Studies in the Theory of Descent, Volume I. Weismann August

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СКАЧАТЬ with Haeckel’s progressive, and the latter with his retrogressive metagenesis. Metagenesis may further originate in various ways. In the first place, from metamorphosis, as for example, in the propagation of the celebrated Cecidomyia with nursing larvæ. The power which these larvæ possess of propagating themselves asexually has evidently been acquired as a secondary character, as appears from the fact that there are many species of the same genus the larvæ of which do not nurse, these larvæ being themselves undoubted secondary forms produced by the adaptation of this stage of phyletic development to a mode of life widely different from that of the later stages. In the form now possessed by these larvæ they could never have represented the final stage of their ontogeny, neither could they have formerly possessed the power of sexual propagation. The conclusion seems inevitable that metagenesis has here proceeded from metamorphosis; that is to say, one stage of the ontogeny, by acquiring asexual propagation, has changed the originally existing metamorphosis into metagenesis.

      Lubbock39 is undoubtedly correct when, for cases like that just mentioned, he attempts to derive alternation of generations from metamorphosis. But if we exclude heterogenesis there still remain a large number of cases of true metagenesis which cannot be explained from this point of view.

      It must be admitted, with Haeckel, that the alternation of generations in the Hydromedusæ and Trematoda does not depend, as in the case of Cecidomyia, upon the larvæ having acquired the power of nursing, but that the inferior stages of these species always possessed this power which they now only preserve. The nursing Trematode larvæ now existing may possibly have been formerly able to propagate themselves also sexually, this mode of propagation having at the present time been transferred to a later phyletic stage. In this case, therefore, metagenesis was not properly produced by metamorphosis, but arose therefrom in the course of the phyletic development, the earlier phyletic stages abandoning the power of sexual reproduction, and preserving the asexual mode of propagation. A third way in which metagenesis might originate is through polymorphosis. When the latter is combined with asexual reproduction, as is especially the case with the Hydrozoa, metagenesis may be derived therefrom. The successive stages of transformation of one and the same physiological individual do not in these cases serve as the point of departure for alternation of generation, but the different contemporary forms living gregariously into which the species has become divided through functional differentiation of the various individuals of the same stock. Individuals are here produced which alone acquire the power of sexual reproduction, and metagenesis is thus brought about, these individuals detaching themselves from the stock on which they originated, while the rest of the individuals remain in combination, and retain the asexual mode of propagation. No sharp distinction can be otherwise drawn between this and the cases previously considered.40 The difference consists only in the whole cycle of reproduction being performed by one stock; both classes have the common character that the different phyletic stages never appear in the same individual (metamorphosis), but in the course of further phyletic development metagenesis at the same time arises, i.e. the division of these stages among a succession of individuals. We are therefore able to distinguish this primary metagenesis from the secondary metagenesis arising from metamorphosis.

      It is not here my intention to enter into the ultimate causes of metagenesis; in this subject we should only be able to advance by making vague hypotheses. The phenomenon of seasonal dimorphism, with which this work has mainly to deal, is evidently far removed from metagenesis, and it was to make this clear that the foregoing observations were brought forward. The characters common in the origin of metagenesis are to be found, according to the views previously set forth, in the facts that here the faculty of asexual and of sexual reproduction is always distributed among several phyletic stages of development which succeed each other in an ascending series (progressive metagenesis of Haeckel), whereas I find differences only in the fact that the power of asexual propagation may (in metagenesis) be either newly acquired (larva of Cecidomyia) or preserved from previous ages (Hydroida). It seems that in this process sexual reproduction is without exception lost by the earlier, and remains confined solely to the most recent stages.

      From the investigations on seasonal dimorphism it appears that a cycle of generations can arise in an entirely different way. In this case a series of generations originally alike are made dissimilar by external influences. This appears to me of the greatest importance, since seasonal dimorphism is without doubt closely related to that mode of reproduction which has hitherto been exclusively designated as heterogenesis, and a knowledge of its mode of origination must therefore throw light on the nature and origin of heterogenesis in general.

      In seasonal dimorphism, as I have attempted to show, it is the direct action of climate, and indeed chiefly that of temperature, which brings about the change in some of the generations. Since these generations have been exposed to the alternating influence of the summer and winter temperature a periodical dimorphism has been developed – a regular cycle of dissimilar generations. It has already been asserted that the consecutive generations of a species comport themselves with respect to heredity in a manner precisely similar to that of the ontogenetic stages, and at the same time such succeeding generations point out the parallelism between metamorphosis and heterogenesis. If influences capable of directly or indirectly producing changes operate on any particular stage of development, these changes are always transmitted to the same stage. Upon this metamorphosis depends. In a precisely similar manner changes which operated periodically on certain generations (1, 3, 5, for instance) are transmitted to these generations only, and not to the intermediate ones. Upon this depends heterogenesis. We have just been led to the comprehension of heterogenesis by cyclical heredity, by the fact that a cycle is produced whenever a series of generations exists under regularly alternating influences. In this cycle newly acquired changes, however minute in character at first, are only transmitted to a later, and not to the succeeding generation, appearing only in the one corresponding, i.e. in that generation which exists under similar transforming influences. Nothing can more clearly show the extreme importance which the conditions of life must have upon the formation and further development of species than this fact. At the same time nothing shows better that the action of these conditions is not suddenly and violently exerted, but that it rather takes place by small and slow operations. In these cases the long-continued accumulation of imperceptibly small variations proves to be the magic means by which the forms of the organic world are so powerfully moulded. By the application of even the greatest warmth nobody would be able to change the winter form of A. Levana into the summer form; nevertheless, the summer warmth, acting regularly on the second and third generations of the year, has, in the course of a lengthened period, stamped these two generations with a new form without the first generation being thereby changed. In the same region two different climatic varieties have been produced (just as in the majority of cases climatic varieties occur only in separate regions) which alternate with each other, and thus give rise to a cycle of which each generation propagates itself sexually.

      But even if seasonal dimorphism is to be ascribed to heterogenesis, it must by no means be asserted that those cases of cyclical propagation hitherto designated as heterogenesis are completely identical with seasonal dimorphism. Their identity extends only to their origin and manner of development, but not to the mode of operation of the causes which bring about their transformation. Both phenomena have a common mode of origination, arising from similar (monomorphic) sexual generations and course of development, a cycle of generations with gradually diverging characters coming into existence by the action of alternating influences. On the other hand, the nature of the changes by which the secondary differs from the primary generation may be referred to another mode of action of the exciting causes. In seasonal dimorphism the differences between the two generations are much less than in other cases of heterogenesis. These differences are both quantitatively less, and are likewise qualitative, affecting only characters of biological insignificance.41 The variations in question are mostly restricted to the marking and colouring of the wings and body, occasionally affecting also the form of the wing, and in a few cases the size of the body (Plebeius Amyntas), whilst the bodily structure – so far at least as my investigations extend – appears to be the same СКАЧАТЬ