Evolutionary Novelty: a Philosophical and Historical Investigation
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Evolutionary novelty: a philosophical and historical investigation Submitted by Thibault Racovski to the University of Exeter as a thesis for the degree of Doctor of Philosophy in Philosophy In June 2018 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. Signature: ………………………………………………………….. Acknowledgements I am happy to be able to thank at least some of the many people who accompanied me during my doctoral experience and made the completion of this thesis possible Philippe Huneman, my co-supervisor in Paris, for introducing me to philosophy of biology eleven years ago and accompanying me through all the stages of this research project with his immense support His generosity and his example greatly contributed to my understanding and appreciation of the world of research and academia The mark of his numerous suggestions and of his critical eye is present throughout this thesis John Dupré, my co-supervisor in Exeter, for his astute guidance and his patience during my sometimes tumultuous intellectual journey John transformed my understanding of the writing process and made vivid the value of collaboration in research I benefitted greatly from being part of the project “A Process Ontology for Contemporary Biology” Along with John, I want to thank Dan Nicholson, Stephan Guttinger and Anne Sophie Meincke for fruitful discussions At the same time, I am grateful to John for allowing me to pursue my own line of research (although only after I had the process tattoo done) Paul Griffiths, my second supervisor in Exeter, for his perceptive comments and his help in structuring the thesis, and especially for convincing me to focus on plants Sabina Leonelli, my mentor, for her attention, her advice, and for her example Alan Love and Staffan Müller-Wille, my examiners, who helped shaping this research through their writings and conversation well before the viva 2 I want to thank Jean Gayon, Michel Morange, Daniel Andler and Pascal Ludwig, my professors and supervisors at the universities of Paris 1 and Paris 4 and at the ENS, for their teaching, ideas and encouragements I am grateful to Joe Cain, Tim Lewens, Gregory Radick and Giovanni Boniolo for showing interest in my thesis project at its early stages and for their constructive suggestions During the last few years I also benefitted from discussions with Anouk Barberousse, James DiFrisco, Gerd Müller, Ann-Sophie Barwich, Sarah Samadi, Sébastien Dutreuil, David Chavalarias, Sophia Rousseau-Mermans, Max Dresow, Laura Nuño de la Rosa, Ceslo Neto, Shane Glackin and Adam Toon At my arrival at Egenis in Exeter, Selina Nath, Jo Donaghy, David Wyatt, Nick Binney, Jim Lowe and Tarquin Holmes made me feel welcome despite the clouds’ efforts to the contrary I tried to do the same with Çağlar Karaça, Thomas Bonnin, Jaanika Puusalu, Flavia Fabris, Javier Suarez, Gregor Halfmann, Thijs van Stigt and Mark Canciani who joined me the following years, even though I was not always keeping my office chair in the postgrad room warm They enriched me in many ways, along with Guclan, Nancy, Stathis, Angie, Simon, Andrea, Ljubica Gezim, Niccolò, Chris and others My family and friends, in France and elsewhere, brought love and joy to these years of research I sometimes wonder what all of their versions of the content of my thesis complied together would look like It would surely be a very baroque document, and, to me, a fascinating, if not flattering, read To Sanja Uskliknimo s ljubavlju Svetitelju Sanji 3 Abstract Evolutionary novelty, the origin of new characters such as the turtle shell or the flower, is a fundamental problem for an evolutionary view of life. Accordingly, it is a central research topic in contemporary biology involving input from several biological disciplines and explanations at several levels of organization. As such it raises questions relative to scientific collaboration and multi-level explanations. Novelty is also involved in theoretical debates in evolutionary biology. It has been appropriated by evo-devo, a scientific synthesis linking research on evolution and development. Thanks to its focus on development, evo-devo claims to explain the mechanistic origin of novelties as new forms, while the Modern Synthesis can only provide statistical explanation of evolutionary change. The origin of an evolutionary novelty is a historical emergence of a new character involving form and function. I focus on three neglected dimensions of the problem of novelty, the functional-historical approach to the problem, research on novelty in the Modern Synthesis era and novelty in plants. I compare the evo-devo approach to novelty to a functional-historical approach of novelty. I focus on its origin in Darwin and its presence in the Modern Synthesis. The comparison of the two approaches reveals distance between conceptual frameworks and proximity in explanatory practices. This is partly related to unwarranted conceptual opposition. In particular, I list several ways of distinguishing novelty and adaptation, some of which are not conceptually sound. I then focus on the relation between novelty and adaptation in the Modern Synthesis era, and on the relation of novelty to other fundamental biological problems (speciation, origin of higher taxa, complexity). Pushing this approach further, I challenge the view that the Modern Synthesis excluded development and reached a hardened consensus. Finally, I analyse how Günter Wagner’s developmental theory of novelty applies to novelties in plant. 4 Table of contents List of figures. _________________________________________________________ 7 GENERAL INTRODUCTION _______________________________________________ 8 FOUR DIFFERENT PROBLEMS ________________________________________________ 10 PHILOSOPHICAL AND HISTORICAL APPROACHES TO THE PROBLEM OF THE ORIGIN OF EVOLUTIONARY NOVELTIES _________________________________________________ 14 OUTLINE OF THE THESIS ____________________________________________________ 20 CHAPTER 1: TWO APPROACHES TO THE PROBLEM OF NOVELTY _______________ 23 INTRODUCTION ___________________________________________________________ 23 Outline of the chapter ____________________________________________________________ 24 I – CHARACTERISATION OF THE PROBLEM OF NOVELTY THE VIEW OF EVO-DEVO ______ 24 Overview of the contemporary debate on the content and structure of evolutionary biology ___ 27 The old and the new in evo-devo ___________________________________________________ 30 Evo-devo and the origin of novelties as a new research question __________________________ 32 II – THE FUNCTIONAL-HISTORICAL APPROACH TO NOVELTY _______________________ 36 Three stages in the functional-historical approach _____________________________________ 36 Darwin and Dohrn, conversion and succession of functions ______________________________ 37 The second phase of the functional-historical approach _________________________________ 40 The third phase _________________________________________________________________ 47 III – CAUSES OF THE NEGLECT OF THE FUNCTIONAL-HISTORICAL APPROACH __________ 54 The reduction to population genetics ________________________________________________ 54 Objections to the functional definition _______________________________________________ 55 The neglect of the functional-historical approach is not universal _________________________ 58 IV – NOVELTY AND ADAPTATION _____________________________________________ 60 Illustration: the origin of novelties in the avian hind limb skeleton _________________________ 62 Comparison of the types of explanations developed by Walter Bock and Gerd Müller _________ 63 V – NOVELTY, VARIATION, AND THE CREATIVITY OF NATURAL SELECTION ____________ 66 The generation of variation and the sorting of variation _________________________________ 67 Two meanings of adaptations ______________________________________________________ 68 CONCLUSION _____________________________________________________________ 71 CHAPTER 2 – NOVELTY IN THE MODERN SYNTHESIS ERA Part 1 NOVELTY IN THE CONCEPT AND RESEARCH SPACE OF THE MODERN SYNTHESIS _________________ 76 INTRODUCTION ___________________________________________________________ 76 Clarifications on the Modern Synthesis_______________________________________________ 77 The theoretical content of the Modern Synthesis ______________________________________ 79 Outline of the chapter ____________________________________________________________ 80 I – THE CONCEPT OF NOVELTY ASSOCIATED WITH MUTATIONISM __________________ 82 Homeotic mutations, macromutations and systemic mutations ___________________________ 84 The central debate against mutationism and saltationism the mode of origin of higher taxa ___ 87 II – NOVELTY, SPECIES AND SPECIATION _______________________________________ 91 Novelty and speciation ___________________________________________________________ 91 Experimental taxonomy in botany __________________________________________________ 97 III – NOVELTY AND HIGHER TAXA ____________________________________________ 101 Novelty and the origin of higher taxa _______________________________________________ 101 The origin of angiosperms ________________________________________________________ 103 Differences in emphasis __________________________________________________________