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Writing, printing, speaking: Rhesus blood-group genetics and nomenclatures in the mid-twentieth century


In the 1940s and 1950s, British and American journals published a flood of papers by doctors, pathologists, geneticists and anthropologists debating the virtues of two competing nomenclatures used to denote the Rhesus blood groups. Accounts of this prolonged and often bitter episode have tended to focus on the main protagonists' personalities and theoretical commitments. Here I take a different approach and use the literature generated by the dispute to recover the practical and epistemic functions of nomenclatures in genetics. Drawing on recent work that views inscriptions as part of the material culture of science, I use the Rhesus controversy to think about the ways in which geneticists visualized and negotiated their objects of research, and how they communicated and collaborated with workers in other settings. Extending recent studies of relations between different media, I consider the material forms of nomenclatures, as they were jotted in notebooks, printed in journals, scribbled on blackboards and spoken out loud. The competing Rhesus nomenclatures had different virtues as they were expressed in different media and made to embody commitments to laboratory practices. In exploring the varied practical and epistemic qualities of nomenclatures I also suggest a new understanding of the Rhesus controversy itself.

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1 Boyd W.C., ‘Present status of Rh blood types and nomenclature’, American Journal of Human Genetics (1949) 7, pp. 519527, 519.

2 Box Joan Fisher, R.A. Fisher: The Life of a Scientist, New York: John Wiley and Sons, 1978, p. 367; Edwards A.W.F., ‘Unravelling of the Rhesus blood group system’, Genetics (2007) 175, pp. 471476; Bodmer W.F., ‘Early British discoveries in human genetics: contributions of R.A. Fisher and J.B.S. Haldane to the development of blood groups’, American Journal of Human Genetics (1992) 50, pp. 671676; Reid Marion E., ‘Alexander S. Wiener: the man and his work’, Transfusion Medicine Reviews (2008) 22, pp. 300316; Schmidt Paul, ‘Rh–Hr: Alexander Wiener's last campaign’, Transfusion (1994) 34, pp. 180182.

3 Mazumdar Pauline H., Species and Specificity: An Interpretation of the History of Immunology, Cambridge: Cambridge University Press, 1995, pp. 337378.

4 Mazumdar, op. cit. (3), p. 378.

5 Klein Ursula, Experiments, Models, Paper Tools: Cultures of Organic Chemistry in the Nineteenth Century, Stanford: Stanford University Press, 2003; Warwick Andrew, Masters of Theory: Cambridge and the Rise of Mathematical Physics, Chicago: The University of Chicago Press, 2003.

6 See also, for example: Sismondo Sergio, ‘Models, simulations, and their objects’, Science in Context (1999) 12, pp. 247260; Griesemer James R., ‘Three-dimensional models in philosophical practice’, in de Chadarevian Soraya and Hopwood Nick (eds.), Models: The Third Dimension of Science, Stanford: Stanford University Press, 2004, pp. 433442; Christoph Meinel, ‘Molecules and croquet balls’, in de Chadarevian and Hopwood, op. cit., pp. 242–275; for a much earlier discussion of the ways that formulae, symbols and diagrams function in the production of theory see Bachelard Gaston, Le matérialisme rationnel, Paris: Presses universitaires de France, 1953, pp. 112153.

7 For paper-based practices in research on inheritance see, for example, Müller-Wille Staffan, ‘Early Mendelism and the subversion of taxonomy: epistemological obstacles as institutions’, Studies in History and Philosophy of Biological and Biomedical Sciences (2005) 36, pp. 465487.

8 Klein, op. cit. (5). Early geneticists themselves noted the analogous functions of chemical symbols and genetic symbols, for example plant geneticist Edward Murray East quoted in Carlson E.A., The Gene: A Critical History, Philadelphia: W.B. Saunders Co., 1966, p. 29. Elsewhere the behaviour of genes was likened to the combination, dissociation and recombination of atoms in mathematical proportions: Allen Garland E., ‘A century of evo-devo: the dialectics of analysis and synthesis in twentieth-century life science’, in Laubichler Manfred D. and Maienschein Jane (eds.) From Embryology to Evo-Devo, Cambridge, MA: MIT Press, 2007, pp. 123167, 147.

9 Olby Robert, ‘Mendel no Mendelian?’, History of Science (1979) 18, pp. 5372; James R. Griesemer, ‘Tracking organic processes: representations and research styles in classical embryology and genetics’, in Laubichler and Maienschein, op. cit. (8), pp. 375–424.

10 For reflections on early genetic notations see Carlson, op. cit. (8), pp. 23–38; for Morgan's ‘notational crisis’ and its resolution see Falk Raphael and Schwartz Sara, ‘Morgan's hypothesis of the genetic control of development’, Genetics (1993) 134, pp. 671674; Kohler Robert E., Lords of the Fly: Drosophila Genetics and the Experimental Life, Chicago: The University of Chicago Press, 1994, pp. 5390, especially pp. 56–61.

11 For other kinds of representation in genetics see, for example, Kohler, op. cit. (10), pp. 53–90; and the essays in Gaudillière Jean-Paul and Rheinberger Hans-Jörg (eds.), Classical Genetic Research and Its Legacy: The Mapping Cultures of Twentieth-Century Genetics, London and New York: Routledge, 2004.

12 For blood groups and the study of human heredity see Mazumdar Pauline, Eugenics, Human Genetics and Human Failings: The Eugenics Society, Its Source and Its Critics in Britain, London: Routledge, 1992; Mazumdar, ‘Two models for human genetics: blood grouping and psychiatry in Germany between the world wars’, Bulletin of the History of Medicine (1996) 70, pp. 609657. For blood groups in anthropology see Schneider William H., ‘Blood group research in Great Britain, France and the United States between the world wars’, Yearbook of Physical Anthropology (1995) 38, pp. 87114; Lisa Gannett and James R. Griesemer, ‘The ABO blood groups: mapping the history and geography of genes in Homo sapiens’, in Gaudillière and Rheinberger, op. cit. (11), pp. 119–172. For transfusion, forensics and paternity disputes see Schneider William H., ‘Blood transfusion between the wars’, Journal for the History of Medicine (2003) 58, pp. 187224; Okroi M. and Voswinckel P., ‘“Obviously impossible”: the application of the inheritance of blood groups as a forensic method. The beginning of paternity tests in Germany, Europe and the USA’, International Congress Series (2003) 1239, pp. 711714.

13 On diagrams taking on novel meanings in new domains see Kaiser David, ‘Stick-figure realism: conventions, reification, and the persistence of Feynman diagrams, 1948–1964’, Representations (2000) 70, pp. 4986; Hendry Robin Findlay, ‘Mathematics, representation and molecular structure’, in Klein Ursula (ed.), Tools and Modes of Representation in the Laboratory Sciences, Dordrecht: Kluwer Academic Publishers, 2001, pp. 221236. On the relations between different media see de Chadarevian and Hopwood, op. cit. (6); Bernadette Bensaude-Vincent, ‘Graphical representations of the periodic system of chemical elements’, in Klein, op. cit., pp. 133–162.

14 ‘Oral performance … has been and remains at the heart of the making of knowledge’: Secord James A., ‘How scientific conversation became shop talk’, Transactions of the Royal Historical Society (2007) 17, pp. 129–56. For anthropological and sociological analyses of discourse in the laboratory see Lynch Michael, Art and Artifact in Laboratory Science: A Study of Shop Work and Shop Talk in the Research Laboratory, London: Routledge & Kegan Paul, 1985. Latour Bruno and Woolgar Steve, Laboratory Life: The Construction of Scientific Facts, Princeton: Princeton University Press, 1986.

15 For serology as a taxonomic tool in the early twentieth century see Strasser Bruno J., ‘Laboratories, museums, and the comparative perspective: Alan A. Boyden's quest for objectivity in serological taxonomy, 1924–1962’, Historical Studies in the Natural Sciences (2010) 40, pp. 149182.

16 Landsteiner Karl and Wiener Alexander, ‘An agglutinable factor in human blood recognised by immune sera for Rhesus blood’, Proceedings of the Society for Experimental Biology and Medicine (1940) 43, p. 223.

17 Landsteiner and Wiener, ‘Studies on an agglutinogen (Rh) in human blood reacting with anti-Rhesus sera and with human isoantibodies’, Journal of Experimental Medicine (1941) 74, pp. 309320.

18 Levine P. et al. , ‘The role of isoimmunization in the pathogenesis of Erythroblastosis fetalis’, American Journal of Obstetrics and Gynecology (1941) 42, pp. 925937.

19 Another human trait with Mendelian inheritance was the ability to taste phenylthiocarbamide, but this had no medical significance and so the test was not so widely deployed. For literature on the history of blood group genetics see note 12 above.

20 Consistent with other blood-group genetics at the time, they also quickly began studying the correlation between Rhesus frequencies and race types. See Landsteiner Karl, Wiener Alexander S. and Matson G. Albin, ‘Distribution of the Rh factor in American “Indians”’, Journal of Experimental Medicine (1942) 76, pp. 7378.

21 Knight R.L., Dictionary of Genetics, Waltham, MA: Chronica Botanica Company, 1948.

22 Mazumdar recounts this story in rich detail: Mazumdar, Eugenics, Human Genetics and Human Failings, op. cit. (12), pp. 58–145.

23 Fisher saw three possible ways in which the study of blood groups could benefit the study of heritable disorders: (a) the detection of serological alleles that cause human disease (for example, the detection of late-onset disorders early in life), (b) the identification of individuals who carry recessive deleterious alleles, (c) the determination of linkage between human disorders and genes that cause serological reactions. For physicians, establishing such ‘linkage’ would, Fisher felt, improve prognoses for the children of individuals afflicted by hereditary disorders. Fisher to O'Brien, 18 July 1934, 01.0001/401A/Box 16, Rockefeller Archives.

24 Mollison P.L. and Taylor G.L., ‘Wanted: anti-Rh sera’, British Medical Journal (1943) 1(4243), pp. 561562.

25 Fisher R.A., ‘The Rhesus factor: a study in scientific method’, American Scientist (1947) 35, pp. 95103; Gunson Harold H. and Dodsworth Helen, ‘Towards a National Blood Transfusion Service’, Transfusion Medicine (1996) 6 (Suppl.), pp. 416.

26 Although it was Fisher's idea, authorship of their first paper was attributed only to Race. Race R.R., ‘An “incomplete” antibody in human serum’, Nature (1944) 153, pp. 771772; Edwards, op. cit. (2).

27 ‘Arbitrary’ perhaps, but the letters C, D and E were presumably chosen to follow from the A and B of the ABO nomenclature.

28 Cappell D.F., ‘The blood group Rh. Part I: a review of the antigenic structure and serological reactions of the Rh subtypes’, British Medical Journal (1946) 2(4477), pp. 601605.

29 Fisher to Cappell, 29 September 1944, R.A. Fisher Digital Archive, University of Adelaide Library.

30 Haldane J.B.S., ‘Two new allelomorphs for heterostylism in Primula’, American Naturalist (1933) 67, pp. 559560.

31 Although, in various correspondence and papers, authors often referred to the dispute over ‘notations’, ‘nomenclatures’ or ‘terminologies’.

32 Mazumdar, op. cit. (3), pp. 337–378.

33 Wiener, quoted in Mazumdar, op. cit. (3), p. 366.

34 Mazumdar, op. cit. (3), p. 366; Indeed, for Fisher the correspondence between antigen and allele was so direct that for all intents and purposes he saw the antisera as interacting directly with alleles. He explained, for example, that ‘blood group genotypes’ could be ‘recognised by a test fluid’. Fisher, op. cit. (25), p. 1.

35 Mazumdar, op. cit. (3), pp. 305–336 and 357.

36 The New York Academy of Sciences met in 1946 to discuss the problem, in 1947 the European Committee on Biological Standardization created an ‘Expert Subcommittee on Rh antigens’ and in 1948 the World Health Organization held a meeting on the topic in Geneva.

37 For more general remarks on the functions of inscriptions, signs, diagrams and signs in the biology laboratory see Latour Bruno, ‘Drawing things together’, in Lynch Michael and Woolgar Steve (eds.) Representation in Scientific Practice, Cambridge, MA: MIT Press, 1990, pp. 1968; Rheinberger Hans-Jörg, ‘Scrips and scribbles’, Modern Language Notes (2003) 118, pp. 622636. ‘Listmania’ focus section, Isis (2012) 103, pp. 710–752.

38 For some of the rhetorical and epistemological functions of tables, lists and charts see Becker Peter and Clark William (eds.), Little Tools of Knowledge, Ann Arbor: University of Michigan Press, 2001, especially chapters by William Clark, ‘On the ministerial registers of academic visitations’, pp. 95–140; and Hans Erich Bödeker, ‘On the origins of the “statistical gaze”: modes of perception, forms of knowledge, and ways of writing in the early social sciences’, pp. 169–196.

39 Wiener A.S., ‘Theory and nomenclature of the Hr blood factors’, Science (1945) 102(2654), pp. 479482, 479.

40 R.A. Fisher to R.R. Race, 28 January 1948, SA/BGU/C.1, Wellcome Library; Race to Snyder, 3 February 1948, SA/BGU/C.1, Wellcome Library. Mazumdar has suggested that aside from his theoretical commitments, one reason why Wiener was sceptical of the new system was that he did not initially have such a strong version of the ‘St’ (anti-c) antiserum that was most suggestive of the reciprocal relationships. Mazumdar, op. cit. (3), p. 354.

41 For clarity I have given these antisera the original Greek symbols assigned by Fisher and Race, as well as their later names in brackets.

42 The second – δ (anti-d) – was also reported but its existence was never confirmed.

43 Staffan Müller-Wille and Hans-Jörg Rheinberger both drew attention to this on their papers on the epistemological functions of paper and inscriptions in eighteenth- and nineteenth-century natural history and plant breeding. Rheinberger, op. cit. (37); Müller-Wille, op. cit. (7); Staffan Müller-Wille and Sara Scharf, ‘Indexing nature: Carl Linneaus (1707–1778) and his fact-gathering strategies’, Working Papers on the Nature of Evidence: How Well Do ‘Facts’ Travel 36/08 (2009).

44 Daston Lorraine and Galison Peter, ‘The image of objectivity’, Representations (1992) 40, pp. 81128.

45 Klein, op. cit. (13), p. 17, notes the ‘graphic suggestiveness’ of letters denoting chemical elements.

46 Race R.R., Taylor G.L. and Murray J., ‘Serological reactions caused by the rare human gene Rh z’, Nature (1945) 155, pp. 112114.

47 Ducey Edward and Modica Robert, ‘On the amendment of the nomenclature of the Rh-CDE system’, Science (1950) 111, pp. 466467.

48 Mollison P.L., Mourant A.E. and Race R.R., Medical Research Council Memorandum No. 19: The Rh Blood Groups and Their Clinical Effects, London: His Majesty's Stationery Office, 1948, p. 18.

49 Mollison, Mourant and Race, op. cit. (48).

50 Race R.R. and Sanger R., Blood Groups in Man, Oxford: Blackwell Scientific Publications, 1950, p. 113.

51 ‘Genetic distance’ approximates the nucleotide distance between two genes on a chromosome.

52 Castle William B., Wintrobe Maxwell M. and Snyder Laurence H., ‘On the nomenclature of the anti-Rh typing serums: report of the advisory review board’, Science (1948) 107, pp. 2731, 30.

53 Castle, Wintrobe and Snyder, op. cit. (52), p. 30.

54 Mourant's surviving papers provide valuable evidence of this, although I am unable to reproduce the material here because the files have now been closed for patient confidentiality. Two pieces of paper present identical results from Rhesus blood-grouping tests, one written by hand, one using a typewriter. The handwritten sheet uses the Wiener-modified shorthand (with sub- and superscripts), while the typewritten sheet deploys the Fisher–Race nomenclature (upper- and lower-case letters). This is consistent with the ease with which subscripts and superscripts, lower- and upper-case letters could be articulated in these media.

55 Strandskov Herluf H., ‘Blood group nomenclature’, Journal of Heredity (1948) 39, pp. 108112, 112.

56 For a relevant discussion of what can and cannot be written in chalk on a blackboard see Barany Michael J. and MacKenzie Donald, ‘Chalk: materials and concepts in mathematics research’, in Coopmans Catelijne, Lynch Michael, Vertesi Janet and Woolgar Steve (eds.), New Representation in Scientific Practice, Cambridge, MA: MIT Press, forthcoming.

57 Ducey and Modica, op. cit. (47), p. 467.

58 Cyril Jenkins Productions Ltd, Blood Grouping, Imperial Chemical Industries Limited, 1955, Wellcome Library. View the film at

59 Haberman Sol and Hill Joseph M., ‘Verbal usage of the CDE notation for Rh blood groups’, British Medical Journal (1952) 4736, p. 851.

60 Haberman and Hill, op. cit. (59), p. 841.

61 Race and Sanger, op. cit. (50), pp. 172–173.

62 DeGowin Elmer Louis, Blood Transfusion, Philadelphia: W.B. Saunders Co., 1949, p. 83.

63 Cappell, op. cit. (28), p. 604.

64 Coombs R., ‘Detection of weak and “incomplete” Rh agglutinins: a new test’, The Lancet (1945) 246, pp. 1516; Race, ‘A summary of present knowledge of human blood groups, with special reference to serological incompatibility as a cause of congenital disease’, British Medical Bulletin (1946) 4, pp. 188193; Race R.R., Mourant A.E. and Callender Sheila, ‘Rh antigens and antibodies in man’, Nature (1946) 157, p. 410411.

65 Murray John, ‘A nomenclature of subgroups of the Rh factor’, Nature (1944) 154, 701702.

66 Baar H.S., ‘Anti-Rh serum nomenclature’, British Medical Journal (1948) 4562, pp. 11561157, 1157.

67 Castle, Wintrobe and Snyder, op. cit. (52), p. 30.

68 For example, Schmidt, op. cit. (2); Mazumdar, op. cit. (3).

69 Blood types Rh+ and Rh– corresponded to the D and d antigens of the Fisher–Race system. For clinicians wanting to identify people at risk from haemolytic disease, they would simply use Rh0/anti-D antisera – a reaction would define the blood sample as Rh+.

70 Brewer H.F. and Keynes G., Blood Transfusion, Bristol: John Wright, 1949.

71 Walker William, ‘Refresher course for general practitioners’, British Medical Journal (1951) 4740, pp. 11421146.

72 Mollison P.L., Blood Transfusion in Clinical Medicine, Oxford: Blackwell Scientific Publications, 1951, p. 154.

73 Mollison P.L., ‘Blood Groups’, British Medical Journal (1951) 1(4696), p. 75.

74 Race and Sanger, op. cit. (50), pp. 105–106.

75 Memoranda include e.g. Mollison, Mourant and Race, op. cit. (48).

76 Mazumdar, op. cit. (3), pp. 373–374.

77 Mazumdar, op. cit. (3), p. 698.

78 Race R.R. and Sanger R., Blood Groups in Man, 5th edn, Oxford: Oxford University Press, 1968; Mazumdar, op. cit. (3), p. 377.

79 Race and Sanger, op. cit. (78).

80 For example, Geifman-Holtzman O. et al. , ‘Noninvasive fetal RhCE genotyping from maternal blood’, British Journal of Obstetrics and Gynaecology (2008) 116, pp. 144151; Chou S.T. and Westhoff C.M., ‘Molecular biology of the Rh system: clinical considerations for transfusion in sickle cell disease’, Hematology: American Society of Hematology Education Program Book (2009) 1, pp. 178184.

81 International Congress of Human Genetics, Proceedings of the First International Congress of Human Genetics, Copenhagen, August 1–6, 1956 (ed. Kemp Tage, Hauge Mogens and Harvald Bent), Basel: S. Karger, 1957.

82 Strandskov Herluf H., ‘Blood group nomenclature’, Journal of Heredity (1948) 39, pp. 108112, 108.

83 Ford E.B., ‘A uniform notation for the human blood groups’, Heredity (1954) 9, pp. 135142.

84 E.B. Ford to R.R. Race, 16 October 1957, SA/BGU/E.11, Wellcome Library.

85 For other illustrations of what happens to graphical representation when fields are brought into alignment see Emily Grosholz, ‘Federoff's translation of McClintock: the uses of chemistry in the reorganisation of genetics’, in Klein, op. cit. (13), pp. 199–220.

86 ‘Genetic nomenclature guide’, Trends in Genetics (1998) 14 (Suppl. 1), S1–S49. For present-day enforcement of standards see, for example,; community websites include,, and

87 Kotb Malak et al. , ‘Consensus nomenclature for the mammalian methionine adenosyltransferase genes and gene products’, Trends in Genetics (1997) 13, pp. 5152.

88 ‘About the HGNC’, HUGO Gene Nomenclature Committee, n.d.,, my emphasis.

89 Ford to Diamond, 11 February 1955, SA/BGU/E.6, Wellcome Library.

90 Beurton Peter, Falk Raphael and Rheinberger Hans-Jörg, The Concept of the Gene in Development and Evolution: Historical and Epistemological Perspectives, Cambridge: Cambridge University Press, 2000.

91 Singer Maxine F., ‘SINE and LINE nomenclature’, Trends in Genetics (1990) 6, 204.

93 We have rich resources for studying this; for some early examples see East E.M., ‘The Mendelian notation as a description of physiological facts’, American Naturalist 46 (1912), 633695; Castle William B., ‘Simplification of Mendelian formulae’, American Naturalist (1913) 47, pp. 170182; Little C.C., ‘Report of the committee on genetic form and nomenclature’, American Naturalist (1921) 55, pp. 175178; ‘Report of the international committee on genetic symbols and nomenclature’, Union of International Science Biology Series B (1957) Colloquia No 30.

An earlier version of this paper was presented at the British Society for the History of Science Annual Conference 2010 at the University of Aberdeen. I thank Nick Hopwood, Nick Jardine, Boris Jardine, Staffan Müller-Wille, Gregory Radick and A.W.F. Edwards for their insightful comments. Additional thanks are due to Nick Jardine for help with translation.

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