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Hence it seems that, on the one hand, slight changes in the conditions of life benefit all organic beings, and on the other hand, that slight crosses, that is crosses between the males and females of the same species which have varied and become slightly different, give vigour and fertility to the offspring. But we have seen that greater changes, or changes of a particular nature, often render organic beings in some degree sterile ; and that greater crosses, that is crosses between males and females which have become widely or specifically different, produce hybrids which are generally sterile in some degree. I cannot persuade myself that this parallelism is an accident or an illusion. Both series of facts seem to be connected together by some common but unknown bond, which is essentially related to the principle of life.
Fertility of Varieties when crossed, and of their Mongrel offspring.-It may be urged, as a most forcible argument, that there must be some essential distinction between species and varieties, and that there must be some error in all the foregoing remarks, inasmuch as varieties, however much they may differ from each other in external appearance, cross with perfect facility, and yield perfectly fertile offspring. With some exceptions, presently to be given, I fully admit that this is very generally the rule. But the subject is surrounded by difficulties, for looking to varieties produced under nature, if two forms hitherto reputed 'to be varieties be found in any degree sterile together, they are at once ranked by most naturalists as species. For instance, the blue and red pimpernel, the primrose and cowslip, which are considered by many of our best botanists as varieties, are said by Gärtner not to be quite fertile when crossed, and he consequently ranks them as undoubted species. If we thus argue in a circle, the fertility of all varieties produced under nature will assuredly have to be granted.'
If we turn to varieties, produced, or supposed to have been produced, under domestication, we are still involved in doubt. For when it is stated, for instance, that the German Spitz dog unites more easily than other dogs with foxes, or that certain South American indigenous domestic dogs do not readily cross with European dogs, the explanation which will occur to every one, and probably the true one, is that these dogs have descended from several aboriginally distinct species. Nevertheless the perfect
fertility of so many domestic varieties, differing widely from each other in appearance, for instance of the pigeon or of the cabbage, is a remarkable fact; more especially when we reflect how many species there are, which, though resembling each other most closely, are utterly sterile when intercrossed. Several considerations, however, render this fertility of domestic varieties less remarkable. In the first place we must remember how ignorant we are regarding the precise cause of sterility, both when species are crossed and when species are removed from their natural conditions. On this latter head I have not had space to adduce the many remarkable facts which could have been given; with respect to sterility from crossing, reflect on the difference in the result of reciprocal crosses,--reflect on the singular cases in which a plant can be more easily fertilised by foreign pollen than by its own. When we think over such cases, and on that of the differently coloured varieties of Verbascum presently to be given, we must feel how ignorant we are, and how little likely it is that we should understand why certain forms are fertile and other forms are sterile when crossed. It can, in the second place, be clearly shown that mere external dissimilarity between two species does not determine their greater or lesser degree of sterility when crossed ; and we may apply the same rule to domestic varieties. In the third place, some eminent naturalists believe that a long course of domestication tends to eliminate sterility in the successive generations of hybrids which were at first only slightly sterile ; and if this be so, we surely ought not to expect to find sterility both appearing and disappearing under 'nearly the same domestic conditions of life. Lastly, and this seems to me the most important consideration, new races of animals and plants are produced under domestication chiefly by man's methodical and unconscious power of selection, for his own use and pleasure: he neither wishes to select, nor could select, slight differences in the reproductive system, or other constitutional differences correlated with the reproductive system. Domestic productions are less closely adapted to climate and to the other physical conditions of the countries which they inhabit than are those in a state of nature, for they can generally be removed to other and differently constituted countries with entire impunity. Man supplies his several varieties with the same food; he treats them in nearly the same manner, and does not wish to alter their general habits of life. Nature
acts uniformly and slowly during vast periods of time on the whole organisation, in any way which may be for each creature's own good; and thus she may, either directly, or more probably indirectly, through correlation, modify the reproductive systems of the several descendants from any one species. Seeing this difference in the process of selection, as carried on by man and nature, we need not be surprised at some difference in the result.
I have as yet spoken as if the varieties of the same species were almost invariably fertile when intercrossed. But it is impossible to resist the evidence of the existence of a certain amount of sterility in the few following cases, which I will briefly abstract. The evidence is at least as good as that from which we believe in the sterility of a multitude of species. The evidence is, also, derived from hostile witnesses, who in all other cases consider fertility and sterility as safe criterions of specific distinction. Gärtner kept during several years a dwarf kind of maize with yellow seeds, and a tall variety with red seeds, growing near each other in his garden; and although these plants have separated sexes, they never naturally crossed. He then fertilised thirteen flowers of the one with the pollen of the other ; but only a single head produced any seed, and this one head produced only five grains. Manipulation in this case could not have been injurious, as the plants have separated sexes. No one, I believe, has suspected that these varieties of maize are distinct species ; and it is important to notice that the hybrid plants thus raised were themselves perfectly fertile ; so that even Gärtner did not venture to consider the two varieties as specifically distinct.
Girou de Buzareingues crossed three varieties of gourd, which like the maize has separated sexes, and he asserts that their mutual fertilisation is by so much the less easy as their differences are greater. How far these experiments may be trusted, I know not; but the forms experimented on are ranked by Sagaret, who mainly founds his classification by the test of infertility, as varieties.
The following case is far more remarkable, and seems at first quite incredible ; but it is the result of an astonishing number of experiments made during many years on nine species of Verbascum, by so good an observer and so hostile a witness as Gärtner: namely, that yellow and white varieties of the same species of Verbascum when intercrossed produce less seed, than do either coloured
varieties when fertilised with pollen from their own coloured flowers. Moreover, he asserts that when yellow and white varieties of one species are crossed with yellow and white varieties of a distinct species, more seed is produced by the crosses between the similarly coloured flowers, than between those which are differently coloured. Yet these varieties of Verbascum present no other difference besides the mere colour of the flower; and one variety can sometimes be raised from the seed of the other.
From experiments which I have made on certain varieties of hollyhock, I am inclined to suspect that they present analogous facts.
Kölreuter, whose accuracy has been confirmed by every subsequent observer, has proved the remarkable fact, that one variety of the common tobacco is more fertile, when crossed with a widely distinct species, than are the other varieties. He experimented on five forms, which are commonly reputed to be varieties, and which he tested by the severest trial, namely, by reciprocal crosses, and he found their mongrel offspring perfectly fertile. But one of these five varieties, when used either as father mother, and crossed with the Nicotiana glutinosa, always yielded hybrids not so sterile as those which were produced from the four other varieties when crossed with N. glutinosa. Hence the reproductive system of this one variety must have been in some manner and in some degree modified.
From thes facts ; from the great difficulty of ascertaining the infertility of varieties in a state of nature, for a supposed variety if infertile in any degree would generally be ranked as species ; from man selecting only external characters in the production of the most distinct domestic varieties, and from not wishing or being able to produce recondite and functional differences in the reproductive system ; from these several considerations and facts, I do not think that the very general fertility of varieties can be proved either to be of universal occurrence, or to form a fundamental distinction between varieties and species. The general fertility of varieties, considering how entirely ignorant we are on the causes of both fertility and sterility, does not seem to me sufficient to overthrow the view taken with respect to the very general, but not invariable, sterility of first between species and of their hybrids, namely, that it is
not a special endowment, but is incidental on slowly acquired modifications, more especially in the reproductive systems of the forms which are crossed.
Hybrids and Mongrels compared, independently of their fertility.-Independently of the question of fertility, the offspring of species when crossed and of varieties when crossed may be compared in several other respects, Gärtner, whose strong wish it was to draw a distinct line between species and varieties, could find very few, and, as it seems to me, quite unimportant differences between the so-called hybrid offspring of species, and the so-called mongrel offspring of varieties. And, on the other hand, they agree most closely in very many important respects.
I shall. here discuss this subject with extreme brevity. The most important distinction is, that in the first generation mongrels are more variable than hybrids ; but Gärtner admits that hybrids from species which have long been cultivated are often variable in the first generation; and I have myself seen striking instances of this fact. Gärtner further admits that hybrids between very closely allied species are more variable than those from very distinct species; and this shows that the difference in the degree of variability graduates away. When mongrels and the more fertile hybrids are propagated for several generations an extreme amount of variability in their offspring is notorious ; but some few cases both of hybrids and mongrels long retaining uniformity of character could be given. The variability, however, in the successive generations of mongrelsis, perhaps, greater than in hybrids.
This greater variability of mongrels than of hybrids does not seem to me at all surprising. For the parents of mongrels are varieties, and mostly domestic varieties (very few experiments having been tried on natural varieties), and this implies in most cases that there has been recent variability; and therefore we might expect that such yariability would often continue and be superadded to that arising from the mere act of crossing. . The slight degree of variability in hybrids from the first cross or in the first generation, in contrast with their extreme variability in the succeeding generations, is a curious fact and deserves attention. For it bears on and corroborates the view which I have taken on the cause of ordinary variability; namely, that it is due to the re