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William Keith Brooks and the naturalist’s defense of Darwinism in the late-nineteenth century

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William Keith Brooks was an American zoologist at Johns Hopkins University from 1876 until his death in 1908. Over the course of his career, Brooks staunchly defended Darwinism, arguing for the centrality of natural selection in evolutionary theory at a time when alternative theories, such as neo-Lamarckism, grew prominent in American biology. In his book The Law of Heredity (1883), Brooks addressed problems raised by Darwin’s theory of pangenesis. In modifying and developing Darwin’s pangenesis, Brooks proposed a new theory of heredity that sought to avoid the pitfalls of Darwin’s hypothesis. In so doing he strengthened Darwin’s theory of natural selection by undermining arguments for the inheritance of acquired characteristics. In later attacks on neo-Lamarckism, Brooks consistently defended Darwin’s theory of natural selection on logical grounds, continued to challenge the idea of the inheritance of acquired characteristics, and argued that natural selection best explained a wide range of adaptations. Finally, he critiqued Galton’s statistical view of heredity and argued that Galton had resurrected an outmoded typological concept of species, one which Darwin and other naturalists had shown to be incorrect. Brooks’s ideas resemble the “biological species concept” of the twentieth century, as developed by evolutionary biologist Ernst Mayr and others. The late-nineteenth century was not a period of total “eclipse” of Darwinism, as biologists and historians have hitherto seen it. Although the “Modern Synthesis” refers to the reconciliation of post-Mendelian genetics with evolution by natural selection, we might adjust our understanding of how the synthesis developed by seeing it as the culmination of a longer discussion that extends back to the late-nineteenth century.

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  1. On Brooks’s career in the context of American morphology, see Benson (1979). On his prominent students, see Maienschein (1991). Christine Keiner (2010) has analyzed Brooks’s scientific work on the oyster in the context of Chesapeake aquaculture.

  2. On the naturalist’s perspective, see Allen (1981).

  3. The eclipse metaphor for this period was first proposed by zoologist Julian Huxley (1942, p. 22) in his effort to champion the return of natural selection to prominence during the “Modern Synthesis.” This characterization, which I believe to be overstated, has been most explicit in historian Bowler’s work (1983, 1988).

  4. Largent (2009) has criticized Bowler’s use of the eclipse metaphor as a historical narrative device because of its teleological implications.

  5. Benson (1979, 2010) has previously framed Brooks’s heredity theory as part of a defense of Darwinism. In this article, I aim to show that Brooks’s heredity work was but one strategy of many that he adopted in a grander defense of Darwinism, which took place over the course of his career. His other strategies included attacking both neo-Lamarckism and Galton’s statistical view of heredity.

  6. On the meaning of “Darwinism” in the late-nineteenth century, see philosopher and evolutionist John Fiske’s useful survey (Fiske 1879, pp. 11–12).

  7. An abstract of Brooks’s paper was published the following year (Brooks 1877).

  8. Keith Benson has elsewhere described Brooks’s modification of Darwin’s pangenesis (Benson 1979, 1981, 2010). I will build on Benson’s work and emphasize the role played by the physiological division of labor in Brooks’s theory.

  9. Brooks’s evolutionary thought, like Darwin’s, was informed by metaphorical reasoning. The key metaphor for Brooks was the division of labor, a powerful concept that Darwin had discussed in the Origin of Species. Darwin cited therein the work of Belgian naturalist Henri Milne-Edwards, who developed the concept of physiological division of labor from the domain of economics (Darwin 1859, p. 115). As a metaphorical framework, the division of labor led Brooks to assume that natural selection, like classical economics, yielded increasingly efficient systems.

  10. Brooks often used active verbs such as “to seize” in describing how natural selection operated, attributing to it a fervent agency.

  11. On the reception of Brook’s theory, see Benson (1979).

  12. His wife Amelia became seriously ill in the late 1890s and he devoted a large portion of his time away from campus caring for her until her death in 1899. Brooks’s own health steadily declined as well, and he became less productive in his final years while under the care of his daughter.

  13. Noteworthy examples include: Brooks (1895a, b, 1906).

  14. George Romanes listed Brooks as one of the most prominent American neo-Lamarckians, and Brooks sternly refuted that characterization in the pages of Natural Science. These three articles (two by Brooks, one by a Lamarckian inquisitor) were republished in the Johns Hopkins University Circular (Brooks 1896d, pp. 75–79).

  15. On neo-Lamarckism in American evolutionary thought, see Bowler (1983), Numbers, (1998), and Pfeiffer (1965).

  16. Neo-Lamarckism is distinct from Lamarck’s pre-Darwinian theory of directed evolution. Nevertheless, Brooks referred to these late-nineteenth century proponents of acquired heredity as “Lamarckians.”

  17. Brooks’s criticism of neo-Larmackian speculation seems equally applicable to his own postulation of the gemmule theory put forth earlier in his career. Perhaps developments in experimental biology in the 1880s and 1890s convinced him to be more cautious about developing speculative theories that were prone to experimental disproof.

  18. The three articles—two by Brooks and a riposte by Cunningham—were reprinted from Natural Science (February and May of 1896) in the Johns Hopkins University Circular (Brooks 1896d).

  19. Emphasis in original.

  20. Brooks’s articles on Galton (Brooks 1895b, 1896a, b) were later published, mostly unchanged, in his Foundations of Zoology (Brooks 1899).

  21. According to Michael Bulmer, Galton’s approach to heredity gradually shifted from physiological and mechanistic to statistical and mathematical (Bulmer 1999).

  22. Ruth Cowan (1972) has argued that Galton had settled on this definition of heredity in his 1889 book, Natural Inheritance. An inherently mathematical concept, Galton’s heredity encompassed the several related phenomena of inheritance, reversion (a form of inheritance as he conceived it), and variation. Brooks too saw heredity as encompassing interdependent phenomena, and in fact had argued before Galton that inheritance and variation were indeed two sides of the same coin, both the product of the same cause. Even after Brooks gave up his pangenesis theory, he never wavered from the belief that inheritance and variation were simply two ways of considering a single phenomenon.

  23. In fact Galton limited his study almost entirely to the analysis of traits in male lineages, ignoring females.

  24. As Ruth Cowan has shown, Galton’s goal of mathematizing the study of heredity was directly linked to his interest in creating a new science of eugenics (Cowan 1969, 1972).

  25. The essay was originally given as a lecture to the Baltimore to the Society of Naturalists in 1894. It was later reprinted with some further additions in Brooks’s Foundations of Zoology (1899).

  26. Bionomics was a laboratory method developed by Kellogg to understand evolution by studying organisms in controlled settings that recreated their natural environments.


Primary Sources

  • Brooks, W. K. (1877). A provisional hypothesis of pangenesis. The American Naturalist, 11, 144–147.

    Article  Google Scholar 

  • Brooks, W. K. (1883). The law of heredity. A study of the cause of variation, and the origin of living organisms. Baltimore: J. Murphy.

    Book  Google Scholar 

  • Brooks, W. K. (1887). Francis Galton on the persistency of type. The American Journal of Psychology, 1, 173-179.  

  • Brooks, W. K. (1895a). Tyranny of the monistic creed. Science, 1, 382–384.

    Google Scholar 

  • Brooks, W. K. (1895b). An inherent error in the views of Galton and Weismann on variation. Science, 1, 121–126.

    Article  Google Scholar 

  • Brooks, W. K. (1896a). The study of inheritance: A review of the writings of Francis Galton. Popular Science Monthly, 48, 480–492.

    Google Scholar 

  • Brooks, W. K. (1896b). The study of inheritance: A review of the writings of Francis Galton [Concluded]. Popular Science Monthly, 48, 617–626.

    Google Scholar 

  • Brooks, W. K. (1896c). What is truth? Science, 3, 779.

    Article  Google Scholar 

  • Brooks, W. K. (1896d). Lyell and Lamarck: A consideration for Lamarckians. Johns Hopkins University Circular, 15, 75–79.

    Google Scholar 

  • Brooks, W. K. (1899). The foundations of zoology. London: MacMillan and Co.

    Book  Google Scholar 

  • Brooks, W. K. (1906). Heredity and variation: Logical and biological. Proceedings of the American Philosophical Society, 45, 70–76.

    Google Scholar 

  • Conklin, E. G. (1896). Discussion of the factors of organic evolution from the embryological standpoint. Proceedings of the American Philosophical Society, 35, 78–88.

    Google Scholar 

  • Conklin, E. G. (1909). The life and work of Professor Brooks. The Anatomical Record, 3, 1–13.

    Article  Google Scholar 

  • Darwin, C. (1859). On the origin of species by means of natural selection, or the preservation of favoured races in the struggle for life. London: John Murray.

    Book  Google Scholar 

  • Darwin, C. (1868). The variation of animals and plants under domestication (Vol. 2). London: John Murray.

    Google Scholar 

  • Fiske, J. (1879). Darwinism and other essays. London: MacMillan and Co.

    Book  Google Scholar 

  • Galton, F. (1870). Experiments in pangenesis, by breeding from rabbits of a pure variety, into whose circulation blood taken from other varieties had previously been largely transfused. Proceedings of the Royal Society of London, 19, 393–410.

    Article  Google Scholar 

  • Galton, F. (1889). Natural inheritance. London: MacMillan and Co.

    Book  Google Scholar 

Secondary Sources

  • Allen, G. (1978). Thomas Hunt Morgan: The man and his science. Princeton: Princeton University Press.

    Google Scholar 

  • Allen, G. (1981). Morphology and twentieth-century biology: A response. Journal of the History of Biology, 14, 159–176.

    Article  Google Scholar 

  • Benson, K. (1979). William Keith Brooks (18481908): A case study in morphology and the development of American biology. Ph.D. dissertation, Oregon State University.

  • Benson, K. (1981). Problems of individual development: descriptive embryological morphology in american at the turn of the century. Journal of the History of Biology, 14, 115–128.

    Article  Google Scholar 

  • Benson, K. (1985). American morphology in the late nineteenth century. Journal of the History of Biology, 18, 163–205.

    Article  Google Scholar 

  • Benson, K. (1987). H. Newell Martin, W.K. Brooks, and the reformation of American biology. American Zoologist, 27, 759–771.

    Google Scholar 

  • Benson, K. (2010). William Keith Brooks (1848–1908) and the defense of late-nineteenth century Darwinian evolutionary theory. In A. Barahona, E. Suárez Díaz & H.-J. Rheinberger (Eds.), Hereditary hourglass: Genetics and epigenetics (1868–2000) (pp. 23–34). Max-Planck-Institute for the History of Science. Accessed October 12, 2011 from

  • Bowler, P. J. (1983). The eclipse of Darwinism: Anti-Darwinian evolution Theories in the decades around 1900. Baltimore: Johns Hopkins University Press.

    Google Scholar 

  • Bowler, P. J. (1988). The Non-Darwinian revolution. Baltimore: Johns Hopkins University Press.

    Google Scholar 

  • Bulmer, M. (1999). The development of Francis Galton’s ideas on the mechanism of heredity. Journal of the History of Biology32, 263–292.

  • Cowan, R. (1969). Sir Francis Galton and the Study of Heredity in the Nineteenth Century. Ph.D. dissertation, Johns Hopkins University.

  • Cowan, R. (1972). Francis Galton’s contributions to genetics. Journal of the History of Biology, 5, 389–412.

    Article  Google Scholar 

  • Huxley, J. (1942). Evolution: The modern synthesis. London: G. Allen & Unwin Ltd.

    Google Scholar 

  • Keiner, C. (2010). The oyster question: Scientists, watermen, and the Maryland Chesapeake Bay since 1880. Athens: University of Georgia Press.

    Google Scholar 

  • Largent, M. (1999). Bionomics: Vernon Lyman Kellogg and the defense of Darwinism. Journal of the History of Biology, 32, 465–488.

    Article  Google Scholar 

  • Largent, M. (2009). The so-called ‘Eclipse of Darwinism. In J. Cain & M. Ruse (Eds.), Descended from Darwin: Insights into American evolutionary studies, 19251950 (pp. 3–21). Philadelphia: American Philosophical Society. Retrieved June 8, 2013, from

  • Maienschein, J. (1991). Transforming traditions in American biology: 1890–1915. Baltimore: Johns Hopkins University Press.

    Google Scholar 

  • Mayr, E., & Provine, W. (1980). The evolutionary synthesis: Perspectives on the unification of biology. Cambridge: Harvard University Press.

    Book  Google Scholar 

  • Numbers, R. (1998). Darwinism comes to America. Cambridge: Harvard University Press.

    Google Scholar 

  • Pfeifer, E. J. (1965). The genesis of American Neo-Lamarckism. Isis, 56, 156–167.

    Article  Google Scholar 

  • Rao, V., & Nanjundiah, V. (2011). J. B. S. Haldane, Ernst Mayr and the Beanbag genetics dispute. Journal of the History of Biology, 44, 233–281.

    Article  Google Scholar 

  • Stocking, G. W. (1962). Lamarckianism in American Social Science: 1890–1915. Journal of the History of Ideas, 23, 239–256.

    Article  Google Scholar 

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Nash, R. William Keith Brooks and the naturalist’s defense of Darwinism in the late-nineteenth century. HPLS 37, 158–179 (2015).

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