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|08 - Hybridism||08-07 - Fertility of varieties when crossed and of their mongrel offspring not universal||10||
Fertility of Varieties when crossed, and of their Mongrel off-spring.
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.
I fully admit that this is almost invariably the case.
But if we look to varieties produced under nature, we are immediately involved in hopeless difficulties; for if two hitherto reputed 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 Gaertner 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 everyone, 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 the fertility of domestic varieties less remarkable than at first appears. It can, in the first 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 second 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 conditions of life.
Lastly, and this seems to me by far the most important consideration, new races of animals and plants are produced under domestication 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 difference correlated with the reproductive system.
He supplies his several varieties with the same food; 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 organization, 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 system in 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 invariably fertile when intercrossed.
But it seems to me 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.
Gaertner 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 fertilized 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 Gaertner 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 fertilization 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 experimentised 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 Gaertner: namely, that yellow and white varieties of the same species of Verbascum when intercrossed produce less seed, than do either coloured varieties when fertilized 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 same 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 observations which I have made on certain varieties of hollyhock, I am inclined to suspect that they present analogous facts.
Koelreuter, 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 experimentised 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 or 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 these 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 to be of universal occurrence, or to form a fundamental distinction between varieties and species.
The general fertility of varieties does not seem to me sufficient to overthrow the view which I have taken with respect to the very general, but not invariable, sterility of first crosses and of 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.