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 senseimpressions 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* and pigeons † give an idea quite different from the picture of these creatures which is classically current. Steiner's observations on frogs

* Pflüger's Archiv, vol. 41, p. 75 (1887). Ibid., vol. 44, p. 175 (1889) Untersuchungen über die Physiologie des Froschhirns, 1885.

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.* 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. Steinert and Vulpian have remarked an even greater vivacity in fishes deprived of their hemispheres. Vulpian says of his brainless carpst 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

* Loc. cit. pp. 80, 82-3. Schrader also found a biting-reflex developed when the medulla oblongata is cut through just behind the cerebellum. + Berlin Akad. Sitzungsberichte for 1886.

Comptes Rendus, vol. 102, p. 90.

her mouth." 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."†

Young rabbits deprived of their hemispheres will stand, run, start at noises, avoid obstacles in their path, and give responsive cries of suffering when hurt. Rats will do the same, and throw themselves moreover into an attitude of defence. Dogs never survive such an operation if performed at once. But Goltz's latest dog, mentioned on p. Comptes Rendus de l'Acad. d. Sciences, vol. 102, p. 1530. Loc. cit. p. 216.

70, which is said to have been kept alive for fifty-one days after both hemispheres had been removed by a series of ablations and the corpora striata and thalami had softened away, shows how much the mid-brain centres and the cord can do even in the canine species. Taken together, the number of reactions shown to exist in the lower centres by these observations make out a pretty good case for the Meynert scheme, as applied to these lower animals. That scheme demands hemispheres which shall be mere supplements or organs of repetition, and in the light of these observations they obviously are so to a great extent. But the Meynert scheme also demands that the reactions of the lower centres shall all be native, and we are not absolutely sure that some of those which we have been considering may not have been acquired after the injury; and it furthermore demands that they should be machine-like, whereas the expression of some of them makes us doubt whether they may not be guided by an intelligence of low degree.

Even in the lower animals, then, there is reason to soften down that opposition between the hemispheres and the lower centres which the scheme demands. The hemispheres may, it is true, only supplement the lower centres, but the latter resemble the former in nature and have some small amount at least of 'spontaneity' and choice.

But when we come to monkeys and man the scheme well-nigh breaks down altogether; for we find that the hemispheres do not simply repeat voluntarily actions which the lower centres perform as machines. There are many functions which the lower centres cannot by themselves perform at all. When the motor cortex is injured in a man or a monkey genuine paralysis ensues, which in man is incurable, and almost or quite equally so in the ape. Dr. Seguin knew a man with hemi-blindness, from cortical injury, which had persisted unaltered for twenty-three years. 'Traumatic inhibition' cannot possibly account for this. The blindness must have been an Ausfallserscheinung,' due to the loss of vision's essential organ. It would seem, then, that in these higher creatures the lower centres must be less adequate than they are farther down in the zoological scale; and that even for certain elementary

combinations of movement and impression the co-operation of the hemispheres is necessary from the start. Even in birds and dogs the power of eating properly is lost when the frontal lobes are cut off.*

The plain truth is that neither in man nor beast are the hemispheres the virgin organs which our scheme called them. So far from being unorganized at birth, they must have native tendencies to reaction of a determinate sort.+ These are the tendencies which we know as emotions and instincts, and which we must study with some detail in later chapters of this book. Both instincts and emotions are reactions upon special sorts of objects of perception; they depend on the hemispheres; and they are in the first instance reflex, that is, they take place the first time the exciting object is met, are accompanied by no forethought or deliberation, and are irresistible. But they are modifiable to a certain extent by experience, and on later occasions of meeting the exciting object, the instincts especially have less of the blind impulsive character which they had at first. All this will be explained at some length in Chapter XXIV. Meanwhile we can say that the multiplicity of emotional and instinctive reactions in man, together with his extensive associative power, permit of extensive recouplings of the original sensory and motor partners. The consequences of one instinctive reaction often prove to be the inciters of an opposite reaction, and being suggested on later occasions by the original object, may then suppress the first reaction altogether, just as in the case of the child and the flame. For this education the hemispheres do not need

* Goltz: Pflüger's Archiv, vol. 42, p. 447; Schrader: ibid. vol. 44, p. 219 ff. It is possible that this symptom may be an effect of traumatic inhibition, however.

A few years ago one of the strongest arguments for the theory that the hemispheres are purely supernumerary was Soltmann's often-quoted observation that in new-born puppies the motor zone of the cortex is not excitable by electricity and only becomes so in the course of a fortnight, presumably after the experiences of the lower centres have educated it to motor duties. Paneth's later observations, however, seem to show that Soltmann may have been misled through overnarcotizing his victims (Pflüger's Archiv, vol. 37, p. 202). In the Neurologisches Centralblatt for 1889, p. 513, Bechterew returns to the subject on Soltmann's side with out, however, noticing Paneth's work.