Darwin, and After Darwin, Volume 3 of 3 / Post-Darwinian Questions: Isolation and Physiological Selection

by George John Romanes

Now the forms of discriminate isolation, or homogamy, are very numerous. When, for example, any section of a species adopts somewhat different habits of life, or occupies a somewhat different station in the economy of nature, homogamy arises within that section. There are forms of homogamy on which Darwin has laid great stress, as we shall presently find. Again, when for these or any other reasons a section of a species becomes in any small degree modified as to form or colour, if the species happens to be one where any psychological preference in pairing can be exercised—as is very generally the case among the higher animals—exclusive breeding is apt to ensue as a result of such preference; for there is abundant evidence to show that, both in birds and mammals, sexual selection is usually opposed to the intercrossing of dissimilar varieties. Once more, in the case of plants, intercrossing of dissimilar varieties may be prevented by any slight difference in their seasons of flowering, of topographical stations, or even, in the case of flowers which depend on insects for their fertilization, by differences in the instincts and preferences of their visitors.

But, without at present going into detail with regard to these different forms of discriminate isolation, there are still two others, both of which are of much greater importance than any that I have hitherto named. Indeed, these two forms are of such immeasurable importance, that were it not for their virtually ubiquitous operation, the process of organic evolution could never have begun, nor, having begun, continued.

The first of these two forms is sexual incompatibility—either partial or absolute—between different taxonomic groups. If all hares and rabbits, for example, were as fertile with one another as they are within their own respective species, there can be no doubt that sooner or later, and on common areas, the two types would fuse into one. And similarly, if the bar of sterility could be thrown down as between all the species of a genus, or all the genera of a family, not otherwise prevented from intercrossing, in time all such species, or all such genera, would become blended into a single type. As a matter of fact, complete fertility, both of first crosses and of their resulting hybrids, is rare, even as between species of the same genus; while as between genera of the same family complete fertility does not appear ever to occur; and, of course, the same applies to all the higher taxonomic divisions. On the other hand, some degree of infertility is not unusual as between different varieties of the same species; and, wherever this is the case, it must clearly aid the further differentiation of those varieties. It will be my endeavour to show that in this latter connexion sexual incompatibility must be held to have taken an immensely important part in the differentiation of varieties into species. But meanwhile we have only to observe that wherever such incompatibility is concerned, it is to be regarded as an isolating agency of the very first importance. And as it is of a character purely physiological, I have assigned to it the name Physiological Isolation; while for the particular case where this general principle is concerned in the origination of specific types, I have reserved the name Physiological Selection.

The other most important form of discriminate isolation to which I have alluded is Natural Selection. To some evolutionists it has seemed paradoxical thus to regard natural selection as a form of isolation; but a little thought will suffice to show that such is really the most accurate way of regarding it. For, as Mr. Gulick says, "Natural selection is the exclusive breeding of those better adapted to the environment: ... it is a process in which the fittest are prevented from crossing with the less fitted, by the exclusion of the less fitted." Therefore it is, strictly and accurately, a mode of isolation, where the isolation has reference to adaptation, and is secured in the most effectual of possible ways—i.e. by the destruction of all individuals whose intercrossing would interfere with the isolation. Indeed, the very term "natural selection" shows that the principle is tacitly understood to be one of isolation, because this name was assigned to the principle by Darwin for the express purpose of marking the analogy that obtains between it and the intentional isolation which is practised by breeders, fanciers, and horticulturists. The only difference between "natural selection" and "artificial selection" consists in this—that under the former process the excluded individuals must necessarily perish, while under the latter they need not do so. But clearly this difference is accidental: it is in no way essential to the process considered as a process of discriminate isolation. For, as far as homogamous breeding is concerned, it can matter nothing whether the exclusion of the dissimilar individuals is effected by separation or by death.

Natural selection, then, is thus unquestionably a form of isolation of the discriminate kind; and therefore, notwithstanding its unique importance in certain respects, considered as a principle of organic evolution it is less fundamental—and also less extensive—than the principle of isolation in general. In other words, it is but a part of a much larger whole. It is but a particular form of a general principle, which, as just shown, presents many other forms, not only of the discriminate, but likewise of the indiscriminate kind. Or, reverting to the terminology of logic, it is a sub-species of the species Homogamy, which in its turn is but a constituent part of the genus Isolation.

So much then for homogamy, or isolation of the discriminate order. Passing on now to apogamy, or isolation of the indiscriminate kind, we may well be disposed, at first sight, to conclude that this kind of isolation can count for nothing in the process of evolution. For if the fundamental importance of isolation in the production of organic forms be due to its segregation of like with like, does it not follow that any form of isolation which is indiscriminate must fail to supply the very condition on which all the forms of discriminate isolation depend for their efficacy in the causing of organic evolution? Or, to return to our concrete example, is it not self-evident that the farmer who separated his stock into two or more parts indiscriminately, would not effect any more change in his stock than if he had left them all to breed together?

Well, although at first sight this seems self-evident, it is in fact untrue. For, unless the individuals which are indiscriminately isolated happen to be a very large number, sooner or later their progeny will come to differ from that of the parent type, or unisolated portion of the previous stock. And, of course, as soon as this change of type begins, the isolation ceases to be indiscriminate: the previous apogamy has been converted into homogamy, with the usual result of causing a divergence of type. The reason why progeny of an indiscriminately isolated section of an originally uniform stock—e.g. of a species—will eventually deviate from the original type is, to quote Mr. Gulick, as follows:—"No two portions of a species possess exactly the same average character, and, therefore, the initial differences are for ever reacting on the environment and on each other in such a way as to ensure increasing divergence as long as the individuals of the two groups are kept from intergenerating[5]." Or, as I stated this principle in my essay on Physiological Selection, published but a short time before Mr. Gulick's invaluable contributions to these topics:—

As a matter of fact, we find that no one individual "is like another all in all"; which is another way of saying that a specific type may be regarded as the average mean of all its individual variations, any considerable departure from this average being, however, checked by intercrossing.... Consequently, if from any cause a section of a species is prevented from intercrossing with the rest of its species, we might expect that new varieties should arise within that section, and that in time these varieties should pass into new species. And this is just what we do find[6].

The name which I gave to this cause of specific change was Independent Variability, or variability in the absence of overwhelming intercrossing. But it now appears to me that this cause is really identical with that which was previously enunciated by Delbœuf. Again, in his important essay on The Influence of Isolation, Weismann concludes, on the basis of a large accumulation of facts, that the constancy of any given specific type "does not arise suddenly, but gradually, and is established by the promiscuous intercrossing of all individuals." From which, he says, it follows, that this constancy must cease so soon as the condition which maintains it ceases—i. e. so soon as intercrossing (Panmixia) between all individuals ceases, or so soon as a portion of a species is isolated from its parent stock. To this principle he assigns the name of Amixia. But Weismann's Amixia differs from my Independent Variability in several important particulars; and on this account I have designedly abstained from adopting his term. Here it is enough to remark that it answers to the generic term Isolation, without reference to the kind of isolation as discriminate or indiscriminate, homogamous or apogamous. On the other hand, my Independent Variability is merely a re-statement of the so-called "Law of Delbœuf," which, in his own words, is as follows:—