NATURAL SELECTION depends on the same three factors as ARTIFICIALSELECTION: on VARIABILITY, INHERITANCE, and SELECTION FOR BREEDING, but this last is here carried out not by a breeder but by what Darwin called the "struggle for existence." This last factor is one of the special features of the Darwinian conception of nature. That there are carnivorous animals which take heavy toll in every generation of the progeny of the animals on which they prey, and that there are herbivores which decimate the plants in every generation had long been known, but it is only since Darwin's time that sufficient attention has been paid to the facts that, in addition to this regular destruction, there exists between the members of a species a keen competition for space and food, which limits multiplication, and that numerous individuals of each species perish because of unfavourable climatic conditions. The "struggle for existence," which Darwin regarded as taking the place of the human breeder in free nature, is not a direct struggle between carnivores and their prey, but is the assumed competition for survival between individuals OF THE SAME species, of which, on an average, only those survive to reproduce which have the greatest power of resistance, while the others, less favourably constituted, perish early. This struggle is so keen, that, within a limited area, where the conditions of life have long remained unchanged, of every species, whatever be the degree of fertility, only two, ON AN AVERAGE, of the descendants of each pair survive; the others succumb either to enemies, or to disadvantages of climate, or to accident. A high degree of fertility is thus not an indication of the special success of a species, but of the numerous dangers that have attended its evolution. Of the six young brought forth by a pair of elephants in the course of their lives only two survive in a given area; similarly, of the millions of eggs which two thread-worms leave behind them only two survive. It is thus possible to estimate the dangers which threaten a species by its ratio of elimination, or, since this cannot be done directly, by its fertility.
Although a great number of the descendants of each generation fall victims to accident, among those that remain it is still the greater or lesser fitness of the organism that determines the "selection for breeding purposes," and it would be incomprehensible if, in this competition, it were not ultimately, that is, on an average, the best equipped which survive, in the sense of living long enough to reproduce.
Thus the principle of natural selection is THE SELECTION OF THE BEST FORREPRODUCTION, whether the "best" refers to the whole constitution, to one or more parts of the organism, or to one or more stages of development.
Every organ, every part, every character of an animal, fertility and intelligence included, must be improved in this manner, and be gradually brought up in the course of generations to its highest attainable state of perfection. And not only may improvement of parts be brought about in this way, but new parts and organs may arise, since, through the slow and minute steps of individual or "fluctuating" variations, a part may be added here or dropped out there, and thus something new is produced.
The principle of selection solved the riddle as to how what was purposive could conceivably be brought about without the intervention of a directing power, the riddle which animate nature presents to our intelligence at every turn, and in face of which the mind of a Kant could find no way out, for he regarded a solution of it as not to be hoped for. For, even if we were to assume an evolutionary force that is continually transforming the most primitive and the ******st forms of life into ever higher forms, and the homogeneity of primitive times into the infinite variety of the present, we should still be unable to infer from this alone how each of the numberless forms adapted to particular conditions of life should have appeared PRECISELY AT THE RIGHT MOMENT IN THE HISTORY OF THE EARTH to which their adaptations were appropriate, and precisely at the proper place in which all the conditions of life to which they were adapted occurred: the humming-birds at the same time as the flowers; the trichina at the same time as the pig; the bark-coloured moth at the same time as the oak, and the wasp-like moth at the same time as the wasp which protects it. Without processes of selection we should be obliged to assume a "pre-established harmony" after the famous Leibnitzian model, by means of which the clock of the evolution of organisms is so regulated as to strike in exact synchronism with that of the history of the earth! All forms of life are strictly adapted to the conditions of their life, and can persist under these conditions alone.
There must therefore be an intrinsic connection between the conditions and the structural adaptations of the organism, and, SINCE THE CONDITIONS OFLIFE CANNOT BE DETERMINED BY THE ANIMAL ITSELF, THE ADAPTATIONS MUST BECALLED FORTH BY THE CONDITIONS.
The selection theory teaches us how this is conceivable, since it enables us to understand that there is a continual production of what is non-purposive as well as of what is purposive, but the purposive alone survives, while the non-purposive perishes in the very act of arising.
This is the old wisdom taught long ago by Empedocles.
II. THE LAMARCKIAN PRINCIPLE.