In this and all other cases, the relative fertility of the two kinds of union can, I think, be judged of more truly by the average number of seeds per capsule than by the proportion of flowers which yield capsules. The two methods might have been combined by giving the average number of seeds produced by all the flowers which were fertilised, whether they yielded capsules or not; but I have thought that it would be more instructive always to show separately the proportion of flowers which produced capsules, and the average number of apparently good seeds which the capsules contained.
Flowers legitimately fertilised set seeds under conditions which cause the almost complete failure of illegitimately fertilised flowers. Thus in the spring of 1862 forty flowers were fertilised at the same time in both ways. The plants were accidentally exposed in the greenhouse to too hot a sun, and a large number of umbels perished. Some, however, remained in moderately good health, and on these there were twelve flowers which had been fertilised legitimately, and eleven which had been fertilised illegitimately. The twelve legitimate unions yielded seven fine capsules, containing on an average each 57.3 good seeds; whilst the eleven illegitimate unions yielded only two capsules, of which one contained 39 seeds, but so poor, that I do not suppose one would have germinated, and the other contained 17 fairly good seeds.
From the facts now given the superiority of a legitimate over an illegitimate union admits of not the least doubt; and we have here a case to which no parallel exists in the vegetable or, indeed, in the animal kingdom. The individual plants of the present species, and as we shall see of several other species of Primula, are divided into two sets or bodies, which cannot be called distinct sexes, for both are hermaphrodites; yet they are to a certain extent sexually distinct, for they require reciprocal union for perfect fertility. As quadrupeds are divided into two nearly equal bodies of different sexes, so here we have two bodies, approximately equal in number, differing in their sexual powers and related to each other like males and females. There are many hermaphrodite animals which cannot fertilise themselves, but most unite with another hermaphrodite. So it is with numerous plants; for the pollen is often mature and shed, or is mechanically protruded, before the flower's own stigma is ready; and such flowers absolutely require the presence of another hermaphrodite for sexual union. But with the cowslip and various other species of Primula there is this wide difference, that one individual, though it can fertilise itself imperfectly, must unite with another individual for full fertility; it cannot, however, unite with any other individual in the same manner as an hermaphrodite plant can unite with any other one of the same species; or as one snail or earth-worm can unite with any other hermaphrodite individual. On the contrary, an individual belonging to one form of the cowslip in order to be perfectly fertile must unite with one of the other form, just as a male quadruped must and can unite only with the female.
I have spoken of the legitimate unions as being fully fertile; and I am fully justified in doing so, for flowers artificially fertilised in this manner yielded rather more seeds than plants naturally fertilised in a state of nature. The excess may be attributed to the plants having been grown separately in good soil. With respect to the illegitimate unions, we shall best appreciate their degree of lessened fertility by the following facts. Gartner estimated the sterility of the unions between distinct species, in a manner which allows of a strict comparison with the results of the legitimate and illegitimate unions of Primula. (1/6. 'Versuche uber die Bastarderzeugung' 1849 page 216.) With P. veris, for every 100 seeds yielded by the two legitimate unions, only 64 were yielded by an equal number of good capsules from the two illegitimate unions. With P. Sinensis, as we shall hereafter see, the proportion was nearly the same- -namely, as 100 to 62.