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Reconceptualising Evolution by Natural Selection

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Reconceptualising Evolution by Natural Selection COPYRIGHT AND USE OF THIS THESIS This thesis must be used in accordance with the provisions of the Copyright Act 1968. Reproduction of material protected by copyright may be an infringement of copyright and copyright owners may be entitled to take legal action against persons who infringe their copyright. Section 51 (2) of the Copyright Act permits an authorized officer of a university library or archives to provide a copy (by communication or otherwise) of an unpublished thesis kept in the library or archives, to a person who satisfies the authorized officer that he or she requires the reproduction for the purposes of research or study. The Copyright Act grants the creator of a work a number of moral rights, specifically the right of attribution, the right against false attribution and the right of integrity. You may infringe the author’s moral rights if you: - fail to acknowledge the author of this thesis if you quote sections from the work - attribute this thesis to another author - subject this thesis to derogatory treatment which may prejudice the author’s reputation For further information contact the University’s Director of Copyright Services sydney.edu.au/copyright Reconceptualising Evolution by Natural Selection Pierrick Bourrat A thesis submitted in fulfilment of the requirements for the degree of Doctor of Philosophy Faculty of Arts and Social Sciences The University of Sydney 2015 Abstract This thesis examines the theoretical and philosophical underpinnings of the concept of natural selection which is pervasively invoked in biology and other ‘evolutionary’ domains. Although what constitutes the process of natural selection appears to be very intuitive (natural selection results from entities exhibiting differences in fitness in a population), this conceals a number of theoretical ambiguities and difficulties. Some of these have been pointed out numerous times; others have hardly been noticed. One aim of this work is to unpack these difficulties and ambiguities; another is to provide new solutions and clarifications to them using a range of philosophical and conceptual tools. The result is a concept of natural selection stripped down from its biological specificities. I start by revisiting the entangled debates over whether natural selection is a cause of evolutionary change as opposed to a mere statistical effect of other causes, at what level this putative cause operates and whether it can be distinguished from drift. Borrowing tools from the causal modelling literature, I argue that natural selection is best conceived as a causal process resulting from individual level differences in a population. I then move to the question of whether the process of natural selection requires perfect transmission of types. I show that this question is ambiguous and can find different answers. From there, I distinguish the process of natural selection from some of its possible products, namely, evolution by natural selection and complex adaptation. I argue that reproduction and inheritance are conceptually distinct from natural selection, and using individual-based models, I demonstrate that they can be conceived as evolutionary products of it. This ultimately leads me to generalise the concepts of heritability and fitness used in the formal equations of evolutionary change. Finally, I argue that concepts of fitness and natural selection crucially depend on the grains of description at and temporal scales over which evolutionary explanations are given. These considerations reveal that the metaphysical status of the process of natural selection is problematic and why neglecting them can lead to flawed arguments in the levels of selection debate. Dedication À la mémoire de mon père, Michel, qui bien qu’il soit parti trop tôt m’a transmis sa curiosité Contents Abstract ............................................................................................................................... i Dedication ........................................................................................................................ iii Contents.............................................................................................................................. v List of figures .................................................................................................................... ix List of tables ..................................................................................................................... xi Acknowledgements ......................................................................................................... xii 0. Introduction ................................................................................................................ 1 1. The Causes of Evolutionary Change, an Individual-Level Perspective ................... 21 1.1. Introduction ...................................................................................................... 21 1.2. What fitness is: Bouchard and Rosenberg’s causal account of natural selection revisited 27 1.3. Fitness as a difference in intrinsic-invariable properties ................................. 33 1.3.1. Godfrey Smith on natural selection and drift................................................ 34 1.4. Problems with the intrinsic/extrinsic distinction ............................................ 42 1.4.1. Drift as intrinsic-variable properties ............................................................. 42 1.4.2. The possibility of Drift in an Indeterministic Set-up: Deterministic versus Probabilistic drift ....................................................................................................... 45 1.4.3. Two fundamentally distinct notions of drift ................................................. 47 1.4.4. Applying Strevens’ account of probability to deterministic drift ................. 49 1.5. Natural selection from a functional causal modelling perspective .................. 51 1.6. Teasing apart natural selection from drift ........................................................ 59 1.7. Natural selection and drift: population level causes of evolutionary change? . 65 1.7.1. Reisman and Forber’s arguments ................................................................. 66 1.7.2. Millstein’s argument ..................................................................................... 74 1.8. Conclusion of the chapter ................................................................................. 76 2. Natural Selection and its Products: Evolution by Natural Selection and Complex Adaptations ...................................................................................................................... 77 2.1. Introduction ...................................................................................................... 77 2.2. Minimal and Pure ENS .................................................................................... 81 2.3. Two cases of evolution ...................................................................................... 84 2.3.1. Case 1: Evolution due to differences in variable properties ......................... 87 2.3.2. Case 2: Evolution due to unreliable channel of transmission ...................... 91 vi CONTENTS 2.3.3. No pure ENS without perfect inheritance across generations ..................... 95 2.3.4. The replicator and classical approaches to ENS reconsidered .................... 99 2.4. Inconsistent Definitions of Natural selection ................................................. 101 2.5. Why reproduction is not essential to natural selection: the problem of individuality 105 2.6. Biased inheritance and natural selection: one or two processes? ................... 117 2.7. Conclusion of the chapter ............................................................................... 125 3. From persistors to replicators: Endogenising Reproduction and Inheritance....... 127 3.1. Introduction .................................................................................................... 127 3.2. ENS without reproduction .............................................................................. 130 3.3. Reproduction and inheritance in ENS ........................................................... 137 3.4. Biased inheritance and paradigmatic ENS .................................................... 149 3.5. Assessing the endogenisation-of-inheritance hypothesis .............................. 160 3.6. Conclusion of the chapter ............................................................................... 163 4. Moving from the Classical to the Formal Account of ENS: Heritability and Fitness reconsidered.................................................................................................................... 165 4.1. Introduction .................................................................................................... 165 4.2. The Price equation .......................................................................................... 168 4.3. Heritability in the Price equation ................................................................... 174 4.3.1. Different approaches to inheritance and heritability .................................. 174 4.3.2. Heritability: from recipes to the Price equation ......................................... 180 4.4. A problem for the recipes? .............................................................................. 184 4.5. Reinterpreting Heritability for Recipes .........................................................
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