CHAPTER VIII. HYBRIDISM. Distinction between the sterility of first crosses and of hybridsSterility various in degree, not universal, affected by close interbreeding, removed by domestication-Laws governing the sterility of hybrids-Sterility not a special endowment, but incidental on other differences, not accumulated by natural selectionCauses of the sterility of first crosses and of hybrids-Parallelism between the effects of changed conditions of life and of crossing -Dimorphism and trimorphism-Fertility of varieties when crossed and of their mongrel offspring not universal-Hybrids and mongrels compared independently of their fertility-Summary. THE view generally entertained by naturalists is that species, when intercrossed, have been specially endowed with the quality of sterility, in order to prevent the confusion of all organic forms. This view certainly seems at first probable, for species within the same country could hardly have kept distinct had they been capable of crossing freely. The importance of the fact that first crosses between distinct species and hybrids are very generally sterile, has, I think, been much underrated by some late writers. On the theory of natural selection the case is especially important, inasmuch as this sterility can hardly have been increased by the continued preservation of successive, profitable degrees of sterility. But to this subject I shall have to return, and I hope ultimately to show that this sterility is neither a specially acquired nor endowed quality, but is incidental on other acquired and little-known differences in the reproductive systems of the parent-species. In treating this subject, two classes of facts, to a large extent fundamentally different, have generally been confounded together; namely, the sterility of two species when first crossed, and the sterility of the hybrids produced from them. Pure species have of course their organs of reproduction in a perfect condition, yet when intercrossed they produce either few or no offspring. Hybrids, on the other hand, have their reproductive organs functionally impotent, as may be clearly seen in the state of the male element in both plants and animals; though the formative organs themselves are perfect in structure, as far as the microscope reveals. In the first case the two sexual elements which go to form the embryo are perfect; in the second case they are either not at all developed, or are imperfectly developed. This distinction is important, when the cause of the sterility, which is common to the two cases, has to be considered. The distinction has probably been slurred over, owing to the sterility in both cases being looked on as a special endowment, beyond the province of our reasoning powers. The fertility of varieties, that is of the forms known or believed to have descended from common parents, when intercrossed, and likewise the fertility of their mongrel offspring, is, with reference to my theory, of equal importance with the sterility of species; for it seems to make a broad and clear distinction between varieties and species. Degrees of Sterility.-First, for the sterility of species when crossed and of their hybrid offspring. It is impossible to study the several memoirs and works of those two conscientious and admirable observers, Kölreuter and Gärtner, who almost devoted their lives to this subject, without being deeply impressed with the high generality of some degree of sterility. Kölreuter makes the rule universal; but then he cuts the knot, for in ten cases in which he found two forms, considered by most authors as distinct species, quite fertile together, he unhesitatingly ranks them as varieties. Gärtner, also, makes the rule equally universal; and he disputes the entire fertility of Kölreuter's ten cases. But in these and in many other cases, Gärtner is obliged carefully to count the seeds, in order to show that there is any degree of sterility. He always compares the maximum number of seeds produced by two species when crossed and by their hybrid offspring, with the average number produced by both pure parent-species in a state of nature. But a serious cause of error seems to me to be here introduced: a plant, to be hybridised, must be castrated, and, what is often more important, must be secluded in order to prevent pollen being brought to it by insects from other plants. Nearly all the plants experimented on by Gärtner were potted, and were kept in a chamber in his house. That these processes are often injurious to the fertility of a plant cannot be doubted; for Gärtner gives in his table about a score of cases of plants which he castrated, and artificially fertilised with their own pollen, and (excluding all cases such as the Leguminosa, in which there is an acknowledged difficulty in the manipulation) half of these twenty plants had their fertility in some degree impaired. Moreover, as Gärtner repeatedly crossed some forms, such as the common red and blue pimpernels (Anagallis arvensis and coerulea), which the best botanists rank as varieties, and found them absolutely sterile, we may doubt whether many other species are really so sterile, when intercrossed, as he believed. It is certain, on the one hand, that the sterility of various species when crossed is so different in degree and graduates away so insensibly, and, on the other hand, that the fertility of pure species is so easily affected by various circumstances, that for all practical purposes it is most difficult to say where perfect fertility ends and sterility begins. I think no better evidence of this can be required than that the two most experienced observers who have ever lived, namely, Kölreuter and Gärtner, should have arrived at diametrically opposite conclusions in regard to the very same species. It is also most instructive to compare— but I have not space here to enter on details—the evidence advanced by our best botanists on the question. whether certain doubtful forms should be ranked as species or varieties, with the evidence from fertility adduced by different hybridisers, or by the same author, from experiments made during different years. It can thus be shown that neither sterility nor fertility affords any clear distinction between species and varieties; but that the evidence from this source graduates away, and is doubtful in the same degree as is the evidence derived from other constitutional and structural differences. In regard to the sterility of hybrids in successive generations; though Gärtner was enabled to rear some hybrids, carefully guarding them from a cross with either pure parent, for six or seven, and in one case for ten generations, yet he asserts positively that their fertility never increased, but generally decreased greatly and suddenly. With respect to this decrease, it may first be noticed that, when any deviation in structure or constitution is common to both parents, this is often transmitted in an augmented degree to the offspring; and both sexual elements in hybrid plants are already affected in some degree. But I believe in nearly all these cases, that the fertility has been diminished by an independent cause, namely, by too close interbreeding. I have collected so large a body of facts, showing on the one hand that an occasional cross with a distinct individual or variety increases the vigour and fertility of the offspring, and on the other hand that very close interbreeding lessens their vigour and fertility, that I must admit the correctness of this almost universal belief amongst breeders. Hybrids are seldom raised by experimentalists in great numbers; and as the parentspecies, or other allied hybrids, generally grow in the same garden, the visits of insects must be carefully prevented during the flowering season: hence hybrids will generally have to be fertilised during each generation by their own individual pollen; and this would probably be injurious to their fertility, already lessened by their hybrid origin. I am strengthened in this conviction by a remarkable statement repeatedly made by Gärtner, namely, that if even the less fertile hybrids be artificially fertilised with hybrid pollen of the same kind, their fertility, notwithstanding the frequent ill effects from manipulation, sometimes decidedly increases, and goes on increasing. Now, in the process of artificial fertilisation, pollen is as often taken by chance (as I know from my own experience) from the anthers of another flower, as from the anthers of the flower itself. which is to be fertilised; so that a cross between two flowers, though probably often on the same plant, would be thus effected. Moreover, whenever complicated experiments are in progress, so careful an observer as Gärtner would have castrated his hybrids, and this would have insured in each generation a cross with pollen from a distinct flower, either from the same plant or from another plant of the same hybrid nature. And thus, the strange fact of the increase of fertility in the successive generations of artificially fertilised hybrids |