It may be worth while to give another and more complex illustration of the action of natural selection. Certain plants excrete sweet juice, apparently for the sake of eliminating something injurious from the sap: this is effected, for instance, by glands at the base of the stipules in some Leguminosae, and at the backs of the leaves of the common laurel. This juice, though small in quantity, is greedily sought by insects; but their visits do not in any way benefit the plant. Now, let us suppose that the juice or nectar was excreted from the inside of the flowers of a certain number of plants of any species. Insects in seeking the nectar would get dusted with pollen, and would often transport it from one flower to another. The flowers of two distinct individuals of the same species would thus get crossed; and the act of crossing, as can be fully proved, gives rise to vigorous seedlings, which consequently would have the best chance of flourishing and surviving. The plants which produced flowers with the largest glands or nectaries, excreting most nectar, would oftenest be visited by insects, and would oftenest be crossed; and so in the long-run would gain the upper hand and form a local variety. The flowers, also, which had their stamens and pistils placed, in relation to the size and habits of the particular insect which visited them, so as to favour in any degree the transportal of the pollen, would likewise be favoured. We might have taken the case of insects visiting flowers for the sake of collecting pollen instead of nectar; and as pollen is formed for the sole purpose of fertilisation, its destruction appears to be a simple loss to the plant; yet if a little pollen were carried, at first occasionally and then habitually, by the pollen-devouring insects from flower to flower, and a cross thus effected, although nine-tenths of the pollen were destroyed it might still be a great gain to the plant to be thus robbed; and the individuals which produced more and more pollen, and had larger anthers, would be selected.
When our plant, by the above process long continued, had been rendered highly attractive to insects, they would, unintentionally on their part, regularly carry pollen from flower to flower; and that they do this effectually I could easily show by many striking facts. I will give only one, as likewise illustrating one step in the separation of the sexes of plants. Some holly-trees bear only male flowers, which have four stamens producing a rather small quantity of pollen, and a rudimentary pistil; other holly-trees bear only female flowers; these have a full-sized pistil, and four stamens with shrivelled anthers, in which not a grain of pollen can be detected. Having found a female tree exactly sixty yards from a male tree, I put the stigmas of twenty flowers, taken from different branches, under the microscope, and on all, without exception, there were a few pollen-grains, and on some a profusion. As the wind had set for several days from the female to the male tree, the pollen could not thus have been carried. The weather had been cold and boisterous and therefore not favourable to bees, nevertheless every female flower which I examined had been effectually fertilised by the bees, which had flown from tree to tree in search of nectar. But to return to our imaginary case; as soon as the plant had been rendered so highly attractive to insects that pollen was regularly carried from flower to flower, another process might commence. No naturalist doubts the advantage of what has been called the "physiological division of labour;" hence we may believe that it would be advantageous to a plant to produce stamens alone in one flower or on one whole plant, and pistils alone in another flower or on another plant. In plants under culture and placed under new conditions of life, sometimes the male organs and sometimes the female organs become more or less impotent; now if we suppose this to occur in ever so slight a degree under nature, then, as pollen is already carried regularly from flower to flower, and as a more complete separation of the sexes of our plant would be advantageous on the principle of the division of labour, individuals with this tendency more and more increased, would be continually favoured or selected, until at last a complete separation of the sexes might be effected. It would take up too much space to show the various steps, through dimorphism and other means, by which the separation of the sexes in plants of various kinds is apparently now in progress; but I may add that some of the species of holly in North America are, according to Asa Gray, in an exactly intermediate condition, or, as he expresses it, are more or less dioeciously polygamous.