"There can be no doubt, from the facts given in this chapter, that extremely slight changes in the conditions of life sometimes, probably often, act in a definite manner on our domesticated productions; and, as the action of changed conditions in causing indefinite variability is accumulative, so it may be with their definite action. Hence considerable and definite modifications of structure probably follow from altered conditions acting during long series of generations[5]."
"How, again, can we explain the inherited effects of the use and disuse of particular organs? The domesticated duck flies less and walks more than the wild duck, and its limb bones have become diminished and increased in a corresponding manner in comparison with those of the wild duck. A horse is trained to certain paces, and the colt inherits similar consensual movements. The domesticated rabbit becomes tame from close confinement; the dog, intelligent from associating with man; the retriever is taught to fetch and carry; and these mental endowments and bodily powers are all inherited. Nothing in the whole circuit of physiology is more wonderful. How can the use or disuse of a particular limb or of the brain affect a small aggregate of reproductive cells, seated in a distant part of the body, in such a manner that the being developed from these cells inherits the characters of either one or both parents?... In the chapters devoted to inheritance, it was shown that a multitude of newly acquired characters, whether injurious or beneficial, whether of the lowest or highest vital importance, are often faithfully transmitted[6]."
"When discussing special cases, Mr. Mivart passes over the effects of the increased use and disuse of parts, which I have always maintained to be highly important, and have treated in my 'Variation under Domestication' at greater length than, as I believe, any other writer[7]."
So much for the matured opinion of Darwin touching the validity of the theory of use-inheritance. Turning now to his opinion on the question whether or not there are yet any further factors concerned in the process of organic evolution, I think it will be sufficient to quote a single passage from the Origin of Species. The first paragraph of the "Conclusion" is devoted to a résumé of his views upon this matter, and consists of the following most emphatic words.
"I have now recapitulated the facts and considerations which have thoroughly convinced me that species have been modified, during a long course of descent. This has been effected chiefly through the natural selection of numerous successive, slight, favourable variations; aided in an important manner by the inherited effects of the use and disuse of parts; and in an unimportant manner, that is in relation to adaptive structures, whether past or present, by the direct action of external conditions, and by variations which seem to us in our ignorance to arise spontaneously. It appears that I formerly underrated the frequency and value of these latter forms of variation, as leading to permanent modifications of structure independently of natural selection. But as my conclusions have lately been much misrepresented, and it has been stated that I attribute the modification of species exclusively to natural selection, I may be permitted to remark that in the first edition of this work, and subsequently, I placed in a most conspicuous position—namely, at the close of the Introduction—the following words: 'I am convinced that natural selection has been the main, but not the exclusive means of modification.' This has been of no avail. Great is the power of steady misrepresentation; but the history of science shows that fortunately this power does not long endure."
In the whole range of Darwin's writings there cannot be found a passage so strongly worded as this: it presents the only note of bitterness in all the thousands of pages which he has published. Therefore I do not think it is necessary to supply any further quotations for the purpose of proving the state of his opinion upon the point in question. But, be it carefully noted, from this great or radical difference of opinion between the joint originators of the theory of natural selection, all their other differences of opinion arise; and seeing that since the death of Darwin a large number of naturalists have gone over to the side of Wallace, it seems desirable here to state categorically what these other or sequent points of difference are. Without at present discussing them, therefore, I will merely set them out in a tabular form, in order that a clear perception may be gained of their logical connexion with this primary point of difference.
| The Theory of Natural Selection according to Darwin. | The theory of Natural Selection according to Wallace. |
| Natural Selection has been the main means of modification, not excepting the case of Man. | Natural Selection has been the sole means of modification, excepting in the case of Man. |
| (a) Therefore it is a question of evidence whether the Lamarckian factors have co-operated. | (a) Therefore it is antecedently impossible that the Lamarckian factors can have co-operated. |
| (b) Neither all species, nor, a fortiori, all specific characters, have been due to natural selection. | (b) Not only all species, but all specific characters, must necessarily have been due to natural selection. |
| (c) Thus the principle of Utility is not of universal application, even where species are concerned. | (c) Thus the principle of Utility must necessarily be of universal application, where species are concerned. |
| (d) Thus, also, the suggestion as to Sexual Selection, or any other supplementary cause of modification, may be entertained; and, as in the case of the Lamarckian factors, it is a question of evidence whether, or how far, they have co-operated. | (d) Thus, also, the suggestion as to Sexual Selection, or of any other supplementary cause of modification, must be ruled out; and, as in the case of the Lamarckian factors, their co-operation deemed impossible. |
| (e) No detriment arises to the theory of natural selection as a theory of the origin of species by entertaining the possibility, or the probability, of supplementary factors. | (e) The possibility—and, a fortiori the probability—of any supplementary factors cannot be entertained without serious detriment to the theory of natural selection, as a theory of the origin of species. |
| (f) Cross-sterility in species cannot possibly be due to natural selection. | (f) Cross-sterility in species is probably due to natural selection[8]. |
As it will be my endeavour in the ensuing chapters to consider the rights and the wrongs of these antithetical propositions, I may reserve further quotations from Darwin's works, which will show that the above is a correct epitome of his views as contrasted with those of Wallace and the Neo-Darwinian school of Weismann. But here, where the object is merely a statement of Darwin's theory touching the points in which it differs from those of Wallace and Weismann, it will be sufficient to set forth these points of difference in another and somewhat fuller form. So far then as we are at present concerned, the following are the matters of doctrine which have been clearly, emphatically, repeatedly, and uniformly expressed throughout the whole range of Darwin's writings.
1. That natural selection has been the main means of modification.
2. That, nevertheless, it has not been the only means; but has been supplemented or assisted by the co-operation of other causes.