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.[10]
(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-coordination 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. 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.[11]
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. 
Fig. 6.—Left Hemisphere of Monkey's Brain. Outer Surface. 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. 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.[12] 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.[13] 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.[14]
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. 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.
Fig. 8.—Right Hemisphere of Human Brain. Lateral Surface. 
Fig. 9.—Right Hemisphere of Human Brain. Mesial Surface. (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.
Fig. 10.—Schematic Transverse Section of Brain showing Motor Strand.—After Edinger. 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. 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. 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.
Fig. 11.—Schematic Profile of Left Hemisphere, with the parts shaded whose destruction causes motor ('Broca') and sensory ('Wernicke') Aphasia. 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). 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.[15] 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.[16] 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.