Charles Robert Darwin (1809–1882). Origin of Species.
The Harvard Classics. 1909–14.
On the Intercrossing of Individuals
I
In the first place, I have collected so large a body of facts, and made so many experiments, showing, in accordance with the almost universal belief of breeders, that with animals and plants a cross between different varieties, or between individuals of the same variety but of another strain, gives vigour and fertility to the offspring; and on the other hand, that close interbreeding diminishes vigour and fertility; that these facts alone incline me to believe that it is a general law of nature that no organic being fertilises itself for a perpetuity of generations; but that a cross with another individual is occasionally—perhaps at long intervals of time—indispensable.
On the belief that this is a law of nature, we can, I think, understand several large classes of facts, such as the following, which on any other view are inexplicable. Every hybridizer knows how unfavourable exposure to wet is to the fertilisation of a flower, yet what a multitude of flowers have their anthers and stigmas fully exposed to the weather! If an occasional cross be indispensable, notwithstanding that the plant’s own anthers and pistil stand so near each other as almost to insure self-fertilisation, the fullest freedom for the entrance of pollen from another individual will explain the above state of exposure of the organs. Many flowers, on the other hand, have their organs of fructification closely enclosed, as in the great papilionaceous or pea-family; but these almost invariably present beautiful and curious adaptations in relation to the visits of insects. So necessary are the visits of bees to many papilionaceous flowers, that their fertility is greatly diminished if these visits be prevented. Now, it is
When the stamens of a flower suddenly spring towards the pistil, or slowly move one after the other towards it, the contrivance seems adapted solely to ensure self-fertilisation; and no doubt it is useful for this end: but the agency of insects is often required to cause the stamens to spring forward, as Kölreuter has shown to be the case with the barberry; and in this very genus, which seems to have a special contrivance for self-fertilisation, it is well known that, if closely allied forms or varieties are planted near each other, it is hardly possible to raise pure seedlings, so largely do they naturally cross. In numerous other cases, far from self-fertilisation being favoured, there are special contrivances which effectually prevent the stigma receiving pollen from its own flower, as I could show from the works of Sprengel and others, as well as from my own observations: for instance, in Lobelia fulgens, there is a really beautiful and elaborate contrivance by which all the infinitely numerous pollen-granules are swept out of the conjoined anthers of each flower, before the stigma of that individual flower is ready to receive them; and as this flower is never visited, at least in my garden, by insects, it never sets a seed, though by placing pollen from one flower on the stigma of another, I raise plenty of seedlings. Another species of Lobelia which is visited by bees, seeds freely in my garden. In very many other cases, though there is no special mechanical contrivance to prevent the stigma receiving pollen from the same flower, yet, as Sprengel, and more recently Hildebrand, and others, have shown, and as I can confirm, either the anthers burst before the stigma is ready for fertilisation, or the stigma is
If several varieties of the cabbage, radish, onion, and of some other plants, be allowed to seed near each other, a large majority of the seedlings thus raised turn out, as I have found, mongrels: for instance, I raised 233 seedling cabbages from some plants of different varieties growing near each other, and of these only 78 were true to their kind, and some even of these were not perfectly true. Yet the pistil of each cabbage-flower is surrounded not only by its own six stamens but by those of the many other flowers on the same plant; and the pollen of each flower readily gets on its own stigma without insect agency; for I have found that plants carefully protected from insects produce the full number of pods. How, then, comes it that such a vast number of the seedlings are mongrelized? It must arise from the pollen of a distinct variety having a prepotent effect over the flower’s own pollen; and that this is part of the general law of good being derived from the intercrossing of distinct individuals of the same species. When distinct species are crossed the case is reversed, for a plant’s own pollen is almost always prepotent over foreign pollen; but to this subject we shall return in a future chapter.
In the case of a large tree covered with innumerable flowers, it may be objected that pollen could seldom be carried from tree to tree, and at most only from flower to flower on the same tree; and flowers on the same tree can be considered as distinct individuals only in a limited sense. I believe this objection to be valid, but that nature has largely provided against it by giving to trees a strong tendency to bear flowers with separated sexes. When the sexes are separated,
Turning for a brief space to animals: various terrestrial species are hermaphrodites, such as the land-mollusca and earth-worms; but these all pair. As yet I have not found a single terrestrial animal which can fertilise itself. This remarkable fact, which offers so strong a contrast with terrestrial plants, is intelligible on the view of an occasional cross being indispensable; for owing to the nature of the fertilising element there are no means, analogous to the action of insects and of the wind with plants, by which an occasional cross could be effected with terrestrial animals without the concurrence of two individuals. Of aquatic animals, there are many self-fertilizing hermaphrodites; but here the currents of water offer an obvious means for an occasional cross. As in the case of flowers, I have as yet failed, after consultation with one of the highest authorities, namely, Professor Huxley, to discover a single hermaphrodite animal with the organs of reproduction so perfectly enclosed that access from without, and the occasional influence of a distinct individual, can be shown to be physically impossible. Cirripedes long appeared to me to present, under this point of view, a case of great difficulty; but I have been enabled, by a fortunate chance, to prove that two individuals, though both are self-fertilising hermaphrodites, do sometimes cross.
It must have struck most naturalists as a strange anomaly that, both with animals and plants, some species of the same family and even of the same genus, though agreeing closely
From these several considerations and from the many special facts which I have collected, but which I am unable here to give, it appears that with animals and plants an occasional intercross between distinct individuals is a very general, if not universal, law of nature.