Part 13

8. First female form: the cap-like anterior part has become detached and the egg-cells (_eiz_) are escaping. 9. Second female form: _eiz_ = egg-cells; outside these are the muscular layer (_m_) and the ectoderm (_ekt_). 10 and 11. Two fragments of such a female broken to pieces by spontaneous division: the egg-cells are embedded in a granular mass, and undergo embryonic development in it at a later period; the whole is surrounded by ciliated cells. 12. Male discharging the spermatozoa by the breaking up of the ectoderm (_ekt_); _sp_ spermatozoa; _m_ muscle.]

There is nevertheless no doubt that in the first female form, when sexually mature, the greater part, not only of the endoderm but of the whole body, is made up of ova, so that the animal resembles a thin-walled sac full of eggs. The ova escape by the bursting of the thin ectoderm, and when they have all escaped, the thin disintegrated membrane, composed of ciliated cells, is no longer in a condition to live, and dies at once. This is the course of events as described by Götte, and he is probably correct in his interpretation. This is the real death of the Orthonectides, and if we regard them as low primitive forms (Mesozoa), here for the first time in the ascending series we meet with natural death. But the causes of this are scarcely so clear as Götte seems to think when he ascribes it to the effect of reproduction—an effect which is ‘not only empirically necessary, but absolutely unavoidable.’ Such a necessity is explained by the fact that the endoderm consists entirely of germ-cells. Now the life of the organism, being dependent upon the mutual action of both layers, must cease as soon as the whole endoderm is extruded during reproduction.

Arguments such as these pass over the presence of a mesoderm; but apart from this omission, it does not appear to me so self-evident from a purely physiological standpoint, that the ectodermal sheath with its muscle layer must die after the extrusion of the germ-cells.

In those females to which Götte refers in this passage, the whole sheath remains at first quite uninjured, with the exception of a small cap at the anterior end, which is pushed off to give exit to the ova; and inasmuch as the sheath continues to swim about in the nutritive fluids after this has taken place, the proof is at any rate wanting that it cannot support itself quite as well as before, although it has lost the germ-cells.

Then why does it die? My answer to this is simple:—because it has lived its time; because its length of life is limited to a period which corresponds with the time necessary for complete reproduction. The physical constitution of the body is so regulated that it remains capable of living until the extrusion of the reproductive cells, and then dies, however favourable external conditions may be for its further support.

The correctness of this explanation is shown by a consideration of the males and the second form of females; for in these cases the body falls to pieces, not as a consequence of reproduction, but as a preparation for it!

Götte only mentions the second female form in a note, in which he says, it appears ‘that in the second female form of these animals the whole body breaks into many pieces, and the superficial layer gradually atrophies, so that it dies before the eggs are extruded.’ In Julin’s account[76], upon which Götte bases his statements, there are, however, some not unimportant differences. For instance, the eggs are not extruded at all, but embryonic development takes place within the body of the mother, which has previously undergone spontaneous division into several pieces. In this case, the eggs differ from those of the other female form, inasmuch as they do not constitute the whole of the endoderm, but are embedded (as was stated above) in a fairly voluminous granular mass at the expense of which, or at least by means of which, they are nourished; for they increase considerably in size during their development. But not only this granular mass, but all the layers of the body of the mother, even the ectoderm, persist during the embryonic development of the offspring. Indeed, the ectoderm must continue to grow during the division of the mother animal, for it gradually covers in the products of division on all sides, and, by means of its cilia, causes the animal to swim about in the fluids of its host. After some time the cilia are lost, and the separate parts into which the mother animal has divided, fix themselves upon some part of the body-cavity of the host; the young become free, and the remains of the body of the mother probably disappear by dissolution and resorption[77]. In this case the remains of the mother animal seem to be, to some extent, consumed by the embryos,—a process which sometimes, although very rarely, happens elsewhere. We can scarcely consider this as a primitive arrangement, or look upon it as a proof that ‘reproduction’ has a necessarily fatal effect upon the Polyplastid organism.

In the male, the mass of spermatozoa does not swell out the body to such an extent that its walls must give way and thus permit an exit, but the large ectoderm cells atrophy spontaneously at the time of maturity, and as they fall off, exit is given to the spermatozoa here and there. In this instance also the dissolution of the body is not a consequence of reproduction, but reproduction can only take place when the dissolution of the body has preceded it!

In this remarkable arrangement we cannot discern anything except an evident adaptation of the life of the body-cells to reproductive purposes, and this adaptation was rendered possible because, after the evacuation of the sexual cells, the body ceased to be of any value for the maintenance of the species.

But even if we assume, that the death of the Orthonectides is, in Götte’s sense, a consequence of reproduction, inasmuch as, in the two forms of females as well as in the male, the extrusion of a mass of developed germ-cells or embryos deprives the organism of the physiological possibility of living longer, how can we explain the necessity of death as a direct consequence of reproduction in all Polyplastides? Is the body—the _soma_—of the Metazoa so imperfectly developed, as compared with the reproductive cells, that the extrusion of the latter involves the death of the former? As a matter of fact in the majority of cases the relations are reversed; the number of body-cells usually exceeds the germ-cells a hundred- or a thousand-fold, and the body is, as regards nutrition, so completely independent of the reproductive cells, that it need not be in the least disadvantageously affected by their extrusion. And if the Orthonectid-like ancestors of the Metazoa were compelled to give up their insignificant somatic part after the extrusion of their germ-cells, because it could now no longer support itself, does it therefore follow that the somatic cells had for ever lost the power of surviving, even when their phyletic descendants were surrounded by more favourable conditions? Had they to inherit ‘the necessity of death’ for all time? Whence came this great change in the nature of organisms which, before the differentiation of Homoplastids into Heteroplastids, were endowed with the immortality of unicellular beings?

And it must be remembered that it is only an assumption which places the Orthonectides among the lowest Metazoa (Heteroplastids). I do not intend to greatly emphasize this point, but the formation of the Gastrula by embole, and the absence of a mouth and alimentary canal, shows that these parasites are extremely degenerate, and the same may be said of almost all endoparasites. The Gastrula, as an independent organism, was without doubt primitively provided with both mouth and stomach, and the mass of ova filling the female Orthonectid is an adaptation to a parasitic life, which on the one side renders the possession of a stomach a superfluity, and on the other demands the production of a great number of germ-cells[78]. It is certain that the Orthonectides, as at present constituted, cannot have lived in the free condition, and also that their adaptation to parasitism cannot have arisen at the beginning of the phyletic development of Metazoa, because they inhabit star-fishes and Nemertines—both relatively highly developed Metazoa. Hence it is, at any rate, doubtful whether the Orthonectides can claim to pass as typical forms of the lowest Heteroplastids, and whether their reproduction can be looked upon ‘as typical for the unknown ancestors of all Polyplastids’ (l. c., p. 45). If, however, we accept some organism resembling these Orthonectides as the most ancient Heteroplastid, being a free-living organism, it must have had a stomach, and the cells surrounding it must—as a whole or in part—have possessed the power of digesting; at any rate, they cannot all have been germ-cells, and therefore it is improbable that death would be the direct result of the extrusion of the germ-cells.

Let us now consider the manner in which Götte has endeavoured to explain the transmission of the cause of death—which first appeared in the Orthonectides—from these organisms to all later Metazoa, until the very highest forms are reached. Exact proofs of this supposition are unfortunately wanting, and the evidence is confined to the collection of a number of cases in which death and reproduction take place nearly or quite simultaneously. These would prove nothing, even if _post hoc_ were always _propter hoc_; and there are, opposed to them, a number of cases in which reproduction and death take place at different times. In obtaining evidence for ‘the fatal influence of reproduction,’ is it possible to point to every case of sudden death after the act of oviposition or fertilization? These cases occur among many of the higher animals, especially in Insects, and were collected by me in an earlier work[79]. It is obvious that such cases are exceptional, but in a restricted sense it is quite true, as far as these individual instances are concerned, that death appears as a consequence of reproduction. The male bee, which invariably dies while pairing, is undoubtedly killed in consequence of a very powerful nervous shock; and the female Psychid, which has laid all her eggs at once, dies of ‘exhaustion’—however we may attempt to explain the term on physiological principles.

Can we conclude from these cases that the effects of reproduction are, in Götte’s sense, universally fatal; that reproduction is the positive and ‘exclusive explanation of natural death’? (l. c., p. 32.) I need not linger over these isolated examples, but I turn at once to the foundation of the whole conclusion—a foundation which is obviously unable to support the superstructure erected on it. Götte formally derives the idea that death is a necessary condition of reproduction, from a very heterogeneous collection of facts. When we examine this collection we find that the process which is taken to be death is not the same thing in all these instances, while the same is true of the influence of reproduction by which death is supposed to be caused. The whole conception arises out of the process of encystment, which is regarded as the building-up of reproductive material—that is, as true reproduction; and since, according to Götte’s view, the formation of germs is always intimately connected with an arrest of life, and since, by his own definition, this stand-still of life is equivalent to death, it follows that, with such a theory, reproduction, in its essential nature, must be inseparably connected with death. It is necessary at this juncture to remember what Götte means by the process of rejuvenescence, and to point out that he is dealing with something quite different from ‘the fatal influence of reproduction,’ which was just now mentioned with regard to insects. ‘Rejuvenescence,’ bound up as it is with encystment and reproduction, is, according to Götte, ‘a re-coining of the specific protoplasm, by means of which the identity of its substance is fixed by heredity,’ a ‘marvellous process in which phenomena the most important in the whole life of the animal, and in fact of all organisms—reproduction and death—have their roots’ (l. c., p. 81). Whether such re-coining really takes place or not, at any rate I claim to have shown above that it does not correspond with death in the Metazoa, and—if it is represented at all in these latter—that it ought to be looked for in the reproductive cells; and indeed, in another passage, Götte himself has placed the process in these cells.

While, among the Monoplastids, according to Götte, the causes of the supposed death lie hidden in this mysterious change of the organism into reproductive material, Götte asserts that among the Polyplastids (such as _Magosphaera_, hypothetically perfected so as to form a genuine Polyplastid), the causes of death operate so that the organism breaks up into its component cells, all these being still reproductive cells—a process which, unlike ‘rejuvenescence,’ has nothing mysterious about it, and which is certainly not genuine death. In the Orthonectid-like animals death does not occur as a consequence of the dispersal of the reproductive cells, but rather because the part of the animal which remains is so small and effete that, being unable to support itself, it necessarily dies. From this point at least the object of death and the conception of it remain the same, but now the idea of reproduction undergoes a change. When the Rhabdite females of _Ascaris_ are eaten up by their offspring, is this mode of death connected with the ‘rejuvenescence of protoplasm’? (l. c., p. 34.) Is there any deep underlying relationship between such an end and the essential nature of reproduction? The same question may be asked with regard to the ‘Redia or the Sporocyst of Trematodes, which are converted into slowly dying sacs during the growth of the Cercariae within them.’ We cannot speak of the ‘fatal influence of reproduction’ among tape-worms just because ‘in the ripe segments the whole organization degenerates under the influence of the excessive growth of the uterus.’ It certainly degenerates, but only so far as the developing mass of eggs demands. In fact, at a sufficiently high temperature, death does not occur, and such mature segments of tape-worms creep about of their own accord. We cannot fail to recognize that in this as well as in the above-mentioned cases we have to do with adaptation to certain very special conditions of existence—an adaptation leading to an immense development of reproductive cells in a mother organism which can no longer take in nourishment, or which has become entirely superfluous because its duty to its species is already fulfilled. If this is an example of death inherent in the essential nature of reproduction, then so is the death of a mature segment of a tapeworm in the gastric juices of the pig that eats it.

With Götte, the conception of reproduction, like the conception of death, is a protean form, which he welcomes in any shape, if only he can use it as evidence. If death is a necessary consequence of reproduction, its cause must be always essentially the same, and might be expressed in one of the following suggestions:—(1) in the necessity for a ‘re-coining’ of the protoplasm of the germ-cells; but here death could only affect the germ-cells themselves: (2) perhaps in the withdrawal of nourishment by the mass of developing reproductive material, just as death occurs sometimes among men by the excessive drain on the system caused by morbid tumours: (3) or in consequence of the development of the offspring in the body of the mother; this however would only affect the females, and could therefore have no deep and general significance: (4) from the extrusion of the sexual cells,—ova or spermatozoa,—and in the impossibility of further nourishment which is consequent upon this extrusion—(Orthonectides?): or (5) finally in an excessively powerful nervous shock brought about by the ejection of the reproductive cells.

But no one of these alternatives is the universal and inevitable cause of death. This proves irrefutably that death does not proceed as an intrinsic necessity from reproduction, although it may be connected with the latter, sometimes in one way and sometimes in another. But we must not overlook the fact that in many cases death is not connected with reproduction at all; for many Metazoa survive for a longer or shorter period after the reproductive processes have ceased.

In fact, I believe I have definitely shown that no process exists among unicellular animals which is at all comparable with the natural death of the higher organisms. Natural death first appeared among multicellular beings, and among these first in the Heteroplastids. Furthermore, it was not introduced from any absolute intrinsic necessity inherent in the nature of living matter, but on grounds of utility, that is from necessities which sprang up, not from the general conditions of life, but from those special conditions which dominate the life of multicellular organisms. If this were not so, unicellular beings must also have been endowed with natural death. I have already expressed these ideas elsewhere[80], and have briefly indicated how far, in my opinion, natural death is expedient for multicellular organisms. I found the essential reason for confining the life of the Metazoa to a fixed and limited period, in the wear and tear to which an individual is exposed in the course of a life-time. For this reason, ‘the longer the individual lived, the more defective and crippled it would become, and the less perfectly would it fulfil the purpose of its species’ (l. c., p. 24). Death seemed to me to be expedient since ‘worn-out individuals are not only valueless to the species, but they are even harmful, for they take the place of those which are sound’ (l. c., p. 24).

I still adhere entirely to this explanation; not of course in the sense that an actual physical struggle has ever taken place between the mortal and immortal varieties of any species. If Götte understood me thus, he may be justified by the brief explanations given in the essay to which I have alluded; but when he also attributes to me the opinion that such hypothetically immortal Metazoa had but a very limited period for reproduction, I fail to see what part of the essay in question can be brought forward in support of his statement. Only under some such supposition can I be reproached with having assumed the existence of a process of natural selection which could never be effective, because any advantage which accrued to the species from the shortening of the duration of life could not make itself felt in a more rapid propagation of the short-lived individuals. The statement ‘that in this and in every other case it is a sufficient explanation of the processes of natural selection to render it probable that any kind of advantage is gained’[81] is indeed erroneous. The explanation ought rather to be ‘that the forms in question would for ever transmit their characters to a greater number of descendants than the other forms.’ I have not however as yet attempted to think out in detail such processes of natural selection as would limit the somatic part of the Metazoan body to a short term of existence, and I only wished to emphasize the general principle lying at the basis of the whole process, without stating the precise manner in which it operates.

If I now attempt to take this course, and to reconstruct theoretically the gradual appearance of natural death in the Metazoa, I must begin by again alluding to Götte’s criticisms in reference to the operation of natural selection.

I consider death as an adaptation, and believe that it has arisen by the operation of natural selection. Götte[82], however, concludes from this that ‘the first origin of hereditary and consequently (for the organization in question) necessary death, is not explained but already assumed.’ ‘The operation and significance of the principle of utility consists in selecting the fittest from among the structures and processes which are at hand, and not in directly creating new ones. Every new structure arises at first, quite independently of any utility, from certain material causes present in a number of individuals, and when it has proved useful and is transmitted, it extends, according to the laws of the survival of the fittest, in the group of animals in which it appeared. This extension will undergo further increase with every advance in utility which results from further structural changes, until it extends over the whole group. So that usefulness effects the preservation and the distribution of new structures, but has nothing whatever to do with the causes of their primary origin and their consequent transmission to all other individuals. Indeed, on these hereditary causes the necessity of the structures in question depends, so that their usefulness in no way explains their necessity.’

‘These conclusions, when applied to the origin of natural death called forth by internal causes, would show that it became inevitable and hereditary in a number of the originally immortal Metazoa, before there could be any question as to the benefits derived from its influence. Such influence must have consisted in the fact that more descendants survived the struggle for existence and were able to enter upon reproduction among the individuals which had inherited the predisposition to die than among the potentially immortal beings which would be damaged in the struggle for existence, and would therefore be exposed to still further injuries. The existing necessity for natural death in all Metazoa might therefore be derived in an unbroken line of descent from the first mortal Metazoan, of which the death became inevitable from internal causes, before the principle of utility could operate in favour of its dissemination.’

In reply to this I would urge: that it has been very often maintained that natural selection can produce nothing new, but can only bring to the front something which existed previously to the exercise of choice; but this argument is only true in a very limited sense. The complex world of plants and animals which we see around us contains much that we should call new in comparison with the primitive beings from which, as we believe, everything has developed by means of natural selection. No leaves or flowers, no digestive system, no lungs, legs, wings, bones or muscles were present in the primitive forms, and all these must have arisen from them according to the principle of natural selection. These primitive forms were in a certain sense predestined to develope them, but only as possibilities, and not of necessity; nor were they preformed in them. The course of development, as it actually took place, first became a necessity by the action of natural selection, that is by the choice of various possibilities, according to their usefulness in fitting the organism for its external conditions of life. If we once accept the principle of natural selection, then we must admit that it really can create new structures, instincts, etc., not suddenly or discontinuously, but working by the smallest stages upon the variations that appear. These changes or variations must be looked upon as very insignificant, and are, as I have of late attempted to show[83], quantitative in their nature; and it is only by their accumulation that changes arise which are sufficiently striking to attract our attention, so that we call them ‘new’ organs, instincts, etc.