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Weismann August
Essays Upon Heredity and Kindred Biological Problems

Hence the presence of polar bodies in Aphidae is a fresh confirmation of their great physiological importance. As bearing upon the main question dealt with in this essay, Blochmann’s observations have an especial interest, because only one polar body was found in the parthenogenetic eggs of Aphis, while the sexual eggs normally produce two. The author rightly states that this result is in striking accordance with my results obtained from the summer-eggs of different Daphnidae, and he adds the remark,—‘It would be of great interest to know whether these facts are due to the operation of some general law.’ To this remark I can now reply that there is indeed such a law: not only in the parthenogenetic eggs of Daphnidae, but also, as I have since found, in those of the Ostracoda and Rotifera²⁵¹, only one primary polar body is formed, while two are formed in all eggs destined for fertilization.

Before proceeding to the conclusions which follow from this fact, I will at once remove a difficulty which is apparently presented by the eggs which may develope with or without fertilization. I refer to the well-known case of the eggs of bees. It might be objected to my theory that the same egg cannot be prepared for development in more than one out of the two possible ways; it might be argued that the egg either possesses the power of entering upon two successive nuclear divisions during maturation, and in this case requires fertilization; or the egg may be of such a nature that it can only enter upon one such division and can therefore form only one polar body, and in that case it is capable of parthenogenetic development. Now there is no doubt, as I pointed out in my paper on the nature of parthenogenesis²⁵², that in the bee the very same egg may develope parthenogenetically, which under other circumstances would have been fertilized. Bessel’s²⁵³ experiments, in which young queens were rendered incapable of flight, and were thus prevented from fertilization, have shown that all the eggs laid by such females develope into drones (males) which are well known to result from parthenogenetic development. On the other hand, bee-keepers have long known that young queens which are fertilized in a normal manner continue for a long time to lay eggs which develope into females, that is to say, which have been fertilized. Hence the same eggs, viz. those which are lowest in the oviducts and are therefore laid first, develope parthenogenetically in the mutilated female, but are fertilized in the normal female. The question therefore arises as to the way in which the eggs become capable of adapting themselves to the expulsion of two polar bodies when they are to be fertilized, and of one only when fertilization does not take place.

But perhaps the solution of this problem is not so difficult as it appears to be. If we may assume that in eggs which are capable of two kinds of development the second polar body is not expelled until the entrance of a spermatozoon has taken place, the explanation of the possibility of parthenogenetic development when fertilization does not occur would be forthcoming. Now we know, from the investigations of O. Hertwig and Fol, that in the eggs of Echinus the two polar bodies are even formed in the ovary, and are therefore quite independent of fertilization, but in this and other similar cases a parthenogenetic development of the egg never takes place. There are, however, observations upon other animals which point to the fact that the first only and not the second polar body may be formed before the spermatozoon penetrates into the egg. It can be easily understood why it is that entirely conclusive observations are wanting, for hitherto there has been no reason for any accurate distinction between the first and the second polar body. But in many eggs it appears certain that the second polar body is not expelled until the spermatozoon has penetrated. O. Schultze, the latest observer of the egg of the frog, in fact saw the first polar body alone extruded from the unfertilized egg: a second nuclear spindle was indeed formed, but the second polar body was not expelled until after fertilization had taken place. A very obvious theory therefore suggests itself:—that while the formation of the second polar body is purely a phenomenon of maturation in most animal eggs, and is independent of fertilization,—in the eggs of a number of other animals, on the other hand, and especially among Arthropods, the formation of the second nuclear spindle is the result of a stimulus due to the entrance of a spermatozoon. If this suggestion be confirmed, we should be able to understand why parthenogenesis occurs in certain classes of animals wherever the external conditions of life render its appearance advantageous, and further, why in so many species of insects a sporadic parthenogenesis is observed, viz. the parthenogenetic development of single eggs (Lepidoptera). Slight individual differences in the facility with which the second nuclear spindle is formed independently of fertilization would in such cases decide whether an egg is or is not capable of parthenogenetic development. As soon, however, as the second nuclear spindle is formed, parthenogenesis becomes impossible. The nuclear spindle which gives rise to the second polar body, and that which initiates segmentation, are two entirely different things, and although they contain the same quantity, and the same kind of germ-plasm, a transformation of the one into the other is scarcely conceivable. This conclusion will be demonstrated in the following part of the essay.

II. The Significance of the Second Polar Body

I have already discussed the physiological importance of the first polar body, or rather of the first division undergone by the nucleus of the egg, and I have explained it as the removal of ovogenetic nuclear substance which has become superfluous and indeed injurious after the maturation of the egg. I do not indeed know of any other meaning which can be ascribed to this process, now that we know of the occurrence of a first division of the nucleus in parthenogenetic as well as in sexual eggs. A part of the nucleus must thus be removed from both kinds of eggs, a part which was necessary to complete their growth, and which then became superfluous and at the same time injurious. In this respect the observations of Blochmann²⁵⁴ upon the eggs of Musca vomitoria seem to me to be very interesting. Here the two successive divisions of the nuclear spindle arising from the egg-nucleus take place, but true polar bodies are not expelled, and the two nuclei corresponding to them (one of which divides once more) are placed on the surface of the egg, surrounded by an area free from yolk granules; and they break up at a later period. The essential point is obviously to eliminate from the egg-cell the influence of nucleoplasm which has been separated from the egg-nucleus as the first polar body; and this condition is satisfied whether the elimination is brought about by a process of true cell-division, as is the rule in the eggs of most animals, or by the division and removal of part of the egg-nucleus alone. The occurrence of the latter method of elimination certainly constitutes a still further proof of the physiological importance of the process, and this, taken together with the universal occurrence of polar bodies in all eggs—parthenogenetic and sexual—forces us to conclude that the process must possess a definite significance. No one of the various attempts which have been made to explain the significance of polar bodies generally is applicable to the first polar body except that which I have attempted.

But the case is different with the significance of the second nuclear division, or the second polar body. Here it might perhaps be possible to return to the view brought forward by Minot, Balfour, and van Beneden, and to consider the removal of this part of the nucleus as the expulsion of the male part of the previously hermaphrodite egg-cell. The second polar body is only expelled when the egg is to be fertilized, and at first sight it appears to be quite obvious that such a preparation of the egg for fertilization must depend upon its reduction to the female state. I believe however that this is not the case, and am of opinion that the process has an entirely different and much deeper meaning.

How can we gain any conception of this supposed hermaphroditism of the egg-cell, and its subsequent attainment of the female state? What are the essential characteristics of the male and female states? We know of female and male individuals, among both animals and plants: their differences consist essentially in the fact that they produce different kinds of reproductive cells; in part they are of a secondary nature, being adaptations of the organism to the functions of reproduction; they are intended to attract the other sex, or to ensure the meeting of the two kinds of reproductive cells, or to enable the fertilized egg to develope and sometimes to guide the development of the offspring until it has reached a certain period of growth. But all these differences, however great they may sometimes be, do not alter the essential nature of the organism. The blood corpuscles of man and woman are the same, and so are the cells of their nerves and muscles; and even the sexual cells, so different in size, appearance, and generally also in motile power, must contain the same fundamental substance, the same idioplasm. Otherwise the female germ-cell could not transmit the male characters of the ancestors of the female quite as readily as the female characters, nor could the male germ-cell transmit the female quite as readily as the male characters of the ancestors of the male. It is therefore clear that the nuclear substance itself is not sexually differentiated.

I have already previously pointed out that the above-mentioned facts of heredity contain the disproof of Minot’s theory, inasmuch as the egg-cell transmits male as well as female characters. Strasburger²⁵⁵ has also raised a similar objection. I consider this objection to be quite conclusive, for there does not seem to be any way in which the difficulty can be met by the supporters of the theory. The difficulty could indeed be evaded until we came to know that the essential part of the polar body is nuclear substance, and that the latter must be regarded as idioplasm,—as the substance which is the bearer of heredity. It might have been maintained that the male part, removed from the egg, consists only in a condition, perhaps comparable to positive or negative electricity; and that this condition is present in the substance of the polar body, so that the removal of the latter would merely signify a removal of the unknown condition. I do not mean to imply that any of those who have adopted Minot’s theory have had any such vague ideas concerning this process, but even if any one were ready to adopt it, he would be unable to make any use of the idea. He would not be able to support the theory in this way, for we now know that nuclear substance is removed with the polar body, and this fact requires an explanation which cannot be afforded by the theory, if we are right in believing that the expelled nuclear substance is not merely the indifferent bearer of the unknown principle of the male condition, but hereditary substance. I therefore believe that Minot’s, Balfour’s, and van Beneden’s hypothesis, although an ingenious attempt which was quite justified at the time when it originated, must be finally abandoned.

My opinion of the significance of the second polar body is shortly this,—a reduction of the germ-plasm is brought about by its formation, a reduction not only in quantity, but above all in the complexity of its constitution. By means of the second nuclear division the excessive accumulation of different kinds of hereditary tendencies or germ-plasms is prevented, which without it would be necessarily produced by fertilization. With the nucleus of the second polar body as many different kinds of idioplasm are removed from the egg as will be afterwards introduced by the sperm-nucleus; thus the second division of the egg-nucleus serves to keep constant the number of different kinds of idioplasm, of which the germ-plasm is composed during the course of generations.

In order to make this intelligible a short explanation is necessary.

From the splendid series of investigations on the process of fertilization, commenced by Auerbach and Bütschli, and continued by Hertwig, Fol, Strasburger, van Beneden, and many others, and from the theoretical considerations brought forward by Pflüger, Nägeli, and myself, at least one certain result follows, viz. that there is an hereditary substance, a material bearer of hereditary tendencies, and that this substance is contained in the nucleus of the germ-cell, and in that part of it which forms the nuclear thread, which at certain periods appears in the form of loops or rods. We may further maintain that fertilization consists in the fact that an equal number of loops from either parent are placed side by side, and that the segmentation nucleus is composed in this way. It is of no importance, as far as this question is concerned, whether the loops of the two parents coalesce sooner or later, or whether they remain separate. The only essential conclusion demanded by our hypothesis is that there should be complete or approximate equality between the quantities of hereditary substance derived from either parent. If then the germ-cells of the offspring contain the united germ-plasms of both parents, it follows that such cells can only contain half as much paternal germ-plasm as was contained in the germ-cells of the father, and half as much maternal germ-plasm as was contained in the germ-cells of the mother. This principle is affirmed in a well-known calculation made by breeders of animals, who only differ from us in their use of the term ‘blood’ instead of the term germ-plasm. Breeders say that half of the ‘blood’ of the offspring has been derived from the father and the other half from the mother. The grandchild similarly derives a quarter of its ‘blood’ from each of the four grandparents, and so on.

Let us imagine, for the sake of argument, that sexual reproduction had not been introduced into the animal kingdom, and that asexual reproduction had hitherto existed alone. In such a case, the germ-plasm of the first generation of a species which enters upon sexual reproduction must still be entirely homogeneous; the hereditary substance must, in each individual, consist of many minute units, each of which is exactly like the other, and each of which contains within itself the tendency to transmit, under certain circumstances, the whole of the characters of the parent to a new organism—the offspring. In each of the offspring of such a first generation, the germ-plasms of two parents will be united, and every germ-cell contained in the individuals of this second sexually produced generation will now contain two kinds of germ-plasm—one kind from the father, and the other from the mother. But if the total quantity of germ-plasm present in each cell is to be kept within the pre-determined limits, each of the two ancestral germ-plasms, as I may now call them, must be represented by only half as many units as were contained in the parent germ-cells.

In the third sexually produced generation, two new ancestral germ-plasms would be added by fertilization to the two already present, and the germ-cells of this generation would therefore contain four different ancestral germ-plasms, each of which would constitute a quarter of the total quantity. In each succeeding generation the number of the ancestral germ-plasms is doubled, while their quantities are reduced by one half. Thus in the fifth sexually produced generation, each of the sixteen ancestral germ-plasms will only constitute 1/16 of the total quantity; in the sixth, each of the thirty-two ancestral germ-plasms, only 1/32, and so on. The germ-plasm of the tenth generation would be composed of 1024 different ancestral germ-plasms, and that of the n^th of 2ⁿ. By the tenth generation each single ancestral germ-plasm would only form 1/1024 of the total quantity of germ-plasm contained in a single germ-cell. We know nothing whatever of the length of time over which this process of division of the ancestral germ-plasms may have endured, but even if it had continued to the utmost possible limit—so far indeed that each ancestral germ-plasm was only represented by a single unit—a time would at last come when any further division into halves would cease to be possible; for the very conception of a unit implies that it cannot be divided without the loss of its essential nature, which in this case constitutes it as the hereditary substance.

In the diagram represented in Fig. I. I have tried to render these conclusions intelligible. In generation I. each paternal and maternal germ-plasm is still entirely homogeneous, and does not contain any combination of different hereditary qualities, but the germ-plasm of the offspring is made up of equal parts of two kinds of germ-plasm. In the second generation this latter germ-plasm unites with another derived from other parents, which is similarly composed of two ancestral germ-plasms, and the resulting third generation now contains four different ancestral germ-plasms in its germ-cells, and so on. The diagram only indicates the fusion of ancestral germ-plasms as far as the offspring of the fourth generation, the germ-cells of which contain sixteen different ancestral germ-plasms. If we imagine the germ-plasm units to be so large that there is only room for sixteen of them in the nuclear thread, the limits of division would-be reached in the fifth generation, and any further division into halves of the ancestral germ-plasms would be impossible.

Now however minute the units may be, there is not the least doubt that the limits of possible division have been long since reached by all existing species, for we may safely assume that no one of them has acquired the sexual method of reproduction within a small number of recent generations. All existing species must therefore now contain as many different kinds of ancestral germ-plasms as they are capable of containing; and the question arises,—How can sexual reproduction now proceed without a doubling of the quantity of germ-plasm in each germ-cell, with every new generation?

There is only one possible answer to such a question:—sexual reproduction can proceed by a reduction in the number of ancestral germ-plasms, a reduction which is repeated in every generation.

Fig. I.

This must be so: the only question is, how and when does the supposed reduction take place.

Inasmuch as the germ-plasm is seated, according to our theory, in the nucleus, the necessary reduction can only be produced by nuclear division; and quite apart from any observation which has been already made, we may safely assert that there must be a form of nuclear division in which the ancestral germ-plasms contained in the nucleus are distributed to the daughter-nuclei in such a way that each of them receives only half the number contained in the original nucleus. After Roux’s²⁵⁶ elaborate review of the whole subject, we need no longer doubt that the complex method of nuclear division, hitherto known as karyokinesis, must be considered not merely as a means for the division of the total quantity of nuclear substance, but also for producing a division of the quantity and quality of each of its single elements. In by far the greater number of instances the object of this division is obviously to effect an equal distribution of nuclear substance in the two daughter-nuclei, so that each of the different qualities contained in the mother-nucleus is transferred to the two daughter-nuclei. This interpretation of ordinary karyokinesis is less uncertain than perhaps at first sight it may appear to be. We cannot, it is true, directly see the ancestral germ-plasms, nor do we even know the parts of the nucleus which are to be looked upon as constituting ancestral germ-plasm; but if Flemming’s original discovery of the longitudinal division of the loops lying in the equatorial plane of the nuclear spindle is to have any meaning at all, its object must be to divide and distribute the different kinds of the minutest elements of the nuclear thread as equally as possible. It has been ascertained that the two halves produced by the longitudinal splitting of each loop never pass into the same daughter-nucleus, but always in opposite directions. The essential point cannot therefore be the division of the nucleus into absolutely equal quantities, but it must be the distribution of the different qualities of the nuclear thread, without exception, in both daughter-nuclei. But these different qualities are what I have called the ancestral germ-plasms, i.e. the germ-plasms of the different ancestors, which must be contained in vast numbers, but in very minute quantities, in the nuclear thread. The supposition of a vast number is not only required by the phenomena of heredity but also results from the comparatively great length of the nuclear thread: furthermore it implies that each of them is present in very small quantity. The vast number together with the minute quantity of the ancestral germ-plasms permit us to conclude that they are, upon the whole, arranged in a linear manner in the thin thread-like loops: in fact the longitudinal splitting of these loops appears to me to be almost a proof of the existence of such an arrangement, for without this supposition the process would cease to have any meaning.

This is the only kind of karyokinesis which has been observed until recently; but if the supposed nuclear division leading to a reduction in the number of ancestral germ-plasms has any real existence, there must be yet another kind of karyokinesis, in which the primary equatorial loops are not split longitudinally, but are separated without division into two groups, each of which forms one of the two daughter-nuclei. In such a case the required reduction in the number of ancestral germ-plasms would take place, for each daughter-nucleus would receive only half the number which was contained in the mother-nucleus.

Now there is more evidence for the existence of this second kind of karyokinesis than the fact that it is demanded by my theory; for I believe that it has been already observed, although it has not been interpreted in this sense.

It is very probable that this is true of van Beneden’s²⁵⁷ observation on the egg of Ascaris megalocephala: he found that the nuclear division which led to the formation of the polar body differs from the ordinary course of karyokinesis, in that the plane of division is at right angles to that usually assumed. Carnoy²⁵⁸ has confirmed this observation in its main features, and he has made the further observation that out of the eight nuclear loops which are found at the equator of the spindle, four are removed with the first polar body, and that half of the remaining four are removed with the second polar body. The first of these two divisions would have to be looked upon as a reduction, if it is certain that each of the eight nuclear loops consists of different ancestral germ-plasms; but this assumption is impossible, although on the other hand it cannot be directly disproved: for we are not able to see the ancestral germ-plasms. But it must nevertheless be maintained that the removal of the first four loops does not imply a reduction in the number of ancestral germ-plasms in the nucleus; because, as I have already argued, two successive divisions of the number of ancestral germ-plasms into halves is inconceivable; and because the first polar body is also present in parthenogenetic eggs in which such division into halves cannot take place. But the karyokinetic process can readily be looked upon as a removal of ovogenetic nucleoplasm, for we know from the observations of Flemming and Carnoy, that, under certain circumstances, subsequent divisions may occur, involving an increase in the number of nuclear loops to double their number. These subsequent divisions of course take place in the daughter-nuclei. This fact proves, as I think, that there are nuclei in which the same ancestral germ-plasm occurs in two different loops: but such loops, identical as regards the composition of their ancestral germ-plasms, may very well contain different ontogenetic stages of this substance. This will be the case in the instance alluded to, if four loops of the first nuclear spindle are to be looked upon as ovogenetic nucleoplasm, and the four others as germ-plasm. It is therefore unnecessary to regard the first division of the egg-nucleus as a ‘reducing division’: it may be looked upon as an ‘equal division’²⁵⁹ entirely analogous to the kind of division which, in my opinion, directs the development of the embryo. This conclusion would receive direct proof if it were possible to show that the eight loops of the first division have arisen by the longitudinal splitting of four primary loops: for a longitudinal splitting of the nuclear thread would be the means by which the different ontogenetic stages of the germ-plasm could be separated from one another, without leading to any reduction in the number of ancestral germ-plasms in the daughter-nuclei. Thus I have previously attempted to prove that the ontogenetic development of the egg must be connected with a progressive transformation of the nucleoplasm during successive nuclear divisions, and this transformation will very frequently (but not always) occur in such a way that the different qualities of the nucleoplasm are separated from one another by the nuclear division. The nucleoplasm of the daughter-nuclei will be identical if the two daughter-cells are to potentially contain corresponding parts of the embryo; as for instance the first two segmentation spheres of the egg of the frog, which according to Roux²⁶⁰ correspond to the right and left halves of the future animal. But the nucleoplasm must be unequal if the products of division are to develope into different parts of the embryo. In both cases, however, karyokinesis is connected with a longitudinal splitting of the nuclear threads, and we may conclude from this fact (which is also confirmed by the phenomena of heredity) that all such nuclei, whether they have entered upon the same or different ontogenetic transformations of their nucleoplasm, are identical as regards the ancestral germ-plasm which they contain. During the whole process of segmentation and the entire development of the embryo, the total number of ancestral germ-plasms which were at first contained in the germ-plasm of the fertilized egg-cell must still be contained in each of the succeeding cells.

Thus no objection can be raised against the view that the four loops of the first polar body contain the ovogenetic nucleoplasm, that is to say, an idioplasm which contains the total number of ancestral germ-plasms, but at an advanced and highly specialized ontogenetic stage.

The formation of the second polar body may be rightly considered as a ‘reducing division,’ as a division leading to the expulsion of half the number of the different ancestral germ-plasms, in the form of two nuclear loops, for no reason can be alleged in support of the assumption that the four loops of the second nuclear spindle are made up of identical pairs. Furthermore the facts of heredity require the assumption that the greatest possible number of ancestral germ-plasms is accumulated in the germ-plasm of each germ-cell, and thus that the small number of loops not only means an increase in quantity but a multiplication in the number of different ancestral germ-plasms present in each of them. If this conclusion be correct, there can be no doubt that the second division of the egg-nucleus means a reduction in the above-mentioned sense.

But there are yet other observations which, if correct, must also be considered as ‘reducing divisions.’ I refer to all those cases in which the longitudinal splitting of the loops is either entirely wanting, or does not occur until after the loops have left the equator of the spindle and have moved towards the poles. In both instances the bearing upon the question would be the same, for only half the number of primary loops would reach each pole in either case. If therefore the primary loops are not made up of identical pairs, it follows that the two daughter-nuclei can only contain half the number of ancestral germ-plasms which were contained in the mother-nucleus. Whether the loops divide on their way to the poles or at the poles themselves, no difference will be brought about in the number of ancestral germ-plasms which they contain, for this number can neither increase nor diminish. The quantity of the different ancestral germ-plasms can alone be increased in this way. I am here referring to observations made by Carnoy²⁶¹ on the cells which form the spermatozoa in various Arthropods. It must be admitted, however, that these divisions cannot be regarded as ‘reducing divisions,’ if Flemming’s²⁶² suggestion be confirmed, that in all these observations the fact has been overlooked that the equatorial loops are not primary but secondary, and that they have arisen from the longitudinal splitting of the nuclear thread during previous stages of nuclear division. But this point can only be decided by renewed investigation. Although many excellent results have been obtained in the subject of karyokinesis, there is still very much to be learnt before our knowledge is complete; and this is not to be wondered at when we remember the great difficulties in the way of observation which are chiefly raised by the minute size of the objects to be investigated. Flemming’s most recent publications prove that we are still in the midst of investigation, and that highly interesting and important processes have hitherto escaped attention. A secure basis of facts is only very gradually obtained, and there are still many conflicting opinions upon the details of this process. I should therefore consider it to be entirely useless, from my point of view, to enter into a critical examination of everything known about all the details of karyokinesis. I am quite content to have shown how it may be imagined that the reduction required by my theory takes place during nuclear division; and at the same time to have pointed out that there are already observations which may be interpreted in this sense. But even if I am mistaken in this interpretation, the theoretical necessity for a reduction in the number of ancestral germ-plasms, a reduction repeated in every generation, seems to me to be so securely founded that the processes by which it is effected must take place, even if they are not supplied by the facts already ascertained. There must be two kinds of karyokinesis according to the different physiological effect of the process. First, a karyokinesis by means of which all the ancestral germ-plasms are equally distributed in each of the two daughter-nuclei after having been divided into halves: secondly, a karyokinesis by means of which each daughter-nucleus receives only half the number of ancestral germ-plasms possessed by the mother-nucleus. The former may be called ‘equal division,’ the latter ‘reducing division.’ Of course these two processes, which differ so greatly in their effects, must also be characterized by morphological differences, but we cannot assume that the latter are necessarily visible. Just as, during the division of the first and second nuclear spindle in the egg of Ascaris megalocephala, karyokinesis takes, upon the whole, the same morphological course, although we must ascribe different physiological meanings to the two processes of division,—so it may be in other cases. The ‘reducing division’ must be always accompanied by a reduction of the loops to half their original number, or by a transverse division of the loops (if such division ever occurs); although reduction can only occur when the loops are not made up of identical pairs. And it will not always be easy to decide whether this is the case. On the other hand, the form of karyokinesis in which a longitudinal splitting of the loops takes place before they separate to form the daughter-nuclei must always, as far as I can see, be considered as an ‘equal division.’ In the accompanying figures II and III, diagrams are given illustrating these two forms of karyokinesis, but I do not mean to imply that it is impossible to imagine any other form in which they may occur.