1. Can you doubt that the hitherto insoluble problem of inter-specific sterility would be solved, supposing cross-infertility were proved to arise before or during the process of specific differentiation, instead of after that process had been fully completed?
2. Can you doubt, after duly considering the circumstances under which allied species of plants have been differentiated—viz. in ninety-five per cent. of cases intimately commingled on common areas, and therefore under identical environments—that cross-infertility must have arisen before or during the specific differentiation?
3. Can you doubt, after duly considering the facts of prepotency on the one hand and those of Jordan's physiological varieties on the other, that cross-infertility does arise before or during the specific differentiation?
4. If you cannot express a doubt upon any of these points, can you explain why you refuse to accept the theory of the origin of species by means of physiological selection, together with the explanation which this theory affords of the continued cross-fertility of domesticated varieties?
5. Supposing this theory to be true, can you conceive of any other classes of facts which, either quantitatively or qualitatively, could more directly or more effectually prove its truth than those which have now been adduced?
On these five heads I entertain no doubt. I am convinced that the theory of physiological selection is the only one that can explain the facts of inter-specific sterility on the one hand, and, on the other hand, the contrast which these facts display to the unimpaired fertility of our domesticated varieties.
In conclusion, it seems desirable once more to insist that there is no antagonism or rivalry between the theories of natural and of physiological selection. For which purpose I will quote the final paragraph of my original paper.
So much, then, for the resemblances and the differences between the two theories. It only remains to add that the two are complementary. I have already shown some of the respects in which the newer theory comes to the assistance of the older, and this in the places where the older has stood most in need of assistance. In particular, I have shown that segregation of the fit entirely relieves survival of the fittest from the difficulty under which it has hitherto laboured of explaining why it is that sterility is so constantly found between species, while so rarely found between varieties which differ from one another even more than many species; why so many features of specific distinction are useless to the species presenting them; and why it is that incipient varieties are not obliterated by intercrossing with parent forms. Again, we have seen that physiological selection, by preventing such intercrossing, enables natural selection to promote diversity of character, and thus to evolve species in ramifying branches instead of in linear series—a work which I cannot see how natural selection could possibly perform unless thus aided by physiological selection. Moreover, we have seen that although natural selection alone could not induce sterility between allied types, yet when this sterility is given by physiological selection, the forms which present it would be favoured in the struggle for existence; and thus again the two principles are found playing, as it were, into each other's hands. And here, as elsewhere, I believe that the co-operation enables the two principles to effect very much more in the way of species-making than either of them could effect if working separately. On the one hand, without the assistance of physiological selection, natural selection would, I believe, be all but overcome by the adverse influences of free intercrossing—influences all the more potent under the very conditions which are required for the multiplication of species by divergence of character. On the other hand, without natural selection, physiological selection would be powerless to create any differences of specific type, other than those of mutual sterility and trivial details of structure, form, and colour—differences wholly without meaning from a utilitarian point of view. But in their combination these two principles appear to me able to accomplish what neither can accomplish alone—namely, a full and satisfactory explanation of the origin of species.