_Professor of Zoology in the University of Freiburg_ (_Baden_)

I. THE IDEA OF SELECTION

Many and diverse were the discoveries made by Charles Darwin in the course of a long and strenuous life, but none of them has had so far-reaching an influence on the science and thought of his time as the theory of selection. I do not believe that the theory of evolution would have made its way so easily and so quickly after Darwin took up the cudgels in favour of it if he had not been able to support it by a principle which was capable of solving, in a simple manner, the greatest riddle that living nature presents to us,--I mean the purposiveness of every living form relative to the conditions of its life and its marvellously exact adaptation to these.

Everyone knows that Darwin was not alone in discovering the principle of selection, and that the same idea occurred simultaneously and independently to Alfred Russel Wallace. At the memorable meeting of the Linnean Society on 1st July, 1858, two papers were read (communicated by Lyell and Hooker) both setting forth the same idea of selection. One was written by Charles Darwin in Kent, the other by Alfred Wallace in Ternate, in the Malay Archipelago. It was a splendid proof of the magnanimity of these two investigators, that they thus in all friendliness and without envy, united in laying their ideas before a scientific tribunal: their names will always shine side by side as two of the brightest stars in the scientific sky.

The idea of selection set forth by the two naturalists was at the time absolutely new, but it was also so simple that Huxley could say of it later, "How extremely stupid not to have thought of that." As Darwin was led to the general doctrine of descent, not through the labours of his predecessors in the early years of the century, but by his own observations, so it was in regard to the principle of selection. He was struck by the innumerable cases of adaptation, as, for instance, that of the woodp.e.c.k.e.rs and tree-frogs to climbing, or the hooks and feather-like appendages of seeds, which aid in the distribution of plants, and he said to himself that an explanation of adaptations was the first thing to be sought for in attempting to formulate a theory of evolution.

But since adaptations point to _changes_ which have been undergone by the ancestral forms of existing species, it is necessary, first of all, to inquire how far species in general are _variable_. Thus Darwin's attention was directed in the first place to the phenomenon of variability, and the use man has made of this, from very early times, in the breeding of his domesticated animals and cultivated plants. He inquired carefully how breeders set to work, when they wished to modify the structure and appearance of a species to their own ends, and it was soon clear to him that _selection for breeding purposes_ played the chief part.

But how was it possible that such processes should occur in free nature? Who is here the breeder, making the selection, choosing out one individual to bring forth offspring and rejecting others? That was the problem that for a long time remained a riddle to him.

Darwin himself relates how illumination suddenly came to him. He had been reading, for his own pleasure, Malthus' book on Population, and, as he had long known from numerous observations, that every species gives rise to many more descendants than ever attain to maturity, and that, therefore, the greater number of the descendants of a species perish without reproducing, the idea came to him that the decision as to which member of a species was to perish and which was to attain to maturity and reproduction might not be a matter of chance, but might be determined by the const.i.tution of the individuals themselves, according as they were more or less fitted for survival. With this idea the foundation of the theory of selection was laid.

In _artificial selection_ the breeder chooses out for pairing only such individuals as possess the character desired by him in a somewhat higher degree than the rest of the race. Some of the descendants inherit this character, often in a still higher degree, and if this method be pursued throughout several generations, the race is transformed in respect of that particular character.

_Natural selection_ depends on the same three factors as _artificial selection_: 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 compet.i.tion for s.p.a.ce 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 a.s.sumed compet.i.tion 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 const.i.tuted, 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 succ.u.mb 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 less fitness of the organism that determines the "selection for breeding purposes," and it would be incomprehensible if, in this compet.i.tion, 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 for reproduction_, whether the "best" refers to the whole const.i.tution, 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 a.s.sume an evolutionary force that is continually transforming the most primitive and the simplest forms of life into ever higher forms, and the h.o.m.ogeneity 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 a.s.sume 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 of life cannot be determined by the animal itself, the adaptations must be called 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

Lamarck, as is well known, formulated a definite theory of evolution at the beginning of the nineteenth century, exactly fifty years before the Darwin-Wallace principle of selection was given to the world. This brilliant investigator also endeavoured to support his theory by demonstrating forces which might have brought about the transformations of the organic world in the course of the ages. In addition to other factors, he laid special emphasis on the increased or diminished use of the parts of the body, a.s.suming that the strengthening or weakening which takes place from this cause during the individual life, could be handed on to the offspring, and thus intensified and raised to the rank of a specific character. Darwin also regarded this _Lamarckian principle_, as it is now generally called, as a factor in evolution, but he was not fully convinced of the transmissibility of acquired characters.

As I have here to deal only with the theory of selection, I need not discuss the Lamarckian hypothesis, but I must express my opinion that there is room for much doubt as to the cooperation of this principle in evolution. Not only is it difficult to imagine how the transmission of functional modifications could take place, but, up to the present time, notwithstanding the endeavours of many excellent investigators, not a single actual proof of such inheritance has been brought forward. Semon's experiments on plants are, according to the botanist Pfeffer, not to be relied on, and even the recent, beautiful experiments made by Dr. Kammerer on salamanders, cannot, as I hope to show elsewhere, be regarded as proof, if only because they do not deal at all with functional modifications, that is, with modifications brought about by use, and it is to these _alone_ that the Lamarckian principle refers.

III. OBJECTIONS TO THE THEORY OF SELECTION

(_a_) _Saltatory evolution_

The Darwinian doctrine of evolution depends essentially on _the c.u.mulative augmentation_ of minute variations in the direction of utility. But can such minute variations, which are undoubtedly continually appearing among the individuals of the same species, possess any selection-value; can they determine which individuals are to survive, and which are to succ.u.mb; can they be increased by natural selection till they attain to the highest development of a purposive variation?

To many this seems so improbable that they have urged a theory of evolution by leaps from species to species. Kolliker, in 1872, compared the evolution of species with the processes which we can observe in the individual life in cases of alternation of generations.

But a polyp only gives rise to a medusa because it has itself arisen from one, and there can be no question of a medusa ever having arisen suddenly and _de novo_ from a polyp-bud, if only because both forms are adapted in their structure as a whole, and in every detail to the conditions of their life. A sudden origin, in a natural way, of numerous adaptations is inconceivable. Even the degeneration of a medusoid from a free-swimming animal to a mere brood-sac (gonoph.o.r.e) is not sudden and saltatory, but occurs by imperceptible modifications throughout hundreds of years, as we can learn from the numerous stages of the process of degeneration persisting at the same time in different species.

If, then, the degeneration to a simple brood-sac takes place only by very slow transitions, each stage of which may last for centuries, how could the much more complex _ascending_ evolution possibly have taken place by sudden leaps? I regard this argument as capable of further extension, for wherever in nature we come upon degeneration, it is taking place by minute steps and with a slowness that makes it not directly perceptible, and I believe that this in itself justifies us in concluding that _the same must be true of ascending_ evolution. But in the latter case the goal can seldom be distinctly recognised while in cases of degeneration the starting-point of the process can often be inferred, because several nearly related species may represent different stages.

In recent years Bateson in particular has championed the idea of saltatory, or so-called discontinuous evolution, and has collected a number of cases in which more or less marked variations have suddenly appeared. These are taken for the most part from among domesticated animals which have been bred and crossed for a long time, and it is hardly to be wondered at that their much mixed and much influenced germ-plasm should, under certain conditions, give rise to remarkable phenomena, often indeed producing forms which are strongly suggestive of monstrosities, and which would undoubtedly not survive in free nature, unprotected by man. I should regard such cases as due to an intensified germinal selection--though this is to antic.i.p.ate a little--and from this point of view it cannot be denied that they have a special interest. But they seem to me to have no significance as far as the transformation of species is concerned, if only because of the extreme rarity of their occurrence.

There are, however, many variations which have appeared in a sudden and saltatory manner, and some of these Darwin pointed out and discussed in detail: the copper beech, the weeping trees, the oak with "fern-like leaves," certain garden-flowers, etc. But none of them have persisted in free nature, or evolved into permanent types.

On the other hand, wherever enduring types have arisen, we find traces of a gradual origin by successive stages, even if, at first sight, their origin may appear to have been sudden. This is the case with _seasonal Dimorphism_, the first known cases of which exhibited marked differences between the two generations, the winter and the summer brood. Take for instance the much discussed and studied form _Vanessa_ (_Araschnia_) _levana-prorsa_. Here the differences between the two forms are so great and so apparently disconnected, that one might almost believe it to be a sudden mutation, were it not that old transition-stages can be called forth by particular temperatures, and we know other b.u.t.terflies, as for instance our Garden Whites, in which the differences between the two generations are not nearly so marked; indeed, they are so little apparent that they are scarcely likely to be noticed except by experts. Thus here again there are small initial steps, some of which, indeed, must be regarded as adaptations, such as the green-sprinkled or lightly tinted under-surface which gives them a deceptive resemblance to parsley or to Cardamine leaves.

Even if saltatory variations do occur, we cannot a.s.sume that these _have ever led to forms which are capable of survival under the conditions of wild life_. Experience has shown that in plants which have suddenly varied the power of persistence is diminished.

Korschinsky attributes to them weaknesses of organisation in general; "they bloom late, ripen few of their seeds, and show great sensitiveness to cold." These are not the characters which make for success in the struggle for existence.

We must briefly refer here to the views--much discussed in the last decade--of H. de Vries, who believes that the roots of transformation must be sought for in _saltatory variations arising from internal causes_, and distinguishes such _mutations_, as he has called them, from ordinary individual variations, in that they breed true, that is, with strict in-breeding they are handed on pure to the next generation. I have elsewhere endeavoured to point out the weaknesses of this theory,[33] and I am the less inclined to return to it here that it now appears[34] that the far-reaching conclusions drawn by de Vries from his observations on the Evening Primrose, _Oenothera lamarckiana_, rest upon a very insecure foundation. The plant from which de Vries saw numerous "species"--his "mutations"--arise was not, as he a.s.sumed, a _wild species_ that had been introduced to Europe from America, but was probably a hybrid form which was first discovered in the Jardin des Plantes in Paris, and which does not appear to exist anywhere in America as a wild species.

This gives a severe shock to the "Mutation theory," for the other _actually wild_ species with which de Vries experimented showed no "mutations" but yielded only negative results.

Thus we come to the conclusion that Darwin[35] was right in regarding transformations as taking place by minute steps, which, if useful, are augmented in the course of innumerable generations, because their possessors more frequently survive in the struggle for existence.

(_b_) _Selection-value of the initial steps_

Is it possible that the insignificant deviations which we know as "individual variations" can form 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 a.s.sume 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 sh.e.l.l of the limpets that live among the breakers on the sh.o.r.e. There can be no doubt that the thickness of these sh.e.l.ls, combined with their flat forms, 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 sh.e.l.l, that it must have varied in regard to differences in sh.e.l.l-thickness, and that these differences must have had selection-value,--no proof therefore, but an a.s.sumption 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[36] 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 raindrops. 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 b.u.t.terflies, 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[37] 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 s.e.xual 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 const.i.tuents. 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 gra.s.s caterpillars, not oblique but longitudinal stripes, which are more effective for concealment among gra.s.s 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 of colour.[38]

For the present I need only draw the conclusion that one and the same caterpillar may exhibit the initial stages of both, and that it depends on the manner in which these marking elements are _intensified_ and _combined_ by natural selection whether whitish longitudinal or oblique stripes should result. In this case then the "useful variations" were actually "always there," and we see that in the same group of Lepidoptera, e.g. species of Sphingidae, evolution has occurred in both directions according to whether the form lived among gra.s.s or on broad leaves with oblique lateral veins, and we can observe even now that the species with oblique stripes have longitudinal stripes when young, that is to say, while the stripes have no biological significance. The white places in the skin which gave rise, probably first as small spots, to this protective marking could be combined in one way or another according to the requirements of the species. They must therefore either have possessed selection-value from the first, or, if this was not the case at their earliest occurrence, there must have been _some other factors_ which raised them to the point of selection-value. I shall return to this in discussing germinal selection. But the case may be followed still farther, and leads us to the same alternative on a still more secure basis.

Many years ago I observed in caterpillars of _Smerinthus populi_ (the poplar hawk-moth), which also possess white oblique stripes, that certain individuals showed _red spots_ above these stripes; these spots occurred only on certain segments, and never flowed together to form continuous stripes. In another species (_Smerinthus tiliae_) similar blood-red spots unite to form a line-like coloured seam in the last stage of larval life, while in _S. ocellata_ rust-red spots appear in individual caterpillars, but more rarely than in _S.

populi_, and they show no tendency to flow together.