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distinct, species, when inhabiting distant parts of the world and living under considerably different conditions of life, yet often retaining nearly the same instincts. For instance, we can understand, on the principle of inheritance, how it is that the thrush of tropical South America lines its nest with mud, in the same peculiar manner as does our British thrush; how it is that the Hornbills of Africa and India have the same extraordinary instinct of plastering up and imprisoning the females in a hole in a tree, with only a small hole left in the plaster through which the males feed them and their young when hatched; how it is that the male wrens (Troglodytes) of North America build “cock-nests,” to roost in, like the males of our Kitty-wrens,—a habit wholly unlike that of any other known bird. Finally, it may not be a logical deduction, but to my imagination it is far more satisfactory to look at such instincts as the young cuckoo ejecting its foster-brothers, ants making slaves, the larvae of ichneumonidae feeding within the live bodies of caterpillars, not as specially endowed or created instincts, but as small consequences of one general law leading to the advancement of all organic beings, namely, multiply, vary, let the strongest live and the weakest die.
Distinction between the sterility of first crosses and of hybrids–Sterility 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 selection—Causes 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 o, of their fertility—Summary.
THE view commonly entertained by naturalists is that species, when intercrossed, have been specially endowed with sterility, in order to prevent their confusion. This view certainly seems at first highly probable, for species living together could hardly have been kept distinct had they been capable of freely crossing. The subject is in many ways important for us, more especially as the sterility of species when first crossed, and that of their hybrid offspring, cannot have been acquired, as I shall show, by the preservation of successive profitable degrees of sterility. It is an
incidental result of 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; namely, the sterility of 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 probably has 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 be descended from common parents, when crossed, 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 first crossed, and the maximum produced by their hybrid offspring, with the average number produced by both pure parent-species in a state of nature. But causes of serious error here intervene: 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 Leguminosae, 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 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, arrived at diametrically opposite conclusions in regard to some of the very same forms. It is also most instructive to compare—but I have not space here to enter into 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 observer from experiments made during different years. It can thus be shown that neither sterility nor fertility affords any certain distinction between species and varieties. 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 increases, but generally decreases 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 that their fertility has been diminished in nearly all these cases by an independent cause, namely, by too close interbreeding. I have made so many experiments and collected so many 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 cannot doubt the correctness of this conclusion. Hybrids are seldom raised by experimentalists in great numbers; and as the parent-species, or other allied hybrids, generally grow in the same garden, the visits of insects must be carefully prevented during the flowering season: hence hybrids, if left to themselves, will generally be fertilised during each generation by pollen from the same flower; 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 ensured 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 an increase of fertility in the successive generations of artificially fertilised hybrids, in contrast with those spontaneously self-fertilised, may, as I believe, be accounted for by too close interbreeding having been avoided. Now let us turn to the results arrived at by a third most experienced hybridiser, namely, the Hon. and Rev. W. Herbert. He is as emphatic in his conclusion that some hybrids are perfectly fertile—as fertile as the pure parent-species—as are Kölreuter and Gärtner that some degree of sterility between distinct species is a universal law of nature. He experimented on some of the very same species as did Gärtner. The difference in their results may, I think, be in part accounted for by Herbert's great horticultural skill, and by his having hot-houses at his command. Of his many important statements I will here give only a single one as an example, namely, that “every ovule in a pod of Crinum capense fertilised by C. revolutum produced a plant, which I never saw to occur in a case of its natural fecundation.” So that here we have perfect or even more than commonly perfect fertility, in a first cross between two distinct species.
This case of the Crinum leads me to refer to a singular fact, namely, that individual plants of certain species of Lobelia, Verbascum and Passiflora, can easily be fertilised by pollen from a distinct species, but not by pollen from the same plant, though this pollen can be proved to be perfectly sound by fertilising other plants or species. In the genus Hippeastrum, in Corydalis as shown by Professor Hildebrand, in various orchids as shown by Mr. Scott and Fritz Müller, all the individuals are in this peculiar condition. So that with some species, certain abnormal individuals, and in other species all the individuals, can actually be hybridised much more readily than they can be fertilised by pollen from the same individual plant! To give one instance, a bulb of Hippeastrum aulicum produced four flowers; three were fertilised by Herbert with their own pollen, and the fourth was subsequently fertilised by the pollen of a compound hybrid descended from three distinct species: the result was that “the ovaries of the three first flowers soon ceased to grow, and after a few days perished entirely, whereas the pod impregnated by the pollen of the hybrid made vigorous growth and rapid progress to maturity, and bore good seed, which vegetated freely.” Mr. Herbert tried similar experiments during many years, and always with the same result. These cases serve to show on what slight and mysterious causes the lesser or greater fertility of a species sometimes depends.
The practical experiments of horticulturists, though not made with scientific precision, deserve some notice. It is notorious in how complicated a manner the species of Pelargonium, Fuchsia, Calceolaria, Petunia, Rhododendron, &c., have been crossed, yet many of these hybrids seed freely. For instance, Herbert asserts that a hybrid from Calceolaria integrifolia and plantaginea, species most widely dissimilar in general habit, “reproduces itself as perfectly as if it had been a natural species from the mountains of Chili.” I have taken some pains to ascertain the degree of fertility of some of the complex crosses of Rhododendrons, and I am assured that many of them are perfectly fertile. Mr. C. Noble, for instance, informs me that he raises stocks for grafting from a hybrid between Rhod. ponticum and catawbiense, and that this hybrid “seeds as freely as it is possible to imagine.” Had hybrids, when fairly treated, always gone on decreasing in fertility in each successive generation, as Gärtner believed to be the case, the fact would have been notorious to nurserymen. Horticulturists raise large beds of the same hybrid, and such alone are fairly treated, for by insect agency the several individuals are allowed to cross freely with each other, and the injurious influence of close interbreeding is thus prevented. Any one may readily convince himself of the efficiency of insect-agency by