The Gene

by Siddhartha Mukherjee

  John Henslow, the botanist and geologist: Duane Isely, One Hundred and One Botanists (Ames: Iowa State University, 1994), “John Stevens Henslow (1796–1861).”

  The first, Natural Theology, published in 1802: William Paley, The Works of William Paley . . . Containing His Life, Moral and Political Philosophy, Evidences of Christianity, Natural Theology, Tracts, Horae Paulinae, Clergyman’s Companion, and Sermons, Printed Verbatim from the Original Editions. Complete in One Volume (Philadelphia: J. J. Woodward, 1836).

  The second book, A Preliminary Discourse: John F. W. Herschel, A Preliminary Discourse on the Study of Natural Philosophy. A Facsim. of the 1830 Ed. (New York: Johnson Reprint, 1966).

  “To ascend to the origin of things”: Ibid., 38.

  “Battered relics of past ages”: Martin Gorst, Measuring Eternity: The Search for the Beginning of Time (New York: Broadway Books, 2002), 158.

  “mystery of mysteries”: Charles Darwin, On the Origin of Species by Means of Natural Selection (London: Murray, 1859), 7.

  dominated by so-called parson-naturalists: Patrick Armstrong, The English Parson-Naturalist: A Companionship between Science and Religion (Leominster, MA: Gracewing, 2000), “Introducing the English Parson-Naturalist.”

  In August 1831, two months after his graduation: John Henslow, “Darwin Correspondence Project,” Letter 105, https://www.darwinproject.ac.uk/letter/DCP-LETT-105.xml.

  The Beagle lifted anchor on December 27, 1831: Darwin, Autobiography of Charles Darwin, “Voyage of the ‘Beagle.’ ”

  Charles Lyell’s Principles of Geology: Charles Lyell, Principles of Geology: Or, The Modern Changes of the Earth and Its Inhabitants Considered as Illustrative of Geology (New York: D. Appleton, 1872).

  Lyell had argued (radically, for his time): Ibid., “Chapter 8: Difference in Texture of the Older and Newer Rocks.”

  In September 1832, exploring the gray cliffs: Charles Darwin, Geological Observations on the Volcanic Islands and Parts of South America Visited during the Voyage of H.M.S. “Beagle” (New York: D. Appleton, 1896), 76–107.

  The skull belonged to a megatherium: David Quammen, “Darwin’s first clues,” National Geographic 215, no. 2 (2009): 34–53.

  In 1835, the ship left Lima: Charles Darwin, Charles Darwin’s Letters: A Selection, 1825–1859, ed. Frederick Burkhardt (Cambridge: University of Cambridge, 1996), “To J. S. Henslow 12 [August] 1835,” 46–47.

  On October 20, Darwin returned to sea: G. T. Bettany and John Parker Anderson, Life of Charles Darwin (London: W. Scott, 1887), 47.

  rather than all species radiating out: Duncan M. Porter and Peter W. Graham, Darwin’s Sciences (Hoboken, NJ: Wiley-Blackwell, 2015), 62–63.

  As an afterthought, he added, “I think”: Ibid., 62.

  In the spring of 1838, as Darwin tore into a new journal: Timothy Shanahan, The Evolution of Darwinism: Selection, Adaptation, and Progress in Evolutionary Biology (Cambridge: Cambridge University Press, 2004), 296.

  But the answer that came to him in October 1838: Barry G. Gale, “After Malthus: Darwin Working on His Species Theory, 1838–1859” (PhD diss., University of Chicago, 1980).

  In 1798, writing under a pseudonym, Malthus: Thomas Robert Malthus, An Essay on the Principle of Population (Chicago: Courier Corporation, 2007).

  “sickly seasons, epidemics, pestilence and plague”: Arno Karlen, Man and Microbes: Disease and Plagues in History and Modern Times (New York: Putnam, 1995), 67.

  “It at once struck me”: Charles Darwin, On the Origin of Species by Means of Natural Selection, ed. Joseph Carroll (Peterborough, Canada: Broadview Press, 2003), 438.

  the phrase survival of the fittest was borrowed: Gregory Claeys, “The ‘Survival of the Fittest’ and the Origins of Social Darwinism,” Journal of the History of Ideas 61, no. 2 (2000): 223–40.

  In 1844, he distilled the crucial parts: Charles Darwin, The Foundations of the Origin of Species, Two Essays Written in 1842 and 1844, ed. Francis Darwin (Cambridge: Cambridge University Press, 1909), “Essay of 1844.”

  Alfred Russel Wallace, published a paper: Alfred R. Wallace, “XVIII.—On the law which has regulated the introduction of new species,” Annals and Magazine of Natural History 16, no. 93 (1855): 184–96.

  Wallace had been born to a middle-class family: Charles H. Smith and George Beccaloni, Natural Selection and Beyond: The Intellectual Legacy of Alfred Russel Wallace (Oxford: Oxford University Press, 2008), 10.

  but on the hard-back benches of the free library: Ibid., 69.

  Like Darwin, Wallace had also embarked: Ibid., 12.

  Wallace moved from the Amazon basin: Ibid., ix.

  “The answer was clearly”: Benjamin Orange Flowers, “Alfred Russel Wallace,” Arena 36 (1906): 209.

  In June 1858, Wallace sent Darwin a tentative draft: Alfred Russel Wallace, Alfred Russel Wallace: Letters and Reminiscences, ed. James Marchant (New York: Arno Press, 1975), 118.

  On July 1, 1858, Darwin’s and Wallace’s papers were read: Charles Darwin, The Correspondence of Charles Darwin, vol. 13, ed. Frederick Burkhardt, Duncan M. Porter, and Sheila Ann Dean, et al. (Cambridge: Cambridge University Press, 2003), 468.

  The next May, the president of the society remarked: E. J. Browne, Charles Darwin: The Power of Place (New York: Alfred A. Knopf, 2002), 42.

  “I heartily hope that my Book”: Charles Darwin, The Correspondence of Charles Darwin, vol. 7, ed. Frederick Burkhardt and Sydney Smith (Cambridge: Cambridge University Press, 1992), 357.

  “All copies were sold [on the] first day”: Charles Darwin, The Life and Letters of Charles Darwin (London: John Murray, 1887), 70.

  “The conclusions announced by Mr. Darwin are such”: “Reviews: Darwin’s Origins of Species,” Saturday Review of Politics, Literature, Science and Art 8 (December 24, 1859): 775–76.

  “We imply that his work [is] one of the most important that”: Ibid.

  “light will be thrown on the origin of man”: Charles Darwin, On the Origin of Species, ed. David Quammen (New York: Sterling, 2008), 51.

  “intellectual husks”: Richard Owen, “Darwin on the Origin of Species,” Edinburgh Review 3 (1860): 487–532.

  “One’s imagination must fill up very wide blanks”: Ibid.

  The “Very Wide Blank”

  The “Very Wide Blank”: Darwin, Correspondence of Charles Darwin, Darwin’s letter to Asa Gray, September 5, 1857, https://www.darwinproject.ac.uk/letter/DCP-LETT-2136.xml.

  Now, I wonder if: Alexander Wilford Hall, The Problem of Human Life: Embracing the “Evolution of Sound” and “Evolution Evolved,” with a Review of the Six Great Modern Scientists, Darwin, Huxley, Tyndall, Haeckel, Helmholtz, and Mayer (London: Hall & Company, 1880), 441.

  In Lamarck’s view: Monroe W. Strickberger, Evolution (Boston: Jones & Bartlett, 1990), “The Lamarckian Heritage.”

  “with a power proportional to the length of time”: Ibid., 24.

  driving himself to the brink: James Schwartz, In Pursuit of the Gene: From Darwin to DNA (Cambridge, MA: Harvard University Press, 2008), 2.

  minute particles containing hereditary information—gemmules: Ibid., 2–3.

  blending inheritance—was already familiar: Brian Charlesworth and Deborah Charlesworth, “Darwin and genetics,” Genetics 183, no. 3 (2009): 757–66.

  Darwin dubbed his theory pangenesis: Ibid., 759–60.

  a new manuscript, The Variation of Animals: Charles Darwin, The Variation of Animals and Plants under Domestication, vol. 2 (London: O. Judd, 1868).

  “It is a rash and crude hypothesis”: Darwin, Correspondence of Charles Darwin, vol. 13, “Letter to T. H. Huxley,” 151.

  “Pangenesis will be called a mad dream”: Charles Darwin, The Life and Letters of Charles Darwin: Including Autobiographical Chapter, vol. 2., ed. Francis Darwin (New York: Appleton, 1896), “C. Darwin to Asa Gray,” October 16, 1867, 256.

  “The [variant] will be swamped”: Fleeming Jenkin, “The Origin of Species,” North British Review 47 (1867): 158.


  There was no denying: In fairness to Darwin, he had sensed the problem in “blending inheritance” even without Jenkin’s interjection. “If varieties be allowed freely to cross, such varieties will be constantly demolished . . . any small tendency in them to vary will be constantly counteracted,” he wrote in his notes.

  “Experiments in Plant Hybridization”: G. Mendel, “Versuche über Pflanzen-Hybriden,” Verhandlungen des naturforschenden Vereins Brno 4 (1866): 3–47 (Journal of the Royal Horticultural Society 26 [1901]: 1–32).

  he made extensive handwritten notes on pages 50, 51, 53, and 54: David Galton, “Did Darwin read Mendel?” Quarterly Journal of Medicine 102, no. 8 (2009): 588, doi:10.1093/qjmed/hcp024.

  “Flowers He Loved”

  “Flowers He Loved”: Edward Edelson, Gregor Mendel and the Roots of Genetics (New York: Oxford University Press, 1999), “Clemens Janetchek’s Poem Describing Mendel after His Death,” 75.

  “We want only to disclose the [nature of] matter and its force”: Jiri Sekerak, “Gregor Mendel and the scientific milieu of his discovery,” ed. M. Kokowski (The Global and the Local: The History of Science and the Cultural Integration of Europe, Proceedings of the 2nd ICESHS, Cracow, Poland, September 6–9, 2006).

  “The whole organic world is the result”: Hugo de Vries, Intracellular Pangenesis; Including a Paper on Fertilization and Hybridization (Chicago: Open Court, 1910), “Mutual Independence of Hereditary Characters.”

  Gregor Mendel decided to return to Vienna: Henig, Monk in the Garden, 60.

  “remained constant without exception”: Eric C. R. Reeve, Encyclopedia of Genetics (London: Fitzroy Dearborn, 2001), 62.

  Contrary to later belief: Mendel had several predecessors who had studied plant hybrids just as intensively, except, perhaps, without Mendel’s immersion in numbers and quantification. In the 1820s, English botanists, such as T. A. Knight, John Goss, Alexander Seton, and William Herbert—attempting to breed more vigorous agricultural plants—had performed experiments with plant hybrids that were strikingly similar to Mendel’s. In France, Augustine Sageret’s work on melon hybrids was also similar to Mendel’s work. The most intensive work on plant hybrids immediately preceding Mendel was performed by the German botanist Josef Kölreuter, who had bred Nicotania hybrids. Kölreuter’s work was followed by the work of Karl von Gaertner and Charles Naudin in Paris. Darwin had actually read Sageret’s and Naudin’s studies, both of which suggested the particulate quality of hereditary information, but Darwin had failed to appreciate their importance.

  “the history of the evolution of organic forms”: Gregor Mendel, Experiments in Plant Hybridisation (New York: Cosimo, 2008), 8.

  By the late summer of 1857, the first hybrid peas: Henig, Monk in the Garden, 81. More details in “Chapter 7: First Harvest.”

  “How small a thought it takes to fill”: Ludwig Wittgenstein, Culture and Value, trans. Peter Winch (Chicago: University of Chicago Press, 1984), 50e.

  Mendel termed these overriding traits: Henig, Monk in the Garden, 86.

  In some of these third-generation crosses: Ibid., 130.

  “It requires indeed some courage”: Mendel, Experiments in Plant Hybridization, 8.

  Mendel presented his paper: Henig, Monk in the Garden, “Chapter 11: Full Moon in February,” 133–47. A second portion of Mendel’s paper was read on March 8, 1865.

  Mendel’s paper was published in: Mendel, “Experiments in Plant Hybridization,” www.mendelweb.org/Mendel.html.

  It is likely that he sent one to Darwin: Galton, “Did Darwin Read Mendel?” 587.

  “one of the strangest silences in the history of biology”: Leslie Clarence Dunn, A Short History of Genetics: The Development of Some of the Main Lines of Thought, 1864–1939 (Ames: Iowa State University Press, 1991), 15.

  “only empirical . . . cannot be proved rational”: Gregor Mendel, “Gregor Mendel’s letters to Carl Nägeli, 1866–1873,” Genetics 35, no. 5, pt. 2 (1950): 1.

  “I knew that the results I obtained”: Allan Franklin et al., Ending the Mendel-Fisher Controversy (Pittsburgh, PA: University of Pittsburgh Press, 2008), 182.

  “an isolated experiment might be doubly dangerous”: Mendel, “Letters to Carl Nägeli,” April 18, 1867, 4.

  In November 1873, Mendel wrote his last letter to Nägeli: Ibid., November 18, 1867, 30–34.

  “I feel truly unhappy that I have to neglect”: Gian A. Nogler, “The lesser-known Mendel: His experiments on Hieracium,” Genetics 172, no. 1 (2006): 1–6.

  On January 6, 1884, Mendel died: Henig, Monk in the Garden, 170.

  “Gentle, free-handed, and kindly . . . Flowers he loved”: Edelson, Gregor Mendel, “Clemens Janetchek’s Poem Describing Mendel after His Death,” 75.

  “A Certain Mendel”

  The origin of species is a natural phenomenon: Lucius Moody Bristol, Social Adaptation: a Study in the Development of the Doctrine of Adaptation as a Theory of Social Progress (Cambridge, MA: Harvard University Press, 1915), 70.

  The origin of species is an object of inquiry: Ibid.

  The origin of species is an object of experimental investigation: Ibid.

  In the summer of 1878: Peter W. van der Pas, “The correspondence of Hugo de Vries and Charles Darwin,” Janus 57: 173–213.

  “margin was too small”: Mathias Engan, Multiple Precision Integer Arithmetic and Public Key Encryption (M. Engan, 2009), 16–17.

  “In another work I shall discuss”: Charles Darwin, The Variation of Animals & Plants under Domestication, ed. Francis Darwin (London: John Murray, 1905), 5.

  Darwin died in 1882: “Charles Darwin,” Famous Scientists, http://www.famousscientists.org/charles-darwin/.

  In 1883, with rather grim determination: James Schwartz, In Pursuit of the Gene: From Darwin to DNA (Cambridge, MA: Harvard University Press, 2008), “Pangenes.”

  Weismann called this hereditary material germplasm: August Weismann, William Newton Parker, and Harriet Rönnfeldt, The Germ-Plasm; a Theory of Heredity (New York: Scribner’s, 1893).

  In a landmark paper written in 1897: Schwartz, In Pursuit of the Gene, 83.

  He called these particles “pangenes”: Ida H. Stamhuis, Onno G. Meijer, and Erik J. A. Zevenhuizen, “Hugo de Vries on heredity, 1889–1903: Statistics, mendelian laws, pangenes, mutations,” Isis (1999): 238–67.

  “I know that you are studying hybrids”: Iris Sandler and Laurence Sandler, “A conceptual ambiguity that contributed to the neglect of Mendel’s paper,” History and Philosophy of the Life Sciences 7, no. 1 (1985): 9.

  “Modesty is a virtue”: Edward J. Larson, Evolution: The Remarkable History of a Scientific Theory (New York: Modern Library, 2004).

  That same year de Vries published his monumental study: Hans-Jörg Rheinberger, “Mendelian inheritance in Germany between 1900 and 1910. The case of Carl Correns (1864–1933),” Comptes Rendus de l’Académie des Sciences—Series III—Sciences de la Vie 323, no. 12 (2000): 1089–96, doi:10.1016/s0764-4469(00)01267-1.

  “I too still believed that I had found something new”: Url Lanham, Origins of Modern Biology (New York: Columbia University Press, 1968), 207.

  “by a strange coincidence”: Carl Correns, “G. Mendel’s law concerning the behavior of progeny of varietal hybrids,” Genetics 35, no. 5 (1950): 33–41.

  de Vries stumbled on an enormous, invasive: Schwartz, In Pursuit of the Gene, 111.

  He called them mutants: Hugo de Vries, The Mutation Theory, vol. 1 (Chicago: Open Court, 1909).

  For William Bateson, the English biologist: John Williams Malone, It Doesn’t Take a Rocket Scientist: Great Amateurs of Science (Hoboken, NJ: Wiley, 2002), 23.

  “We are in the presence of a new principle”: Schwartz, In Pursuit of the Gene, 112.

  “I am writing to ask you”: Nicholas W. Gillham, “Sir Francis Galton and the birth of eugenics,” Annual Review of Genetics 35, no. 1 (2001): 83–101.

  First, he independently confirmed Mendel’s work: Other scientists, including Reginald Punnett and Lu
cien Cuenot, provided crucial experimental support for Mendel’s laws. In 1905, Punnett authored Mendelism, considered the first textbook of modern genetics.

  “His linen is foul. I daresay”: Alan Cock and Donald R. Forsdyke, Treasure Your Exceptions: The Science and Life of William Bateson (Dordrecht: Springer Science & Business Media, 2008), 186.

  Nicknamed “Mendel’s bulldog”: Ibid., “Mendel’s Bulldog (1902–1906),” 221–64.

  “man’s outlook on the world”: William Bateson, “Problems of heredity as a subject for horticultural investigation,” Journal of the Royal Horticultural Society 25 (1900–1901): 54.

  “No single word in common use”: William Bateson and Beatrice (Durham) Bateson, William Bateson, F.R.S., Naturalist; His Essays & Addresses, Together with a Short Account of His Life (Cambridge: Cambridge University Press, 1928), 93.

  In 1905, still struggling for an alternative: Schwartz, In Pursuit of the Gene, 221.

  “What will happen when . . . enlightenment actually comes to pass”: Bateson and Bateson, William Bateson, F.R.S., 456.

  Eugenics

  Improved environment and education: Herbert Eugene Walter, Genetics: An Introduction to the Study of Heredity (New York: Macmillan, 1938), 4.

  Most Eugenists are Euphemists: G. K. Chesterton, Eugenics and Other Evils (London: Cassell, 1922), 12–13.

  In 1883, one year after Charles Darwin’s death: Francis Galton, Inquiries into Human Faculty and Its Development (London: Macmillan, 1883).

  “We greatly want a brief word to express”: Roswell H. Johnson, “Eugenics and So-Called Eugenics,” American Journal of Sociology 20, no. 1 (July 1914): 98–103, http://www.jstor.org/stable/2762976.

  “at least a neater word . . . than viriculture”: Ibid., 99.

  “Believing, as I do, that human eugenics”: Galton, Inquiries into Human Faculty, 44.

  A child prodigy, Galton: Dean Keith Simonton, Origins of Genius: Darwinian Perspectives on Creativity (New York: Oxford University Press, 1999), 110.

  He tried studying medicine, but then switched: Nicholas W. Gillham, A Life of Sir Francis Galton: From African Exploration to the Birth of Eugenics (New York: Oxford University Press, 2001), 32–33.