Mendel in America: Theory and Practice, 1900-1919

Copyright © 1988 by the University of Pennsylvania Press.

(This article originally appeared in The American Development of Biology, edited by Ronald Rainger, Keith R. Benson and Jane Maienschein, (University of Pennsylvania Press, 1988), pp. 281-310. It appears at MendelWeb, for non-commercial educational use only, with the kind permission of the authors and the University of Pennsylvania Press. The entire volume has been reprinted by, and is currently available from, Rutgers University Press ((800) 446-9323). Although you are welcome to download this text, please do not reproduce it without the permission of the authors and the University of Pennsylvania Press.)

"Mendel in America: Theory and Practice, 1900-1919"

Notes:

1 Willet M. Hays, "Address by Chairman of Organization Committee," Proceedings of the American Breeders Association, 1905, 1: 9-15.

2 William Bateson, "Hybridisation and Cross-breeding as a Method of Scientific Investigation," Journal of the Royal Horticultural Society, 1900, 24: 59-66.

3 Charles Davenport, "Report of the Committee on Eugenics," The American Breeders Magazine, 1910, 1: 126-129. See also Barbara A. Kimmelman, "The American Breeders Association: Genetics and Eugenics in an Agricultural Context," Social Studies of Science, 1983, 13: 163-204.

4 Deborah K. Fitzgerald, "The Business of Breeding: Public and Private Development of Hybrid Corn in Illinois, 1890-1940" (Ph.D. dissertations, University of Pennsylvania, 1985).

5 During this period, investigators generally used the terms hybridization, crossbreeding, and cross-fertlization to mean, respectively, interspecific crosses, intervarietal crosses, and intraspecific crosses between male and female of different plants. Darwin based his crucial analogy between interspecific hybrids and intervarietal mongrels on such distinctions. In practice, the analogy proved more powerful than the distinction, and many investigators used the terms interchangeably. We will see that, after 1910, hybrids came to acquire a more precise, technical meaning.

6 Walter A. Cannon, "Review of Proceedings: International Conference on Plant Breeding and Hybridization," Torreya, 1905, 1: 12-14.

7 C. W. Ward, "Improvement of Carnations," Memoirs of the Horticultural Society of New York, 1904, 1: 151-155.

8 William Bateson, "Practical Aspects of the New Discoveries in Heredity," Mems. Hort. Soc. N.Y., 1904, 1: 1-8.

9 William Bateson, Mendel's Principles of Heredity: A Defence (Cambridge: Cambridge University Press; New York: MacMillan Co.; 1902) Alan Cock notes that the book was published on 3 or 4 June 1902 in Britain and must have appeared almost simultaneously in the United States (private communication).

10 Liberty Hyde Bailey, "Comments," Mems. Hort. Soc. N.Y., 1904, 1: 8.

11 William Bateson to Beatrice Bateson, 3 October 1902, William Bateson papers, University of Cambridge Library, letter G-3D-05; typed transcript at G8G01D. We gratefully acknowledge the Library's permission to quote the Bateson letter. The originals of the collection prefaced G have recently been transferred to the Library from the John Innes Institute, which now has photocopies. Mrs. Rosemary Harvey, archivist at the John Innes, kindly supplied copies of Bateson's 1902 letters from America.

12 Garland Allen, Life Science in the Twentieth Century (New York: John Wiley and Sons, 1975), p. 52.

13 Ibid.

14 "Seedsmen," or "practical breeders," were primarily farmers who also bred particular varieties of seed/grain for purpose of sale. A few of these farmers (such as the Funks) engaged in breeding for sale as a major enterprise but for most it was a minor activity. Some seedsmen (mostly urban horticulturalists) were businessmen simpliciter.

15 Barbara A. Kimmelman examines the founding and early years of research departments in genetics within the agricultural colleges at Ithaca, N.Y., Madison, Wis., and Berkeley, Calif., in "A Progressive Era Discipline: Genetics at American Agricultural Colleges and Experiment Stations, 1890-1920," (Ph.D. dissertations, University of Pennsylvania, 1987).

16 Bateson, "Practical Aspects," p. 3.

17 C. C. Hurst, "Notes on Mendel's Methods of Cross Breeding," Mems. Hort. Soc. N.Y., 1904, 1: 11-16. Hugo de Vries, "On Artificial Atavism," Mems. Hort. Soc. N.Y., 1904, 1: 16-23.

18 This point is illustrated by the diverse character of articles on Mendelism published in American journals between 1901 and 1903. The first to appear was Charles Davenport's "Mendel's Laws of Dichotomy," in the Biological Bulletin, 1901, 2: 307-310. It was quickly followed by E. B. Wilson, "Mendel's Principles of Heredity and the Maturation of the Germ-cells," Science, 1902, 16: 991-992; Walter Sutton, "On the Morphology of the Chromosome Group in Brachystola magna," Biol. Bull., 1902, 3: 24-39; W. J. Spillman, "Exceptions to Mendel's Law," Sci., 1902, 16: 709-710 and 784-796; R. A. Emerson, "Preliminary Account of Variation in Bean Hybrids," 15th Annual Report of the Nebraska Experiment Station, 1902; and Walter A. Cannon, "A Cytological Basis for Mendelian Cases," Bulletin of the Torrey Botanical Club, 1902. Other early accounts include Liberty Hyde Bailey, "A Discussion of Mendel's Law and its Bearings," Address before the Society for Plant Morphology and Physiology, Washington, D. C., 29 Dec. 1902, published as "Some Recent Ideas on the Evolution of Plants," Sci., 1903, 17: 441-454; Walter Sutton, "The Chromosomes in Heredity," Biol. Bull., 1902, 4: 231-251; and William Castle, "The Laws of Heredity of Galton and Mendel and some Laws Governing Race Improvement by Selection," Proceedings of the American Academy of Arts and Sciences, 1903, 38: 535-548; reprinted as "Mendel's Law of Heredity," in Sci., 1903, 18: 396-406. Compare with the lack of interest expressed by the Botanical Gazette. The first mention of Mendel is a dismissive comment by the editor, John Merle Coulter, in a review of the third edition of Liberty Hyde Bailey's Plant Breeding (Botanical Gazette, 1904, 37: 471-472). The American Naturalist was also unimpressed. Other than a passing reference in a Botanical Note of 1902, there is no mention of Mendelism until 1904, and then only in Charles Davenport's book reviews. Editorial notes and articles first appear in 1907.

19 Jan Sapp, "The Struggle for Authority in the Field of Heredity, 1900-1932: New Perspectives on the Rise of Genetics," Journal of the History of Biology, 1983, 16: 311-342.

20 Willet M. Hays, "Address by Chairman of Organization Committee," p. 10.

21 W. J. Spillman, "Mendel's Law in Relation to Animal Breeding," Proc. ABA, 1905, 1: 171-176; H. J. Webber, "Explanation of Mendel's Law of Hybrids," Proc. ABA, 1905, 1: 138-143.

22 William Bateson, "G. Mendel: Experiments in Plant Hybridisation," J. Royal Hort. Soc., 1901, 26: 1-32.

23 U.S. Department of Agriculture, Office of Experiment Stations, Abstract of G. Mendel, "Experiments in Plant Hybridization," Experiment Station Record, 1901-1902, vol. 13 (Washington, D. C. : Government Printing Office, 1902), p. 744.

24 Ibid., p. 745.

25 Ibid., p. 1004.

26 See, for example, U. S. Department of Agriculture, Office of Experiment Stations, Abstracts of W. F. R. Weldon, "Mendel's Law of Alternative Inheritance in Peas," C. C. Hurst, "Mendel's Law Applied to Orchid Hybrids," and Carl Correns, "Apparent Exceptions to Mendel's Law of Dissociation in Hybrids," Experiment Station Record, 1902-1903, vol. 14 (Washington, D. C. : Government Printing Office, 1903), pp. 466-467, 569. Also see U. S. Department of Agriculture, Office of Experiment Stations, Abstracts of A. D. Darbishire, "On the Bearing of Mendelian Principles of Heredity on Current Theories of the Origin of Species," Erich von Tschermak, "Further Studies in Crossing Peas, Stocks and Beans," and David Starr Jordan, "Some Experiments of Luther Burbank," Experiment Station Record, 1904-1905, vol. 16 (Washington, D. C. : Government Printing Office, 1905), pp. 232, 263, 773-774. Jordan's study indicated that Burbank's results did not generally conform to Mendel's laws.

27 Bailey, "A Discussion of Mendel's Law," pp. 445-446.

28 Margaret Rossiter, "The Organization of the Agricultural Sciences," in Alexandra Oleson and John Voss, eds., The Organization of Knowledge in Modern America, 1860-1920 (Baltimore: Johns Hopkins University Press, 1979), pp. 211-248.

29 Our interpretation of the Progressive Era accords with the work of Sidney Fine, Laissez Faire and the General Welfare State: A Study of Conflict in American Thought 1865-1901 (Ann Arbor: University of Michigan Press, 1956); Samuel P. Hays, The Response to Industrialism 1885-1914 (Chicago: University of Chicago Press, 1947); Harold U. Faulkner, Politics, Reform, and Expansion 1890-1900 (New York: Harper and Row, 1959); Robert H. Weibe, The Search for Order 1877-1920 (New York: Hill and Wang, 1967); and James Weinstein, The Corporate Ideal in the Liberal State 1890-1918 (Boston: Beacon Press, 1968). See also Samuel P. Hays, "Introduction: The New Organizational Society," in Jerry Israel, ed. Building the Organizational Society (New York: The Free Press, 1972), pp. 1-15.

30 U. S. Department of Agriculture, Report to the Commissioner of Agriculture for the year 1885 "Report of the Statistician," by J. R. Dodge (Washington, D. C.: Government Printing Office, 1885), p. 372.

31 For example, see U. S. Department of Agriculture, Report to the Commissioner of Agriculture for the year 1880, "Report of the Statistician," by Charles Worthington (Washington, D. C.: Government Printing Office, 1880), p. 210; Dodge, "Report of the Statistician," pp. 372-376; and U. S. Department of Agriculture, First Report of the Secretary of Agriculture, 1889, "Report of the Secretary of Agriculture," by J. M. Rusk (Washington, D. C.: Government Printing Office, 1889), p. 14.

32 Dodge, "Report of the Statistician," pp. 379-380.

33 Ibid., p. 373.

34 A valuable discussion of the growing importance of U. S. international markets (although, like its subjects, it disparages the significance of the agricultural sector) is Walter LeFeber, The New Empire: An Interpretation of American Expansion 1860-1898 (Ithaca: Cornell University Press, 1967). Works addressing this theme in agriculture include Fred A. Shannon, The Farmer's Last Frontier: Agriculture, 1860-1897 (New York: Farrar and Rinehart, 1945); Alan I. Marcus, Agricultural Science and the Quest for Legitimacy: Farmers, Agricultural Colleges, and Experiment Stations, 1870-1890 (Ames: Iowa State University Press, 1985); and David Danbom, The Resisted Revolution: Urban America and the Industrialization of Agriculture, 1900-1930 (Ames: Iowa State University Press, 1979), although it deals with a slightly later period.

35 U. S. Department of Agriculture, Report to the Commissioner of Agriculture for the year 1887, "Report of the Commissioner of Agriculture," by Norman J. Colman (Washington D. C. : Government Printing Office, 1888), p. 8.

36 U. S. Department of Agriculture, Report to the Commissioner of Agriculture for the year 1888, "Report of the Statistician," by J. R. Dodge (Washington D. C. : Government Printing Office, 1889), p. 201.

37 Ibid., pp. 201-202. See also U. S. Department of Agriculture, Report to the Commissioner of Agriculture for the year 1891, "Report of the Statistician," by J. R. Dodge (Washington D. C. : Government Printing Office, 1892), pp. 301-306.

38 U. S. Department of Agriculture, Report to the Commissioner of Agriculture for the year 1892, "Report of the Secretary of Agriculture," by J. M. Rusk (Washington D. C. : Government Printing Office, 1893), p. 10; also U. S. Department of Agriculture, Report to the Commissioner of Agriculture for the year 1893, "Report of the Secretary of Agriculture," by J. Sterling Morton (Washington D. C. : Government Printing Office), p. 16.

39 The agricultural appropriation act of 1881 statutorily established a divisional organization for the USDA, at which time the divisions of Seed, Gardens and Grounds and of Botany were founded. The Division of Pomology was established in 1886; the Division of Vegetable Pathology achieved independence from the Division of Botany in 1891, and became the Division of Vegetable Physiology and Pathology in 1895; in that year, the Division of Agrostology was also founded. The Section Foreign Seed and Plant Introduction was established in 1897. See U. S. Department of Agriculture, Division of Publications, Historical Sketch of the U. S. Department of Agriculture: Its Objects and Present Organization, by Charles H. Greathouse, Bulletin no. 3 (Washington D. C. : Government Printing Office, 1898), pp. 33-40; Fred Wilbur Powell, The Bureau of Plant Industry, its History, Activities and Organizations, Institute for Government Research, Service Monographs of the United States Government, no. 47 (Baltimore: Johns Hopkins University Press, 1927), pp. 1-9; and David Fairchild, The World Was My Garden, Travels of a Plant Explorer (New York: Charles Scribner's Sons, 1939), pp. 105-107.

40 U. S. Department of Agriculture, Report to the Commissioner of Agriculture for the year 1885, "Report of the Chief of the Seed Division," by William M. King (Washington D. C. : Government Printing Office, 1885), p. 47.

41 Ibid., p. 51.

42 Ibid., p. 53.

43 U. S. Department of Agriculture, Report to the Commissioner of Agriculture for the year 1886, "Report of the Pomologist," by H. E. Van Deman (Washington D. C. : Government Printing Office, 1886), p. 260.

44 U. S. Department of Agriculture, Report to the Commissioner of Agriculture for the year 1887, "Report of the Pomologist," by H. E. Van Deman (Washington D. C. : Government Printing Office, 1888), pp. 627-628.

45 U. S. Department of Agriculture, Report to the Commissioner of Agriculture for the year 1892," "Report of the Superintendent of Gardens and Grounds," by William Saunders (Washington D. C. : Government Printing Office, 1893).

46 U. S. Department of Agriculture, Report to the Commissioner of Agriculture for the year 1892," "Report of the Chief of Vegetable Pathology," by B. T. Galloway (Washington D. C. : Government Printing Office, 1893), p. 246.

47 B. T. Galloway, "Division of Vegetable Physiology and Pathology," Yearbook U.S.D.A. 1897 (Washington D. C. : Government Printing Office, 1898), p. 104.

48 Ibid., p. 106.

49 Albert F. Woods, "Work in Vegetable Physiology and Pathology," Yearbook U.S.D.A. 1898 (Washington D. C. : Government Printing Office, 1899), pp. 264-266. See also U. S. Department of Agriculture, Report to the Commissioner of Agriculture for the year 1899," "Report of the Secretary of Agriculture," by James Wilson (Washington D. C. : Government Printing Office, 1899), p. 15.

50 As illustrative of the division's work during this period and of the intellectual problems addressed by its researchers, see Walter T. Swingle and Herbert J. Webber, "Hybrids and Their Utilization in Plant Breeding," Yearbook U.S.D.A. 1897, pp. 383-421.

51 For example, see U. S. Department of Agriculture, Report to the Commissioner of Agriculture for the year 1889," "Report of the Botanist," by George Vasey (Washington D. C. : Government Printing Office, 1889), pp. 377-381; and U. S. Department of Agriculture, Report to the Commissioner of Agriculture for the year 1890, "Special Report of the Assistant Secretary: The Scientific Work of the Department in its Relations to Practical Agriculture," by Edwin Willits (Washington D. C. : Government Printing Office, 1890), pp. 59-73.

52 Rossiter, "The Organization of the Agricultural Sciences," pp. 213-215.

53 U. S. Department of Agriculture, Report to the Commissioner of Agriculture for the year 1898," "Report of the Secretary of Agriculture," by James Wilson (Washington D. C. : Government Printing Office, 1898), p. 47.

54 U. S. Department of Agriculture, Office of Experiment Stations, Organization of the Agricultural Experiment Stations of the United States, Norman J. Colman, Bulletin no. 1 (Washington D. C. : Government Printing Office, 1884).

55 See Charles E. Rosenberg, No Other Gods: On Science and American Social Thought (Baltimore: Johns Hopkins University Press, 1976), pp. 174-175.

56 This group included William James Beal, Charles E. Bessey, and Liberty Hyde Bailey. On their efforts, see Andrew Denny Rodgers, John Merle Coulter, Missionary in Science (Princeton: Princeton University Press, 1944), esp. pp. 52-64, and idem, Liberty Hyde Bailey, A Story of American Plant Sciences (Princeton: Princeton University Press, 1949), esp. pp. 22-181, 242-247. See also Richard Overfield, "Charles E. Bessey: The Impact of the 'New Botany' in American Agriculture, 1880-1910," Technology and Culture, 1975, 16: 162-181.

57 See U. S. Department of Agriculture, Office of Experiment Stations, Digest of the Annual Reports of the Agricultural Experiment Stations in the United States for 1888, Bulletin no. 2, pt. 1 (Washington D. C. : Government Printing Office, 1884). For a discussion of constraints on scientific work at the stations during this period, see Rosenberg, "Science, Technology, and Economic Growth: The Case of the Agricultural Experiment Station Scientist, 1875-1914," in Rosenberg, No Other Gods, pp. 153-172.

58 The stations were South Dakota, Maryland, Massachusetts, and Mississippi. See U. S. Department of Agriculture, Office of Experiment Stations, List of Horticulturalists of the Agricultural Experiment Stations in the United States, with an Outline of Work in Horticulture at the Several Stations, by W. B. Alwood, Bulletin no. 4 (Washington D. C. : Government Printing Office, 1889).

59 Ibid. The stations were Colorado, Florida, Kansas, Michigan, New York (Geneva), Missouri, New Jersey and Ohio.

60 U. S. Department of Agriculture, Office of Experiment Stations, List of Botanists of the Agricultural Experiment Stations in the United States, with an Outline of the Work in Botany at the Several Stations, Bulletin no. 6 (Washington D. C. : Government Printing Office, 1890), p. 6.

61 Ibid., pp. 7-23, for responses from the stations in alphabetical order by state. The states with strong programs planned in crossbreeding were Illinois, Indiana, Kansas, Michigan, New Jersey and Ohio.

62 See H. J. Webber and E. A. Bessey, "The Progress of Plant Breeding in the United States," Yearbook U.S.D.A. 1989 (Washington D. C. : Government Printing Office, 1899), esp. pp. 478, 480-481, 487-488.

63 Office of Experiment Stations, List of Botanists, p. 6.

64 The Illinois and Kansas stations' corn work is summarized in the U.S. Department of Agriculture, Office of Experiment Stations, Handbook of Experimental Station Work: A Popular Digest of the Agricultural Experiment Stations in the United States, Bulletin no. 15 (Washington D. C. : Government Printing Office, 1893), pp. 81-89 (entry under "Corn, crossing").

65 William Bateson, Materials for the Study of Variation (London: 1894); the culmination of de Vries'' late-nineteenth-century work was his Die Mutationstheorie, vol. 1 (Leipzig: 1901) and vol. 2 (Leipzig: 1903).

66 Wilfred Mark Webb, "The International Conference on Hybridisation and Cross-breeding," Nature, 1899, 60:305-307. Others invited but not present (in addition to Bailey) were David Fairchild of the USDA, Luther Burbank, and J. M. MacFarlane of the University of Pennsylvania.

67 William Bateson, "Hybridisation and Cross-breeding a Method of Scientific Investigation."

68 Hugo de Vries, "Hybridisation as a Mean of Pangenic Infection," J. Roy. Hort. Soc., 1900, 24: 69-75.

69 C. C. Hurst, "Experiments in Hybridisation and Cross-breeding," J. Roy. Hort. Soc., 1900, 24: 90-127. "Prepotency" suggests a sire's ability to impress his traits on his progeny. It generally referred to an overall "type," rather than single characters.

70 Ibid., p. 90.

71 Herbert J. Webber, "Work of the United States Department of Agriculture in Plant Hybridisation," J. Roy. Hort. Soc. , 1900, 24: 128-145; see also L. H. Bailey, "Progress of Hybridisation in the United States of America," J. Roy. Hort. Soc. , 1900, 24: 209-213.

72 Willet M. Hays, "Breeding Staple Food Plants," J. Roy. Hort. Soc. , 1900, 24: 257-265.

73 See Webb, "International Conference on Hybridisation and Cross-breeding," p. 307.

74 Office of Experiment Stations, Experiment Station Record (1902), p. 205.

75 W. M. Hays, "Address by Chairman of Organization Committee."

76 W. J. Spillman, Mems. N.Y. Hort. Soc., 1904, 1: 155.

77 Bailey, "A Discussion of Mendel's Law," p. 450.

78 Bateson, "Practical Aspects," p. 2.

79 Bailey, "A Discussion of Mendel's Law," p. 450.

80 L. C. Dunn, A Short History of Genetics (New York: McGraw-Hill, 1965), p. 125. The following list provides a few representation comments: Paul Mangelsdorf writes that the increased yields resulting from hybrids "contributed not only to this country's war effort but also the rehabilitation of Europe after the war," in "George Harrison Shull," Genetics, 1955, 60: 2-3. L. J. Stadler claims that "It is ... no exaggeration to say, speaking in terms of the overall national economy, that the dividend on our research investment in hybrid corn, during the war years alone, was enough to pay the money cost of the development of the atomic bomb, " quoted in G. Shull, "Hybrid Seed Corn," Sci., 1946, 103: 547-550. Bentley Glass states that the increased in yield was "enough to pay for the Manhattan Project" and even "most significantly, increased production [which] permitted the United States to ship vast quantities of food abroad after the war, thus preventing famine and pestilence," in "Shull, George Harrison," Dictionary of American Biography, suppl. 5 (New York: Scribners, 1977), p. 629. Richard Crabb writes that hybrid corn helped "tip the scales in our favor in a war to the finish," in Richard Crabb, The Hybrid-Corn Makers: Prophets of Plenty (New Brunswick, N. J.: Rutgers University Press, 1948), p. 13.

81 Jean-Pierre Berlan and Richard Lewontin, "The Political Economy of Hybrid Corn," Monthly Review, 1986, 38: 35-47; and Berlan and Lewontin, "Breeders' Rights and Patenting Life Forms," Nat., 1986, 322, 785-788, esp. pp. 787-788.

82 They should do better, rather than equally well, because of the ubiquity of partial dominance (discussed below).

83 G. H. Shull, "Beginnings of the Heterosis Concept," in John Gowan, ed., Heterosis (Ames: Iowa State College Press, 1952), pp. 14-48.

84 Shull, "A Pure-line Method of Corn Breeding," Proc. ABA, 1909, 5: 51-59.

85 Edward M. East and Donald F. Jones, Inbreeding and Outbreeding: Their Genetic and Sociological Significance (Philadelphia: J. B. Lippincott, 1919), p. 168.

86 Shull, "A Pure-line Method of Corn Breeding," p. 52.

87 Charles Darwin, The Effects of Cross and Self-fertilization in the Vegetable Kingdom (London: John Murray, 1876).

88 E. M. East, "The Distinction between Development and Heredity in Inbreeding," Am. Nat., 1909, 43: 173-181.

89 Ibid., p. 177.

90 John Farley, Gametes and Spores (Baltimore: Johns Hopkins University Press, 1982), pp. 203-208.

91 G. H. Shull, "Duplicate Genes for Capsule Form in Bursa bursa-pastoris," Zeitschrift für Induktive Abstammungs- und Vererbungslehre, 1914, 12: 97-149, on p. 127.

92 Ibid., p. 126; also quoted in G. H. Shull, "What is Heterosis?" Genetics, 1948, 33: 439-446, on p. 440.

93 G. H. Shull, "The Composition of a Field of Maize," Proc. ABA, 1908, 4: 296-301, on p. 301.

94 Shull, "Hybrid Seed Corn," p. 549.

95 Conway Zirkle, "Early Ideas of Inbreeding and Crossbreeding," in Gowan, Heterosis, pp. 1-13.

96 See Frederick D. Richey, "Hybrid Vigor and Corn Breeding," Journal of the American Society of Agronomists, 1946, 38: 833-841, and G. F. Sprague, "Heterosis in Maize: Theory and Practice," in R. Frankel, ed., Heterosis: Reappraisal of Theory and Practice (New York: Springer-Verlag, 1982), p. 50.

97 E. M. East, "Heterosis," Genet., 1936, 21: 375-397, on p. 375.

98 E. M. East and D. F. Jones, Inbreeding and Outbreeding.

99 The inbred plants used to produce hybrids were so depressed that they generated little (hence expensive) seed. In the double-cross, the plants that produce the seed are themselves hybrid.

100 "Homozygosity, when obtained with the combination of all the most favorable characters, is the most effective condition for the purpose of growth and reproductions," East and Jones, Inbreeding and Outbreeding, p. 187.

101 Fred Hull, "Recurrent Selection for Specific Combining Ability in Corn," J. Am. Soc. Agron., 1945, 37: 134-145.

102 J. L. Lush, Animal Breeding Plans, 3rd ed. (Ames, Iowa: The Collegiate Press, 1945). Lush's influence is noted in James F. Crow, "Muller, Dobzhansky, and Overdominance," J. Hist. Biol., 1987, 20: 351-380.

103 By 1945, 99.9 percent of corn acreage in Iowa and 98.1 percent in Illinois and Indiana had been planted in hybrids. USDA, Agricultural Statistics, 1947 (Washington, D. C. : Government Printing Office, 1949), table 48, p. 43.

104 East and Jones, Inbreeding and Outbreeding, pp. 177-178.

105 Such a response occurs when there are large numbers of already existing inbred lines, maximizing the chance of finding a favorable cross.

106 East and Jones, Inbreeding and Outbreeding, p. 182.

107 Ibid., p. 182.

108 Ibid., p. 169, 181. Jones characterizes hybrids as a "makeshift measure" in "Selection in Self-Fertilized Lines as the Basis of Corn Improvement," J. Am. Soc. Agron., 1920, 12: 77-100, on p. 95.

109 East and Jones, Inbreeding and Outbreeding, p. 224.

110 George S. Carter to Henry A. Wallace, 12 May 1925, quoted in Jack Kloppenburg, First the Seed: The Political Economy of Plant Biotechnology (Cambridge: Cambridge University Press, 1988). The Wallace papers are deposited at the University of Iowa Library and are available on microfilm.

111 Compare Jones, "Selection in Self-fertilized Lines," p. 87, and Willet Hays, "Distributing Valuable New Varieties and Breeds," Proc. ABA, 1905, 1: 58-65.

112 See Fitzgerald, "The Business of Breeding," esp. chap. 5 and conclusion.

113 See Kloppenburg, First the Seed, and Fitzgerald, "The Business of Breeding."

(This article originally appeared in The American Development of Biology, edited by Ronald Rainger, Keith R. Benson and Jane Maienschein, (University of Pennsylvania Press, 1988), pp. 281-310. It appears at MendelWeb, for non-commercial educational use only, with the kind permission of the authors and the University of Pennsylvania Press. The entire volume has been reprinted by, and is currently available from, Rutgers University Press ((800) 446-9323). Although you are welcome to download this text, please do not reproduce it without the permission of the authors and the University of Pennsylvania Press.)