H

longata, thereby cutting off the brain from all connection with the rest of the body, the frog will still continue to live, but with a very peculiarly modified activity. It ceases to breathe or swallow; it lies flat on its belly, and does not, like a normal frog, sit up on its forepaws, though its hind-legs are kept, as usual, folded against its body and immediately resume this position if drawn out. If thrown on its back it lies there quietly, without turning over like a normal frog. Locomotion and voice seem entirely abolished. If we suspend it by the nose, and irritate different portions of its skin by acid, it performs a set of remarkable ́defensive' movements calcu

Cb.

M

0 Th

FIG. 39.—C, H, cere

bral hemispheres; O Th, optic tha

lami;

L, optic

lobes; Cb, cerebellum; M O, medulla

oblongata; S C, spi

nal cord.

lated to wipe away the irritant. Thus, if the breast be touched, both fore-paws will rub it vigorously; if we touch the outer side of the elbow, the hind-foot of the same side will rise directly to the spot and wipe it. The back of the foot will rub the knee if that be attacked, whilst if the foot be cut away, the stump will make ineffectual movements, and then, in many frogs, a pause will come, as if for deliberation, succeeded by a rapid passage of the opposite unmutilated foot to the acidulated spot.

The most striking character of all these movements, after their teleological appropriateness, is their precision. They vary, in sensitive frogs and with a proper amount of irritation, so little as almost to resemble in their machinelike regularity the performances of a jumping-jack, whose legs must twitch whenever you pull the string. The spinal cord of the frog thus contains arrangements of cells and fibres fitted to convert skin-irritations into movements of defence. We may call it the centre for defensive movements in this animal. We may indeed go farther than this, and by cutting the spinal cord in various places find that its

separate segments are independent mechanisms, for appropriate activities of the head and of the arms and legs respectively. The segment governing the arms is especially active, in male frogs, in the breeding season; and these members alone, with the breast and back appertaining to them, and everything else cut away, will actively grasp a finger placed between them and remain hanging to it for a considerable time.

Similarly of the medulla oblongata, optic lobes, and other centres between the spinal cord and the hemispheres of the frog. Each of them is proved by experiment to contain a mechanism for the accurate execution, in response to definite stimuli, of certain special acts. Thus with the medulla the animal swallows; with the medulla and cerebellum together he jumps, swims, and turns over from his back; with his optic lobes he croaks when pinched; etc. A frog which has lost his cerebral hemispheres alone is by an unpractised observer indistinguishable from a normal animal.

Not only is he capable, on proper instigation, of all the acts already mentioned, but he guides himself by sight, so that if an obstacle be set up between him and the light, and he be forced to move forward, he either jumps over it or swerves to one side. He manifests the sexual instinct at the proper seasons, and discriminates between male and female individuals of his own species. He is, in short, so similar in every respect to a normal frog that it would take a person very familiar with these animals to suspect anything wrong or wanting about him; but even then such a person would soon remark the almost entire absence of spontaneous motion-that is, motion unprovoked by any present incitation of sense. The continued movements of swimming, performed by the creature in the water, seem to be the fatal result of the contact of that fluid with its skin. They cease when a stick, for example, touches his hands. This is a sensible irritant towards which the feet are automatically drawn by reflex action, and on which the animal remains sitting. He manifests no hunger, and will

suffer a fly to crawl over his nose unsnapped at. Fear, too, seems to have deserted him. In a word, he is an extremely complex machine whose actions, so far as they go, tend to self-preservation; but still a machine, in this sense-that it seems to contain no incalculable element. By applying the right sensory stimulus to him we are almost as certain of getting a fixed response as an organist is of hearing a certain tone when he pulls out a certain stop.

But now if to the lower centres we add the cerebral hemispheres, or if, in other words, we make an intact animal the subject of our observations, all this is changed. In addition to the previous responses to present incitements of sense, our frog now goes through long and complex acts of locomotion spontaneously, or as if moved by what in ourselves we should call an idea. His reactions to outward stimuli vary their form, too. Instead of making simple defensive movements with his hind-legs, like a headless frog, if touched; or of giving one or two leaps and then sitting still like a hemisphereless one, he makes persistent and varied efforts of escape, as if, not the mere contact of the physiologist's hand, but the notion of danger suggested by it were now his spur. Led by the feeling of hunger, too, he goes in search of insects, fish, or smaller frogs, and varies his procedure with each species of victim. The physiologist cannot by manipulating him elicit croaking, crawling up a board, swimming or stopping, at will. His conduct has become incalculable-we can no longer foretell it exactly. Effort to escape is his dominant reaction, but he may do anything else, even swell up and become perfectly passive in our hands.

Such are the phenomena commonly observed, and such the impressions which one naturally receives. Certain general conclusions follow irresistibly. First of all the following:

The acts of all the centres involve the use of the same muscles. When a brainless frog's hind-leg wipes the acid,

he calls into play all the leg-muscles which a frog with his full medulla oblongata and cerebellum uses when he turns from his back to his belly. Their contractions are, however, combined differently in the two cases, so that the results vary widely. We must consequently conclude that specific arrangements of cells and fibres exist in the cord for wiping, in the medulla for turning over, etc. Similarly they exist in the thalami for jumping over seen obstacles and for balancing the moved body; in the optic lobes for creeping backwards, or what not. But in the hemispheres, since the presence of these organs brings no new elementary form of movement with it, but only determines differently the occasions on which the movements shall occur, making the usual stimuli less fatal and machine-like, we need suppose no such machinery directly coördinative of muscular contractions to exist. We may rather assume, when the mandate for a wiping-movement is sent forth by the hemispheres, that a current goes straight to the wiping-arrangement in the spinal cord, exciting this arrangement as a whole. Similarly, if an intact frog wishes to jump, all he need do is to excite from the hemispheres the jumping-centre in the thalami or wherever it may be, and the latter will provide for the details of the execution. It is like a general ordering a colonel to make a certain movement, but not telling him how it shall be done.

The same muscle, then, is repeatedly represented at different heights; and at each it enters into a different combination with other muscles to coöperate in some special form of concerted movement. At each height the movement is discharged by some particular form of sensorial stimulus, whilst the stimuli which discharge the hemispheres would seem not so much to be elementary sorts of sensation, as groups of sensations forming determinate objects or things.

The Pigeon's Lower Centres.-The results are just the same if, instead of a frog, we take a pigeon, cut out his hemispheres carefully and wait till he recovers from the

operation. There is not a movement natural to him which this brainless bird cannot execute; he seems, too, after some days to execute movements from some inner irritation, for he moves spontaneously. But his emotions and instincts exist no longer. In Schrader's striking words:

"The hemisphereless animal moves in a world of bodies which . . . are all of equal value for him. . . . He is, to use Goltz's apt expression, impersonal. . . . Every object is for him only a space-occupying mass, he turns out of his path for an ordinary pigeon no otherwise than for a stone. He may try to climb over both. All authors agree that they never found any difference, whether it was an inanimate body, a cat, a dog, or a bird of prey which came in their pigeon's way. The creature knows neither friends nor enemies, in the thickest company it lives like a hermit. The languishing cooing of the male awakens no more impression than the rattling of the peas, or the call-whistle which in the days before the injury used to make the birds hasten to be fed. Quite as little as the earlier observers have I seen hemisphereless she-birds answer the courting of the male. A hemisphereless male will coo all day long and show distinct signs of sexual excitement, but his activity is without any object, it is entirely indifferent to him whether the she-bird be there or not. If one is placed near him, he leaves her unnoticed. . . . As the male pays no attention to the female, so she pays none to her young. The brood may follow the mother ceaselessly calling for food, but they might as well ask it from a stone. .. The hemisphereless pigeon is in the highest degree tame, and fears man as little as cat or bird of prey."

General Notion of Hemispheres.-All these facts lead us, when we try to formulate them broadly, to some such conception as this: The lower centres act from present sensational stimuli alone; the hemispheres act from considerations, the sensations which they may receive serving only as suggesters of these. But what are considerations but expectations, in the fancy, of sensations which will be felt one way or another according as action takes this course or