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
