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in any country, it might be a great advantage to the plant to have a shorter or more deeply divided corolla, so that the hive-bees should be enabled to suck its flowers. Thus I can understand how a flower and a bee might slowly become, either simultaneously or one after the other, modified and adapted to each other in the most perfect manner, by the continued preservation of all the individuals which presented slight deviations of structure mutually favourable to each other. I am well aware that this doctrine of natural selection, exemplified in the above imaginary instances, is open to the same objections which were first urged against Sir Charles Lyell's noble views on “the modern changes of the earth, as illustrative of geology;” but we now seldom hear the agencies which we see still at work, spoken of as trifling or insignificant, when used in explaining the excavation of the deepest valleys or the formation of long lines of inland cliffs. Natural selection acts only by the preservation and accumulation of small inherited modifications, each profitable to the preserved being; and as modern geology has almost banished such views as the excavation of a great valley by a single diluvial wave, so will natural selection banish the belief of the continued creation of new organic beings, or of any great and sudden modification in their structure.
On the Intercrossing of Individuals.
I must here introduce a short digression. In the case of animals and plants with separated sexes, it is of course obvious that two individuals must always (with the exception of the curious and not well understood cases of parthenogenesis) unite for each birth; but in the case of hermaphrodites this is far from obvious. Nevertheless there is reason to believe that with all hermaphrodites two individuals, either occasionally or habitually, concur for the reproduction of their kind. This view was long ago doubtfully suggested by Sprengel, Knight and Kölreuter. We shall presently see its importance; but I must here treat the subject with extreme brevity, though I have the materials prepared for an ample discussion. All vertebrate animals, all insects, and some other large groups of animals, pair for each birth. Modern research has much diminished the number of supposed hermaphrodites, and of real hermaphrodites a large number pair; that is, two individuals regularly unite for reproduction, which is all that concerns us. But still there are many hermaphrodite animals which certainly do not habitually pair, and a vast majority of plants are hermaphrodites. What reason, it may be asked, is there for supposing in these cases that two individuals ever concur in reproduction? As it is impossible here to enter on details, I must trust to some general considerations alone.
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. 10
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 scarcely possible for insects to fly from flower to flower, and not to carry pollen from one to the other, to the great good of the plant. Insects act like a camel-hair pencil, and it is sufficient, to ensure fertilisation, just to touch with the same brush the anthers of one flower and then the stigma of another; but it must not be supposed that bees would thus produce a multitude of hybrids between distinct species; for if a plant's own pollen and that from another species are placed on the same stigma, the former is so prepotent that it invariably and completely destroys, as has been shown by Gärtner, the influence of the foreign pollen.
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 ready before the pollen of that flower is ready, so that these so-named dichogamous plants have in fact separated sexes, and must habitually be crossed. So it is with the reciprocally dimorphic and trimorphic plants previously alluded to. How strange are these facts! How strange that the pollen and stigmatic surface of the same flower, though placed so close together, as if for the very purpose of self-fertilisation, should be in so many cases mutually useless to each other? How simply are these facts explained on the view of an occasional cross with a distinct individual being advantageous or indispensable! 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