Inhibition is a vera causa, of that there can be no doubt. The pneumogastric nerve inhibits the heart, the splanchnic inhibits the intestinal movements, and the superior laryngeal those of inspiration. The nerve-irritations which may inhibit the contraction of arterioles are innumerable, and reflex actions are often repressed by the simultaneous excitement of other sensory nerves. For all such facts the reader must consult the treatises on physiology. What concerns us here is the inhibition exerted by different parts of the nerve-centres, when irritated, on the activity of distant parts. The flaccidity of a frog from 'shock,' for a minute or so after his medulla oblongata is cut, is an inhibition from the seat of injury which quickly passes away.
What is known as 'surgical shock '(unconsciousness, pallor, dilatation of splanchnic blood-vessels, and general syncope and collapse) in the human subject is an inhibition which lasts a longer time. Goltz, Freusberg, and others, cutting the spinal cord in dogs, proved that there were functions inhibited still longer by the wound, but which re-established themselves ultimately if the animal was kept alive. The lumbar region of the cord was thus found to contain independent vaso-motor centres, centres for erection, for control of the sphincters, etc., which could be excited to activity by tactile stimuli and as readily reinhibited by others simultaneously applied.[70] We may therefore plausibly suppose that the rapid reappearance of motility, vision, etc., after their first disappearance in consequence of a cortical mutilation, is due to the passing off of inhibitions exerted by the irritated surface of the wound. The only question is whether all restorations of function must be explained in this one simple way, or whether some part of them may not be owing to the formation of entirely new paths in the remaining centres, by which they become 'educated' to duties which they did not originally possess. In favor of an indefinite extension of the inhibition theory facts may be cited such as the following: In dogs whose disturbances due to cortical lesion have disappeared, they may in consequence of some inner or outer accident reappear in all their intensity for 24 hours or so and then disappear again.[71] In a dog made half blind by an operation, and then shut up in the dark, vision comes back just as quickly as in other similar dogs whose sight is exercised systematically every day.[72] A dog which has learned to beg before the operation recommences this practice quite spontaneously a week after a double-sided ablation of the motor zone.[73] Occasionally, in a pigeon (or even, it is said, in a dog) we see the disturbances less marked immediately after the operation than they are half an hour later.[74] This would be impossible were they due to the subtraction of the organs which normally carried them on. Moreover the entire drift of recent physiological and pathological speculation is towards enthroning inhibition as an ever-present and indispensable condition of orderly activity. We shall see how great is its importance, in the chapter on the Will. Mr. Charles Mercier considers that no muscular contraction, once begun, would ever stop without it, short of exhaustion of the system;[75] and Brown-Séquard has for years been accumulating examples to show how far its influence extends.[76] Under these circumstances it seems as if error might more probably lie in curtailing its sphere too much than in stretching it too far as an explanation of the phenomena following cortical lesion.[77]
On the other hand, if we admit no re-education of centres, we not only fly in the face of an a priori probability, but we find ourselves compelled by facts to suppose an almost incredible number of functions natively lodged in the centres below the thalami or even in those below the corpora quadrigemina. I will consider the a priori objection after first taking a look at the facts which I have in mind. They confront us the moment we ask ourselves just which are the parts which perform the functions abolished by an operation after sufficient time has elapsed for restoration to occur?
The first observers thought that they must be the corresponding parts of the opposite or intact hemisphere. But as long ago as 1875 Carville and Duret tested this by cutting out the fore-leg-centre on one side, in a dog, and then, after waiting till restitution had occurred, cutting it out on the opposite side as well. Goltz and others have done the same thing.[78] If the opposite side were really the seat of the restored function, the original palsy should have appeared again and been permanent. But it did not appear at all; there appeared only a palsy of the hitherto unaffected side. The next supposition is that the parts surrounding the cut-out region learn vicariously to perform its duties. But here, again, experiment seems to upset the hypothesis, so far as the motor zone goes at least; for we may wait till motility has returned in the affected limb, and then both irritate the cortex surrounding the wound without exciting the limb to movement, and ablate it, without bringing back the vanished palsy.[79] It would accordingly seem that the cerebral centres below the cortex must be the seat of the regained activities. But Goltz destroyed a dog's entire left hemisphere, together with the corpus striatum and the thalamus on that side, and kept him alive until a surprisingly small amount of motor and tactile disturbance remained.[80] These centres cannot here have accounted for the restitution. He has even, as it would appear,[81] ablated both the hemispheres of a dog, and kept him alive 51 days, able to walk and stand. The corpora striata and thalami in this dog were also practically gone. In view of such results we seem driven, with M. François-Franck,[82] to fall back on the ganglia lower still, or even on the spinal cord as the 'vicarious' organ of which we are in quest. If the abeyance of function between the operation and the restoration was due exclusively to inhibition, then we must suppose these lowest centres to be in reality extremely accomplished organs. They must always have done what we now find them doing after function is restored, even when the hemispheres were intact. Of course this is conceivably the case; yet it does not seem very plausible. And the a priori considerations which a moment since I said I should urge, make it less plausible still.
For, in the first place, the brain is essentially a place of currents, which run in organized paths. Loss of function can only mean one of two things, either that a current can no longer run in, or that if it runs in, it can no longer run out, by its old path. Either of these inabilities may come from a local ablation; and 'restitution' can then only mean that, in spite of a temporary block, an inrunning current has at last become enabled to flow out by its old path again—e.g., the sound of 'give your paw' discharges after some weeks into the same canine muscles into which it used to discharge before the operation. As far as the cortex itself goes, since one of the purposes for which it actually exists is the production of new paths,[83] the only question before us is: Is the formation of these particular 'vicarious' paths too much to expect of its plastic powers? It would certainly be too much to expect that a hemisphere should receive currents from optic fibres whose arriving-place within it is destroyed, or that it should discharge into fibres of the pyramidal strand if their place of exit is broken down. Such lesions as these must be irreparable within that hemisphere. Yet even then, through the other hemisphere, the corpus callosum, and the bilateral connections in the spinal cord, one can imagine some road by which the old muscles might eventually be innervated by the same incoming currents which innervated them before the block. And for all minor interruptions, not involving the arriving-place of the 'cortico-petal' or the place of exit of the 'cortico-fugal' fibres, roundabout paths of some sort through the affected hemisphere itself must exist, for every point of it is, remotely at least, in potential communication with every other point. The normal paths are only paths of least resistance. If they get blocked or cut, paths formerly more resistant become the least resistant paths under the changed conditions. It must never be forgotten that a current that runs in has got to run out somewhere; and if it only once succeeds by accident in striking into its old place of exit again, the thrill of satisfaction which the consciousness connected with the whole residual brain then receives will reinforce and fix the paths of that moment and make them more likely to be struck into again. The resultant feeling that the old habitual act is at last successfully back again, becomes itself a new stimulus which stamps all the existing currents in. It is matter of experience that such feelings of successful achievement do tend to fix in our memory whatever processes have led to them; and we shall have a good deal more to say upon the subject when we come to the Chapter on the Will.
My conclusion then is this: that some of the restitution of function (especially where the cortical lesion is not too great) is probably due to genuinely vicarious function on the part of the centres that remain; whilst some of it is due to the passing off of inhibitions. In other words, both the vicarious theory and the inhibition theory are true in their measure. But as for determining that measure, or saying which centres are vicarious, and to what extent they can learn new tricks, that is impossible at present.
FINAL CORRECTION OF THE MEYNERT SCHEME.
And now, after learning all these facts, what are we to think of the child and the candle-flame, and of that scheme which provisionally imposed itself on our acceptance after surveying the actions of the frog? (Cf. [pp. 25-6], supra.) It will be remembered that we then considered the lower centres en masse as machines for responding to present sense-impressions exclusively, and the hemispheres as equally exclusive organs of action from inward considerations or ideas; and that, following Meynert, we supposed the hemispheres to have no native tendencies to determinate activity, but to be merely superadded organs for breaking up the various reflexes performed by the lower centres, and combining their motor and sensory elements in novel ways. It will also be remembered that I prophesied that we should be obliged to soften down the sharpness of this distinction after we had completed our survey of the farther facts. The time has now come for that correction to be made.
Wider and completer observations show us both that the lower centres are more spontaneous, and that the hemispheres are more automatic, than the Meynert scheme allows. Schrader's observations in Goltz's Laboratory on hemisphereless frogs[84] and pigeons[85] give an idea quite different from the picture of these creatures which is classically current. Steiner's[86] observations on frogs already went a good way in the same direction, showing, for example, that locomotion is a well-developed function of the medulla oblongata. But Schrader, by great care in the operation, and by keeping the frogs a long time alive, found that at least in some of them the spinal cord would produce movements of locomotion when the frog was smartly roused by a poke, and that swimming and croaking could sometimes be performed when nothing above the medulla oblongata remained.[87] Schrader's hemisphereless frogs moved spontaneously, ate flies, buried themselves in the ground, and in short did many things which before his observations were supposed to be impossible unless the hemispheres remained. Steiner[88] and Vulpian have remarked an even greater vivacity in fishes deprived of their hemispheres. Vulpian says of his brainless carps[89] that three days after the operation one of them darted at food and at a knot tied on the end of a string, holding the latter so tight between his jaws that his head was drawn out of water. Later, "they see morsels of white of egg; the moment these sink through the water in front of them, they follow and seize them, sometimes after they are on the bottom, sometimes before they have reached it. In capturing and swallowing this food they execute just the same movements as the intact carps which are in the same aquarium. The only difference is that they seem to see them at less distance, seek them with less impetuosity and less perseverance in all the points of the bottom of the aquarium, but they struggle (so to speak) sometimes with the sound carps to grasp the morsels. It is certain that they do not confound these bits of white of egg with other white bodies, small pebbles for example, which are at the bottom of the water. The same carp which, three days after operation, seized the knot on a piece of string, no longer snaps at it now, but if one brings it near her, she draws away from it by swimming backwards before it comes into contact with her mouth."[90] Already on [pp. 9-10], as the reader may remember, we instanced those adaptations of conduct to new conditions, on the part of the frog's spinal cord and thalami, which led Pflüger and Lewes on the one hand and Goltz on the other to locate in these organs an intelligence akin to that of which the hemispheres are the seat.
When it comes to birds deprived of their hemispheres, the evidence that some of their acts have conscious purpose behind them is quite as persuasive. In pigeons Schrader found that the state of somnolence lasted only three or four days, after which time the birds began indefatigably to walk about the room. They climbed out of boxes in which they were put, jumped over or flew up upon obstacles, and their sight was so perfect that neither in walking nor flying did they ever strike any object in the room. They had also definite ends or purposes, flying straight for more convenient perching places when made uncomfortable by movements imparted to those on which they stood; and of several possible perches they always chose the most convenient. "If we give the dove the choice of a horizontal bar (Reck) or an equally distant table to fly to, she always gives decided preference to the table. Indeed she chooses the table even if it is several meters farther off than the bar or the chair." Placed on the back of a chair, she flies first to the seat and then to the floor, and in general "will forsake a high position, although it give her sufficiently firm support, and in order to reach the ground will make use of the environing objects as intermediate goals of flight, showing a perfectly correct judgment of their distance. Although able to fly directly to the ground, she prefers to make the journey in successive stages.... Once on the ground, she hardly ever rises spontaneously into the air."[91]