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.

OM

The

FIG. 11.-Schematic Profile of Left Hemisphere, with the parts shaded whose destruction causes motor ( Broca ') and sensory (* Wernicke') Aphasia.

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 righthandedness 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, how

ever, 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.* 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.† Under the heads of sight and hearing I shall have a little more to say.

*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 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:*

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 cœlo in these respects with certain simply blinded pigeons who were kept with

*The history up to 1885 may be found in A. Christiani: Zur Physiologie des Gehirnes (Berlin, 1885).

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.*

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

Pflüger's Archiv, vol. 44, p. 176. Munk (Berlin Academy Sitzsungsberichte, 189, XXXI) returns to the charge, denying the extirpations of Schrader to be complete: "Microscopic portions of the Sehsphäre must remain."