This does not apply to the individuals of the same species when cultivated in cleared ground in the same garden. But if their flowers are visited by insects, they will intercross; and this will give to their sexual elements during a considerable number of generations a sufficient amount of differentiation for a cross to be beneficial. Moreover, seeds are frequently exchanged or procured from other gardens having a different kind of soil; and the individuals of the same cultivated species will thus be subjected to a change of conditions. If the flowers are not visited by our native insects, or very rarely so, as in the case of the common and sweet pea, and apparently in that of the tobacco when kept in a hothouse, any differentiation in the sexual elements caused by intercrosses will tend to disappear. This appears to have occurred with the plants just mentioned, for they were not benefited by being crossed one with another, though they were greatly benefited by a cross with a fresh stock.

I have been led to the views just advanced with respect to the causes of the differentiation of the sexual elements and of the variability of our garden plants, by the results of my various experiments, and more especially by the four cases in which extremely inconstant species, after having been self-fertilised and grown under closely similar conditions for several generations, produced flowers of a uniform and constant tint. These conditions were nearly the same as those to which plants, growing in a garden clear of weeds, are subjected, if they are propagated by self-fertilised seeds on the same spot. The plants in pots were, however, exposed to less severe fluctuations of climate than those out of doors; but their conditions, though closely uniform for all the individuals of the same generation, differed somewhat in the successive generations. Now, under these circumstances, the sexual elements of the plants which were intercrossed in each generation retained sufficient differentiation during several years for their offspring to be superior to the self-fertilised, but this superiority gradually and manifestly decreased, as was shown by the difference in the result between a cross with one of the intercrossed plants and with a fresh stock. These intercrossed plants tended also in a few cases to become somewhat more uniform in some of their external characters than they were at first. With respect to the plants which were self-fertilised in each generation, their sexual elements apparently lost, after some years, all differentiation, for a cross between them did no more good than a cross between the flowers on the same plant. But it is a still more remarkable fact, that although the seedlings of Mimulus, Ipomoea, Dianthus, and Petunia which were first raised, varied excessively in the colour of their flowers, their offspring, after being self-fertilised and grown under uniform conditions for some generations, bore flowers almost as uniform in tint as those on a natural species. In one case also the plants themselves became remarkably uniform in height.

The conclusion that the advantages of a cross depend altogether on the differentiation of the sexual elements, harmonises perfectly with the fact that an occasional and slight change in the conditions of life is beneficial to all plants and animals. (12/13. I have given sufficient evidence on this head in my 'Variation under Domestication' chapter 18 volume 2 2nd edition page 127.) But the offspring from a cross between organisms which have been exposed to different conditions, profit in an incomparably higher degree than do young or old beings from a mere change in the conditions. In this latter case we never see anything like the effect which generally follows from a cross with another individual, especially from a cross with a fresh stock. This might, perhaps, have been expected, for the blending together of the sexual elements of two differentiated beings will affect the whole constitution at a very early period of life, whilst the organisation is highly flexible.

Charles Darwin

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