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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 protected in the ovarium at its base. The pistil consists of a stigma supported on the style ; but in some Compositae,
the male florets, which of course cannot be fecundated, bave a pistil, which is in a rudimentary state, for it is not crowned with a stigma ; but the style remains well developed, and is clothed with hairs as in other Compositæ, for the purpose of brushing the pollen out of the surrounding anthers. Again, an organ may become rudimentary for its proper purpose, and be used for a distinct object : in certain fish the swim-bladder seems to be nearly rudimentary for its proper function of giving buoyancy, but has become converted into a nascent breathing organ or lung. Other similar instances could be given.
Organs, however little developed, if of use, should not be called rudimentary ; they cannot properly be said to be in an atrophied condition ; they may be called nascent, and may hereafter be developed to any extent by natural selection. Rudimentary organs, on the other hand, are essentially useless, as teeth which never cut through the gums; in a still less developed condition, they would be of still less use. They cannot, therefore, under their present condition, have been formed by natural selection, which acts solely by the preservation of useful modifications; they have been retained, as we shall see, by inheritance, and relate to a former condition of their possessor,
It is difficult to know what are nascent organs ; looking to the future, we cannot of course tell how any part will be developed, and whether it is now nascent; looking to the past, creatures with an organ in a nascent condition will generally have been supplanted and exterminated by their successors with the organ in a more perfect and developed condition. The wing of the penguin is of high service, and acts as a fin ; it may, therefore, represent the nascent state of the wings of birds ; 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 is useless, and is truly rudimentary. The mammary glands of the Ornithorhynchus may, perhaps, be considered, in comparison with the udder of a cow, as in a nascent state. The ovigerous frena of certain cirripedes, which are only slightly developed and which have ceased to give attachment to the ova, are nascent branchiæ.
Rudimentary organs in the individuals of the same species are very liable to vary in degree of development and in other respects. Moreover, in closely allied species, the degree to which the same organ has been rendered rudimentary occasionally differs much. This latter fact is well exemplified in the state of the wings of the female moths in certain groups. Rudimentary organs may be utterly aborted; and this implies, that we find in an animal or plant no trace of an organ, which analogy would lead us to expect to find, and which is occasionally found in monstrous individuals of the species. Thus in the snapdragon (antirrhinum) we generally do not find a rudiment of a fifth stamen; but this may sometimes be seen. In tracing the homologies of the same part in different members of
a class, nothing is more common, or more necessary, than the use and discovery of rudiments. This is well shown in the drawings given by Owen of the bones of the leg of the horse, ox, and rhinoceros.
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 or organ is of greater size relatively to the adjoining parts in the embryo, 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, also, a rudimentary organ in the adult is often said to have retained its embryonic condition.
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 reason. ing power which tells us plainly 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 seems to me no explanation, merely a re-statement of the fact. Would it be thought sufficient to say that because planets revolve in elliptic courses round the sun, satellites follow the same course round the planets, for the sake of symmetry, and to complete the scheme of nature ? An eminent physiologist accounts for the presence of rudimentary organs, by supposing that they serve to excrete matter in excess, or injurious to the system ; but can we suppose that the minute papilla, which often represents the pistil in male flowers, and which is formed merely of cellular tissue, can thus act? Can we suppose that the formation of rudimentary teeth, which are subsequently absorbed, can be of any service to the rapidly growing embryonic calf by the excretion of precious phosphate of lime? When a man's fingers have been amputated, imperfect nails sometimes appear on the stumps: I could as soon believe that these vestiges of nails have appeared, not from unknown laws of growth, but in order to excrete horny matter, as that the rudimentary nails on the fin of the manatee were formed for this purpose.
On my view of descent with modification, the origin of rudimentary organs is simple. 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,-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 I doubt whether species under nature ever undergo abrupt changes. I believe that disuse has been the main agency ; that it has led in successive generations to the gradual reduction of various organs, until they have become 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 to take fight, 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 would continue slowly to reduce the organ, until it was rendered harmless and rudimentary.
Any change in function, which can be effected by insensibly small steps, is within the power of natural selection ; so that an organ rendered, during changed habits of life, useless or injurious for one purpose, might be modified and used for another purpose. Or an organ might be retained for one alone of its former functions. An organ, when rendered useless, may well be variable, for its variations cannot be checked by natural selection. At whatever period of life disuse or selection reduces an organ, and this will generally be when the being has come to maturity and to its full powers of action, the principle of inheritance at corresponding ages will reproduce the organ in its reduced state at the same age, and consequently will seldom affect or reduce it in the embryo. Thus we can understand the greater relative size of rudimentary organs in the embryo, and their lesser relative size in the adult. But if each step of the process of reduction were to be inherited, not at the corresponding age, but at an extremely early period of life (as we have good reason to believe to be possible), the rudimentary part would tend to be wholly lost, and we should have a case of complete abortion. The principle, also, of economy, explained in a former chapter, by which the materials forming any part or structure, if not useful to the
possessor, will be saved as far as is possible, will probably often come into play; and this will tend to cause the entire obliteration of a rudimentary organ.
As the presence of rudimentary organs is thus due to the tendency in every part of the organisation, which has long existed, to be inherited-we can understand, on the genealogical view of classification, how it is that systematists have 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 in seeking 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 ordinary doctrine of creation, might even have been anticipated, and can be accounted for by the laws of inheritance.
Summary.—In this chapter I have attempted to show, that the subordination of group to group in all organisms throughout all time; that the nature of the relationship, by which all living and extinct beings are united by complex, radiating, and circuitous lines of affinities into one grand system; the rules followed and the difficulties encountered by naturalists in their classifications; the value set upon characters, if constant and prevalent, whether of high vital importance, or of the most trifling importance, or, as in rudimentary organs, of no importance; the wide opposition in value between analogical or adaptive characters, and characters of true affinity; and other such rules ;-all naturally follow on the view of the common parentage of those forms which are considered by naturalists as allied, together with their modification through natural selection, with its contingencies of extinction and divergence of character. In considering this view of classification, it should be