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14 - Recapitulation and Conclusion 14-02 - Recapitulation of the general and special circumstances in its favour 10 Now let us turn to the other side of the argument.

Under domestication we see much variability.

This seems to be mainly due to the reproductive system being eminently susceptible to changes in the conditions of life so that this system, when not rendered impotent, fails to reproduce offspring exactly like the parent-form.

Variability is governed by many complex laws, -- by correlation of growth, by use and disuse, and by the direct action of the physical conditions of life.

There is much difficulty in ascertaining how much modification our domestic productions have undergone; but we may safely infer that the amount has been large, and that modifications can be inherited for long periods.

As long as the conditions of life remain the same, we have reason to believe that a modification, which has already been inherited for many generations, may continue to be inherited for an almost infinite number of generations.

On the other hand we have evidence that variability, when it has once come into play, does not wholly cease; for new varieties are still occasionally produced by our most anciently domesticated productions.

Man does not actually produce variability; he only unintentionally exposes organic beings to new conditions of life, and then nature acts on the organisation, and causes variability.

But man can and does select the variations given to him by nature, and thus accumulate them in any desired manner.

He thus adapts animals and plants for his own benefit or pleasure.

He may do this methodically, or he may do it unconsciously by preserving the individuals most useful to him at the time, without any thought of altering the breed.

It is certain that he can largely influence the character of a breed by selecting, in each successive generation, individual differences so slight as to be quite inappreciable by an uneducated eye.

This process of selection has been the great agency in the production of the most distinct and useful domestic breeds.

That many of the breeds produced by man have to a large extent the character of natural species, is shown by the inextricable doubts whether very many of them are varieties or aboriginal species.
14 - Recapitulation and Conclusion 14-01 - Recapitulation of the difficulties on the theory of Natural Selection 10 As this whole volume is one long argument, it may be convenient to the reader to have the leading facts and inferences briefly recapitulated.

That many and grave objections may be advanced against the theory of descent with modification through natural selection, I do not deny.

I have endeavoured to give to them their full force.

Nothing at first can appear more difficult to believe than that the more complex organs and instincts should have been perfected not by means superior to, though analogous with, human reason, but by the accumulation of innumerable slight variations, each good for the individual possessor.

Nevertheless, this difficulty, though appearing to our imagination insuperably great, cannot be considered real if we admit the following propositions, namely, -- that gradations in the perfection of any organ or instinct, which we may consider, either do now exist or could have existed, each good of its kind, -- that all organs and instincts are, in ever so slight a degree, variable, -- and, lastly, that there is a struggle for existence leading to the preservation of each profitable deviation of structure or instinct.

The truth of these propositions cannot, I think, be disputed.
13 - Mutual Affinities of Organic Beings: Morphology: Embryology: Rudimentary Or 13-12 - Summary 10 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 borne in mind that the element of descent has been universally used in ranking together the sexes, ages, and acknowledged varieties of the same species, however different they may be in structure.

If we extend the use of this element of descent, the only certainly known cause of similarity in organic beings, we shall understand what is meant by the natural system: it is genealogical in its attempted arrangement, with the grades of acquired difference marked by the terms varieties, species, genera, families, orders, and classes.

On this same view of descent with modification, all the great facts in Morphology become intelligible, whether we look to the same pattern displayed in the homologous organs, to whatever purpose applied, of the different species of a class; or to the homologous parts constructed on the same pattern in each individual animal and plant.

On the principle of successive slight variations, not necessarily or generally supervening at a very early period of life, and being inherited at a corresponding period, we can understand the great leading facts in Embryology; namely, the resemblance in an individual embryo of the homologous parts, which when matured will become widely different from each other in structure and function; and the resemblance in different species of a class of the homologous parts or organs, though fitted in the adult members for purposes as different as possible.

Larvae are active embryos, which have become specially modified in relation to their habits of life, through the principle of modifications being inherited at corresponding ages.

On this same principle and bearing in mind, that when organs are reduced in size, either from disuse or selection, it will generally be at that period of life when the being has to provide for its own wants, and bearing in mind how strong is the principle of inheritance the occurrence of rudimentary organs and their final abortion, present to us no inexplicable difficulties; on the contrary, their presence might have been even anticipated.

The importance of embryological characters and of rudimentary organs in classification is intelligible, on the view that an arrangement is only so far natural as it is genealogical.

Finally, the several classes of facts which have been considered in this chapter, seem to me to proclaim so plainly, that the innumerable species, genera, and families of organic beings, with which this world is peopled, have all descended, each within its own class or group, from common parents, and have all been modified in the course of descent, that I should without hesitation adopt this view, even if it were unsupported by other facts or arguments.
13 - Mutual Affinities of Organic Beings: Morphology: Embryology: Rudimentary Or 13-11 - RUDIMENTARY ORGANS; their origin explained 10 Organs or parts in this strange condition, bearing the stamp of inutility, are extremely common throughout nature.

For instance, rudimentary mammae are very general in the males of mammals: I presume that the `bastard-wing' in birds may be safely considered as a digit in a rudimentary state: in very many snakes one lobe of the lungs is rudimentary; in other snakes there are rudiments of the pelvis and hind limbs.

Some of the cases of rudimentary organs are extremely curious; for instance, the presence of teeth in foetal whales, which when grown up have not a tooth in their heads; and the presence of teeth, which never cut through the gums, in the upper jaws of our unborn calves.

It has even been stated on good authority that rudiments of teeth can be detected in the beaks of certain embryonic birds.

Nothing can be plainer than that wings are formed for flight, yet in how many insects do we see wings so reduced in size as to be utterly incapable of flight, and not rarely lying under wing-cases, firmly soldered together!

The meaning of rudimentary organs is often quite unmistakeable: for instance there are beetles of the same genus (and even of the same species) resembling each other most closely in all respects, one of which will have full-sized wings, and another mere rudiments of membrane; and here it is impossible to doubt, that the rudiments represent wings.

Rudimentary organs sometimes retain their potentiality, and are merely not developed: this seems to be the case with the mammae of male mammals, for many instances are on record of these organs having become well developed in full-grown males, and having secreted milk.

So again there are normally four developed and two rudimentary teats in the udders of the genus Bos, but in our domestic cows the two sometimes become developed and give milk.

In individual plants of the same species the petals sometimes occur as mere rudiments, and sometimes in a well-developed state.

In plants with separated sexes, the male flowers often have a rudiment of a pistil; and Kölreuter found that by crossing such male plants with an hermaphrodite species, the rudiment of the pistil in the hybrid offspring was much increased in size; and this shows that the rudiment and the perfect pistil are essentially alike in nature.

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 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, have 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 compositae, 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 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.

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 reasoning 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 restatement 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 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 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 easily 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.