done excellent service by insisting repeatedly on this latter point.
It often occurred to me that it would be advisable to try whether seedlings from cross-fertilised flowers were in any way superior to those from self-fertilised flowers. But as no instance was known with animals of any evil appearing in a single generation from the closest possible interbreeding, that is between brothers and sisters, I thought that the same rule would hold good with plants; and that it would be necessary at the sacrifice of too much time to self-fertilise and intercross plants during several successive generations, in order to arrive at any result. I ought to have reflected that such elaborate provisions favouring cross-fertilisation, as we see in innumerable plants, would not have been acquired for the sake of gaining a distant and slight advantage, or of avoiding a distant and slight evil. Moreover, the fertilisation of a flower by its own pollen corresponds to a closer form of interbreeding than is possible with ordinary bi-sexual animals; so that an earlier result might have been expected.
I was at last led to make the experiments recorded in the present volume from the following circumstance. For the sake of determining certain points with respect to inheritance, and without any thought of the effects of close interbreeding, I raised close together two large beds of self-fertilised and crossed seedlings from the same plant of Linaria vulgaris. To my surprise, the crossed plants when fully grown were plainly taller and more vigorous than the self-fertilised ones. Bees incessantly visit the flowers of this Linaria and carry pollen from one to the other; and if insects are excluded, the flowers produce extremely few seeds; so that the wild plants from which my seedlings were raised must have been intercrossed during all previous generations. It seemed therefore quite incredible that the difference between the two beds of seedlings could have been due to a single act of self-fertilisation; and I attributed the result to the self-fertilised seeds not having been well ripened, improbable as it was that all should have been in this state, or to some other accidental and inexplicable cause. During the next year, I raised for the same purpose as before two large beds close together of self-fertilised and crossed seedlings from the carnation, Dianthus caryophyllus. This plant, like the Linaria, is almost sterile if insects are excluded; and we may draw the same inference as before, namely, that the parent-plants must have been intercrossed during every or almost every previous generation. Nevertheless, the self-fertilised seedlings were plainly inferior in height and vigour to the crossed.
My attention was now thoroughly aroused, for I could hardly doubt that the difference between the two beds was due to the one set being the offspring of crossed, and the other of self-fertilised flowers. Accordingly I selected almost by hazard two other plants, which happened to be in flower in the greenhouse, namely, Mimulus luteus and Ipomoea purpurea, both of which, unlike the Linaria and Dianthus, are highly self-fertile if insects are excluded. Some flowers on a single plant of both species were fertilised with their own pollen, and others were crossed with pollen from a distinct individual; both plants being protected by a net from insects. The crossed and self-fertilised seeds thus produced were sown on opposite sides of the same pots, and treated in all respects alike; and the plants when fully grown were measured and compared. With both species, as in the cases of the Linaria and Dianthus, the crossed seedlings were conspicuously superior in height and in other ways to the self-fertilised. I therefore determined to begin a long series of experiments with various plants, and these were continued for the following eleven years; and we shall see that in a large majority of cases the crossed beat the self-fertilised plants. Several of the exceptional cases, moreover, in which the crossed plants were not victorious, can be explained.
It should be observed that I have spoken for the sake of brevity, and shall continue to do so, of crossed and self-fertilised seeds, seedlings, or plants; these terms implying that they are the product of crossed or self-fertilised flowers. Cross-fertilisation always means a cross between distinct plants which were raised from seeds and not from cuttings or buds. Self-fertilisation always implies that the flowers in question were impregnated with their own pollen.
My experiments were tried in the following manner. A single plant, if it produced a sufficiency of flowers, or two or three plants were placed under a net stretched on a frame, and large enough to cover the plant (together with the pot, when one was used) without touching it. This latter point is important, for if the flowers touch the net they may be cross-fertilised by bees, as I have known to happen;
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