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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.
06 - Difficutiles in Theory 06-11 - Organs of Small Importance 30 Organs now of trifling importance have probably in some cases been of high importance to an early progenitor, and, after having been slowly perfected at a former period, have been transmitted to existing species in nearly the same state, although now of very slight use; but any actually injurious deviations in their structure would of course have been checked by natural selection.

Seeing how important an organ of locomotion the tail is in most aquatic animals, its general presence and use for many purposes in so many land animals, which in their lungs or modified swimbladders betray their aquatic origin, may perhaps be thus accounted for.
cheetah
cheetah


A well-developed tail having been formed in an aquatic animal, it might subsequently come to be worked in for all sorts of purposes,- as a fly-flapper, an organ of prehension, or as an aid in turning, as in the case of the dog, though the aid in this latter respect must be slight, for the hare, with hardly any tail, can double still more quickly.
11 - Geographical Distribution 11-04 - Centres of creation 10 On these views, it is obvious, that the several species of the same genus, though inhabiting the most distant quarters of the world, must originally have proceeded from the same source, as they have descended from the same progenitor.

In the case of those species, which have undergone during whole geological periods but little modification, there is not much difficulty in believing that they may have migrated from the same region; for during the vast geographical and climatal changes which will have supervened since ancient times, almost any amount of migration is possible.

But in many other cases, in which we have reason to believe that the species of a genus have been produced within comparatively recent times, there is great difficulty on this head.

It is also obvious that the individuals of the same species, though now inhabiting distant and isolated regions, must have proceeded from one spot, where their parents were first produced: for, as explained in the last chapter, it is incredible that individuals identically the same should ever have been produced through natural selection from parents specifically distinct.

We are thus brought to the question which has been largely discussed by naturalists, namely, whether species have been created at one or more points of the earth's surface.

Undoubtedly there are very many cases of extreme difficulty, in understanding how the same species could possibly have migrated from some one point to the several distant and isolated points, where now found.

Nevertheless the simplicity of the view that each species was first produced within a single region captivates the mind.

He who rejects it, rejects the vera causa of ordinary generation with subsequent migration, and calls in the agency of a miracle.

It is universally admitted, that in most cases the area inhabited by a species is continuous; and when a plant or animal inhabits two points so distant from each other, or with an interval of such a nature, that the space could not be easily passed over by migration, the fact is given as something remarkable and exceptional.

The capacity of migrating across the sea is more distinctly limited in terrestrial mammals, than perhaps in any other organic beings; and, accordingly, we find no inexplicable cases of the same mammal inhabiting distant points of the world.

No geologist will feel any difficulty in such cases as Great Britain having been formerly united to Europe, and consequently possessing the same quadrupeds.

But if the same species can be produced at two separate points, why do we not find a single mammal common to Europe and Australia or South America? The conditions of life are nearly the same, so that a multitude of European animals and plants have become naturalised in America and Australia; and some of the aboriginal plants are identically the same at these distant points of the northern and southern hemispheres?

The answer, as I believe, is, that mammals have not been able to migrate, whereas some plants, from their varied means of dispersal, have migrated across the vast and broken interspace.

The great and striking influence which barriers of every kind have had on distribution, is intelligible only on the view that the great majority of species have been produced on one side alone, and have not been able to migrate to the other side.

Some few families, many sub-families, very many genera, and a still greater number of sections of genera are confined to a single region; and it has been observed by several naturalists, that the most natural genera, or those genera in which the species are most closely related to each other, are generally local, or confined to one area.

What a strange anomaly it would be, if, when coming one step lower in the series, to the individuals of the same species, a directly opposite rule prevailed; and species were not local, but had been produced in two or more distinct areas!

Hence it seems to me, as it has to many other naturalists, that the view of each species having been produced in one area alone, and having subsequently migrated from that area as far as its powers of migration and subsistence under past and present conditions permitted, is the most probable.

Undoubtedly many cases occur, in which we cannot explain how the same species could have passed from one point to the other.

But the geographical and climatal changes, which have certainly occurred within recent geological times, must have interrupted or rendered discontinuous the formerly continuous range of many species.

So that we are reduced to consider whether the exceptions to continuity of range are so numerous and of so grave a nature, that we ought to give up the belief, rendered probable by general considerations, that each species has been produced within one area, and has migrated thence as far as it could.

It would be hopelessly tedious to discuss all the exceptional cases of the same species, now living at distant and separated points; nor do I for a moment pretend that any explanation could be offered of many such cases.

But after some preliminary remarks, I will discuss a few of the most striking classes of facts; namely, the existence of the same species on the summits of distant mountain-ranges, and at distant points in the arctic and antarctic regions; and secondly (in the following chapter), the wide distribution of freshwater productions; and thirdly, the occurrence of the same terrestrial species on islands and on the mainland, though separated by hundreds of miles of open sea.

If the existence of the same species at distant and isolated points of the earth's surface, can in many instances be explained on the view of each species having migrated from a single birthplace; then, considering our ignorance with respect to former climatal and geographical changes and various occasional means of transport, the belief that this has been the universal law, seems to me incomparably the safest.

In discussing this subject, we shall be enabled at the same time to consider a point equally important for us, namely, whether the several distinct species of a genus, which on my theory have all descended from a common progenitor, can have migrated (undergoing modification during some part of their migration) from the area inhabited by their progenitor.

If it can be shown to be almost invariably the case, that a region, of which most of its inhabitants are closely related to, or belong to the same genera with the species of a second region, has probably received at some former period immigrants from this other region, my theory will be strengthened; for we can clearly understand, on the principle of modification, why the inhabitants of a region should be related to those of another region, whence it has been stocked.

A volcanic island, for instance, upheaved and formed at the distance of a few hundreds of miles from a continent, would probably receive from it in the course of time a few colonists, and their descendants, though modified, would still be plainly related by inheritance to the inhabitants of the continent.

Cases of this nature are common, and are, as we shall hereafter more fully see, inexplicable on the theory of independent creation.

This view of the relation of species in one region to those in another, does not differ much (by substituting the word variety for species) from that lately advanced in an ingenious paper by Mr Wallace, in which he concludes, that `every species has come into existence coincident both in space and time with a pre-existing closely allied species.' And I now know from correspondence, that this coincidence he attributes to generation with modification.

Alfred Russel Wallace
Alfred Russel Wallace


The previous remarks on `single and multiple centres of creation' do not directly bear on another allied question, namely whether all the individuals of the same species have descended from a single pair, or single hermaphrodite, or whether, as some authors suppose, from many individuals simultaneously created.

With those organic beings which never intercross (if such exist), the species, on my theory, must have descended from a succession of improved varieties, which will never have blended with other individuals or varieties, but will have supplanted each other; so that, at each successive stage of modification and improvement, all the individuals of each variety will have descended from a single parent.

But in the majority of cases, namely, with all organisms which habitually unite for each birth, or which often intercross, I believe that during the slow process of modification the individuals of the species will have been kept nearly uniform by intercrossing; so that many individuals will have gone on simultaneously changing, and the whole amount of modification will not have been due, at each stage, to descent from a single parent.

To illustrate what I mean: our English racehorses differ slightly from the horses of every other breed; but they do not owe their difference and superiority to descent from any single pair, but to continued care in selecting and training many individuals during many generations.

Race Horse
Race Horse


Before discussing the three classes of facts, which I have selected as presenting the greatest amount of difficulty on the theory of `single centres of creation,' I must say a few words on the means of dispersal.
05 - Laws of Variation 05-03 - Acclimatisation 40 On the whole, we may conclude that habit, or use and disuse, have, in some cases, played a considerable part in the modification of the constitution and structure; but that the effects have often been largely combined with, and sometimes overmastered by the natural selection of innate variations.