Agricultural Science, Plant Breeding and the Emergence of a Mendelian System in Britain, 1880-1930 Berris Charnley Submitted in accordance with the requirements of the degree of PhD The University of Leeds Department of Philosophy September 2011 ii The candidate confirms that the work submitted is his own, except where work which has formed part of jointly-authored publications has been included. The contribution of the candidate and the other authors to this work has been explicitly indicated below. The candidate confirms that appropriate credit has been given within the thesis where reference has been made to the work of others. The collaborative paper, Berris Charnley and Gregory Radick (2010). “Plant Breeding and Intellectual Property Before and After the Rise of Mendelism: The Case of Britain”, in D. J. Kevles et al. (eds.), Living Properties: Making Knowledge and Controlling Ownership in Modern Biology, pp. 51-55. Berlin: Max Planck Institute for the History of Science, submitted alongside this thesis as loose sheets, was produced in collaboration with Prof. Gregory Radick. Material dealing with W. F. R. Weldon and the ancestral law of heredity was produced by Prof. Radick. The introduction was planned collaboratively and written by Prof. Radick. All other material was produced by Berris Charnley. Material from this paper appears in Chapter 3 of this thesis. 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. © 2011 The University of Leeds and Berris Charnley. iii Acknowledgements First and foremost I would like to thank Gregory Radick, my lead supervisor. Greg has been tireless in keeping my often wayward prose on the straight and narrow, and in leading by example, his intellectual curiosity and enthusiasm have inspired me on too many occasions to count. Secondly, I would like to thank Graeme Gooday, my co-supervisor, and the other members of the Owning and Disowning Invention project; Stathis Arapostathis, Graham Dutfield, Jonathan Hopwood-Lewis and Christine Macleod. On the many happy occasions when I shared my work with this group their feedback was insightful and always helpful. The Centre for History and Philosophy of Science, my institutional home over the last four years, has always been a hospitable place to work. I hope the centre’s staff and students see their ideas flatteringly reflected in my thinking in this thesis. These people have also been my friends, so I should thank them twice, along with other friends, and my family, for their caring support along the way. One other group of friends deserve a special mention; Sam Bacha, Dom Berry, Chris Renwick and Paulo Wylis formed a team who proof read this thesis. Their patience and perseverance with this un-enjoyable task is only matched by my luck in having such good friends. Many of the ideas presented here were born and shaped at conferences and workshops. I would like to take this opportunity to thank several groups which have been unstintingly generous in creating opportunities for me to present my work: Daniel J. Kevles, Hans-Jörg Rheinberger and Jean-Paul Gaudillière who organised the Living Properties and Plants, Animals and Ownership workshops held at the Max Planck Institute for the History of Science and Yale; Alessandro Nuvolari, Kristine Bruland and Liliane Pérez who organised the Innovation without Patents workshops in Geneva, Eindhoven and Utrecht; Staffan Müller-Wille and the other iv organisers of the Egenis Seminar series; and the organisers of ISHPSSB 2009 in Brisbane; the British Council’s Darwin Now conference; WEHC 2009 in Utrecht and SHOT 2008 in Lisbon. The participants of these events have been equally generous with their ideas, feedback and critical engagement with my work. Collecting the evidence for this thesis involved a great deal of time spent in archives, in some cases, literally on the other side of the world. The members of staff at the archives I visited were unanimously patient, professional and friendly. Kate West at the John Innes and Tricia Cullimore at NIAB were both particularly helpful in bringing new material to my attention. My archive research was funded by the Department of Philosophy at the University of Leeds, the Royal Historical Society and the AHRC. The AHRC also provided financial assistance with maintenance and fees, without which writing this thesis would not have been possible. Finally, I would like to dedicate the thesis to Marija and Julian. Marija, my wife, has never given up on me, no matter how high the drift of papers has piled up in our front room. I will always be thankful for her loving support over the past few years. Julian’s help runs throughout these pages. I often stayed with him when I was working in London or traveling back and forth to conferences and workshops. Julian’s curiosity about my work and willingness to engage with, and challenge, what I was telling him always inspired me. Julian’s life held the world around him to a higher standard, he will be sorely missed. Julian Brown (1977-2011) il miglior fabbro. v Abstract Following Thomas P. Hughes’s systems approach in the history of technology, and making use of previously unexamined sources, this dissertation seeks to show that the development of British Mendelism may be explained, and the success it enjoyed more accurately gauged, by analysing the emergence of a system whose elements justified the theory, protected it, made it useful, and slowly territorialized the world. Accordingly, the analysis will cover the principle elements of this system: the system builders, institutes, ideas and varieties that were, in one way or another, Mendelian. The first of the Mendelian system builders, William Bateson, is already well known for his introduction of Mendelism to Britain in the years after 1901 and his coinage of a new name for the discipline; Genetics. He was joined by two colleagues, Rowland Biffen and Thomas Wood, both of whom collaborated with Bateson in creating a string of institutes concerned with changing agriculture by using the new Mendelian theory. The proponents of the new theory often talked of their new found ability to transfer characters and build up new varieties of agricultural value. These claims were welcomed by politicians and the popular press and the idea that the new genetics would lead to a beneficial revolution in agriculture became a popular cause of the day. However, the release of the first of these new Mendelian varieties in 1910 in Britain is far less well known than the almost simultaneous development of the chromosome theory at Columbia University by Thomas Hunt Morgan. On one view of the history of genetics, the discipline, which had been born in Moravia, and popularised in Britain, was from 1910 most fruitfully developed in Morgan’s fly room. From this perspective it might be thought that the British School, under Bateson, became a disciplinary backwater, at least in part because Bateson refused to accept chromosome theory. This thesis argues that far from being in a genetic backwater, Bateson along with Mendelian allies Biffen and Wood were at the cutting edge of a wide ranging movement to improve agriculture through the introduction of new Mendelian varieties. vi Table of Contents Acknowledgements iii Abstract v Table of Contents vi List of Illustrations ix Introduction 1 British Genetics and Agriculture in the Twentieth Century 5 Mendelian Success in Britain and Further Afield 13 A New Historiographical Approach 20 The Thesis Plan 27 1. A New Institutional Map: Creating Space for Mendelism 31 1.1 Agricultural Science and Mendelism at Cambridge 36 1.1.1 The Cambridge Department of Agriculture before 1901 36 1.1.2 Mendelism at the Department 41 1.1.3 The Cambridge School of Agriculture 48 1.2 Other Mendelian Organisations 51 1.2.1 The Home Grown Wheat Committee 52 1.2.2 The John Innes Horticultural Research Institute 55 1.2.3 The Development Commission 60 1.3 New Funding and New Institutes 62 1.3.1 The Plant Breeding Institute 62 1.3.2 The British Seed Corn Association 65 1.3.3 The National Institute for Agricultural Botany 67 1.3.4 Popular Reception of the Mendelian System 75 vii 2. A New Theoretical Map: Purity as a Theoretical Problem 81 2.1 The Arrival of Mendelian Theory 85 2.1.1 Pre-Mendelian Inheritance and the Rogue Problem 85 2.1.2 The New Theory: Gametic Purity and Segregation 89 2.1.3 William Bateson and the Heterozygote Explanation of Rogues 93 2.2 The New Theory in an Agricultural Context 99 2.2.1 Pure Line Theory and the Statistical Solution 99 2.2.2 Biffen’s Rogue Wheat and Unwieldy Ratios 105 2.2.3 Bateson’s Rogue Peas and the Physiological Solution 114 3. Managing Mendelian Varieties: From Experimental Plot to Market (and back again) 123 3.0 Prelude: The Moral Economy of Plant Breeding 126 3.1 Mendelism and Wheat Breeding 133 3.1.1 Breeding and the Mendelian Discovery 133 3.1.2 Biffen on Mendelian Plant Breeding 138 3.2 The Mendelian Innovation Process 143 3.2.1 Biffen the Mendelian Wheat Breeder 143 3.2.2 Marketing the New Varieties: The Case of Yeoman 145 3.2.3 Harvesting Success 151 3.2.4 Protecting the New Varieties: From Yeoman to Yeoman II 154 4. Remaking the Field: How the New Varieties Meshed with the New Farming 163 4.1 The Importance of Agricultural History 167 4.1.1 British Agriculture in the Doldrums, 1880-1930 167 4.2 Mendelian Strategies for Agricultural Development 178 viii 4.2.1 Disease Resistance 179 4.2.2 Strength and the All-English Campaign 181 4.2.3 Intensification 186 4.3 A Reassessment of the Success of Biffen’s Varieties 189 4.3.1 The Popularity of Mendelian Varieties 189 4.3.2 The Use of Mendelian Varieties 194 4.3.3 Biological Innovation Prior to 1950 201 5.