Darwin, and After Darwin, Volume 2 of 3 / Post-Darwinian Questions: Heredity and Utility

by George John Romanes

How, then, we may well ask, is it that so able a naturalist and so clear a thinker as Weismann can have so far departed from the inductive methods as to have not merely propounded the question touching Continuity and its degrees, or even of Continuity as absolute; but to have straightway assumed the latter possibility as a basis on which to run a system of branching and ever-changing speculations concerning evolution, variation, the ultimate structure of living material, the intimate mechanism of heredity, or, in short, such a system of deductive conjectures as has never been approached in the history of science? The answer to this question is surely not far to seek. Must it not be the answer already given? Must it not have been for the sake of rearing this enormous structure of speculation that Weismann has adopted the assumption of Continuity as absolute? As we have just seen, Galton had well shown how a theory of heredity could be founded on the general doctrine of Continuity, without anywhere departing from the inductive methods—even while fully recognizing the possibility of such continuity as absolute. But Galton's theory was a "Theory of Heredity," and nothing more. Therefore, while clearly perceiving that the Continuity in question may be absolute, he saw no reason, either in fact or in theory, for concluding that it must be. On the contrary, he saw that this question is, for the present, necessarily unripe for profitable discussion—and, a fortiori, for the shedding of clouds of seed in all the directions of "Weismannism."

Hence, what I desire to be borne in mind throughout the following discussion is, that it will have exclusive reference to the question of fact already stated, without regard to any superjacent theories; and, still more, that there is a vast distinction between any question touching the degrees in which acquired characters are transmitted to progeny, and the question as to whether they are ever transmitted in any degree at all. Now, the latter question, being of much greater importance than the former, is the one which will mainly occupy our attention throughout the rest of this Section.

We have already seen that before the subject was taken up by Weismann the difference between acquired and congenital characters in respect to transmissibility was generally taken to be one of degree; not one of kind. It was usually supposed that acquired characters, although not so fully and not so certainly inherited as congenital characters, nevertheless were inherited in some lesser degree; so that if the same acquired character continued to be successively acquired in a number of sequent generations, what was at first only a slight tendency to be inherited would become by summation a more and more pronounced tendency, till eventually the acquired character might become as strongly inherited as a congenital one. Or, more precisely, it was supposed that an acquired character, in virtue of such a summation of hereditary influence, would in time become congenital. Now, if this supposition be true, it is evident that more or less assistance must be lent to natural selection in its work of evolving adaptive modifications[28]. And inasmuch as we know to what a wonderful extent adaptive modifications are secured during individual life-times—by the direct action of the environment on the one hand, and by increased or diminished use of special organs and mental faculties on the other—it becomes obvious of what importance even a small measure of transmissibility on their part would be in furnishing to natural selection ready-made variations in required directions, as distinguished from promiscuous variations in all directions. Contrariwise, if functionally-produced adaptations and adaptations produced by the direct action of the environment are never transmitted in any degree, not only would there be an incalculable waste, so to speak, of adaptive modifications—these being all laboriously and often most delicately built up during life-times of individuals only to be thrown down again as regards the interest of species—but so large an additional burden would be thrown upon the shoulders of natural selection that it becomes difficult to conceive how even this gigantic principle could sustain it, as I shall endeavour to show more fully in future chapters. On the other hand, however, Weismann and his followers not only feel no difficulty in throwing overboard all this ready-made machinery for turning out adaptive modifications when and as required; but they even represent that by so doing they are following the logical maxim, Entia non sunt multiplicanda praeter necessitatem—which means, in its relation to causality, that we must not needlessly multiply hypothetical principles to explain given results. But when appeal is here made to this logical principle—the so-called Law of Parsimony—two things are forgotten.

In the first place, it is forgotten that the very question in debate is whether causes of the Lamarckian order are unnecessary to explain all the phenomena of organic nature. Of course if it could be proved that the theory of natural selection alone is competent to explain all these phenomena, appeal to the logical principle in question would be justifiable. But this is precisely the point which the followers of Darwin refuse to accept; and so long as it remains the very point at issue, it is a mere begging the question to represent that a class of causes which have hitherto been regarded as necessary are, in fact, unnecessary. Or, in other words, when Darwin himself so decidedly held that these causes are necessary as supplements to natural selection, the burden of proof is quite as much on the side of Weismann and his followers to show that Darwin's opinion was wrong, as it is on the side of Darwin's followers to show that it was right. Yet, notwithstanding the elaborate structure of theory which Weismann has raised, there is nowhere one single fact or one single consideration of much importance to the question in debate which was not perfectly well known to Darwin. Therefore I say that all this challenging of Darwinists to justify their "Lamarckian assumptions" really amounts to nothing more than a pitting of opinion against opinion, where there is at least as much call for justification on the one side as on the other.

Again, when these challenges are thrown down by Weismann and his followers, it appears to be forgotten that the conditions of their own theory are such as to render acceptance of the gauge a matter of great difficulty. The case is very much like that of a doughty knight pitching his glove into the sea, and then defying any antagonist to take it up. That this is the case a very little explanation will suffice to show.

The question to be settled is whether acquired characters are ever transmitted by heredity. Now suppose, for the sake of argument, that acquired characters are transmitted by heredity—though not so fully and not so certainly as congenital characters—how is this fact to be proved to the satisfaction of Weismann and his followers? First of all they answer,—Assuredly by adducing experimental proof of the inheritance of injuries, or mutilations. But in making this answer they appear to forget that Darwin has already shown its inefficiency. That the self-styled Neo-Lamarckians have been much more unguarded in this respect, I fully admit; but it is obviously unfair to identify Darwin's views with those of a small section of evolutionists, who are really as much opposed to Darwin's teaching on one side as is the school of Weismann on the other. Yet, on reading the essays of Weismann himself—and still more those of his followers—one would almost be led to gather that it is claimed by him to have enunciated the distinction between congenital and acquired characters in respect of transmissibility; and therefore also to have first raised the objection which lies against the theory of Pangenesis in respect of the non-transmissibility of mutilations. In point of fact, however, Darwin is as clear and decided on these points as Weismann. And his answer to the obvious difficulty touching the non-transmissibility of mutilations is, to quote his own words, "the long-continued inheritance of a part which has been removed during many generations is no real anomaly, for gemmules formerly derived from the part are multiplied and transmitted from generation to generation[29]." Therefore, so far as Darwin's theory is concerned, the challenge to produce evidence of the transmission of injuries is irrelevant: it is no more a part of Darwin's theory than it is of Weismann's to maintain that injuries are transmitted.

There is, however, one point in this connexion to which allusion must here be made. Although Darwin did not believe in the transmissibility of mutilations when these consist merely in the amputation of parts of an organism, he did believe in a probable tendency to transmission when removal of the part is followed by gangrene. For, as he says, in that case, all the gemmules of the mutilated or amputated part, as they are gradually attracted to that part (in accordance with the law of affinity which the theory assumes), will be successively destroyed by the morbid process. Now it is of importance to note that Darwin made this exception to the general rule of the non-transmissibility of mutilations, not because his theory of pangenesis required it, but because there appeared to be certain very definite observations and experiments—which will be mentioned later on—proving that when mutilations are followed by gangrene they are apt to be inherited: his object, therefore, was to reconcile these alleged facts with his theory, quite as much as to sustain his theory by such facts.

So much, then, for the challenge to produce direct evidence of the transmissibility of acquired characters, so far as mutilations are concerned: believers in Darwin's theory, as distinguished from Weismann's, are under no obligation to take up such a challenge. But the challenge does not end here. Show us, say the school of Weismann, a single instance where an acquired character of any kind (be it a mutilation or otherwise) has been inherited: this is all that we require: this is all that we wait for: and surely, unless it be acknowledged that the Lamarckian doctrine reposes on mere assumption, at least one such case ought to be forthcoming. Well, nothing can sound more reasonable than this in the first instance; but as soon as we begin to cast about for cases which will satisfy the Neo-Darwinians, we find that the structure of their theory is such as to preclude, in almost every conceivable instance, the possibility of meeting their demand. For their theory begins by assuming that natural selection is the one and only cause of organic evolution. Consequently, what their demand amounts to is throwing upon the other side the burden of disproving this assumption—or, in other words, of proving the negative that in any given case of transmitted adaptation natural selection has not been the sole agent at work. Now, it must obviously be in almost all cases impossible to prove this negative among species in a state of nature. For, even supposing that among such species Lamarckian principles have had a large share in the formation of hereditary and adaptive characters, how would Weismann himself propose that we should set about the proof of such a fact, where the proof demanded by his assumption is, that the abstract possibility of natural selection having had anything to do with the matter must be excluded? Obviously this is impossible in the case of inherited characters which are also adaptive characters. How then does it fare with the case of inherited characters which are not also adaptive? Merely that this case is met by another and sequent assumption, which constitutes an integral part of the Neo-Darwinian creed—namely, that in nature there can be no such characters. Seeing that natural selection is taken to be the only possible cause of change in species, it follows that all changes occurring in species must necessarily be adaptive, whether or not we are able to perceive the adaptations. In this way apparently useless characters, as well as obviously useful ones, are ruled out of the question: that is to say, all hereditary characters of species in a state of nature are assumed to be due to natural selection, and then it is demanded that the validity of this assumption should be disproved by anybody who doubts it. Yet Weismann himself would be unable to suggest any conceivable method by which it can be disproved among species in a state of nature—and this even supposing that the assumption is entirely false[30].

Consequently, the only way in which these speciously-sounding challenges can be adequately met is by removing some individuals of a species from a state of nature, and so from all known influences of natural selection; then, while carefully avoiding artificial selection, causing these individuals and their progeny through many generations unduly to exercise some parts of their bodies, or unduly to fail in the exercise of others. But, clearly, such an experiment is one that must take years to perform, and therefore it is now too early in the day to reproach the followers of Darwin with not having met the challenges which are thrown down by the followers of Weismann[31].