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Charles Robert Darwin (1809–1882).  Origin of Species.
The Harvard Classics.  1909–14.
 
XIV. Mutual Affinities of Organic Beings: Morphology—Embryology—Rudimentary Organs
 
Rudimentary, Atrophied, and Aborted Organs
 
 
ORGANS or parts in this strange condition, bearing the plain stamp of inutility, are extremely common, or even general, throughout nature. It would be impossible to name one of the higher animals in which some part or other is not in a rudimentary condition. In the mammalia, for instance, the males possess rudimentary mammæ; in snakes one lobe of the lungs is rudimentary; in birds the “bastard-wing” may safely be considered as a rudimentary digit, and in some species the whole wing is so far rudimentary that it cannot be used for flight. What can be more curious than the presence of teeth in fœtal whales, which when grown up have not a tooth in their heads; or the teeth, which never cut through the gums, in the upper jaws of unborn calves?  1
  Rudimentary organs plainly declare their origin and meaning in various ways. There are beetles belonging to closely allied species, or even to the same identical species, which have either full-sized and perfect wings, or mere rudiments of membrane, which not rarely lie under wing-covers firmly soldered together; and in these cases it is impossible to doubt, that the rudiments represent wings. Rudimentary organs sometimes retain their potentiality: this occasionally occurs with the mammae of male mammals, which have been known to become well developed and to secrete milk. So again in the udders in the genus Bos, there are normally four developed and two rudimentary teats; but the latter in our domestic cows sometimes become well developed and yield milk. In regard to plants the petals are sometimes rudimentary, and sometimes well-developed in the individuals of the same species. In certain plants having separated sexes Kölreuter found that by crossing a species, in which the male flowers included a rudiment of a pistil, with an hermaphrodite species, having of course a well-developed pistil, the rudiment in the hybrid offspring was much increased in size; and this clearly shows that the rudimentary and perfect pistils are essentially alike in nature. An animal may possess various parts in a perfect state, and yet they may in one sense be rudimentary, for they are useless: thus the tadpole of the common salamander or water-newt, as Mr. G. H. Lewes remarks, “has gills, and passes its existence in the water; but the Salamandra atra, which lives high up among the mountains, brings forth its young full-formed. This animal never lives in the water. Yet if we open a gravid female, we find tadpoles inside her with exquisitely feathered gills; and when placed in water they swim about like the tadpoles of the water-newt. Obviously this aquatic organisation has no reference to the future life of the animal, nor has it any adaptation to its embryonic condition; it has solely reference to ancestral adaptations, it repeats a phase in the development of its progenitors.”  2
  An organ, serving for two purposes, may become rudimentary or utterly aborted for one, even the more important purpose, and remain perfectly efficient for the other. Thus in plants, the office of the pistil is to allow the pollen-tubes to reach the ovules within the ovarium. The pistil consists of a stigma supported on a style; but in some Compositæ, the male florets, which of course cannot be fecundated, have a rudimentary pistil, for it is not crowned with a stigma; but the style remains well developed and is clothed in the usual manner with hairs, which serve to brush the pollen out of the surrounding and conjoined anthers. Again, an organ may become rudimentary for its proper purpose, and be used for a distinct one: in certain fishes the swim-bladder seems to be rudimentary for its proper function of giving buoyancy, but has become converted into a nascent breathing organ or lung. Many similar instances could be given.  3
  Useful organs, however little they may be developed, unless we have reason to suppose that they were formerly more highly developed, ought not to be considered as rudimentary. They may be in a nascent condition, and in progress towards further development. Rudimentary organs, on the other hand, are either quite useless, such as teeth which never cut through the gums, or almost useless, such as the wings of an ostrich, which serve merely as sails. As organs in this condition would formerly, when still less developed, have been of even less use than at present, they cannot formerly have been produced through variation and natural selection, which acts solely by the preservation of useful modifications. They have been partially retained by the power of inheritance, and relate to a former state of things. It is, however, often difficult to distinguish between rudimentary and nascent organs; for we can judge only by analogy whether a part is capable of further development, in which case alone it deserves to be called nascent. Organs in this condition will always be somewhat rare; for beings thus provided will commonly have been supplanted by their successors with the same organ in a more perfect state, and consequently will have become long ago extinct. The wing of the penguin is of high service, acting as a fin; it may, therefore, represent the nascent state of the wing: not that I believe this to be the case; it is more probably a reduced organ, modified for a new function: the wing of the Apteryx, on the other hand, is quite useless, and is truly rudimentary. Owen considers the simple filamentary limbs of the lepidosiren as the “beginnings of organs which attain full functional development in higher vertebrates”; but, according to the view lately advocated by Dr. Günther, they are probably remnants, consisting of the persistent axis of a fin, with the lateral rays of branches aborted. The mammary glands of the Ornithorhynchus may be considered, in comparison with the udders of a cow, as in a nascent condition. The ovigerous frena of certain cirripedes, which have ceased to give attachment to the ova and are feebly developed, are nascent branchiæ.  4
  Rudimentary organs in the individuals of the same species are very liable to vary in the degree of their development and in other respects. In closely allied species, also, the extent to which the same organ has been reduced occasionally differs much. This latter fact is well exemplified in the state of the wings of female moths belonging to the same family. Rudimentary organs may be utterly aborted; and this implies, that in certain animals or plants, parts are entirely absent which analogy would lead us to expect to find in them, and which are occasionally found in monstrous individuals. Thus in most of the Scrophulariaceæ the fifth stamen is utterly aborted; yet we may conclude that a fifth stamen once existed, for a rudiment of it is found in many species of the family, and this rudiment occasionally becomes perfectly developed, as may sometimes be seen in the common snap-dragon. In tracing the homologies of any part in different members of the same class, nothing is more common, or, in order fully to understand the relations of the parts, more useful than the discovery of rudiments. This is well shown in the drawings given by Owen of the leg-bones of the horse, ox, and rhinoceros.  5
  It is an important fact that rudimentary organs, such as teeth in the upper jaws of whales and ruminants, can often be detected in the embryo, but afterwards wholly disappear. It is also, I believe, a universal rule, that a rudimentary part is of greater size in the embryo relatively to the adjoining parts, than in the adult; so that the organ at this early age is less rudimentary, or even cannot be said to be in any degree rudimentary. Hence rudimentary organs in the adult are often said to have retained their embryonic condition.  6
  I have now given the leading facts with respect to rudimentary organs. In reflecting on them, every one 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 re-statement 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.  7
  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.  8
  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 ultimatelylost 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.  9
  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 has 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 effect 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.  10
  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 towards augmentation of size, then we should be able to understand how an organ which has become useless would be rendered, independently of the effects of disuse, rudimentary and would at last be wholly suppressed; for the variations towards diminished size would no longer be checked by natural selection. The principle of the economy of growth, explained in a former chapter, by which the materials forming any part, if not useful to the possessor, are saved as far as possible, will perhaps come into play in rendering a useless part rudimentary. But this principle will almost necessarily be confined to the earlier stages of the process of reduction; for we cannot suppose that a minute papilla, for instance, representing in a male flower the pistil of the female flower, and formed merely of cellular tissue, could be further reduced or absorbed for the sake of economising nutriment.  11
  Finally, as rudimentary organs, by whatever steps they may have been degraded into their present useless condition, are the record of a former state of things, and have been retained solely through the power of inheritance,—we can understand, on the genealogical view of classification, how it is that systematists, in placing organisms in their proper places in the natural system, have often found rudimentary parts as useful as, or even sometimes more useful than, parts of high physiological importance. Rudimentary organs may be compared with the letters in a word, still retained in the spelling, but become useless in the pronunciation, but which serve as a clue for its derivation. On the view of descent with modification, we may conclude that the existence of organs in a rudimentary, imperfect, and useless condition, or quite aborted, far from presenting a strange difficulty, as they assuredly do on the old doctrine of creation, might even have been anticipated in accordance with the views here explained.  12
 

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