Weismann August
Essays Upon Heredity and Kindred Biological Problems
In Figure II a nuclear spindle is seen at A, and at its equatorial zone there are twelve primary loops. The transverse cross-lines and other markings on the loops indicate that they are composed of different ancestral germ-plasms. The loops are shaded differently in order to render the diagram clear. At B six of the loops are seen to have moved to either pole, so that the figure is a representation of the ‘reducing division.’ Figure III is a diagrammatic representation of ‘equal division.’ The six loops at the equatorial zone of A are shown by different cross-lining and shading to be composed of different ancestral germ-plasms. The loops split longitudinally in a direction indicated by the longitudinal line upon each of them. In B the halves of the loops are seen to have moved to the opposite poles of the spindle, so that there are not only six loops at each pole, but also all the six combinations of ancestral germ-plasms.
Figs. II, III.
Perhaps some may be inclined to look upon direct nuclear division as a ‘reducing division,’ but I believe that such a view would be incorrect. It is only approximately true that the nuclear thread is divided into two halves of equal quantity by direct division, and exact equality would only happen as it were accidentally; so that we cannot speak of a perfectly equal distribution of the ancestral germ-plasm in the two daughter-nuclei. But the ‘reducing division’ must obviously effect an exactly regular and uniform distribution of the ancestral germ-plasms, although this does not imply that every ancestral germ-plasm of the mother-nucleus would be represented in each of the two daughter-nuclei. But if out of e.g. eight nuclear loops at the equatorial plane, four pass into one, and the other four into the other daughter-nucleus, each of the latter will contain an equal number of ancestral germ-plasms, although different ones. This is indeed part of the foundation of the theory, for the ‘reducing division’ must remove exactly half of the original number of ancestral germ-plasms, and precisely the same number must be replaced at a later period by the sperm-nucleus. This could hardly be achieved with sufficient precision by direct nuclear division.
I now come to inquire whether the expulsion of the second polar body is in reality, as I have already maintained, a reduction in the number of ancestral germ-plasms present in the nucleus of the egg. The view itself is sufficiently obvious, and it would supply an explanation of the meaning of the process which is still greatly wanted; but it will nevertheless be not entirely useless to consider other possible theories.
It would be quite conceivable to suppose that the youngest egg-cells, which multiply by division, may undergo one ‘reducing division’ in addition to the ordinary process. Of course this should occur once only, for if repeated, the number of ancestral idioplasms in the nucleus of the germ-cell would undergo a decrease greater than could be afterwards compensated by the increase due to fertilization. Thus the number of ancestral germ-plasms would continually decrease in the course of generations,—a process which would necessarily end with their complete reduction to a single kind, viz. to the paternal or the maternal germ-plasm. But the occurrence of such a result is disproved by the facts of heredity. Although such an early occurrence of the ‘reducing division’ would offer advantages in that nothing would be lost, for both daughter-nuclei would become eggs, instead of one of them being lost as a polar body, nevertheless I do not believe that it really occurs: weighty reasons can be alleged against it.
Above all, the facts of parthenogenesis are against it. If the number of ancestral germ-plasms received from the parents were reduced to half in the ovary of the young animal, how then could parthenogenetic development ever take place? It is true that we cannot at once assert the impossibility of an early ‘reducing division’ on this account, for as I have shown above, the power to develope parthenogenetically depends upon the quantity of germ-plasm contained in the mature egg; the necessary amount might be produced by growth, quite independently of the number of different kinds of ancestral germ-plasms which form its constituents. The size of a heap of grains may depend upon the number of grains, and not upon the number of different kinds of grains. But in another respect such a supposition would lead to an unthinkable conclusion. In the first place, the number of ancestral germ-plasms in the germ-cells would be diminished by one half in each new generation arising by the parthenogenetic method; thus after ten generations only 1/1024 of the original number of ancestral germ-plasms would be present.
Now, it might be supposed that the ‘reducing division’ of the young egg-cells was lost at the time when the parthenogenetic mode of reproduction was assumed by a species; but this suggestion cannot hold, because there are certain species in which the same eggs can develope either sexually or parthenogenetically (e.g. the bee). It seems to me that such cases distinctly point to the fact that the reduction in the number of ancestral germ-plasms must take place immediately before the commencement of embryonic development, or, in other words, at the time of maturation of the egg. It is only decided at this time whether the egg of the bee is to develope into an embryo by the parthenogenetic or the sexual method; such decision being brought about, as was shown above, by the fact that only one polar body is expelled in the first case, while two are expelled in the second. But if we are obliged to assume that reproduction by means of fertilization, necessarily implies a reduction to one half of the number of ancestral germ-plasms inherited from the parents,—the further conclusion is obvious, that the second division of the egg-nucleus and the expulsion of the second polar body represent such a reduction, and that this second division of the egg-nucleus is unequal in the sense mentioned above, viz. one half of the ancestral germ-plasms remains in the egg-nucleus, the original number being subsequently restored by conjugation with a sperm-nucleus; while the other half is expelled in the polar body and perishes.
I may add that observations, so far as they have extended to such minute processes, do indeed prove that the number of loops is reduced to one half. It has been already mentioned that, according to Carnoy, such reduction occurs in Ascaris megalocephala, but the same author also describes the process of the formation of polar bodies in a large number of other Nematodes263, and his descriptions show that the process occurs in such a way that the number of ancestral germ-plasms must be reduced by half. Sometimes half the number of primary loops pass into the nucleus of the polar body, while the other half remains in the egg. In other cases, as in Ophiostomum mucronatum, the primary nuclear rods divide transversely,—a process which must produce the same effect. It is true that these observations require confirmation, and since, with unfavourable objects, the difficulties of observation are extremely great, there may have been errors of detail; but I do not think that there is any reason for doubting the accuracy of the essential point. And this essential point is the fact that the number of primary loops is divided into half by the formation of the polar body.
But even if we could not admit that such a conclusion is securely founded, it cannot be doubted that the formation of the second polar body reduces to one half the quantity of the nucleus which would have become the segmentation-nucleus in the parthenogenetic development of the egg. This is a simple logical conclusion from the two following facts: first, parthenogenetic eggs expel only one polar body; secondly, there are eggs (such as those of the bee) in which it is absolutely certain that the same half of the nucleus—which is expelled as the second polar body in the egg requiring fertilization—remains in the egg when it is to develope parthenogenetically, and acts as half of the segmentation-nucleus. But this proves that the expelled half of the nucleus must consist of true germ-plasm, and thus a secure foundation is laid for the assumption that the formation of the nucleus of the second polar body must be considered as a ‘reducing division.’
I was long ago convinced that sexual reproduction must be connected with a reduction in the number of ancestral germ-plasms to one half, and that such reduction was repeated in each generation. When, in 1885, I brought forward my theory of the continuity of the germ-plasm, I had long before that time considered whether the formation and expulsion of polar bodies must not be interpreted in this sense. But the two divisions of the egg-nucleus caused me to hesitate. The two divisions did not seem to admit of such an interpretation, for by it the quantity of the nucleus is not divided into halves, but into quarters. But a division of the number of ancestral germ-plasms into quarters would have caused, as was shown above, a continuous decrease, leading to their complete disappearance; and such a conclusion is contradicted by the facts of heredity. For this reason I was led at that time to oppose Strasburger’s view that the expulsion of the polar bodies means a reduction of the quantity of nuclear substance by only half. My objection to such a view was valid when I said that the quantity of idioplasm contained in the egg-nucleus is not, as a matter of fact, reduced to one half, but to one quarter, inasmuch as two successive divisions take place. I may add that I had also considered whether the two successive divisions might not possess an entirely different meaning,—whether one of them led to the removal of ovogenetic nucleoplasm, while the other resulted in a reduction in the number of ancestral germ-plasms. But at that time there were no ascertained facts which supported the supposition of such a difference, and I did not wish to bring forward the idea, even as a suggestion, when there was no secure foundation for it. The morphological aspects of the formation of the first and second polar bodies are so extremely similar that such a supposition might have been considered as a mere effort of the imagination.
Hensen264 also rejected the second part of the supposition that reduction must take place in the number of the hereditary elements of the egg, and that such reduction is caused by the expulsion of polar bodies, because he believed it to be incompatible with the fact, which had just been discovered, that polar bodies are formed by parthenogenetic eggs. He concludes with these words: ‘If this striking fact be confirmed, the hypothesis which assumes that the egg must be divided into half before maturation, is refuted, and there only remains the rather vague explanation that a process of purification must precede the development of the embryo.’ Nevertheless Hensen is the only writer who has hitherto taken into consideration the idea that sexual reproduction causes a regularly occurring ‘diminution in the hereditary elements of the egg.’
III. The Foregoing Considerations Applied To the Male Germ-Cells
If the result of the previous considerations be correct, and if the number of ancestral germ-plasms contained in the nucleus of the egg-cell destined for fertilization must be reduced by one half, there can be no doubt that a similar reduction must also take place, at some time and by some means, in the germ-plasms of the male germ-cells. This must be so if we are correct in maintaining that the young germ-cells of a new individual contain the same nuclear substance, the same germ-plasm, which was contained in the fertilized egg-cell from which the individual has been developed. The young germ-cells of the offspring must contain this substance if my theory of the continuity of the germ-plasm be well founded, for this theory supposes that, during the development of a fertilized egg, the whole quantity of germ-plasm does not pass through the various stages of ontogenetic development, but that a small part remains unchanged, and at a later period forms the germ-cells of the young organism, after having undergone an increase in quantity. According to this supposition therefore the germ-plasm of the parents must be found unchanged in the germ-cells of the offspring. If this theory were false, if the germ-plasm of the germ-cells were formed anew by the organism, perhaps from Darwin’s ‘gemmules’ which pour into the germ-cells from all sides, it would be impossible to understand why it has not been long ago arranged that each germ-cell should receive only half the number of the ancestral gemmules present in the body of the parent. Hence the expulsion of the second polar body—assuming the validity of my interpretation—is an indirect proof of the soundness of the theory of the continuity of the germ-plasm, when contrasted with the theory of pangenesis. If furthermore, a kind of cyclical development of the idioplasm took place, as supposed by Strasburger, and if its final ontogenetic stage resulted in the re-appearance of the initial condition of the germ-plasm, we should fail to understand how any of the ancestral germ-plasms could be lost during such a course of development.
Whichever view, the latter or the theory of the continuity of the germ-plasm, be correct, in either case the male germ-cells of the young animal must contain the same germ-plasm as that which existed in the fertilized maternal egg, that is to say, they must contain all the ancestral germ-plasms of the father and the mother. Here therefore a reduction must occur, for otherwise the number of ancestral germ-plasms would be increased by one half at every fertilization. The egg-cell would furnish 1/2, but the sperm-cell 2/2 of the total quantity of germ-plasm present in the germ-cells of the parents. But there is no reason for believing that the reduction of germ-plasm in the sperm-cell must proceed in precisely the same way as in the egg-cell, viz. by the expulsion of a polar body. On the contrary, the processes of spermatogenesis are so remarkably different from those of ovogenesis that we may expect to find that reduction is also brought about in a different manner.
The egg-cell does not expel the superfluous ancestral germ-plasms until the end of its development, and in a form which induces the destruction of the separated portion. This is certainly remarkable, for germ-plasm is a most important substance, and although it seems to be wasted in the production of enormous quantities of sperm- and egg-cells, such waste is only apparent, and is in reality the means which renders the species capable of existence. It may perhaps be possible to prove that in this case also the waste is only apparent. Such proof would be forthcoming if it could be shown that the means by which reduction is brought about in eggs is advantageous, and therefore also, ceteris paribus, necessary. We see that everywhere, as far as our observation extends, the useful is also the actual, unless indeed it is impossible of attainment or can only be attained by the aid of processes which are injurious to the species. And if it be asked why germ-plasm is wasted in the maturation of egg-cells, the following may perhaps be a satisfactory answer.
Let us suppose that the necessary reduction of the germ-plasm does not take place by the separation of the second polar body, but that it happens during the first division of the first primitive-germ-cell which is found in the embryo, so that the two first egg-cells resulting from this division would already contain only half the number of ancestral germ-plasms from the father and the mother, contained in the fertilized egg-cell. In this case the main object, the reduction of the ancestral germ-plasms, would be gained by a single division, and all the succeeding nuclear divisions, causing the multiplication of these two first germ-cells, might take place by the ordinary form of nuclear division, viz. ‘equal division.’ But perhaps nature not only cares for this one main object alone, but also secures certain secondary advantages at the same time. In the case which we have supposed the egg-cells of the mature ovary would only contain two different combinations of germ-plasm, which we may call combinations A and B. Even if millions of egg-cells were formed, every one of them would contain either A or B, and hence (at least as far as the female pronucleus is concerned) only two kinds of individuals could arise from such eggs—viz. offspring A’ and B’. All the offspring A’ would be as similar to one another as identical twins, and the same would be true of offspring B’.
But if the 100th instead of the 1st embryonic germ-cell entered upon the ‘reducing division,’ a hundred cells would undergo this division at the same time, and thus two hundred different combinations of ancestral germ-plasm would arise, and two hundred different kinds of germ-cells would be found in the mature ovary. A still greater number of different combinations of hereditary tendencies would arise if the ‘reducing division’ occurred still later; but undoubtedly the diversity in the composition of the germ-plasm must be greatest of all when the ‘reducing division’ does not take place during the period in which the germ-cells undergo multiplication, but at the end of the entire course of ovarian development, and separately in each full-grown mature egg ready for embryonic development. In such a case there will be as many different combinations of ancestral germ-plasms as there are eggs, for, as I have shown above, it is hardly conceivable that such a complex body as the nuclear substance of the egg-cell—composed of innumerable different units—would ever divide twice in precisely the same manner. Every egg will therefore contain a somewhat different combination of hereditary tendencies, and thus the offspring which arise from the different germ-cells of the same mother can never be identical. Hence by the late occurrence of the ‘reducing division’ the greatest possible variability in the offspring is secured.
If my interpretation of the second polar body be accepted, it is obvious that the late occurrence of the ‘reducing division’ is proved. At the same time we receive an explanation of the advantage gained by the postponement of the reduction of the germ-plasm until the end of the ovarian development of the egg; because the greatest possible number of individual variations in the offspring are produced in this way.
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 built265.
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 Julin266 stated in 1884 that direct and karyokinetic nuclear divisions alternate with each other in the spermatogenesis of Ascaris megalocephala. Again, Carnoy267 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.’ Platner268, 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. George269 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.
