Read Ebook: Darwin and After Darwin Volume 3 of 3 Post-Darwinian Questions: Isolation and Physiological Selection by Romanes George John

Font size: 10 px 15 px 20 px 25 px 30 px 35 px

Background color: BlackWhiteGrayLight GrayDark GrayDim Gray

Text color: BlackWhiteGrayLight GrayDark GrayDim Gray

Apply/Save settings

Ebook has 358 lines and 60439 words, and 8 pages

It will be seen, then, that both Darwin, and Wallace, fully recognize the necessity of finding some explanation of the infertility of allied species, over and above the mere reaction of morphological differentiation on the physiology of the reproductive system, and they both agree in suggesting additional causes, though they entirely disagree as to what these causes are. Now, the theory of physiological selection likewise suggests an additional cause--or, rather, a new explanation--and one which is surely the most probable. For what is to be explained? The very general association of a certain physiological peculiarity with that amount of morphological change which distinguishes species from species, of whatever kind the change may be, and in whatever family of the animal or vegetable kingdom it may occur. Well, the theory of physiological selection explains this very general association by the simple supposition that, at least in a large number of cases, it was the physiological peculiarity which first of all led to the morphological divergence, by interposing the bar of sterility between two sections of a previously uniform species; and by thus isolating the two sections one from another, started each upon a subsequently independent course of divergent evolution.

Or, to put it in another way, if the occurrence of this physiological peculiarity has been often the only possible means of isolating two sections of a species occupying a common area, and thus giving rise to a divergence of specific type , it is nothing less than a necessary consequence that many allied species should now present the physiological peculiarity in question. Thus the association between the physiological peculiarity and the morphological divergence is explained by the simple hypothesis, that the former has acted as a necessary condition to the occurrence of the latter. In the absence of other forms of isolation, the morphological divergence could not have taken place at all, had not the physiological peculiarity arisen; and hence it is that we now meet with so many cases where such divergence is associated with this peculiarity.

Suppose two divergent local varieties were to arise in adjacent areas, and were subsequently driven together into a single area.... If their unions be fertile, the isolation will be annulled by intercrossing--the two varieties will form one mean or average variety. But if the unions be infertile, the isolation will be preserved, and the two varieties will continue separate. Suppose now, and the supposition is by no means an improbable one, that this has taken place again and again in the evolution of species; then it is clear that those varietal forms which had continued to be fertile together would be swamped by intercrossing; while those varietal forms which had become infertile would remain isolated. Hence, in the long run, isolated forms occupying a common area would be infertile,

If then cross-sterility may thus arise even in association with geographical isolation, may it not also arise in its absence? And may it not thus give rise to the differentiation of varieties on account of this physiological isolation alone?

The first is, that, as Mr. Wallace remarks, "every species has come into existence coincident both in space and time with a pre-existing and closely allied species." I regard this as important evidence that physiological selection is one of the natural causes concerned. For the general fact implied is that every species has come into existence on an area occupied by its parent type, and therefore under circumstances which render it imperative that intercrossing with that type should be prevented. In the case of monotypic evolution by natural selection alone, intercrossing with the parent type is prevented through the gradual extinction of that type by successive generations of the developing type. But in the case of polytypic evolution, intercrossing with the parent type can only be prevented by some form of isolation other than natural selection; and here it is evident that cross-infertility with the parent type must be as efficient to that end as any other form of isolation that can be imagined. Consequently we might almost have expected beforehand that in a large proportional number of cases cross-infertility should have been the means employed. And the fact that this is actually the case so far corroborates the only theory which is able to explain it.

The second point is this.

It appears to be comparatively rare for any cause of specific divergence to prove effectual on common areas, unless it sooner or later becomes associated with some degree of cross-infertility. But through this association, the segregating influence of both the causes concerned is, as Mr. Gulick has shown, greatly increased. For instance, if the segregating influence of some degree of cross-infertility be associated with that of any other form of isolation, then, not only will the two segregating influences be added, but multiplied together. And thus, by their mutual action and reaction, divergent evolution is promoted at a rapidly increasing rate.

I will now summarize the main points of the theory of physiological isolation in a categorical form.

To prevent, if possible, the continuance of certain misunderstandings with regard to my original statement of the new theory, let me here disclaim some views which have been assigned to me. They are:

EVIDENCES OF PHYSIOLOGICAL SELECTION.

I will now give an outline sketch of the evidences in favour of the theory which has been set forth in the preceding chapter, stating first what is the nature of the verification which it requires.

The antecedent standing or logical basis of the theory has already been in large measure displayed in the preceding chapter; for it was impossible to state the theory without thereby showing in how considerable a degree it is self-evident. A brief recapitulation is therefore all that is here necessary.

It has been shown that divergent or polytypic evolution on common areas is inexplicable by natural selection alone. Hence the question arises: What form of isolation has, under such circumstances, rendered possible divergent evolution? In answer to this question the theory of physiological selection suggests that variations in the reproductive function occur in such a way as to isolate more or less perfectly from each other different sections of a species. While cross-fertility remains unimpaired among the members of each section, there is more or less cross-infertility when members of either section mate with those of the other. Thus a physiological barrier is interposed between the two sections; and any divergences of structure, colouring, or instinct arising in the members of either section will not in any way be affected by such divergences as arise among the members of the other.

In support of this suggestion, it has been shown in the preceding chapter that the very general association of cross-infertility with specific differentiation points most strongly to the inference that the former has usually been an indispensable condition to the occurrence of the latter. It cannot be denied that in many cases the specific distinction is now maintained by means of that sexual isolation which cross-infertility confers: it is therefore probable that such isolation has been instrumental in securing its initial attainment.

This probability is strengthened by the observed fact that the general association in question is conspicuously absent in the case of domesticated varieties, notwithstanding that their multitudinous and diverse varietal characters usually equal, and frequently surpass, specific characters in their degrees of divergence.

Since, then, it would seem to be impossible for divergent evolution on common areas to take place in the absence of some mode of isolation; since cross-infertility appears to be the only possible mode under the given circumstances; and since among domesticated varieties, where isolation is otherwise secured by artificial means, cross-infertility is usually absent, the logical foundations of the theory of physiological selection would seem to be securely laid.

We may therefore pass to more special lines of evidence.

Darwin has adduced very good evidence to show that large areas, notwithstanding the disadvantages which must arise from free intercrossing, are what he terms better manufactories of species than smaller areas, such as oceanic islands. On the other hand, as a matter of fact, oceanic islands are comparatively rich in peculiar species. These two statements, however, are not incompatible. Smaller areas are, as a rule, rich in peculiar species relatively to the number of their inhabitants; but it does not follow that they are rich in species as contrasted with larger areas containing very many more inhabitants. Therefore, the rules are that large areas turn out an absolutely greater number of specific types than small areas; although, relatively to the number of individuals or amount of population, the small areas turn out a larger number of species than the large areas.

The two subsidiary facts, that very closely allied species have restricted ranges, and that dominant species are rich in varieties, both seem to tell more in favour of physiological than of natural selection. For "very closely allied species" is but another name for species which scarcely differ from one another at all except in their reproductive systems; and, therefore, the more restricted their ranges, the more certainly would they have become fused by intercrossing with one another, had it not been for the barrier of sterility imposed by the primary distinction. Or rather, I should say, had it not been for the original occurrence of this barrier, these now closely-allied species could never have become species. Again, that dominant species should be rich in varieties is what might have been expected; for the greater the number of individuals in a species, the greater is the chance of variations taking place in all parts of the organic type, and particularly in the reproductive system, seeing that this system is the most sensitive to small changes in the conditions of life, and that the greater the number of individuals composing a specific type, the more certainty there is of some of them encountering such changes. Hence, the richness of dominant species in varieties is, I believe, mainly due to the greater opportunity which such species afford of some degree of cross-infertility arising between their constituent members.

Here is another general fact, also first noticed by Darwin, and one which he experiences some difficulty an explaining on the theory of natural selection. He says:--

In travelling from north to south over a continent, we generally meet at successive intervals with closely-allied or representative species, evidently filling the same place in the economy of the land. These representative species often meet and interlock, and as one becomes rarer and rarer, the other becomes more and more frequent, till the one replaces the other. But if we compare these species where they intermingle, they are generally as absolutely distinct from each other in every detail of structure as are specimens taken from the metropolis of each.... In the intermediate region, having intermediate conditions of life, why do we not now find closely-linking intermediate varieties? This difficulty for a long time quite confounded me. But I think it can in large part be explained.

That in all such cases of species-distribution these principles have played an important part in the species-formation, appears to be rendered further probable from the suddenness of transition on the area occupied by contiguous species, as well as from the completeness of it--i. e. the absence of connecting forms. For these facts combine to testify that the transition was originally due to that particular change in the reproductive systems of the forms concerned, which still enables those forms to "interlock" without intercrossing. On the other hand, neither of these facts appears to me compatible with the theory of species-formation by natural selection alone.

But this leads us to another general fact, also mentioned by Darwin, and well recognized by all naturalists, namely, that closely allied species, or species differing from one another in trivial details, usually occupy contiguous areas; or, conversely stated, that contiguity of geographical position is favourable to the appearance of species closely allied to one another. Now, the large body of facts to which I here allude, but need not at present specify, appear to me to constitute one of the strongest of all my arguments in favour of physiological selection. Take, for instance, a large continental area, and follow across it a chain of species, each link of which differs from those on either side of it by the minute and trivial distinctions of a secondary kind, but all the links of which differ from one another in respect of the primary distinction, so that no one member of the series is perfectly fertile with any other member. Can it be supposed that in every case this constant primary distinction has been superinduced by the secondary distinctions, distributed as they are over different parts of all these kindred organisms, and yet nowhere presenting any but a trifling amount of morphological change?

For my own part, I cannot believe--any more than Darwin could believe--that all these numerous, diverse, and trivial changes have always had the accidental effect of inducing the same peculiar change in the reproductive system, and so producing it without any reference to the process of specific divergence. Nor can I believe, as Darwin incidentally and provisionally suggested, that prolonged exposure to uniform conditions of life have so generally induced an equally meaningless result. I can only believe that all the closely allied species inhabiting our supposed continent, and differing from one another in so many and such divers points of small detail, are merely so many records of the fact that selective fertility has arisen among their ancestry, and has thus given as many opportunities for the occurrence of morphological differentiations as it has furnished cases of efficient isolation. Of course, I do not deny that many, or probably most, of these trivial morphological differentiations have been produced by natural selection on account of their utility: I merely deny that they could have been so produced on this common area, but for the sexual isolation with which every distinct set of them is now found to be associated.

It will be at once apparent that a study of the topographical distribution of organic types is of even more importance for us than a study of their geographical distribution. For while the former study is conducted, as it were, with a low power of our observing microscope, the latter is conducted with a high power. The larger facts of geographical distribution yield, indeed, all the general characters which we might expect them to yield, on the theory that divergence of specific types on common areas has been in chief part determined by physiological conditions. But for the purpose of testing this theory in a still more exacting manner, it is of the first importance to consider the more detailed facts of topographical distribution, since we here come to closer quarters with the problem of specific differentiation. Therefore, as we have already considered this problem under the most general points of view, we will now consider it under more special points of view.

Hence, where the question is whether selective fertility has played any large or general part in the differentiation of specific types, the best criterion we can apply is to ascertain whether it is a general rule that closely allied species occur in intimate association, so that their individual members constitute, as it were, a single population, or, on the other hand, whether they occur rather on different sides of physical barriers. If they occur intimately associated, the form of homogamy to which their differentiation was due must have presumably been the physiological form; whereas, if they are proved to be correlated with physical barriers, the form of homogamy which was concerned in their differentiation must presumably have been the geographical form.

Now, at first this consideration was a trouble to me, because Moritz Wagner had strenuously argued--and supported his argument by a considerable wealth of illustration--that allied species are always found correlated with physical barriers or discontinuous areas. Weismann's answer, indeed, had shown that Wagner's statement was much too general: nevertheless, I was disappointed to find that so much could be said in favour of the geographical form of isolation where closely allied species are concerned. Subsequently, however, I read the writings of N?geli on this subject, and in them I find a very different state of matters represented.

But if the idea of selective fertility had ever occurred to N?geli as a form of segregation which gives rise to specific differentiation, I can have no doubt that so astute and logical a thinker would have perceived that his whole indictment against natural selection was answered. For it is incredible that he should not have perceived how this physiological form of homogamy would perform exactly the same function on a continuous area, as he allowed that "isolation" does on a discontinuous one.

After stating the theory of natural selection, he says that if the theory is a true explanation of the origin of specific forms, it ought to follow that

two closely allied forms, derived the one from the other, would necessarily occupy two different geographical areas , since otherwise they would soon become blended. Until they had already become sufficiently consolidated as distinct species to render mutual intercrossing highly improbable, they could not be intermingled without disadvantage . Had Darwin endeavoured to support his hypothesis by facts, he would, at least in the vegetable kingdom, have found little to favour his cause. I can cite many hundreds of cases, in which species in every stage of development have been found closely mingling with one another, and not in any way isolated. Therefore, I do not think that one can rightly speak of natural selection in the Darwinian sense in the vegetable kingdom; and, in my estimation, there is a great difference between the formation of species by nature and the production of stock by a breeder.... .

Of the two kinds of distribution , Synoicy is by far the most usual in nature. I reckon that out of a hundred allied vegetable forms, at least ninety-five would be found to be synoical .

FURTHER EVIDENCES OF PHYSIOLOGICAL SELECTION.

I shall begin with some further quotations from N?geli.

Species may be synoical at all stages of relationship. We come across varieties, scarcely distinguishable from one another, growing in the same locality ; again, we meet other varieties more accentuated , other varieties even more marked, which might almost be elevated to the rank of species , or forms still more distinct, up to well-defined species. I could enumerate endless examples at all stages.

Not to multiply quotations to the same general effect, I will supply but one other, referring to a particular case.

Now it is clear, without comment, that all this is exactly as it ought to be, if allied species have been differentiated on common areas by selective fertility. For if, as N?geli elsewhere says, "one meets forms in nature associated with one another, and severally distinguished by every possible degree of differentiation," not only as N?geli adds, does this general fact lead to the inference that species are developed when plants grow intimately associated together; but as certainly it leads to the further inference that such development must be due to a prior development of cross-infertility between the diverging varietal forms, cross-infertility which is therefore afterwards so characteristic of the allied species, when these are found, in their fully differentiated condition, still occupying the same area in large and intimately mingled populations.

To my mind there could not be any inference more strongly grounded than this, because, with the one exception of the physiological form, no other form of homogamy can be conceived which shall account for the origin and permanence of these synoical varieties, in all degrees of differentiation up to well-defined synoical species. Least of all, as we have seen, can natural selection alone have had anything to do with such a state of matters; while, as we have likewise seen, in all its details it is exactly the state of matters which the theory of physiological selection requires.

The points which are of importance to evolutionists--and of the first order of importance in the present connexion--may be briefly summarized as follows:--

Now, it is self-evident that every one of these seven points is exactly what the theory of physiological selection requires, while there is not one of them which it does not require. For if the theory be sound, we should expect to find large numbers of species belonging to numerous and varied orders of plants presenting constant varieties on common areas; we should expect this to be a highly general, though not a universal, rule; and we should expect it to apply only to species which are indigenous. Moreover, we should expect these varieties, although but slightly differentiated morphologically, to present a great differentiation physiologically--and this in the special direction of selective fertility, combined, of course, with heredity.

We have now to consider the bearing of what is called "prepotency" on the theory of physiological selection.

Speaking of the vast number of species of Compositae, Darwin says:--

There can be no doubt that if the pollen of all these species could be simultaneously or successively placed on the stigma of any one species, this one would elect with unerring certainty its own pollen. This elective capacity is all the more wonderful, as it must have been acquired since the many species of this great group of plants branched off from a common progenitor.

Darwin is here speaking of elective affinity in its fully developed form, as absolute cross-sterility between fully differentiated species. But we meet with all lower degrees of cross-infertility--sometimes between "incipient species," or permanent varieties, and at other times between closely allied species. It is then known as "prepotency" of the pollen belonging to the same variety or species over the pollen of the other variety or species, when both sets of pollen are applied to the same stigma. Although in the absence of the prepotent pollen the less potent will fertilize the seed, yet, such is the appetency for the more appropriate pollen, that even if this be applied to the stigma some considerable time after the other, it will outstrip or overcome the other in fertilizing the ovules, and therefore produce the same result on the next generation as if it had been applied to the mother plant without any admixture of the less potent pollen, although in some cases such incipient degrees of cross-infertility are further shown by the number or quality of the seeds being fewer or inferior.

Now, in different varieties and in different allied species, all degrees of such prepotency have been noticed by many observers, from the faintest perceptible amount up to complete impotency of the alien pollen--when, of course, there is absolute sterility between the two varieties or allied species. The inference is obvious. In this graduated scale of prepotency--beginning with an experimentally almost imperceptible amount of sexual differentiation between two varieties, and ending in an absolute partitioning of two allied species--we have the only remaining fact that is required to complete the case in favour of the present theory. We are here brought back to the very earliest stages of physiological differentiation or to the stages which lie behind Jordan's "Physiological Species"; and therefore, when taken in conjunction with his results, the phenomena of prepotency may be said to give us the complete and final demonstration of one continuous development, which, beginning in an almost imperceptible amount of cross-infertility, ends in absolute cross-sterility. The "elective capacity" to which Darwin alludes as having been "acquired" by all the species of Compositae since they "branched off from a common progenitor," is thus seen among innumerable other species actually in process of acquisition; and so we can perfectly well understand, what is otherwise unintelligible, that closely allied species of plants occur, in ninety-five per cent. of cases, intimately associated on common areas, while exhibiting towards one another the character of mutual sterility.

But more than this. The importance of the widespread phenomena of prepotency to the theory of physiological selection does not consist merely in thus supplying the last link in the chain of evidence touching the origin of species by selective fertility, or "elective capacity." These phenomena are of further importance as showing how in plants, at all events, physiological selection appears to be frequently capable of differentiating specific types without the necessary assistance of any other form of homogamy. In my original statement of the theory, I was careful to insist upon the great value, as differentiating agents, of even small degrees of other forms of homogamy when co-operating with physiological selection. But I also stated my belief that in many cases selective fertility is presumably of itself capable of splitting a specific type; and the reason why I still believe this is, that I do not otherwise understand these phenomena of prepotency. I cannot believe that in all the innumerable cases where they arise, they have been super-induced by some prior morphological changes going on in some other part of the organism, or by "prolonged exposure to uniform conditions of life," on the part of two well-nigh identical forms which have arisen intimately commingled in exactly the same environment, and under the operation of a previously universal intercrossing. Even if such a thing could be imagined as happening occasionally, I feel it difficult to imagine that it can happen habitually, and yet this view must be held by those who would attribute prepotency to natural selection.

It must never be forgotten that the relatively enormous changes as to size, structure, habit, &c., which are presented by our domesticated plants as results of artificial selection, do not entail the physiological character of cross-sterility in any degree, save possibly in some small number of cases. Although in wild species any correspondingly small percentage of cases would doubtless be the ones to survive on common areas, still it is surely incredible that such an accidental association between natural selection and cross-infertility is so habitually the means of specific differentiation as the facts of prepotency would necessarily demand.

Moreover, this view of the matter is still further corroborated by certain other facts and considerations. For example, the phenomena of prepotency are found to occur when the two forms occupy a common area, i.e. are growing intermingled with one another. Therefore, but for this physiological differentiation, there could be absolutely nothing to prevent free intercrossing. Yet the fact that hybrids are so comparatively rare in a state of nature--a fact which Sir Joseph Hooker has pointed out to me as otherwise inexplicable--proves the efficacy of even a low degree of such differentiation in preventing the physiologically-differentiated forms from intercrossing. Even in cases where there is no difficulty in producing artificial hybrids or mongrels between species or varieties growing on common areas, it is perfectly astonishing what an extremely small percentage of the hybrid or mongrel forms are found to occur in nature. And there can be no question that this is due to the very efficient manner in which prepotency does its work--efficient, I mean, from the point of view of the new theory; for upon any other theory prepotency is a meaningless phenomenon, which, notwithstanding its frequent occurrence, plays no part whatever in the process of organic evolution.

I attach considerable importance to the phenomena of prepotency in view of the contrast which is presented between plants and animals in the relation of their species to physical barriers. For animals--and especially the higher animals--appear to depend for their specific differentiations upon such barriers much more than in the case with plants. This is no more than we should expect; for, in accordance with our theory, selective fertility is not so likely to work alone in the case of the higher animals which mate together, as in plants which are fertilized through the agency of wind or insects. In the former case there is no opportunity given for the first rise of cross-infertility, in the form of prepotency; and even where selective fertility has gained a footing in other ways, the chances against the suitable mating of "physiological complements" must be much greater than it is in the latter case. Hence, among the higher animals, selective fertility ought much more frequently to be found in association with other forms of homogamy than it is among plants. And this is exactly what we find. Thus it seems to me that this contrast between the comparative absence and presence of physical barriers, where allied species of plants and of higher animals are respectively concerned, is entitled to be taken as a further corroboration of our theory. For while it displays exactly such a general correlation as this theory would expect, the correlation is one which cannot possibly be explained on any other theory. It is just where physiological selection can be seen to have the best opportunity of acting that we find the most unequivocal evidence of its action; while, on the other hand, it is just where it can be seen to have the least opportunity of asserting itself that we find it most associated with, and therefore assisted by, other forms of homogamy, i. e. not only geographical isolation, but also by sexual preference in pairing, and the several other forms of homogamy, which Mr. Gulick has shown to arise in different places as the result of intelligence.

Hitherto I have been considering, from the most general point of view, the most widespread facts and broadest principles which serve to substantiate the theory of physiological selection. I now pass to the consideration of one of those special cases in which the theory appears to have been successfully applied.

Professor Le Conte has adduced the fossil snails of Steinheim as serving to corroborate the theory of physiological selection.

The facts are these. The snail population of this lake remain for a long time uniform and unchanged. Then a small percentage of individuals suddenly began to vary as regards the form of their shells, and this in two or three directions at the same time, each affected individual, however, only presenting one of the variations. But after all these variations had begun to affect a proportionally large number of individuals, some individuals occur in which two or more of the variations are blended together, evidently, as Weismann says, by intercrossing of the varieties so blended. Later still, both the separate varieties and their blended progeny became more and more numerous, and eventually a single blended type, comprising in itself all the initial varieties, supplanted the parent form. Then another long period of stability ensued until another eruption of new variations took place; and these variations, after having affected a greater and greater number of individuals, eventually blended together by intercrossing and supplanted their parent form. So the process went on, comparatively short periods of variation alternating with comparatively long periods of stability, the variations, moreover, always occurring suddenly in crops, then multiplying, blending together, and in their finally blended type eventually supplanting their parent form.

Share on: WhatsApp @Hash Facebook Tweet