'Die Bastardbefruchtung im Pflanzenreich' 1865.); and we have seen exactly parallel cases with the illegitimate seedlings of Lythrum and Primula. Many hybrids are the most persistent and profuse flowerers, as are some illegitimate plants. When a hybrid is crossed by either pure parent-form, it is notoriously much more fertile than when crossed inter se or by another hybrid; so when an illegitimate plant is fertilised by a legitimate plant, it is more fertile than when fertilised inter se or by another illegitimate plant. When two species are crossed and they produce numerous seeds, we expect as a general rule that their hybrid offspring will be moderately fertile; but if the parent species produce extremely few seeds, we expect that the hybrids will be very sterile. But there are marked exceptions, as shown by Gartner, to these rules. So it is with illegitimate unions and illegitimate offspring. Thus the mid- styled form of Lythrum salicaria, when illegitimately fertilised with pollen from the longest stamens of the short-styled form, produced an unusual number of seeds; and their illegitimate offspring were not at all, or hardly at all, sterile. On the other hand, the illegitimate offspring from the long-styled form, fertilised with pollen from the shortest stamens of the same form, yielded few seeds, and the illegitimate offspring thus produced were very sterile; but they were more sterile than might have been expected relatively to the difficulty of effecting the union of the parent sexual elements. No point is more remarkable in regard to the crossing of species than their unequal reciprocity. Thus species A will fertilise B with the greatest ease; but B will not fertilise A after hundreds of trials. We have exactly the same case with illegitimate unions; for the mid-styled Lythrum salicaria was easily fertilised by pollen from the longest stamens of the short-styled form, and yielded many seeds; but the latter form did not yield a single seed when fertilised by the longest stamens of the mid-styled form.

Another important point is prepotency. Gartner has shown that when a species is fertilised with pollen from another species, if it be afterwards fertilised with its own pollen, or with that of the same species, this is so prepotent over the foreign pollen that the effect of the latter, though placed on the stigma some time previously, is entirely destroyed. Exactly the same thing occurs with the two forms of a heterostyled species. Thus several long-styled flowers of Primula veris were fertilised illegitimately with pollen from another plant of the same form, and twenty-four hours afterwards legitimately with pollen from a short- styled dark-red polyanthus which is a variety of P. veris; and the result was that every one of the thirty seedlings thus raised bore flowers more or less red, showing plainly how prepotent the legitimate pollen from a short-styled plant was over the illegitimate pollen from a long-styled plant.

In all the several foregoing points the parallelism is wonderfully close between the effects of illegitimate and hybrid fertilisation. It is hardly an exaggeration to assert that seedlings from an illegitimately fertilised heterostyled plant are hybrids formed within the limits of one and the same species. This conclusion is important, for we thus learn that the difficulty in sexually uniting two organic forms and the sterility of their offspring, afford no sure criterion of so-called specific distinctness. If any one were to cross two varieties of the same form of Lythrum or Primula for the sake of ascertaining whether they were specifically distinct, and he found that they could be united only with some difficulty, that their offspring were extremely sterile, and that the parents and their offspring resembled in a whole series of relations crossed species and their hybrid offspring, he might maintain that his varieties had been proved to be good and true species; but he would be completely deceived. In the second place, as the forms of the same trimorphic or dimorphic heterostyled species are obviously identical in general structure, with the exception of the reproductive organs, and as they are identical in general constitution (for they live under precisely the same conditions), the sterility of their illegitimate unions and that of their illegitimate offspring, must depend exclusively on the nature of the sexual elements and on their incompatibility for uniting in a particular manner.

Charles Darwin

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