the beginning of a process of selection? Can they decide which is to perish and which to survive? To use a phrase of Romanes, can they have SELECTION-VALUE?
Darwin himself answered this question, and brought together many excellent examples to show that differences, apparently insignificant because very small, might be of decisive importance for the life of the possessor. But it is by no means enough to bring forward cases of this kind, for the question is not merely whether finished adaptations have selection-value, but whether the first beginnings of these, and whether the small, I might almost say minimal increments, which have led up from these beginnings to the perfect adaptation, have also had selection-value. To this question even one who, like myself, has been for many years a convinced adherent of the theory of selection, can only reply: WE MUST ASSUME SO, BUT WE CANNOT PROVE IT IN ANY CASE. It is not upon demonstrative evidence that we rely when we champion the doctrine of selection as a scientific truth; we base our argument on quite other grounds. Undoubtedly there are many apparently insignificant features, which can nevertheless be shown to be adaptations—for instance, the thickness of the basin-shaped shell of the limpets that live among the breakers on the shore. There can be no doubt that the thickness of these shells, combined with their flat form, protects the animals from the force of the waves breaking upon them—but how have they become so thick? What proportion of thickness was sufficient to decide that of two variants of a limpet one should survive, the other be eliminated? We can say nothing more than that we infer from the present state of the shell, that it must have varied in regard to differences in shell-thickness, and that these differences must have had selection-value—no proof therefore, but an assumption which we must show to be convincing.
For a long time the marvellously complex RADIATE and LATTICE-WORK skeletons of Radiolarians were regarded as a mere outflow of "Nature's infinite wealth of form," as an instance of a purely morphological character with no biological significance. But recent investigations have shown that these, too, have an adaptive significance (Hacker). The same thing has been shown by Schutt in regard to the lowly unicellular plants, the Peridineae, which abound alike on the surface of the ocean and in its depths. It has been shown that the long skeletal processes which grow out from these organisms have significance not merely as a supporting skeleton, but also as an extension of the superficial area, which increases the contact with the water-particles, and prevents the floating organisms from sinking. It has been established that the processes are considerably shorter in the colder layers of the ocean, and that they may be twelve times as long (Chun, "Reise der Valdivia", Leipzig, 1904.) in the warmer layers, thus corresponding to the greater or smaller amount of friction which takes place in the denser and less dense layers of the water.
The Peridineae of the warmer ocean layers have thus become long-rayed, those of the colder layers short-rayed, not through the direct effect of friction on the protoplasm, but through processes of selection, which favoured the longer rays in warm water, since they kept the organism afloat, while those with short rays sank and were eliminated. If we put the question as to selection-value in this case, and ask how great the variations in the length of processes must be in order to possess selection-value; what can we answer except that these variations must have been minimal, and yet sufficient to prevent too rapid sinking and consequent elimination? Yet this very case would give the ideal opportunity for a mathematical calculation of the minimal selection-value, although of course it is not feasible from lack of data to carry out the actual calculation.
But even in organisms of more than microscopic size there must frequently be minute, even microscopic differences which set going the process of selection, and regulate its progress to the highest possible perfection.
Many tropical trees possess thick, leathery leaves, as a protection against the force of the tropical rain drops. The DIRECT influence of the rain cannot be the cause of this power of resistance, for the leaves, while they were still thin, would simply have been torn to pieces. Their toughness must therefore be referred to selection, which would favour the trees with slightly thicker leaves, though we cannot calculate with any exactness how great the first stages of increase in thickness must have been. Our hypothesis receives further support from the fact that, in many such trees, the leaves are drawn out into a beak-like prolongation (Stahl and Haberlandt) which facilitates the rapid falling off of the rain water, and also from the fact that the leaves, while they are still young, hang limply down in bunches which offer the least possible resistance to the rain. Thus there are here three adaptations which can only be interpreted as due to selection. The initial stages of these adaptations must undoubtedly have had selection-value.
But even in regard to this case we are reasoning in a circle, not giving "proofs," and no one who does not wish to believe in the selection-value of the initial stages can be forced to do so. Among the many pieces of presumptive evidence a particularly weighty one seems to me to be THE SMALLNESS OF THE STEPS OF PROGRESS which we can observe in certain cases, as for instance in leaf-imitation among butterflies, and in mimicry generally. The resemblance to a leaf, for instance of a particular Kallima, seems to us so close as to be deceptive, and yet we find in another individual, or it may be in many others, a spot added which increases the resemblance, and which could not have become fixed unless the increased deceptiveness so produced had frequently led to the overlooking of its much persecuted possessor. But if we take the selection-value of the initial stages for granted, we are confronted with the further question which I myself formulated many years ago: How does it happen THAT THE NECESSARY BEGINNINGS OF A USEFUL VARIATION ARE ALWAYS PRESENT? How could insects which live upon or among green leaves become all green, while those that live on bark become brown? How have the desert animals become yellow and the Arctic animals white? Why were the necessary variations always present? How could the green locust lay brown eggs, or the privet caterpillar develop white and lilac-coloured lines on its green skin?
It is of no use answering to this that the question is wrongly formulated (Plate, "Selektionsprinzip u. Probleme der Artbildung" (3rd edition), Leipzig, 1908.) and that it is the converse that is true; that the process of selection takes place in accordance with the variations that present themselves. This proposition is undeniably true, but so also is another, which apparently negatives it: the variation required has in the majority of cases actually presented itself. Selection cannot solve this contradiction; it does not call forth the useful variation, but simply works upon it. The ultimate reason why one and the same insect should occur in green and in brown, as often happens in caterpillars and locusts, lies in the fact that variations towards brown presented themselves, and so also did variations towards green: THE KERNEL OF THE RIDDLE LIES IN THE VARYING, and for the present we can only say, that small variations in different directions present themselves in every species. Otherwise so many different kinds of variations could not have arisen. I have endeavoured to explain this remarkable fact by means of the intimate processes that must take place within the germ-plasm, and I shall return to the problem when dealing with "germinal selection."
We have, however, to make still greater demands on variation, for it is not enough that the necessary variation should occur in isolated individuals, because in that case there would be small prospect of its being preserved, notwithstanding its utility. Darwin at first believed, that even single variations might lead to transformation of the species, but later he became convinced that this was impossible, at least without the cooperation of other factors, such as isolation and sexual selection.
In the case of the GREEN CATERPILLARS WITH BRIGHT LONGITUDINAL STRIPES, numerous individuals exhibiting this useful variation must have been produced to start with. In all higher, that is, multicellular organisms, the germ-substance is the source of all transmissible variations, and this germ-plasm is not a simple substance but is made up of many primary constituents. The question can therefore be more precisely stated thus: How does it come about that in so many cases the useful variations present themselves in numbers just where they are required, the white oblique lines in the leaf-caterpillar on the under surface of the body, the accompanying coloured stripes just above them? And, further, how has it come about that in grass caterpillars, not oblique but longitudinal stripes, which are more effective for concealment among grass and plants, have been evolved? And finally, how is it that the same Hawk-moth caterpillars, which to-day show oblique stripes, possessed longitudinal stripes in Tertiary times? We can read this fact from the history of their development, and I have before attempted to show the biological significance of this change