On the Form of Sea-urchins
As a corollary to the problem of the bird’s egg, we may consider for a moment the forms assumed by the shells of the sea-urchins. These latter are commonly divided into two classes, the Regular and the Irregular Echinids. The regular sea-urchins, save in {662} slight details which do not affect our problem, have a complete radial symmetry. The axis of the animal’s body is vertical, with mouth below and the intestinal outlet above; and around this axis the shell is built as a symmetrical system. It follows that in horizontal section the shell is everywhere circular, and we shall have only to consider its form as seen in vertical section or projection. The irregular urchins (very inaccurately so-called) have the anal extremity of the body removed from its central, dorsal situation; and it follows that they have now a single plane of symmetry, about which the organism, shell and all, is bilaterally symmetrical. We need not concern ourselves in detail with the shapes of their shells, which may be very simply interpreted, by the help of radial co-ordinates, as deformations of the circular or “regular” type.
The sea-urchin shell consists of a membrane, stiffened into rigidity by calcareous deposits, which constitute a beautiful skeleton of separate, neatly fitting “ossicles.” The rigidity of the shell is more apparent than real, for the entire structure is, in a sluggish way, plastic; inasmuch as each little ossicle is capable of growth, and the entire shell grows by increments to each and all of these multitudinous elements, whose individual growth involves a certain amount of freedom to move relatively to one another; in a few cases the ossicles are so little developed that the whole shell appears soft and flexible. The viscera of the animal occupy but a small part of the space within the shell, the cavity being mainly filled by a large quantity of watery fluid, whose density must be very near to that of the external sea-water.
Apart from the fact that the sea-urchin continues to grow, it is plain that we have here the same general conditions as in the egg-shell, and that the form of the sea-urchin is subject to a similar equilibrium of forces. But there is this important difference, that an external muscular pressure (such as the oviduct administers during the consolidation of egg-shell), is now lacking. In its place we have the steady continuous influence of gravity, and there is yet another force which in all probability we require to take into consideration.
While the sea-urchin is alive, an immense number of delicate “tube-feet,” with suckers at their tips, pass through minute pores {663} in the shell, and, like so many long cables, moor the animal to the ground. They constitute a symmetrical system of forces, with one resultant downwards, in the direction of gravity, and another outwards in a radial direction; and if we look upon the shell as originally spherical, both will tend to depress the sphere into a flattened cake. We need not consider the radial component, but may treat the case as that of a spherical shell symmetrically depressed under the influence of gravity. This is precisely the condition which we have to deal with in a drop of liquid lying on a plate; the form of which is determined by its own uniform surface-tension, plus gravity, acting against the uniform internal hydrostatic pressure. Simple as this system is, the full mathematical investigation of the form of a drop is not easy, and we can scarcely hope that the systematic study of the Echinodermata will ever be conducted by methods based on Laplace’s differential equation; but we have no difficulty in seeing that the various forms represented in a series of sea-urchin shells are no other than those which we may easily and perfectly imitate in drops.
In the case of the drop of water (or of any other particular liquid) the specific surface-tension is always constant, and the pressure varies inversely as the radius of curvature; therefore the smaller the drop the more nearly is it able to conserve the spherical form, and the larger the drop the more does it become flattened under gravity. We can represent the phenomenon by using india-rubber balls filled with water, of different sizes; the little ones will remain very nearly spherical, but the larger will fall down “of their own weight,” into the form of more and more flattened cakes; and we see the same thing when we let drops of heavy oil (such as the orthotoluidene spoken of on p. ), fall through a tall column of water, the little ones remaining round, and the big ones getting more and more flattened as they sink. In the case of the sea-urchin, the same series of forms may be assumed to occur, irrespective of size, through variations in T, the specific tension, or “strength,” of the enveloping shell. Accordingly we may study, entirely from this point of view, such a series as the following (Fig. ). In a very few cases, such as the fossil Palaeechinus, we have an approximately spherical {664} shell, that is to say a shell so strong that the influence of gravity becomes negligible as a cause of deformation. The ordinary species of Echinus begin to display a pronounced depression, and this reaches its maximum in such soft-shelled flexible forms as Phormosoma. On the general question I took the opportunity of consulting Mr C. R. Darling, who is an acknowledged expert in drops, and he at once agreed with me that such forms as are represented in Fig. are no other than diagrammatic illustrations
Fig. 328. Diagrammatic vertical outlines of various Sea-urchins: A, Palaeechinus; B, Echinus acutus; C, Cidaris; D, D ′ Coelopleurus; E, E ′ Genicopatagus; F, Phormosoma luculenter; G, P. tenuis; H, Asthenosoma; I, Urechinus.
of various kinds of drops, “most of which can easily be reproduced in outline by the aid of liquids of approximately equal density to water, although some of them are fugitive.” He found a difficulty in the case of the outline which represents Asthenosoma, but the reason for the anomaly is obvious; the flexible shell has flattened down until it has come in contact with the hard skeleton of the jaws, or “Aristotle’s lantern,” within, and the curvature of the outline is accordingly disturbed. The elevated, conical shells such as those of Urechinus and Coelopleurus evidently call for some further explanation; for there is here some cause at work {665} to elevate, rather than to depress the shell. Mr Darling tells me that these forms “are nearly identical in shape with globules I have frequently obtained, in which, on standing, bubbles of gas rose to the summit and pressed the skin upwards, without being able to escape.” The same condition may be at work in the sea-urchin; but a similar tendency would also be manifested by the presence in the upper part of the shell of any accumulation of substance lighter than water, such as is actually present in the masses of fatty, oily eggs.