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Weismann August
Essays Upon Heredity and Kindred Biological Problems

If we trace all the permanent hereditary variations from generation to generation back to the quantitative variations of the germ, as I have sought to do, the question naturally occurs as to the source from which these variations arose in the germ itself. I will not enter into this subject at any length on the present occasion, for I have already expressed my opinion upon it⁵⁸.

I believe however that they can be referred to the various external influences to which the germ is exposed before the commencement of embryonic development. Hence we may fairly attribute to the adult organism influences which determine the phyletic development of its descendants. For the germ-cells are contained in the organism, and the external influences which affect them are intimately connected with the state of the organism in which they lie hid. If it be well nourished, the germ-cells will have abundant nutriment; and, conversely, if it be weak and sickly, the germ-cells will be arrested in their growth. It is even possible that the effects of these influences may be more specialized; that is to say, they may act only upon certain parts of the germ-cells. But this is indeed very different from believing that the changes of the organism which result from external stimuli can be transmitted to the germ-cells and will re-develope in the next generation at the same time as that at which they arose in the parent, and in the same part of the organism.

We have an obvious means by which the inheritance of all transmitted peculiarities takes place, in the continuity of the substance of the germ-cells, or germ-plasm. If, as I believe, the substance of the germ-cells, the germ-plasm, has remained in perpetual continuity from the first origin of life, and if the germ-plasm and the substance of the body, the somatoplasm, have always occupied different spheres, and if changes in the latter only arise when they have been preceded by corresponding changes in the former, then we can, up to a certain point, understand the principle of heredity; or, at any rate, we can conceive that the human mind may at some time be capable of understanding it. We may at least maintain that it has been rendered intelligible, for we can thus trace heredity back to growth; we can thus look upon reproduction as an overgrowth of the individual, and can thus distinguish between a succession of species and a succession of individuals, because in the latter succession the germ-plasm remains similar, while in the succession of the former it becomes different. Thus individuals, as they arise, are always assuming new and more complex forms, until the interval between the simple unicellular protozoon and the most complex of all organisms—man himself—is bridged over.

I have not been able to throw light upon all sides of the question which we are here discussing. There are still some essential points which I must leave for the present; and, furthermore, I am not yet in a position to explain satisfactorily all the details which arise at every step of the argument. But it appeared to me to be necessary to state this weighty and fundamental question, and to formulate it concisely and definitely; for only in this way will it be possible to arrive at a true and lasting solution of the problem. We must however be clear on this point—that the understanding of the phenomena of heredity is only possible on the fundamental supposition of the continuity of the germ-plasm. The value of experiment in relation to this question is somewhat doubtful. A careful collection and arrangement of facts is far more likely to decide whether, and to what extent, the continuity of germ-plasm is reconcilable with the assumption of the transmission of acquired characters from the parent body to the germ, and from the germ to the body of the offspring. At present such transmission is neither proved as a fact, nor has its assumption been shown to be unquestionably necessary.

III.
LIFE AND DEATH.
1883

LIFE AND DEATH.
PREFACE

The following paper was first printed as an academic lecture in the summer of the present year (1883), with the title ‘Upon the Eternal Duration of Life’ (‘Über die Ewigkeit des Lebens’). In now bringing it before a larger public in an expanded and improved form, I have chosen a title which seemed to me to correspond better with the present contents of the paper.

The stimulus which led to this biological investigation was given in a memoir by Götte, in which this author opposes views which I had previously expressed. Although such an origin has naturally caused my paper to take the form of a reply, my intention was not merely to controvert the views of my opponent, but rather—using those opposed views as a starting-point—to throw new light upon certain questions which demand consideration; to give additional support to thoughts which I have previously expressed, and to penetrate, if possible, more deeply into the problem of life and death.

If, in making this attempt, the views of my opponent have been severely criticized, it will be acknowledged that the criticisms do not form the purpose of my paper, but only the means by which the way to a more correct understanding of the problems before us may be indicated.

A. W.

Freiburg i. Breisgau,

Oct. 18, 1883.

III.
LIFE AND DEATH

In the previous essay, entitled ‘The Duration of Life,’ I have endeavoured to show that the limitation of life in single individuals by death is not, as has been hitherto assumed, an inevitable phenomenon, essential to the very nature of life itself; but that it is an adaptation which first appeared when, in consequence of a certain complexity of structure, an unending life became disadvantageous to the species. I pointed out that we could not speak of natural death among unicellular animals, for their growth has no termination which is comparable with death. The origin of new individuals is not connected with the death of the old; but increase by division takes place in such a way that the two parts into which an organism separates are exactly equivalent one to another, and neither of them is older or younger than the other. In this way countless numbers of individuals arise, each of which is as old as the species itself, while each possesses the capability of living on indefinitely, by means of division.

I suggested that the Metazoa have lost this power of unending life by being constructed of numerous cells, and by the consequent division of labour which became established between the various cells of the body. Here also reproduction takes place by means of cell-division, but every cell does not possess the power of reproducing the whole organism. The cells of the organism are differentiated into two essentially different groups, the reproductive cells—ova or spermatozoa, and the somatic cells, or cells of the body, in the narrower sense. The immortality of the unicellular organism has only passed over to the former; the others must die, and since the body of the individual is chiefly composed of them, it must die also.

I have endeavoured to explain this fact as an adaptation to the general conditions of life. In my opinion life became limited in its duration, not because it was contrary to its very nature to be unlimited, but because an unlimited persistence of the individual would be a luxury without a purpose. Among unicellular organisms natural death was impossible, because the reproductive cell and the individual were one and the same: among multicellular animals it was possible, and we see that it has arisen.

Natural death appeared to me to be explicable on the principle of utility, as an adaptation.

These opinions, to which I shall return in greater detail in a later part of this paper, have been opposed by Götte⁵⁹, who does not attribute death to utility, but considers it to be a necessity inherent in life itself. He considers that it occurs not only in the Metazoa or multicellular animals, but also in unicellular forms of life, where it is represented by the process of encystment, which is to be regarded as the death of the individual. This encystment is a process of rejuvenescence, which, after a longer or shorter interval, interrupts multiplication by means of fission. According to Götte, this process of rejuvenescence consists in the dissolution of the specific structure of the individual, or in the retrogression of the individual to a form of organic matter which is no longer living but which is comparable to the yolk of an egg. This matter is, by means of its internal energy, and in consequence of the law of growth which is inherent in its constitution, enabled to give rise to a new individual of the same species. Furthermore, the process of rejuvenescence among unicellular beings corresponds to the formation of germs in the higher organisms. The phenomena of death were transmitted by heredity from the unicellular forms to the Metazoa when they arose. Death does not therefore appear for the first time in the Metazoa, but it is an extremely ancient process which ‘goes back to the first origin of organic beings’ (l. c., p. 81).

It is obvious, from this short résumé, that Götte’s view is totally opposed to mine. Inasmuch as only one of these views can be fundamentally right, it is worth while to compare the two; and although we cannot at present hope to explain the ultimate physiological processes which involve life and death, I think nevertheless that it is quite possible to arrive at definite conclusions as to the general causes of these phenomena. At any rate, existing facts have not been so completely thought out that it is useless to consider them once more.

The question—what do we understand by death? must be decided before we can speak of the origin of death. Götte says, ‘we are not able to explain this general expression quite definitely and in all its details, because the moment of death, or perhaps more exactly the moment when death is complete, can in no case be precisely indicated. We can only say that in the death of the higher animals, all those phenomena which make up the life of the individual cease, and further that all the cells and elements of tissue which form the dead organism, die, and are resolved into their elements.’

This definition would suffice if it did not include that which is to be defined. For it assumes that under the expression ‘dead organism’ we must include those organisms which have brought to an end the whole of their vital functions, but of which the component cells and elements may still be living. This view is afterwards more accurately explained, and in fact there is no doubt that the cessation of the activity of life in the multicellular organism rarely implies any direct connection with the cessation of vital functions in all its constituents. The question however arises, whether it is right or useful to limit the conception of death to the cessation of the functions of the organism. Our conceptions of death have been derived from the higher organisms alone, and hence it is quite possible that the conception may be too limited. The limitation might perhaps be removed by accurate and scientific comparison with the somewhat corresponding phenomena among unicellular organisms, and we might then arrive at a more comprehensive definition. Science has without doubt the right to make use of popular terms and conceptions, and by a more profound insight to widen or restrict them. But the main idea must always be retained, so that nothing quite new or strange may appear in the widened conception. The conception of death, as it has been expressed with perfect uniformity in all languages, has arisen from observations on the higher animals alone; and it signifies not only the cessation of the vital functions of the whole organism, but at the same time the cessation of life in its single parts, as is shown by the impossibility of revival. The post-mortem death of the cells is also part of death, and was so, long before science established the fact that an organism is built up of numerous very minute living elements, of which the vital processes partially continue for some time after the cessation of those of the whole organism. It is precisely this incapacity on the part of the organism to reproduce the phenomena of life anew, which distinguishes genuine death from the arrest of life or trance; and the incapacity depends upon the fact that the death of the cells and tissues follows upon the cessation of the vital functions as a whole. I would, for this reason, define death as an arrest of life, from which no lengthened revival, either of the whole or any of its parts, can take place; or, to put it concisely, as a definite arrest of life. I believe that in this definition I have expressed the exact meaning of the conception which language has sought to convey in the word death. For our present purpose, the cause which gives rise to this phenomenon is of no importance,—whether it is simultaneous or successive in the various parts of the organism, whether it makes its appearance slowly or rapidly. For the conception itself it is also quite immaterial whether we are able to decide if death has really taken place in any particular case; however uncertain we might be, the state which we call death would be not less sharply and definitely limited. We might consider the caterpillar of Euprepia flavia to be dead when frozen in ice, but if it recovered after thawing and became an imago, we should say that it had only been apparently dead, that life stood still for a time, but had not ceased for ever. It is only the irretrievable loss of life in an organism which we call death, and we ought to hold fast to this conception, so that it will not slip from us, and become worthless, because we no longer know what we mean by it.

We cannot escape this danger if we look upon the post-mortem death of the cells of the body as a phenomenon which may accompany death, but which may sometimes be wanting. An experiment might be made in which some part of a dead animal, such as the comb of a cock, might be transplanted, before the death of the cells, to some other living animal: such a part might live in its new position, thus showing that single members may survive after the appearance of death, as I understand it. But the objection might be raised that in such a case the cock’s comb has become a member of another organism, so that it would be lost labour to insert a clause in our definition of death which would include this phenomenon. The same objection might be raised if the transplantation took place a day or even a year before the death of the cock.

Götte is decidedly in error when he considers that the idea of death merely expresses an ‘arrest of the sum of vital actions in the individual,’ without at the same time including that definite arrest which involves the impossibility of any revival. Decomposition is not quite essential to our definition, inasmuch as death may be followed by drying-up⁶⁰, or by perpetual entombment in Siberian ice (as in the well-known case of the mammoth), or by digestion in the stomach of a beast of prey. But the notion of a dead body is indeed inseparably connected with that of death, and I believe that I was right in distinguishing between the division of an Infusorian into two daughter-cells, and the death of a Metazoon, which leaves offspring behind it, by calling attention to the absence of a dead body in the process of fission among Infusoria (See below.). The real proof of death is that the organized substance which previously gave rise to the phenomena of life, for ever ceases to originate such phenomena. This, and this alone, is what mankind has hitherto understood by death, and we must start from this definition if we wish to retain a firm basis for our considerations.

We must now consider whether this definition, derived from observation of higher animals, may be also applied without alteration to the lower, or whether the corresponding phenomena which arise in these latter, differ in detail from those of the higher animals, so that a narrower limitation of the above definition is rendered necessary.

Götte believes the process of encystment which takes place in so many unicellular animals (Monoplastides) to be the analogue of death. According to this authority, the individuals in question, not only undergo a kind of winter sleep—a period of latent life—but when surrounded by the cyst they lose their former specific organization; they become a ‘homogeneous substance,’ and are resolved into a germ, from which, by a process of development, a new individual of the same species once more arises. The division of the contents of the cyst, viz. its multiplication, is, according to this view, of secondary importance, and the essential feature in the process is the rejuvenescence of the individual. This rejuvenescence however is said to not only consist in the simple transformation of the old individual, but in its death, followed by the building up anew of another individual. ‘The parent organism and its offspring are two successive living stages of the same substance—separated, and at the same time connected, by the condition of rejuvenescence which lies between them’ (l. c., p. 79). An ‘absolute continuity of life does not exist’; it is only the dead organic matter which establishes the connection, and the ‘identity of this matter ensures heredity.’

It is certainly surprising that Götte should identify encystment with a cessation of life, and we may well inquire for the evidence which is believed to support such a view. The only evidence lies in a certain degree of degeneration in the structure of the individual, and in the cessation of the visible external phenomena of life, such as feeding and moving. Does Götte really believe that it is an incorrect interpretation of the facts to assume that a vita minima continues to exist in the protoplasm, after its complexity has diminished? Are we compelled to invoke a mystical explanation of the facts, by an appeal to such an indefinite principle as Götte’s rejuvenescence? Would not the oxygen, dissolved in the water, affect the organic substance the life of which it formerly maintained, and would it not cause its decomposition, if it were in reality dead?

I, too, hold that the division of the encysted mass is of secondary importance, and that the encystment itself, without the resulting multiplication, is the original and essential part of the phenomenon. But it does not follow from this that the encystment should be considered as a process of rejuvenescence. What is there to be rejuvenated? Certainly not the substance of the animal, for nothing is added to it, and it can therefore acquire no new energy; and the forms of energy which it manifests cannot be changed, since the form of the matter is just the same after quitting the cyst as it was before. Rejuvenescence has also been mentioned in connection with the process of conjugation, but this is quite another thing. It is quite reasonable, at least in a certain sense, to maintain the connection of rejuvenescence with conjugation; for a fusion of the substance of two individuals takes place, to a greater or lesser extent, in conjugation, and the matter which composes each individual is therefore really altered. But in simple encystment, rejuvenescence can only be understood in the sense in which we speak of the fable of the Phœnix, which, when old, was believed to be consumed by fire, and to rise again from its own ashes as a young bird. I doubt whether this idea is in agreement with the physiology of to-day, or with the laws of the conservation of energy. It is easy to pull down an old house with rotten beams and crumbling walls, but it would be impossible to build it anew with the old material, even if we used new mortar, represented in Götte’s hypothesis by water and oxygen. For these reasons I consider the idea of rejuvenescence of the encysted individual to be contrary to our present physiological knowledge.

It is much more simple and natural to regard encystment as adapted for the protection of certain individuals in a colony from destruction by being dried up or frozen, or for the protection of the individual during multiplication by division, when it is helpless, and would easily fall a prey to enemies, or to secure advantages in some other way⁶¹. The case of Actinosphaerium, mentioned by Götte, clearly demonstrates that rejuvenescence of the individual is not the only event which happens during encystment, for this would scarcely require six months. The long duration of latent life, from summer to the next spring, clearly proves that encystment is of the highest importance for the species, in order to maintain the life of the individual through the dangers of an unfavourable season⁶².

When in this case, the specific organization degenerates to a certain extent, such changes depend in part upon the endeavour to diminish as far as possible the size of the organism—the pseudopodia being drawn in, while the vacuoles contract and completely disappear. The degeneration may also, perhaps, depend in part upon the secretion of the cyst itself, which implies a certain loss of substance⁶³. But degeneration chiefly depends upon the fact that the encystment is accompanied by reproduction in the way of fission, which seems to begin with a simplification of the organization, that is, with a fusion of the numerous nuclei. It is well known that many unicellular animals contain several nuclei—in other words, that the nuclear substance is scattered in small parts throughout the whole cell. But when the animal prepares for division, these pieces of nuclear substance fuse into a single nucleus which itself undergoes division into two equal parts⁶⁴ during the division of the animal. It is evident that the equal division of the whole nuclear substance only becomes possible in this way.

There are, however, numerous cases which prove that the bodies of encysted animals may retain, during the whole process, exactly the same structure and differentiation, which were previously characteristic of them. Thus the large Infusorian Tillina magna, described by Gruber, can be seen through the thin-walled cyst to retain the characteristic structure of its ectoplasm, and the whole of its organization. Even the movements of the enclosed animal do not cease; it continues to rotate actively in the narrow cyst, as do the two or four parts into which it subsequently divides. Such observations prove that Götte’s view that ‘every characteristic of the previous organization is lost,’ is quite out of the question⁶⁵ (l. c., p. 62).

For this reason I must strongly oppose Götte’s view that an encysted individual is a germ, viz. an organic mass still unorganized which can only become an adult individual by means of a process of development. I believe that an encysted individual is one possessing a protective membrane, in structure more or less simplified as an adaptation to the narrow space within the cyst, and to a possible subsequent increase by division, in short one in which active life is reduced to a minimum, and sometimes even completely in abeyance, as happens when it is frozen.

It is evident from the above considerations that encystment in no way corresponds with that which every one, including myself, understands by death, because during encystment one and the same being is first apparently dead and then again alive; and we merely witness a condition of rest, from which active life will again emerge. This would remain true even if it were proved that life is, in reality, suspended for a time. But such proof is still wanting, and Götte was apparently only influenced by theoretical considerations, when he imagined that death intervened where unprejudiced observers have only recognised a condition of rest. He apparently entirely overlooked the fact that it is possible to test his views; for all unicellular beings are in reality capable of dying: we can kill them, for example, by boiling, and they are then really dead and cannot be revived. But this state of the organism differs chemically and physically from the encysted condition, although we do not know all the details of the difference. The encysted animal, when placed in fresh water, presently originates a living individual, but the one killed by boiling only results in decomposition of the dead organic matter. Hence we see that the same external conditions give rise to different results in two different states of the organism. It cannot be right to apply the same term to two totally different states. There is only one phenomenon which can be called death, although it may be produced by widely different causes. But if the encysted condition is not identical with the death which we can produce at will, then natural death, viz. that arising from internal causes, does not exist at all among unicellular organisms.

These facts refute Götte’s peculiar view, which depends on the existence of natural death among the Monoplastid organisms; upon proof of the contradictory, his whole theory collapses. But there is nevertheless a certain interest in following it further, for we shall thus reach many ideas worthy of consideration.

First, the question arises as to how death could have been transmitted from the Monoplastides⁶⁶ to the Polyplastides, a process which must have taken place according to Götte. I will for the present omit the fact that I cannot accept the supposition that the process of encystment represents death. We may then inquire whether death has taken the place of encystment among the Polyplastides, or, if this is not the case, whether any process comparable to encystment exists among the Polyplastides.

Götte believes that death is always connected with reproduction, and is a consequence of the latter in both Protozoa and Metazoa. Reproduction has, in his opinion, a directly ‘fatal effect,’ and the reproducing individual must die. Thus the may-fly and the butterfly die directly after laying their eggs, and the male bee dies immediately after pairing; the Orthonectides expire after expelling their germ-cells, while Magosphaera resolves itself into germ-cells, and nothing persists except these elements. It is but a step from this latter organism to the unicellular animals which transform themselves as a whole into germ-cells; but in order to achieve this they must undergo the process of rejuvenescence, which Götte assumes to be the same as death.

These views contain many fallacies quite apart from the soundness or unsoundness of their foundation. The process of encystment, as Götte thinks, represents, in the Monoplastides, true reproduction to which multiplication by means of division has been secondarily added. This encystment cannot be dispensed with, for internal causes determine that it must occasionally interrupt the process of multiplication by simple division. But, on the other hand, Götte also considers the division of the contents of the cyst to be a secondary process. The essential characteristic of encystment is a simple process of rejuvenescence without multiplication. Hence we are forced to accept a primitive condition in which simple division as well as the division of the encysted individual were absent, and in which reproduction consisted only in an often-repeated process of rejuvenescence among existing individuals, without any increase in their number. Such a condition is inconceivable because it would involve a rapid disappearance of the species, and the whole consideration clearly shows us that division of un-encysted individuals must have existed from the first, and that this, and not a vague and mysterious rejuvenescence, has always been the real and primitive reproduction of the Monoplastides. The fact that encystment does not always lead to the division of the contents of the cyst proves, in my opinion, that not reproduction but preservation against injury from without, was the primitive meaning of encystment. It is possible that at the present time there are but few Monoplastides which are able to go through an infinite number of divisions without the interposition of the resting condition implied by encystment; although it has not yet been demonstrated for all species⁶⁷. But it is not right to conclude from this that there is an internal necessity which leads to encystment, that is to say to identify this process with rejuvenescence. It is much more probable that encystment is merely an adaptation to continual changes in the external conditions of life, such as drought and frost, and perhaps also the want of food which arises from the over-population of small areas. The same phenomenon is known in certain low Crustacea—the Daphnidae—which possess an ephippium or protective case for their winter-eggs. This case is only developed after a certain definite number of generations has been run through, an event which may happen at any time in the year in species living in pools which are liable to be often dried-up; but only in the autumn in such as live in lakes which are never dry. No one ever doubted that the periodical formation of the ephippium in certain generations was an adaptation to changes in the external conditions of life.

Even if the process of rejuvenescence in the Monoplastides were really equivalent to the death of the higher animals, we could not conclude from this that it is necessarily associated with reproduction. Encystment alone is not reproduction, and it first becomes a form of reproduction when it is associated with the division of the encysted animal. Simple division was the true and original form of reproduction in Monoplastides, and even now it is the principal and fundamental form.

Hence we see that among the Monoplastides reproduction is not connected with death, even if we accept Götte’s view and allow that encystment represents death. I shall return later on to the relation between death and reproduction in the Metazoa; but the question first arises whether encystment, if it is not death, has any analogue in the higher animals, and further whether death takes that place in their development which is occupied by encystment in the Monoplastides.