Darwin and Modern Science

by A. C. Seward

But a deeper reason than this has been assigned for the importance of embryology in classification. It has been asserted, and is implied by Darwin in the passage quoted, that the ancestral history is repeated in a condensed form in the embryonic, and that a study of the latter enables us to form a picture of the stages of structure through which the organism has passed in its evolution. It enables us on this view to reconstruct the pedigrees of animals and so to form a genealogical tree which shall be the true expression of their natural relations.

The real question which we have to consider is to what extent the embryological studies of the last 50 years have confirmed or rendered probable this "theory of recapitulation." In the first place it must be noted that the recapitulation theory is itself a deduction from the theory of evolution. The facts of embryology, particularly of vertebrate embryology, and of larval history receive, it is argued, an explanation on the view that the successive stages of development are, on the whole, records of adult stages of structure which the species has passed through in its evolution. Whether this statement will bear a critical verbal examination I will not now pause to inquire, for it is more important to determine whether any independent facts can be alleged in favour of the theory. If it could be shown, as was stated to be the case by L. Agassiz, that ancient and extinct forms of life present features of structure now only found in embryos, we should have a body of facts of the greatest importance in the present discussion. But as Huxley (See Huxley's "Scientific Memoirs", London, 1898, Vol. I. page 303: "There is no real parallel between the successive forms assumed in the development of the life of the individual at present, and those which have appeared at different epochs in the past." See also his Address to the Geological Society of London (1862) 'On the Palaeontological Evidence of Evolution', ibid. Vol. II. page 512.) has shown and as the whole course of palaeontological and embryological investigation has demonstrated, no such statement can be made. The extinct forms of life are very similar to those now existing and there is nothing specially embryonic about them. So that the facts, as we know them, lend no support to theory.

But there is another class of facts which have been alleged in favour of the theory, viz. the facts which have been included in the generalisation known as the Law of v. Baer. The law asserts that embryos of different species of animals of the same group are more alike than the adults and that, the younger the embryo, the greater are the resemblances. If this law could be established it would undoubtedly be a strong argument in favour of the "recapitulation" explanation of the facts of embryology. But its truth has been seriously disputed. If it were true we should expect to find that the embryos of closely similar species would be indistinguishable from one another, but this is notoriously not the case. It is more difficult to meet the assertion when it is made in the form given above, for here we are dealing with matters of opinion. For instance, no one would deny that the embryo of a dogfish is different from the embryo of a rabbit, but there is room for difference of opinion when it is asserted that the difference is less than the difference between an adult dogfish and an adult rabbit. It would be perfectly true to say that the differences between the embryos concern other organs more than do the differences between the adults, but who is prepared to affirm that the presence of a cephalic coelom and of cranial segments, of external gills, of six gill slits, of the kidney tubes opening into the muscle-plate coelom, of an enormous yolk-sac, of a neurenteric canal, and the absence of any trace of an amnion, of an allantois and of a primitive streak are not morphological facts of as high an import as those implied by the differences between the adults? The generalisation undoubtedly had its origin in the fact that there is what may be called a family resemblance between embryos and larvae, but this resemblance, which is by no means exact, is largely superficial and does not extend to anatomical detail.

It is useless to say, as Weismann has stated ("The Evolution Theory", by A. Weismann, English Translation, Vol. II. page 176, London, 1904.), that "it cannot be disputed that the rudiments [vestiges his translator means] of gill-arches and gill-clefts, which are peculiar to one stage of human ontogeny, give us every ground for concluding that we possessed fish-like ancestors." The question at issue is: did the pharyngeal arches and clefts of mammalian embryos ever discharge a branchial function in an adult ancestor of the mammalia? We cannot therefore, without begging the question at issue in the grossest manner, apply to them the terms "gill-arches" and "gill-clefts". That they are homologous with the "gill-arches" and "gill-clefts" of fishes is true; but there is no evidence to show that they ever discharged a branchial function. Until such evidence is forthcoming, it is beside the point to say that it "cannot be disputed" that they are evidence of a piscine ancestry.

It must, therefore, be admitted that one outcome of the progress of embryological and palaeontological research for the last 50 years is negative. The recapitulation theory originated as a deduction from the evolution theory and as a deduction it still remains.

Let us before leaving the subject apply another test. If the evolution theory and the recapitulation theory are both true, how is it that living birds are not only without teeth but have no rudiments of teeth at any stage of their existence? How is it that the missing digits in birds and mammals, the missing or reduced limb of snakes and whales, the reduced mandibulo-hyoid cleft of elasmobranch fishes are not present or relatively more highly developed in the embryo than in the adult? How is it that when a marked variation, such as an extra digit, or a reduced limb, or an extra segment, makes its appearance, it is not confined to the adult but can be seen all through the development? All the clear evidence we can get tends to show that marked variations, whether of reduction or increase, of organs are manifest during the whole of the development of the organ and do not merely affect the adult. And on reflection we see that it could hardly be otherwise. All such evidence is distinctly at variance with the theory of recapitulation, at least as applied to embryos. In the case of larvae of course the case will be different, for in them the organs are functional, and reduction in the adult will not be accompanied by reduction in the larva unless a change in the conditions of life of the larva enables it to occur.

If after 50 years of research and close examination of the facts of embryology the recapitulation theory is still without satisfactory proof, it seems desirable to take a wider sweep and to inquire whether the facts of embryology cannot be included in a larger category.

As has been pointed out by Huxley, development and life are co-extensive, and it is impossible to point to any period in the life of an organism when the developmental changes cease. It is true that these changes take place more rapidly at the commencement of life, but they are never wholly absent, and those which occur in the later or so-called adult stages of life do not differ in their essence, however much they may differ in their degree, from those which occur during the embryonic and larval periods. This consideration at once brings the changes of the embryonic period into the same category as those of the adult and suggests that an explanation which will account for the one will account for the other. What then is the problem we are dealing with? Surely it is this: Why does an organism as soon as it is established at the fertilisation of the ovum enter upon a cycle of transformations which never cease until death puts an end to them? In other words what is the meaning of that cycle of changes which all organisms present in a greater or less degree and which constitute the very essence of life? It is impossible to give an answer to this question so long as we remain within the precincts of Biology—and it is not my present purpose to penetrate beyond those precincts into the realms of philosophy. We have to do with an ultimate biological fact, with a fundamental property of living matter, which governs and includes all its other properties. How may this property be stated? Thus: it is a property of living matter to react in a remarkable way to external forces without undergoing destruction. The life-cycle, of which the embryonic and larval periods are a part, consists of the orderly interaction between the organism and its environment. The action of the environment produces certain morphological changes in the organism. These changes enable the organism to come into relation with new external forces, to move into what is practically a new environment, which in its turn produces further structural changes in the organism. These in their turn enable, indeed necessitate, the organism to move again into a new environment, and so the process continues until the structural changes are of such a nature that the organism is unable to adapt itself to the environment in which it finds itself. The essential condition of success in this process is that the organism should always shift into the environment to which its new structure is suited—any failure in this leading to the impairment of the organism. In most cases the shifting of the environment is a very gradual process (whether consisting in the very slight and gradual alteration in the relation of the embryo as a whole to the egg-shell or uterine wall, or in the relations of its parts to each other, or in the successive phases of adult life), and the morphological changes in connection with each step of it are but slight. But in some cases jumps are made such as we find in the phenomena known as hatching, birth, and metamorphosis.

This property of reacting to the environment without undergoing destruction is, as has been stated, a fundamental property of organisms. It is impossible to conceive of any matter, to which the term living could be applied, being without it. And with this property of reacting to the environment goes the further property of undergoing a change which alters the relation of the organism to the old environment and places it in a new environment. If this reasoning is correct, it necessarily follows that this property must have been possessed by living matter at its first appearance on the earth. In other words living matter must always have presented a life-cycle, and the question arises what kind of modification has that cycle undergone? Has it increased or diminished in duration and complexity since organisms first appeared on the earth? The current view is that the cycle was at first very short and that it has increased in length by the evolutionary creation of new adult phases, that these new phases are in addition to those already existing and that each of them as it appears takes over from the preceding adult phase the functional condition of the reproductive organs. According to the same view the old adult phases are not obliterated but persist in a more or less modified form as larval stages. It is further supposed that as the life-history lengthens at one end by the addition of new adult phases, it is shortened at the other by the abbreviation of embryonic development and by the absorption of some of the early larval stages into the embryonic period; but on the whole the lengthening process has exceeded that of shortening, so that the whole life-history has, with the progress of evolution, become longer and more complicated.

Now there can be no doubt that the life-history of organisms has been shortened in the way above suggested, for cases are known in which this can practically be seen to occur at the present day. But the process of lengthening by the creation of new stages at the other end of the life-cycle is more difficult to conceive and moreover there is no evidence for its having occurred. This, indeed, may have occurred, as is suggested below, but the evidence we have seems to indicate that evolutionary modification has proceeded by ALTERING and not by SUPERSEDING: that is to say that each stage in the life-history, as we see it to-day, has proceeded from a corresponding stage in a former era by the modification of that stage and not by the creation of a new one. Let me, at the risk of repetition, explain my meaning more fully by taking a concrete illustration. The mandibulo-hyoid cleft (spiracle) of the elasmobranch fishes, the lateral digits of the pig's foot, the hind-limbs of whales, the enlarged digit of the ostrich's foot are supposed to be organs which have been recently modified. This modification is not confined to the final adult stage of the life-history but characterises them throughout the whole of their development. A stage with a reduced spiracle does not proceed in development from a preceding stage in which the spiracle shows no reduction: it is reduced at its first appearance. The same statement may be made of organs which have entirely disappeared in the adult, such as bird's teeth and snake's fore-limbs: the adult stage in which they have disappeared is not preceded by embryonic stages in which the teeth and limbs or rudiments of them are present. In fact the evidence indicates that adult variations of any part are accompanied by precedent variations in the same direction in the embryo. The evidence seems to show, not that a stage is added on at the end of the life-history, but only that some of the stages in the life-history are modified. Indeed, on the wider view of development taken in this essay, a view which makes it coincident with life, one would not expect often to find, even if new stages are added in the course of evolution, that they are added at the end of the series when the organism has passed through its reproductive period. It is possible of course that new stages have been intercalated in the course of the life-history, though it is difficult to see how this has occurred. It is much more likely, if we may judge from available evidence, that every stage has had its counterpart in the ancestral form from which it has been derived by descent with modification. Just as the adult phase of the living form differs, owing to evolutionary modification, from the adult phase of the ancestor from which it has proceeded, so each larval phase will differ for the same reason from the corresponding larval phase in the life-history of the ancestor. Inasmuch as the organism is variable at every stage of its independent existence and is exposed to the action of natural selection there is no reason why it should escape modification at any stage.