Form and Function Part 24

It was not that men's minds were not ripe for evolution, for in the early decades of the 19th century evolution was in the air. There were few of von Baer's contemporaries who had not read Lamarck;[346] Erasmus Darwin's _Zoonomia_ ran through three editions, and was translated into German, French and Italian;[247] German philosophy was full of the idea of evolution.

There was no unreadiness to accept the derivation of present-day species from a primordial form--if only some solid evidence for such derivation were forthcoming. Cuvier and von Baer, as we have seen, combated the current evolution theories on the ground that the evidence was insufficient, but von Baer at least had no rooted objection to evolution. In an essay of 1834, ent.i.tled _The Most General Law of Nature in all Development_,[348] von Baer expressed belief in a limited amount of evolution. In this paper he did not admit that all animals have developed from one parent form, and he refused to believe that man has descended from an ape; but, basing his supposition upon the facts of variability and upon the evidence of palaeontology, he went so far as to maintain that many species have evolved from parent stocks. In the absence of conclusive proofs he did not commit himself to a belief in any extended or comprehensive process of evolution.

Imbued as he was with the idea of development von Baer saw in evolution a process essentially of the same nature as the development of the individual. Evolution, like development, was due to a _Bildungskraft_ or formative force. The ultimate law of all becoming was that "the history of Nature is nothing but the history of the ever-advancing victory of spirit over matter" (p. 71). In a later essay (1835) in the same volume he says that all natural science is nothing but a long commentary on the single phrase _Es werde!_. (p. 86).

As we shall see, von Baer adopted in later years the same att.i.tude to Darwinism as he did to the evolution theories in vogue in his youth.

Although in the twenty or thirty years before the publication of the _Origin of Species_ (1859) no evolution theory of any importance was published, and although the great majority of biologists believed in the constancy of species, there were not wanting some who, like von Baer, had an open mind on the subject, or even believed in the occurrence of evolutionary processes of small scope. Isidore Geoffroy St Hilaire, the son of the great Etienne Geoffroy St Hilaire, seems to have held that species might be formed from varieties. The law which L. Aga.s.siz thought he could establish,[349] of the parallelism between palaeontological succession, systematic rank, and embryological development, tended to help the progress of evolutionary ideas. J. V. Carus, who afterwards became a supporter of Darwin, seems already, in 1853, to have inferred from Aga.s.siz's law the probability of evolution.[350]

But no evolution theory was taken very seriously before 1859, when the _Origin of Species_ was published.

Like Lamarck, Charles Darwin was, neither by inclination nor by training, a morphologist. In his youth he was a collector, a sportsman and a field geologist. His voyage round the world on the _Beagle_ aroused in him keen interest in the problem of species--their variety, their variation according to place and time, their adaptedness to environment. The conviction gradually took possession of his mind that the puzzling facts of geographical range and geological succession which he observed wherever he went were explicable only on the hypothesis that species change. He was not satisfied with the theories of evolution that had been proposed by his grandfather, by Lamarck, and by E. Geoffroy St Hilaire--he did not indeed understand these theories any too well. He resolved to work out the problem in his own way, for his own satisfaction. He tells us all this very clearly in his autobiography.

"During the voyage of the _Beagle_ I had been deeply impressed by discovering in the Pampean formation great fossil animals covered with armour like that on the existing armadillos; secondly, by the manner in which closely allied animals replace one another in proceeding southwards over the continent; and thirdly, by the South American character of most of the productions of the Galapagos archipelago, and more especially by the manner in which they differ slightly on each island of the group; some of the islands appearing to be very ancient in a geological sense.

"It was evident that such facts as these, as well as many others, could only be explained on the supposition that species gradually become modified; and the subject haunted me. But it was equally evident that neither the action of the surrounding conditions, nor the will of the organisms (especially in the case of plants) could account for the innumerable cases in which organisms of every kind are beautifully adapted to their habits of life--for instance, a woodp.e.c.k.e.r or a tree-frog to climb trees, or a seed for dispersal by hooks or plumes. I had always been much struck by such adaptations, and until these could be explained it seemed to me almost useless to endeavour to prove by indirect evidence that species have been modified."[351]

All Darwin's varied subsequent work revolved round these, for him, essential problems--How do species change, and how do they become adapted to their environment? He never ceased to be essentially a field naturalist, and his theory of natural selection would have been an empty and abstract thing if his vast knowledge and understanding of the "web of life" had not given it colour and form. He never lost touch with the living thing in its living, breathing reality--even plants he rightly regarded as active things, full of tricks and contrivances for making their way in the world. No one ever realised more vividly than he the delicacy and complexity of the adaptations to environment which are the necessary condition of success in the struggle for existence. Almost his greatest service to biology was that he made biologists realise as they never did before the vast importance of environment. He took biology into the open air, away from the museum and the dissecting-room.

Naturally this att.i.tude was not without its drawbacks. It led him to take only a lukewarm interest in the problems of morphology. It is true he used the facts of morphology with great effect as powerful arguments for evolution, but it was not from such facts that he deduced his theory to account for evolution. It is questionable indeed whether the theory of natural selection is properly applicable to the problems of form. It was invented to account for the evolution of specific differences and of ecological adaptations; it was not primarily intended as an explanation of the more wonderful and more mysterious facts of the _convenance des parties_ and the interaction of structure and function. Perhaps Darwin did not realise this inner aspect of adaptation quite so vividly as he did the more superficial adaptation of organisms to their environment.

It was, perhaps, his lack of morphological training and experience that led him to disregard the problems of form, or at least to realise very insufficiently their difficulty.

It is in any case very significant that only a small part of his _Origin of Species_ is devoted to the discussion of morphological questions--only one chapter out of the fourteen contained in the first edition.

Though the theory of natural selection took little account of the problems of form, Darwin's masterly vindication of the theory of evolution was of immense service to morphology, and Darwin himself was the first to point out what a great light evolution threw upon all morphological problems. In a few pages of the _Origin_ he laid the foundations of evolutionary morphology.

We have here to consider his interpretation of morphological facts and its relation to the current morphology of his time.

The sketch of his theory, written in 1842,[352] shows a very significant division into two parts--the first dealing with the positive facts of variability and the theory of natural selection, the second with the general evidence for evolution. It is in the second part that the paragraphs on morphological matters occur. In paragraph 7, on affinities and cla.s.sification, Darwin points out that on the theory of evolution h.o.m.ological relationship would be real relationship, and the natural system would really be genealogical. In the next paragraph he notes that evolution would account for the unity of type in the great cla.s.ses, for the metamorphosis of organs, and for the close resemblance which early embryos show to one another. It is of special interest to note that he definitely rejects the Meckel-Serres theory of recapitulation. "It is not true," he writes, "that one pa.s.ses through the form of a lower group, though no doubt fish more nearly related to foetal state" (p.

42). The greater divergence which adults show seems to him to be due to the fact that selection acts more on the later than on the embryonic stages. He realises very clearly how illuminative the theory of evolution is when applied to the puzzling facts of embryonic development. "The less differences of foetus--this has obvious meaning on this view: otherwise how strange that a horse, a man, a bat should at one time of life have arteries, running in a manner which is only intelligibly useful in a fish! The natural system being on theory genealogical, we can at once see why foetus, retaining traces of the ancestral form, is of the highest value in cla.s.sification" (p. 45).

Abortive organs, too, gain significance on the evolutionary hypothesis.

"The affinity of different groups, the unity of types of structure, the representative forms through which foetus pa.s.ses, the metamorphosis of organs, the abortion of others, cease to be metaphorical expressions and become intelligible facts" (p. 50).

In general, organisms can be understood only if we take into account the cardinal fact that they are historical beings. "We must look at every complicated mechanism and instinct as the summary of a long history of useful contrivances much like a work of art" (p. 51).[353]

Already in 1842 Darwin had seized upon the main principles of evolutionary morphology: the indications then given are elaborated in the thirteenth chapter of the _Origin of Species_ (1st ed., 1859). A good part of this chapter is given up to a discussion of the principles of cla.s.sification, only a few pages dealing with morphology proper. But, as Darwin rightly saw, the two things are inseparable.

We note first that there is no hint of the "scale of beings"--Darwin conceives the genealogical tree as many branched. Animals can be cla.s.sed in "groups under groups," and cannot be arranged in one single series.

He discusses first what kind of characters have the greatest cla.s.sificatory value. Certain empirical rules have been recognised, more or less consciously, by systematists--that a.n.a.logical characters are less valuable than h.o.m.ological, that characters of great physiological importance are not always valuable for cla.s.sificatory purposes, that rudimentary organs are often very useful, and so on. He finds that as a general rule "the less any part of the organisation is concerned with special habits, the more important it becomes for cla.s.sification" (p.

414), and adduces in support Owen's remark that the generative organs afford very clear indications of affinities, since they are unlikely to be modified by special habits. These rules of cla.s.sification can be explained "on the view that the natural system is founded on descent with modification; that the characters which naturalists consider as showing true affinity ... are those which have been inherited from a common parent, and, in so far, all true cla.s.sification is genealogical; that community of descent is the hidden bond which naturalists have been unconsciously seeking, and not some unknown plan of creation, or the enunciation of general propositions, and the mere putting together and separating objects more or less alike" (p. 420).

In general, then, h.o.m.ological characters are more valuable for cla.s.sificatory purposes because they have a longer pedigree than a.n.a.logical characters, which represent recent acquirements of the race.

Coming to morphology proper, Darwin takes up the question of the unity of type, and the h.o.m.ology of parts, for which the unity of type is but a general expression.

He treats this on the same lines as E. Geoffroy St Hilaire, and Owen, referring indeed specifically to Geoffroy's law of connections. "What can be more curious," he asks, "than that the hand of a man, formed for grasping, that of a mole for digging, the leg of a horse, the paddle of the porpoise, and the wing of the bat, should all be constructed on the same pattern, and should include similar bones, in the same relative positions? Geoffroy St Hilaire has strongly insisted on the high importance of relative position or connection in h.o.m.ologous parts; they may differ to almost any extent in form and size, and yet remain connected together in the same invariable order" (p. 434).

The unity of plan cannot be explained on teleological grounds, as Owen has admitted in his _Nature of Limbs_, nor is it explicable on the hypothesis of special creation (p. 435). It can be understood only on the theory that animals are descended from one another and retain for innumerable generations the essential organisation of their ancestors.

"The explanation is to a large extent simple on the theory of the selection of successive slight modifications--each modification being profitable in some way to the modified form, but often affecting by correlation other parts of the organisation. In changes of this nature, there will be little or no tendency to alter the original pattern or to transpose the parts.... If we suppose that the ancient progenitor, the archetype as it may be called, of all animals, had its limbs constructed on the existing general pattern, for whatever purpose they served, we can at once perceive the plain significance of the h.o.m.ologous construction of the limbs throughout the whole cla.s.s" (p. 435).

We may note three important points in this pa.s.sage--first, the identification of the archetype with the common progenitor; second, the view that progressive evolution is essentially adaptive, and dominated by natural selection; and third, the _pet.i.tio principii_ involved in the a.s.sumption that adaptive modification brings inevitably in its train the necessary correlative changes.

In his section on morphology Darwin shows clearly the influence of Owen, and through him of the transcendental anatomists. He refers to the transcendental idea of "metamorphosis," as exemplified in the vertebral theory of the skull and the theory of the plant appendage, and shows how, on the hypothesis of descent with modification, "metamorphosis" may now be interpreted literally, and no longer figuratively merely (p.

439).

Very great interest attaches to Darwin's treatment of development, for post-Darwinian morphology was based to a very large extent on the presumed relation between the development of the individual and the evolution of the race. Just as he kept clear of the notion of the scale of beings, so he avoided the snare of the Meckel-Serres theory of recapitulation, according to which the embryo of the highest animal, man, during its development climbs the ladder upon the rungs of which the whole animal series is distributed, in its gradual progression from simplicity to complexity. The law of development which he adopts is that of von Baer, which states that development is essentially differentiation, and that as a result embryos belonging to the same group resemble one another the more the less advanced they are in development. There can be little doubt that he was indebted to von Baer for the idea, and in the later editions of the _Origin_ he acknowledges this by quoting the well-known pa.s.sage in which von Baer tells how he had two embryos in spirit which he was unable to refer definitely to their proper cla.s.s among Vertebrates.[354]

Not only are embryos more alike than adults, because less differentiated, but it is in points not directly connected with the conditions of existence, not strictly adaptive, that their resemblance is strongest (p. 440)--think, for instance, of the arrangement of aortic arches common to all vertebrate embryos. Larval forms are to some extent exceptions to this rule, for they are often specially adapted to their particular mode of life, and convergence of structure may accordingly result. All these facts require an explanation. "How, then, can we explain these several facts in embryology--namely, the very general, but not universal, difference in structure between the embryo and the adult--of parts in the same individual embryo, which ultimately become very unlike and serve for different purposes, being at this early period of growth alike--of embryos of different species within the same cla.s.s, generally but not universally, resembling each other--of the structure of the embryo not being closely related to its conditions of existence, except when the embryo becomes at any period of life active and has to provide for itself--of the embryo apparently having sometimes a higher organisation than the mature animal, into which it is developed" (pp.

442-3). Obviously all these facts are formally explained by the doctrine of descent. But Darwin goes further, he tries to show exactly how it is that the embryos resemble one another more than the adults. He thinks that the phenomenon results from two principles--first, that modifications usually supervene late in the life of the individual; and second, that such modifications tend to be inherited by the offspring at a corresponding, not early, age (p. 444).

Thus, applying these principles to a hypothetical case of the origin of new species of birds from a common stock, he writes:--"... from the many slight successive steps of variation having supervened at a rather late age and having been inherited at a corresponding age, the young of the new species of our supposed genus will manifestly tend to resemble each other much more closely than do the adults, just as we have seen in the case of pigeons"[355] (pp. 446-7).

Since the embryo shows the generalised type, the structure of the embryo is useful for cla.s.sificatory purposes. "For the embryo is the animal in its less modified state; and in so far it reveals the structure of its progenitor" (p. 449)--the embryological archetype reveals the ancestral form. "Embryology rises greatly in interest, when we thus look at the embryo as a picture, more or less complete, of the parent form of each great cla.s.s of animals" (p. 450)--a prophetic remark, in view of the enormous subsequent development of phylogenetic speculation.

We may sum up by saying that Darwin interpreted von Baer's law phylogenetically.

The rest of the chapter is devoted to a discussion of abortive and vestigial organs, whose existence Darwin naturally turns to great advantage in his argument for evolution. Throughout the whole chapter Darwin's preoccupation with the problems of cla.s.sification is clearly manifest.

On the question as to whether descent was monophyletic or polyphyletic Darwin expressed no dogmatic opinion. "I believe that animals have descended from at most only four or five progenitors, and plants from an equal or lesser number.... I should infer from a.n.a.logy that probably all the organic beings which have ever lived on this earth have descended from one primordial form, into which life was first breathed" (p. 484).

Darwin rightly laid much stress upon the morphological evidence for evolution,[356] which he considered to be weighty. It probably contributed greatly to the success of his theory. Though he himself did little or no work in pure morphology, he was alive to the importance of such work,[357]

and followed with interest the progress of evolutionary morphology, incorporating some of its results in later editions of the _Origin_, and in his _Descent of Man_ (1871).

In his morphology Darwin was hardly up to date. He does not seem to have known at first hand the splendid work of the German morphologists, such as Rathke and Reichert; he pays no attention to the cell-theory, nor to the germ-layer theory. His sources are, in the main, Geoffroy St Hilaire, Owen, von Baer, Aga.s.siz, Milne-Edwards, and Huxley.

Perhaps his greatest omission was that he did not give any adequate treatment of the problem of functional adaptation and the correlation of parts. It is not too much to say that Darwin not only disregarded these problems almost entirely, but by his insistence upon ecological adaptation and upon certain superficial aspects of correlation, succeeded in giving to the words "adaptation" and "correlation" a new signification, whereby they lost to a large extent their true and original functional meaning.

It is true that Darwin himself, as well as his successors, believed that natural selection was all-powerful to account for the evolution of the most complicated organs, but it may be questioned whether he realised all the conditions of the problem of which he thus easily disposed. He says, rightly, in an important pa.s.sage, that "It is generally acknowledged that all organic beings have been formed on two great laws--Unity of Type, and the Conditions of Existence. By unity of type is meant that fundamental agreement in structure which we see in organic beings of the same cla.s.s, and which is quite independent of their habits of life. On my theory, unity of type is explained by unity of descent.

The expression of conditions of existence, so often insisted upon by the ill.u.s.trious Cuvier, is fully embraced by the principle of natural selection. For natural selection acts by either now adapting _the varying parts of each being to its organic and inorganic conditions of life_:[358] or by having adapted them during past periods of time: the adaptations being aided in many cases by the increased use or disuse of parts, being affected by the direct action of the external conditions of life, and subjected in all cases to the several laws of growth and variation. Hence, in fact, the law of the Conditions of Existence is the higher law; as it includes, through the inheritance of former variations and adaptations, that of Unity of Type" (_Origin_, 6th ed., Pop.

Impression, pp. 260-1). It is clear that Darwin took the phrase "Conditions of Existence" to mean the environmental conditions, and the law of the Conditions of Existence to mean the law of adaptation to environment. But that is not what Cuvier meant by the phrase: he understood by it the principle of the co-ordination of the parts to form the whole, the essential condition for the existence of any organism whatsoever (see above, Chap. III., p. 34).

Of this thought there is in Darwin little trace, and that is why he did not sufficiently appreciate the weight of the argument brought against his theory that it did not account for the correlation of variations.

Darwin's conception of correlation was singularly incomplete. As examples of correlation he advanced such trivial cases as the relation between albinism, deafness and blue eyes in cats, or between the tortoise-sh.e.l.l colour and the female s.e.x. He used the word only in connection with what he called "correlated variation," meaning by this expression "that the whole organisation is so tied together during its growth and development, that when slight variations in any one part occur, and are acc.u.mulated through natural selection, other parts become modified" (6th ed., p. 177). He took it for granted that the "correlated variations" would be adapted to the original variation which was acted upon by natural selection, and he saw no difficulty in the gradual evolution of a complicated organ like the eye if only the steps were small enough. "It has been objected," he writes, "that in order to modify the eye and still preserve it as a perfect instrument, many changes would have to be effected simultaneously, which, it is a.s.sumed, could not be done through natural selection; but as I have attempted to show in my work on the variation of domestic animals, it is not necessary to suppose that the modifications were all simultaneous, if they were extremely slight and gradual" (6th ed., p. 226).

In post-Darwinian speculation the difficulty of explaining correlated variation by natural selection alone became more acutely realised, and it was chiefly this difficulty that led Weismann to formulate his hypothesis of germinal selection as a necessary supplement to the general selection theory.

The change in the conception of correlation which Darwin's influence brought about has been very clearly stated by E. von Hartmann,[359] from whom the following is taken:--"While the correlation of parts in the organism was before Darwin regarded exclusively from the standpoint of morphological systematics, Darwin tried to look at it from the standpoint of physiological and genealogical development, and in so doing he put the standpoint of morphological systematics in the shade.

But the more we are now beginning to realise that systematic relationship does not necessarily imply genetic affinity the more must the correlation of parts come back into favour as a systematic principle. While Darwin only, as it were, against his will, relied on the law of correlation as a last resort when all other help failed, this law must be regarded, from the standpoint of the orderly inner determination of all organic form-change, as having the rank of the highest principle of all, a principle which rules parallel, divergent and convergent evolution" (pp. 47-8).

Further on, following Radl, he characterises Darwin's att.i.tude to the law of correlation in these terms:--"Darwin's interest is entirely focussed on the variation, the function, the causes of form-production, in short, upon evolution. Accordingly he regards correlation essentially as correlative variation in the sense of a _departure_ from the given type. With morphological correlation in _different_ types Darwin troubles himself not at all, nor with correlation in the normal development of a type" (p. 49).

Cuvier's conception of the _convenance des parties_, essential to all biology, remained on the whole foreign to Darwin's thought, and to the thought of his successors.

It was indeed one of their boasts that they had finally eliminated all teleology from Nature. The great and immediate success which Darwinism had among the younger generation of biologists and among scientific men in general was due in large part to the fact that it fitted in well with the prevailing materialism of the day, and gave solid ground for the hope that in time a complete mechanistic explanation of life would be forthcoming. "Darwinismus" became the battle-cry of the militant spirits of that time.

It was precisely this element in Darwinism that was repugnant to most of Darwin's opponents, in whose ranks were found the majority of the morphologists of the old school. They found it impossible to believe that evolution could have come about by fortuitous variation and fortuitous selection; they objected to Darwin that he had enunciated no real _Entwickelungsgesetz_, or law governing evolution. They were not unwilling to believe that evolution was a real process, though many drew the line at the derivation of man from apes, but they felt that if evolution had really taken place, it must have been under the guidance of some principle of development, that there must have been manifested in evolution some definite and orderly tendency towards perfection.[360]