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05 - Laws of Variation 05-01 - Effects of External Conditions 10 I HAVE hitherto sometimes spoken as if the variations- so common and multiform with organic beings under domestication, and in a lesser degree with those under nature- were due to chance.

This, of course, is a wholly incorrect expression, but it serves to acknowledge plainly our ignorance of the cause of each particular variation.

Some authors believe it to be as much the function of the reproductive system to produce individual differences, or slight deviations of structure, as to make the child like its parents.

But the fact of variations and monstrosities occurring much more frequently under domestication than under nature, and the greater variability of species having wider ranges than of those with restricted ranges, lead to the conclusion that variability is generally related to the conditions of life to which each species has been exposed during several successive generations.

In the first chapter I attempted to show that changed conditions act in two ways, directly on the whole organisation or on certain parts alone, and indirectly through the reproductive system.

In all cases there are two factors, the nature of the organism, which is much the most important of the two, and the nature of the conditions.

The direct action of changed conditions leads to definite or indefinite results. In the latter case the organisation seems to become plastic, and we have much fluctuating variability.

In the former case the nature of the organism is such that it yields readily, when subjected to certain conditions, and all, or nearly all the individuals become modified in the same way.

It is very difficult to decide how far changed conditions, such as of climate, food, &c., have acted in a definite manner.

There is reason to believe that in the course of time the effects have been greater than can be proved by clear evidence.

But we may safely conclude that the innumerable complex co-adaptations of structure, which we see throughout nature between various organic beings, cannot be attributed simply to such action.

In the following cases the conditions seem to have produced some slight definite effect: E. Forbes asserts that shells at their southern limit, and when living in shallow water, are more brightly coloured than those of the same species from further north or from a greater depth; but this certainly does not always hold good.

Sea Shell
Sea Shell
10 - On The Geological Succession of Organic Beings 10-10 - Summary of preceding and present chapters 10 I have attempted to show that the geological record is extremely imperfect; that only a small portion of the globe has been geologically explored with care; that only certain classes of organic beings have been largely preserved in a fossil state; that the number both of specimens and of species, preserved in our museums, is absolutely as nothing compared with the incalculable number of generations which must have passed away even during a single formation; that, owing to subsidence being necessary for the accumulation of fossiliferous deposits thick enough to resist future degradation, enormous intervals of time have elapsed between the successive formations; that there has probably been more extinction during the periods of subsidence, and more variation during the periods of elevation, and during the latter the record will have been least perfectly kept; that each single formation has not been continuously deposited; that the duration of each formation is, perhaps, short compared with the average duration of specific forms; that migration has played an important part in the first appearance of new forms in any one area and formation; that widely ranging species are those which have varied most, and have oftenest given rise to new species; and that varieties have at first often been local.

All these causes taken conjointly, must have tended to make the geological record extremely imperfect, and will to a large extent explain why we do not find interminable varieties, connecting together all the extinct and existing forms of life by the finest graduated steps.

He who rejects these views on the nature of the geological record, will rightly reject my whole theory.

For he may ask in vain where are the numberless transitional links which must formerly have connected the closely allied or representative species, found in the several stages of the same great formation.

He may disbelieve in the enormous intervals of time which have elapsed between our consecutive formations; he may overlook how important a part migration must have played, when the formations of any one great region alone, as that of Europe, are considered; he may urge the apparent, but often falsely apparent, sudden coming in of whole groups of species.

europe
europe


He may ask where are the remains of those infinitely numerous organisms which must have existed long before the first bed of the Silurian system was deposited: I can answer this latter question only hypothetically, by saying that as far as we can see, where our oceans now extend they have for an enormous period extended, and where our oscillating continents now stand they have stood ever since the Silurian epoch; but that long before that period, the world may have presented a wholly different aspect; and that the older continents, formed of formations older than any known to us, may now all be in a metamorphosed condition, or may lie buried under the ocean.

Passing from these difficulties, all the other great leading facts in palaeontology seem to me simply to follow on the theory of descent with modification through natural selection.

We can thus understand how it is that new species come in slowly and successively; how species of different classes do not necessarily change together, or at the same rate, or in the same degree; yet in the long run that all undergo modification to some extent.

The extinction of old forms is the almost inevitable consequence of the production of new forms.

We can understand why when a species has once disappeared it never reappears.

Groups of species increase in numbers slowly, and endure for unequal periods of time; for the process of modification is necessarily slow, and depends on many complex contingencies.

The dominant species of the larger dominant groups tend to leave many modified descendants, and thus new sub-groups and groups are formed.

As these are formed, the species of the less vigorous groups, from their inferiority inherited from a common progenitor, tend to become extinct together, and to leave no modified offspring on the face of the earth.

Earth
Earth


But the utter extinction of a whole group of species may often be a very slow process, from the survival of a few descendants, lingering in protected and isolated situations.

When a group has once wholly disappeared, it does not reappear; for the link of generation has been broken.

We can understand how the spreading of the dominant forms of life, which are those that oftenest vary, will in the long run tend to people the world with allied, but modified, descendants; and these will generally succeed in taking the places of those groups of species which are their inferiors in the struggle for existence.

Hence, after long intervals of time, the productions of the world will appear to have changed simultaneously.

We can understand how it is that all the forms of life, ancient and recent, make together one grand system; for all are connected by generation.

We can understand, from the continued tendency to divergence of character, why the more ancient a form is, the more it generally differs from those now living.

Why ancient and extinct forms often tend to fill up gaps between existing forms, sometimes blending two groups previously classed as distinct into one; but more commonly only bringing them a little closer together.

The more ancient a form is, the more often, apparently, it displays characters in some degree intermediate between groups now distinct; for the more ancient a form is, the more nearly it will be related to, and consequently resemble, the common progenitor of groups, since become widely divergent.

Extinct forms are seldom directly intermediate between existing forms; but are intermediate only by a long and circuitous course through many extinct and very different forms.

We can clearly see why the organic remains of closely consecutive formations are more closely allied to each other, than are those of remote formations; for the forms are more closely linked together by generation: we can clearly see why the remains of an intermediate formation are intermediate in character.

The inhabitants of each successive period in the world's history have beaten their predecessors in the race for life, and are, in so far, higher in the scale of nature; and this may account for that vague yet ill-defined sentiment, felt by many palaeontologists, that organisation on the whole has progressed.

If it should hereafter be proved that ancient animals resemble to a certain extent the embryos of more recent animals of the same class, the fact will be intelligible.

embryo
embryo


The succession of the same types of structure within the same areas during the later geological periods ceases to be mysterious, and is simply explained by inheritance.

If then the geological record be as imperfect as I believe it to be, and it may at least be asserted that the record cannot be proved to be much more perfect, the main objections to the theory of natural selection are greatly diminished or disappear.

On the other hand, all the chief laws of palaeontology plainly proclaim, as it seems to me, that species have been produced by ordinary generation: old forms having been supplanted by new and improved forms of life, produced by the laws of variation still acting round us, and preserved by Natural Selection.

fossil
fossil
08 - Hybridism 08-08 - Hybrids and mongrels compared independently of their fertility 10 Hybrids and Mongrels compared, independently of their fertility.

Independently of the question of fertility, the offspring of species when crossed and of varieties when crossed may be compared in several other respects.

Gaertner, whose strong wish was to draw a marked line of distinction between species and varieties, could find very few and, as it seems to me, quite unimportant differences between the so-called hybrid offspring of species, and the so-called mongrel offspring of varieties.

And, on the other hand, they agree most closely in very many important respects.

I shall here discuss this subject with extreme brevity.

The most important distinction is, that in the first generation mongrels are more variable than hybrids; but Gaertner admits that hybrids from species which have long been cultivated are often variable in the first generation; and I have myself seen striking instances of this fact.

Gaertner further admits that hybrids between very closely allied species are more variable than those from very distinct species; and this shows that the difference in the degree of variability graduates away.

When mongrels and the more fertile hybrids are propagated for several generations an extreme amount of variability in their offspring is notorious; but some few cases both of hybrids and mongrels long retaining uniformity of character could be given.

The variability, however, in the successive generations of mongrels is, perhaps, greater than in hybrids.

This greater variability of mongrels than of hybrids does not seem to me at all surprising.

For the parents of mongrels are varieties, and mostly domestic varieties (very few experiments having been tried on natural varieties), and this implies in most cases that there has been recent variability; and therefore we might expect that such variability would often continue and be super-added to that arising from the mere act of crossing

The slight degree of variability in hybrids from the first cross or in the first generation, in contrast with their extreme variability in the succeeding generations, is a curious fact and deserves attention.

For it bears on and corroborates the view which I have taken on the cause of ordinary variability; namely, that it is due to the reproductive system being eminently sensitive to any change in the conditions of life, being thus often rendered either impotent or at least incapable of its proper function of producing offspring identical with the parent-form.

Now hybrids in the first generation are descended from species (excluding those long cultivated) which have not had their reproductive systems in any way affected, and they are not variable; but hybrids themselves have their reproductive systems seriously affected, and their descendants are highly variable.

But to return to our comparison of mongrels and hybrids: Gaertner states that mongrels are more liable than hybrids to revert to either parent-form; but this, if it be true, is certainly only a difference in degree.

Gaertner further insists that when any two species, although most closely allied to each other, are crossed with a third species, the hybrids are widely different from each other; whereas if two very distinct varieties of one species are crossed with another species, the hybrids do not differ much.

But this conclusion, as far as I can make out, is founded on a single experiment; and seems directly opposed to the results of several experiments made by Koelreuter.

These alone are the unimportant differences, which Gaertner is able to point out, between hybrid and mongrel plants.

On the other hand, the resemblance in mongrels and in hybrids to their respective parents, more especially in hybrids produced from nearly related species, follows according to Gaertner the same laws.

When two species are crossed, one has sometimes a prepotent power of impressing its likeness on the hybrid; and so I believe it to be with varieties of plants.

With animals one variety certainly often has this prepotent power over another variety.

Hybrid plants produced from a reciprocal cross, generally resemble each other closely; and so it is with mongrels from a reciprocal cross.

Both hybrids and mongrels can be reduced to either pure parent-form, by repeated crosses in successive generations with either parent.

These several remarks are apparently applicable to animals; but the subject is here excessively complicated, partly owing to the existence of secondary sexual characters; but more especially owing to prepotency in transmitting likeness running more strongly in one sex than in the other, both when one species is crossed with another, and when one variety is crossed with another variety.

For instance, I think those authors are right, who maintain that the ass has a prepotent power over the horse, so that both the mule and the hinny more resemble the ass than the horse; but that the prepotency runs more strongly in the male-ass than in the female, so that the mule, which is the offspring of the male-ass and mare, is more like an ass, than is the hinny, which is the offspring of the female-ass and stallion.

ass
ass

horse
horse

mule
mule

hinny
hinny


Much stress has been laid by some authors on the supposed fact, that mongrel animals alone are born closely like one of their parents; but it can be shown that this does sometimes occur with hybrids; yet I grant much less frequently with hybrids than with mongrels.

Looking to the cases which I have collected of cross-bred animals closely resembling one parent, the resemblances seem chiefly confined to characters almost monstrous in their nature, and which have suddenly appeared such as albinism, melanism, deficiency of tail or horns, or additional fingers and toes; and do not relate to characters which have been slowly acquired by selection.

Consequently, sudden reversions to the perfect character of either parent would be more likely to occur with mongrels, which are descended from varieties often suddenly produced and semi-monstrous in character, than with hybrids, which are descended from species slowly and naturally produced.

On the whole I entirely agree with Dr Prosper Lucas, who, after arranging an enormous body of facts with respect to animals, comes to the conclusion, that the laws of resemblance of the child to its parents are the same, whether the two parents differ much or little from each other, namely in the union of individuals of the same variety, or of different varieties, or of distinct species.

Laying aside the question of fertility and sterility, in all other respects there seems to be a general and close similarity in the offspring of crossed species, and of crossed varieties.

If we look at species as having been specially created, and at varieties as having been produced by secondary laws, this similarity would be an astonishing fact.

But it harmonizes perfectly with the view that there is no essential distinction between species and varieties.
04 - Natural Selection 04-01 - Natural Selection 10 How will the struggle for existence, briefly discussed in the last chapter, act in regard to variation?

Can the principle of selection, which we have seen is so potent in the hands of man, apply under nature?

I think we shall see that it can act most efficiently.

Let the endless number of slight variations and individual differences occurring in our domestic productions, and, in a lesser degree, in those under nature, be borne in mind; as well as the strength of the hereditary tendency.

Under domestication, it may be truly said that the whole organisation becomes in some degree plastic.

clay
clay


But the variability, which we almost universally meet with in our domestic productions, is not directly produced, as Hooker and Asa Gray have well remarked, by man; he can neither originate varieties, nor prevent their occurrence; he can preserve and accumulate such as do occur.

Unintentionally he exposes organic beings to new and changing conditions of life, and variability ensues; but similar changes of conditions might and do occur under nature.

Let it also be borne in mind how infinitely complex and close-fitting are the mutual relations of all organic beings to each other and to their physical conditions of life; and consequently what infinitely varied diversities of structure might be of use to each being under changing conditions of life.

Can it, then, be thought improbable, seeing that variations useful to man have undoubtedly occurred, that other variations useful in some way to each being in the great and complex battle of life, should occur in the course of many successive generations?

If such do occur, can we doubt (remembering that many more individuals are born than can possibly survive) that individuals having any advantage, however slight, over others, would have the best chance of surviving and of procreating their kind?

On the other hand, we may feel sure that any variation in the least degree injurious would be rigidly destroyed.

This preservation of favourable individual differences and variations, and the destruction of those which are injurious, I have called Natural Selection.