Evolution and the Levels of Selection

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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 attention is paid to the concept of causality as it relates to the levels of selection, in particular the idea that natural selection at one hierarchical level can have effects that 'filter' up or down to other levels. Unlike previous work in this area by philosophers of science, full account is taken of the recent biological literature on 'major evolutionary transitions' and the recent resurgence of interest in multi-level selection theory among biologists. Other biological topics discussed include Price's equation, kin and group selection, the gene's eye view, evolutionary game theory, outlaws and selfish genetic elements, species and clade selection, and the evolution of individuality. Philosophical topics discussed include reductionism and holism, causation and correlation, the nature of hierarchical organization, and realism and pluralism.
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Editorial Reviews

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

"The current volume provides an exceptionally lucid and analyitically rigorous review of the main conceptual challenges facing biologists and philosophers who have engaged in this work."--Mark E. Borrello, The Quarterly Review of Biology

"Major contribution toward putting this controversial area on a coherent conceptual and philosophical footing. ... I can't imagine anyone working on multilevel selection-or attempting to dismiss it-without reading this book."--Science

"Every philosopher of biology interested in aspects of the levels of selection debates ought to confront this material, and should think seriously about how the positions he or she ahs staked out fits into the frameworks Okasha outlines. Okasha has written an extremely important book."--Jonathan Michael Kaplan, Notre Dame Philosophical Reivews

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Product Details

  • ISBN-13: 9780199556717
  • Publisher: Oxford University Press, USA
  • Publication date: 1/15/2009
  • Pages: 288
  • Product dimensions: 5.40 (w) x 8.40 (h) x 0.70 (d)

Meet the Author

Samir Okasha is Professor of Philosophy at Bristol University. Before that he taught at the University of York for 3 years, and was a Jacobsen Research Fellow at the London School of Economics for 2 years. He was a Postdoctoral Research Fellow at the National Univeristy of Mexico for 1 year and received his doctorate in 1998 from the University of Oxford.

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Table of Contents

Introduction     1
Natural Selection in the Abstract     10
Introduction     10
Abstract Formulations of Darwinian Principles     13
Price's Equation     18
Interpretation of Price's Equation     23
Statistical versus Causal Decomposition     25
Random Drift and Causal Decomposition     31
Price's Equation and the Lewontin Conditions     34
Selection at Multiple Levels: Concepts and Methods     40
Introduction     40
Hierarchical Organization     40
Selection at Multiple Levels: Key Concepts     46
Particle Characters and Collective Characters     48
Life Cycles     49
Particle Fitness and Collective Fitness     53
The Two Types of Multi-Level Selection     56
Particle Heritability and Collective Heritability     59
Price's Equation in a Hierarchical Setting     62
The Price Approach to MLS1     62
Applications     66
Heritability in MLS1 Revisited     71
The Price Approach to MLS2     74
Causality and Multi-Level Selection     76
Introduction     76
Causes, Correlations, and Cross-LevelBy-Products     76
Selection on Correlated Characters     80
Cross-Level By-Products in MLS1     84
Contextual Analysis: Further Remarks     89
Contextual Analysis versus Price's Equation     93
Cross-Level By-Products in MLS2     100
Particle[RightArrow]Collective By-Products     100
Collective[RightArrow]Particle By-Products     107
Philosophical Issues in the Levels-of-Selection Debate     112
Introduction     112
Emergence and Additivity     112
The Emergent Character Requirement     112
Additivity and the Wimsatt/Lloyd Approach     114
Emergent Relations and the Damuth-Heisler Approach     119
Screening Off and the Levels of Selection     121
Realism versus Pluralism about the Levels of Selection     125
Pluralism and Causality     128
Pluralism and Hierarchical Organization     130
Pluralism and Multiple Representations     133
Reductionism     139
The Gene's-Eye View and its Discontents     143
Introduction     143
The Origins of Gene's-Eye Thinking     143
Genic Selection and the Gene's-Eye View: Process versus Perspective      146
Outlaws and Genetic Conflicts     149
Price's Equation versus Contextual Analysis Revisited     154
Bookkeeping and Causality     158
The Limits of Genic Accounting     158
Sober and Lewontin's Heterosis Argument     162
Context-Dependence and the Gene's-Eye View     166
Reductionism and Pluralism Revisited     169
The Group Selection Controversy     173
Introduction     173
Origins of the Group Selection Controversy     174
Group Selection and the MLS1/MLS2 Distinction     178
Kin Selection, Reciprocal Altruism, and Evolutionary Game Theory     180
Maynard Smith versus Sober and Wilson on Group Heritability     185
The Averaging Fallacy     189
Random versus Assortative Grouping, Strong versus Weak Altruism     192
Contextual Analysis versus the Neighbour Approach     198
Species Selection, Clade Selection, and Macroevolution     203
Introduction     203
Origins of Species Selection     203
Genuine Species Selection versus 'Causation from Below'     206
Species versus Avatars: Damuth's Challenge     210
The Concept of Clade Selection     212
Levels of Selection and the Major Evolutionary Transitions     218
Introduction     218
The Transformation of the Levels-of-Selection Question     219
Genic versus Hierarchical Approaches to the Transitions     225
MLS1 versus MLS2 in Relation to Evolutionary Transitions     229
Michod on Fitness Decoupling and the Emergence of Individuality     233
Concluding Remarks     236
Bibliography     241
Index     257
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