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The embryological origins of the gene theory

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References

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  23. Ibid.

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  37. E. B.Wilson, The Cell in Development and Inheritance (New York: MacMillan, 1896), pp. 250–251. Claude Bernard claimed that the nucleus was the site of synthetic activity while the cytoplasm was the site of degradative metabolism. Wilson also expressed this notion in appendix to one of his graduate students' papers on cytoplasmic localization. See E. B. Wilson, “On Cleavage and Mosaic Work” (Appendix to H. E. Crampton, Jr.) Arch. Entwicklungsmech., 3 (1896), 19: “Cytoplasmic organization, while affording the immediate conditions for development, is itself a result in the last analysis of the nature of the nuclear substance which represents by its inherent composition the totality of hereditable potence.”

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  38. E. B.Wilson, The Cell in Development and Inheritance (New York: MacMillan, 1896), pp. 262. The concept of “formitive energy” was influential during this period, when much of what was known in physiology centered on energy metabolism. C. O. Whitman introduces it into his description of fertilization (“Clepsine”, p. 252) stating this process to be “a re-union, not of exhausted, but of complementary energies.” To those who favored the environmental determination of sex, these energies were manifest in the anabolism-catabolism ratios which determined the character. Wilson used this concept extensively, linking the “formitive power” directly to Bernard's notions of “chemical synthesis.” Wilson's view that sex differences were caused by differences in the intensity or energy, not substance, of the chromosomes, is consistent with his belief that “the nucleus is the formitive centre of the cell in the chemical sense, and through this is the especial seat of the formitive energy in a morphological sense.” (Cell, p. 261).

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  40. Wilson, “Mosaic Theory,” p. 320. Throughout his research, Wilson maintained this view that nuclear constancy directed epigenetic events through physiological reactions. Davidson (Gene Activity, p. 309) has stated Wilson's position as the idea that “apparently preformed characters can only be regarded as the product of an earlier epigenetic process originating in the oocyte nucleus during oogenesis.” Just as Wilson abolished the dichotomy between mosaic and regulative cleavages, so Wilson integrated apparent preformationism into an epigenetic framework. While Morgan was viewing the problems of heredity and development as identical, Wilson separated the two processes (cf. the quotations referred to in notes 117. and 119, below). Thus, like Roux, he could envision a hereditarily stable nucleus exempt from the epigenetic processes of development. Hereditary factors could be preformed and direct epigenetic developmental processes: “Heredity is effected by the transmission of a nuclear preformation which in the course of development finds expression in a process of cytoplasmic epigenesis” (Wilson, The Cell in Heredity and Development, 3rd ed., [New York: MacMillan, 1925]). This is the same idea Wilson details in the 1896 edition of The Cell in Development and Inheritance, p. 320. Wilson is always emphatic that development is epigenetic; and even at the onset of the chromosome controversy, he repeats this position: “Early in its development the egg contains only a few of these specific stuffs... and that as development goes forward, new stuffs are formed and distributed... The actual progressive development of the protoplasm must be conceived as a process of epigenesis, not of preformation or evolution (E. B. Wilson, “Mosaic Development in the Annelid Egg,” Science, 20 [1904], 750).

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  44. As Wilson once put the argument (“The Problem of Development”, Science, 21 [1905], 288): “The protoplasmic stuffs appear to be only the immediate means or the efficient cause of differentiation, and we still seek its primary determination in the causes that lie more deeply.”

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  50. Ibid.

  51. Science is being used to justify social mores here, since implicit in this theory is that an active woman is unnatural. A similar degree of male supremacy can be seen in the basic argument of the theory, which holds that when times get rough, only the males survive. In 1914, Geddes and Thomson will restate this argument, with the following additon: “We may speak of women's constitution and temper as more conservative, of man's as more unstable... We regard the woman as relatively more anabolic, man as relatively katabolic; and whether this biological hypothesis be a good one or not, it certainly does no social harm” (Problems of sex [Moffat, N.Y., 1914], pp. 205–206). Geddes's views of sex in society are of further value since he was also one of the leading sociologists of his time (and Lewis Mumford's mentor). Geddes sees a distinct sexual dimorphism in the bodies, sensibilities, and aptitudes of men and women. They are not equal but complementary, and he welcomes the advances of women as having the potential of transforming the masculine neotechnic age into a totally human eutechnic era by their humanitarian ideals and inspiration. (I am indebted to David Cahan for suggesting the relationship of Geddes the botanist with Geddes the social theorist).

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  93. Morgan, “Sex Determining Factors,” p. 383. This view recalls van Beneden's hypothesis that each cell is hermaphroditic, containing both male and female chemical determinants. Each unfertilized egg, however, would exclude the male material in its polar bodies. After a similar process occured in the formation of the sperm, the union of gametes would yield a new hermaphroditic assembly whose component materials would interact physiologically to produce a new organism. By the time Morgan was writing, however, Hertwig had already shown that a process analogous to polar body extrusion did not happen in the male, so the resulting union might well be expected to create “double-barreled sexhybrids.”

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  100. Wilson, Cell. p. 320. It should be noted that even at this time, Wilson viewed metabolism as entailing a specific series of reactions, the sex differences being caused by quantitative rather than qualitative variation in cell metabolism. Such hypotheses show Wilson's inclination to be physiological rather than morphological, epigenetic rather than preformationist.

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  122. N. C.Mullins, “The Development of a Scientific Specialty: The Phage Group and the Origins of Molecular Biology,” Minerva, 10, (1972), 51–82. It is interesting to note that disciplines seem to change according to their own vocabularies, as if sensitive to their own metaphors. Copernicus's theory of celestial orbits occasioned a “revolution” in astronomy, while the history of evolutionary thought lends itself readily to terms of natural selection. In embryology, change did not occur as a revolution nor were new ideas selected by a changing social environment. Rather, embryology underwent a metamorphosis. There was continuity of substance between the old and the new disciplines, a small portion of the old structure expanding rapidly to produce a new one from within. During this period, embryology underwent two metamorphoses—once in the group of physiological embryologists who created developmental mechanics out of the descriptive embryology, and again when the group of developmental physiologists concerned with nucleus-cytoplasm relationships created the gene theory.

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    Scott F. Gilbert

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Gilbert, S.F. The embryological origins of the gene theory. Journal of the History of Biology 11, 307–351 (1978). https://doi.org/10.1007/BF00389303

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