Part 16

It will be noted that although certain regions of the cortex are found connected closely with certain of the main sense organs, there are important receptive organs which do not appear to have any special region of cortex assigned to their sensual products. Thus, there is the "vestibular labyrinth" of the ear. This great receptive organ, so closely connected in function with the movements and adjustment of the postures of the head and eyes, and indeed of the whole body, is prominent in the co-ordination necessary for the equilibrium of the body, an essential part of the fundamental acts of progression, standing, &c. Yet neither structural nor functional connexion with any special region of the cortex has been traced as yet for the labyrinthine receptors. Perceptions of the position of the head and of the body are of course part of our habitual and everyday experience. It may perhaps be that these perceptions are almost entirely obtained through sense organs which are not labyrinthine, but visual, muscular, tactual, and so on. The labyrinth may, though it controls and adjusts the muscular

## activities which maintain the balance of the body, operate reflexly

without in its operation exciting of itself sensations. The results of the unconscious reflexes it initiated and guided would be perceptible through other organs of sense. But against this purely unconscious functioning of the labyrinth and its nervous apparatus stands the fact that galvanic stimulation of the labyrinth is accompanied by well-known distinctive sensations--including giddiness, &c. Moreover, the prominent factor in sea-sickness, a disorder richly suffused with sensations, is probably the labyrinth. Yet there is marked absence of evidence of any special and direct connexion between the _cortex cerebri_ and the labyrinth organs.

Also there is curiously little evidence of connexion of the cortex with the nervous paths of conduction concerned with pain. As far as the present writer can find from reference to books and from the clinical experience of others, "pain" is unknown as an _aura_ in cortical epilepsy, or at most is of equivocal occurrence.

The preceding brief exposition of some of the main features of the localization of function in the _cortex cerebri_, gradually deciphered by patient inquiry, shows that the scheme of partition of function so far perceptible does not follow the quaint lines of analysis of the phrenologists with their supposed mental entities, so-called "faculties." On the contrary it is based, as some of those who early favoured a differential arrangement of function in the cerebrum had surmised, on the _separateness of the incoming channels from peripheral organs of sense_. These organs fall into groups separate one from another not only by reason of their spatial differentiation at the surface and in the thickness of the body, but also because each group generates sensations which introspection tells us are of a species unbridgeably separate from those generated by the other groups. Between sensations of hearing and sensations of sight there is a dissimilarity across which no intermediate series of sensual phenomena extend. The two species of sensations are wholly disparate. Similarly there is a total and impassable gap between sensations of touch and sensations of sight and sound. In other words the sensations fall into groups which are wholly disparate and are hence termed species. But within each species there exist multifold varieties of the specific sensation, e.g. sensations of red, of yellow, &c. We should expect, therefore, that the conducting paths from the receptive organs which in their function as sense-organs yield wholly disparate sensations would in so far as subserving sensation diverge and pass to separate neural mechanisms. That these sense-organs should in fact be found to possess in the cortex of the cerebrum separate fields for their sensual nervous apparatus is, therefore, in harmony with what would be the _a priori_ supposition.

But, as emphasized at the beginning of this article, the receptive organs belonging to the surfaces and the depths of the body and forming the starting-points for the whole system of the afferent nerves, have two functions more or less separate. One of these functions is to excite sensations and the other is to excite movements, by reflex action, especially in glands and muscles. In this latter function, namely the reflexifacient, all that the receptive organs effect is effected by means of the efferent nerves. They all have to use the efferent, especially the motor, nerves of the body. So rich is the connexion of the receptive organs with the efferent nerves that it is not improbable that, through the central nervous organ, each receptive organ is connected with every motor nerve of the whole nervous system,--the facts of strychnine poisoning show that if this is not literally true it is at least approximately so. Hence one of the goals to which each afferent fibre from a receptive organ leads is a number of motor nerves. Their conducting paths must, therefore, converge in passing to the starting-points of the motor nerves; because these latter are instruments common to the use of a number of different receptive organs in so far as they excite reflex actions. On the other hand those of their conducting paths which are concerned in the genesis of sensation, instead of converging, diverge, at least as far as the _cortex cerebri_, or if not divergent, remain separate. These considerations would make it appear likely that the conducting path from each receptive organ divides in the central nervous system into two main lines, one of which goes off to its own particular region of the _cortex cerebri_ whither run conductors only of similar sensual species to itself, while the other main line passes with many others to a great motor station where, as at a telephone exchange, coordinate use of the outgoing lines is assured to them all. Now there is in fact a portion of the cortex in mammals the functions of which are so pre-eminently motor, as judged by our present methods, that it is commonly designated the _motor cortex_ (see fig. 24). This region of the cortex occupies in the Primates, including Man, the pre-central gyrus. Among the items of evidence which reveal its motor capabilities are the following.

[Illustration: FIG. 24.--Diagram of the Topography of the Main Groups of Foci in the Motor Field of Chimpanzee.]

_The Precentral or Motor Region of the Cortex._--The application to it of electric currents excites movements in the skeletal muscles. The movements occur in the half of the body of the side crossed from that of the hemisphere excited. The "motor representation," as it is termed, is in the cortex better described as a representation of definite actions than of particular muscles. The actions "represented" in the top part of the gyrus, namely next the great longitudinal fissure, move the leg; those in the lowest part of the gyrus belong to the tongue and mouth. The topical distribution along the length of the gyrus may be described in a general way as following a sequence resembling that of the motor representation in the spinal cord, the top of the gyrus being taken as corresponding with the caudal end of the spinal cord. The sequence as the gyrus is followed downwards runs: perineum, foot, knee, hip, abdomen, chest, shoulder, elbow, wrist, hand, eyelids and ear, nose, mouth and tongue. The nature of the movement is very fairly constant for separate points of this motor cortex as observed both in the same and in similar experiments. Thus flexion of the arm will be excitable from one set of points, and extension of the arm from another set of points; opening of the jaw from one set and closure from another, and so on. These various movements if excited strongly tend to have characters like those of the movements seen in an epileptic convulsion. Strong stimulation excites in fact a convulsion like that of epilepsy, beginning with the movement usual for the point stimulated and spreading so as to assume the proportions of a convulsion affecting the entire skeletal musculature of one half or even of the whole body. The resemblance to an epileptic seizure is the closer because the movement before it subsides becomes clonic (rhythmic) as in epilepsy. The determination of the exact spots of cortex in which are represented the various movements of the body has served a useful practical purpose in indicating the particular places in the cortex which are the seat of disease. These the physician can localize more exactly by reason of this knowledge. Hence the surgeon, if the nature of the disease is such as can be dealt with by surgical means, can without unnecessarily damaging the skull and brain, proceed directly to the point which is the seat of the mischief.

The motor representation of certain parts of the body is much more liberal than is that of others. There is little correspondence between the mere mass of musculature involved and the area of the cortex devoted to its representation. Variety of movement rather than force or energy of movement seems to demand extent of cortex. The cortical area for the thumb is larger than those for the whole abdomen and chest combined. The cortical area for the tongue is larger than that for the neck. Different movements of one and the same part are very unequally represented in the cortex. Thus, flexion of the leg is more extensively represented than is extension, opening of the jaw has a much larger cortical area than has closure of the jaws. It is interesting that certain agents, for instance strychnine, and the poison of the bacilli which cause the disease known as tetanus or lock-jaw, upset this normal topography, and replace in the cortex flexion of the limb by extension of the limb, and opening of the jaw by closure of the jaw. There is, however, no evidence that they do this by changing in any way the cortical mechanisms themselves. It is more likely that their action is confined to the lower centres, bulbar and spinal, upon which the discharge excited from the cortex plays. The change thus induced in the movement excited by the cortex does, however, show that the point of cortex which causes for instance opening of the mouth is connected with the motor nerves to the closing muscles as well as with those of the opening muscles. This is an item of evidence that the "centres" of the cortex are connected with the motor nerves of antagonistic muscles in such a way that when the "centre" excites one set of the muscles to contract, it simultaneously under normal circumstances causes inhibition of the motor neurones of the opposed set of muscles (reciprocal innervation). In the great majority of movements excited from the motor cortex of a single hemisphere of the cerebrum, the movement evoked is confined to one side of the body, namely to that opposite to the hemisphere stimulated. There are, however, important exceptions to this. Thus, adduction of both vocal cords is excited from the cortex of either hemisphere. The movement of closure of the eyelids is usually bilateral, unless the stimulation be very weak; then the movement is of the eyelids of the opposite side only. The same holds true for the movements of the jaw. It, therefore, seems clear that with many movements which are usually bilaterally performed in ordinary life, such as opening of the jaw, blinking, &c., the symmetrical areas of the motor regions of both hemispheres are simultaneously in action.

In regard to all these movements elicitable by artificial stimuli from the motor cortex it is obvious that were there clearer evidence that the pallial region from which they are elicitable is fairly directly connected with corticopetal paths subserving cutaneous sensation or "muscular sense," the movements might be regarded as falling into the category of higher reflexes connected with the organs of touch, muscular sense, &c., just as the movements of the eyeball excitable from the visual cortex may be regarded as higher reflexes connected with vision. The evidence of the connexion of the reactions of the motor cortex with cutaneous and muscular senses appears, however, scarcely sufficient to countenance at present this otherwise plausible view, which has on general grounds much to commend it.

It is remarkable that movements of the eyeball itself, i.e. apart from movement of the lids, are not in the category of movements elicitable from the precentral gyrus, the "motor" cortex. They are found represented in a region farther forward, namely in front of the precentral gyrus altogether, and occupying a scattered set of points in the direction frontal from the areas for movements of arm and face. This frontal area yields on excitation conjugate movements of both eyeballs extremely like if not exactly similar to those yielded by excitation of the occipital (visual) region of the cortex. It is supposed by some that this frontal area yielding eye-movements has its function in this respect based upon afferent conductors from other parts of the eyeball than the retina, for instance upon kinaesthetic (Bastian) impressions or upon sensual impressions derived from the cornea and the coats of the eyeball including the ciliary and iris muscles. The ocular muscles are certainly a source of centripetal impulses, but their connexion with the cortex is not clear as to either their nature or their seat. The question seems for the present to allow no clearer answer. It is certain, however, that the frontal area of eye movements has corticofugal paths descending from it to the lower motor centres of the eyeballs quite independent of those descending from the occipital (visual) area of eye-movements. Further, it seems clear that in many animals there is another cortical region, a third region, the region which we saw above might be considered auditory, where movements of the eyeball similar to those elicitable in the occipital and frontal cortex can be provoked. A. Tschermak is inclined to give the eyeball movements of the frontal region the significance of reflex movements which carry the visual field in various directions in answer to demands made by sensory data derived from touch, &c., as for instance from the hand. The movements of the eyeballs elicitable from the occipital region of the cortex he regards as probably concerned with directing the gaze toward something seen, for instance, in the peripheral field of vision. The occipital movement would, therefore, be excited through the retina, and would result in bringing the yellow spot region of the retinae of both eyes to bear upon the object. This view has much to justify it. The movements of the eyeballs excited from the cortex of the auditory region would in a similar way be explicable as bringing the gaze to bear upon a direction in which a sound had been located, auditory initiation replacing the visual and tactual of the occipital and the frontal regions respectively.

Turning from these still speculative matters to others less suggestive but of actual ascertainment, we find that the motor nature of the precentral cortex as ascertained by electric stimuli is further certified by the occurrence of disturbance and impairment of motor power and adjustment following destruction of that region of the cortex. The movements which such a part as a limb executes are of course manifold in purpose. The hind limb of a dog is used for standing, for stepping, for scratching, for squatting, and, where a dog, for instance, has been trained to stand or walk on its hind legs alone, for skilled acts requiring a special training for their acquisition. It is found that when the motor area of the brain has been destroyed, the limb is at first paralysed for all these movements, but after a time the limb recovers the ability to execute some of them, though not all. The scratching movement suffers little, and rapid improvement after cerebral injury soon effaces the impairment, at first somewhat pronounced, in the use of the limb for walking, running, &c., and ordinary movements of progression. Even when both hemispheres have been destroyed the dog can still stand and walk and run. Destruction of the motor region of the cortex renders the fore limbs of the dog unable to execute such skilled movements as the steadying of a bone for gnawing or the trained act of offering the paw in answer to the command of the master. Skilled acts of the limb, apart from conjoined movements in which it, together with all the other limbs, takes part, assume of course a larger share of the office of the limb in the Primates than in the dog; and this is especially true for the fore limb. It is when the fore-foot becomes a hand that opportunity is given for its more skilled individual use and for its training in movements as a tool, or for the handling of tools and weapons. It is these movements which suffer most heavily and for the longest period after injury of the motor region of the cortex. Hence the disablement ensuing upon injury to the cortex would be expected to be most apparent in the Primates; and it is so, and most of all in Man. Further, in Man there ensues a condition called "contracture," which is not so apparent or frequent a result in other animals,--indeed, does not occur at all in other animals except the monkey. In contracture the muscles of the paretic limb are not flaccid, as they are usually in paralysis, but they are tense and the limb is more or less rigidly fixed by them in a certain position, usually one of flexion at elbow and wrist. This condition does not occur at first, but gradually supervenes in the course of a number of weeks. In Man the destruction of the motor area of the cortex cripples the limb even for the part it should play in the combined limb movements of walking, &c., and cripples it to an extent markedly contrasting with the slight disturbances seen in the lower mammals, e.g. the dog.

As regards the recovery of motor power after lesions of the motor cortex, two processes seem at work which are termed respectively _restitution_ and _compensation_. By the former is understood the recovery obtained when a part of a "centre" is destroyed, and the rest of the centre, although thrown out of function at first, recovers and supplements the deficiency later. An example of restitution would be the recovery from temporary hemianopia caused by a small injury in one occipital lobe. By compensation is understood the improvement of an impaired nervous function, traceable to other centres different from those destroyed supplying means to compass the reaction originally dependent on the centres subsequently destroyed. Instances of such compensation are the recovery of taxis for equilibrium subsequent to destruction of the labyrinth of the ear, where the recovery is traceable to assistance obtained through the eye. It will be noted that these instances of recovery by restitution and by compensation respectively are taken, from cases of injury inflicted on receptive rather than on motor centres. It is doubtful how far they really apply to the undoubted improvement that does within certain limits progress and succeed in

## partially effacing the paresis immediately consequent on lesions of the

motor area. It has to be remembered that in all cases of traumatic injury to the nervous system, especially where the trauma implicates the central nervous organ, the first effects and impairment of function resulting are due to a mixed cause, namely on the one hand the mechanical rupture of conducting paths actually broken by solution of their continuity, and on the other hand the temporary interruption of conducting paths by "shock." Shock effects are not permanent: they pass off. They are supposed to be due to a change at the synapses connecting neurone with neurone in the grey matter. They amount in effect to a long-lasting and gradually subsiding inhibition.

For diseases of the brain see NEUROPATHOLOGY, INSANITY, SKULL (_Surgery_), &c. (C. S. S.)

FOOTNOTE:

[1] The literature of the pineal region is enormous. Studnicka (in _Oppels Vergleichende mikrosk. Anat._ Teile 4-5, 1904, 1905) gives 285 references. The present conception of the generalized arrangement is: ([alpha]) A single glandular median organ from the fore-brain called the paraphysis. ([beta]) A pouch of the ependymal roof of the ventricle called the dorsal sac. ([gamma]) A right and left epiphysis, one of which may be wholly or partially suppressed. These may change their position to anterior and posterior in some animals.

BRAINERD, DAVID (1718-1747), American missionary among the Indians, was born at Haddam, Connecticut, on the 20th of April 1718. He was orphaned at fourteen, and studied for nearly three years (1739-1742) at Yale. He then prepared for the ministry, being licensed to preach in 1742, and early in 1743 decided to devote himself to missionary work among the Indians. Supported by the Scottish "Society for Promoting Christian Knowledge," he worked first at Kaunaumeek, an Indian settlement about 20 m. from Stockbridge, Massachusetts, and subsequently, until his death, among the Delaware Indians in Pennsylvania (near Easton) and New Jersey (near Cranbury). His heroic and self-denying labours, both for the spiritual and for the temporal welfare of the Indians, wore out a naturally feeble constitution, and on the 19th of October 1747 he died at the house of his friend, Jonathan Edwards, in Northampton, Massachusetts.

His _Journal_ was published in two parts in 1746 by the Scottish Society for Promoting Christian Knowledge; and in 1749, at Boston, Jonathan Edwards published _An Account of the Life of the Late Rev. David Brainerd, chiefly taken from his own Diary and other Private Writings_, which has become a missionary classic. A new edition, with the _Journal_ and Brainerd's letters embodied, was published by Sereno E. Dwight at New Haven in 1822; and in 1884 was published what is substantially another edition, _The Memoirs of David Brainerd_, edited by James M. Sherwood.

BRAINERD, a city and the county-seat of Crow Wing county, Minnesota, U.S.A., on the E. bank of the Mississippi river, about 127 m. N.W. of Minneapolis. Pop. (1890) 5703; (1900) 7524, of whom 2193 were foreign-born; (1905) 8133; (1910) 8526. It is served by the Minnesota & International and the Northern Pacific railways. The latter maintains here large car and repair shops, and a sanatorium for its employees. There are also the Sisters of St Joseph hospital, a county court house, a public library and a Y.M.C.A. building. A dam across the Mississippi provides water power (about 60,000 H.P.) which is utilized extensively for manufacturing purposes. Lumbering is an important industry, and there are saw mills and planing mills, and an extensive creosote plant for treating railway ties and timber. There are also flour mills, paper and pulp mills, cigar factories, a brewery, a large foundry and a grain elevator. In 1906 large quantities of iron ore were discovered in the vicinity, the new range, the Cuyuna, running through the city from north-east to south-west. Brainerd, named in honour of David Brainerd, was settled in 1870, and chartered as a city in 1883.

BRAINTREE, a market town in the Maldon parliamentary division of Essex, England; 45 m. N.E. of London by a branch line from Witham of the Great Eastern railway. Pop. of urban district, 5330. The parish church of St Michael is a fine edifice of Early English work with later additions. A corn exchange, mechanics' institute and public hall may also be mentioned. The bishops of London had formerly a palace in the town, but there are no remains of the building. The manufactures of silk and crape have superseded that of woollen cloth, which was introduced by the Flemings who fled to England to escape the persecution of the duke of Alva. Matting and brushes are also made. On the north lies the large village of BOCKING, with the Perpendicular parish church of St Mary, similar industries, and a population of 3347.