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.
Let us now turn to the nectar-feeding insects; we may suppose the plant of which we have been slowly increasing the nectar by continued selection, to be a common plant; and that certain insects depended in main part on its nectar for food. I could give many facts showing how anxious bees are to save time: for instance, their habit of cutting holes and sucking the nectar at the bases of certain flowers, which with a very little more trouble they can enter by the mouth. Bearing such facts in mind, it may be believed that under certain circumstances individual differences in the curvature or length of the proboscis, etc., too slight to be appreciated by us, might profit a bee or other insect, so that certain individuals would be able to obtain their food more quickly than others; and thus the communities to which they belonged would flourish and throw off many swarms inheriting the same peculiarities. The tubes of the corolla of the common red or incarnate clovers (Trifolium pratense and incarnatum) do not on a hasty glance appear to differ in length; yet the hive-bee can easily suck the nectar out of the incarnate clover, but not out of the common red clover, which is visited by humble-bees alone; so that whole fields of the red clover offer in vain an abundant supply of precious nectar to the hive-bee. That this nectar is much liked by the hive-bee is certain; for I have repeatedly seen, but only in the autumn, many hive-bees sucking the flowers through holes bitten in the base of the tube by humble bees. The difference in the length of the corolla in the two kinds of clover, which determines the visits of the hive-bee, must be very trifling; for I have been assured that when red clover has been mown, the flowers of the second crop are somewhat smaller, and that these are visited by many hive-bees. I do not know whether this statement is accurate; nor whether another published statement can be trusted, namely, that the Ligurian bee, which is generally considered a mere variety of the common hive-bee, and which freely crosses with it, is able to reach and suck the nectar of the red clover. Thus, in a country where this kind of clover abounded, it might be a great advantage to the hive-bee to have a slightly longer or differently constructed proboscis. On the other hand, as the fertility of this clover absolutely depends on bees visiting the flowers, if humble-bees were to become rare 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 and 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.
4. 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 Kolreuter. 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
