Hybridism

“ And now we see that this same conclusion is independently and strongly fortified by the consideration of the illegitimate unions of trimorphic and dimorphic heterostyled plants. In so complex and obscure a subject as hybridism it is no slight gain to arrive at a definite conclusion, namely, that we must look exclusively to functional differences in the sexual elements, as the cause of the sterility of species when first crossed and of their hybrid offspring. ”

“ The very important and interesting phenomenon of hybridism also belongs to this law of mixed or amphigonous211 transmission. It alone, when rightly estimated, is quite sufficient to refute the prevailing dogma of the constancy of species. Plants, as well as animals, belonging to quite different species, may sexually mingle with one another and produce descendants which in many cases can again propagate themselves, and that indeed either (more frequently) by mingling with one of the two parental species, or (more rarely) by pure in-breeding, hybrid mixing with hybrid. ”

“ It throws light on the origin of the two sexes and on their separation or union in the same individual, and lastly on the whole subject of hybridism, which is one of the greatest obstacles to the general acceptance and progress of the great principle of evolution. ”

“ A closer scrutiny of the facts, however, makes the term hybridism less isolated and more vague. The words species and genus, and still more subspecies and variety, do not correspond with clearly marked and sharply defined zoological categories, and no exact line can be drawn between the various kinds of crossings from those between individuals apparently identical to those belonging to genera universally recognized as distinct. ”

“ In nearly the same manner as it is admitted by naturalists that hybridism, from inducing sterility, is of service in keeping the forms of life distinct and fitted for their proper places; so, when species are rendered highly variable by changed conditions of life, the free inter-crossing of the varying individuals will tend to keep each form fitted for its proper place in nature ”

“ In the other part of the area, however, where hybridism occurs with perfect freedom, hybrids of various degrees may increase till they equal or even exceed in number the pure species—that is, the incipient species will be liable to be swamped by intercrossing. ”

“ But such failure is usually due to social factors though these writers attribute it to the inborn traits of the half-breeds. Biologists regard hybridism in general as a good thing in the animate world at large and as an important factor in organic evolution. ”

“ It is possible that many wild species have originated in this way, as even Linnæus assumed. At all events, these hybrid species, which can maintain and propagate themselves as well as pure species, prove that hybridism cannot serve in any way to give an absolute definition to the idea of species. ”

“ The chapter on hybridism is most ingenious, able, and instructive. If sterility of crosses is a special original arrangement to prevent the confusion of species by mingling, as is generally assumed, then, since varieties cross readily and their offspring is fertile inter se, there is a fundamental distinction between varieties and species. Mr. Darwin therefore labors to show that it is not a special endowment, but an incidental acquirement. He does show that the sterility of crosses is of all degrees ”

“ The great body of facts of which a condensed account has been given in the present chapter, although from an experimental point of view very insufficient, all point to the general conclusion we have now reached, and afford us a not unsatisfactory solution of the great problem of hybridism in relation to the origin of species by means of natural selection. ”

“ It may well be that the length of time required before an artificially-selected race will exhibit, when interbred with the parent species, phenomena of hybridism similar to those which are witnessed when distinct natural species are interbred—will be fatal to the production of this important line of evidence. ”

“ But if you take members of two distinct species, however similar they may be to each other and make them breed together, you will find a check, with some modifications and exceptions, however, which I shall speak of presently. If you cross two such species with each other, then,—although you may get offspring in the case of the first cross, yet, if you attempt to breed from the products of that crossing, which are what are called HYBRIDS—that is, if you couple a male and a female hybrid—then the result is that in ninety-nine cases out of a hundred you will get no offspring at all ”

“ Nor is it surprising that the facility of effecting a first cross, the fertility of hybrids produced from it, and the capacity of being grafted together—though this latter capacity evidently depends on widely different circumstances—should all run to a certain extent parallel with the systematic affinity of the forms which are subjected to experiment; for systematic affinity attempts to express all kinds of resemblance between all species. ”

“ Perhaps on close inspection you may find in some cases a slight difference, some character being not as fully developed in the hybrid as in the species. But when such is not the case, or where the opportunity for such a closer examination is wanting, a hybrid may easily be taken for a specimen of the pure race. Now take the seeds of your plant and sow them. If you had not supposed it to be hybrid you will be astonished at finding among its progeny some of a wholly different type. You will be led to conclude that you are observing a sudden change in structure such as is usually called a sport. ”

“ But it is not as yet proved that a race ever exhibits, when crossed with another race of the same species, those phenomena of hybridization which are exhibited by many species when crossed with other species. On the other hand, not only is it not proved that all species give rise to hybrids infertile inter se, but there is much reason to believe that, in crossing, species exhibit every gradation from perfect sterility to perfect fertility. ”

“ Why should there often be so great a difference in the result of a reciprocal cross between the same two species? Why, it may even be asked, has the production of hybrids been permitted? to grant to species the special power of producing hybrids, and then to stop their further propagation by different degrees of sterility, not strictly related to the facility of the first union between their parents, seems to be a strange arrangement. ”

“ There are real obstacles (of which I will say more below) in natural conditions to hybrid-breeding between any two species which occur naturally in the same territory. Thus the idea of a species is expanded so as to be not merely "a group of individuals of constant likeness in form and characteristics," but we add to that definition a living or constitutional quality expressed by the words, "which produce fertile offspring by pairing with one another, but do not pair with the members of other species." ”

“ Hybridization differs from ordinary crossing only in degree. A species is only a race which has assumed greater fixity. The purposes and results of crossing within the species and of hybridization of different species are essentially alike. The formation of the new individual by the sexual relation of two parents is in itself a species of crossing, giving each new individual in its degree new traits or new combinations. ”

“ Hybrids from two species which are very difficult to cross, and which rarely produce any offspring, are generally very sterile; but the parallelism between the difficulty of making a first cross, and the sterility of the hybrids thus produced—two classes of facts which are generally confounded together—is by no means strict. There are many cases, in which two pure species can be united with unusual facility, and produce numerous hybrid-offspring, yet these hybrids are remarkably sterile. ”

“ That true hybrids raised from entirely distinct species, though they lose in fertility, often gain in size and constitutional vigour, is certain. It would be superfluous to quote any facts; for all experimenters, Kölreuter, Gärtner, Herbert, Sageret, Lecoq, and Naudin, have been struck with the wonderful vigour, height, size, tenacity of life, precocity, and hardiness of their hybrid productions. ”

“ I well know the large amount of natural as well as artificial hybridisation that goes on. But, on the other hand, there are so many species which either will not cross at all, or produce sterile hybrids, that, taking a general view of all species together, mutual sterility does become by far the most generally distributed single peculiarity—i.e. is the one peculiarity which, more than any other that can be named, is common to numberless species. ”

“ Independently of 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 harmonises perfectly with the view that there is no essential distinction between species and varieties. ”

“ The intermixture of distinct species is guarded against by the aversion of the individuals composing them to sexual union, or by the sterility of the mule offspring. It does not appear that true hybrid races have ever been perpetuated for several generations, even by the assistance of man; for the cases usually cited relate to the crossing of mules with individuals of pure species, and not to the intermixture of hybrid with hybrid. ”

“ The occurrence of hybrids in nature explains the difficulty experienced by botanists in deciding on what is a species, and the widely different limitations of the term adopted by different observers in the case of willows, roses, brambles, &c. The artificial process is practically the same in hybridization as in cross-fertilization, but usually requires more care. ”

“ Others, on the contrary, of our most acute observers, having acquired convincing evidence of natural hybridity in a few cases, have generalized their conclusions; they have supposed natural hybrids to be of constant and frequent occurrence; and they have ascribed to this cause alone the majority of variations from the supposed typical forms of species, or even attributed to original hybridisations the multitude of nearly allied, but constant species, in several of the largest genera. ”

“ This notion, indeed, is wholly at variance with all that we know of hybrid generation; for the phenomena entitle us to affirm, that had the types been at first somewhat distinct, no cross-breeds would ever have been produced, much less those prolific races which we now recognize as distinct species. ”

“ So long as you breed together descendants from a common stock they continue fruitful to each other, without any check. Now, if naturalists cannot produce by selective breeding varieties from a common parentage that are infertile to each other, then it is quite clear that species did not come from varieties by the process of variation preserved by natural selection, since here is a phenomenon existing in connection with species which cannot, to all appearances, be produced by breeding together varieties. ”

“ It is said that such a test is to be found in the phenomena of hybridization—in the results of crossing races as compared with the results of crossing species.
So far as the evidence goes at present, individuals, of what are certainly known to be mere races produced by selection, however distinct they may appear to be, not only breed freely together, but the offspring of such crossed races are also perfectly fertile with one another. ”

“ But is it not possible to apply a test whereby a true species may be known from a mere variety? Is there no criterion of species? Great authorities affirm that there is—that the unions of members of the same species are always fertile, while those of distinct species are either sterile, or their offspring, called hybrids, are so. ”