The infertility which may be observed in various dimorphic and trimorphic plants, when they are illegitimately fertilised, that is by pollen taken from stamens not corresponding in height with the pistil, differs much in degree, up to absolute and utter sterility; just in the same manner as occurs in crossing distinct species. As the degree of sterility in the latter case depends in an eminent degree on the conditions of life being more or less favourable, so I have found it with illegitimate unions. It is well known that if pollen of a distinct species be placed on the stigma of a flower, and its own pollen be afterwards, even after a considerable interval of time, placed on the same stigma, its action is so strongly prepotent that it generally annihilates the effect of the foreign pollen; so it is with the pollen of the several forms of the same species, for legitimate pollen is strongly prepotent over illegitimate pollen, when both are placed on the same stigma. I ascertained this by fertilising several flowers, first illegitimately, and twenty-four hours afterwards legitimately, with pollen taken from a peculiarly coloured variety, and all the seedlings were similarly coloured; this shows that the legitimate pollen, though applied twenty-four hours subsequently, had wholly destroyed or prevented the action of the previously applied illegitimate pollen. Again, as in making reciprocal crosses between the same two species, there is occasionally a great difference in the result, so the same thing occurs with trimorphic plants; for instance, the mid-styled form of Lythrum salicaria was illegitimately fertilised with the greatest ease by pollen from the longer stamens of the short-styled form, and yielded many seeds; but the latter form did not yield a single seed when fertilised by the longer stamens of the mid-styled form.

In all these respects, and in others which might be added, the forms of the same undoubted species, when illegitimately united, behave in exactly the same manner as do two distinct species when crossed. This led me carefully to observe during four years many seedlings, raised from several illegitimate unions. The chief result is that these illegitimate plants, as they may be called, are not fully fertile. It is possible to raise from dimorphic species, both long-styled and short-styled illegitimate plants, and from trimorphic plants all three illegitimate forms. These can then be properly united in a legitimate manner. When this is done, there is no apparent reason why they should not yield as many seeds as did their parents when legitimately fertilised. But such is not the case. They are all infertile, in various degrees; some being so utterly and incurably sterile that they did not yield during four seasons a single seed or even seed-capsule. The sterility of these illegitimate plants, when united with each other in a legitimate manner, may be strictly compared with that of hybrids when crossed inter se. If, on the other hand, a hybrid is crossed with either pure parent-species, the sterility is usually much lessened: and so it is when an illegitimate plant is fertilised by a legitimate plant. In the same manner as the sterility of hybrids does not always run parallel with the difficulty of making the first cross between the two parent-species, so that sterility of certain illegitimate plants was unusually great, while the sterility of the union from which they were derived was by no means great. With hybrids raised from the same seed- capsule the degree of sterility is innately variable, so it is in a marked manner with illegitimate plants. Lastly, many hybrids are profuse and persistent flowerers, while other and more sterile hybrids produce few flowers, and are weak, miserable dwarfs; exactly similar cases occur with the illegitimate offspring of various dimorphic and trimorphic plants.

Altogether there is the closest identity in character and behaviour between illegitimate plants and hybrids. It is hardly an exaggeration to maintain that illegitimate plants are hybrids, produced within the limits of the same species by the improper union of certain forms, while ordinary hybrids are produced from an improper union between so-called distinct species. We have also already seen that there is the closest similarity in all respects between first illegitimate unions and first crosses between distinct species. This will perhaps be made more fully apparent by an illustration; we may suppose that a botanist found two well-marked varieties (and such occur) of the long-styled form of the trimorphic Lythrum salicaria, and that he determined to try by crossing whether they were specifically distinct. He would find that they yielded only about one-fifth of the proper number of seed, and that they behaved in all the other above specified respects as if they had been two distinct species. But to make the case sure, he would raise plants from his supposed hybridised seed, and he would find that the seedlings were miserably dwarfed and utterly sterile, and that they behaved in all other respects like ordinary hybrids. He might then maintain that he had actually proved, in accordance with the common view, that his two varieties were as good and as distinct species as any in the world; but he would be completely mistaken.

Charles Darwin

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