Essays Upon Heredity and Kindred Biological Problems

The recent observations of Born274 upon the eggs of the frog also make it very probable that a double development is produced by the entrance of two spermatozoa into the egg, but here also only monstrosities, and not twins, were produced. On the other hand, it has been shown that in birds twins may be produced from the same egg, and there is no reason for the belief that their production is due to multiple impregnation. But if it may be assumed that human twins, when identical, have been derived from a single egg, it seems to me to be extremely probable that fertilization was also effected by a single sperm-cell. We cannot understand how such a high degree of similarity could have been produced if two sperm-cells had been made use of, for we are compelled to assume that two such cells would very rarely contain identical germ-plasms.

It is most probable that the egg-nucleus coalesces with the nucleus of a single spermatozoon, but the resulting segmentation-nucleus divides together with the cell-body itself, without the occurrence of those ontogenetic changes in the germ-plasm which normally take place. The nucleoplasm of the two daughter-cells still remains in the condition of germ-plasm, and its ontogenetic transformation begins afterwards—a transformation which must of course proceed in the same way in both cells, and must lead to the production of identical offspring. This is at least a possible explanation which we may retain until it has been either confirmed or disproved by fresh observations,—an explanation which is moreover supported by the well-known process of budding in the eggs of lower animals.

VI. Recapitulation

To bring together shortly the results of this essay:—the fundamental fact upon which everything else is founded is the fact that two polar bodies are expelled, as a preparation for embryonic development, from all animal eggs which require fertilization, while only one such body is expelled from all parthenogenetic eggs.

This fact in the first place refutes every purely morphological explanation of the process. If it were physiologically valueless, such a phyletic reminiscence of the two successive divisions of the egg-nucleus must have been also retained by the parthenogenetic egg.

In my opinion the expulsion of the first polar body implies the removal of ovogenetic nucleoplasm when it has become superfluous after the maturation of the egg has been completed. The expulsion of the second polar body can only mean the removal of part of the germ-plasm itself, a removal by which the number of ancestral germ-plasms is reduced to one half. This reduction must also take place in the male germ-cells, although we are not able to associate it confidently with any of the histological processes of spermatogenesis which have been hitherto observed.

Parthenogenesis takes place when the whole of the ancestral germ-plasms, inherited from the parents, are retained in the nucleus of the egg-cell. Development by fertilization makes it necessary that half the number of these ancestral germ-plasms must be first expelled from the egg, the original quantity being again restored by the addition of the sperm-nucleus to the remaining half.

In both cases the beginning of embryogenesis depends upon the presence of a certain, and in both cases equal, quantity of germ-plasm. This certain quantity is produced by the addition of the sperm-nucleus to the egg requiring fertilization, and the beginning of embryogenesis immediately follows fertilization. The parthenogenetic egg contains within itself the necessary quantity of germ-plasm, and the latter enters upon active development as soon as the single polar body has removed the ovogenetic nucleoplasm. The question which I have raised on a previous occasion—‘When is the parthenogenetic egg capable of development?’—now admits of the precise answer—‘Immediately after the expulsion of the polar body.’

From the preceding facts and considerations the important conclusion results that the germ-cells of any individual do not contain the same hereditary tendencies, but are all different, in that no two of them contain exactly the same combinations of hereditary tendencies. On this fact the well-known differences between the children of the same parents depend.

But the deeper meaning of this arrangement must doubtless be sought for in the individual variability which is thus continuously kept up and is always being forced into new combinations. Thus sexual reproduction is to be explained as an arrangement which ensures an ever-varying supply of individual differences.

VII.

ON THE SUPPOSED BOTANICAL PROOFS

OF THE

TRANSMISSION OF ACQUIRED CHARACTERS.

1888

VII.

ON THE SUPPOSED BOTANICAL PROOFS

OF THE

TRANSMISSION OF ACQUIRED CHARACTERS

In a lecture on heredity, delivered in 1883275, I first brought forward the opinion that acquired characters cannot be transmitted; and I then stated that there are no proofs of such transmission, that its occurrence is theoretically improbable, and that we must attempt to explain the transformation of species without its aid. Since that time many biologists have expressed their opinions upon the subject, some of them agreeing with me, while others have taken the opposite side. It is unnecessary to allude to those who have attacked my opinions without first understanding the real point in dispute, which turns upon the true meaning of the phrase ‘acquired character.’ I think it is now generally admitted that a very important problem is involved in this question, the solution of which will contribute in a decisive manner towards the formation of ideas as to the causes which have produced the transformation of species. For if acquired characters cannot be transmitted, the Lamarckian theory completely collapses, and we must entirely abandon the principle by which alone Lamarck sought to explain the transformation of species,—a principle of which the application has been greatly restricted by Darwin in the discovery of natural selection, but which was still to a large extent retained by him. Even the apparently powerful factors in transformation—the use and disuse of organs, the results of practice or neglect—cannot now be regarded as possessing any direct transforming influence upon a species. And the same is true of all the other direct influences, such as nutrition, light, moisture, and that combination of different influences which we call climate. All these, with use and disuse, may perhaps produce great effects upon the body (soma) of the individual, but cannot produce any effect in the transformation of the species, simply because they can never reach the germ-cells from which the succeeding generation arises. But if—as it seems to me—the facts of the case compel us to reject the assumption of the transmission of acquired characters, there only remains one principle by which we can explain the transformation of species—the direct alteration of the germ-plasm, however we may imagine that such alterations have been produced and combined to form useful modifications of the body.

The difficulty of understanding these processes of transformation is by no means lightened by abandoning the Lamarckian theory. The difficulty in fact becomes much greater, for we are now compelled to seek a different explanation of many phenomena which were previously believed to be understood. But this can hardly be regarded as a reason for not accepting the view: for we are in want of a correct explanation rather than one which is easy and convenient. We seek truth, and when we recognize that our path is leading in a wrong direction, we must leave it and take another road even if it presents more difficulties.

My theory rests, on the one hand, upon certain theoretical considerations which will be mentioned below, and which I have attempted to develope in previous papers276. On the other hand, it rests upon the want of any actual proof of the transmission of acquired characters. My theory might be disproved in two ways,—either by actually proving that acquired characters are transmitted, or by showing that certain classes of phenomena admit of absolutely no explanation unless such characters can be transmitted. It will be admitted, however, that we must be very cautious in accepting proofs of this latter kind, for the impossibility of explaining a given phenomenon may be merely temporary, and may disappear with the progress of science. No one could have explained the useful adaptations so common in animals and plants, before the light of the theory of natural selection had fallen on these phenomena; at that time we should have been far from right if we had assumed that organisms possess a power which causes them to respond to external influences by useful modifications, a power unknown elsewhere, entirely unproved and only supported by the fact that at that time it did not seem possible to explain the phenomena in any other way.

Although my theory has not been disproved, I will nevertheless attempt to bring into further accordance with it certain phenomena which seem at first sight to oppose it. I first began to take this course in my paper ‘On Heredity277.’ In that paper I attempted to show how the fact that disused organs become rudimentary may be readily explained without assuming the transmission of acquired characters; and also that the origin of instincts may in all cases be referred to the process of natural selection278, although many observers had followed Darwin in explaining them as inherited habits,—a view which becomes untenable if the habits adopted and practised in a single life cannot be transmitted.

Other phenomena which appeared to present difficulties were also considered and brought into accordance with the theory, and I think that I have been successful in showing that adequate and simple explanations may be given.

There certainly remain many phenomena which seem to be opposed to my theory and for which a new explanation must be found. Thus Romanes279, following Herbert Spencer280, has recently pointed to the phenomena of correlation as a proof of the transmission of acquired characters; but, at no distant time, I hope to be able to consider this objection, and to show that the apparent support given to the old idea is in reality insecure and breaks down as soon as it is critically examined. I believe that I shall be able to prove that correlation cannot be used as the indirect proof of an hypothesis, of which all direct evidence is still completely wanting. It must not be forgotten that the onus probandi rests with my opponents: they defend the assertion that acquired characters can be transmitted, and they ought therefore to bring forward actual proofs; for the mere fact that the assertion has been hitherto accepted as a matter of course by almost everyone, and has only been doubted by a very few (such as His, du Bois-Reymond, and Pflüger), cannot be taken as any proof of its validity. Not a single fact hitherto brought forward can be accepted as a proof of the assumption. Such proofs ought to be found: facts ought to be discovered which can only be understood with the aid of this hypothesis. If, for instance, it could be shown that artificial mutilation spontaneously re-appears in the offspring with sufficient frequency to exclude all possibilities of chance, then such proof would be forthcoming. The transmission of mutilations has been frequently asserted, and has been even recently again brought forward, but all the supposed instances have broken down when carefully examined. I think I may here safely omit all further reference to the proofs dependent upon transmitted mutilations, especially as Döderlein281 has already, in the most convincing manner, disposed of the argument derived from the tailless cats which were so triumphantly exhibited at the last meeting of the Association of German Naturalists282.

I now come to the real subject of this paper—the supposed botanical proofs of the transmission of acquired changes. The botanist Detmer has recently brought forward certain phenomena in vegetable physiology283, as a support for the transmission of such changes, and although I do not believe that they will bear this interpretation, the discussion of them may perhaps be useful. I am even inclined to think that these and a few other phenomena in vegetable physiology, upon which I shall also touch, are very likely to throw new light upon the whole question which has been so frequently misunderstood. I should have preferred to leave this discussion to a botanist, but I do not know whether my views will meet with any support from the followers of this subject, and I must therefore attempt the discussion myself. And perhaps it is of some assistance in clearing up the question, for one who is not accustomed to the usual botanical views, and is more conversant with other classes of biological knowledge, to consider the facts brought to light by modern botany, from a general point of view. Of course I shall not attempt to question the validity of the observations, nor even the accuracy with which the facts have been interpreted. I shall only deal with the conclusions which may be drawn from the facts, and I do not think that it is absolutely necessary that such criticism should be made by a botanist. Questions of general biological significance such as that of heredity cannot be entirely solved within the single domain of either zoological or botanical facts. Both botanists and zoologists must give due weight to the facts of the province which is not their own, and must see whether the views which they have chiefly gained in the one province can be applied to the other, or whether phenomena occur in the latter which are in opposition to their previously formed views and which cause them to be abandoned or modified.

Detmer begins by bringing forward certain facts which prove, as he believes, that rather important changes in the organism can be directly produced by external influences. He is of opinion that I under-estimate the weight of these influences, and that I make light of the changes which may thus arise in a single individual life. But obviously, it is of no importance for the question of the transmission of acquired characters, whether the changes directly produced by external influences upon the soma of an individual are greater or smaller: the only question is whether they can be transmitted. If they can be transmitted, the smallest changes might be increased by summation in the course of generations, into characters of the highest degree of importance. It is in this way that Lamarck and Darwin have supposed that an organism is transformed by external influences. It is therefore interesting to see what Detmer considers to be a change which has been directly effected. We can in this way gain a very distinct appreciation of the difference in views which is caused by the different spheres of experience which belong to botany and zoology. It will be useful to gain a clear idea of the differences which are thus caused.

Detmer first alludes to the dorso-ventral structure of the shoots of Thuja occidentalis, chiefly shown in the fact that the upper sides of these shoots contain the green palisade cells, while the under sides which are turned away from the light possess green spheroidal (isodiametric) cells. If the branches of Thuja are turned upside down and fixed in this position before the production of new shoots, it is found that the anatomical structure of the latter, when developed, is reversed. The side of the shoot which was destined to become the under side, but which was artificially compelled to become the upper side, assumes the structure of the upper side and developes the characteristic palisade parenchyma; and on the other hand, the under side which was intended to become the upper side developes the spongy parenchyma which is characteristic of the under side. From these facts Detmer concludes that the dorso-ventral structure of the shoots of Thuja has resulted from the continual operation of an external force, and that the light must be considered as the cause of the structural change.

But such a conclusion obviously depends upon a confusion of ideas. No one will doubt that the light was the stimulus which led to the reversal of the structures in the shoot, but this is a very different thing from maintaining that it was the cause which conferred upon the Thuja-shoot the power of producing palisade and spongy parenchyma. When a phenomenon only occurs under certain conditions, it does not follow that these conditions are the cause of the phenomenon. A certain temperature is necessary for the development of a bird in the egg, but surely no one will maintain that the temperature is the cause of the capacity for such development. It is obvious that the egg has acquired the power of producing a bird chiefly as the result of a long phyletic course of development which has led to such a chemical and physical structure in the egg and the fertilizing sperm-cell, that after their union and development, a bird, and only a bird of a particular species, must be produced. But of course certain conditions must be fulfilled in order that such development may take place; and a definite temperature is one of these conditions of development. Thus we may briefly say that the physical nature of the egg is the cause of its development into a bird, and we may similarly maintain that the physical nature of a Thuja-shoot, and not the influence of light, is the cause of the development of tissues which are characteristic of the species. In the development of such a shoot the light plays precisely the same part which is played by temperature in the development of a bird: it is one of the conditions of development.

There is nevertheless a difference between these two cases in that the Thuja-shoot possesses the possibility of development in two different ways instead of only one. The upper side of the shoot can assume the structure of the under side and vice versa, and this structural reversal depends upon the way in which the light is thrown upon the shoot. But even if the light causes the structural reversal, does this justify us in assuming that the structure itself is also the direct consequence of the influence of light? I see no reason for rejecting the supposition that the physical nature of part of a plant may be of such a kind that this or that structure may be produced according as this or that condition of development prevails. Thus with stronger light the structure of the upper side of the shoot developes; with weaker light, the structure of the under side. But this physical nature of the Thuja-bud depends, like that of a bird’s egg, upon its phyletic history, as we must assume to be the case with the germs producing all individual developments. It is therefore quite impossible to interpret the reversal of the structure in the Thuja-shoot as the result of modification produced by the direct influence of external conditions. It is an instance of double adaptation—one of those cases in which the specific nature of a germ, an organism, or a part of an organism, possesses such a constitution that it reacts differently under the incidence of different stimuli. An entirely analogous example of reversal occurs in the climbing shoots of the Ivy, and is described in Sachs’ lectures on the physiology of plants. Such shoots produce leaves only on the side directed towards the light, and roots (which are made use of in climbing) only upon the opposite side. If however the position of the plant be altered so that the root-bearing side is turned towards the light, while the leafy side is shaded, a reversal occurs, so that from that time the former only produces leaves, and the latter nothing but roots. In other words, the Ivy-shoot reacts under strong light with the production of leaves and under weak light with the production of roots, just as litmus-paper becomes red with an acid and blue with an alkali. The physical nature of the Ivy-shoot was present before the production of either structure, and was no more due to the action of light itself, than the physical nature of litmus-paper is due to an acid or an alkali. But this is quite consistent with the possession of a physical nature which reacts differently under the two different conditions afforded by light and shade.

No one would think of bringing forward the changes in the colour of the green frog (Hyla) as a proof of the power of direct influences in causing structural modifications in the animal body. The frog is light green when it is resting upon green leaves, but it becomes dark brown or nearly black when transferred to dark surroundings. This is an obvious instance of adaptation, for the changes in the colour of the frog depend upon a complex reflex mechanism. The changes in the shape of the chromatophores of the skin are not produced by the direct influence of the different rays of light upon the body-surface, but in consequence of the action of these rays upon the retina. Blind frogs do not react under the changes of light. Hence it is impossible that any one can maintain that the skin of the frog has gained its green colour as the direct result of the green light reflected from its usual surroundings. It must be admitted that in this and in all similar cases, there is only one possible explanation, viz. an appeal to the operation of natural selection. It may be objected that we are not here dealing, as in the Thuja and Ivy, with changes in the course of ontogenetic development following upon the occurrence of this or that external condition, but only with the different reactions of a mature organism. But nevertheless, cases of the former kind appear to be also present in the animal kingdom.

Thus the very careful and extensive investigations of Poulton284 upon the colours of certain caterpillars have distinctly shown that some species possess the possibility of development in two directions, and that the actual direction taken by the individual is decided by the influence of external conditions. Poulton surrounded certain larvae of Geometrae with an abundance of dark branches, in addition to the leaves upon which they fed. When such conditions prevailed from the beginning of larval life, the caterpillars as they developed, gradually assumed the dark colour of the twigs and branches upon which they rested. When other larvae of the same species (and in many experiments hatched from the same batch of eggs) were similarly exposed to the green leaves of the same food-plant, they did not indeed become bright green like the leaves, but were invariably of a much lighter colour than the other larvae, while many of them gained a brownish-green tint. The larvae of Smerinthus ocellatus285 also possess the power of assuming different shades of green and of thus approaching, to some extent, the green of the plant upon which they happen to live. It is quite impossible to explain the phyletic development of the green colour of these and other caterpillars as due to the direct action upon the skin of the green light reflected from the leaves upon which they sit. The impossibility of such an effect was pointed out long ago by Darwin, and also followed from my own investigations. Here, as in the other cases, the only possible solution is afforded by natural selection. The colour of the caterpillars has become gradually more and more perfectly adapted to the colour of the leaves,—and often to the particular side of the leaves upon which these animals rest,—not by the direct effect of reflected light, but by the selection of those individuals which were best protected. Poulton’s experiments quoted above prove that certain species which occur upon different plants with different colours (or even in some cases upon the differently coloured parts of the same plant), present us with a further complication in the process of adaptation, inasmuch as each individual has acquired the power of assuming a lighter or darker colour286. The light which falls upon a single individual caterpillar during the course of its growth determines whether the lighter or darker colour shall be developed. Here therefore we have a case exactly parallel to that of the Thuja-shoot in which the palisade or spongy parenchyma is developed according to the position in which the shoot is fixed.

As far as it is possible in the present condition of our knowledge to offer any opinion upon the origin of sex in bisexual animals, it may be suggested that this problem is also capable of an essentially similar solution. Each germ-cell may possess the possibility of developing in either of two directions, the one resulting in a male individual, and the other resulting in a female, while the decision as to which of the two possible alternatives is actually taken may rest with the external conditions. We must, however, include among the external circumstances everything which is not germ-plasm. Moreover, this explanation is by no means certain, and I only mention it as an instance which, if we assume it to be correct, further illustrates my views upon the phenomena presented by the Thuja-shoot.