attaches separate areas of cortex to the several species of sense, namely the visual, the auditory, the olfactory, and so on. This being so, we should expect to find the sensual representation in the cortex especially marked for the organs of the great distance-receptors, the organs which--considered as _sense_ organs--initiate sensations having the quality of projicience into the sensible environment. The organs of distance-receptors are the olfactory, the visual and the auditory. The environmental agent which acts as stimulus in the case of the first named is chemical, in the second is radiant, and in the last is mechanical.

_Olfactory Region of Cortex._--There is phylogenetic evidence that the development of the _cortex cerebri_ first occurred in connexion with the distance-receptors for chemical stimuli--that is, expressed with reference to psychosis, in connexion with olfaction. The olfactory apparatus even in mammals still exhibits a neural architecture of primitive pattern. The cell which conducts impulses to the brain from the olfactory membrane in the nose resembles cells in the skin of the earthworm, in that its cell-body lies actually amid the epithelium of the skin-surface and is not deeply buried near or in the central nervous organ. Further, it has at its external end tiny hairlets such as occur in specially receptive-cells but not usually in purely nervous cells. Hence we must think that one and the same cell by its external end receives the environmental stimulus and by its deep end excites the central nervous organ. The cell under the stimulation of the environmental agent will therefore generate in itself a nervous impulse. This is the clearest instance we have of a neurone being actually excited under natural circumstances by an agent of the environment _directly_, not indirectly. The deep ends of these olfactory neurones having entered the central nervous organ come into contact with the dendrites of large neurones, called, from their shape, mitral. In the dog, an animal with high olfactory sense, the axone of each olfactory neurone is connected with five or six mitral cells. In man each olfactory neurone is connected with a single mitral cell only. We may suppose that the former arrangement conduces to intensification of the central reaction by summation. At the same time it is an arrangement which could tend to smother sharp differentiation of the central reaction in respect to locality of stimulus at the receptive surface. Considering the diffuse way in which olfactory stimuli are applied in comparison, for instance, with visual, the exact localization of the former can obviously yield little information of use for locating the exact position of their source. On the other hand, in the case of visual stimuli the locus of incidence, owing to the rectilinear propagation of light, can serve with extraordinary exactitude for inferences as to the position of their source. The adaptation of the neural connexions of the two organs in this respect is therefore in accord with expectation.

The earliest cerebral cortex is formed in connexion with the neurone-chains coming into the central nervous organ from the patch of olfactory cells on the surface of the head. The region of cerebrum thus developed is the so-called olfactory lobe and hippocampal formation. The greater part of the cerebral hemisphere is often termed the _pallium_, because as its development extends it folds cloak-wise over the older structures at the base of the brain. The olfactory lobe, from its position, is sometimes called the _pallium basale_, and the hippocampal formation the _pallium marginale_; and these two parts of the pallium form what, on account of their phylogenetic history, Elliott Smith well terms the _archipallium_. A fissure, the limbic fissure, marks off more or less distinctly this archipallium from the rest of the pallium, a remainder which is of later development and therefore designated by Elliott Smith the _neopallium_. Of the archipallium, the portion which constitutes the olfactory lobe is well formed in the selachian fish. In the reptilian cerebrum the hippocampal region, the pallium marginale, coexists in addition. These are both of them olfactory in function. Even so high up in the animal scale as the lowest mammals they still form one half of the entire pallium. But in the higher apes and in man the olfactory portion of the pallium is but a small fraction of the pallium as a whole. It is indeed so relatively dwarfed and obscured as to be invisible when the brain is regarded from the side or above. The olfactory part of the pallium exhibits little variation in form as traced up through the higher animals. It is of course small in such animals as Cetaceans, which are _anosmatic_. In highly osmatic such as the dog it is large. The _uncus_, and _subiculum cornu ammonis_ of the human brain, belong to it. Disease of these parts has been accompanied by disturbance of the sense of smell. When stimulated electrically (in the rabbit) the olfactory pallium occasions peculiar torsion of the nose and lips (Ferrier), and change, often slowing or arrested, of the respiratory rhythm. P.E. Flechsig has shown that the nerve-fibres of this part of the pallium attain the final stage of their growth, that is to say, acquire their sheaths of myelin, early in the ontogenetic development of the brain. In the human brain they are myelinate before birth. This is significant from the point of view of function, for reasons which have been made clear especially by the researches of Flechsig himself.

The completion of the growth of the nerve-fibres entering and leaving the cortex occurs at very various periods in the growth of the brain. Study of the development of the fibres entering and leaving the various regions of the pallium in the human brain, discovers that the regions may be conveniently grouped into those whose fibres are perfected before birth and those whose fibres are perfected during the first post-natal month, and those whose fibres are perfected after the first but before the end of the fourth post-natal month. The regions thus marked out by completion before birth are five in number, and are each connected, as also shown by collateral evidence, with one or other particular species of sense-organ. And these regions have another character in common recognizable in the nerve-fibres entering and leaving them, namely, they possess fibres projected to or from parts of the nervous system altogether outside the cortex itself. These fibres are termed "projection" fibres. Other regions of the cortex possess fibres coming from or going to various regions of the cortex itself, but do not possess in addition, as do the five primitive cortical fields, the fibres of projection. So that the facts established by Flechsig for the regions of pallium, which other evidence already indicated as connected with the sense-organ of smell, support that evidence and bring the olfactory region of cortex into line with certain other regions of cortex similarly primarily connected with organs of sense.

It will be noted that what has been achieved by these various means of study in regard to the region of the cortex to which olfactory functions are attributed amounts at present to little more than the bare ascertainment of the existence there of nervous mechanisms connected with olfaction, and to the delimiting roughly of their extent and of their ability to influence certain movements, and in man sensations, habitually associated with exercise of the olfactory organ. As to what part the cortical mechanism has in the elaboration or association of mental processes to which olfaction contributes, no evidence worth the name seems as yet forthcoming. In this respect our knowledge, or rather our want of knowledge, of the functions of the olfactory region of the cortex, is fairly typical of that to which we have to confess in regard to the other regions of the cortex, even the best known.