Essays Upon Heredity and Kindred Biological Problems

If I am not mistaken, this argument lends additional support to the idea which I have previously propounded,—that the most important duty of sexual reproduction is to preserve and continually call forth individual variability, the foundation upon which the transformation of species is built[265 - See the preceding Essay (#x22_x_22_i94) on ‘The Significance of Sexual Reproduction in the theory of Natural Selection.’].

But if it be asked whether the postponement of the ‘reducing division’ to the end of the ovarian development of the egg is inconsistent with the preservation of the other half of the dividing nucleus, I should be inclined to reply that a ‘reducing division’ of the mature egg, resulting in the production of two eggs, was probably the phyletic precursor of the present condition. I imagine that the division of the mature egg-cell—although it is now so extremely unequal—was equal in very remote times; but that for reasons of utility, connected with the specialization of the eggs of animals, it gradually became more and more unequal. It is now hardly possible to give in detail the various reasons of utility which have brought about this condition, but it may be assumed that the enormous size attained by many animal egg-cells has been especially potent in producing the change.

A careful consideration of this last point seems to me to be demanded by a comparison of the egg-cells with the male germ-cells. Just as the female germ-cells of animals are distinguished by the attainment of a large size, the male germ-cells are generally remarkable for their minute proportions. In most cases it would be physiologically impossible for a large egg-cell, rich in yolk, to attain double its specific size in order to undergo division into two equal halves and yet to remain of the characteristic size. Even without the additional difficulties imposed by the necessity for such a division, all means—such as cells used as food, or the passage of food from follicular cells into the ovum, etc.—are employed in order to bring the egg-cell to the greatest attainable size. Furthermore, the ‘reducing division’ of the nucleus cannot take place before the egg has attained its full size, because the ovogenetic nucleoplasm still controls the egg-cell, and must be removed before the germ-plasm can regulate its development. By arguments such as these I should attempt to render the whole subject intelligible.

But the case is entirely different with the sperm-cells, which are generally minute: here it is quite conceivable that a ‘reducing division’ of the nuclei may take place by an equal division of the sperm-cells, occurring towards the end of the period of their formation; that is to say, in such a way that both products of division remain sperm-cells, and neither of them perishes like the polar bodies. But the other possibility also demands consideration, viz. that the reducing division may occur at an earlier stage in the development of sperm-cells. At all events, the arguments adduced above, which proved that the consequence would be a want of variability in the egg-cells, would not apply to an equal extent in the case of the male germ-cells. Among the egg-cells it may be very important that each one should have its special individual character, produced by a somewhat different composition of its germ-plasm, inasmuch as a considerable proportion of the eggs frequently developes, although this is never the case with all of them. But the production of sperm-cells is in most animals so enormous that only a very small percentage can be used for fertilization. If, therefore, e. g. ten or a hundred spermatozoa contained germ-plasm with exactly the same composition, so that, as far as the paternal influence is concerned, ten or a hundred identical individuals would result if they were all used in fertilization, such an arrangement would be practically harmless, for only one spermatozoon out of an immense number would be employed for this purpose. From this point of view we might expect that the ‘reducing division’ of the sperm-nucleus would not take place at the end of the development of the sperm-cell, but at some earlier period. There is no necessary reason for the assumption that this division must take place at the end of development, and without some cause natural selection cannot operate. It is, of course, conceivable that the causes of other events may also involve the occurrence of this division at the end of development; but we do not at present know of any such causes. I should not consider the influence of the specific histogenetic nucleoplasm, i.e. the spermatogenetic nucleoplasm, to be such a cause, because the quantitative proportions are very different from those which obtain in the formation of egg-cells, and because it is not inconceivable that the small quantity of true germ-plasm which must be present in the nuclei of the sperm-cells at every stage in their formation might enter upon a ‘reducing division’ with the spermatogenetic nucleoplasm, even when the latter preponderated.

As soon as we can recognize with certainty the forms of nuclear division which are ‘reducing divisions,’ the question will be settled as far as spermatogenesis is concerned. It has been already established that various forms of nuclear division occur at different periods of spermatogenesis. I make this assertion, not only from my own observations, but also from observations which have been made and insisted upon by others. Thus, van Beneden and Julin[266 - E. van Beneden and Julin, ‘La Spermatogénèse chez l’Ascaride mégalocéphale.’ Brussels, 1884.] stated in 1884 that direct and karyokinetic nuclear divisions alternate with each other in the spermatogenesis of Ascaris megalocephala. Again, Carnoy[267 - Carnoy, ‘La Cytodiérèse chez les Arthropodes.’] distinctly states that the different cell-generations in the same testis may not uncommonly exhibit considerable differences as regards karyokinesis. ‘This may go so far that direct and indirect division may proceed simultaneously.’ Platner[268 - Gustav Platner, ‘Die Karyokinese bei den Lepidopteren als Grundlage für eine Theorie der Zelltheilung.’ Internation. Monatsschrift f. Anatomie und Histologie, Bd. III. Heft 10. Leipzig, 1886.], in his excellent paper on karyokinesis in Lepidoptera, also points out that the karyokinesis of the spermatocytes is essentially different from that of the spermatogonia. According to his description, the latter form may be very well interpreted as a ‘reducing division,’ for no equatorial plate is formed, and the chromatin rods (or granules, as they are better called in this case) remain from the first on both sides of the equatorial plane, and finally unite at the opposite poles to form the two daughter-nuclei. Furthermore, if Carnoy has correctly observed, the form of karyokinesis which I have previously interpreted as a ‘reducing division’ occurs in the sperm-mother-cells—a karyokinesis in which the chromatin rods either do not divide longitudinally, or else divide in this way after they have left the equatorial plate and are proceeding towards the poles. Carnoy does not himself attach any special importance to these observations, for he only considers them as proofs that the longitudinal splitting of the loops may occur at various periods in different species—either at the equator, or on the way towards the poles, or even at the poles themselves. We cannot conclude from the author’s statements whether this form of nuclear division only occurs in a single cell-generation during spermatogenesis, as it must do if it really represents a ‘reducing division.’ Until this point is settled, we cannot decide with certainty whether the described form of karyokinesis is to be considered as the ‘reducing division’ for which we are seeking. Fresh investigations, undertaken from these points of view, are necessary in order to settle the question. It would be useless to seek further support for the theory by going into further details, and by critically examining the numerous observations upon spermatogenesis which have now been recorded.

I will only mention that among the various nuclei and other bodies in different animals which have been considered by different observers as the polar bodies of the sperm-cells, or the cells which form the latter—in my opinion the paranucleus (‘Nebenkern’) of the ‘spermatides’ described by La Valette St. George[269 - La Valette St. George, ‘Ueber die Genese der Samenkörper.’ Fünfte Mittheilung. Die Spermatogenese bei den Säugethieren und dem Menschen,’ Archiv f. mikrosk. Anat. Bd. XV. 1878.] has the highest claim to be considered as the homologue of a polar body. But I am inclined to identify it with the first rather than the second polar body of the egg-cells, and to regard it as the histogenetic part of the nucleoplasm which has been expelled or rendered powerless by internal transformations. There are two reasons which lead me to this conclusion: first, as I have tried to show above, it is probable that the ancestral germ-plasms are not removed by expulsion, but by means of equal cell-division; secondly, my theory asserts that the histogenetic nucleoplasm cannot be rendered powerless until the close of histological differentiation.

The whole question of the details of the transformations undergone by the nucleus of the male germ-cells is not ready for the expression of a mature opinion. From the very numerous and mostly minute and careful observations which have been hitherto recorded, we cannot conclude with any degree of certainty when and how the ‘reducing division’ of the nucleus takes place, nor can we decide upon the processes which signify the purification of the germ-plasm from the merely histogenetic part of the nucleoplasm. But perhaps it has not been without value as regards future investigation that I have tried to apply to the male germ-cells the views gained from our more certain knowledge of the corresponding structures in the female, and thus to indicate the problems which now chiefly demand solution.

IV. The Foregoing Considerations applied To Plants

It remains to briefly consider the case of plants. Obviously, the ‘reducing division’ of the germ-nuclei, if it takes place at all, cannot be restricted to the germ-cells of animals. There must be a corresponding process in plants, for sexual reproduction is essentially the same in both kingdoms; and if fertilization must be preceded by the expulsion of half the number of ancestral germ-plasms from the eggs of animals, the same necessity must hold in the case of plants.

But whether the process always takes place in the form of polar bodies, and not perhaps principally, or at any rate frequently, in the form of equal cell-division, is another question. It is true that polar bodies occur in numerous plants, as we chiefly know from Strasburger’s researches[270 - Weismann, ‘Studien zur Descendenztheorie,’ ii. p. 306, Leipzig, 1876, translated by Meldola; see ‘Studies in the Theory of Descent,’ p. 680.]. Strasburger shows that cells are separated by division from the germ-cells, and perish. But it seems to me doubtful whether we must always regard their formation as the removal of half the number of ancestral germ-plasms rather than the histogenetic nucleoplasm of the germ-cell. It appears to me that histogenetic nucleoplasm must be present in the highly differentiated vegetable germ-cells, especially in the male cells, and also that it must be removed during the maturation of the cell, if my idea of the histogenetic nucleoplasm be accepted. It is very possible, as I have already mentioned, that there may be quite indifferent germ-cells, viz. cells which are entirely without specific histological structure, and in such cases histogenetic nucleoplasm would be absent; and during the maturation of such germ-cells no polar body would be formed for its removal. This view accords with the fact that polar bodies are absent in many plants. Furthermore, I am far from maintaining that in the cases where polar bodies occur, they must have the above-mentioned significance. I only wish to point out that the reduction assumed to be necessary for the nucleus of the vegetable germ-cells is not necessarily to be sought for at the close of their maturation, but perhaps even more frequently in an equal division of the germ-cells during some period of their development.

It also seems to me to be not impossible that a number of these vegetative ‘polar bodies’ may have an entirely different significance, viz. to perform some special function accessory to fertilization, as in the so-called ‘ventral canal-cells’ of the higher cryptogams and conifers. As we know that even the two polar bodies of the animal egg are not identical—although externally they are extremely similar, and although they arise in a precisely similar manner—I am even more inclined than before to consider that the very various ‘polar bodies’ of plants possess very different meanings.

But I do not feel justified in criticizing in detail the results of botanical investigation. I must leave the decision of such questions to botanists, and I only desire to state distinctly that a ‘reducing division’ of the nuclei of germ-cells must occur in plants as well as in animals.

V. Conclusions with regard to Heredity

The ideas developed in the preceding paragraphs lead to remarkable conclusions with regard to the theory of heredity,—conclusions which do not harmonize with the ideas on this subject which have been hitherto received. For if every egg expels half the number of its ancestral germ-plasms during maturation, the germ-cells of the same mother cannot contain the same hereditary tendencies, unless of course we make the supposition that corresponding ancestral germ-plasms are retained by all eggs—a supposition which cannot be sustained. For when we consider how numerous are the ancestral germ-plasms which must be contained in each nucleus, and further how improbable it is that they are arranged in precisely the same manner in all germ-cells, and finally how incredible it is that the nuclear thread should always be divided in exactly the same place to form corresponding loops or rods,—we are driven to the conclusion that it is quite impossible for the ‘reducing division’ of the nucleus to take place in an identical manner in all the germ-cells of a single ovary, so that the same ancestral germ-plasms would always be removed in the polar bodies. But if one group of ancestral germ-plasms is expelled from one egg, and a different group from another egg, it follows that no two eggs can be exactly alike as regards their contained hereditary tendencies: they must all differ. In many cases the differences will only be slight, that is, when the eggs contain very similar combinations of ancestral germ-plasms. Under other circumstances the differences will be very great, viz. when the combinations of ancestral germ-plasms retained in the egg are very different. I might here mention various other considerations; but this would lead me too far from my subject, into new theories of heredity. I hope to be able at some later period to develope further the theoretical ideas which are merely indicated in the present essay. I only wish to show that the consequences which follow from my theory upon the second division of the egg-nucleus, and the formation of the second polar body, are by no means opposed to the facts of heredity, and even explain them better than has hitherto been possible.

The fact that the children of the same parents are never entirely identical could hitherto only be rendered intelligible by the vague suggestion that the hereditary tendencies of the grandfather predominate in one, and those of the grandmother in another, while the tendencies of the great-grandfather predominate in a third, and so on. Any further explanation as to why this should happen was entirely wanting. Others even looked for an explanation to the different influences of nutrition, to which it is perfectly true that the egg is subjected in the ovary during its later development, according to its position and immediate surroundings. I had myself referred to these influences as a partial explanation[271 - l. c., p. 92.], before I recognized clearly how extremely feeble and powerless are the influences of nourishment, as compared with hereditary tendencies. According to my theory, the differences between the children of the same parents become intelligible in a simple manner from the fact that each maternal germ-cell (I shall speak of the paternal germ-cells later on) contains a peculiar combination of ancestral germ-plasms, and thus also a peculiar combination of hereditary tendencies. These latter by their co-operation also produce a different result in each case, viz. the offspring, which are characterized by more or less pronounced individual peculiarities.

But the theory which explains individual differences by referring to the inequality of germ-cells, may be proved with a high degree of probability by an appeal to facts of an opposite kind, viz. by showing that identity between offspring only occurs when they have arisen from the same egg-cell. It is well known that occasionally some of the children of the same parents appear to be almost exactly alike, but such children are without exception twins, and there is every reason to believe that they have been derived from the same egg. In other words, the two children are exactly alike because they have arisen from the same egg-cell, which could of course only contain a single combination of ancestral germ-plasms, and therefore of hereditary tendencies[272 - [The similar conclusion that identical ova lead to the appearance of identical individuals was drawn from the same data by Francis Galton in 1875. See ‘The history of the Twins, as a criterion of the relative powers of Nature and Nurture,’ by Francis Galton, F.R.S., Journal of the Anthropological Institute, 1875, p. 391; also by the same author, ‘Short Notes on Heredity, etc. in Twins,’ in the same Journal, 1875, p. 325.The author investigated about eighty cases of close similarity between twins, and was able to obtain instructive details in thirty-five of these. Of the latter there were no less than seven cases ‘in which both twins suffered from some special ailment or had some exceptional peculiarity;’ in nine cases it appeared that ‘both twins are apt to sicken at the same time;’ in eleven cases there was evidence for a remarkable association of ideas; in sixteen cases the tastes and dispositions were described as closely similar. These points of identity are given in addition to the more superficial indications presented by the failure of strangers or even parents to distinguish between the twins. A very interesting part of the investigation was concerned with the after-lives of the thirty-five twins. ‘In some cases the resemblance of body and mind had continued unaltered up to old age, notwithstanding very different conditions of life,’ in the other cases ‘the parents ascribed such dissimilarity as there was, wholly, or almost wholly, to some form of illness.’The conclusions of the author are as follows: ‘Twins who closely resembled each other in childhood and early youth, and were reared under not very dissimilar conditions, either grow unlike through the development of natural characteristics which had lain dormant at first, or else they continue their lives, keeping time like two watches, hardly to be thrown out of accord except by some physical jar. Nature is far stronger than nurture within the limited range that I have been careful to assign to the latter.’ And again, ‘where the maladies of twins are continually alike, the clocks of their two lives move regularly on, and at the same rate, governed by their internal mechanism. Necessitarians may derive new arguments from the life histories of twins.’The above facts and conclusions held for twins of the same sex, of which at any rate the majority are shown by Kleinwächter’s observations to have been enclosed in the same embryonic membranes, and therefore presumably to have been derived from a single ovum; but in rarer cases the twins, although also invariably of the same sex, were marked by remarkable differences, greater than those which usually distinguish children of the same family. Mr. Galton met with twenty of these cases. In such twins the conditions of training, etc. had been as similar as possible, so that the evidence of the power of nature over nurture is strongly confirmed. Mr. Galton writes, ‘I have not a single case in which my correspondents speak of originally dissimilar characters having become assimilated through identity of nurture. The impression that all this evidence leaves on the mind is one of wonder whether nurture can do anything at all, beyond giving instruction and professional training.’The fact that twins produced from a single ovum seem to be invariably of the same sex is in itself extremely interesting, for it proves that the sex of the individual is predetermined in the fertilized ovum.—E. B. P.]]. The factors which by their co-operation controlled the construction of the organism were the same, and consequently the results were also the same. Twins derived from a single egg are identical: this is a statement which, although not mathematically proved, may be looked upon as nearly certain. But there are also twins which do not possess this high degree of similarity, and these are even far commoner than the others. The explanation is to be found in the fact that the latter were derived from two egg-cells which were fertilized at the same time. In most cases, indeed, each twin is enclosed in its own embryonic membranes, while much less frequently both twins are enclosed in the same membranes. In one point only the proof is incomplete; for it has not yet been shown that identical twins are always derived from a single egg, since such an origin, together with a high degree of similarity, could only be established as occurring together in a small proportion of the cases. We therefore see that under conditions of nutriment which are as identical as possible, two egg-cells develope into unlike twins, one into identical twins; although we cannot yet affirm that the latter result invariably follows. It is conceivable that the stimulus for the production of two eggs from one may be afforded by the entrance of two spermatozoa, but these latter, as was shown above, could hardly contain identical hereditary tendencies, and thus two identical twins would not arise. It appears indeed that some cases have been observed in which differences have been exhibited by twins which were enclosed in the same embryonic membranes; but nevertheless I believe that two spermatozoa are not necessary to cause the formation of twins by a single egg. We know, it is true, from the investigations of Fol[273 - Fol, Recherches sur la fécondation et le commencement de l’hénogénie: Genève, Bâle, Lyon. 1879.], that multiple impregnation produces the simultaneous beginning of several embryos in the eggs of star-fishes. But several embryos and young animals are not developed in this way, for embryonic development soon ceases, and the egg dies.

The recent observations of Born[274 - Born, ‘Ueber Doppelbildungen beim Frosch und deren Entstehung.’ Breslauer ärztl. Zeitschrift, 1882.] 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 1883[275 - See the second Essay (#x9_x_9_i90).], 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 papers[276 - Consult ‘Ueber die Vererbung,’ Jena, 1883; ‘Die Kontinuität des Keimplasmas,’ Jena, 1885; ‘Ueber die Zahl der Richtungskörper und über ihre Bedeutung für die Vererbung,’ Jena, 1887. These papers are translated as the second (#x9_x_9_i90), fourth (#x16_x_16_i30) and sixth (#x28_x_28_i58) Essays in the present volume.]. 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 Heredity[277 - See the second (#x9_x_9_i90) Essay.].’ 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 selection[278 - [See R. Meldola in Ann. and Mag. Nat. Hist., 1878, vol. i. pp. 158-161. The author discusses many cases among insects in which instinct is related to protective structure or colouring: he also considers that instinct is to be explained by the principle of natural selection which accounts for the other protective features.—E. B. P.]], 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 Romanes[279 - [See ‘Nature,’ vol. 36, pp. 491-507.—E. B. P.]], following Herbert Spencer[280 - [See ‘The Factors of organic Evolution’ in ‘The Nineteenth Century’ for April and May 1886.—E. B. P.]], 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 Döderlein[281 - See ‘Biol. Centralbl.’ Bd. VII. No. 23.] 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 Naturalists[282 - See the next Essay (VIII) (#x35_x_35_i25).].

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 physiology[283 - Detmer, ‘Zum Problem der Vererbung,’ Pflüger’s Archiv f. Physiologie, Bd. 41, (1887), p. 203.], 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 Poulton[284 - [Dr. Weismann is here alluding to experiments upon the larvae of Rumia Crataegata. A short account of the results will be found in the Report of the British Association at Manchester (1887), and in ‘Nature,’ vol. 36, p. 594. I have now obtained similar results with many other species (see Trans. Ent. Soc., Lond. 1888, p. 553); but many of the results are as yet unpublished.—E. B. P.]] 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 ocellatus[285 - [See the editorial notes by Raphael Meldola, in his translation of Weismann’s ‘Studies in the Theory of Descent’ (the Essay on ‘The Origin of the Markings of Caterpillars,’ pp. 241 and 306): also E. B. Poulton, in ‘Proc. Roy. Soc.,’ vol. xxxviii. pp. 296-314; and in ‘Proc. Roy. Soc.,’ vol. xl. p. 135.—E. B. P.]] 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 colour[286 - [Professor Meldola first called attention to the scattered instances of the kind here alluded to by Professor Weismann, in 1873: see ‘Proc. Zool. Soc.,’ 1873, p. 153. The author explains the relation of this ‘variable protective colouring’ to other protective appearances, and he is strongly of the opinion that the former as well as the latter is to be explained by the action of the ‘survival of the fittest.’The validity of Dr. Weismann’s interpretation of these effects as due to adaptation, through the operation of natural selection, is conclusively proved by the following facts. The light reflected from green leaves becomes the stimulus for the production of dark brown pigment in those cases in which the leaves constitute the surroundings for many months. Under these circumstances the leaves of course become brown at a relatively early date, and protection is thus afforded for the remainder of the period, although the dark pigment is produced before the change in the colour of the leaf. Instances of this kind are seen in the colours of cocoons spun among leaves by certain lepidopterous larvae (see ‘Proc. Ent. Soc. Lond.,’ 1887, pp. l, li, and 1888, p. xxviii), the cocoons of the same species being of a creamy white colour when spun upon white paper.Conversely, the light reflected from the same surfaces serves as the stimulus for withholding pigment in the cases alluded to by Dr. Weismann (larvae of R. Crataegata, &c.), in all of which the organism only remains in contact with the leaves while they are green, viz. at a time when the dark colour would be disadvantageous.Hence precisely opposite effects are produced by the operation of the same force; the nature of the effect which actually follows in any case being solely determined by the advantage afforded to the organism.—E. B. P.]]. 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.

The two other facts brought forward by Detmer as proofs of the transforming power of external influences can be explained in precisely the same manner. These instances are—the fact that Tropaeolum when grown in moist air produces leaves with anatomical characters different from those produced when the plant is grown in dry air; and the differences in the structure of the leaves of many plants, according as they have been grown in the sun or shade respectively. Such differences do not by any means afford proof of the direct production of structural changes by means of external influences. How would such an explanation be consistent with the fact that the leaves are, in all these cases, changed in a highly purposeful manner? Or is it assumed that these organs were so constituted from the beginning, that they are compelled to respond to external conditions by the production of useful changes? Any one who made such an assertion nowadays, or who even thought of such a thing as a possibility, would prove that he is entirely ignorant of the facts of organic nature, and that he has no claim to be heard upon the question of the transformation of species. The very first necessity in any scientific question is to gain acquaintance with that which has been thought and said upon the subject. And it has been frequently shown that whole groups of useful characters cannot by any possibility have been produced by the direct action of external influences. If a caterpillar, which hides itself by day in the crevices of the bark, possesses the same colour as the latter, while other caterpillars which rest on leaves are of a green colour, these facts cannot be explained as the results of the direct influence of the bark and leaves. And it would be even less possible to explain upon the same principle all the details of marking and colour by which these animals gain still further protection. If the upper side of the upper wings of certain moths is grey like the stone on which they rest by day, while in butterflies the under side of both wings which are exposed during rest, exhibits analogous protective colours, these facts cannot be due to the direct influence of the surroundings which are resembled, but, if they have arisen in any natural manner, they must have been indirectly produced by the surroundings. One may reasonably complain when compelled to repeat again and again these elements of knowledge and of thought upon the causes of transformation!