Domesticating the Wild Type: a Historical Investigation of the Role of the Domestic-Wild Divide in Scientific Knowledge Production
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Domesticating the Wild Type: A Historical Investigation of the Role of the Domestic-Wild Divide in Scientific Knowledge Production Submitted by Tarquin Holmes to the University of Exeter as a thesis for the degree of Doctor of Philosophy in philosophy, in September 2015. This thesis is available for Library use on the understanding that it is copyright material and that no quotation from the thesis may be published without proper acknowledgement. I certify that all material in this thesis which is not my own work has been identified and that no material has previously been submitted and approved for the award of a degree by this or any other University. Page -1- Acknowledgements This thesis started life as a relatively diminutive 20,000 word MA dissertation on the origins of the wild type concept in classical genetics. It was then introduced into an artificially open PhD environment where, subjected to continual bombardment by all manner of unusual academic influences not found in its original habitation, it mutated (or should I say degenerated?) and dramatically expanded, requiring rigorous selection in order to be trimmed into shape and fully domesticated. As it is, it's still something of a monster, but one I find slightly less terrifying now that it's at least partially been tamed. Certainly, it's not the cute little mogwai it started as, but through a process of continual re-engineering, I think I've at least managed to find and remove most of the gremlins. Joking aside, this has been a difficult project which has taken longer than hoped and has taken more twists and turns than I could have expected. It's certainly been no linear evolution, but rather a thicket of branching enquiries, many of which have ended up being dead ends. I certainly could not have managed through if it had not been for the support of others. I will start by thanking my uncle Garth for funding the costs of my fees, as well as his secretary Mandy and my uncle and aunt Earle and Eve, who helped arrange the necessary transactions. I would then like to extend thanks to all the staff and faculty at Byrne House for helping create such a congenial and invigorating environment for research. This of course includes my supervisors Staffan and Sabina, who have been incredibly patient over the years with what has often been an ill-disciplined and sprawling project, as well as my mentor Nigel. I am also indebted to my fellow graduate students, especially Ann-Sophie, David, Jean, Jim, Jo, Lou, Kai, Mila, Nick and Trijsje, as well as post-doc and research fellow colleagues such as Berris, Chris, Dan, Daniele and Ginny, who together helped make Egenis a welcoming community of friends as well as minds (this list is far from exhaustive!). Further gratitude for their loving care goes to my immediate family, Ben, Zsuzsa, Marcus and Patti, who have always believed in my potential even when I have been less optimistic. But the one person I owe most to, and without whose undiminished and loving support I could not have done without, is my wife Kay, and it is to her that I dedicate this thesis. Page -2- Abstract This thesis focuses on the role and historical development of strategies of experimental domestication in scientific knowledge production, with a particular focus on the function of the laboratory strains known as 'wild types' in the model organism systems of classical genetics, where they play the role of standing in for the 'natural' instance of the species so that variation may be measured. As part of establishing how lab wild types came to assume this role, I have situated them within a much longer historical trajectory that tracks how changes in the manner that European intellectual traditions conceptualised the domestic-wild divide were linked to the development of new forms of scientific domestication and knowledge production. These new developments required that existing domesticating practices be intensified, expanded and analogised in order to better control, capture and comprehend 'wild' nature. My first two chapters introduce the domestic-wild divide by discussing both contemporary and ancient interpretations of it. In my third and fourth chapter, I explore the roots of the knowledge regime of European scientific domestication. I highlight Francis Bacon's campaign to use knowledge of domesticating practices to restore human dominion, before showing how Linnaeus later re-conceptualised the natural economy as an autonomous order and original order, with domestication reinterpreted as an artful transformation of nature requiring human maintenance to prevent reversion to its wild 'natural state'. I identify this idea of the wild as original and the domestic as derivative and artificially maintained as the basis of the original wild type concept. In my fifth chapter, I discuss Darwin's attempt to unite the domestic and wild under common laws of variation and selection, including his argument that reversion was simply a product of a return to ancestral conditions of existence. I observe that Darwin's theory of variation was problematic for the effort to bring wild nature under controlled conditions for study, so in my sixth and seventh chapters discuss how this difficulty was resolved, first by experimental naturalists both before and after Darwin who utilised vivaria and microscopes to bring pieces of nature indoors, and then by Weismann and Galton's sequestration of heredity, which helped persuade scientists that domestication was not in itself a cause of germinal variation. In my eighth and ninth chapter, I detail how sequestration led the early Mendelians de Vries and Bateson to assume that wild types could be brought into the lab from nature and purified into true-breeding strains. I discuss their differing atomist and interactionist perspectives on wild type, with de Vries favouring 'elementary species' as units of nature, whereas Bateson held wild types and mutants to Page -3- represent normal and abnormal forms of the species respectively. In my last chapter, I cover the replacement of Bateson's interactionist genetics by the reductionist genetics of the Morgan group and argue that this led to a disintegration of wild types into their component genes. I conclude with a discussion of what wild type strains in classical genetics were meant to be representative of, and end by establishing that whilst these strains may not wholly be representative of their species, they are nonetheless useful tools for scientific knowledge production. Page -4- Table of Contents Introduction – Introducing Wild Types and the Domestic-Wild Divide: pp. 6-16. Chapter 1 – The Domestic and the Wild: pp. 17-49. Chapter 2 – The Domestic-Wild Divide in Pre-Modern and Early Modern Western Thought: pp. 50-71. Chapter 3 – Gardens and Fields: Baconian Natural Philosophy and the Role of Hybrid Spaces for Scientific Knowledge Production & the Restoration of Human Dominion: pp. 72-95. Chapter 4 – Gardens of Knowledge, Economies of the Domestic and Wild: Linnaeus and the Origins of the Wild Type Concept: pp. 96-121. Chapter 5 – The Concept of Wild Type amongst Darwin and his Contemporaries: pp. 122- 148. Chapter 6 – Expanding Domestication, Encapsulating the Wild: the New Indoor Nature and the Birth of the Laboratory Lab Sciences, c. 1800-1880: pp 149-177. Chapter 7 – Darwinism, the Late 19th Century Problem of Variation, and the 'Move out of History': pp. 178-197. Chapter 8 – ‘Revisionary Darwinians’: De Vries & Bateson, their Influences & their Solutions to the Problem of Variation: pp. 198-224. Chapter 9 – Bateson and de Vries after Mendelism: pp. 225-249. Chapter 10 – The Chromosome Theory of Mendelian Heredity, the Downfall of Interactionist Models of Wild Type & Its Disintegration Into Genes: pp. 250-263. Conclusion – The Place of the Wild Type in Model Organism Systems in Classical Genetics: pp. 264-274. Bibliography: pp. 274-300. Figures 1 and 2: p. 249. Figure 3: p. 257. Page -5- Introduction – Introducing Wild Types and the Domestic-Wild Divide What are Wild Types? The term ‘wild type’ is generally used in genetics to describe individuals deemed ‘normal’ or typical of their species and also to refer to the ‘normal’ alleles of genes associated with these typical individuals. But more concretely wild type is what the standard lab strains of experimental organisms in genetics are commonly referred to as. The role these wild type strains play in model organism systems in genetics is one of acting as a control against which deviation is measured. As the philosopher Rachel Ankeny has observed, without first establishing a ‘wild type’ “it is not possible to have a ‘norm’ against which ‘abnormal’ (or more precisely, that which is variant) can be compared”. Selection of a wild type is therefore “the first step in the underlying strategy [of classical genetics]”.1 The necessity of undertaking this first step was recognised early on in genetics. A ‘Report of the Committee on Genetic form and Nomenclature’ from 1921, written up by the mouse geneticist C.C. Little, comments on how “In most animals and plants it is convenient to settle on a standard type, preferably the wild type, when this is known. The effects of the various genetic factors are in general to be measured by the departure from type which they bring about”. It was further noted that “This recommendation involves no real departure from the system now in use by most geneticists”.2 The importance of the wild type’s role as an instrument against which to measure variation in the laboratory has therefore been acknowledged both by early 20th century geneticists and some of today’s philosophers of science. It has commonly been emphasised that the historical selection of original model organism species and stocks often had a strongly arbitrary element where the principle concern was readiness to hand, familiarity with life history and known tolerance of captive conditions.