Evolution and the Levels of Selection
Does natural selection act primarily on individual organisms, on groups, on genes, or on whole species? Samir Okasha provides a comprehensive analysis of the debate in evolutionary biology over the levels of selection, focusing on conceptual, philosophical and foundational questions. A systematic framework is developed for thinking about natural selection acting at multiple levels of the biological hierarchy; the framework is then used to help resolve outstanding issues. Considerable...
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Does natural selection act primarily on individual organisms, on groups, on genes, or on whole species? The question of levels of selection - on which biologists and philosophers have long disagreed - is central to evolutionary theory and to the philosophy of biology. Samir Okasha's comprehensive analysis gives a clear account of the philosophical issues at stake in the current debate.
Introduction 1Natural Selection in the Abstract 10Introduction 10Abstract Formulations of Darwinian Principles 13Price's Equation 18Interpretation of Price's Equation 23Statistical versus Causal Decomposition 25Random Drift and Causal Decomposition 31Price's Equation and the Lewontin Conditions 34Selection at Multiple Levels: Concepts and Methods 40Introduction 40Hierarchical Organization 40Selection at Multiple Levels: Key Concepts 46Particle Characters and Collective Characters 48Life Cycles 49Particle Fitness and Collective Fitness 53The Two Types of Multi-Level Selection 56Particle Heritability and Collective Heritability 59Price's Equation in a Hierarchical Setting 62The Price Approach to MLS1 62Applications 66Heritability in MLS1 Revisited 71The Price Approach to MLS2 74Causality and Multi-Level Selection 76Introduction 76Causes, Correlations, and Cross-LevelBy-Products 76Selection on Correlated Characters 80Cross-Level By-Products in MLS1 84Contextual Analysis: Further Remarks 89Contextual Analysis versus Price's Equation 93Cross-Level By-Products in MLS2 100Particle[RightArrow]Collective By-Products 100Collective[RightArrow]Particle By-Products 107Philosophical Issues in the Levels-of-Selection Debate 112Introduction 112Emergence and Additivity 112The Emergent Character Requirement 112Additivity and the Wimsatt/Lloyd Approach 114Emergent Relations and the Damuth-Heisler Approach 119Screening Off and the Levels of Selection 121Realism versus Pluralism about the Levels of Selection 125Pluralism and Causality 128Pluralism and Hierarchical Organization 130Pluralism and Multiple Representations 133Reductionism 139The Gene's-Eye View and its Discontents 143Introduction 143The Origins of Gene's-Eye Thinking 143Genic Selection and the Gene's-Eye View: Process versus Perspective 146Outlaws and Genetic Conflicts 149Price's Equation versus Contextual Analysis Revisited 154Bookkeeping and Causality 158The Limits of Genic Accounting 158Sober and Lewontin's Heterosis Argument 162Context-Dependence and the Gene's-Eye View 166Reductionism and Pluralism Revisited 169The Group Selection Controversy 173Introduction 173Origins of the Group Selection Controversy 174Group Selection and the MLS1/MLS2 Distinction 178Kin Selection, Reciprocal Altruism, and Evolutionary Game Theory 180Maynard Smith versus Sober and Wilson on Group Heritability 185The Averaging Fallacy 189Random versus Assortative Grouping, Strong versus Weak Altruism 192Contextual Analysis versus the Neighbour Approach 198Species Selection, Clade Selection, and Macroevolution 203Introduction 203Origins of Species Selection 203Genuine Species Selection versus 'Causation from Below' 206Species versus Avatars: Damuth's Challenge 210The Concept of Clade Selection 212Levels of Selection and the Major Evolutionary Transitions 218Introduction 218The Transformation of the Levels-of-Selection Question 219Genic versus Hierarchical Approaches to the Transitions 225MLS1 versus MLS2 in Relation to Evolutionary Transitions 229Michod on Fitness Decoupling and the Emergence of Individuality 233Concluding Remarks 236Bibliography 241Index 257
\ From the Publisher"Samir Okasha's wonderful new book...is a philosophical examination of the conceptual framework that multi-level selection theory deploys...It is gratifying that his book engages the details of mathematical models and at the same time connects those details with broader philosophical questions."--Elliott Sober, Bioscience\ "Okasha has written an extremely important book. required reading for anyone working on the levels of selection question.."--Jonathan Michael Kaplan, Notre Dame Philosophical Reviews\ "Evolution and the Levels of Selection is a major contribution toward putting this controversial area on a coherent conceptual and philosophical footing...Okasha has greatly clarified many of the central issues. I can't imagine anyone working on multilevel selection - or attempting to dismiss it - without reading this book."--David Jablonski, Science\ "...a major conceptual contribution to evolutionary theory...Okasha's book makes the sort of contribution that will not be able to be ignored by anyone interested in this field for many years to come."--Massimo Pigliucci, Biology and Philosophy\ "...an extremely thought-provoking and important book about a difficult and highly technical topic."--Matt Haber, Mind\ "...a clearly written, unique and useful book."--Elizabeth Lloyd, Trends in Ecology and Evolution\ \ \
