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in the great majority of cases, this touchstone for species is wholly inapplicable. The constitution of many wild animals is so altered by confinement that they will not breed even with their own females, so that the negative results obtained from crosses are of no value; and the antipathy of wild animals of different species for one another, or even of wild and tame members of the same species, is ordinarily so great, that it is hopeless to look for such unions in Nature. The hermaphrodism of most plants, the difficulty in the way of insuring the absence of their own or the proper working of other pollen, are obstacles of no less magnitude in applying the test to them. And, in both animals and plants, is superadded the further difficulty, that experiments must be continued over a longtime for the purpose of ascertaining the fertility of the mongrel or hybrid progeny, as well as of the first crosses from which they spring. Not only do these great practical difficulties lie in the way of applying the hybridisation test, but even when this oracle can be questioned, its replies are sometimes as doubtful as those of Delphi. For example, cases are cited by Mr. Darwin, of plants which are more fertile with the pollen of another species than with their own; and there are others, such as certain Fuci, the male element of which will fertilise the ovule of a plant of distinct species, while the males of the latter species are ineffective with the females of the first. So that, in the last-named instance, a physiologist, who should cross the two species in one way, would decide that they were true species; while another, who should cross them in the reverse way, would, with equal justice, according to the rule, pronounce them to be mere races. Several plants, which there is great reason to believe are mere varieties, are almost sterile when crossed; while both animals and plants, which have always been regarded by naturalists as of distinct species, turn out, when the test is applied, to be perfectly fertile. Again, the sterility or fertility of crosses seems to bear no relation to the structural resemblances or differences of the members of any two groups.
Mr. Darwin has discussed this question with singular ability and circumspection, and his conclusions are summed up as follows, at page 276 of his work:
“First crosses between forms sufficiently distinct to be ranked as species, and their hybrids, are very generally, but not universally, sterile. The sterility is of all degrees, and is often so slight that the two most careful experimentalists who have ever lived have come to diametrically opposite conclusions in ranking forms by this test. The sterility is innately variable in individuals of the same species, and is eminently susceptible of favourable and unfavourable conditions. The degree of sterility does not strictly follow systematic affinity, but is governed by several curious and complex laws. It is generally different and sometimes widely disserent, in reciprocal crosses
between the same two species. It is not always equal in degree in a first cross, and in the hybrid produced from this cross. “In the same manner as in grafting trees, the capacity of one species or variety to take on another is incidental on generally unknown differences in their vegetative systems; so in crossing, the greater or less facility of one species to unite with another is incidental on unknown differences in their reproductive systems. There is no more reason to think that species have been specially endowed with various degrees of sterility to prevent them crossing and breeding in Nature, than to think that trees have been specially endowed with various and somewhat analogous degrees of difficulty in being grafted together, in order to prevent them becoming inarched in our forests. “The sterility of first crosses between pure species, which have their reproductive systems perfect, seems to depend on several circumstances; in some cases largely on the early death of the embryo. The sterility of hybrids which have their reproductive systems imperfect, and which have had this system and their whole organisation disturbed by being compounded of two distinct species, seems closely allied to that sterility which so frequently affects pure species when their natural conditions of life have been disturbed. This view is supported by a parallelism of another kind: namely, that the crossing of forms, only slightly different, is favourable to the vigour and fertility of the offspring ; and that slight changes in the conditions of life are apparently favourable to the vigour and fertility of all organic beings. It is not surprising that the degree of difficulty in uniting two species, and the degree of sterility of their hybrid offspring, should generally correspond, though due to distinct causes; for both depend on the amount of difference of some kind between the species which are crossed. 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. “First crosses between forms known to be varieties, or sufficiently alike to be considered as varieties, and their mongrel offspring, are very generally, but not quite universally, fertile. Nor is this nearly general and perfect fertility surprising, when we remember how liable we are to argue in a circle with respect to varieties in a state of Nature; and when we remember that the greater number of varieties have been produced under domestication by the selection of mere external differences, and not of differences in the reproductive system. In all other respects, excluding fertility, there is a close general resemblance between hybrids and mongrels.”—PP. 276–8.
We fully agree with the general tenor of this weighty passage; but forcible as are these arguments, and little as the value of fertility or infertility as a test of species may be, it must not be forgotten that the really important fact, so far as the inquiry into the origin of species goes, is, that there are such things in Nature as groups of animals and of plants, the members of which are incapable offertile union with those of other groups; and that there are such things as hybrids, which are absolutely sterile when crossed with other hybrids. For, if such phaenomena as these were exhibited by only two of those assemblages of living objects, to which the name of species (whether it be used in its physiological or in its morphological sense) is given, it would have to be accounted for by any theory of the origin of species, and every theory which could not account for it would be, so far, imperfect.
Up to this point, we have been dealing with matters of fact, and the statements which we have laid before the reader would, to the best of our knowledge, be admitted to contain a fair exposition of what is at present known respecting the essential properties of species, by all who have studied the question. And whatever may be his theoretical views, no naturalist will probably be disposed to demur to the following summary of that exposition —
Living beings, whether animals or plants, are divisible into multitudes of distinctly definable kinds, which are morphological species. They are also divisible into groups of individuals, which breed freely together, tending to reproduce their like, and are physiological species. Normally resembling their parents, the offspring of members of these species are still liable to vary; and the variation may be perpetuated by selection, as a race, which race, in many cases, presents all the characteristics of a morphological species. But it is not as yet proved that a race ever exhibits, when crossed with another race of the same species, those phaenomena of hybridisation 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.