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Animal breeding in the age of biotechnology: the investigative pathway behind the cloning of Dolly the sheep

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Abstract

This paper addresses the 1996 cloning of Dolly the sheep, locating it within a long-standing tradition of animal breeding research in Edinburgh. Far from being an end in itself, the cell-nuclear transfer experiment from which Dolly was born should be seen as a step in an investigative pathway that sought the production of medically relevant transgenic animals. By historicising Dolly, I illustrate how the birth of this sheep captures a dramatic redefinition of the life sciences, when in the 1970s and 1980s the rise of neo-liberal governments and the emergence of the biotechnology market pushed research institutions to show tangible applications of their work. Through this broader interpretative framework, the Dolly story emerges as a case study of the deep transformations of agricultural experimentation during the last third of the twentieth century. The reorganisation of laboratory practice, human resources and institutional settings required by the production of transgenic animals had unanticipated consequences. One of these unanticipated effects was that the boundaries between animal and human health became blurred. As a result of this, new professional spaces emerged and the identity of Dolly the sheep was reconfigured, from an instrument for livestock improvement in the farm to a more universal symbol of the new cloning age.

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Notes

  1. Apart from popular and autobiographic literature (Wilmut et al. 2000; Kolata 2011), the most comprehensive scholarly investigation of Dolly is a monograph written by sociologist Sarah Franklin. In it, she explores the long-standing commercial exchange of sheep across Britain and its former Empire, as the basis of scientific research around this animal which led ultimately to the cloning experiments (Franklin 2007b). Other academics have addressed the public and regulatory debates around Dolly (Suk et al. 2007), as well as the models of biomedical innovation in which the cloning experiments were conducted (Fransman 2001; Clay and Goldberg 1997).

  2. http://news.bbc.co.uk/1/hi/scotland/2930537.stm (last accessed June 2015).

  3. http://www.archives.lib.ed.ac.uk/towardsdolly/ (last accessed June 2015).

  4. Agricultural historian E. J. Russell has documented a long tradition of trade, exchange and mating of different domestic and farm animals that can be traced back, at least to the Early Modern period. As Sarah Franklin has shown, from the late eighteenth century onwards, English sheep were introduced into Australia and played an important role in the wool trade, setting up international connections that later became visible in the work at the Roslin Institute (Franklin 2007b, Chaps. 3–4; Russell 1966). For other studies of the political and economic use of agriculture in France, Germany, Italy, the US, Mexico, Portugal and Spain see Harwood (2005), Matchett (2006), Bonneuil and Thomas (2009), Saraiva (2010), Camprubí (2010), Santesmases (2013), von Schwerin (2013) and Barahona et al. (2005).

  5. The lectureship was awarded to the then promising geneticist A.D. Darbishire, who was killed in World War One (Ankeny 2000, p. 339ff). The role of head of the Department—and later director of the Institute of Animal Genetics—fell therefore to F.A.E. Crew, a charismatic but inexperienced researcher at that time, who became a widely consulted expert in the field of animal genetics (Hogben 1974). See also http://libraryblogs.is.ed.ac.uk/towardsdolly/2013/11/11/remembering-arthur-dukinfield-darbishire-1879-1916/ (last accessed June 2015).

  6. ABRO was earlier known as NABGBRO (National Animal Breeding and Genetics Research Organisation) and was located in London. It moved to Edinburgh when Waddington was appointed in 1947. Another emerging site of animal agricultural science after World War Two was Cambridge, where the ARC established research centres in animal reproduction and physiology which would be merged with the PRC and ABRO in the 1980s (Wilmot 2007, p. 425ff; Polge 2007; see below). See also Bulfield (1999) “Eighty years ago…”, Roslin Institute Annual Report, 1998–1999, pp. 24–25. Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/5/1. On the development of breeding science after World War Two and rise of pig as a research object see Woods (2012), Brassley (2007).

  7. This autonomy and distance from basic genetics mirrors the gap between scientific theory and practice that other historians have shown in early twentieth-century agriculture (Wieland 2006; Theunissen 2008).

  8. “Current research projects”, ABRO Report—January 1970, pp. 45–48. Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/1/1. The academic literature on the early work of ABRO and the PRC is surprisingly thin. However, the Towards Dolly project is developing a blog, curated by the archivist Clare Button, with substantial information on these two institutions, as well as the Institute of Animal Genetics and Ewart’s early work, see http://libraryblogs.is.ed.ac.uk/towardsdolly/ (last accessed June 2015).

  9. H.P. Donald: “New tasks for livestock geneticists”, ABRO Report—January 1974, quotes from pp. 7–8. Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/1/1.

  10. A similar debate developed in the US during the mid-to-late 1970s, with some scientists, commentators and policy-makers suggesting that State-funded laboratories should only produce new animal or plant varieties at the request of industrial actors (Hightower 1978; Kloppenburg 2005; Clarke 2007, p. 320ff).

  11. This perception of industrial underperformance in the UK was to a large extent fuelled by prior examples of alleged British failure to patent biological discoveries that were then commercially exploited in the United States, such as penicillin and monoclonal antibodies—the latter developed in 1975. Historians have critically addressed this notion of a failure (de Chadarevian 2011; Liebenau 1987) and placed it within a long history of perceived decline of the British industrial potential when compared to the US (Edgerton 1996).

  12. “Comment”, ABRO Annual Report—January 1983, p. 3. Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/1/1.

  13. “Preface”, ABRO Annual Report—January 1986, p. 1. Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/1/1. The IAPGR’s Cambridge Research Station was based in Babraham Hall, a nineteenth-century estate surrounded by vast amounts of land that had been bought by the ARC in 1948, as part of the post-World War Two establishment of breeding research centres (see above). Since then, it had housed the Institute of Animal Physiology, an institution with substantial expertise in biochemistry, animal health and reproductive biology. See “Origins”, IAPGR—Report for 1988–89, p. 8. Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/4/1.

  14. “Comment”, ABRO Annual Report—January 1984, p. 3. Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/1/1.

  15. Peter Harper’s interview with Nick Hastie, Cardiff University’s Genetics and Medicine Historical Network, 26th May 2004, transcript available at http://www.genmedhist.info/interviews/Hastie (last accessed June 2015).

  16. R. Lathe, M.P. Kieny, R. Drillien, J.P. Lecocq: “Vaccin contre la rage et proceed pour sa preparation”, European patent number EP0162757B1, filed in 1985 and awarded in 1990, available at http://www.freepatentsonline.com/EP0162757.pdf, last accessed June 2015.

  17. “Comment”, ABRO Annual Report—January 1985, p. 4, both from Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/1/1.

  18. Research programme number 10: “Investigations of genome engineering using the techniques of molecular biology and its potential in animal breeding”, ABRO Annual Report—January 1984, p. 42. Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/1/1. A scheme of the technique that Lathe, Clark and Wilmut developed to produce transgenic sheep, as described in ABRO’s 1986 report, can be found at http://libraryblogs.is.ed.ac.uk/towardsdolly/files/2012/06/immunogeneticsabroreport1986v3.jpg (last accessed June 2015).

  19. I. Wilmut, D.I. Sales, C. Manson, G. Newell: “Non-surgical transfer of cattle embryos: a field trial”, ABRO Annual Report—January 1978, p. 41 and I. Wilmut, D.I. Sales: “Does hormonal imbalance cause death of embryos in sheep?”, ABRO Annual Report—January 1982, pp. 25–30. Both from Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/1/1. As Franklin has shown, another offshoot of these embryo transfer techniques was animal and human IVF, which was substantially developed in Australia during the early 1980 s by researchers who had also worked at the Cambridge Animal Research Station (Franklin 2007a).

  20. “Research programme”, ABRO Annual Report—January 1984, pp. 40 and 42. Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/1/1.

  21. Biopharming referred to the use of plants and animals as living factories to produce substances of commercial utility. The field was viewed with interest at the time by the scientific and industrial communities (Milne 2012).

  22. R. Lathe: “Molecular tailoring of the farm animal germlines”, ABRO Annual Report—January 1985, pp. 7–10; J. O. Bishop, A. L. Archibald, A. J. Clark, R. F. Lathe, J.P. Simons, I. Wilmut: “Germline manipulation of livestock”, ABRO Annual Report—January 1986, pp. 22–26. Both from Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/1/1. The animosities that resulted from the merger (see above) meant that the Edinburgh researchers initiated little interaction with the Institute of Animal Physiology, despite the extensive expertise in recombinant DNA and reproductive biology available there. Wilmut’s former institution, the Animal Research Station, had become part of the Institute of Animal Physiology shortly before his move to Edinburgh. Following the merger, both institutions were housed in Babraham—the base of IAPGR’s Cambridge Research Station. See Polge (2007) and B. A. Cross: “Foreword”, IAPGR—Edinburgh Research Station Report for 1986–87, p. I. Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/4/1.

  23. M. McClenaghan, A. L. Archibald, J. P. Simons, C. B. A. Whitelaw, I. Wilmut, A. J. Clark: “The mammary gland as an organ for protein production”, IAPGR Report for 1988–89, pp. 74–75; “Pharmaceutical Proteins”, Roslin Institute Annual Report 1993–94, pp. 16–17. Both from Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference numbers Coll-1506/4/1 and Coll-1506/5/1.

  24. “Comment”, ABRO Report—January 1984, pp. 3–4. Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/1/1. See also Land (1985).

  25. “Preface”, ABRO Report—January 1986, p. 1. Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/1/1.

  26. A. J. Clark, A. L. Archibald, M. McClenaghan, J. P. Simons, C. B. A. Whitelaw, I. Wilmut: “High level expression of biomedical proteins in the milk of transgenic animals”, IAPGR Report for 1990–91, p. 53. Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/4/1.

  27. K. Campbell: “Multiplying farm animal embryos by nuclear transfer”, Roslin Institute Annual Report 1994–95, pp. 24–27; I. Wilmut: “New developments in embryo transfer”, Roslin Institute Annual Report 1995–96, pp. 26–30. Both from Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/5/1. Megan celebrated her tenth birthday in 2005 and was the oldest cloned animal at the time. For a scheme of the cloning technique used in Dolly see http://www.roslin.ed.ac.uk/public-interest/dolly-the-sheep/technical-aspects-of-cloning/ (last accessed June 2015).

  28. One year afterwards, in 1994, the AFRC acquired its current denomination as Biotechnology and Biological Sciences Research Council (BBSRC).

  29. Campbell always remained behind the scenes in the public dissemination despite, according to Wilmut, deserving 66 % of the credit of the cloning experiments. A lack of recognition may have had an emotional impact on Campbell, who left Edinburgh in 1999 and died in violent circumstances in 2012: http://www.telegraph.co.uk/news/uknews/1512377/I-didnt-clone-Dolly-the-sheep-says-prof.html and http://www.telegraph.co.uk/news/uknews/10065584/Scientist-behind-Dolly-the-sheep-killed-himself-by-mistake-in-drunken-fury.html (last accessed June 2015).

  30. “Potential benefits from cloning/nuclear transfer”, “Uses of cloning in farm animal production” and “Moral and ethical concerns”, Roslin Institute Annual Report, 1996–97, pp. 22–23, 24–25 and 26–27. Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/5/1.

  31. “Nuclear transfer: a brief history”, Roslin Institute Annual Report, 1996–97, pp. 18–19; J. Clark: “Genetic modification of livestock”, Roslin Institute Annual Report, 1997–98, pp. 32–41. Both from Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/5/1.

  32. By that time, the Roslin Institute had already expanded its influence to the US, given that in 1993 PPL merged with a company in Virginia that produced transgenic cows and pigs. Following Geron’s operation, PPL retained the part of the patent of the transgenic technologies that related to biopharming. However, it faced persisting problems in developing commercially viable products and around 2004 PPL went to liquidation and dissolved. “Stock market values: PPL therapeutics at £110 m”, Roslin Institute Report, 1995–96, pp. 18–19; “Roslin BioMed”, Roslin Institute Annual Report, 1997–98, pp. 20–21; “Roslin signs six year deal with Geron”, Roslin Institute Annual Report, 1998–99, pp. 14–17. All from Wellcome Trust Towards Dolly archival project, Edinburgh University Library Special Collections, provisional reference number Coll-1506/5/1. On the commercial activity and liquidation of PPL see Reid and Smith (2006).

  33. For a similar approach to rhetoric in the history of molecular biology see Abir-Am (1985).

  34. On the absence of farmers in the history of agriculture see Palladino (1990), Olby (1991), Vernon (1997), DeJager (1993), Woods (2012).

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Acknowledgments

This paper benefitted from useful comments from the journal and special issue editors, Staffan Müler-Wille and Giuditta Parolini, an anonymous referee and the members of a panel at the 2014 Annual Meeting of the British Society for the History of Science. Institutional and professional support at the University of Edinburgh and Roslin Institute was excellent, especially from Steve Sturdy, Ann Bruce, Grahame Bulfield and Clare Button. The investigations reported in this paper were funded by a Chancellor’s Fellowship and internal conference and research grants awarded by the University of Edinburgh. The final stages of my work were supported by a BBSRC research grant that will enable me to expand the project’s scope.

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  1. University of Edinburgh, Edinburgh, UK

    Miguel García-Sancho

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  1. Miguel García-Sancho
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Correspondence to Miguel García-Sancho.

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García-Sancho, M. Animal breeding in the age of biotechnology: the investigative pathway behind the cloning of Dolly the sheep. HPLS 37, 282–304 (2015). https://doi.org/10.1007/s40656-015-0078-6

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  • DOI: https://doi.org/10.1007/s40656-015-0078-6

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