Essays Upon Heredity and Kindred Biological Problems. Weismann August

into the former, cause such changes in their nature that at the next succeeding cell-division they are compelled to break up into dissimilar parts.

      At first sight this hypothesis seems to be quite reasonable. It is not only conceivable that particles might proceed from the somatic to the reproductive cells, but the very nutrition of the latter at the expense of the former is a demonstration that such a passage actually takes place. But a closer examination reveals immense difficulties. In the first place, the molecules of the body devoured are never simply added to those of the feeding individual without undergoing any change, but as far as we know, they are really assimilated38, that is, converted into the molecules of the latter. We cannot therefore gain much by assuming that a number of molecules can pass from the growing somatic cells into the growing reproductive cells, and can be deposited unchanged in the latter, so that, at their next division, the molecules are separated to become the somatic cells of the following generation. How can such a process be conceivable, when the colony becomes more complex, when the number of somatic cells becomes so large that they surround the reproductive cells with many layers, and when at the same time by an increasing division of labour a great number of different tissues and cells are produced, all of which must originate de novo from a single reproductive cell? Each of these various elements must, ex hypothesi, give up certain molecules to the reproductive cells; hence those which are in immediate contact with the latter would obviously possess an advantage over those which are more remote. If then any somatic cell must send the same number of molecules to each reproductive cell39, we are compelled to suspend all known physical and physiological conceptions, and must make the entirely gratuitous assumption of an affinity on the part of the molecules for the reproductive cells. Even if we admit the existence of this affinity, its origin and means of control remain perfectly unintelligible if we suppose that it has arisen from differentiation of the complete colony. An unknown controlling force must be added to this mysterious arrangement, in order to marshal the molecules which enter the reproductive cell in such a manner that their arrangement corresponds with the order in which they must emerge as cells at a later period. In short, we become lost in unfounded hypotheses.

      It is well known that Darwin has attempted to explain the phenomena of heredity by means of a hypothesis which corresponds to a considerable extent with that just described. If we substitute gemmules for molecules we have the fundamental idea of Darwin’s provisional hypothesis of pangenesis. Particles of an excessively minute size are continually given off from all the cells of the body; these particles collect in the reproductive cells, and hence any change arising in the organism, at any time during its life, is represented in the reproductive cell40. Darwin believed that he had by this means rendered the transmission of acquired characters intelligible, a conception which he held to be necessary in order to explain the development of species. He himself pointed out that the hypothesis was merely provisional, and that it was only an expression of immediate, and by no means satisfactory knowledge of these phenomena.

      It is always dangerous to invoke some entirely new force in order to understand phenomena which cannot be readily explained by the forces which are already known.

      I believe that an explanation can in this case be reached by an appeal to known forces, if we suppose that characters acquired (in the true sense of the term) by the parent cannot appear in the course of the development of the offspring, but that all the characters exhibited by the latter are due to primary changes in the germ.

      This supposition can obviously be made with regard to the above-mentioned colony with its constituent elements differentiated into somatic and reproductive cells. It is conceivable that the differentiation of the somatic cells was not primarily caused by a change in their own structure, but that it was prepared for by changes in the molecular structure of the reproductive cell from which the colony arose.

      The generally received idea assumes that changes in the external conditions can, in connection with natural selection, call forth persistent changes in an organism; and if this view be accepted it must be as true of all Metazoa as it is of unicellular or of homogeneous multicellular organisms. Supposing that the hypothetical colonies, which were at first entirely made up of similar cells, were to gain some advantages, if in the course of development, the molecules of the reproductive cells, from which each colony arose became distributed irregularly in the resulting organism, there would be a tendency towards the perpetuation of such a change, wherever it appeared as the result of individual variability. As a result of this change the colony would no longer remain homogeneous, and its cells would become dissimilar from the first, because of the altered arrangement of the molecules in the reproductive cells. Nothing prevents us from assuming that, at the same time, the nature of a part of the molecule may undergo still further change, for the molecules are by nature complex, and may split up or combine together.

      If then the reproductive cells have undergone such changes that they can produce a heterogeneous colony as the result of continual division, it follows that succeeding generations must behave in exactly the same manner, for each of them is developed from a portion of the reproductive cell from which the previous generation arose, and consists of the same reproductive substance as the latter.

      From this point of view the exact manner in which we imagine the subsequent differentiation of the colony to be potentially present in the reproductive cell, becomes a matter of comparatively small importance. It may consist in a different molecular arrangement, or in some change of chemical constitution, or it may be due to both these causes combined. The essential point is that the differentiation was originally due to some change in the reproductive cells, just as this change itself produces all the differentiations which appear in the ontogeny of all species at the present day. No one doubts that the reason why this or that form of segmentation takes place, or why this or that species finally appears, is to be found in the ultimate structure of the reproductive cells. And, as a matter of fact, molecular differentiation and grouping, whether present from the beginning or first appearing in the course of development, plays a rôle which can be almost directly observed in certain species. The first segmentation furrow divides the egg of such species into an opaque and a clear half, or, as is often the case among Medusae, into a granular outer layer and a clear central part, corresponding respectively with the ectoderm and endoderm which are formed at a later period. Such early differentiations are only the visible proofs of certain highly complex molecular rearrangements in the cells, and the fact appears to indicate that we cannot be far wrong in maintaining that differentiations which appear in the course of ontogeny depend upon the chemical and physical constitution of the molecules in the reproductive cell.

      At the first appearance of the earliest Metazoa alluded to above, only two kinds of cells, somatic and reproductive, arose from the segmentation of the reproductive cell. The reproductive cells thus formed must have possessed exactly the same molecular structure as the mother reproductive cell, and would therefore pass through precisely the same developmental changes. We can easily imagine that all the succeeding stages in the development of the Metazoa have been due to the same causes which were efficient at the earliest period. Variations in the molecular structure of the reproductive cells would continue to appear, and these would be increased and rendered permanent by means of natural selection, when their results, in the alteration of certain cells in the body, were advantageous to the species. The only condition necessary for the transmission of such changes is that a part of the reproductive substance (the germ-plasm) should always remain unchanged during segmentation and the subsequent building up of the body, or in other words, that such unchanged substance should pass into the organism, and after the lapse of a variable period, should reappear as the reproductive cells. Only in this way can we render to some extent intelligible the transmission of those changes which have arisen in the phylogeny of the species; only thus can we imagine the manner in which the first somatic cells gradually developed in numbers and in complexity.

      It is only by supposing that these changes arose from molecular alterations in the reproductive cell that we can understand how the reproductive cells of the next generation can originate the same changes in the cells which are developed from them; and it is impossible to imagine any way in which the transmission of changes, produced by the direct action of external forces upon the somatic cells, can be brought about

38

Or is an exception perhaps afforded by the nutritive cells of the egg, which occur in many animals?

39

Or more precisely, they must give up as many molecules as would correspond to the number of the kind of cell in question found in the mature organism.

40

See Darwin, ‘The Variation of Animals and Plants under Domestication,’ 1875, vol. ii. chapter xxvii. pp. 349-399.