CHAPTER II.
THE FUNCTIONS OF THE BRAIN.
If I begin chopping the foot of a tree, its branches are unmoved by my act, and its leaves murmur as peacefully as ever in the wind. If, on the contrary, I do violence to the foot of a fellow-man, the rest of his body instantly responds to the aggression by movements of alarm or defence. The reason of this difference is that the man has a nervous system whilst the tree has none; and the function of the nervous system is to bring each part into harmonious co-operation with every other. The afferent nerves, when excited by some physical irritant, be this as gross in its mode of operation as a chopping axe or as subtle as the waves of light, conveys the excitement to the nervous centres. The commotion set up in the centres does not stop there, but discharges itself, if at all strong, through the efferent nerves into muscles and glands, exciting movements of the limbs and viscera, or acts of secretion, which vary with the animal, and with the irritant applied. These acts of response have usually the common character of being of service. They ward off the noxious stimulus and support the beneficial one; whilst if, in itself indifferent, the stimulus be a sign of some distant circumstance of practical importance, the animal's acts are addressed to this circumstance so as to avoid its perils or secure its benefits, as the case may be. To take a common example, if I hear the conductor calling 'All aboard!' as I enter the depot, my heart first stops, then palpitates, and my legs respond to the air-waves falling on my tympanum by quickening their movements. If I stumble as I run, the sensation of falling provokes a movement of the hands towards the direction of the fall, the effect of which is to shield the body from too sudden a shock. If a cinder enter my eye, its lids close forcibly and a copious flow of tears tends to wash it out.
These three responses to a sensational stimulus differ, however, in many respects. The closure of the eye and the lachrymation are quite involuntary, and so is the disturbance of the heart. Such involuntary responses we know as 'reflex' acts. The motion of the arms to break the shock of falling may also be called reflex, since it occurs too quickly to be deliberately intended. Whether it be instinctive or whether it result from the pedestrian education of childhood may be doubtful; it is, at any rate, less automatic than the previous acts, for a man might by conscious effort learn to perform it more skilfully, or even to suppress it altogether. Actions of this kind, into which instinct and volition enter upon equal terms, have been called 'semi-reflex.' The act of running towards the train, on the other hand, has no instinctive element about it. It is purely the result of education, and is preceded by a consciousness of the purpose to be attained and a distinct mandate of the will. It is a 'voluntary act.' Thus the animal's reflex and voluntary performances shade into each other gradually, being connected by acts which may often occur automatically, but may also be modified by conscious intelligence.
An outside observer, unable to perceive the accompanying consciousness, might be wholly at a loss to discriminate between the automatic acts and those which volition escorted. But if the criterion of mind's existence be the choice of the proper means for the attainment of a supposed end, all the acts seem to be inspired by intelligence, for appropriateness characterizes them all alike. This fact, now, has led to two quite opposite theories about the relation to consciousness of the nervous functions. Some authors, finding that the higher voluntary ones seem to require the guidance of feeling, conclude that over the lowest reflexes some such feeling also presides, though it may be a feeling of which we remain unconscious. Others, finding that reflex and semi-automatic acts may, notwithstanding their appropriateness, take place with an unconsciousness apparently complete, fly to the opposite extreme and maintain that the appropriateness even of voluntary actions owes nothing to the fact that consciousness attends them. They are, according to these writers, results of physiological mechanism pure and simple. In a near chapter we shall return to this controversy again. Let us now look a little more closely at the brain and at the ways in which its states may be supposed to condition those of the mind.
THE FROG'S NERVE-CENTRES.
Both the minute anatomy and the detailed physiology of the brain are achievements of the present generation, or rather we may say (beginning with Meynert) of the past twenty years. Many points are still obscure and subject to controversy; but a general way of conceiving the organ has been reached on all hands which in its main feature seems not unlikely to stand, and which even gives a most plausible scheme of the way in which cerebral and mental operations go hand in hand.
The best way to enter the subject will be to take a lower creature, like a frog, and study by the vivisectional method the functions of his different nerve-centres.
Fig. 1.—C H, Cerebral Hemispheres; O Th, Optic Thalami; O L, Optic Lobes; Cb, Cerebellum; M O, Medulla Oblongata; S C, Spinal Cord.
The frog's nerve-centres are figured in the accompanying diagram, which needs no further explanation. I will first proceed to state what happens when various amounts of the anterior parts are removed, in different frogs, in the way in which an ordinary student removes them; that is, with no extreme precautions as to the purity of the operation. We shall in this way reach a very simple conception of the functions of the various centres, involving the strongest possible contrast between the cerebral hemispheres and the lower lobes. This sharp conception Will have didactic advantages, for it is often very instructive to start with too simple a formula and correct it later on. Our first formula, as we shall later see, will have to be softened down somewhat by the results of more careful experimentation both on frogs and birds, and by those of the most recent observations on dogs, monkeys, and man. But it will put us, from the outset, in clear possession of some fundamental notions and distinctions which we could otherwise not gain so well, and none of which the later more completed view will overturn.
If, then, we reduce the frog's nervous system to the spinal cord alone, by making a section behind the base of the skull, between the spinal cord and the medulla oblongata, thereby cutting off the brain from all connection with the rest of the body, the frog will still continue to live, but with a very peculiarly modified activity. It ceases to breathe or swallow; it lies flat on its belly, and does not, like a normal frog, sit up on its fore paws, though its hind legs are kept, as usual, folded against its body and immediately resume this position if drawn out. If thrown on its back, it lies there quietly, without turning over like a normal frog. Locomotion and voice seem entirely abolished. If we suspend it by the nose, and irritate different portions of its skin by acid, it performs a set of remarkable 'defensive' movements calculated to wipe away the irritant. Thus, if the breast be touched, both fore paws will rub it vigorously; if we touch the outer side of the elbow, the hind foot of the same side will rise directly to the spot and wipe it. The back of the foot will rub the knee if that be attacked, whilst if the foot be cut away, the stump will make ineffectual movements, and then, in many frogs, a pause will come, as if for deliberation, succeeded by a rapid passage of the opposite unmutilated foot to the acidulated spot.
The most striking character of all these movements, after their teleological appropriateness, is their precision. They vary, in sensitive frogs and with a proper amount of irritation, so little as almost to resemble in their machine-like regularity the performances of a jumping-jack, whose legs must twitch whenever you pull the string. The spinal cord of the frog thus contains arrangements of cells and fibres fitted to convert skin irritations into movements of defence. We may call it the centre for defensive movements in this animal. We may indeed go farther than this, and by cutting the spinal cord in various places find that its separate segments are independent mechanisms, for appropriate activities of the head and of the arms and legs respectively. The segment governing the arms is especially active, in male frogs, in the breeding season; and these members alone with the breast and back appertaining to them, everything else being cut away, will then actively grasp a finger placed between them and remain hanging to it for a considerable time.
The spinal cord in other animals has analogous powers. Even in man it makes movements of defence. Paraplegics draw up their legs when tickled; and Robin, on tickling the breast of a criminal an hour after decapitation, saw the arm and hand move towards the spot. Of the lower functions of the mammalian cord, studied so ably by Goltz and others, this is not the place to speak.
If, in a second animal, the cut be made just behind the optic lobes so that the cerebellum and medulla oblongata remain attached to the cord, then swallowing, breathing, crawling, and a rather enfeebled jumping and swimming are added to the movements previously observed.[1] There are other reflexes too. The animal, thrown on his back, immediately turns over to his belly. Placed in a shallow bowl, which is floated on water and made to rotate, he responds to the rotation by first turning his head and then waltzing around with his entire body, in the opposite direction to the whirling of the bowl. If his support be tilted so that his head points downwards, he points it up; he points it down if it be pointed upwards, to the right if it be pointed to the left, etc. But his reactions do not go farther than these movements of the head. He will not, like frogs whose thalami are preserved, climb up a board if the latter be tilted, but will slide off it to the ground.
If the cut be made on another frog between the thalami and the optic lobes, the locomotion both on land and water becomes quite normal, and, in addition to the reflexes already shown by the lower centres, he croaks regularly whenever he is pinched under the arms. He compensates rotations, etc., by movements of the head, and turns over from his back; but still drops off his tilted board. As his optic nerves are destroyed by the usual operation, it is impossible to say whether he will avoid obstacles placed in his path.
When, finally, a frog's cerebral hemispheres alone are cut off by a section between them and the thalami which preserves the latter, an unpractised observer would not at first suspect anything abnormal about the animal. Not only is he capable, on proper instigation, of all the acts already described, but he guides himself by sight, so that if an obstacle be set up between him and the light, and he be forced to move forward, he either jumps over it or swerves to one side. He manifests sexual passion at the proper season, and, unlike an altogether brainless frog, which embraces anything placed between his arms, postpones this reflex act until a female of his own species is provided. Thus far, as aforesaid, a person unfamiliar with frogs might not suspect a mutilation; but even such a person would soon remark the almost entire absence of spontaneous motion—that is, motion unprovoked by any present incitation of sense. The continued movements of swimming, performed by the creature in the water, seem to be the fatal result of the contact of that fluid with its skin. They cease when a stick, for example, touches his hands. This is a sensible irritant towards which the feet are automatically drawn by reflex action, and on which the animal remains sitting. He manifests no hunger, and will suffer a fly to crawl over his nose unsnapped at. Fear, too, seems to have deserted him. In a word, he is an extremely complex machine whose actions, so far as they go, tend to self-preservation; but still a machine, in this sense—that it seems to contain no incalculable element. By applying the right sensory stimulus to him we are almost as certain of getting a fixed response as an organist is of hearing a certain tone when he pulls out a certain stop.
But now if to the lower centres we add the cerebral hemispheres, or if, in other words, we make an intact animal the subject of our observations, all this is changed. In addition to the previous responses to present incitements of sense, our frog now goes through long and complex acts of locomotion spontaneously, or as if moved by what in ourselves we should call an idea. His reactions to outward stimuli vary their form, too. Instead of making simple defensive movements with his hind legs like a headless frog if touched, or of giving one or two leaps and then sitting still like a hemisphereless one, he makes persistent and varied efforts at escape, as if, not the mere contact of the physiologist's hand, but the notion of danger suggested by it were now his spur. Led by the feeling of hunger, too, he goes in search of insects, fish, or smaller frogs, and varies his procedure with each species of victim. The physiologist cannot by manipulating him elicit croaking, crawling up a board, swimming or stopping, at will. His conduct has become incalculable. We can no longer foretell it exactly. Effort to escape is his dominant reaction, but he may do anything else, even swell up and become perfectly passive in our hands.
Such are the phenomena commonly observed, and such the impressions which one naturally receives. Certain general conclusions follow irresistibly. First of all the following:
The acts of all the centres involve the use of the same muscles. When a headless frog's hind leg wipes the acid, he calls into play all the leg-muscles which a frog with his full medulla oblongata and cerebellum uses when he turns from his back to his belly. Their contractions are, however, combined differently in the two cases, so that the results vary widely. We must consequently conclude that specific arrangements of cells and fibres exist in the cord for wiping, in the medulla for turning over, etc. Similarly they exist in the thalami for jumping over seen obstacles and for balancing the moved body; in the optic lobes for creeping backwards, or what not. But in the hemispheres, since the presence of these organs brings no new elementary form of movement with it, but only determines differently the occasions on which the movements shall occur, making the usual stimuli less fatal and machine-like; we need suppose no such machinery directly co-ordinative of muscular contractions to exist. We may rather assume, when the mandate for a wiping-movement is sent forth by the hemispheres, that a current goes straight to the wiping-arrangement in the spinal cord, exciting this arrangement as a whole. Similarly, if an intact frog wishes to jump over a stone which he sees, all he need do is to excite from the hemispheres the jumping-centre in the thalami or wherever it may be, and the latter will provide for the details of the execution. It is like a general ordering a colonel to make a certain movement, but not telling him how it shall be done.[2]
The same muscle, then, is repeatedly represented at different heights; and at each it enters into a different combination with other muscles to co-operate in some special form of concerted movement. At each height the movement is discharged by some particular form of sensorial stimulus. Thus in the cord, the skin alone occasions movements; in the upper part of the optic lobes, the eyes are added; in the thalami, the semi-circular canals would seem to play a part; whilst the stimuli which discharge the hemispheres would seem not so much to be elementary sorts of sensation, as groups of sensations forming determinate objects or things. Prey is not pursued nor are enemies shunned by ordinary hemisphereless frogs. Those reactions upon complex circumstances which we call instinctive rather than reflex, are already in this animal dependent on the brain's highest lobes, and still more is this the case with animals higher in the zoological scale.
The results are just the same if, instead of a frog, we take a pigeon, and cut out his hemispheres as they are ordinarily cut out for a lecture-room demonstration. There is not a movement natural to him which this brainless bird cannot perform if expressly excited thereto; only the inner promptings seem deficient, and when left to himself he spends most of his time crouched on the ground with his head sunk between his shoulders as if asleep.
GENERAL NOTION OF HEMISPHERES.
All these facts lead us, when we think about them, to some such explanatory conception as this: The lower centres act from present sensational stimuli alone; the hemispheres act from perceptions and considerations, the sensations which they may receive serving only as suggesters of these. But what are perceptions but sensations grouped together? and what are considerations but expectations, in the fancy, of sensations which will be felt one way or another according as action takes this course or that? If I step aside on seeing a rattlesnake, from considering how dangerous an animal he is, the mental materials which constitute my prudential reflection are images more or less vivid of the movement of his head, of a sudden pain in my leg, of a state of terror, a swelling of the limb, a chill, delirium, unconsciousness, etc., etc., and the ruin of my hopes. But all these images are constructed out of my past experiences. They are reproductions of what I have felt or witnessed. They are, in short, remote sensations; and the difference between the hemisphereless animal and the whole one may be concisely expressed by saying that the one obeys absent, the other only present, objects.
The hemispheres would then seem to be the seat of memory. Vestiges of past experience must in some way be stored up in them, and must, when aroused by present stimuli, first appear as representations of distant goods and evils; and then must discharge into the appropriate motor channels for warding off the evil and securing the benefits of the good. If we liken the nervous currents to electric currents, we can compare the nervous system, C, below the hemispheres to a direct circuit from sense-organ to muscle along the line S . . . C . . . M of Fig. 2 (p. 21). The hemisphere, H, adds the long circuit or loop-line through which the current may pass when for any reason the direct line is not used.
Thus, a tired wayfarer on a hot day throws himself on the damp earth beneath a maple-tree.
Fig. 2.
The sensations of delicious rest and coolness pouring themselves through the direct line would naturally discharge into the muscles of complete extension: he would abandon himself to the dangerous repose. But the loop-line being open, part of the current is drafted along it, and awakens rheumatic or catarral reminiscences, which prevail over the instigations of sense, and make the man arise and pursue his way to where he may enjoy his rest more safely. Presently we shall examine the manner in which the hemispheric loop-line may be supposed to serve as a reservoir for such reminiscences as these. Meanwhile I will ask the reader to notice some corollaries of its being such a reservoir.
First, no animal without it can deliberate, pause, postpone, nicely weigh one motive against another, or compare. Prudence, in a word, is for such a creature an impossible virtue. Accordingly we see that nature removes those functions in the exercise of which prudence is a virtue from the lower centres and hands them over to the cerebrum. Wherever a creature has to deal with complex features of the environment, prudence is a virtue. The higher animals have so to deal; and the more complex the features, the higher we call the animals. The fewer of his acts, then, can such an animal perform without the help of the organs in question. In the frog many acts devolve wholly on the lower centres; in the bird fewer; in the rodent fewer still; in the dog very few indeed; and in apes and men hardly any at all.
The advantages of this are obvious. Take the prehension of food as an example and suppose it to be a reflex performance of the lower centres. The animal will be condemned fatally and irresistibly to snap at it whenever presented, no matter what the circumstances may be; he can no more disobey this prompting than water can refuse to boil when a fire is kindled under the pot. His life will again and again pay the forfeit of his gluttony. Exposure to retaliation, to other enemies, to traps, to poisons, to the dangers of repletion, must be regular parts of his existence. His lack of all thought by which to weigh the danger against the attractiveness of the bait, and of all volition to remain hungry a little while longer, is the direct measure of his lowness in the mental scale. And those fishes which, like our cunners and sculpins, are no sooner thrown back from the hook into the water, than they automatically seize the hook again, would soon expiate the degradation of their intelligence by the extinction of their type, did not their exaggerated fecundity atone for their imprudence. Appetite and the acts it prompts have consequently become in all higher vertebrates functions of the cerebrum. They disappear when the physiologist's knife has left the subordinate centres alone in place. The brainless pigeon will starve though left on a corn-heap.
Take again the sexual function. In birds this devolves exclusively upon the hemispheres. When these are shorn away the pigeon pays no attention to the billings and cooings of its mate. And Goltz found that a bitch in heat would excite no emotion in male dogs who had suffered large loss of cerebral tissue. Those who have read Darwin's 'Descent of Man' know what immense importance in the amelioration of the breed in birds this author ascribes to the mere fact of sexual selection. The sexual act is not performed until every condition of circumstance and sentiment is fulfilled, until time, place, and partner all are fit. But in frogs and toads this passion devolves on the lower centres. They show consequently a machine-like obedience to the present incitement of sense, and an almost total exclusion of the power of choice. Copulation occurs per fas aut nefas, occasionally between males, often with dead females, in puddles exposed on the highway, and the male may be cut in two without letting go his hold. Every spring an immense sacrifice of batrachian life takes place from these causes alone.
No one need be told how dependent all human social elevation is upon the prevalence of chastity. Hardly any factor measures more than this the difference between civilization and barbarism. Physiologically interpreted, chastity means nothing more than the fact that present solicitations of sense are overpowered by suggestions of æsthetic and moral fitness which the circumstances awaken in the cerebrum; and that upon the inhibitory or permissive influence of these alone action directly depends.
Within the psychic life due to the cerebrum itself the same general distinction obtains, between considerations of the more immediate and considerations of the more remote. In all ages the man whose determinations are swayed by reference to the most distant ends has been held to possess the highest intelligence. The tramp who lives from hour to hour; the bohemian whose engagements are from day to day; the bachelor who builds but for a single life; the father who acts for another generation; the patriot who thinks of a whole community and many generations; and finally, the philosopher and saint whose cares are for humanity and for eternity,—these range themselves in an unbroken hierarchy, wherein each successive grade results from an increased manifestation of the special form of action by which the cerebral centres are distinguished from all below them.
In the 'loop-line' along which the memories and ideas of the distant are supposed to lie, the action, so far as it is a physical process, must be interpreted after the type of the action in the lower centres. If regarded here as a reflex process, it must be reflex there as well. The current in both places runs out into the muscles only after it has first run in; but whilst the path by which it runs out is determined in the lower centres by reflections few and fixed amongst the cell-arrangements, in the hemispheres the reflections are many and instable. This, it will be seen, is only a difference of degree and not of kind, and does not change the reflex type. The conception of all action as conforming to this type is the fundamental conception of modern nerve-physiology. So much for our general preliminary conception of the nerve-centres! Let us define it more distinctly before we see how well physiological observation will bear it out in detail.
THE EDUCATION OF THE HEMISPHERES.
Nerve-currents run in through sense-organs, and whilst provoking reflex acts in the lower centres, they arouse ideas in the hemispheres, which either permit the reflexes in question, check them, or substitute others for them. All ideas being in the last resort reminiscences, the question to answer is: How can processes become organized in the hemispheres which correspond to reminiscences in the mind?[3]
Nothing is easier than to conceive a possible way in which this might be done, provided four assumptions be granted. These assumptions (which after all are inevitable in any event) are:
1) The same cerebral process which, when aroused from without by a sense-organ, gives the perception of an object, will give an idea of the same object when aroused by other cerebral processes from within.
2) If processes 1, 2, 3, 4 have once been aroused together or in immediate succession, any subsequent arousal of any one of them (whether from without or within) will tend to arouse the others in the original order. [This is the so-called law of association.]
3) Every sensorial excitement propagated to a lower centre tends to spread upwards and arouse an idea.
4) Every idea tends ultimately either to produce a movement or to check one which otherwise would be produced.
Suppose now (these assumptions being granted) that we have a baby before us who sees a candle-flame for the first time, and, by virtue of a reflex tendency common in babies of a certain age,
Fig. 3.
extends his hand to grasp it, so that his fingers get burned. So far we have two reflex currents in play: first, from the eye to the extension movement, along the line 1—1—1—1 of Fig. 3; and second, from the finger to the movement of drawing back the hand, along the line 2—2—2—2. If this were the baby's whole nervous system, and if the reflexes were once for all organic, we should have no alteration in his behavior, no matter how often the experience recurred. The retinal image of the flame would always make the arm shoot forward, the burning of the finger would always send it back. But we know that 'the burnt child dreads the fire,' and that one experience usually protects the fingers forever. The point is to see how the hemispheres may bring this result to pass.
We must complicate our diagram (see Fig. 4). Let the current 1—1, from the eye, discharge upward as well as downward when it reaches the lower centre for vision, and arouse the perceptional process s1 in the hemispheres;
Fig. 4.—The dotted lines stand for afferent paths, the broken lines for paths between the centres; the entire lines for efferent paths.
let the feeling of the arm's extension also send up a current which leaves a trace of itself, m1; let tha burnt finger leave an analogous trace, s2; and let the movement of retraction leave m2. These four processes will now, by virtue of assumption 2), be associated together by the path s1—m1—s2—m2, running from the first to the last. so that if anything touches off s1, ideas of the extension, of the burnt finger, and of the retraction will pass in rapid succession through the mind. The effect on the child's conduct when the candle-flame is next presented is easy to imagine. Of course the sight of it arouses the grasping reflex; but it arouses simultaneously the idea thereof, together with that of the consequent pain, and of the final retraction of the hand; and if these cerebral processes prevail in strength over the immediate sensation in the centres below, the last idea will be the cue by which the final action is discharged. The grasping will be arrested in mid-career, the hand drawn back, and the child's fingers saved.
In all this we assume that the hemispheres do not natively couple any particular sense-impression with any special motor discharge. They only register, and preserve traces of, such couplings as are already organized in the reflex centres below. But this brings it inevitably about that, when a chain of experiences has been already registered and the first link is impressed once again from without, the last link will often be awakened in idea long before it can exist in fact. And if this last link were previously coupled with a motion, that motion may now come from the mere ideal suggestion without waiting for the actual impression to arise. Thus an animal with hemispheres acts in anticipation of future things; or, to use our previous formula, he acts from considerations of distant good and ill. If we give the name of partners to the original couplings of impressions with motions in a reflex way, then we may say that the function of the hemispheres is simply to bring about exchanges among the partners. Movement mn, which natively is sensation sn's partner, becomes through the hemispheres the partner of sensation s1, s2 or s3. It is like the great commutating switch-board at a central telephone station. No new elementary process is involved; no impression nor any motion peculiar to the hemispheres; but any number of combinations impossible to the lower machinery taken alone, and an endless consequent increase in the possibilities of behavior on the creature's part.
All this, as a mere scheme,[4] is so clear and so concordant with the general look of the facts as almost to impose itself on our belief; but it is anything but clear in detail. The brain-physiology of late years has with great effort sought to work out the paths by which these couplings of sensations with movements take place, both in the hemispheres and in the centres below.
So we must next test our scheme by the facts discovered in this direction. We shall conclude, I think, after taking them all into account, that the scheme probably makes the lower centres too machine-like and the hemispheres not quite machine-like enough, and must consequently be softened down a little. So much I may say in advance. Meanwhile, before plunging into the details which await us, it will somewhat clear our ideas if we contrast the modern way of looking at the matter with the phrenological conception which but lately preceded it.
THE PHRENOLOGICAL CONCEPTION.
In a certain sense Gall was the first to seek to explain in detail how the brain could subserve our mental operations. His way of proceeding was only too simple. He took the faculty-psychology as his ultimatum on the mental side, and he made no farther psychological analysis. Wherever he found an individual with some strongly-marked trait of character he examined his head; and if he found the latter prominent in a certain region, he said without more ado that that region was the 'organ' of the trait or faculty in question. The traits were of very diverse constitution, some being simple sensibilities like 'weight' or 'color;' some being instinctive tendencies like 'alimentiveness' or 'amativeness;' and others, again, being complex resultants like 'conscientiousness,' 'individuality.' Phrenology fell promptly into disrepute among scientific men because observation seemed to show that large
faculties and large 'bumps' might fail to coexist; because the
scheme of Gall was so vast as hardly to admit of accurate
determination at all—who of us can say even of his own
brothers whether their perceptions of weight and of time are well developed or not?—because the followers of Gall and Spurzheim were unable to reform these errors in any appreciable degree; and, finally, because the whole analysis of faculties was vague and erroneous from a psychologic point of view. Popular professors of the lore have nevertheless continued to command the admiration of popular audiences; and there seems no doubt that Phrenology, however little it satisfy our scientific curiosity about the functions of different portions of the brain, may still be, in the hands of intelligent practitioners, a useful help in the art of reading character. A hooked nose and a firm jaw are usually signs of practical energy; soft, delicate hands are signs of refined sensibility. Even so may a prominent eye be a sign of power over language, and a bull-neck a sign of sensuality. But the brain behind the eye and neck need no more be the organ of the signified faculty than the jaw is the organ of the will or the hand the organ of refinement. These correlations between mind and body are, however, so frequent that the 'characters' given by phrenologists are often remarkable for knowingness and insight.
Phrenology hardly does more than restate the problem. To answer the question, "Why do I like children?" by saying, "Because you have a large organ of philoprogenitiveness," but renames the phenomenon to be explained. What is my philoprogenitiveness? Of what mental elements does it consist? And how can a part of the brain be its organ? A science of the mind must reduce such complex manifestations as 'philoprogenitiveness' to their elements. A science of the brain must point out the functions of its elements. A science of the relations of mind and brain must show how the elementary ingredients of the former correspond to the elementary functions of the latter. But phrenology, except by occasional coincidence, takes no account of elements at all. Its 'faculties,' as a rule, are fully equipped persons in a particular mental attitude. Take, for example, the 'faculty' of language. It involves in reality a host of distinct powers. We must first have images of concrete things and ideas of abstract qualities and relations; we must next have the memory of words and then the capacity so to associate each idea or image with a particular word that, when the word is heard, the idea shall forthwith enter our mind. We must conversely, as soon as the idea arises in our mind, associate with it a mental image of the word, and by means of this image we must innervate our articulatory apparatus so as to reproduce the word as physical sound. To read or to write a language other elements still must be introduced. But it is plain that the faculty of spoken language alone is so complicated as to call into play almost all the elementary powers which the mind possesses, memory, imagination, association, judgment, and volition. A portion of the brain competent to be the adequate seat of such a faculty would needs be an entire brain in miniature,—just as the faculty itself is really a specification of the entire man, a sort of homunculus.
Yet just such homunculi are for the most part the phrenological organs. As Lange says:
"We have a parliament of little men together, each one of whom, as happens also in a real parliament, possesses but a single idea which he ceaselessly strives to make prevail"—benevolence, firmness, hope, and the rest. "Instead of one soul, phrenology gives us forty, each alone as enigmatic as the full aggregate psychic life can be. Instead of dividing the latter into effective elements, she divides it into personal beings of peculiar character. . . . 'Herr Pastor, sure there be a horse inside,' called out the peasants to X after their spiritual shepherd had spent hours in explaining to them the construction of the locomotive. With a horse inside truly everything becomes clear, even though it be a queer enough sort of horse—the horse itself calls for no explanation! Phrenology takes a start to get beyond the point of view of the ghost-like soul entity, but she ends by populating the whole skull with ghosts of the same order."[5]
Modern Science conceives of the matter in a very different way. Brain and mind alike consist of simple elements, sensory and motor. "All nervous centres," says Dr. Hughlings Jackson,[6] "from the lowest to the very highest (the substrata of consciousness), are made up of nothing else than nervous arrangements, representing impressions and movements. . . . I do not see of what other materials the brain can be made." Meynert represents the matter similarly when he calls the cortex of the hemispheres the surface of projection for every muscle and every sensitive point of the body. The muscles and the sensitive points are represented each by a cortical point, and the brain is nothing but the sum of all these cortical points, to which, on the mental side, as many ideas correspond. Ideas of sensation, ideas of motion are, on the other hand, the elementary factors out of which the mind is built up by the associationists in psychology. There is a complete parallelism between the two analyses, the same diagram of little dots, circles, or triangles joined by lines symbolizes equally well the cerebral and mental processes: the dots stand for cells or ideas, the lines for fibres or associations. We shall have later to criticise this analysis so far as it relates to the mind; but there is no doubt that it is a most convenient, and has been a most useful, hypothesis, formulating the facts in an extremely natural way.
If, then, we grant that motor and sensory ideas variously associated are the materials of the mind, all we need do to get a complete diagram of the mind's and the brain's relations should be to ascertain which sensory idea corresponds to which sensational surface of projection, and which motor idea to which muscular surface of projection. The associations would then correspond to the fibrous connections between the various surfaces. This distinct cerebral localization of the various elementary sorts of idea has been treated as a 'postulate' by many physiologists (e.g. Munk); and the most stirring controversy in nerve-physiology which the present generation has seen has been the localization-question.
THE LOCALIZATION OF FUNCTIONS IN THE HEMISPHERES.
Up to 1870, the opinion which prevailed was that which the experiments of Flourens on pigeons' brains had made plausible, namely, that the different functions of the hemispheres were not locally separated, but carried on each by the aid of the whole organ. Hitzig in 1870 showed, however, that in a dog's brain highly specialized movements could be produced by electric irritation of determinate regions of the cortex; and Ferrier and Munk, half a dozen years later, seemed to prove, either by irritations or excisions or both, that there were equally determinate regions connected with the senses of sight, touch, hearing, and smell. Munk's special sensorial localizations, however, disagreed with Ferrier's; and Goltz, from his extirpation-experiments, came to a conclusion adverse to strict localization of any kind. The controversy is not yet over. I will not pretend to say anything more of it historically, but give a brief account of the condition in which matters at present stand.
The one thing which is perfectly well established is this, that the 'central' convolutions, on either side of the fissure of Rolando, and (at least in the monkey) the calloso-marginal convolution (which is continuous with them on the mesial surface where one hemisphere is applied against the other), form the region by which all the motor incitations which leave the cortex pass out, on their way to those executive centres in the region of the pons, medulla, and spinal cord from which the muscular contractions are discharged in the last resort. The existence of this so-called 'motor zone' is established by the lines of evidence successively given below:
(1) Cortical Irritations. Electrical currents of small intensity applied to the surface of the said convolutions in dogs, monkeys, and other animals, produce well-defined movements in face, fore-limb, hind-limb, tail, or trunk, according as one point or another of the surface is irritated. These movements affect almost invariably the side opposite to the brain irritations: If the left hemisphere be excited, the movement is of the right leg, side of face, etc. All the objections at first raised against the validity of these experiments have been overcome. The movements are certainly not due to irritations of the base of the brain by the downward spread of the current, for: a) mechanical irritations will produce them, though less easily than electrical; b) shifting the electrodes to a point close by on the surface changes the movement in ways quite inexplicable by changed physical conduction of the current; c) if the cortical 'centre' for a certain movement be cut under with a sharp knife but left in situ, although the electric conductivity is physically unaltered by the operation, the physiological conductivity is gone and currents of the same strength no longer produce the movements which they did; d) the time-interval between the application of the electric stimulus to the cortex and the resultant movement is what it would be if the cortex acted physiologically and not merely physically in transmitting the irritation. It is namely a well-known fact that when a nerve-current has to pass through the spinal cord to excite a muscle by reflex action, the time is longer than if it passes directly down the motor nerve: the cells of the cord take a certain time to discharge. Similarly, when a stimulus is applied directly to the cortex the muscle contracts two or three hundredths of a second later than it does when the place on the cortex is cut away and the electrodes are applied to the white fibres below.[7]
(2) Cortical Ablations. When the cortical spot which is found to produce a movement of the fore-leg, in a dog, is excised (see spot 5 in Fig. 5), the leg in question becomes peculiarly affected. At first it seems paralyzed. Soon, however, it is used with the other legs, but badly. The animal does not bear his weight on it, allows it to rest on its dorsal surface, stands with it crossing the other leg, does not remove it if it hangs over the edge of a table, can no longer 'give the paw' at word of command if able to do so before the operation, does not use it for scratching the ground, or holding a bone as formerly, lets it slip out when running on a smooth surface or when shaking himself, etc., etc. Sensibility of all kinds seems diminished as well as motility, but of this I shall speak later on. Moreover the dog tends in voluntary movements to swerve towards the side of the brain-lesion instead of going straight forward. All these symptoms gradually decrease, so that even with a very severe brain-lesion the dog may be outwardly indistinguishable from a well dog after eight or ten weeks. Still, a slight chloroformization will reproduce the disturbances, even then. There is a certain appearance of ataxic in-coördination in the movements—the dog lifts his fore-feet high and brings them down with more strength than usual, and yet the trouble is not ordinary lack of co-ordination. Neither is there paralysis.
Fig. 5.—Left Hemisphere of Dog's Brain, after Ferrier. A. the fissure of Sylvius. B, the crucial sulcus. O, the olfactory bulb. I, II, III, IV, indicate the first, second, third, and fourth external convolutions respectively. (1), (4), and (5) are on the sigmoid gyrus.
The strength of whatever movements are made is as great as ever—dogs with extensive destruction of the motor zone can jump as high and bite as hard as ever they did, but they seem less easily moved to do anything with the affected parts. Dr. Loeb, who has studied the motor disturbances of dogs more carefully than any one, conceives of them en masse as effects of an increased inertia in all the processes of innervation towards the side opposed to the lesion. All such movements require an unwonted effort for their execution; and when only the normally usual effort is made they fall behind in effectiveness.[8]
Even when the entire motor zone of a dog is removed, there is no permanent paralysis of any part, but only this curious sort of relative inertia when the two sides of the body are compared; and this itself becomes hardly noticeable after a number of weeks have elapsed. Prof. Goltz has described a dog whose entire left hemisphere was destroyed, and who retained only a slight motor inertia on the right half of the body.
Fig. 6.—Left Hemisphere of Monkey's Brain. Outer Surface.
In particular he could use his right paw for holding a bone whilst gnawing it, or for reaching after a piece of meat. Had he been taught to give his paw before the operations, it would have been curious to see whether that faculty also came back. His tactile sensibility was permanently diminished on the right side.[9] In monkeys a genuine paralysis follows upon ablations of the cortex in the motor region. This paralysis affects parts of the body which vary with the brain-parts removed. The monkey's opposite arm or leg hangs flaccid, or at most takes a small part in associated movements. When the entire region is removed there is a genuine and permanent hemiplegia in which the arm is more affected than the leg; and this is followed months later by contracture of the muscles, as in man after inveterate hemiplegia.[10] According to Schaefer and Horsley, the trunk-muscles also become paralyzed after destruction of the marginal convolution on both sides (see Fig. 7). These differences between dogs and monkeys show the danger of drawing general conclusions from experiments done on any one sort of animal. I subjoin the figures given by the last-named authors of the motor regions in the monkey's brain.[11]
Fig. 7.—Left Hemisphere of Monkey's Brain. Mesial Surface.
In man we are necessarily reduced to the observation post-mortem of cortical ablations produced by accident or disease (tumor, hemorrhage, softening, etc.). What results during life from such conditions is either localized spasm, or palsy of certain muscles of the opposite side. The cortical regions which invariably produce these results are homologous with those which we have just been studying in the dog, cat, ape, etc. Figs. 8 and 9 show the result of 169 cases carefully studied by Exner. The parts shaded are regions where lesions produced no motor disturbance.
Fig. 8.—Right Hemisphere of Human Brain. Lateral Surface.
Fig. 9.—Right Hemisphere of Human Brain. Mesial Surface.
Those left white were, on the contrary, never injured without motor disturbances of some sort. Where the injury to the cortical substance is profound in man, the paralysis is permanent and is succeeded by muscular rigidity in the paralyzed parts, just as it may be in the monkey.
(3) Descending degenerations show the intimate connection of the rolandic regions of the cortex with the motor tracts of the cord. When, either in man or in the lower animals, these regions are destroyed, a peculiar degenerative change known as secondary sclerosis is found to extend downwards through the white fibrous substance of the brain in a perfectly definite manner, affecting certain distinct strands which pass through the inner capsule, crura, and pons, into the anterior pyramids of the medulla oblongata, and from thence (partly crossing to the other side) downwards into the anterior (direct) and lateral (crossed) columns of the spinal cord.
(4) Anatomical proof of the continuity of the rolandic regions with these motor columns of the cord is also clearly given. Flechsig's 'Pyramidenbahn' forms an uninterrupted strand (distinctly traceable in human embryos, before its fibres have acquired their white 'medullary sheath') passing upwards from the pyramids of the medulla, and traversing the internal capsule and corona radiata to the convolutions in question (Fig. 10). None of the inferior gray matter of the brain seems to have any connection with this important fibrous strand. It passes directly from the cortex to the motor arrangements in the cord, depending for its proper nutrition (as the facts of degeneration show) on the influence of the cortical cells, just as motor nerves depend for their nutrition on that of the cells of the spinal cord. Electrical stimulation of this motor strand in any accessible part of its course has been shown in dogs to produce movements analogous to those which excitement of the cortical surface calls forth.
One of the most instructive proofs of motor localization in the cortex is that furnished by the disease now called aphemia, or motor Aphasia. Motor aphasia is neither loss of voice nor paralysis of the tongue or lips. The patient's voice is as strong as ever, and all the innervations of his hypoglossal and facial nerves, except those necessary for speaking, may go on perfectly well. He can laugh and cry, and even sing; but he either is unable to utter any words at all; or a few meaningless stock phrases form his only speech; or else he speaks incoherently and confusedly, mispronouncing, misplacing, and misusing his words in various degrees. Sometimes his speech is a mere broth of unintelligible syllables. In cases of pure motor aphasia the patient recognizes his mistakes and suffers acutely from them.
Fig. 10.—Schematic Transverse Section of Brain showing Motor Strand.—After Edinger.
Now whenever a patient dies in such a condition as this, and an examination of his brain is permitted, it is found that the lowest frontal gyrus (see Fig. 11) is the seat of injury. Broca first noticed this fact in 1861, and since then the gyrus has gone by the name of Broca's convolution.
Fig. 11.—Schematic Profile of Left Hemisphere, with the parts shaded whose destruction causes motor ('Broca') and sensory ('Wernicke') Aphasia.
The injury in right-handed people is found on the left hemisphere, and in left-handed people on the right hemisphere. Most people, in fact, are left-brained, that is, all their delicate and specialized movements are handed over to the charge of the left hemisphere. The ordinary right-handedness for such movements is only a consequence of that fact, a consequence which shows outwardly on account of that extensive decussation of the fibres whereby most of those from the left hemisphere pass to the right half of the body only. But the left-brainedness might exist in equal measure and not show outwardly. This would happen wherever organs on both sides of the body could be governed by the left hemisphere; and just such a case seems offered by the vocal organs, in that highly delicate and special motor service which we call speech. Either hemisphere can innervate them bilaterally, just as either seems able to innervate bilaterally the muscles of the trunk, ribs, and diaphragm. Of the special movements of speech, however, it would appear (from the facts of aphasia) that the left hemisphere in most persons habitually takes exclusive charge. With that hemisphere thrown out of gear, speech is undone; even though the opposite hemisphere still be there for the performance of less specialized acts, such as the various movements required in eating.
It will be noticed that Broca's region is homologous with the parts ascertained to produce movements of the lips, tongue, and larynx when excited by electric currents in apes (cf. Fig. 6, p. 34). The evidence is therefore as complete as it well can be that the motor incitations to these organs leave the brain by the lower frontal region.
Victims of motor aphasia generally have other disorders. One which interests us in this connection has been called agraphia: they have lost the power to write. They can read writing and understand it; but either cannot use the pen at all or make egregious mistakes with it. The seat of the lesion here is less well determined, owing to an insufficient number of good cases to conclude from.[12] There is no doubt, however, that it is (in right-handed people) on the left side, and little doubt that it consists of elements of the hand-and-arm region specialized for that service. The symptom may exist when there is little or no disability in the hand for other uses. If it does not get well, the patient usually educates his right hemisphere, i.e. learns to write with his left hand. In other cases of which we shall say more a few pages later on, the patient can write both spontaneously and at dictation, but cannot read even what he has himself written! All these phenomena are now quite clearly explained by separate brain-centres for the various feelings and movements and tracts for associating these together. But their minute discussion belongs to medicine rather than to general psychology, and I can only use them here to illustrate the principles of motor localization.[13] Under the heads of sight and hearing I shall have a little more to say.
The different lines of proof which I have taken up establish conclusively the proposition that all the motor impulses which leave the cortex pass out, in healthy animals, from the convolutions about the fissure of Rolando.
When, however, it comes to defining precisely what is involved in a motor impulse leaving the cortex, things grow more obscure. Does the impulse start independently from the convolutions in question, or does it start elsewhere and merely flow through? And to what particular phase of psychic activity does the activity of these centres correspond? Opinions and authorities here divide; but it will be better, before entering into these deeper aspects of the problem, to cast a glance at the facts which have been made out concerning the relations of the cortex to sight, hearing, and smell.
Sight.
Ferrier was the first in the field here. He found, when the angular convolution (that lying between the 'intra parietal' and 'external occipital' fissures, and bending round the top of the fissure of Sylvius, in Fig. 6) was excited in the monkey, that movements of the eyes and head as if for vision occurred; and that when it was extirpated, what he supposed to be total and permanent blindness of the opposite eye followed. Munk almost immediately declared total and permanent blindness to follow from destruction of the occipital lobe in monkeys as well as dogs, and said that the angular gyrus had nothing to do with sight, but was only the centre for tactile sensibility of the eyeball. Munk's absolute tone about his observations and his theoretic arrogance have led to his ruin as an authority. But he did two things of permanent value. He was the first to distinguish in these vivisections between sensorial and psychic blindness, and to describe the phenomenon of restitution of the visual function after its first impairment by an operation; and the first to notice the hemiopic character of the visual disturbances which result when only one hemisphere is injured. Sensorial blindness is absolute insensibility to light; psychic blindness is inability to recognize the meaning of the optical impressions, as when we see a page of Chinese print but it suggests nothing to us. A hemiopic disturbance of vision is one in which neither retina is affected in its totality, but in which, for example, the left portion of each retina is blind, so that the animal sees nothing situated in space towards its right. Later observations have corroborated this hemiopic character of all the disturbances of sight from injury to a single hemisphere in the higher animals; and the question whether an animal's apparent blindness is sensorial or only psychic has, since Munk's first publications, been the most urgent one to answer, in all observations relative to the function of sight.
Goltz almost simultaneously with Ferrier and Munk reported experiments which led him to deny that the visual function was essentially bound up with any one localized portion of the hemispheres. Other divergent results soon came in from many quarters, so that, without going into the history of the matter any more, I may report the existing state of the case as follows:[14]
In fishes, frogs, and lizards vision persists when the hemispheres are entirely removed. This is admitted for frogs and fishes even by Munk, who denies it for birds.
All of Munk's birds seemed totally blind (blind sensorially) after removal of the hemispheres by his operation. The following of a candle by the head and winking at a threatened blow, which are ordinarily held to prove the retention of crude optical sensations by the lower centres in supposed hemisphereless pigeons, are by Munk ascribed to vestiges of the visual sphere of the cortex left behind by the imperfection of the operation. But Schrader, who operated after Munk and with every apparent guarantee of completeness, found that all his pigeons saw after two or three weeks had elapsed, and the inhibitions resulting from the wound had passed away. They invariably avoided even the slightest obstacles, flew very regularly towards certain perches, etc., differing toto cælo in these respects with certain simply blinded pigeons who were kept with them for comparison. They did not pick up food strewn on the ground, however. Schrader found that they would do this if even a small part of the frontal region of the hemispheres was left, and ascribes their non-self-feeding when deprived of their occipital cerebrum not to a visual, but to a motor, defect, a sort of alimentary aphasia.[15]
In presence of such discord as that between Munk and his opponents one must carefully note how differently significant is loss, from preservation, of a function after an operation on the brain. The loss of the function does not necessarily show that it is dependent on the part cut out; but its preservation does show that it is not dependent: and this is true though the loss should be observed ninety-nine times and the preservation only once in a hundred similar excisions. That birds and mammals can be blinded by cortical ablation is undoubted; the only question is, must they be so? Only then can the cortex be certainly called the 'seat of sight.' The blindness may always be due to one of those remote effects of the wound on distant parts, inhibitions, extensions of inflammation,—interferences, in a word,—upon which Brown-Séquard and Goltz have rightly insisted, and the importance of which becomes more manifest every day. Such effects are transient; whereas the symptoms of deprivation (Ausfallserscheinungen, as Goltz calls them) which come from the actual loss of the cut-out region must from the nature of the case be permanent. Blindness in the pigeons, so far as it passes away, cannot possibly be charged to their seat of vision being lost, but only to some influence which temporarily depresses the activity of that seat. The same is true mutatis mutandis of all the other effects of operations, and as we pass to mammals we shall see still more the importance of the remark.
In rabbits loss of the entire cortex seems compatible with the preservation of enough sight to guide the poor animals' movements, and enable them to avoid obstacles. Christiani's observations and discussions seem conclusively to have established this, although Munk found that all his animals were made totally blind.[16]
In dogs also Munk found absolute stone-blindness after ablation of the occipital lobes. He went farther and mapped out determinate portions of the cortex thereupon, which he considered correlated with definite segments of the two retinæ, so that destruction of given portions of the cortex produces blindness of the retinal centre, top, bottom, or right or left side, of the same or opposite eye. There seems little doubt that this definite correlation is mythological. Other observers, Hitzig, Goltz, Luciani, Loeb, Exner, etc., find, whatever part of the cortex may be ablated on one side, that there usually results a hemiopic disturbance of both eyes, slight and transient when the anterior lobes are the parts attacked, grave when an occipital lobe is the seat of injury, and lasting in proportion to the latter's extent. According to Loeb, the defect is a dimness of vision ('hemiamblyopia') in which (however severe) the centres remain the best seeing portions of the retina, just as they are in normal dogs. The lateral or temporal part of each retina seems to be in exclusive connection with the cortex of its own side. The centre and nasal part of each seems, on the contrary, to be connected with the cortex of the opposite hemispheres. Loeb, who takes broader views than any one, conceives the hemiamblyopia as he conceives the motor disturbances, namely, as the expression of an increased inertia in the whole optical machinery, of which the result is to make the animal respond with greater effort to impressions coming from the half of space opposed to the side of the lesion. If a dog has right hemiamblyopia, say, and two pieces of meat are hung before him at once, he invariably turns first to the one on his left. But if the lesion be a slight one, shaking slightly the piece of meat on his right (this makes of it a stronger stimulus) makes him seize upon it first. If only one piece of meat be offered, he takes it, on whichever side it be.
When both occipital lobes are extensively destroyed total blindness may result. Munk maps out his 'Sehsphäre' definitely, and says that blindness must result when the entire shaded part, marked A, A, in Figs. 12 and 13, is involved in the lesion. Discrepant reports of other observations he explains as due to incomplete ablation.
Fig. 12.Fig. 13.
The Dog's visual centre according to Munk, the entire striated region, A, A, being the exclusive seat of vision, and the dark central circle, A', being correlated with the retinal centre of the opposite eye.
Luciani, Goltz, and Lannegrace, however, contend that they have made complete bilateral extirpations of Munk's Sehsphäre more than once, and found a sort of crude indiscriminating sight of objects to return in a few weeks.[17] The question whether a dog is blind or not is harder to solve than would at first appear; for simply blinded dogs, in places to which they are accustomed, show little of their loss and avoid all obstacles; whilst dogs whose occipital lobes are gone may run against things frequently and yet see notwithstanding. The best proof that they may see is that which Goltz's dogs furnished: they carefully avoided, as it seemed, strips of sunshine or paper on the floor, as if they were solid obstacles. This no really blind dog would do. Luciani tested his dogs when hungry (a condition which sharpens their attention) by strewing pieces of meat and pieces of cork before them. If they went straight at them, they saw; and if they chose the meat and left the cork, they saw discriminatingly. The quarrel is very acrimonious; indeed the subject of localization of functions in the brain seems to have a peculiar effect on the temper of those who cultivate it experimentally. The amount of preserved vision which Goltz and Luciani report seems hardly to be worth considering, on the one hand; and on the other, Munk admits in his penultimate paper that out of 85 dogs he only 'succeeded' 4 times in his operation of producing complete blindness by complete extirpation of his 'Sehsphäre.'[18]
Fig. 14.—Distribution of the Visual Function in the Cortex, according to Luciani.
The safe conclusion for us is that Luciani's diagram, Fig. 14, represents something like the truth. The occipital lobes are far more important for vision than any other part of the cortex, so that their complete destruction makes the animal almost blind. As for the crude sensibility to light which may then remain, nothing exact is known either about its nature or its seat.
In the monkey, doctors also disagree. The truth seems, however, to be that the occipital lobes in this animal also are the part connected most intimately with the visual function. The function would seem to go on when very small portions of them are left, for Ferrier found no 'appreciable impairment' of it after almost complete destruction of them on both sides. On the other hand, he found complete and permanent blindness to ensue when they and the angular gyri in addition were destroyed on both sides. Munk, as well as Brown and Schaefer, found no disturbance of sight from destroying the angular gyri alone, although Ferrier found blindness to ensue. This blindness was probably due to inhibitions exerted in distans, or to cutting of the white optical fibres passing under the angular gyri on their way to the occipital lobes. Brown and Schaefer got complete and permanent blindness in one monkey from total destruction of both occipital lobes. Luciani and Seppili, performing this operation on two monkeys, found that the animals were only mentally, not sensorially, blind. After some weeks they saw their food, but could not distinguish by sight between figs and pieces of cork. Luciani and Seppili seem, however, not to have extirpated the entire lobes. When one lobe only is injured the affection of sight is hemiopic in monkeys: in this all observers agree. On the whole, then, Munk's original location of vision in the occipital lobes is confirmed by the later evidence.[19]
In man we have more exact results, since we are not driven to interpret the vision from the outward conduct On the other hand, however, we cannot vivisect, but must wait for pathological lesions to turn up. The pathologists who have discussed these (the literature is tedious ad libitum) conclude that the occipital lobes are the indispensable part for vision in man. Hemiopic disturbance in both eyes comes from lesion of either one of them, and total blindness, sensorial as well as psychic, from destruction of both.
Hemiopia may also result from lesion in other parts, especially the neighboring angular and supra-marginal gyri, and it may accompany extensive injury in the motor region of the cortex. In these cases it seems probable that it is due to an actio in distans, probably to the interruption of fibres proceeding from the occipital lobe. There seem to be a few cases on record where there was injury to the occipital lobes without visual defect. Ferrier has collected as many as possible to prove his localization in the angular gyrus.[20] A strict application of logical principles would make one of these cases outweigh one hundred contrary ones. And yet, remembering how imperfect observations may be, and how individual brains may vary, it would certainly be rash for their sake to throw away the enormous amount of positive evidence for the occipital lobes. Individual variability is always a possible explanation of an anomalous case. There is no more prominent anatomical fact than that of the 'decussation of the pyramids,' nor any more usual pathological fact than its consequence, that left-handed hemorrhages into the motor region produce right-handed paralyses. And yet the decussation is variable in amount, and seems sometimes to be absent altogether.[21] If, in such a case as this last, the left brain were to become the seat of apoplexy, the left and not the right half of the body would be the one to suffer paralysis.
The schema on the opposite page, copied from Dr. Seguin, expresses, on the whole, the probable truth about the regions concerned in vision. Not the entire occipital lobes, but the so-called cunei, and the first convolutions, are the cortical parts most intimately concerned. Nothnagel agrees with Seguin in this limitation of the essential tracts.[22]
A most interesting effect of cortical disorder is mental blindness. This consists not so much in insensibility to optical impressions, as in inability to understand them. Psychologically it is interpretable as loss of associations between optical sensations and what they signify; and any interruption of the paths between the optic centres and the centres for other ideas ought to bring it about. Thus, printed letters of the alphabet, or words, signify certain sounds and certain articulatory movements. If the connection between the articulating or auditory centres, on the one hand, and the visual centres on the other, be ruptured,
Fig. 15.—Scheme of the mechanism of vision, after Seguin. The cuneus convolution (Cu) of the right occipital lobe is supposed to be injured, and all the parts which lead to it are darkly shaded to show that they fail to exert their function. F. O. are the intra-hemispheric optical fibres. P. O. C. is the region of the lower optic centres (corpora geniculata and quadrigemina). T. O. D. is the right optic tract; C, the chiasma; F. L. D. are the fibres going to the lateral or temporal half T of the right retina; and F. C. S. are those going to the central or nasal half of the left retina. O. D. is the right, and O. S. the left eyeball. The rightward half of each is therefore blind: in other words, the right nasal field, R. N. F., and the left temporal field, L. T. F., have become invisible to the subject with the lesion at Cu.
we ought a priori to expect that the sight of words would fail to awaken the idea of their sound, or the movement for pronouncing them. We ought, in short, to have alexia, or inability to read: and this is just what we do have in many cases of extensive injury about the fronto-temporal regions, as a complication of aphasic disease. Nothnagel suggests that whilst the cuneus is the seat of optical sensations, the other parts of the occipital lobe may be the field of optical memories and ideas, from the loss of which mental blindness should ensue. In fact, all the medical authors speak of mental blindness as if it must consist in the loss of visual images from the memory. It seems to me, however, that this is a psychological misapprehension. A man whose power of visual imagination has decayed (no unusual phenomenon in its lighter grades) is not mentally blind in the least, for he recognizes perfectly all that he sees. On the other hand, he may be mentally blind, with his optical imagination well preserved; as in the interesting case published by Wilbrand in 1887.[23] In the still more interesting case of mental blindness recently published by Lissauer,[24] though the patient made the most ludicrous mistakes, calling for instance a clothes-brush a pair of spectacles, an umbrella a plant with flowers, an apple a portrait of a lady, etc. etc., he seemed, according to the reporter, to have his mental images fairly well preserved. It is in fact the momentary loss of our non-optical images which makes us mentally blind, just as it is that of our non-auditory images which makes us mentally deaf. I am mentally deaf if, hearing a bell, I can't recall how it looks; and mentally blind if, seeing it, I can't recall its sound or its name. As a matter of fact, I should have to be not merely mentally blind, but stone-blind, if all my visual images were lost. For although I am blind to the right half of the field of view if my left occipital region is injured, and to the left half if my right region is injured, such hemianopsia does not deprive me of visual images, experience seeming to show that the unaffected hemisphere is always sufficient for production of these. To abolish them entirely I should have to be deprived of both occipital lobes, and that would deprive me not only of my inward images of sight, but of my sight altogether.[25] Recent pathological annals seem to offer a few such cases.[26] Meanwhile there are a number of cases of mental blindness, especially for written language, coupled with hemianopsia, usually of the rightward field of view. These are all explicable by the breaking down, through disease, of the connecting tracts between the occipital lobes and other parts of the brain, especially those which go to the centres for speech in the frontal and temporal regions of the left hemisphere. They are to be classed among disturbances of conduction or of association; and nowhere can I find any fact which should force us to believe that optical images need[27] be lost in mental blindness, or that the cerebral centres for such images are locally distinct from those for direct sensations from the eyes.[28]
Where an object fails to be recognized by sight, it often happens that the patient will recognize and name it as soon as he touches it with his hand. This shows in an interesting way how numerous the associative paths are which all end by running out of the brain through the channel of speech. The hand-path is open, though the eye-path be closed. When mental blindness is most complete, neither sight, touch, nor sound avails to steer the patient, and a sort of dementia which has been called asymbolia or apraxia is the result. The commonest articles are not understood. The patient will put his breeches on one shoulder and his hat upon the other, will bite into the soap and lay his shoes on the table, or take his food into his hand and throw it down again, not knowing what to do with it, etc. Such disorder can only come from extensive brain-injury.[29]
The method of degeneration corroborates the other evidence localizing the tracts of vision. In young animals one gets secondary degeneration of the occipital regions from destroying an eyeball, and, vice versa, degeneration of the optic nerves from destroying the occipital regions. The corpora geniculata, thalami, and subcortical fibres leading to the occipital lobes are also found atrophied in these cases. The phenomena are not uniform, but are indisputable;[30] so that, taking all lines of evidence together, the special connection of vision with the occipital lobes is perfectly made out. It should be added that the occipital lobes have frequently been found shrunken in cases of inveterate blindness in man.
Hearing.
Hearing is hardly as definitely localized as sight. In the dog, Luciani's diagram will show the regions which directly or indirectly affect it for the worse when injured. As with sight, one-sided lesions produce symptoms on both sides. The mixture of black dots and gray dots in the diagram is meant to represent this mixture of 'crossed' and 'uncrossed' connections, though of course no topographical exactitude is aimed at. Of all the region, the temporal lobe is the most important part; yet permanent absolute deafness did not result in a dog of Luciani's, even from bilateral destruction of both temporal lobes in their entirety.[31]
Fig. 16.—Luciani's Hearing Region.
In the monkey, Ferrier and Yeo once found permanent deafness to follow destruction of the upper temporal convolution (the one just below the fissure of Sylvius in Fig. 6) on both sides. Brown and Schaefer found, on the contrary, that in several monkeys this operation failed to noticeably affect the hearing. In one animal, indeed, both entire temporal lobes were destroyed. After a week or two of depression of the mental faculties this beast recovered and became one of the brightest monkeys possible, domineering over all his mates, and admitted by all who saw him to have all his senses, including hearing, 'perfectly acute.'[32] Terrible recriminations have, as usual, ensued between the investigators, Ferrier denying that Brown and Schaefer's ablations were complete,[33] Schaefer that Ferrier's monkey was really deaf.[34] In this unsatisfactory condition the subject must be left, although there seems no reason to doubt that Brown and Schaefer's observation is the more important of the two.
In man the temporal lobe is unquestionably the seat of the hearing function, and the superior convolution adjacent to the sylvian fissure is its most important part. The phenomena of aphasia show this. We studied motor aphasia a few pages back; we must now consider sensory aphasia. Our knowledge of this disease has had three stages: we may talk of the period of Broca, the period of Wernicke, and the period of Charcot. What Broca's discovery was we have seen. Wernicke was the first to discriminate those cases in which the patient can not even understand speech from those in which he can understand, only not talk; and to ascribe the former condition to lesion of the temporal lobe.[35] The condition in question is word-deafness, and the disease is auditory aphasia. The latest statistical survey of the subject is that by Dr. Allen Starr,[36] In the seven cases of pure word-deafness which he has collected, cases in which the patient could read, talk, and write, but not understand what was said to him, the lesion was limited to the first and second temporal convolutions in their posterior two thirds. The lesion (in right-handed, i.e. left-brained, persons) is always on the left side, like the lesion in motor aphasia. Crude hearing would not be abolished, even were the left centre for it utterly destroyed; the right centre would still provide for that. But the linguistic use of hearing appears bound up with the integrity of the left centre more or less exclusively. Here it must be that words heard enter into association with the things which they represent, on the one hand, and with the movements necessary for pronouncing them, on the other. In a large majority of Dr. Starr's fifty cases, the power either to name objects or to talk coherently was impaired. This shows that in most of us (as Wernicke said) speech must go on from auditory cues; that is, it must be that our ideas do not innervate our motor centres directly, but only after first arousing the mental sound of the words. This is the immediate stimulus to articulation; and where the possibility of this is abolished by the destruction of its usual channel in the left temporal lobe, the articulation must suffer. In the few cases in which the channel is abolished with no bad effect on speech we must suppose an idiosyncrasy. The patient must innervate his speech-organs either from the corresponding portion of the other hemisphere or directly from the centres of ideation, those, namely, of vision, touch, etc., without leaning on the auditory region. It is the minuter analysis of the facts in the light of such individual differences as these which constitutes Charcot's contribution towards clearing up the subject.
Every namable thing, act, or relation has numerous properties, qualities, or aspects. In our minds the properties of each thing, together with its name, form an associated group. If different parts of the brain are severally concerned with the several properties, and a farther part with the hearing, and still another with the uttering, of the name, there must inevitably be brought about (through the law of association which we shall later study) such a dynamic connection amongst all these brain-parts that the activity of any one of them will be likely to awaken the activity of all the rest. When we are talking as we think, the ultimate process is that of utterance. If the brain-part for that be injured, speech is impossible or disorderly, even though all the other brain-parts be intact: and this is just the condition of things which, on page 37, we found to be brought about by limited lesion of the left inferior frontal convolution. But back of that last act various orders of succession are possible in the associations of a talking man's ideas. The more usual order seems to be from the tactile, visual, or other properties of the things thought-about to the sound of their names, and then to the latter's utterance. But if in a certain individual the thought of the look of an object or of the look of its printed name be the process which habitually precedes articulation, then the loss of the hearing centre will pro tanto not affect that individual's speech. He will be mentally deaf, i.e. his understanding of speech will suffer, but he will not be aphasic. In this way it is possible to explain the seven cases of pure word-deafness which figure in Dr. Starr's table.
If this order of association be ingrained and habitual in that individual, injury to his visual centres will make him not only word-blind, but aphasic as well. His speech will become confused in consequence of an occipital lesion. Naunyn, consequently, plotting out on a diagram of the hemisphere the 71 irreproachably reported cases of aphasia which he was able to collect, finds that the lesions concentrate themselves in three places: first, on Broca's centre; second, on Wernicke's; third, on the supra-marginal and angular gyri under which those fibres pass which connect the visual centres with the rest of the brain[37] (see Fig. 17). With this result Dr. Starr's analysis of purely sensory cases agrees.
Fig. 17.
In a later chapter we shall again return to these differences in the effectiveness of the sensory spheres in different individuals. Meanwhile few things show more beautifully than the history of our knowledge of aphasia how the sagacity and patience of many banded workers are in time certain to analyze the darkest confusion into an orderly display.[38] There is no 'centre of Speech' in the brain any more than there is a faculty of Speech in the mind. The entire brain, more or less, is at work in a man who uses language. The subjoined diagram, from Ross, shows the four parts most critically concerned, and, in the light of our text, needs no farther explanation (see Fig. 18).
Fig. 18.
Smell.
Taste
we know little that is definite. What little there is points to the lower temporal regions again. Consult Ferrier as below.
Fig. 19.—Luciani's Olfactory Region in the Dog.
Touch.
Interesting problems arise with regard to the seat of tactile and muscular sensibility. Hitzig, whose experiments on dogs' brains fifteen years ago opened the entire subject which we are discussing, ascribed the disorders of motility observed after ablations of the motor region to a loss of what he called muscular consciousness. The animals do not notice eccentric positions of their limbs, will stand with their legs crossed, with the affected paw resting on its back or hanging over a table's edge, etc.; and do not resist our bending and stretching of it as they resist with the unaffected paw. Goltz, Munk, Schiff, Herzen, and others promptly ascertained an equal defect of cutaneous sensibility to pain, touch, and cold. The paw is not withdrawn when pinched, remains standing in cold water, etc. Farrier meanwhile denied that there was any true anæsthesia produced by ablations in the motor zone, and explains the appearance of it as an effect of the sluggish motor responses of the affected side.[40] Munk[41] and Schiff[42], on the contrary, conceive of the 'motor zone' as essentially sensory, and in different ways explain the motor disorders as secondary results of the anæsthesia which is always there. Munk calls the motor zone the Fühlsphäre of the animal's limbs, etc., and makes it coördinate with the Sehsphäre, the Hörsphäre, etc., the entire cortex being, according to him, nothing but a projection-surface for sensations, with no exclusively or essentially motor part. Such a view would be important if true, through its bearings on the psychology of volition. What is the truth? As regards the fact of cutaneous anaesthesia from motor-zone ablations, all other observers are against Ferrier, so that he is probably wrong in denying it. On the other hand, Munk and Schiff are wrong in making the motor symptoms depend on the anæsthesia, for in certain rare cases they have been observed to exist not only without insensibility, but with actual hyperæsthesia of the parts.[43] The motor and sensory symptoms seem, therefore, to be independent variables.
In monkeys the latest experiments are those of Horsley and Schaefer,[44] whose results Ferrier accepts. They find that excision of the hippocampal convolution produces transient insensibility of the opposite side of the body, and that permanent insensibility is produced by destruction of its continuation upwards above the corpus callosum, the so-called gyrus fornicatus (the part just below the 'calloso-marginal fissure' in Fig. 7). The insensibility is at its maximum when the entire tract comprising both convolutions is destroyed. Ferrier says that the sensibility of monkeys is 'entirely unaffected' by ablations of the motor zone,[45] and Horsley and Schaefer consider it by no means necessarily abolished.[46] Luciani found it diminislied in his three experiments on apes.[47]
Fig. 20.—Luciani's Tactile Region in the Dog.
In man we have the fact that one-sided paralysis from disease of the opposite motor zone may or may not be accompanied with anæsthesia of the parts. Luciani, who believes that the motor zone is also sensory, tries to minimize the value of this evidence by pointing to the insufficiency with which patients are examined. He himself believes that in dogs the tactile sphere extends backwards and forwards of the directly excitable region, into the frontal and parietal lobes (see Fig. 20). Nothnagel considers that pathological evidence points in the same direction;[48] and Dr. Mills, carefully reviewing the evidence, adds the gyri fornicatus and hippocampi to the cutaneo-muscular region in man.[49] If one compare Luciani's diagrams together (Figs. 14, 16, 19, 20) one will see that the entire parietal region of the dog's skull is common to the four senses of sight, hearing, smell, and touch, including muscular feeling. The corresponding region in the human brain (upper parietal and supra-marginal gyri—see Fig. 17, p. 56) seems to be a somewhat similar place of conflux. Optical aphasias and motor and tactile disturbances all result from its injury, especially when that is on the left side.[50] The lower we go in the animal scale the Page:Principles of Psychology (1890) v1.djvu/81 Page:Principles of Psychology (1890) v1.djvu/82 Page:Principles of Psychology (1890) v1.djvu/83 Page:Principles of Psychology (1890) v1.djvu/84 Page:Principles of Psychology (1890) v1.djvu/85 Page:Principles of Psychology (1890) v1.djvu/86 Page:Principles of Psychology (1890) v1.djvu/87 Page:Principles of Psychology (1890) v1.djvu/88 Page:Principles of Psychology (1890) v1.djvu/89 Page:Principles of Psychology (1890) v1.djvu/90 Page:Principles of Psychology (1890) v1.djvu/91 Page:Principles of Psychology (1890) v1.djvu/92 Page:Principles of Psychology (1890) v1.djvu/93 Page:Principles of Psychology (1890) v1.djvu/94 Page:Principles of Psychology (1890) v1.djvu/95 Page:Principles of Psychology (1890) v1.djvu/96 Page:Principles of Psychology (1890) v1.djvu/97 Page:Principles of Psychology (1890) v1.djvu/98 Page:Principles of Psychology (1890) v1.djvu/99 Page:Principles of Psychology (1890) v1.djvu/100
- ↑ It should be said that this particular cut commonly proves fatal. The text refers to the rare cases which survive.
- ↑ I confine myself to the frog for simplicity's sake. In higher animals, especially the ape and man, it would seem as if not only determinate combinations of muscles, but limited groups or even single muscles could be innervated from the hemispheres.
- ↑ I hope that the reader will take no umbrage at my so mixing the physical and mental, and talking of reflex acts and hemispheres and reminiscences in the same breath, as if they were homogeneous quantities and factors of one causal chain. I have done so deliberately; for although I admit that from the radically physical point of view it is easy to conceive of the chain of events amongst the cells and fibres as complete in itself, and that whilst so conceiving it one need make no mention of 'ideas,' I yet suspect that point of view of being an unreal abstraction. Reflexes in centres may take place even where accompanying feelings or ideas guide them. In another chapter I shall try to show reasons for not abandoning this common-sense position; meanwhile language lends itself so much more easily to the mixed way of describing, that I will continue to employ the latter. The more radical-minded reader can always read 'ideational process' for 'idea.'
- ↑ I shall call it hereafter for shortness 'the Meynert scheme;' for the child-and-flame example, as well as the whole general notion that the hemispheres are a supernumerary surface for the projection and association of sensations and movements natively coupled in the centres below, is due to Th. Meynert, the Austrian anatomist. For a popular account of his views, see his pamphlet 'Zur Mechanik des Gehirnbaues,' Vienna, 1874. His most recent development of them is embodied in his 'Psychiatry,' a clinical treatise on diseases of the forebrain, translated by B. Sachs, New York, 1885.
- ↑ Geschichte des Materialismus, 2d ed., II. p. 345.
- ↑ West Riding Asylum Reports, 1876, p. 267.
- ↑ For a thorough discussion of the various objections, see Ferrier's 'Functions of the Brain,' 2d ed., pp. 227–334. and François-Franck's 'Leçons sur les Fonctions Motrices du Cerveau' (1887), Leçon 31. The most minutely accurate experiments on irritation of cortical points are those of Paneth, in Pflüger's Archiv, vol 37, p. 528.—Recently the skull has been fearlessly opened by surgeons, and operations upon the human brain performed, sometimes with the happiest results. In some of these operations the cortex has been electrically excited for the purpose of more exactly localizing the spot, and the movements first observed in dogs and monkeys have then been verified in men.
- ↑ J. Loeb: 'Beiträge zur Physiologie des Grosshirns;' Pflüger's Archiv, xxxix. 293. I simplify the author's statement.
- ↑ Goltz: Pflüger's Archiv, XLII. 419.
- ↑ 'Hemiplegia' means one-sided palsy.
- ↑ Philosophical Transactions, vol. 179, pp. 6, 10 (1888). In a later paper (ibid. p. 305) Messrs. Beevor and Horsley go into the localization still more minutely, showing spots from which single muscles or single digits can be made to contract.
- ↑ Nothnagel und Naunyn: Die Localization in den Gehirnkrankheiten (Wiesbaden, 1887), p. 34.
- ↑ An accessible account of the history of our knowledge of motor aphasia is in W. A. Hammond's 'Treatise on the Diseases of the Nervous System,' chapter VII.
- ↑ The history up to 1885 may be found in A. Christiani: Zur Physiologie des Gehirnes (Berlin, 1885).
- ↑ Pflüger's Archiv, vol. 44, p. 176.Munk (Berlin Academy Sitzsungsberichte, 1889, XXXI) returns to the charge, denying the extirpations of Schrader to be complete: "Microscopic portions of the Sehsphäre must remain."
- ↑ A. Christiani: Zur Physiol. d. Gehirnes (Berlin, 1885), chaps, II, III, IV. H. Munk: Berlin Akad. Stzgsb. 1884, XXIV.
- ↑ Luciani und Seppili: Die Functions-Localization auf der Grosshirnrinde (Deutsch von Fraenkel), Leipzig, 1886, Dogs M, N, and S.Goltz in Pflüger's Archiv, vol. 34, pp. 490–6; vol. 42, p. 454.Cf. also Munk: Berlin Akad. Stzgsb. 1886, VII, VIII, pp. 113–121, and Loeb: Pflüger's Archiv, vol. 39, p. 337.
- ↑ Berlin Akad. Sitzungsberichte, 1886, VII, VIII, p. 124.
- ↑ H. Munk: Functionen der Grosshirnrinde (Berlin, 1881), pp. 36–40.Ferrier: Functions, etc., 2d ed., chap, IX, pt. I. Brown and Schaefer: Philos. Transactions, vol. 179, p. 321. Luciani u. Seppili, op. cit. pp. 131–138. Lannegrace found traces of sight with both occipital lobes destroyed, and in one monkey even when angular gyri and occipital lobes were destroyed altogether. His paper is in the Archives de Médecine Expérimentale for January and March, 1889. I only know it from the abstract in the Neurologisches Centralblatt, 1889, pp. 108–420. The reporter doubts the evidence of vision in the monkey. It appears to have consisted in avoiding obstacles and in emotional disturbance in the presence of men.
- ↑ Localization of Cerebral Disease (1878), pp. 117–8.
- ↑ For cases see Flechsig: Die Leitungsbahnen in Gehirn u. Rückenmark (Leipzig, 1876), pp. 112, 272; Exner's Untersuchungen, etc., p. 83; Ferrier's Localization, etc., p. 11; François-Franck's Cerveau Moteur, p. 63, note.
- ↑ E. C. Seguin: Hemianopsia of Cerebral Origin, in Journal of Nervous and Mental Disease, vol. XIII. p. 30.Nothnagel und Naunyn: Ueber die Localization der Gehirnkrankheiten (Wiesbaden, 1887), p. 10.
- ↑ Die Seelenblindheit, etc., p. 51 ff. The mental blindness was in this woman's case moderate in degree.
- ↑ Archiv f. Psychiatrie, vol. 21, p. 222.
- ↑ Nothnagel (loc. cit. p. 23) says: "Dies trifft aber nicht zu." He gives, however, no case in support of his opinion that double-sided cortical lesion may make one stone-blind and yet not destroy one's visual images; so that I do not know whether it is an observation of fact or an a priori assumption.
- ↑ In a case published by C. S. Freund: Archiv f. Psychiatrie, vol. XX, the occipital lobes were injured, but their cortex was not destroyed, on both sides. There was still vision. Cf. pp. 291–5.
- ↑ I say 'need,' for I do not of course deny the possible coexistence of the two symptoms. Many a brain-lesion might block optical associations and at the same time impair optical imagination, without entirely stopping vision. Such a case seems to have been the remarkable one from Charcot which I shall give rather fully in the chapter on Imagination.
- ↑ Freund (in the article cited above 'Ueber optische Aphasie und Seelenblindheit') and Bruns ('Ein Fall von Alexie,' etc., in the Neurologisches Centralblatt for 1888, pp. 581, 509) explain their cases by broken-down conduction. Wilbrand, whose painstaking monograph on mental blindness was referred to a moment ago, gives none but a priori reasons for his belief that the optical 'Erinnerungsfeld' must be locally distinct from the Wahrnehmungsfeld (cf. pp. 84, 93). The a priori reasons are really the other way. Mauthner ('Gehirn u. Auge' (1881), p. 487 ff.) tries to show that the 'mental blindness' of Munk's dogs and apes after occipital mutilation was not such, but real dimness of sight. The best case of mental blindness yet reported is that by Lissauer, as below. The reader will also do well to read Bernard: De l'Aphasie (1885) chap, V; Ballet: Le Langage Intérieur (1886), chap, VIII; and Jas. Ross's little book on Aphasia (1887), p. 74.
- ↑ For a case see Wernicke's Lehrb. d. Gebirnkrankheiten, vol. II. p. 554 (1881).
- ↑ The latest account of them is the paper 'Über die optiscben Centren u. Bahnen' by von Monakow in the Archiv für Psychiatrie, vol. XX. p. 714.
- ↑ Die Functions-Localization, etc., Dog X; see also p. 161.
- ↑ Philos. Trans., vol. 179, p. 312.
- ↑ Brain, vol. XI. p. 10.
- ↑ Ibid. p. 147.
- ↑ Der aphasische Symptomencomplex (1874). See in Fig. 11 the convolution marked Wernicke.
- ↑ 'The Pathology of Sensory Aphasia,' 'Brain,' July, 1889.
- ↑ Nothnagel und Naunyn: op. cit., plates.
- ↑ Ballet's and Bernard's works cited on p. 51 are the most accessible documents of Charcot's school. Bastian's book on the Brain as an Organ of Mind (last three chapters) is also good.
- ↑ For details, see Ferrier's 'Functions,' chap. IX. pt. III, and Chas. K. Mills: Transactions of Congress of American Physicians and Surgeons, 1888, vol. I. p. 278.
- ↑ Functions of the Brain, chap. X. § 14.
- ↑ Ueber die Functionen d. Grosshirnrinde (1881), p. 50.
- ↑ Lezioni di Fisiologia sperimentale sul sistema nervoso encefalico (1873), p. 537 ff. Also 'Brain,' vol. IX. p. 298.
- ↑ Bechterew (Pflüger's Archiv., vol. 35, p. 137) found no anæsthesia in a cat with motor symptoms from ablation of sigmoid gyrus. Luciani got hyperæsthesia coexistent with cortical motor defect in a dog, by simultaneously hemisecting the spinal cord (Luciani u. Seppili, op. cit. p. 234). Goltz frequently found hyperæsthesia of the whole body to accompany motor defect after ablation of both frontal lobes, and he once found it after ablating the motor zone (Pflüger's Archiv, vol. 34, p. 471).
- ↑ Philos. Transactions, vol. 179, p. 20 ff.
- ↑ Functions, p. 375.
- ↑ Pp. 15–17.
- ↑ Luciani u. Seppili, op. cit. pp. 275–288.
- ↑ Op. cit. p. 18.
- ↑ Trans. of Congress, etc., p. 272.
- ↑ See Exner's Unters. üb. Localization, plate XXV.