This point is well illustrated by the tusk of an Indian elephant which I lately found in a London sale-room. This tusk is by some unknown cause, presumably a chronic inflammation, thrown up into thirteen well-marked ridges which closely simulate a series of segments (Fig. 1). Whatever the cause the condition shows how easily a normally unsegmented structure may be converted into a series of repeated parts.
The spread of segmentation through tissues normally unsegmented is very clearly exemplified in the skates' jaws shown in in Fig. 2. The right side of the upper figure shows the normal arrangement in the species Rhinoptera jussieui, but the structure on the left side is very different. The probable relations of the several rows of teeth to the normal rows is indicated by the lettering, but it is evident that by the appearance of new planes of division constituting separate centers of growth, the series has been recast. The pattern of the left side is so definite that had the variation affected the right side also, no systematist would have hesitated to give the specimen a new specific name. The other two drawings show similar variations of a less extensive kind, the nature of which is explained by the lettering of the rows of teeth.
Fig. 2. Jaws of Skates (Rhinoptera) showing meristic variation.
(For a detailed discussion see Materials for the Study of Variation, p. 259.)
This power to divide is a fundamental attribute of life, and of that power cell-division is a special example. In regard to almost all the chief vital phenomena we can say with truth that science has made some progress. If I mention respiration, metabolism, digestion, each of these words calls to mind something more than a bare statement that such acts are performed by an animal or a plant. Each stands for volumes of successful experiment and research, But the expression cell-division, the fundamental act which typifies the rest, and on which they all depend, remains a bare name. We can see with the microscope the outward symptoms of division, but we have no surmise as to the nature of the process by which the division is begun or accomplished. I know nothing which to a man well trained in scientific knowledge and method brings so vivid a realisation of our ignorance of the nature of life as the mystery of cell-division. What is a living thing? The best answer in few words that I know is one which my old teacher, Michael Foster, used to give in his lectures introductory to biology. "A living thing is a vortex of chemical and molecular change." This description gives much, if not all, that is of the essence of life. The living thing is unlike ordinary matter in the fact that, through it, matter is always passing. Matter is essential to it; but, provided that the flow in and out is unimpeded, the life-process can go on so far as we know indefinitely. Yet the living "vortex" differs from all others in the fact that it can divide and throw off other "vortices," through which again matter continually swirls.
We may perhaps take the parallel a stage further. A simple vortex, like a smoke-ring, if projected in a suitable way will twist and form two rings. If each loop as it is formed could grow and then twist again to form more loops, we should have a model representing several of the essential features of living things.
It is this power of spontaneous division which most sharply distinguishes the living from the non-living. In the excellent book dealing with the problems of development, lately published by Mr. Jenkinson a special emphasis is very properly laid on the distinction between the processes of division, and those of differentiation. Too often in discussions of the developmental processes the distinction is obscured. He regards differentiation as the "central difficulty." "Growth and division of the nucleus and the cells," he tells us, are side-issues. This view is quite defensible, but I suspect that the division is the central difficulty, and that if we could get a rationale of what is happening in cell-division we should not be long before we had a clue to the nature of differentiation. It may be self-deception, but I do not feel it impossible to form some hypothesis as to the mode of differentiation, but in no mood of freest speculation are we ever able to form a guess as to the nature of the division. We see differentiations occurring in the course of chemical action, in some phenomena of vibration and so forth: but where do we see anything like the spontaneous division of the living cell? Excite a gold-leaf electroscope, and the leaves separate, but we know that is because they were double before. In electrolysis various substances separate out at the positive and negative poles respectively. Now if in cell-division the two daughter-cells were always dissimilar—that is to say, if differentiation always occurred—we could conceive some rough comparison with such dissociations. But we know the dissimilarity between daughter-cells is not essential. In the reproduction of unicellular organisms and many other cases, the products formed at the two poles are, so far as we can tell, identical. Any assumption to the contrary, if we were disposed to make it, would involve us in difficulties still more serious. At any rate, therefore, if differentiation be really the central difficulty in development, it is division which is the essential problem of heredity.
Sir George Darwin and Professor Jeans tell us that "gravitational instability" consequent on the condensation of gases is "the primary agent at work in the actual evolution of the universe," which has led to the division of the heavenly bodies. The greatest advance I can conceive in biology would be the discovery of the nature of the instability which leads to the continual division of the cell. When I look at a dividing cell I feel as an astronomer might do if he beheld the formation of a double star: that an original act of creation is taking place before me. Enigmatical as the phenomenon seems, I am not without hope that, if it were studied for its own sake, dissociated from the complications which obscure it when regarded as a mere incident in development, some hint as to the nature of division could be found. It is I fear a problem rather for the physicist than for the biologist. The sentiment may not be a popular one to utter before an assembly of biologists, but looking at the truth impersonally I suspect that when at length minds of first rate analytical power are attracted to biological problems, some advance will be made of the kind which we are awaiting.
The study of the phenomena of bodily symmetry offers perhaps the most hopeful point of attack. The essential fact in reproduction is cell-division, and the essential basis of hereditary resemblance is the symmetry of cell-division. The phenomena of twinning provide a convincing demonstration that this is so. By twinning we mean the production of equivalent structures by division. The process is one which may affect the whole body of an animal or plant, or certain of its parts. The term twin as ordinarily used refers to the simultaneous birth of two individuals. Those who are naturalists know that such twins are of two kinds, (1) twins that are not more alike than any other two members of the same family, and (2) twins that are so much alike that even intimate friends mistake them. These latter twins, except in imaginative literature, are always of the same sex.
It is scarcely necessary for me to repeat the evidence from which it has been concluded that without doubt such twins arise by division of the same fertilised ovum. There is a perfect series of gradations connecting them with the various forms of double monsters united by homologous parts. They have been shown several times to be enclosed in the same chorion, and the proofs of experimental embryology show that in several animals by the separation of the two first hemispheres of a dividing egg twins can be produced. Lastly we have recently had the extraordinarily interesting demonstration of Loeb, to which I may specially refer. Herbst some years ago found that in sea water, from which all lime salts had been removed, the segments of the living egg fall apart as they are formed. Using this method Loeb has shown that a temporary immersion in lime-free sea water may result in the production of 90 per cent. of twins. We are therefore safe in regarding the homologous or "identical" twins as resulting from the divisions of one fertilised egg, while the non-identical or "fraternal" twins, as they are called, arise by the fertilisation of two separate ova.[3]