Origin of Species Variorum

Comparison with 1869

somewhat analogous degrees of difficulty in being grafted together in order to prevent their inarching in our forests.

The sterility of first crosses and of their hybrid progeny has not, as far as we can judge, been acquired through natural selection. With first crosses between pure species, in which the reproductive system is in a perfect condition, the sterility seems to depend on several circumstances; in some cases largely on the early death of the embryo, but this apparently depends on some imperfection in the original act of impregnation. ↑ In the case of hybrids, it perhaps depends on their whole organisation having been disturbed by being compounded from two distinct forms; the sterility being closely allied to that .. which so frequently affects pure species, when exposed to unnatural conditions of life. .. .. .. This view is supported by a parallelism of another kind: namely, that, first, the crossing of forms only slightly differentiated favours the vigour and fertility of their offspring, whilst close interbreeding is injurious; and secondly, that slight changes in the conditions of life apparently add to the vigour and fertility of all organic beings, whilst greater changes are often injurious. But the facts given on the sterility of the illegitimate unions of dimorphic and trimorphic plants and of their illegitimate progeny, render it probable that there is some unknown bond connecting in all cases the degree of fertility of first unions with that of their offspring. The consideration of these facts on dimorphism, as well as the results of reciprocal crosses, drive us to conclude that in all cases the primary cause of sterility, both in the parents and in the offspring, is confined to differences in their reproductive systems. But why in numerous species, descended from a common parent-form, the reproductive system should in all have become more or less modified, leading to their mutual infertility, we do not know in the least; nor whether this has been effected directly, or in correlation with other structural and functional modifications. ↑

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, even if 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, and the fertility of the hybrids thus produced, 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, as is so often stated, invariably

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 their 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 the 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 uni- versally,