GENESIS, HEREDITY, AND VARIATION § 97a Most if not all of them proceed on the assumption, shown in §66 to be needful, that the structural characters of organisms are determined by the special natures of units which are intermediate between the chemical units and the morphological units—between the invisible molecules of proteid-substances and the visible tissue-components called cells. Four years after the first edition of this volume was published, appeared Mr. Darwin's work, The Variation of Animals and Plants under Domestication; and in this he set forth his doctrine of Pangenesis. Referring to the doctrine of physiological units which the preceding chapters work out, he at first expressed a doubt whether his own was or was not the same, but finally concluded that it was different. He was right in so concluding. Throughout my argument the implication everywhere is that the physiological units are all of one kind; whereas Mr. Darwin regards his component units, or "gemmules," as being of innumerable unlike kinds. He supposes that every cell of every tissue gives off gemmules Much more recently have been enunciated the hypotheses of Prof. Weismann, differing from the foregoing hypotheses in two respects. In the first place it is assumed that the fragment of matter out of which each organism arises consists of two portions—one of them, the germ-plasm, reserved within the generative organ of the incipient individual, representing in its components the structure of the species, and gives origin to the germs of future individuals; and the other of them, similarly representative of the specific structure, giving origin to the rest of the body, or soma, but contains in its components none of those latent powers possessed by those of the germ-plasm. In the second place the germ-plasm, in common with the soma-plasm, consists of multitudinous kinds of units portioned out to originate the various organs. Of these there are groups, sub-groups, and sub-sub-groups. The largest of them, called "idants," are supposed each to contain a number of "ids"; within each id there are numerous "determinants"; and each determinant is made up of many "biophors"—the smallest elements possessing vitality. Passing over details, the essential assumption is that there exists a separate determinant for each part of the organism capable of independent variation; and Prof. Weismann infers that while there may be but one for the blood and but one for a considerable area of skin (as a stripe of the zebra) there must be a determinant for each scale on a butterfly's wing: the number on the four wings being over two Here it is needless to specify the modifications of these hypotheses espoused by various biologists—all of them assuming that the structural traits of each species are expressed in certain units intermediate between morphological units and chemical units. § 97b Already in §77 I have illustrated the power which some plants possess of developing new individuals from mere fragments of leaves and even from detached scales. Striking as are the facts there instanced, they are scarcely more significant than some which are familiar. The formation of cauline buds, presently growing into shoots, shows us a kind of inheritance which a true theory must explain. As described by Kerner, such buds arise in Pimpernel, Toad-flax, etc., below the seed-leaves, even while yet there are no axils in which buds usually grow; and in many plants they arise from intermediate places on the stem: that is, without definite relations to pre-existing structures. How fortuitous is their origin is shown when a branch is induced to bud by keeping it wrapped round with a wet cloth. Even still better proved is the absence of any relation between cauline buds and normal germs by the frequent growth of them out of "callus"—the tissue which spreads over wounds and the cut ends of branches. It is not easy to reconcile these facts Still less does the hypothesis of Prof. Weismann harmonize with the evidence as plants display it. Plant-embryogeny yields no sign of separation between germ-plasm and soma-plasm; and, indeed, the absence of such separation is admitted. After instancing cases among certain of the lower animals, in which no differentiation of the two arises in the first generation resulting from a fertilized ovum, Prof. Weismann continues:— "The same is true as regards the higher plants, in which the first shoot arising from the seed never contains germ-cells, or even cells which subsequently become differentiated into germ cells. In all these last-mentioned cases the germ-cells are not present in the first person arising by embryogeny as special cells, but are only formed in much later cell-generations from the offspring of certain cells of which this first person was composed." (Germ-Plasm, p. 185.) How this admission consists with the general theory it is difficult to understand. The units of the soma-plasm are here recognized as having the same generative powers as the units of the germ-plasm. In so far as one organic kingdom and a considerable part of the other are concerned the doctrine is relinquished. Relinquishment is, indeed, necessitated even by the ordinary facts, and still more by the The hypothesis of physiological units, then, remains outstanding. For cauline buds imply that throughout the plant-tissue, where not unduly differentiated, the local physiological units have a power of arranging themselves into the structure of the species. But this hypothesis, too, as it now stands, is inadequate. Under the form thus far given to it, it fails to explain some accompanying facts. For if the branch just instanced as producing a cauline bud be cut off and its end stuck in the ground, or if it be bent down and a portion of it covered with earth, there will grow from it rootlets and presently roots. The same portion of tissue which otherwise would have produced a shoot with all its appendages, constituting an individual, now produces only a special part of an individual. § 97c The often-cited reproduction of a crab's lost claw or a lizard's tail, Mr. Darwin thought explicable by his hypothesis of diffused gemmules, representing all organs or their component cells. But though, after simple amputation, regrowth of the proximate part of the tail is conceivable as hence resulting, it is not easy to understand how the remoter part, the components of which are now absent from the organism, can arise afresh from gemmules no longer originated in due proportion. Prof. Weismann's hypothesis, again, implies that there must exist at the place of separation, a ready-provided supply of determinants, previously latent, able to reproduce the missing tail in all its details—nay, even to do this several times over: a strong supposition! Various annulose animals display anomalies of development difficult to explain on any hypothesis. We have creatures like the fresh-water Nais which, though it has advanced structures, including a vascular system, branchiÆ, and a nervous cord ending with cephalic ganglia, nevertheless shows us an ability like that of the Hydra to reproduce the whole from a small part: nearly forty pieces into which a Nais was cut having severally grown into complete animals. Again we have, in the order PolychÆtÆ, types like Myrianida, in which by longitudinal budding a string of individuals, sometimes numbering even thirty, severally develop certain segments into heads, while increasing their segments in number. In yet other types there occurs not longitudinal gemmation only, but lateral gemmation: a segment will send out sideways a bud which presently becomes a complete worm. Once more, Syllis ramosa is a species in which the individual worms growing from lateral buds, while remaining attached to the parent, themselves give origin to buds; and so produce a branched aggregate of worms. How shall we explain the reparative and reproductive powers thus exemplified? It seems undeniable that each portion has an ability to produce, according to circumstances, the whole creature or a missing part of the creature. When we read of Sir J. Dalyell that he "cut a Dasychone into three pieces; the Still less explicable in any way thus far proposed are certain remedial actions seen in animals. An example of them was furnished in §67, where "false joints" were described—joints formed at places where the ends of a broken bone, failing to unite, remain moveable one upon the other. According to the character of the habitual motions there results a rudely formed hinge-joint or a ball-and-socket joint, either having the various constituent parts—periosteum, fibrous tissue, capsule, ligaments. Now Mr. Darwin's hypothesis, contemplating only normal structures, fails to account for this formation of an abnormal structure. Neither can we ascribe this local development to determinants: there were no appropriate ones in the germ-plasm, since no such structure was provided for. Nor does the hypothesis of physiological units, as presented in preceding chapters, yield an interpretation. These could have no other tendency than to restore the normal form of the limb, and might be expected to oppose the genesis of these new parts. Thus we have to seek, if not another hypothesis, then some such qualification of an existing hypothesis as will harmonize it with various exceptional phenomena. § 97d My purpose at that time being the elucidation of sociological truths, it did not concern me to carry further the biological half of this comparison. Otherwise there might have been named the case in which a supernumerary finger, beginning to bud out, completes itself as a local organ with bones, muscles, skin, nail, etc., in defiance of central control: even repeating itself when cut off. There might also have Now let us contemplate the essential fact in the analogy. The men in an Australian mining-camp, as M. Pierre Leroy Beaulieu points out, fall into Anglo-Saxon usages different from those which would characterize a French mining-camp. Emigrants to a far West settlement in America quickly establish post-office, bank, hotel, newspaper, and other urban institutions. We are thus shown that along with certain traits leading to a general type of social organization, there go traits which independently produce fit local organizations. Individuals are led into occupations and official posts, often quite new to them, by the wants of those around—are now influenced and now coerced into social arrangements which, as shown perhaps by gambling saloons, by shootings at sight, and by lynchings, are scarcely at all affected by the central government. Now the physiological units in each species appear to have a similar combination of capacities. Besides their general proclivity towards the specific organization, they show us abilities to organize themselves locally; and these abilities are in some cases displayed in defiance of the general control, as in the supernumerary finger or the false joint. Apparently each physiological unit, while having in a manner the whole organism as the structure which, along with the rest, it tends to form, has also an aptitude to take part in forming any local structure, and to assume its place in that structure under the influence of adjacent physiological units. A familiar fact supports this conclusion. Everyone has at hand, not figuratively but literally, an illustration. Let him compare the veins on the backs of his two hands, either with one another or with the veins on another person's hands, and he will see that the branchings and inosculations do not § 97e The independent vitality shown by separated portions of ventral skin from frog-larvÆ may be named as the first illustration. With their attached yolk-cells these lived for days, and underwent such transformations as proved some structural proclivity, though of course the product was amorphous. Detached portions of tails of larvÆ went on developing their component parts in much the same ways as they would have done if remaining attached. More striking still was the evidence furnished by experiments in grafting. These proved that the undifferentiated rudiment of an organ will, when cut off and joined to a non-homologous place in another individual, develop itself as it would have done if left in its original place. In brief, then, we may say that each part is in chief measure autogenous. These strange facts presented by small aggregates of organic matter, which are the seats of extremely complex forces, will seem less incomprehensible if we observe what has taken Having thus seen that in a way we have not hitherto recognized the same general principles pervade inorganic and organic evolution, let us revert to the case of super-organic evolution from which a parallel was drawn above. As analogous to the germinal mass of units out of which a new organism is to evolve, let us take an assemblage of colonists not yet socially organized but placed in a fertile region—men derived from a society (or rather a succession of societies) of long-established type, who have in their adapted natures the proclivity towards that type. In passing from its wholly unorganized state to an organized state, what will be the first step? Clearly this assemblage, though it may have § 97f Doubtless this is true. Still we have a consensus of proofs that the component units of organisms have constitutions of extremely involved kinds. Contemplate the facts and their implications. (1) Here is some large division of the animal kingdom—say the Vertebrata. The component units of all its members have certain fundamental traits in common: all But unimaginable as all this is, the facts may nevertheless in some way answer to it. As before remarked, progressing science reveals complexity within complexity—tissues made up of cells, cells containing nuclei and cytoplasm, cytoplasm Whatever be their supposed natures we are compelled to ascribe extreme complexity to these unknown somethings which have the power of organizing themselves into a structure of this or that species. If gemmules be alleged, then the ability of every organ and part of an organ to vary, implies that the gemmules it gives off are severally capable of receiving minute modifications of their ordinary structures: they must have many parts admitting of innumerable relations. Supposing determinants be assumed, then in addition to the complexity which each must have to express in itself the structure of the part evolved from it, it must have the further complexity implied by every superposed modification which causes a variation of that part. And, as One more assumption seems necessary if we are to imagine how changes of structure caused by changes of function can be transmitted. Reverting to §54d, where an unceasing circulation of protoplasm throughout an organism was inferred, we must conceive that the complex forces of which each constitutional unit is the centre, and by which it acts on other units while it is acted on by them, tend continually to remould each unit into congruity with the structures around: superposing on it modifications answering to the modifications which have arisen in those structures. Whence is to be drawn the corollary that in course of time all the circulating units,—physiological, or constitutional if we prefer so to call them—visiting all parts of the organism, are severally made bearers of traits expressing local modifications; and that those units which are eventually gathered into sperm-cells and germ-cells also bear these superposed traits. If against all this it be urged that such a combination of structures and forces and processes is inconceivably involved, then the reply is that so astonishing a transformation as that which an unfolding organism displays cannot possibly be effected by simple agencies. § 97g What are we to say of the familiar fact that certain small organs which, with the approach to maturity, become active, entail changes of structure in remote parts—that after the testes have undergone certain final developments, the hairs on the chin grow and the voice deepens? It has been contended that certain concomitant modifications in the fluids throughout the body may produce correlated sexual traits; The inadequacy of our explanations we at once see in presence of a structure like a peacock's tail-feather. Mr. Darwin's hypothesis is that all parts of every organ are continually giving off gemmules, which are consequently everywhere present in their due proportions. But a completed feather is an inanimate product and, once formed, can add to the circulating fluids no gemmules representing all its parts. If we follow Prof. Weismann we are led into an astounding supposition. He admits that every variable part must have a special determinant, and that this results in the assumption of over two hundred thousand for the four wings of a butterfly. Let us ask what must happen in the case of a peacock's feather. On looking at the eye near its end, we see that the minute processes on the edge of each lateral thread must have been in some way exactly adjusted, in colour and position, so as to fall into line with the processes on adjacent threads: otherwise the symmetrical arrangement of coloured rings would be impossible. Each of these processes, then, being an independent variable, must have had its particular determinant. Now there are about 300 threads on the shaft of a large feather, and each of them bears on the average 1,600 processes, making for the whole feather 480,000 of these processes. For one feather alone there must have been 480,000 determinants, and for the whole tail many millions. And these, along with the determinants for the detailed parts of all the other feathers, and for the variable components of all organs forming the body at large, must have been contained At last then we are obliged to admit that the actual organizing process transcends conception. It is not enough to say that we cannot know it; we must say that we cannot even conceive it. And this is just the conclusion which might have been drawn before contemplating the facts. For if, as we saw in the chapter on "The Dynamic Element in Life," it is impossible for us to understand the nature of this element—if even the ordinary manifestations of it which a living body yields from moment to moment are at bottom incomprehensible; then, still more incomprehensible must be that astonishing manifestation of it which we have in the initiation and unfolding of a new organism. Thus all we can do is to find some way of symbolizing the process so as to enable us most conveniently to generalize its phenomena; and the only reason for adopting the hypothesis of physiological units or constitutional units is that it best serves this purpose. |