I have now given the leading facts with respect to rudimentary organs. In reflecting on them, everyone must be struck with astonishment; for the same reasoning power which tells us that most parts and organs are exquisitely adapted for certain purposes, tells us with equal plainness that these rudimentary or atrophied organs are imperfect and useless. In works on natural history, rudimentary organs are generally said to have been created “for the sake of symmetry,” or in order “to complete the scheme of nature.” But this is not an explanation, merely a restatement of the fact. Nor is it consistent with itself: thus the boa-constrictor has rudiments of hind limbs and of a pelvis, and if it be said that these bones have been retained “to complete the scheme of nature,” why, as Professor Weismann asks, have they not been retained by other snakes, which do not possess even a vestige of these same bones? What would be thought of an astronomer who maintained that the satellites revolve in elliptic courses round their planets “for the sake of symmetry,” because the planets thus revolve round the sun? An eminent physiologist accounts for the presence of rudimentary organs, by supposing that they serve to excrete matter in excess, or matter injurious to the system; but can we suppose that the minute papilla, which often represents the pistil in male flowers, and which is formed of mere cellular tissue, can thus act? Can we suppose that rudimentary teeth, which are subsequently absorbed, are beneficial to the rapidly growing embryonic calf by removing matter so precious as phosphate of lime? When a man’s fingers have been amputated, imperfect nails have been known to appear on the stumps, and I could as soon believe that these vestiges of nails are developed in order to excrete horny matter, as that the rudimentary nails on the fin of the manatee have been developed for this same purpose.
On the view of descent with modification, the origin of rudimentary organs is comparatively simple; and we can understand to a large extent the laws governing their imperfect development. We have plenty of cases of rudimentary organs in our domestic productions, as the stump of a tail in tailless breeds, the vestige of an ear in earless breeds of sheep—the reappearance of minute dangling horns in hornless breeds of cattle, more especially, according to Youatt, in young animals—and the state of the whole flower in the cauliflower. We often see rudiments of various parts in monsters; but I doubt whether any of these cases throw light on the origin of rudimentary organs in a state of nature, further than by showing that rudiments can be produced; for the balance of evidence clearly indicates that species under nature do not undergo great and abrupt changes. But we learn from the study of our domestic productions that the disuse of parts leads to their reduced size; and that the result is inherited.
It appears probable that disuse has been the main agent in rendering organs rudimentary. It would at first lead by slow steps to the more and more complete reduction of a part, until at last it became rudimentary—as in the case of the eyes of animals inhabiting dark caverns, and of the wings of birds inhabiting oceanic islands, which have seldom been forced by beasts of prey to take flight, and have ultimately lost the power of flying. Again, an organ, useful under certain conditions, might become injurious under others, as with the wings of beetles living on small and exposed islands; and in this case natural selection will have aided in reducing the organ, until it was rendered harmless and rudimentary.
Any change in structure and function, which can be effected by small stages, is within the power of natural selection; so that an organ rendered, through changed habits of life, useless or injurious for one purpose, might be modified and used for another purpose. An organ might, also, be retained for one alone of its former functions. Organs, originally formed by the aid of natural selection, when rendered useless may well be variable, for their variations can no longer be checked by natural selection. All this agrees well with what we see under nature. Moreover, at whatever period of life either disuse or selection reduces an organ, and this will generally be when the being has come to maturity and to exert its full powers of action, the principle of inheritance at corresponding ages will tend to reproduce the organ in its reduced state at the same mature age, but will seldom affect it in the embryo. Thus we can understand the greater size of rudimentary organs in the embryo relatively to the adjoining parts, and their lesser relative size in the adult. If, for instance, the digit of an adult animal was used less and less during many generations, owing to some change of habits, or if an organ or gland was less and less functionally exercised, we may infer that it would become reduced in size in the adult descendants of this animal, but would retain nearly its original standard of development in the embryo.
There remains, however, this difficulty. After an organ has ceased being used, and has become in consequence much reduced, how can it be still further reduced in size until the merest vestige is left; and how can it be finally quite obliterated? It is scarcely possible that disuse can go on producing any further effect after the organ has once been rendered functionless. Some additional explanation is here requisite which I cannot give. If, for instance, it could be proved that every part of the organisation tends to vary in a greater degree towards diminution than toward augmentation of size,