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 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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1 Natural Selection in the Abstract
Concepts and Methods
3 Causality and MultiLevel Selection
4 Philosophical Issues in the LevelsofSelection Debate
5 The GenesEye View and its Discontents
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allele altruism argue argument average particle fitness causal causal decomposition cells Chapter character value character-fitness covariance clade clade selection collective character collective fitness collective level collective-level selection components concept contextual analysis contextual approach correlation Cov wi Covk w,z cross-level by-products Dawkins defined differential diploid direct selection discussed effect emergent character evolution evolutionary change evolutionary game theory evolutionary transitions evolve example fitness differences fitness2 focal units gene’s-eye perspective gene’s-eye view genic selection genotype global population group selection heterosis hierarchical levels individual selection interactions kin selection levels of selection levels-of-selection question Lewontin lower-level Maynard Smith means MLS1 multi-level selection multicelled organisms natural selection neighbourhood character number of offspring Nunney organismic fitness part–whole particle character particle-level selection phenotypic pluralism population genetics Price approach Price’s equation propagules regression relation replicators reproduction selection process selfish Smith and Szathmáry Sober and Wilson species selection Sterelny transmission bias types units of selection variance Vrba