Essays Upon Heredity and Kindred Biological Problems. Weismann August

Essays Upon Heredity and Kindred Biological Problems - Weismann August


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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.

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      1

      Humboldt’s ‘Ausichten der Natur.’

      2

      This estimate is derived from observation of the time during which these insects are to be seen upon the wing. Direct observations upon the duration of life in this species are unknown to me.

      3

      [Sir John Lubbock has now kept a queen ant alive for nearly 15 years. See note 2 {note 18 below} on p. 51.—E. B. P.]

      4

      [After reading these proofs Dr. A. R. Wallace kindly sent me an unpublished note upon the production of death by means of natural selection, written by him some time between 1865 and 1870. The note contains some ideas on the subject, which were jotted down for further elaboration, and were then forgotten until recalled by the argument of this Essay. The note is of great interest in relation to Dr. Weismann’s suggestions, and with Dr. Wallace’s permission I print it in full below.

      ‘The Action of Natural Selection in Producing Old Age, Decay, and Death.

      ‘Supposing organisms ever existed that had not the power of natural reproduction, then since

1

Humboldt’s ‘Ausichten der Natur.’

2

This estimate is derived from observation of the time during which these insects are to be seen upon the wing. Direct observations upon the duration of life in this species are unknown to me.

3

[Sir John Lubbock has now kept a queen ant alive for nearly 15 years. See note 2 {note 18 below} on p. 51.—E. B. P.]

4

[After reading these proofs Dr. A. R. Wallace kindly sent me an unpublished note upon the production of death by means of natural selection, written by him some time between 1865 and 1870. The note contains some ideas on the subject, which were jotted down for further elaboration, and were then forgotten until recalled by the argument of this Essay. The note is of great interest in relation to Dr. Weismann’s suggestions, and with Dr. Wallace’s permission I print it in full below.

‘The Action of Natural Selection in Producing Old Age, Decay, and Death.

‘Supposing organisms ever existed that had not the power of natural reproduction, then since the absorptive surface would only increase as the square of the dimensions while the bulk to be nourished and renewed would increase as the cube, there must soon arrive a limit of growth. Now if such an organism did not produce its like, accidental destruction would put an end to the species. Any organism therefore that, by accidental or spontaneous fission, could become two organisms, and thus multiply itself indefinitely without increasing in size beyond the limits most favourable for nourishment and existence, could not be thus exterminated: since the individual only could be accidentally destroyed,—the race would survive. But if individuals did not die they would soon multiply inordinately and would interfere with each other’s healthy existence. Food would become scarce, and hence the larger individuals would probably decompose or diminish in size. The deficiency of nourishment would lead to parts of the organism not being


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<p>62</p>

The views of most biologists who have worked at this subject agree in all essentials with that expressed above. Bütschli says (Bronn’s ‘Klassen und Ordnungen des Thierreichs,’ Protozoa, p. 148): ‘The process of encystment does not appear to have originally borne any direct relation to reproduction: it appears on the contrary to have taken place originally,—as it frequently does at the present day,—either for the protection of the organism against injurious external influences, such as desiccation or the fatal effects of impure water, etc.; and also to enable the organism, after taking up an unusually abundant supply of food, to assimilate it in safety.’ Balbiani (‘Journ. de Micrographie,’ Tom. V. 1881, p. 293) says in reference to the Infusoria, ‘Un petit nombre d’espèces, au lieu de se multiplier à l’état de vie active, se reproduisent dans une sorte d’état de repos, dit état d’enkystement. Ces sortes de kystes peuvent être désignés sous le nom de kystes de reproduction, par opposition avec d’autres kystes, dans lesquels les Infusoires se renferment pour se soustraire à des conditions devenues défavorables du milieu qu’ils habitent, le manque d’air, le dessèchement, etc.—ceux-ci sont des kystes de conservation....’

<p>63</p>

This is of importance in so far as single individuals might be thus compelled to encyst even when the existing external conditions of life do not require it. The substance which Actinosphaerium, for example, employs in the secretion of its thick siliceous cyst must have been gradually accumulated by means of a process peculiar to the species. We can scarcely be in error if we assume that the silica accumulated in the organism cannot increase to an unlimited extent without injury to the other vital processes and that the secretion of the cyst must take place as soon as the accumulation has exceeded a certain limit. Thus we can understand that encystment may occur without any external necessity. Similarly, certain Entomostraca (e. g. Moina) produce winter-eggs in a particular generation, and these are formed even when the animals are kept in a room protected from cold and desiccation.

<p>64</p>

Upon this point Professor Gruber intends to publish an elaborate memoir.

<p>65</p>

This view has not even been proved for Actinosphaerium, upon which Götte chiefly relies. The observations which we now possess merely indicate that the animal contracts to the smallest volume possible. Compare F. E. Schulze, ‘Rhizopodenstudien,’ I, Arch. f. mikr. Anat. Bd. 10, p. 328; and Karl Brandt, ‘Ueber Actinosphaerium Eichhornii,’ Inaug. Diss.; Halle, 1877.