From Biological Revolution to Biotech Age: Plant Biotechnology in British Agriculture since 1950 Matthew Robert Holmes Submitted in accordance with the requirements for the degree of PhD The University of Leeds School of Philosophy, Religion, and the History of Science Centre for History and Philosophy of Science September 2017 ii iii The candidate confirms that the work submitted is his own and that appropriate credit has been given when reference has been made to the work of others. The sole paper, Matthew Holmes, ‘Crops in a Machine: Industrialising Barley Breeding in Twentieth-Century Britain’ in Jon Agar and Jacob Ward (Eds.), Technology, Environment and Modern Britain (London: UCL Press, 2018), is submitted alongside this thesis as loose sheets. Material from this paper appears in Chapters 1 and 2. The sole paper, Matthew Holmes ‘Somatic Hybridization: The Rise and Fall of a Mid- Twentieth-Century Biotechnology’, Historical Studies in the Natural Sciences 48 (2018), is submitted alongside this thesis as loose sheets. Material from this paper appears in Chapter 3. The sole paper, Matthew Holmes, ‘Changing Techniques in Crop Plant Classification: Molecularization at the National Institute of Agricultural Botany during the 1980s’ Annals of Science 74 (2017): 149-164, is submitted alongside this thesis as loose sheets, Material from this paper appears in Chapter 4. This copy has been supplied on the understanding that it is copyright material and that no quotation from the thesis may be published without proper acknowledgement. ©2017 The University of Leeds and Matthew Robert Holmes iv v Acknowledgements I would first like to thank my supervisors, Greg Radick and Tina Barsby. Greg has been a constant source of encouragement and inspiration throughout the writing of this thesis. He has urged me on every step of the way and has helped and supported me in all my endeavours. There really is no way of thanking him enough for all that he has done for me these past four years. Tina has been generous with her time and knowledge, taking this thesis down fascinating avenues that would have otherwise have been left unexplored. I would also like to thank my academic advisor, Graeme Gooday, who asked the questions that needed answering and helped me think through difficult conceptual problems. My thanks also go to Joan Fujimura for hosting me during a two month stay at the University of Wisconsin-Madison and guiding me through the intricacies of interviewing. Throughout my time at Leeds, the staff and students at the Centre for History and Philosophy of Science have always been welcoming and supportive. I would like to thank all of them for making my time there so enjoyable. This thesis could not have been written without the help of staff at a number of archives across the UK. I would like to thank all of them for hosting me. In particular, I would like to thank Tricia Cullimore at the NIAB, who went out of her way to make me welcome and point me in the direction of new sources. Over the course of writing this thesis I attended and presented at a number of conferences and workshops, which encouraged and shaped the ideas in this thesis. I would like to thank all of the organisers of these events and the British Society for the History of Science for generously providing me with travel grants on numerous occasions. The AHRC provided the financial assistance which made the writing of this thesis possible. Finally, I would like to thank my family for their care and reassurance. My brother Jonathan endured a barrage of doubts and complaints. My partner Annie not only provided me with unwavering love and support, but proofread everything: her kindness made the burden of writing much lighter. vi vii Abstract The modern Biotech Age possesses a very particular set of characteristics: the use of recombinant DNA technology, a close relationship between academic science and industry and, in Britain, public hostility to genetically modified crops. Yet despite increasingly widespread recognition among historians of science that biotechnology has a long and multi-faceted history, there is no thorough account of the history of plant biotechnology in British agriculture. Harnessing previously unexamined archival sources at the National Institute of Agricultural Botany (NIAB), John Innes Centre (JIC) and the Science Museum, this thesis uncovers a number of largely unexamined plant biotechnologies and discusses their uptake in British agriculture since the mid- twentieth century. In doing so, it raises several new insights for historians. Chapters One and Two demonstrate how two commercially successful biotechnologies, industrial hybridization and mutation breeding, found agricultural applications by careful integration with existing industrial systems. Chapter Three shows how plant cell fusion became a genuine alternative to recombinant DNA technology during the 1960s and ‘70s. Chapter Four counters the standard narrative of a move from the morphological to the molecular in biological analysis with a case study of electrophoresis and other classificatory technologies. Chapter Five demonstrates the importance of Cold War ideology on the development of biotechnology with a case study of the graft hybrid in British horticulture. Finally, Chapter Six examines the GM controversy in Britain and considers what broader lessons about public attitudes to biotechnology can be taken from the debate. Taken together, this thesis demonstrates that a unique combination of plant biotechnologies emerged in mid- twentieth-century Britain. These biotechnologies succeeded or failed to influence British agriculture thanks to a combination of technological, economic and ideological factors. The Biotech Age could, at many points since 1950, have emerged in a very different way with very different characteristics. viii ix Table of Contents Acknowledgements ....................................................................................................... v Abstract ...................................................................................................................... vii Table of Contents ......................................................................................................... ix List of Figures ............................................................................................................... xi List of Abbreviations .................................................................................................... xiii Introduction .................................................................................................................. 1 1. State of the Historiography: The Long History of Biotechnology, in Britain and Beyond ........................................................................................................................... 6 2. State of the Historiography: The Long History of Envirotechnology, in Britain and Beyond ......................................................................................................................... 11 3. State of the Historiography: The Politics of Post-War Plant Breeding, in Britain and Beyond ................................................................................................................. 16 4. Overview of the Thesis ........................................................................................... 22 1. Industrial Hybridization: British Barley as Cold War Biotechnology ...................... 29 1. Engineering Hybrid Barley ...................................................................................... 32 2. Industrial Crosstalk with the Brewing Industry ..................................................... 36 3. The NIAB’s 1970 Hybrid Conference ..................................................................... 39 4. Malthus’s Shallow Grave ........................................................................................ 42 5. Economic Concerns in British Agriculture ............................................................. 47 6. Hybrid Monocultures and Rise of Plant Disease ................................................... 50 Conclusions ................................................................................................................. 53 2. Mutation Breeding: Chromosome Breakage in the Atomic Age ........................... 57 1. Chromosome Breakage: A New Branch of Cold War Science .............................. 59 2. Gamma Radiation and British Barley Breeding ..................................................... 67 3. Farming and Fallout: Radiation Botany at Wantage ............................................. 74 Conclusions ................................................................................................................. 81 3. Cell Fusion: Biotechnological Optimism in Plant Physiology ................................. 85 1. Plant Physiologists and the Cell.............................................................................. 87 2. Enzymes and Protoplast Production ...................................................................... 92 3. Somatic Hybridization and Plant Breeding ............................................................ 96 4. Technical Difficulties and Competition ................................................................ 101 5. Biotechnological Collaboration and Cybrids........................................................ 106 x Conclusions ..............................................................................................................