Chapter 18

The problem which was to be Darwin's chief subject of occupation for the rest of his life thus presented itself, at first, mainly under its distributional aspect. Why do species present certain relations in space and in time? Why are the animals and plants of the Galapagos Archipelago so like those of South America and yet different from them? Why are those of the several islets more or less different from one another? Why are the animals of the latest geological epoch in South America similar in _facies_ to those which exist in the same region at the present day, and yet specifically or generically different?

The reply to these questions, which was almost universally received fifty years ago, was that animals and plants were created such as they are; and that their present distribution, at any rate so far as terrestrial organisms are concerned, has been effected by the migration of their ancestors from the region in which the ark stranded after the subsidence of the deluge. It is true that the geologists had drawn attention to a good many tolerably serious difficulties in the way of the diluvial part of this hypothesis, no less than to the supposition that the work of creation had occupied only a brief space of time. But even those, such as Lyell, who most strenuously argued in favour of the sufficiency of natural causes for the production of the phenomena of the inorganic world, held stoutly by the hypothesis of creation in the case of those of the world of life.

For persons who were unable to feel satisfied with the fashionable doctrine, there remained only two alternatives--the hypothesis of spontaneous generation, and that of descent with modification. The former was simply the creative hypothesis with the creator left out; the latter had already been propounded by De Maillet and Erasmus Darwin, among others; and, later, systematically expounded by Lamarck. But in the eyes of the naturalist of the "Beagle" (and, probably, in those of most sober thinkers), the advocates of transmutation had done the doctrine they expounded more harm than good.

Darwin's opinion of the scientific value of the "Zoonomia" has already been mentioned. His verdict on Lamarck is given in the following passage of a letter to Lyell (March, 1863):--

"Lastly, you refer repeatedly to my view as a modification of Lamarck's doctrine of development and progression. If this is your deliberate opinion there is nothing to be said, but it does not seem so to me. Plato, Buffon, my grandfather, before Lamarck and others, propounded the _obvious_ view that if species were not created separately they must have descended from other species, and I can see nothing else in common between the "Origin" and Lamarck. I believe this way of putting the case is very injurious to its acceptance, as it implies necessary progression, and closely connects Wallace's and my views with what I consider, after two deliberate readings, as a wretched book, and one from which (I well remember to my surprise) I gained nothing."

"But," adds Darwin with a little touch of banter, "I know you rank it higher, which is curious, as it did not in the least shake your belief." (III. p. 14; see also p. 16, "to me it was an absolutely useless book.")

Unable to find any satisfactory theory of the process of descent with modification in the works of his predecessors, Darwin proceeded to lay the foundations of his own views independently; and he naturally turned, in the first place, to the only certainly known examples of descent with modification, namely, those which are presented by domestic animals and cultivated plants. He devoted himself to the study of these cases with a thoroughness to which none of his predecessors even remotely approximated; and he very soon had his reward in the discovery "that selection was the keystone of man's success in making useful races of animals and plants." (I. p. 83.)

This was the first step in Darwin's progress, though its immediate result was to bring him face to face with a great difficulty. "But how selection could be applied to organisms living in a state of nature remained for some time a mystery to me." (I. p. 83.)

The key to this mystery was furnished by the accidental perusal of the famous essay of Malthus "On Population" in the autumn of 1838. The necessary result of unrestricted multiplication is competition for the means of existence. The success of one competitor involves the failure of the rest, that is, their extinction; and this "selection" is dependent on the better adaptation of the successful competitor to the conditions of the competition. Variation occurs under natural, no less than under artificial, conditions. Unrestricted multiplication implies the competition of varieties and the selection of those which are relatively best adapted to the conditions.

Neither Erasmus Darwin, nor Lamarck, had any inkling of the possibility of this process of "natural selection"; and though it had been foreshadowed by Wells in 1813, and more fully stated by Matthew in 1831, the speculations of the latter writer remained unknown to naturalists until after the publication of the "Origin of Species."

Darwin found in the doctrine of the selection of favourable variations by natural causes, which thus presented itself to his mind, not merely a probable theory of the origin of the diverse species of living forms, but that explanation of the phenomena of adaptation, which previous speculations had utterly failed to give. The process of natural selection is, in fact, dependent on adaptation--it is all one, whether one says that the competitor which survives is the "fittest" or the "best adapted." And it was a perfectly fair deduction that even the most complicated adaptations might result from the summation of a long series of simple favourable variations.

Darwin notes as a serious defect in the first sketch of his theory that he had omitted to consider one very important problem, the solution of which did not occur to him till some time afterwards. "This problem is the tendency in organic beings descended from the same stock to diverge in character as they become modified.... The solution, as I believe, is that the modified offspring of all dominant and increasing forms tend to become adapted to many and highly diversified places in the economy of nature." (I. p. 84.)

It is curious that so much importance should be attached to this supplementary idea. It seems obvious that the theory of the origin of species by natural selection necessarily involves the divergence of the forms selected. An individual which varies, _ipso facto_ diverges from the type of its species; and its progeny, in which the variation becomes intensified by selection, must diverge still more, not only from the parent stock, but from any other race of that stock starting from, a variation of a different character. The selective process could not take place unless the selected variety was either better adapted to the conditions than the original stock, or adapted to other conditions than the original stock. In the first case, the original stock would be sooner or later extirpated; in the second, the type, as represented by the original stock and the variety, would occupy more diversified stations than it did before.

The theory, essentially such as it was published fourteen years later, was written out in 1844, and Darwin was so fully convinced of the importance of his work, as it then stood, that he made special arrangements for its publication in case of his death. But it is a singular example of reticent fortitude, that, although for the next fourteen years the subject never left his mind, and during the latter half of that period he was constantly engaged in amassing facts bearing upon it from wide reading, a colossal correspondence, and a long series of experiments, only two or three friends were cognisant of his views. To the outside world he seemed to have his hands quite sufficiently full of other matters. In 1844, he published his observations on the volcanic islands visited during the voyage of the "Beagle." In 1845, a largely remodelled edition of his "Journal" made its appearance, and immediately won, as it has ever since held, the favour of both the scientific and the unscientific public. In 1846, the "Geological Observations in South America" came out, and this book was no sooner finished than Darwin set to work upon the Cirripedes. He was led to undertake this long and heavy task, partly by his desire to make out the relations of a very anomalous form which he had discovered on the coast of Chili; and partly by a sense of "presumption in accumulating facts and speculating on the subject of variation without having worked out my due share of species." (II. p. 31.) The eight or nine years of labour, which resulted in a monograph of first-rate importance in systematic zoology (to say nothing of such novel points as the discovery of complemental males), left Darwin no room to reproach himself on this score, and few will share his "doubt whether the work was worth the consumption of so much time." (I. p. 82.)

In science no man can safely speculate about the nature and relation of things with which he is unacquainted at first hand, and the acquirement of an intimate and practical knowledge of the process of species-making and of all the uncertainties which underlie the boundaries between species and varieties, drawn by even the most careful and conscientious systematists [Footnote: "After describing a set of forms as distinct species, tearing up my MS., and making them one species, tearing that up and making them separate, and then making them one again (which has happened to me), I have gnashed my teeth, cursed species, and asked what sin I had committed to be so punished." (II. p. 40.) Is there any naturalist provided with a logical sense and a large suite of specimens, who has not undergone pangs of the sort described in this vigorous paragraph, which might, with advantage, be printed on the title-page of every systematic monograph as a warning to the uninitiated?] were of no less importance to the author of the "Origin of Species" than was the bearing of the Cirripede work upon "the principles of a natural classification." (I. p. 81.) No one, as Darwin justly observes, has a "right to examine the question of species who has not minutely described many." (II. p. 39.)

In September, 1854, the Cirripede work was finished, "ten thousand barnacles" had been sent "out of the house, all over the world," and Darwin had the satisfaction of being free to turn again to his "old notes on species." In 1855, he began to breed pigeons, and to make observations on the effects of use and disuse, experiments on seeds, and so on, while resuming his industrious collection of facts, with a view "to see how far they favour or are opposed to the notion that wild species are mutable or immutable. I mean with my utmost power to give all arguments and facts on both sides. I have a _number_ of people helping me every way, and giving me most valuable assistance; but I often doubt whether the subject will not quite overpower me." (II. p. 49.)

Early in 1856, on Lyell's advice, Darwin began to write out his views on the origin of species on a scale three or four times as extensive as that of the work published in 1859. In July of the same year he gave a brief sketch of his theory in a letter to Asa Gray; and, in the year 1857, his letters to his correspondents show him to be busily engaged on what he calls his "big book." (II. pp. 85, 94.) In May, 1857, Darwin writes to Wallace: "I am now preparing my work [on the question how and in what way do species and varieties differ from each other] for publication, but I find the subject so very large, that, though I have written many chapters, I do not suppose I shall go to press for two years." (II. p. 95.) In December, 1857, he writes, in the course of a long letter to the same correspondent, "I am extremely glad to hear that you are attending to distribution in accordance with theoretical ideas. I am a firm believer that without speculation there is no good and original observation." (II. p. 108.) [Footnote: The last remark contains a pregnant truth, but it must be confessed it hardly squares with the declaration in the _Autobiography_, (I. p. 83), that he worked on "true Baconian principles."] In June, 1858, he received from Mr. Wallace, then in the Malay Archipelago, an "Essay on the tendency of varieties to depart indefinitely from the original type," of which Darwin says, "If Wallace had my MS. sketch written out in 1842 he could not have made a better short abstract! Even his terms stand now as heads of my chapters. Please return me the MS., which he does not say he wishes me to publish, but I shall, of course, at once write and offer to send it to any journal. So all my originality, whatever it may amount to, will be smashed, though my book, if ever it will have any value, will not be deteriorated; as all the labour consists in the application of the theory." (II. p. 116.)

Thus, Darwin's first impulse was to publish Wallace's essay without note or comment of his own. But, on consultation with Lyell and Hooker, the latter of whom had read the sketch of 1844, they suggested, as an undoubtedly more equitable course, that extracts from the MS. of 1844 and from the letter to Dr. Asa Gray should be communicated to the Linnean Society along with Wallace's essay. The joint communication was read on July 1, 1858, and published under the title "On the Tendency of Species to form Varieties; and on the Perpetuation of Varieties and Species by Natural Means of Selection." This was followed, on Darwin's part, by the composition of a summary account of the conclusions to which his twenty years' work on the species question had led him. It occupied him for thirteen months, and appeared in November, 1859, under the title "On the Origin of Species by means of Natural Selection or the Preservation of Favoured Races in the Struggle of Life."

It is doubtful if any single book, except the "Principia," ever worked so great and so rapid a revolution in science, or made so deep an impression on the general mind. It aroused a tempest of opposition and met with equally vehement support, and it must be added that no book has been more widely and persistently misunderstood by both friends and foes. In 1861, Darwin remarks to a correspondent, "You understand my book perfectly, and that I find a very rare event with my critics." (I. p. 313.) The immense popularity which the "Origin" at once acquired was no doubt largely due to its many points of contact with philosophical and theological questions in which every intelligent man feels a profound interest; but a good deal must be assigned to a somewhat delusive simplicity of style, which tends to disguise the complexity and difficulty of the subject, and much to the wealth of information on all sorts of curious problems of natural history, which is made accessible to the most unlearned reader. But long occupation with the work has led the present writer to believe that the "Origin of Species" is one of the hardest of books to master; [Footnote: He is comforted to find that probably the best qualified judge among all the readers of the _Origin_ in 1859 was of the same opinion. Sir J. Hooker writes, "It is the very hardest book to read, to full profit, that I ever tried." (II. p. 242.)] and he is justified in this conviction by observing that although the "Origin" has been close on thirty years before the world, the strangest misconceptions of the essential nature of the theory therein advocated are still put forth by serious writers.

Although, then, the present occasion is not suitable for any detailed criticism of the theory, or of the objections which have been brought against it, it may not be out of place to endeavour to separate the substance of the theory from its accidents; and to show that a variety not only of hostile comments, but of friendly would-be improvements lose their _raison d'être_ to the careful student. Observation proves the existence among all living beings of phenomena of three kinds, denoted by the terms heredity, variation, and multiplication. Progeny tend to resemble their parents; nevertheless all their organs and functions are susceptible of departing more or less from the average parental character; and their number is in excess of that of their parents. Severe competition for the means of living, or the struggle for existence, is a necessary consequence of unlimited multiplication; while selection, or the preservation of favourable variations and the extinction of others, is a necessary consequence of severe competition. "Favourable variations" are those which are better adapted to surrounding conditions. It follows, therefore, that every variety which is selected into a species is so favoured and preserved in consequence of being, in some one or more respects, better adapted to its surroundings than its rivals. In other words, every species which exists, exists in virtue of adaptation, and whatever accounts for that adaptation accounts for the existence of the species.

To say that Darwin has put forward a theory of the adaptation of species, but not of their origin, is therefore to misunderstand the first principles of the theory. For, as has been pointed out, it is a necessary consequence of the theory of selection that every species must have some one or more structural or functional peculiarities, in virtue of the advantage conferred by which, it has fought through the crowd of its competitors and achieved a certain duration. In this sense, it is true that every species has been "originated" by selection.

There is another sense, however, in which it is equally true that selection originates nothing. "Unless profitable variations ... occur natural selection can do nothing" ("Origin," Ed. I. p. 82). "Nothing can be effected unless favourable variations occur" (_ibid_., p. 108). "What applies to one animal will apply throughout time to all animals--that is, if they vary--for otherwise natural selection can do nothing. So it will be with plants" (_ibid_., p. 113). Strictly speaking, therefore, the origin of species in general lies in variation; while the origin of any particular species lies, firstly, in the occurrence, and secondly, in the selection and preservation of a particular variation. Clearness on this head will relieve one from the necessity of attending to the fallacious assertion that natural selection is a _deus ex machinâ_, or occult agency.

Those, again, who confuse the operation of the natural causes which bring about variation and selection with what they are pleased to call "chance" can hardly have read the opening paragraph of the fifth chapter of the "Origin" (Ed. I, p. 131): "I have sometimes spoken as if the variations ... had been due to chance. This is of course a wholly incorrect expression, but it seems to acknowledge plainly our ignorance of the cause of each particular variation."

Another point of great importance to the right comprehension of the theory, is, that while every species must needs have some adaptive advantageous characters to which it owes its preservation by selection, it may possess any number of others which are neither advantageous nor disadvantageous, but indifferent, or even slightly disadvantageous. (_Ibid_., p. 81.) For variations take place, not merely in one organ or function at a time, but in many; and thus an advantageous variation, which gives rise to the selection of a new race or species, may be accompanied by others which are indifferent, but which are just as strongly hereditary as the advantageous variations. The advantageous structure is but one product of a modified general constitution which may manifest itself by several other products; and the selective process carries the general constitution along with the advantageous special peculiarity. A given species of plant may owe its existence to the selective adaptation of its flowers to insect fertilisers; but the character of its leaves may be the result of variations of an indifferent character. It is the origin of variations of this kind to which Darwin refers in his frequent reference to what he calls "laws of correlation of growth" or "correlated variation."

These considerations lead us further to see the inappropriateness of the objections raised to Darwin's theory on the ground that natural selection does not account for the first commencements of useful organs. But it does not pretend to do so. The source of such commencements is necessarily to be sought in different variations, which remain unaffected by selection until they have taken such a form as to become utilisable in the struggle for existence.

It is not essential to Darwin's theory that anything more should be assumed than the facts of heredity, variation, and unlimited multiplication; and the validity of the deductive reasoning as to the effect of the last (that is, of the struggle for existence which it involves) upon the varieties resulting from the operation of the former. Nor is it essential that one should take up any particular position in regard to the mode of variation, whether, for example, it takes place _per saltum_ or gradually; whether it is definite in character or indefinite. Still less are those who accept the theory bound to any particular views as to the causes of heredity or of variation.

That Darwin held strong opinions on some or all of these points may be quite true; but, so far as the theory is concerned, they must be regarded as _obiter dicta_. With respect to the causes of variation, Darwin's opinions are, from first to last, put forward altogether tentatively. In the first edition of the "Origin," he attributes the strongest influence to changes in the conditions of life of parental organisms, which he appears to think act on the germ through the intermediation of the sexual organs. He points out, over and over again, that habit, use, disuse, and the direct influence of conditions have some effect, but he does not think it great, and he draws attention to the difficulty of distinguishing between effects of these agencies and those of selection. There is, however, one class of variations which he withdraws from the direct influence of selection, namely, the variations in the fertility of the sexual union of more or less closely allied forms. He regards less fertility, or more or less complete sterility, as "incidental to other acquired differences." (_Ibid_., p. 245.)