Evolution and Adaptation

by Thomas Hunt Morgan

Darwin again makes the statement that under the term variation it must never be forgotten that mere individual differences are meant. “As a man can produce a great result with his domestic animals and plants by adding up in any given direction individual differences, so could natural selection, but far more easily from having incomparably longer time for action.” Too much emphasis cannot be laid on the fact that Darwin believed that selection takes place amongst the small individual differences that we find in animals and plants. Some of his followers, as we shall see, are apt to put into the background this fundamental conception of Darwin’s view. His constant comparison between the results of artificial and natural selection leaves no room for doubt as to his meaning. Darwin himself seems, at times, not unconscious of the weakness of this comparison. He says: “How fleeting are the wishes and efforts of man! how short his time! and consequently how poor will be his results, compared with those accumulated by Nature during whole geological periods. Can we wonder then that Nature’s productions should be far ‘truer’ in character than man’s productions; that they should be infinitely better adapted to the most complex conditions of life, and should plainly bear the stamp of far higher workmanship?” We should not lose sight of the fact that even after the most rigorous selective process has been brought to bear on organisms, namely, by isolation under domestication, we do not apparently find ourselves gradually approaching nearer and nearer to the formation of new species, but we find, on the contrary, that we have produced something quite different. In the light of this truth, the relation between the two selective theories may appear quite different from the interpretation that Darwin gives of it. We may well doubt whether nature does select so much better than does man, and whether she has ever made new species in this way.

We come now to a point that touches the theory of natural selection in a very vital spot.

“It may be well here to remark that with all beings there must be much fortuitous destruction, which can have little or no influence on the course of natural selection. For instance, a vast number of eggs or seeds are annually devoured, and these could be modified through natural selection only if they varied in some manner which protected them from their enemies. Yet many of these eggs or seeds would perhaps, if not destroyed, have yielded individuals better adapted to their conditions of life than any of those which happened to survive. So again a vast number of mature animals and plants, whether or not they be the best adapted to their conditions, must be annually destroyed by accidental causes, which would not be in the least degree mitigated by certain changes of structure or constitution which would in other ways be beneficial to the species. But let the destruction of the adults be ever so heavy, if the number which can exist in any district be not wholly kept down by such causes,—or again let the destruction of eggs or seeds be so great that only a hundredth or a thousandth part are developed,—yet of those which do survive, the best adapted individuals, supposing that there is any variability in a favorable direction, will tend to propagate their kind in larger numbers than the less well adapted. If the numbers be wholly kept down by the causes just indicated, as will often have been the case, natural selection will be powerless in certain beneficial directions; but this is no valid objection to its efficiency at other times and in other ways; for we are far from having any reason to suppose that many species ever undergo modification and improvement at the same time in the same area.”

Some of the admissions made in this paragraph have an important bearing on the theory of natural selection. Far from supposing that fortuitous destruction would have no influence on the course of natural selection, it can be shown that it would have a most disastrous effect. In many cases the destruction comes in the form of a catastrophe to the individuals, so that small differences in structure, whether advantageous or not, are utterly unavailing. Our experience shows us that a destruction of this sort is going on around us all the time, and accounts in large part for the way in which the majority of animals and plants are destroyed. Unless, for example, a seed happen to fall on a place suitable for its growth, it will perish without respect to a slight advantage it may have over other seeds of its kind. Of the thousands of eggs laid by one starfish, chance alone will decide whether one or another embryo is destroyed by larger animals, or if they escape this danger, the majority of them may be carried out to sea, where it will not be of the least avail if one individual has a slight advantage over the others. Darwin admits this, but adds that, if only a thousandth part is developed, yet of those that do survive the best adapted individuals will tend to propagate their kind in larger numbers than the less well adapted. The argument is not, however, so simple as it appears to be on the surface. I pass over, for the present, the apparent inconsequence in this statement that the best adapted individuals will tend to propagate their kind in larger numbers. It is not by any means certain that this is the case. Darwin’s meaning is, however, fairly clear, and can be interpreted to mean this: after the fortuitous destruction has finished, there will be a further competition of the survivors amongst themselves and with the surrounding conditions. In this higher competition, which is less severe, small individual differences suffice to determine the survival of certain individuals. These are, therefore, selected.

In this argument it is assumed that a second competition takes place after the first destruction of individuals has occurred, and this presupposes that more individuals reach maturity than there is room for in the economy of nature. But we do not know to what extent this takes place. If only as many mature as can survive, then the second competition does not take place. If, on the other hand, fewer mature than there is room for, then again competition does not take place. And if at all times selection is not rigorously carried out, everything may be lost that has been so laboriously gained. We see then that the result that Darwin imagines would take place, can be carried out only when more individuals reach maturity than there is room for (if it is a case of competition with one another), or that escape their enemies (if it is a question of competition with other forms).

It is instructive to consider some of the examples that Darwin has given to illustrate how the process of natural selection is carried out. The first example is the imaginary case of a species of wolf, the individuals of which secure their prey sometimes by craft, sometimes by strength, and sometimes by fleetness. If the prey captured by the first two methods should fail, then all the wolves would be obliged to capture their food by fleetness, and consequently the fleetest alone would survive. “I can see no more reason to doubt that this would be the result than that man should improve the fleetness of his greyhounds.” But even if the fleetness of the race could be kept up in this way, it does not follow that a new species of wolf would be formed in consequence, as Darwin implies. His own comment on this illustration is, perhaps, the best criticism that can be made.

“It should be observed that, in the above illustration, I speak of the slimmest individual wolves, and not of any single strongly marked variation having been preserved. In former editions of this work I sometimes spoke as if this latter alternative had frequently occurred. I saw the great importance of individual differences, and this led me fully to discuss the results of unconscious selection by man, which depends on the preservation of all the more or less valuable individuals, and on the destruction of the worst. I saw, also, that the preservation in a state of nature of any occasional deviation of structure, such as a monstrosity, would be a rare event; and that, if at first preserved, it would generally be lost by subsequent intercrossing with ordinary individuals. Nevertheless, until reading an able and valuable article in the North British Review (1867), I did not appreciate how rarely single variations, whether slight or strongly marked, could be perpetuated. The author takes the case of a pair of animals, producing during their lifetime two hundred offspring, of which, from various causes of destruction, only two on an average survive to procreate their kind. This is rather an extreme estimate for most of the higher animals, but by no means so for many of the lower organisms. He then shows that if a single individual were born, which varied in some manner, giving it twice as good a chance of life as that of the other individuals, yet the chances would be strongly against its survival. Supposing it to survive and to breed, and that half its young inherited the favourable variation; still, as the reviewer goes on to show, the young would have only a slightly better chance of surviving and breeding; and this chance would go on decreasing in the succeeding generations. The justice of these remarks cannot, I think, be disputed. If, for instance, a bird of some kind could procure its food more easily by having its beak curved, and if one were born with its beak strongly curved, and which consequently flourished, nevertheless there would be a very poor chance of this one individual perpetuating its kind to the exclusion of the common form; but there can hardly be a doubt, judging by what we see taking place under domestication, that this result would follow from the preservation during many generations of a large number of individuals with more or less strongly curved beaks, and from the destruction of a still larger number with the straightest beaks.”

There then follows what, I believe, is one of the most significant admissions in the “Origin of Species”:—

“It should not, however, be overlooked that certain rather strongly marked variations, which no one would rank as mere individual differences, frequently recur owing to a similar organization being similarly acted on—of which fact numerous instances could be given with our domestic productions. In such cases, if the varying individual did not actually transmit to its offspring its newly acquired character, it would undoubtedly transmit to them, as long as the existing conditions remained the same, a still stronger tendency to vary in the same manner. There can also be little doubt that the tendency to vary in the same manner has often been so strong that all the individuals of the same species have been similarly modified without the aid of any form of selection. Or only a third, fifth, or tenth part of the individuals may have been thus affected, of which fact several instances could be given. Thus Graba estimates that about one-fifth of the guillemots in the Faroe Islands consist of a variety so well marked, that it was formerly ranked as a distinct species under the name of Uria lacrymans. In cases of this kind, if the variation were of a beneficial nature, the original form would soon be supplanted by the modified form, through the survival of the fittest.”

Do not the admissions in this paragraph almost amount to a withdrawal of much that has preceded in regard to the survival of fluctuating, individual differences? In the last edition, from which we have just quoted, Darwin, in response to the criticisms which his book met, inserted here and there statements that are in many ways in contradiction to the statements in the first edition, and yet the earlier statements have been allowed to stand for the most part.