Jelly-Fish, Star-Fish, and Sea-Urchins: Being a Research on Primitive Nervous Systems

by George John Romanes

The only other experiments in nerve-section to which the simple anatomy of a Star-fish exposes itself is that of dividing the nerve-ring in the disc; or, which is virtually the same thing, while leaving this intact, dividing all the nerves where they pass from it into the rays. In specimens mutilated by severing the nerves at the base of each of the five rays, or by dividing the nerve-ring between all the rays, the animal loses all power of co-ordination among its rays. When a common Star-fish is so mutilated it does not crawl in the same determinate manner as an unmutilated animal, but, if it moves at all, it moves slowly and in various directions. When inverted, the power of effecting the righting manœuvre is seen to be gravely impaired, although eventually success is always achieved. There is a marked tendency, as compared with unmutilated specimens, to a promiscuous distribution of spirals and doublings, so that instead of a definite plan of the manœuvre being formed from the first, as is usually the case with unmutilated specimens, such a plan is never formed at all; among the five rays there is a continual change of un-coördinated movements, so that the righting seems to be eventually effected by a mere accidental prepotency of some of the righting movements over others. Appended is a sketch of such un-coördinated movement, taken from a specimen which for more than an hour had been twisting its rays in various directions (Fig. 57). Another sketch is appended to show a form of bending which specimens mutilated as described are very apt to manifest, especially just after the operation. When placed upon their dorsal surface, they turn up all their rays with a peculiar and exactly similar curve in each, which gives to the animal a somewhat tulip-like form (Fig. 58). This form is never assumed by unmutilated specimens, and in mutilated ones, although it may last for a long time, it is never permanent. In detached rays this peculiar curve is also frequently exhibited; but if the nerve of such a ray is divided at any point in its length, the curve is restricted to the distal portion of the ray, and it stops abruptly at the line of nerve-section. When entire Star-fish are mutilated by a section of each nerve-trunk half-way up each ray, and the animal is then placed upon its back, the tetanic contraction of the muscles in the rays before mentioned as occurring under this form of section in detached rays, has the effect, when now occurring in all the rays, of elevating the disc from the floor of the tank. This opisthotonous-like spasm is not, however, permanent; and the distal ends of the rays forming adhesions to the floor of the tank, thy animal eventually rights itself, though much more slowly than unmutilated specimens. After it has righted itself, although it twists about the distal portions of the rays, it does not begin to crawl for a long time, and when it does so, it crawls in a slow and indeterminate manner. Star-fish so mutilated, however, can ascend perpendicular surfaces.

Fig. 57. Un-coördinated movements of a Star-fish, in which the nerves of all the rays have been divided.

Fig. 58. Form frequently assumed by Star-fish under similar circumstances.

The loss of co-ordination between the rays caused by division of the nerve-ring in the disc is rendered most conspicuous in Brittle-stars, from the circumstance that in locomotion and in righting so much here depends upon co-ordinated muscular contraction of the rays. Thus, for instance, when a Brittle-star has its nerve-ring severed between each ray, an interesting series of events follows. First, there is a long period of profound shock—spontaneity, and even irritability, being almost suspended, and the rays appearing to be rigid, as if in tetanic spasm. After a time, feeble spontaneity returns—the animal, however, not moving in any determinate direction. Irritability also returns, but only for the rays immediately irritated, stimulation of one ray causing active writhing movements in that ray, but not affecting, or only feebly affecting, the other rays. The animal, therefore, is quite unable to escape from the source of irritation, the aimless movements of the rays now forming a very marked contrast to the instantaneous and vigorous leaping movements of escape which are manifested by unmutilated specimens. Moreover, unmutilated specimens will vigorously leap away, not only from stimulation of the rays, but also from that of the disc; but those with their nerve-ring cut make no attempts to escape, even from the most violent stimulation of the disc. In other words, the disc is entirely severed from all physiological connection with the rays.

If the nerve-ring be divided at two points, one on either side of a ray, that ray becomes physiologically separated from the rest of the organism. If the two nerve-divisions are so placed as to include two adjacent rays—i.e. if one cut is on one side of a ray and the other on the further side of an adjacent ray—then these two rays remain in physiological continuity with one another, although they suffer physiological separation from the other three. When a Brittle-star is completely divided into two portions, one portion having two arms and the other three, both portions begin actively to turn over on their backs, again upon their faces, again upon their backs, and so on alternately for an indefinite number of times. These movements arise from the rays, under the influence of stimulation caused by the section, seeking to perform their natural movements of leaping, which however end, on account of the weight of the other rays being absent, in turning themselves over. An entire Brittle-star when placed on its back after division of its nerve-ring is not able to right itself, owing to the destruction of co-ordination among its rays. Astropecten, under similar circumstances, at first bends its rays about in various ways, with a preponderant disposition to the tulip form, and keeps its ambulacral feet in active movement. But after half an hour, or an hour, the feet generally become retracted and the rays nearly motionless—the animal, like a Brittle-star, remaining permanently on its back. In this, as in other species, the effect of dividing the nerve-ring on either side of a ray is that of destroying its physiological connection with the rest of the animal, the feet in that ray, although still remaining feebly active, no longer taking part in any co-ordinated movement—that ray, therefore, being merely dragged along by the others.

Under this division it only remains further to be said, that section of the nerve-ring in the disc, or the nerve-trunks of the rays, although, as we have seen, so completely destroying physiological continuity in the rows of ambulacral feet and muscular system of the animal, does not destroy physiological continuity in the external nerve-plexus; for however much the nerve-ring and nerve-trunks may be injured, stimulation of the dorsal surface of the animal throws all the ambulacral feet and all the muscular system of the rays into active movement. This fact proves that the ambulacral feet and the muscles are all held in nervous connection with one another by the external plexus, without reference to the integrity of the main nerve-trunks.

2. Echini.—Section of external surface of shell.—If a cork-borer be applied to the external surface of the shell of an Echinus, and rotated there till the calcareous substance of the shell is reached, and therefore a continuous circular section of the over-lying tissues effected, it is invariably found that the spines and pedicellariæ within the circular area are physiologically separated from the contiguous spines and pedicellariæ, as regards local reflex excitability. That is to say, if any part of this circular area be stimulated, all the spines and pedicellariæ within that area immediately respond to the stimulation in the ordinary way; while none of the spines or pedicellariæ surrounding the area are affected. Similarly, if any part of the shell external to the circumscribed area be stimulated, the spines and pedicellariæ within that area are not affected. These facts prove that the function which is manifested by these appendages of localizing and gathering round a seat of stimulation, is exclusively dependent upon the external nerve-plexus. It is needless to add that in this experiment it does not signify of what size or shape or by what means the physiological island is made, so long as the destruction of the nervous plexus by a closed curve of injury is rendered complete. In order to ascertain whether, in the case of an unclosed curve of injury, any irradiation of a stimulus would take place round the ends of the curve, we made sundry kinds of section. It is, however, needless to describe these, for they all showed that, after injury of a part of the plexus, there is no irradiation of the stimulus round the ends of the injury. Thus, for instance, if a short straight line of injury be made, by drawing the point of a scalpel over the shell, say along the equator of the animal, and if a stimulus be afterwards applied on either side of that line, even quite close to one of its ends, no effect will be exerted on the spines or pedicellariæ on the other side of the line. This complete inability of a stimulus to escape round the ends of an injury, forms a marked contrast to the almost unlimited degree in which such escape takes place in the more primitive nervous plexus of the Medusæ.

Although the nervous connections on which the spines and pedicellariæ depend for their function of localizing and closing round a seat of stimulation are thus shown to be completely destroyed by injury of the external plexus, other nervous connections, upon which another function of the spines depends, are not in the smallest degree impaired by such injury. The other function to which I allude is that which brings about the general co-ordinated action of all the spines for the purposes of locomotion. That this function is not impaired by injury of the external plexus is proved by the fact that if the area within a closed line of injury on the surface of the shell be strongly irritated, all the spines over the whole surface begin to manifest their peculiar bristling movements, and by this co-ordinated action rapidly move the animal in a straight line of escape from the source of irritation; the injury to the external plexus, although completely separating the spines enclosed by it from their neighbouring spines as regards what may be called their local function of seizing the instrument of stimulation, nevertheless leaves them in undisturbed connection with all the other spines in the organism as regards what may be called their universal function of locomotion.