(2) The second great fact is that among all the young born to a pair of parents, no two are exactly alike, nor are any exactly like their parents; nor are any two taken from all produced by all parents of that species exactly alike. They all resemble their parents at the corresponding age, in a general way and even very closely; but the resemblance is far from amounting to identity. This is called “variation.” It is familiar to us all in the case of the organism which we know best, and observe most closely, namely, man. It is also a matter of common observation in the case of dogs, cats, horses, and other domesticated animals. Many of these “variations” are exhibited in points of size, proportion, and colour, which are easily noted at once by the eye. But “variation” is really a deep-seated thing, and depends on causes which lie below the surface. We know that the offspring of men and of animals and of plants, give evidence of variations in what we call constitution, tendency, temperament, aptitude, strength, and that the colour, and even size of this or that part, are really only indications of a deep-seated difference in the living chemistry, the forces of nutrition and growth which reside in the living substance. The fact that many thousands of a species may be born and only a few survive, means therefore that many thousand varieties, often varieties not readily measured by the eye, are produced in each generation, from which a few individuals are in some way “selected” for survival.

(3) The third great fact is that though there is variation, amongst all the offspring in each generation, there is also a continual and definite inheritance by offspring of the qualities and structure of their parents to a degree which altogether preponderates over the variations. To put it in another way, we all know that every parental organism transmits to its young not only the qualities and structure of the species, or of the race, or of the family, but also transmits its own peculiarities or variations in which it departed from its parents, and from its brothers and sisters. This is best illustrated by our daily experience of human families.

These facts being admitted, and abundantly illustrated and traced in detail by years of observation and experimental breeding in all kinds of living things by hundreds of careful observers who have published the records of their studies, we come to the step where Darwin makes use of supposition or hypothesis. The question is, “Does the one which, out of the thousands of slightly different varieties, survives—do so by haphazard? or is there a necessarily acting state of things which selects that one special variety for survival?” Gardeners and breeders of pigeons, dogs, and cattle deliberately select the variations which they desire, breed from them, and so carry on by inheritance the special variation—whilst they ruthlessly destroy or restrain from breeding the numerous other variations in their “stock” which they do not desire. “If,” said Darwin, “there is any necessarily selective mechanism in Nature which could act as the breeder does, new varieties might be ‘naturally’ selected, and changes of form and appearance naturally established, which in the course of long ages would amount to such marked differences as separate what we call one species from another.” He showed that there is a natural mechanism of the required kind. “Since,” he says, “the competition among the members of any one kind or species for a place in life is so very severe, and the hostile circumstances so varied, and since all the competing offspring differ by ‘variation’ ever so little from one another, those varieties which are better suited in even the smallest degree to hold their own not merely in fighting with the others, but in withstanding injurious influences, in escaping enemies, and in procuring food, will be the ones which will survive, when a large number of cases, many thousands, extending over a large area and many years, are considered. Those which are ‘best fitted’ to get through the exceedingly numerous dangers and difficulties of life will be the survivors.” Hence we get the survival of the fit—the fit variations—by natural selection in the struggle for life. This, it will be observed, is an inference, and not a direct observation.

So long as the conditions remain practically or effectively unchanged, the animal or plant already “fitted” to them will be succeeded by those of its offspring which most resemble it in the essential points of “fitness.” But we know that in the course of ages, more or less rapidly, climates change, land emerges from the sea, islands join continents, continents become scattered islands, animals and plants migrate into regions previously uninhabited by them. As such changes gradually come on, the natural selection of favoured varieties will necessarily lead to the survival of others than those previously favoured, other variations better suited to the new conditions will survive.

The natural selection of favoured variations would not amount to much, were the variations not perpetuated by transmission to the young which they produce. This, it is common knowledge [see [(3)]], does take place. It is known also that a variation so established is as a result of the regular process of variation presented in larger volume or emphasised in character in some individuals of subsequent generations, and by continued “natural selection” it may become more and more a prominent or dominant feature of the race.

So far, the only assumption made by Mr. Darwin is that any or some of the endless variations which occur in all the offspring of wild plants and animals, in various combinations and degree in each individual, can be sufficiently important to determine the survival or non-survival of the organisms possessing them. That is a matter which has been largely studied and discussed. The verdict of those who have studied on the spot (as Darwin himself did) the teeming life of the tropics, the insects, birds, and plants of those regions, is that we are justified in considering that small variations are sufficiently important to turn the scale in favour of survival or non-survival. It is not easy for a man who is not a determined naturalist, constantly observing the ways of wild living things, to appreciate the evidence as to the efficacy of small variations, even were I able here to submit it to him. It is to be found in the published works of an army of investigators. In any case it is granted that effective variations—whether small or great—occur in nature, and that natural selection favours and perpetuates the new and fitter variety to the exclusion of the less fit.

The real difficulty to most people comes in the supposition next made by Mr. Darwin—namely, that this slow process of change by natural selection of favoured variations and their transmission and perpetuation by inheritance is sufficient to effect by its continued operation through enormous ages of time the conversion of a race of ancestral three-toed zebras into the one-toed horse of to-day; before that, of five-toed beasts into three-toed; at an earlier stage of fishlike creatures into four-footed land animals, and so on. You have to picture the whole series of animals and of plants which are now or ever have been, as two gigantic family trees or pedigrees, meeting in common ancestors of the simplest grade of microscopic life. All the diverging branches and twigs of these great “family trees” have been determined by the adaptation of living form to the endlessly varied conditions of life on this planet, by the natural selection or survival of variations and the transmission and accumulation of those variations from parent to offspring. This is a tremendous demand on the imagination. It is, however, not a difficult one to concede, when one is acquainted with the facts and conclusions of geology. The history of the crust of the earth was explained twenty years before the date of Darwin’s theory by Charles Lyell as due to the continued action through immense periods of time of the same natural forces which are now at work. And, moreover, the examination of the successive stratified deposits of the earth’s crust has yielded the remains of whole series of animals and of plants (simpler in character the older and deeper the rock in which they occur), which can be satisfactorily explained and interpreted as the ancestral forms from which present organisms have been developed.

The theory of the natural selection of variations as the moving spring in the gradual development of living forms from simplest living matter is Darwin’s theory. It is not possible to find any naturalist of consideration who does not accept it. There are various views held and discussed as to the cause of variation, as to the importance of small and of big variations, as to the non-transmissibility of some kinds of variation, and as to various peculiarities in regard to inheritance. They do not for the most part touch the main features of Mr. Darwin’s theory. No doubt we are learning and shall learn more about the facts of variation and the details of the process of hereditary transmission, but such increase of knowledge has not tended to undermine Mr. Darwin’s theory, and does not seem at all likely to do so.

On the occasion of the celebration at Cambridge in 1909 of the centenary of Darwin’s birth, I was invited by the Vice-Chancellor, on behalf of the University, to deliver in the Senate-house an address, others being given by representatives of the United States (Prof. Osborne), of Germany (Prof. Hertwig), and of Russia (Prof. Metchnikoff). The following is the text of that address:—

“I feel it a great honour to be called upon to speak here to-day, and to stand, on behalf of the naturalists of the British Empire, by the side of the distinguished men whose orations you have just heard.