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Darwin, and After Darwin (Vol. 1 and 3, of 3)
by George John Romanes
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Another and closely allied objection is, that the theory of natural selection "personifies an abstraction." Or, as the Duke of Argyll states it, the theory is "essentially the image of mechanical necessity concealed under the clothes, and parading in the mask, of mental purpose. The word 'natural' suggests Matter, and the physical forces. The word 'selection' suggests Mind, and the powers of choice." This, however, is a mere quarrelling about words. Darwin called the principle which he had discovered by the name natural selection in order to mark the analogy between it and artificial selection. No doubt in this analogy there is not necessarily supposed to be in nature any counterpart to the mind of the breeder, nor, therefore, to his powers of intelligent choice. But there is no need to limit the term selection (se and lego, Gr. [Greek: lego]) to powers of intelligent choice. As previously remarked, a bank of sea-weed on the sea-shore may be said to have been selected by the waves from all the surrounding sand and stones. Similarly, we may say that grain is selected from chaff by the wind in the process of winnowing corn. Or, if it be thought that there is any ambiguity involved in such a use of the term in the case of "Natural Selection," there is no objection to employing the phrase which has been coined by Mr. Spencer as its equivalent—namely, "Survival of the Fittest." The point of the theory is, that those organisms which are best suited to their surroundings are allowed to live and to propagate, while those which are less suited are eliminated; and whether we call this process a process of selection, or call it by any other name, is clearly immaterial.

A material question is raised only when it is asked whether the process is one that can be ascribed to causation strictly natural. It is often denied that such is the case, on the ground that natural selection does not originate the variations which it favours, but depends upon the variations being supplied by some other means. For, it is said, all that natural selection does is to preserve the suitable variations after they have arisen. Natural selection does not cause these suitable variations; and therefore, it is argued, Darwin and his followers are profoundly mistaken in representing the principle as one which produces adaptations. Now, although this objection has been put forward by some of the most intelligent minds in our generation, it appears to me to betoken some extraordinary failure to appreciate the very essence of Darwinian doctrine. No doubt it is perfectly true that natural selection does not produce variations of any kind, whether beneficial or otherwise. But if it be granted that variations of many kinds are occurring in every generation, and that natural selection is competent to preserve the more favourable among them, then it appears to me unquestionable that this principle of selection deserves to be regarded as, in the full sense of the word, a natural cause. The variations being expressly regarded by the theory as more or less promiscuous[42], survival of the fittest becomes the winnowing fan, whose function it is to eliminate all the less fit in each generation, in order to preserve the good grain, out of which to constitute the next generation. And as this process is supposed to be continuous through successive generations, its action is supposed to be cumulative, till from the eye of a worm there is gradually developed the eye of an eagle. Therefore it follows from these suppositions (which are not disputed by the present objection), that if it had not been for the process of selection, such development would never have been begun; and that in the exact measure of its efficiency will the development proceed. But any agency without the operation of which a result cannot take place may properly be designated the cause of that result: it is the agency which, in co-operation with all the other agencies in the cosmos, produces that result.

[42] The degree in which variability is indefinite, or, on the contrary, determinate, is a question which is not yet ripe for decision—nor even, in my opinion, for discussion. But I may here state the following general principles with regard to it.

(1) It is evident that up to some point or another variations must be pre-determined in definite lines. Men do not gather grapes from thorns, figs from thistles, nor even moss-roses from sweet-briars. In other words, "the nature of the organism" in all cases necessitates the limiting of variations within certain bounds.

(2) But when the question is as to what these bounds may be, we can only answer in a general way that, according to the general theory of evolution, they must be such as are imposed by heredity, coupled with the degree to which external conditions of life (and possibly also use-inheritance) are capable, in given cases, of modifying congenital characters. These are the only causes which the theory of descent can consistently recognise as producing variations in determinate directions.

(3) Inasmuch as variation presupposes the existence of parts that vary, and inasmuch as the variation of parts can only be in the alternative directions of increase or decrease around an average, it follows that, in the first instance at all events, every variation, if determinate, must be so only in one or other of these two opposite directions.

(4) In as far as variations are summated in successive generations, so as eventually to give rise to new structures, organs, mechanisms, &c., natural selection is theoretically competent to explain the facts, without our having to postulate the operation of unknown causes producing variations in determinate lines,—or not further than is stated in paragraphs 1 and 2.

(5) Nevertheless, it does not follow that there are not such other unknown causes; and, if there are, of course the importance of natural selection as a cause of adaptive modification would be limited in proportion to their number and the extent of their operation. But it is for those who, like the late Professors Asa Gray and Naegeli, maintain the existence of such causes, to substantiate their belief by indicating them.

Take any analogous case. The selective agency of specific gravity which is utilised in gold-washing does not create the original differences between gold-dust and dust of all other kinds. But these differences being presented by as many different bodies in nature, the gold-washer takes advantage of the selective agency in question, and, by using it as a cause of segregation, is enabled to separate the gold from all the earths with which it may happen to be mixed. So far as the objects of the gold-washer are concerned, it is immaterial with what other earths the gold-dust may happen to be mixed. For although gold-dust may occur in intimate association with earths of various kinds in various proportions, and although in each case the particular admixture which occurs must have been due to definite causes, these things, in relation to the selective process of the washer, are what is called accidental: that is to say, they have nothing to do with the causative action of the selective process. Now, in precisely the same sense Darwin calls the multitudinous variations of plants and animals accidental. By so calling them he expressly says he does not suppose them to be accidental in the sense of not all being due to definite causes. But they are accidental in relation to the sifting process of natural selection: all that they have to do is to furnish the promiscuous material on which this sifting process acts.

Or let us take an even closer analogy. The power of selective breeding by man is so wonderful, that in the course of successive generations all kinds of peculiarities as to size, shape, colour, special appendages or abortions, &c., can be produced at pleasure, as we saw in the last chapter. Now all the promiscuous variations which are supplied to the breeder, and out of which, by selecting only those that are suited to his purpose, he is able to produce the required result—all those promiscuous variations, in relation to that purpose, are accidental. Therefore the selective agency of the breeder deserves to be regarded as the cause of that which it produces, or of that which could not have been produced but for the operation of such agency. But where is the difference between artificial and natural selection in this respect? And, if there is no difference, is not natural selection as much entitled to be regarded as a true cause of the origin of natural species, as artificial selection is to be regarded as a true cause of our domesticated races? Here, as in the case of the previous illustration, if there be any ambiguity in speaking of variations as accidental, it arises from the incorrect or undefined manner in which the term "accidental" is used by Darwin's critics. In its original and philosophically-correct usage, the term "accident" signifies a property or quality not essential to our conception of a substance: hence, it has come to mean anything that happens as a result of unforeseen causes—or, lastly, that which is causeless. But, as we know that nothing can happen without causes of some kind, the term "accident" is divested of real meaning when it is used in the last of these senses. Yet this is the sense that is sought to be placed upon it by the objection which we are considering. If the objectors will but understand the term in its correct philosophical sense—or in the only sense in which it presents any meaning at all,—they will see that Darwinians are both logically and historically justified in employing the word "accidental" as the word which serves most properly to convey the meaning that they intend—namely, variations due to causes accidental to the struggle for existence. Similarly, when it is said that variations are "spontaneous," or even "fortuitous," nothing further is meant than that we do not know the causes which lead to them, and that, so far as the principle of selection is concerned, it is immaterial what these causes may be. Or, to revert to our former illustration, the various weights of different kinds of earths are no doubt all due to definite causes; but, in relation to the selective action of the gold-washer, all the different weights of whatever kinds of earth he may happen to include in his washing-apparatus are, strictly speaking, accidental. And as at different washings he meets with different proportions of heavy earths with light ones, and as these "variations" are immaterial to him, he may colloquially speak of them as "fortuitous," or due to "chance," even though he knows that at each washing they must have been determined by definite causes.

More adequately to deal with this merely formal objection, however, would involve more logic-chopping than is desirable on the present occasion. But I have already dealt with it fully elsewhere,—viz. in The Contemporary Review for June, 1888, to which therefore I may refer any one who is interested in dialectics of this kind[43].

[43] Within the last few months this objection has been presented anew by Mr. D. Syme, whose book On the Modification of Organisms exhibits a curious combination of shrewd criticisms with almost ludicrous misunderstandings. One of the latter it is necessary to state, because it pervades the quotation which I am about to supply. He everywhere compares "natural selection" with "the struggle for existence," uses them as convertible terms, and while absurdly stating that "Darwin defines natural selection as the struggle for existence," complains of "the liability of error, both on his own part and on the part of his readers," which arises from his not having everywhere adhered to this definition! (p. 8).

"Darwin has put forth two distinct and contradictory theories of the functions of natural selection. According to the one theory natural selection is selective or preservative, and nothing more. According to the other theory natural selection creates the variations(!) ... It certainly seems absurd to speak of natural selection, or the struggle for existence, as selective or preservative, for the struggle for existence does not preserve at all, not even the fit variations, as both the fit and the unfit struggle for existence, the unfit naturally more than the fit, and the fit are preserved, not in consequence of the struggle, but in consequence of their fitness. Suppose two varieties of the same species are driven, by an increase of their numbers, to seek for subsistence in a colder region than they have been accustomed to, and that one of these varieties had a hardier constitution than the other; and let us suppose that the former withstood the severe climate better than the latter, and consequently survived, while the other perished. In this case the hardier survived, not because of the struggle, but because it had a constitution better adapted to the climate. I wish to ascertain if a certain metal in my possession is gold or some baser metal, and I apply the usual test; but the mere fact of my testing this metal would not make it gold or any other kind of metal."

I have thought it worth while to quote this passage for the sake of showing the extraordinary confusion of mind which still prevails on the part of Darwin's critics, even with reference to the very fundamental parts of his theory. For, as I have said, the writer of this passage shows himself a shrewd critic in some other parts of his essay, where he is not engaged especially on the theory of natural selection.

I will now pass on to consider another misconception of the Darwinian theory, which is very prevalent in the public mind. It is virtually asked, If some species are supposed to have been improved by natural selection, why have not all species been similarly improved? Why should not all invertebrated animals have risen into vertebrated? Or why should not all monkeys have become men?

The answers are manifold. In the first place, it by no means follows that because an advance in organization has proved itself of benefit in the case of one form of life, therefore any or every other form would have been similarly benefited by a similar advance. The business of natural selection is to bring this and that form of life into the closest harmony with its environment that all the conditions of the case permit. Sometimes it will happen that the harmony will admit of being improved by an improvement of organization. But just as often it will happen that it will be best secured by leaving matters as they are. If, therefore, an organism has already been brought into a tolerably full degree of harmony with its environment, natural selection will not try to change it so long as the environment remains unchanged; and this, no doubt, is the reason why some species have survived through enormous periods of geological time without having undergone any change. Again, as we saw in a previous chapter, there are yet other cases where, on account of some change in the environment or even in the habits of the organisms themselves, adaption will be best secured by an active reversal of natural selection, with the result of causing degeneration.

But, it is sometimes further urged, there are cases where we cannot doubt that improvement of organization would have been of benefit to species; and yet such improvement has not taken place—as, for instance, in the case all monkeys not turning into men. Here, however, we must remember that the operation of natural selection in any case depends upon a variety of highly complex conditions; and, therefore, that the fact of all those conditions having been satisfied in one instance is no reason for concluding that they must also have been satisfied in other instances. Take, for example, the case of monkeys passing into men. The wonder to me appears to be that this improvement should have taken place in even one line of descent; not that, having taken place in one line, it should not also have taken place in other lines. For how enormously complex must have been the conditions—physical, anatomical, physiological, psychological, sociological—which by their happy conjunction first began to raise the inarticulate cries of an ape into the rational speech of a man. Therefore, the more that we appreciate the superiority of a man to an ape, the less ought we to countenance this supposed objection to Darwin's theory—namely, that natural selection has not effected the change in more than one line of descent.

Even in the case of two races of mankind where one has risen higher in the scale of civilization than another, it is now generally impossible to assign the particular causes of the difference; much more, then, must this be impossible in the case of still more remote conditions which have led to the divergence of species. The requisite variations may not have arisen in the one line of descent which did arise in the other; or if they did arise in both, some counterbalancing disadvantages may have attended their initial development in the one case which did not obtain in the other. In short, where so exceedingly complex a play of conditions are concerned, the only wonder would be if two different lines of descent had happened to present two independent and yet perfectly parallel lines of history.

These general considerations would apply equally to the great majority of other cases where some types have made great advances upon others, notwithstanding that we can see no reason why the latter should not in this respect have imitated the former. But there is yet a further consideration which must be taken into account. The struggle for existence is always most keen between closely allied species, because, from the similarity of their forms, habits, needs, &c., they are in closest competition. Therefore it often happens that the mere fact of one species having made an advance upon others of itself precludes the others from making any similar advance: the field, so to speak, has already been occupied as regards that particular improvement, and where the struggle for existence is concerned possession is emphatically nine points of the law. For example, to return to the case of apes becoming men, the fact of one rational species having been already evolved (even if the rational faculty were at first but dimly nascent) must make an enormous change in the conditions as regards the possibility of any other such species being subsequently evolved—unless, of course, it be by way of descent from the rational one. Or, as Sir Charles Lyell has well put it, two rational species can never coexist on the globe, although the descendants of one rational species may in time become transformed into another single rational species[44].

[44] Principles of Geology, vol. ii. p. 487 (11th ed.).

In view of such considerations, another and exactly opposite objection has sometimes been urged—viz. that we ought never to find inferior forms of organization in company with superior, because in the struggle for existence the latter ought to have exterminated the former. Or, to quote the most recent expression of this view, "in every locality there would only be one species, and that the most highly organized; and thus a few superior races would partition the earth amongst them to the entire exclusion of the innumerable varieties, species, genera, and orders which now inhabit it[45]." Of course to this statement it would be sufficient to enquire, On what would these few supremely organized species subsist? Unless manna fell from heaven for their especial benefit, it would appear that such forms could under no circumstances be the most improved forms; in exterminating others on such a scale as this, they would themselves be quickly, and very literally, improved off the face of the earth. But even when the statement is not made in so extravagant a form as this, it must necessarily be futile as an objection unless it has first been shown that we know exactly all the conditions of the complex struggle for existence between the higher and lower forms in question. And this it is impossible that we ever can know. The mere fact that one form has been changed in virtue of this struggle must in many cases of itself determine a change in the conditions of the struggle. Again, the other and closely allied forms (and these furnish the best grounds for the objection) may also have undergone defensive changes, although these may be less conspicuous to our observation, or perhaps less suggestive of "improvement" to our imperfect means of judging. Lastly, not to continue citing an endless number of such considerations, there is the broad fact that it is only to those cases where, for some reason or another, the lower forms have not been exposed to a struggle of fatal intensity, that the objection applies. But we know that in millions of other cases the lower (i. e. less fitted) forms have succumbed, and therefore I do not see that the objection has any ground to stand upon. That there is a general tendency for lower forms to yield their places to higher is shown by the gradual advance of organization throughout geological time; for if all the inferior forms had survived, the earth could not have contained them, unless she had been continually growing into something like the size of Jupiter. And if it be asked why any of the inferior forms have survived, the answer has already been given, as above.

[45] Syme, on the Modification of Organisms, p. 46.

There is only one other remark to be made in this connexion. Mr. Syme chooses two cases as illustrations of the supposed difficulty. These are sufficiently diverse—viz. Foraminifera and Man. Touching the former, there is nothing that need be added to the general answer just given. But with regard to the latter it must be observed that the dominion of natural selection as between different races of mankind is greatly restricted by the presence of rationality. Competition in the human species is more concerned with wits and ideas than with nails and teeth; and therefore the "struggle" between man and man is not so much for actual being, as for well-being. Consequently, in regard to the present objection, the human species furnishes the worst example that could have been chosen.

* * * * *

Hitherto I have been considering objections which arise from misapprehensions of Darwin's theory. I will now go on to consider a logically sound objection, which nevertheless is equally futile, because, although it does not depend on any misapprehension of the theory, it is not itself supported by fact.

The objection is the same as that which we have already considered in relation to the general theory of descent—namely, that similar organs or structures are to be met with in widely different branches of the tree of life. Now this would be an objection fatal to the theory of natural selection, supposing these organs or structures in the cases compared are not merely analogous, but also homologous. For it would be incredible that in two totally different lines of descent one and the same structure should have been built up independently by two parallel series of variations, and that in these two lines of descent it should always and independently have ministered to the same function. On the other hand, there would be nothing against the theory of natural selection in the fact that two structures, not homologous, should come by independent variation in two different lines of descent to be adapted to perform the same function. For it belongs to the very essence of the theory of natural selection that a useful function should be secured by favourable variations of whatever structural material may happen to be presented by different organic types. Flying, for instance, is a very useful function, and it has been developed independently in at least four different lines of descent—namely, the insects, reptiles, birds, and mammals. Now if in all, or indeed in any, of these four cases the wings had been developed on the same anatomical pattern, so as not only to present the analogical resemblance which it is necessary that they should present in order to discharge their common function of flying, but likewise an homologous or structural resemblance, showing that they had been formed on the same anatomical "plan,"—if such has been the case, I say, the theory of natural selection would certainly be destroyed.

Now it has been alleged by competent naturalists that there are several such cases in organic nature. We have already noticed in a previous chapter (pp. 58, 59), that Mr. Mivart has instanced the eye of the cuttle-fish as not only analogous to, but also homologous with, the eye of a true fish—that is to say, the eye of a mollusk with the eye of a vertebrate. And he has also instanced the remarkable resemblance of a shrew to a mouse—that is, of an insectivorous mammal to a rodent—not to mention other cases. In the chapter alluded to these instances of homology, alleged to occur in different branches of the tree of life, were considered with reference to the process of organic evolution as a fact: they are now being considered with reference to the agency of natural selection as a method. And just as in the former case it was shown, that if any such alleged instances could be proved, the proof would be fatal to the general theory of organic evolution by physical causes, so in the present case, if this could be proved, it would be equally fatal to the more special theory of natural selection. But, as we have before seen, no single case of this kind has ever been made out; and, therefore, not only does this supposed objection fall to the ground, but in so doing it furnishes an additional argument in favour of natural selection. For in the earlier chapter just alluded to I showed that this great and general fact of our nowhere being able to find two homologous structures in different branches of the tree of life, was the strongest possible testimony in favour of the theory of evolution. And, by parity of reasoning, I now adduce it as equally strong evidence of natural selection having been the cause of adaptive structures, independently developed in all the different lines of descent. For the alternative is between adaptations having been caused by natural selection or by supernatural design. Now, if adaptations were caused by natural selection, we can very well understand why they should never be homologous in different lines of descent, even in cases where they have been brought to be so closely analogous as to have deceived so good a naturalist as Mr. Mivart. Indeed, as I have already observed, so well can we understand this, that any single instance to the contrary would be sufficient to destroy the theory of natural selection in toto, unless the structure be one of a very simple type. But on the other hand, it is impossible to suggest any rational explanation why, if all adaptations are due to supernatural design, such scrupulous care should have been taken never to allow homologous adaptations to occur in different divisions of the animal or vegetable kingdoms. Why, for instance, should the eye of a cuttle-fish not have been constructed on the same ideal pattern as that of vertebrate? Or why, among the thousands of vertebrated species, should no one of their eyes be constructed on the ideal pattern that was devised for the cuttle-fish? Of course it may be answered that perhaps there was some hidden reason why the design should never have allowed an adaptation which it had devised for one division of organic nature to appear in another—even in cases where the new design necessitated the closest possible resemblance in everything else, save in the matter of anatomical homology. Undoubtedly such may have been the case—or rather such must have been the case—if the theory of special design is true. But where the question is as to the truth of this theory, I think there can be no doubt that its rival gains an enormous advantage by being able to explain why the facts are such as they are instead of being obliged to take refuge in hypothetical possibilities of a confessedly unsubstantiated and apparently unsubstantial kind.

Therefore, as far as this objection to the theory of natural selection is concerned—or the allegation that homologous structures occur in different divisions of organic nature—not only does it fall to the ground, but positively becomes itself converted into one of the strongest arguments in favour of the theory. As soon as the allegation is found to be baseless, the very fact that it cannot be brought to bear upon any one of all the millions of adaptive structures in organic nature becomes a fact of vast significance on the opposite side.

* * * * *

The next difficulty to which I shall allude is that of explaining by the theory of natural selection the preservation of the first beginnings of structures which are then useless, though afterwards, when more fully developed, they become useful. For it belongs to the very essence of the theory of natural selection, that a structure must be supposed already useful before it can come under the influence of natural selection: therefore the theory seems incapable of explaining the origin and conservation of incipient organs, or organs which are not yet sufficiently developed to be of any service to the organisms presenting them.

This objection is one that has been advanced by all the critics of Darwinism; but has been presented with most ability and force by the Duke of Argyll. I will therefore state it in his words.

If the doctrine of evolution be true—that is to say, if all organic creatures have been developed by ordinary generation from parents—then it follows of necessity that the primaeval germs must have contained potentially the whole succeeding series. Moreover, if that series has been developed gradually and very slowly, it follows, also as a matter of necessity, that every modification of structure must have been functionless at first, when it began to appear.... Things cannot be selected until they have first been produced. Nor can any structure be selected by utility in the struggle for existence until it has not only been produced, but has been so far perfected as to actually be used.

The Duke proceeds to argue that all adaptive structures must therefore originally have been due to special design: in the earlier stages of their development they must all have been what he calls "prophetic germs." Not yet themselves of any use, and therefore not yet capable of being improved by natural selection, both in their origin and in the first stages (at all events) of their development, they must be regarded as intentionally preparatory to the various uses which they subsequently acquire.

Now this argument, forcible as it appears at first sight, is really at fault both in its premiss and in its conclusion. By which I mean that, in the first place the premiss is not true, and, in the next place, that even if it were, the conclusion would not necessarily follow. The premiss is, "that every modification of structure must have been functionless at first, when it began to appear;" and the conclusion is, that, qua functionless, such a modification cannot have been caused by natural selection. I will consider these two points separately.

First as to the premiss, it is not true that every modification of structure must necessarily be functionless when it first begins to appear. There are two very good reasons why such should not be the case in all instances, even if it should be the case in some. For, as a matter of observable fact, a very large proportional number of incipient organs are useful from the very moment of their inception. Take, for example, what is perhaps the most wonderful instance of refined mechanism in nature—the eye of a vertebrated animal. Comparative anatomy and embryology combine to testify that this organ had its origin in modifications of the endings of the ordinary nerves of the skin. Now it is evident that from the very first any modification of a cutaneous nerve whereby it was rendered able, in however small a degree, to be differently affected by light and by darkness would be of benefit to the creature presenting it; for the creature would thus be able to seek the one and shun the other according to the requirements of its life. And being thus useful from the very moment of its inception, it would afterwards be gradually improved as variations of more and more utility presented themselves, until not only would finer and finer degrees of difference between light and shade become perceptible, but even the outlines of solid bodies would begin to be appreciated. And so on, stage by stage, till from an ordinary nerve-ending in the skin is evolved the eye of an eagle.

Moreover, in this particular instance there is very good reason to suppose that the modification of the cutaneous nerves in question began by a progressive increase in their sensitiveness to temperature. Wherever dark pigment happened to be deposited in the skin—and we know that in all animals it is apt to be deposited in points and patches, as it were by accident, or without any "prophecy" as to future uses,—the cutaneous nerves in its vicinity would be better able to appreciate the difference between sun and shade in respect of temperature, even though as yet there were no change at all in these cutaneous nerves tending to make them responsive to light. Now it is easy to see how, from such a purely accidental beginning, natural selection would have had from the first sufficient material to act upon. It being of advantage to a lowly creature that it should distinguish with more and more delicacy, or with more and more rapidity, between light and darkness by means of its thermal sensations, the pigment spots in the skin would be rendered permanent by natural selection, while the nerves in that region would by the same agency be rendered more and more specialized as organs adapted to perceive changes of temperature, until from the stage of responding to the thermal rays of the non-luminous spectrum alone, they become capable of responding also to luminous.

So much, then, for the first consideration which serves to invalidate the Duke's premiss. The second consideration is, that very often an organ which began by being useful for the performance of one function, after having been fully developed for the performance of that function, finds itself, so to speak, accidentally fitted to the performance of some other and even more important function, which it thereupon begins to discharge, and so to undergo a new course of adaptive development. In such cases, and so far as the new function is concerned, the difficulty touching the first inception of an organ does not apply; for here the organ has already been built up by natural selection for one purpose, before it begins to discharge the other. As an example of such a case we may take the lung of an air-breathing animal. Originally the lung was a swim-bladder, or float, and as such it was of use to the aquatic ancestors of terrestrial animals. But as these ancestors gradually became more and more amphibious in their habits, the swim-bladder began more and more to discharge the function of a lung, and so to take a wholly new point of departure as regards its developmental history. But clearly there is here no difficulty with regard to the inception of its new function, because the organ was already well developed for one purpose before it began to serve another. Or, to take only one additional example, there are few structures in the animal kingdom so remarkable in respect of adaptation as is the wing of a bird or a bat; and at first sight it might well appear that a wing could be of no conceivable use until it had already acquired enormous proportional dimensions, as well as an immense amount of special elaboration as to its general form, size of muscle, amount of blood-supply, and so on. For, obviously, not until it had attained all these things could it even begin to raise the animal in the air. But observe how fallacious is this argument. Although it is perfectly true that a wing could be of no use as a wing until sufficiently developed to serve the purpose of flight, this is merely to say that until it has become a wing it is no use as a wing. It does not, however, follow that on this account it was of no prior use for any other purpose. The first modifications of the fore-limb which ended in its becoming an organ of flight may very well have been due to adapting it as an organ for increased rapidity of locomotion of other kinds—whether on land as in the case of its now degenerated form in the ostrich, or in water as in the case of the expanded fins of fish. Indeed, we may see the actual process of transition from the one function to the other in the case of "flying-fish." Here the progressive expansion of the pectoral fins must certainly have been always of use for continuously promoting rapidity of locomotion through water; and thus natural selection may have continuously increased their development until they now begin to serve also as wings for carrying the animal a short distance through air. Again, in the case of the so-called flying squirrels we find the limbs united to the body by means of large extensions of the skin, so-that when jumping from one tree to another the animal is able to sustain itself through a long distance in the air by merely spreading out its limbs, and thus allowing the skin-extensions to act after the manner of a parachute. Here, of course, we have not yet got a wing, any more than we have in the case of the flying-fish; but we have the foundations laid for the possible development of a future wing, upon a somewhat similar plan as that which has been so wonderfully perfected in the case of bats. And through all the stages of progressive expansion which the skin of the squirrel has undergone, the expansion has been of use, even though it has not yet so much as begun to acquire the distinctive functions of a wing. Here, then, there is obviously nothing "prophetic" in the matter, any more than there was in the case of the swim-bladder and the lung, or in that of the nerve-ending and the eye. In short, it is the business of natural selection to secure the highest available degree of adaptation for the time being; and, in doing this, it not unfrequently happens that an extreme development of a structure in one direction (produced by natural selection for the sake of better and better adapting the structure to perform some particular function) ends by beginning to adapt it to the performance of some other function. And, whenever this happens to be the case, natural selection forthwith begins to act upon the structure, so to speak, from a new point of departure.

So much, then, for the Duke's premiss—namely, that "every modification of structure must have been functionless at first, when it began to appear." This premiss is clearly opposed to observable fact. But now, the second position is that, even if this were not so, the Duke's conclusion would not follow. This conclusion, it will be remembered, is, that if incipient structures are useless, it necessarily follows that natural selection can have had no part whatever in their inception. Now, this is a conclusion which does not "necessarily" follow. Even if it be granted that there are structures which in their first beginnings are not of any use at all for any purpose, it is still possible that they may owe their origin to natural selection—not indeed directly, but indirectly. This possibility arises from the occurrence in nature of a principle which has been called the Correlation of Growth.

Mr. Darwin, who has paid more attention to this matter than any other writer, has shown, in considerable detail, that all the parts of any given organism are so intimately bound together, or so mutually dependent upon each other, that when one part is caused to change by means of natural selection, some other parts are very likely to undergo modification as a consequence. For example, there are several kinds of domesticated pigeons and fowls, which grow peculiar wing-like feathers on the feet. These are quite unlike all the other feathers in the animal, except those of the wing, to which they bear a very remarkable resemblance. Mr. Darwin records the case of a bantam where these wing-like feathers were nine inches in length, and I have myself seen a pigeon where they reproduced upon the feet a close imitation of the different kinds of feathers which occupy homologous positions in the wing—primaries, secondaries, and tertiaries all being distinctly repeated in their proper anatomical relations. Furthermore, in this case, as in most cases where such wing-feathers occur upon the feet, the third and fourth toes were partly united by skin; and, as is well known, in the wing of a bird the third and fourth digits are completely united by skin; "so that in feather-footed pigeons, not only does the exterior surface support a row of long feathers, like wing-feathers [which, as just stated, may in some cases be obviously differentiated into primaries, secondaries and tertiaries], but the very same digits which in the wing are completely united by skin become partially united by skin in the feet; and thus by the law of correlated variation of homologous parts, we can understand the curious connexion of feathered legs and membrane between the two outer toes[46]." The illustration is drawn from the specimen to which I have referred.

[46] Variation of Plants and Animals, vol. ii. p. 315.



Many similar instances of the same law are to be met with throughout organic nature; and it is evident that in this principle we find a conceivable explanation of the origin of such adaptive structures as could not have been originated by natural selection acting directly upon themselves: they may have been originated by natural selection developing other adaptive structures elsewhere in the organism, the gradual evolution of which has entailed the production of these by correlation of growth. And, if so, when once started in this way, these structures, because thus accidentally useful, will now themselves come under the direct action of natural selection, and so have their further evolution determined with or without the correlated association which first led to their inception.

Of course it must be understood that in thus applying the principle of correlated growth, to explain the origin of adaptive structures where it is impossible to explain such origin by natural selection having from the first acted directly upon these structures themselves, Darwinists do not suppose that in all—or even in most—cases of correlated growth the correlated structures are of use. On the contrary, it is well known that structures due to correlated growth are, as a rule, useless. Being only the by-products of adaptive changes going on elsewhere, in any given case the chances are against these correlated effects being themselves of any utilitarian significance; and, therefore, as a matter of fact, correlated growths appear to be usually meaningless from the point of view of adaptation. Still, on the doctrine of chances, it is to be expected that sometimes a change of structure which has thus been indirectly produced by correlation of growth might happen to prove useful for some purpose or another; and in as many cases as such indirectly produced structures do prove useful, they will straightway begin to be improved by the direct action of natural selection. In all such cases, therefore, we should have an explanation of the origin of such a structure, which is the only point that we are now considering.

I think, then, that all this effectually disposes of the doctrine of "prophetic germs." But, before leaving the subject, I should like to make one further statement of greater generality than any which I have hitherto advanced. This statement is, that we must remember how large a stock of meaningless structures are always being produced in the course of specific transmutations, not only by correlation of growth, which we have just been considering, but also by the direct action of external conditions, together with the constant play of all the many and complex forces internal to organisms themselves. In other words, important as the principle of correlation undoubtedly is, we must remember that even this is very far from being the only principle which is concerned in the origination of structures that may or may not chance to be useful. Therefore, it is not only natural selection when operating indirectly through the correlation of growth that is competent to produce new structures without reference to utility. In all the complex action and reaction of internal and external forces, new variations are perpetually arising without any reference to utility, either present or future. Among all this multitude of promiscuous variations, the chances must be that some percentage will prove of some service, either from the first moment of their appearance, or else after they have undergone some amount of development. Such development prior to utility may be due, either to correlation of growth, to the structure having previously performed some other function, as already explained, or else to a continued operation of the causes which were concerned in the first appearance of originally useless characters. In a series of chapters which will be devoted to the whole question of utility in the next volume, I shall hope to give very good reasons for concluding that useless characters are not only of highly frequent occurrence, but are due to a variety of other causes besides correlation of growth. And, if so, the possibility of originally useless characters happening in some cases to become, by increased development, useful characters, is correspondingly increased. Among a hundred varietal or specific characters which are directly produced in as many different species by a change of climate, for example, some five or six may be potentially useful: that is to say, characters thus adventitiously produced in an incipient form may only require to be further developed by a continuance of the same causes as first originated them, in order that some percentage of the whole number shall become of some degree of use. Those professed followers of Darwin, therefore, who without any reason—or, as it appears to me, against all reason—deny the possibility of useless specific characters in any case or in any degree (unless correlated with useful characters), are playing into the hands of Darwin's critics by indirectly countenancing the difficulty which we are now considering. For, if correlation of growth is unreasonably supposed to be the only possible cause of the origin of incipient structures which are not useful from the first moment of their inception, clearly the field is greatly narrowed as regards the occurrence of incipient characters sufficient in amount—and, still more, in constancy of appearance and persistency of transmission—to admit of furnishing material for the working of natural selection. But in the measure that incipient characters—whether varietal or specific—are recognised as not always or "necessarily" useful from the moment of their inception, and yet capable of being developed to a certain extent by the causes which first led to their occurrence, in that measure is this line of criticism closed. For of all the variations which thus occur, it is only those which afterwards prove of any use that are laid hold upon and wrought up by natural selection into adaptive structures, or working organs. And, therefore, what we see in organic nature is the net outcome of the development of all the happy chances. So it comes that the appearance presented by organic nature as a whole is that of a continual fulfilment of structural prophecies, when, in point of fact, if we had a similar record of all the other variations it would be seen that possibly not one such prophecy in a thousand is ever destined to be fulfilled.

* * * * *

Here, then, I feel justified in finally taking leave of the difficulty from the uselessness of incipient organs, as this difficulty has been presented, in varying degrees of emphasis, by the Duke of Argyll, Mr. Mivart, Professors Naegeli, Bronn, Broca, Eimer, and, indeed, by all other writers who have hitherto advanced it. For, as thus presented, I think I have shown that it admits of being adequately met. But now, I must confess, to me individually it does appear that behind this erroneous presentation of the difficulty there lies another question, which is deserving of much more serious attention. For although it admits of being easily shown—as I have just shown—that the difficulty as ordinarily presented fails on account of its extravagance, the question remains whether, if stated with more moderation, a real difficulty might not be found to remain.

My quarrel with the conclusion, like my quarrel with the premiss, is due to its universality. By saying in the premiss that all incipient organs are necessarily useless at the time of their inception, these writers admit of being controverted by fact; and by saying in the conclusion that, if all incipient organs are useless, it necessarily follows that in no case can natural selection have been the cause of building up an organ until it becomes useful, they admit of being controverted by logic. For, even if the premiss were true in fact—namely, that all incipient organs are useless at the time of their inception,—it would not necessarily follow that in no case could natural selection build up a useless structure into a useful one; because, although it is true that in no case can natural selection do this by acting on a useless structure directly, it may do so by acting on the useless structure indirectly, through its direct action on some other part of the organism with which the useless structure happens to be correlated. Moreover, as I believe, and will subsequently endeavour to prove, there is abundant evidence to show that incipient characters are often developed to a large extent by causes other than natural selection (or apart from any reference to utility), with the result that some of them thus happen to become of use, when, of course, the supposed difficulty is at an end.

But although it is thus easy to dispose of both the propositions in question, on account of their universality, stated more carefully they would require, as I have said, more careful consideration. Thus, if it had been said that some incipient organs are presumably useless at the time of their inception, and that in some of these cases it is difficult, or impossible, to conceive how the principle of correlation, or any other principle hitherto suggested, can apply—then the question would have been raised from the sphere of logical discussion to that of biological fact. And the new question thus raised would have to be debated, no longer on the ground of general or abstract principles, but on that of special or concrete cases. Now until within the last year or two it has not been easy to find such a special or concrete case—that is to say, a case which can be pointed to as apparently excluding the possibility of natural selection having had anything to do with the genesis of an unquestionably adaptive structure. But eventually such a case has arisen, and the Duke of Argyll has not been slow in perceiving its importance. This case is the electric organ in the tail of the skate. No sooner had Professor Cossar Ewart published an abstract of his first paper on this subject, than the Duke seized upon it as a case for which, as he said, he had long been waiting—namely, the case of an adaptive organ the genesis of which could not possibly be attributed to natural selection, and must therefore be attributed to supernatural design. Now, I do not deny that he is here in possession of an admirable case—a case, indeed, so admirable that it almost seems to have been specially designed for the discomfiture of Darwinians. Therefore, in order to do it full justice, I will show that it is even more formidable than the Duke of Argyll has represented.

Electric organs are known to occur in several widely different kinds of fish—such as the Gymnotus and Torpedo. Wherever these organs do occur, they perform the function of electric batteries in storing and discharging electricity in the form of more or less powerful shocks. Here, then, we have a function which is of obvious use to the fish for purposes both of offence and defence. These organs are everywhere composed of a transformation of muscular, together with an enormous development of nervous tissue; but inasmuch as they occupy different positions, and are also in other respects dissimilar in the different zoological groups of fishes where they occur, no difficulty can be alleged as to these analogous organs being likewise homologous in different divisions of the aquatic vertebrata.

Now, in the particular case of the skate, the organ is situated in the tail, where it is of a spindle-like form, measuring, in a large fish, about two feet in length by about an inch in diameter at the middle of the spindle. Although its structure is throughout as complex and perfect as that of the electric organ in Gymnotus or Torpedo, its smaller size does not admit of its generating a sufficient amount of electricity to yield a discharge that can be felt by the hand. Nevertheless, that it does discharge under suitable stimulation has been proved by Professor Burdon Sanderson by means of a telephone; for he found that every time he stimulated the animal its electrical discharge was rendered audible by the telephone. Here, then, the difficulty arises. For of what conceivable use is such an organ to its possessor? We can scarcely suppose that any aquatic animal is more sensitive to electric shocks than is the human hand; and even if such were the case, a discharge of so feeble a kind taking place in water would be short-circuited in the immediate vicinity of the skate itself. So there can be no doubt that such weak discharges as the skate is able to deliver must be wholly imperceptible alike to prey and to enemies. Yet for the delivery of such discharges there is provided an organ of such high peculiarity and huge complexity, that, regarded as a piece of living mechanism, it deserves to rank as at once the most extremely specialized and the most highly elaborated structure in the whole animal kingdom. Thousands of separately formed elements are ranged in row after row, all electrically insulated one from another, and packed away into the smallest possible space, with the obvious end, or purpose, of conspiring together for the simultaneous delivery of an electric shock. Nevertheless, the shock when delivered is, as we have just seen, too slight to be of any conceivable use to the skate. Therefore it appears impossible to suggest how this astonishing structure—much more astonishing, in my opinion, than the human eye or the human hand—can ever have been begun, or afterwards developed, by means of natural selection. For if it be not even yet of any conceivable use to its possessor, clearly thus far survival of the fittest can have had nothing to do with its formation. On the other hand, seeing that electric organs when of larger size, as in the Gymnotus and Torpedo, are of obvious use to their possessors, the facts of the case, so far as the skate is concerned, assuredly do appear to sanction the doctrine of "prophetic germs." The organ in the skate seems to be on its way towards becoming such an organ as we meet with in these other animals; and, therefore, unless we can show that it is now, and in all previous stages of its evolution has throughout been, of use to the skate, the facts do present a serious difficulty to the theory of natural selection, while they readily lend themselves to the interpretation of a disposing or fore-ordaining mind, which knows how to construct an electric battery by thus transforming muscular tissue into electric tissue, and is now actually in process of constructing such an apparatus for the prospective benefit of future creatures.

Should it be suggested that possibly the electric organ of the skate may be in process of degeneration, and therefore that it is now the practically functionless remnant of an organ which in the ancestors of the skate was of larger size and functional use—against so obvious a suggestion there lie the whole results of Professor Ewart's investigations, which go to indicate that the organ is here not in a stage of degeneration, but of evolution. For instance, in Raia radiata, it does not begin to be formed out of the muscular tissue until some time after the animal has left the egg-capsule, and assumed all the normal proportions (though not yet the size) of the adult creature. The organ, therefore, is one of the very latest to appear in the ontogeny of R. radiata; and, moreover, it does not attain its full development (i. e. not merely growth, but transforming of muscular fibres into electrical elements) till the fish attains maturity. Read in the light of embryology, these facts prove, (1) that the electric organ of R. radiata must be one of the very latest products of the animal's phylogeny; and, (2) that as yet, at all events, it has not begun to degenerate. But, if not, it must either be at a stand-still, or it must be in course of further evolution; and, whichever of these alternatives we adopt, the difficulty of accounting for its present condition remains. In this connexion also it is worth while to remark that the electric organ, even after it has attained its full development, continues its growth with the growth of the fish, and this in a much higher ratio, either than the tail alone, or the whole animal. Lastly, Prof. Burdon Sanderson finds that section for section the organ in the skate is as efficient as it is in Torpedo. It is evident that these facts also point to the skate's organ being in course of phylogenetic evolution.

.

[Illustration: Fig. 119.—Electric organ of the skate. The left-hand drawing (i) represents the entire organ (natural size) of a full-grown r. radiata. This is a small skate, which rarely exceeds 50 centms. in length; but in the large r. batis, the organ may exceed two feet in length. The other drawings represent single muscle-fibres in successive stages of transition. In the first of the series (ii) the motor plate, and the nerves connected with it, have already been considerably enlarged. In the other three specimens, the fibre becomes more and more club-like, and eventually cup-like. These changes of shape are expressive of great changes of structure, as may be seen in the last of the series (v), where the shallow cup is seen in partial section. The electric plate lines the concavity of the cup, and is richly supplied with nerves (only a few of which are represented in the last drawing); the thick walls of the cup are composed of muscular fibres, the striation of which is distinctly visible.]



Again, it cannot be answered that the principle of correlation may be drawn upon in mitigation of the difficulty. The structure of the electric organ is far too elaborate, far too specialized, and far too obviously directed to a particular end, to admit of our conceivably supposing it due to any accidental correlation with structural changes going on elsewhere. Even as regards the initial changes of muscle-elements into electrical-elements, I do not think the principle of correlation can be reasonably adduced by way of explanation; for, as shown in the illustrations, even this initial change is most extraordinarily peculiar, elaborate, and specialized. But, be this as it may, I am perfectly certain that the principle of correlation cannot possibly be adduced to explain the subsequent association of these electrical elements into an electric battery, actuated by a special nervous mechanism of enormous size and elaboration—unless of course, the progress of such a structure were assumed to have been throughout of some utility. Under this supposition, however, the principle of correlation would be forsaken in favour of that of natural selection; and we should again be in the presence of the same difficulty as that with which we started.

But now, and further, if we do thus abandon correlation in favour of natural selection, and therefore if for the sake of saving an hypothesis we assume that the organ as it now stands must be of some use to the existing skate, we should still have to face the question—Of what conceivable use can those initial stages of its formation have been, when first the muscle-elements began to be changed into the very different electrical-elements, and when therefore they became useless as muscles while not yet capable of performing even so much of the electrical function as they now perform?

Lastly, we must remember that not only have we here the most highly specialized, the most complex, and altogether the most elaboratively adaptive organ in the animal kingdom; but also that in the formation of this structure there has been needed an altogether unparalleled expenditure of the most physiologically expensive of all materials—namely, nervous tissue. Whether estimated by volume or by weight, the quantity of nervous tissue which is consumed in the electric organ of the skate is in excess of all the rest of the nervous system put together. It is needless to say that nowhere else in the animal kingdom—except, of course, in other electric fishes—is there any approach to so enormous a development of nervous tissue for the discharge of a special function. Therefore, as nervous tissue is, physiologically speaking, the most valuable of all materials, we are forced to conclude that natural selection ought strongly to have opposed the evolution of such organs, unless from the first moment of their inception, and throughout the whole course of their development, they were of some such paramount importance as biologically to justify so unexampled an expenditure. Yet this paramount importance does not admit of being so much as surmised, even where the organ has already attained the size and degree of elaboration which it presents in the skate.

In view of all these considerations taken together, I freely confess that the difficulty presented by this case appears to me of a magnitude and importance altogether unequalled by that of any other single case—or any series of cases—which has hitherto been encountered by the theory of natural selection. So that, if there were many other cases of the like kind to be met with in nature, I should myself at once allow that the theory of natural selection would have to be discarded. But inasmuch as this particular case stands so far entirely by itself, and therefore out of analogy with thousands, or even millions, of other cases throughout the whole range of organic nature, I am constrained to feel it more probable that the electric organ of the skate will some day admit of being marshalled under the general law of natural selection—in just the same way as proved to be the case with the conspicuous colouring of those caterpillars, which, as explained in the last chapter, at one time seemed to constitute a serious difficulty to the theory, and yet, through a better knowledge of all the relations involved, has now come to constitute one of the strongest witnesses in its favour.

* * * * *

I have now stated all the objections of any importance which have hitherto been brought against the theory of natural selection, excepting three, which I left to be dealt with together because they form a logically connected group. With a brief consideration of these, therefore, I will bring this chapter to a close.

The three objections to which I allude are, (1) that a large proportional number of specific, as well as of higher taxonomic characters, are seemingly useless characters, and therefore do not lend themselves to explanation by the Darwinian theory; (2) that the most general of all specific characters—viz. cross-infertility between allied species—cannot possibly be due to natural selection, as is demonstrated by Darwin himself; (3) that the swamping effects of free intercrossing must always render impossible by natural selection alone any evolution of species in divergent (as distinguished from serial) lines of change.

These three objections have been urged from time to time by not a few of the most eminent botanists and zoologists of our century; and from one point of view I cannot myself have the smallest doubt that the objections thus advanced are not only valid in themselves, but also by far the most formidable objections which the theory of natural selection has encountered. From another point of view, however, I am equally convinced that they all admit of absolute annihilation. This strong antithesis arises, as I have said, from differences of standpoint, or from differences in the view which we take of the theory of natural selection itself. If we understand this theory to set forth natural selection as the sole cause of organic evolution, then all the above objections to the theory are not merely, as already stated, valid and formidable, but as I will now add, logically insurmountable. On the other hand, if we take theory to consist merely in setting forth natural selection as a factor of organic evolution, even although we believe it to have been the chief factor or principal cause, all the three objections in question necessarily vanish. For in this case, even if it be satisfactorily proved that the theory of natural selection is unable to explain the three classes of facts above mentioned, the theory is not thereby affected: facts of each and all of these classes may be consistently left by the theory to be explained by causes other than natural selection—whether these be so far capable or incapable of hypothetical formulation. Thus it is evident that whether the three objections above named are to be regarded as logically insurmountable by the theory, or as logically non-existent in respect to it, depends simply upon the manner in which the theory itself is stated.

In the next volume a great deal more will have to be said upon these matters—especially with regard to the causes other than natural selection which in my opinion are capable of explaining these so-called "difficulties." In the present connexion, however, all I have attempted to show is, that, whatever may be thought touching the supplementary theories whereby I shall endeavour to explain the facts of inutility, cross-sterility, and non-occurrence of free intercrossing, no one of these facts is entitled to rank as an objection against the theory of natural selection, unless we understand this theory to claim an exclusive prerogative in the field of organic evolution. This, as we have previously seen, is what Mr. Wallace does claim for it; while on the other hand, Mr. Darwin expressly—and even vehemently—repudiates the claim: from which it follows that all the three main objections against the theory of natural selection are objections which vitally affect the theory only as it has been stated and upheld by Wallace. As the theory has been stated and upheld by Darwin, all these objections are irrelevant. This is a fact which I had not myself perceived at the time when I mentioned these objections in a paper entitled Physiological Selection, which was published in 1886. The discussions to which that paper gave rise, however, led me to consider these matters more closely; and further study of Darwin's writings, with these matters specially in view, has led me to see that none of the objections in question are relevant to his theory, as distinguished from that of Mr. Wallace. This, I acknowledge, I ought to have perceived before I published the paper just alluded to; but in those days I had had no occasion to follow out the differences between Darwin and Wallace to all their consequences, and therefore adopted the prevalent view that their theories of evolution were virtually identical. Now, however, I have endeavoured to make it clear that the points wherein they differ involve the important consequences above set forth. All these the most formidable objections against the theory of natural selection arise simply and solely from what I conceive to be the erroneous manner in which the theory has been presented by Darwin's distinguished colleague.

* * * * *

I have now considered, as impartially as I can, all the main criticisms and objections which have been brought against the theory of natural selection; and the result is to show that, neither singly nor collectively, are they entitled to much weight. On the other hand, as we have seen in the preceding chapter, there is a vast accumulation of evidence in favour of the theory. Hence, it is no wonder that the theory has now been accepted by all naturalists, with scarcely any one notable exception, as at any rate the best working hypothesis which has ever been propounded whereby to explain the facts of organic evolution. Moreover, in the opinion of those most competent to judge, the theory is entitled to be regarded as something very much more than a working hypothesis: it is held to be virtually a completed induction, or, in other words, the proved exhibition of a general law, whereby the causation of organic evolution admits of being in large part—if not altogether—explained.

Now, whether or not we subscribe to this latter conclusion ought, I think, to depend upon what we mean by an explanation in the case which is before us. If we mean only that, given the large class of known facts and unknown causes which are conveniently summarized under the terms Heredity and Variability, then the further facts of Struggle and Survival serve, in some considerable degree or another, to account for the phenomena of adaptive evolution, I cannot see any room to question that the evidence is sufficient to prove the statement. But it is clear that by taking for granted these great facts of Heredity and Variability, we have assumed the larger part of the problem as a whole. Or, more correctly, by thus generalizing, in a merely verbal form, all the unknown causes which are concerned in these two great factors of the process in question, we are not so much as attempting to explain the precedent causation which serves as a condition to the process. Much more than half the battle would already have been won, had Darwin's predecessors been able to explain the causes of Heredity and Variation; hence it is but a very partial victory which we have hitherto gained in our recent discovery of the effects of Struggle and Survival.

Yet partial though it be in relation to the whole battle, in itself, or considered absolutely, there can be no reasonable doubt that it constitutes the greatest single victory which has ever been gained by the science of Biology. For this very reason, however, it behoves us to consider all the more carefully the extent to which it goes. But my discussion of this matter must be relegated to the next volume, where I hope to give abundant proof of the soundness of Darwin's judgment as conveyed in the words:—"I am convinced that natural selection has been the main, but not the exclusive, means of modification."



CHAPTER X.

THE THEORY OF SEXUAL SELECTION, AND CONCLUDING REMARKS.

Although the explanatory value of the Darwinian theory of natural selection is, as we have now seen, incalculably great, it nevertheless does not meet those phenomena of organic nature which perhaps more than any other attract the general attention, as well as the general admiration, of mankind: I mean all that class of phenomena which go to constitute the Beautiful. Whatever value beauty as such may have, it clearly has not a life-preserving value. The gorgeous plumage of a peacock, for instance, is of no advantage to the peacock in his struggle for life, and therefore cannot be attributed to the agency of natural selection. Now this fact of beauty in organic structures is a fact of wide generality—almost as wide, indeed, as is the fact of their utility. Mr. Darwin, therefore, suggested another hypothesis whereby to render a scientific explanation of this fact. Just as by his theory of natural selection he sought to explain the major fact of utility, so did he endeavour to explain the minor fact of beauty by a theory of what he termed Sexual Selection.

It is a matter of observation that the higher animals do not pair indiscriminately; but that the members of either sex prefer those individuals of the opposite sex which are to them most attractive. It is important to understand in limine that nobody has ever attempted to challenge this statement. In other words, it is an unquestionable fact that among many of the higher animals there literally and habitually occurs a sexual selection; and this fact is not a matter of inference, but, as I have said, a matter of observation. The inference only begins where, from this observable fact, it is argued,—1st, that the sexual selection has reference to an aesthetic taste on the part of the animals themselves; and 2nd, that, supposing the selection to be determined by such a taste, the cause thus given is adequate to explain the phenomena of beauty which are presented by these animals. I will consider these two points separately.

From the evidence which Darwin has collected, it appears to me impossible to doubt that an aesthetic sense is displayed by many birds, and not a few mammals. This of course does not necessarily imply that the standards of such a sense are the same as our own; nor does it necessarily imply that there is any constant relation between such a sense and high levels of intelligence in other respects. In point of fact, such is certainly not the case, because the best evidence that we have of an aesthetic sense in animals is derived from birds, and not from mammals. The most cogent cases to quote in this connexion are those of the numerous species of birds which habitually adorn their nests with gaily coloured feathers, wool, cotton, or any other gaudy materials which they may find lying about the woods and fields. In many cases a marked preference is shown for particular objects—as, for instance, in the case of the Syrian nut-hatch, which chooses the iridescent wings of insects, or that of the great crested fly-catcher, which similarly chooses the cast-off skins of snakes. But no doubt the most remarkable of these cases is that of the baya-bird of Asia, which after having completed its bottle-shaped and chambered nest[47], studs it over with small lumps of clay, both inside and out, upon which the cock-bird sticks fire-flies, apparently for the sole purpose of securing a brilliantly decorative effect. Other birds, such as the hammer-head of Africa, adorn the surroundings of their nests (which are built upon the ground) with shells, bones, pieces of broken glass and earthenware, or any objects of a bright and conspicuous character which they may happen to find. The most consummate artists in this respect are, however, the bower-birds; for the species of this family construct elaborate play-houses in the form of arched tunnels, built of twigs upon the ground. Through and around such a tunnel they chase one another; and it is always observable that not only is the floor paved with a great collection of shells, bones, coloured stones, and any other brilliant objects which they are able to carry in their beaks, but also that the walls are decorated with the most gaudy articles which the birds can find. There is one genus, in Papua, which even goes so far as to provide the theatre with a surrounding garden. A level piece of ground is selected as a site for the building. The latter is about two feet high, and constructed round the growing stalk of a shrub, which therefore serves as a central pillar to which the frame-work of the roof is attached. Twigs are woven into this frame-work until the whole is rendered rain-proof. The tent thus erected is about nine feet in circumference at its base, and presents a large arch as an entrance. The central pillar is banked up with moss at its base, and a gallery is built round the interior of the edifice. This gallery is decorated with flowers, fruits, fungi, &c. These are also spread over the garden, which covers about the same area as the play-house. The flowers are said to be removed when they fade, while fresh ones are gathered to supply their places. Thus the garden is always kept bright with flowers, as well as with the brilliant green of mosses, which are collected and distributed in patches, resembling tiny lawns.

[47] The chambers are three in number. The two upper ones are occupied respectively by the male and the sitting female. The lower one serves as a general living room when the young are hatched.



Now these sundry cases alone seem to prove a high degree of the aesthetic sense as occurring among birds; for, it is needless to say, none of the facts just mentioned can be due to natural selection, seeing that they have no reference to utility, or the preservation of life. But if an aesthetic sense occurs in birds, we should expect, on a priori grounds, that it would probably be exercised with reference to the personal appearance of the sexes. And this expectation is fully realized. For it is an observable fact that in most species of birds where the males are remarkable for the brilliancy of their plumage, not only is this brilliancy most remarkable during the pairing season, but at this season also the male birds take elaborate pains to display their charms before the females. Then it is that the peacock erects his tail to strut round and round the hens, taking care always to present to them a front view, where the coloration is most gorgeous. And the same is true of all other gaily coloured male birds. During the pairing season they actively compete with one another in exhibiting their attractiveness to the females; and in many cases there are added all sorts of extraordinary antics in the way of dancings and crowings. Again, in the case of all song-birds, the object of the singing is to please the females; and for this purpose the males rival one another to the best of their musical ability.

Thus there can be no question that the courtship of birds is a highly elaborate business, in which the males do their best to surpass one another in charming the females. Obviously the inference is that the males do not take all this trouble for nothing; but that the females give their consent to pair with the males whose personal appearance, or whose voice, proves to be the most attractive. But, if so, the young of the male bird who is thus selected will inherit his superior beauty; and thus, in successive generations, a continuous advance will be made in the beauty of plumage or of song, as the case may be,—both the origin and development of beauty in the animal world being thus supposed due to the aesthetic taste of animals themselves.

Such is the theory of sexual selection in its main outlines; and with regard to it we must begin by noting two things which are of most importance. In the first place, it is a theory wholly and completely distinct from the theory of natural selection; so that any truth or error in the one does not in the least affect the other. The second point is, that there is not so great a wealth of evidence in favour of sexual selection as there is in favour of natural selection; and, therefore, that while all naturalists nowadays accept natural selection as a (whether or not the) cause of adaptive, useful, or life-preserving structures, there is no such universal—but only a very general—agreement with reference to sexual selection as a cause of decorative, beautiful, or life-embellishing structures. Nevertheless, the evidence in favour of sexual selection is both large in amount and massive in weight.

Our consideration of this evidence will bring us to the second division of our subject, as previously marked out for discussion—namely, granting that an aesthetic sense occurs in certain large divisions of the animal kingdom, what is the proof that such a sense is a cause of the beauty which is presented by the animals in question?

Before proceeding to state this proof, however, it is desirable to observe that under the theory of sexual selection Darwin has included two essentially different classes of facts. For besides the large class of facts to which I have thus far been alluding,—i. e. the cases where two sexes of the same species differ from one another in respect of ornamentation,—there is another class of facts equally important, namely, the cases where the two sexes of the same species differ from one another in respect of size, strength, and the possession of natural weapons, such as spurs, horns, &c. In most of these cases it is the males which are thus superiorly endowed; and it is a matter of observation that in all cases where they are so endowed they use their superior strength and natural weapons for fighting together, in order to secure possession of the females. Hence results what Mr. Darwin has called the Law of Battle between males of the same species; and this law of battle he includes under his theory of sexual selection. But it is evident that the principle which is operative in the law of battle differs from the principle which is concerned in the form of sexual selection that has to do with embellishment, and consequent charm. The law of battle, in fact, more nearly approaches the law of natural selection; seeing that it expresses the natural advantages of brute force in the struggling of rival animals, and so frequently results in death of the less fitted, as distinguished from a mere failure to propagate. Now against this doctrine of the law of battle, and the consequences to which it leads in the superior fighting powers of male animals, no objection has been raised in any quarter. It is only with regard to the other aspect of the theory of sexual selection—or that which is concerned with the superior embellishment of male animals—that any difference of opinion obtains. I will now proceed to give the main arguments on both sides of this question, beginning with a resume of the evidences in favour of sexual selection.

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