ASSEMBLING LIFE. Models, the cell, and the reformations of biological science, 1920-1960 Max Stadler Centre for the History of Science, Technology and Medicine Imperial College, London University of London PhD dissertation 1 I certify that all the intellectual contents of this thesis are of my own, unless otherwise stated. London, October 2009 Max Stadler 2 Acknowledgments My thanks to: my friends and family; esp. my mother and Julia who didn't mind their son and brother becoming a not-so-useful member of society, helped me survive in the real world, and cheered me up when things ; my supervisor, Andrew Mendelsohn, for many hours of helping me sort out my thoughts and infinite levels of enthusiasm (and Germanisms-tolerance); my second supervisor, David Edgerton, for being the intellectual influence (I thought) he was; David Munns, despite his bad musical taste and humour, as a brother-in-arms against disciplines; Alex Oikonomou, for being a committed smoker; special thanks (I 'surmise') to Hermione Giffard, for making my out-of-the-suitcase life much easier, for opening my eyes in matters of Frank Whittle and machine tools, and for bothering to proof-read parts of this thesis; and thanks to all the rest of CHoSTM; thanks also, for taking time to read and respond to over-length drafts and chapters: Cornelius Borck; Stephen Casper; Michael Hagner; Rhodrie Hayward; Henning Schmidgen; Fabio de Sio; Sktili Sigurdsson; Pedro Ruiz Castell; Andrew Warwick; Abigail Woods; to Anne Harrington for having me at the History of Science Department, Harvard, and to Hans- Joerg Rheinberger for having me at the Max-Planck-Institute for the History of Science in Berlin; thanks, finally, to all those who in some way or another encouraged, accompanied and/or enabled the creation and completion of this thing, especially: whoever invented the internet; Hanna Rose Shell; and the Hans Rausing Fund. 3 Abstract The subject of this thesis are the fundamental, bioelectrical expressions of life from the interwar period into the 1950s. Or rather — at issue are very elusive manifestations of life — it is a history of models of the cellular life, and the things, materials and practices surrounding them. This living cell, modelled or not, indeed is largely absent from the narratives we tell of twentieth century biology. The big pictures we have revolve around even smaller entities: genes, molecules, and enzymes. But the cell was still there, this thesis shows. And its presence, this thesis argues, must affect the stories of life science we tell. The historical material covered here thus deliberately encompasses a range of fairly obscure forays into the nature of bio-electricity — from exercise physiology to colloid science to medical physics - as well as such well-known advances as the Hodgkin-Huxley model of the neuron. For, the central aim of this thesis is to show that not only was this living cell central to shaping biological science in the twentieth century, we can uncover it at very mundane and unexpected places. Science, this thesis shows, knew the elusive cell mainly through other and mundane things — as models. These models were assembled from a fabric that was not living, organic and natural, but fabricated, processed, made-up and hence, known, controlled and transparent: things ranging from soap-films to electrical circuits to calculation machines. It follows that this science of life was not in fact life science but something still broader which belongs to histories far beyond that of biological specialities, model- organisms, academic research and disciplines, or indeed, that of the progressive molecularization of life. Cellular life took shape — mediated via models — within broad- scale technological and scientific projects that coalesced around the macroscopic materialities that defined this modern, industrial age. 4 Sources and abbreviations ADM Records of the Admiralty, Naval Forces, Royal Marines, Coastguard, and related bodies, UK National Archives, Kew AE Personal papers of Sir Alfred Charles Glyn Egerton, Royal Society, London AIR Records created or inherited by the Air Ministry, the Royal Air Force, and related bodies, UK National Archives, Kew AVHL The papers of Professor A.V. 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Rockefeller Archives Center, Sleepy Hollow, NY RF/RG.303 Detlev W Bronk papers, Rockefeller Archives Center, Sleepy Hollow, NY RNDL The papers of Sir John Randall, Churchill Archives Centre, Churchill College, Cambridge ROUGHTON/APS Francis Roughton Papers, American Philosophical Society Library, Philadelphia ROUGHTON/CUL The papers of Francis Roughton, Cambridge University Library, Cambridge Minutes of the Colloid and Biophysics Committee, Royal Society of Chemistry, Royal Institution, London SCHEMINZKY Personalakte Scheminzky, Med.12 Nr.4, Universitatsarchiv Wien, Vienna T Records created and inherited by HM Treasury, UK National Archives, Kew UCC Records Office, University College, London UGC Records created or inherited by the Higher Education Funding Council for England, UK National Archives, Kew WILKINSON Private papers of F J. Wilkinson, Imperial War Museum, London WO Records created or inherited by the War Office, Armed Forces, Judge Advocate General, and related bodies, UK National Archives, Kew YOUNG Papers of John Z. Young, UCL Special Collections, London 7 Figures Figure 1: Reversal effects, 1921 Fischer, Hooker and McLaughlin (1921), Figure 111 Figure 2: Emulsion reversal, 1916 Clowes (1916), p.421 Figure 3: Beutner oil-systems ....Haynes (1922), p.96 Figure 4: Brown's substance-table, 1915 Brown (1915), p.598 Figure 5: Substance-table, 1926 Collander (1927), p.216 Figure 6: The paucimolecular model, 1935 Danielli & Dayson (1935), Figure 1 Figure 7: Ergostol and irradiation products, 1934 Danielli & Adam (1934), p.1584 Figure 8: Subjected to 'supersonics', 1932 Harvey & Loomis (1932), p.151 Figure 9: Flattening oil globules, 1934 Harvey & Shapiro (1934), p.259 Figure 10: Proposed 'schema', 1934 Danielli & E.N. Harvey (1934), p.491 Figure 11: Bubble formations, 1935/36 Danielli (1936), p.399 Figure 12: Black film, 1929 Lawrence (1929), Plate X Figure 13: The 'black'. Sandwich model, 1929 Lawrence (1929), Figure 59 Figure 14: The 'black'. Sandwich model, 1930 Adam (1930), Figure 28 Figure 15: Phases of muscular heat production, 1920 Hill & Hartree (1920),