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Proc Biol Sci. 2015 Aug 22;282(1813):20151019. doi: 10.1098/rspb.2015.1019.

The extended evolutionary synthesis: its structure, assumptions and predictions.

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School of Biology, University of St Andrews, St Andrews, Fife, UK
Edward Grey Institute, Department of Zoology, University of Oxford, Oxford, UK Department of Biology, University of Lund, Lund, Sweden.
Department of Biology, Stanford University, Herrin Hall, Stanford, CA 94305, USA.
School of Philosophy, Australian National University, Canberra, Australia School of History, Philosophy, Political Science and International Relations, Victoria University of Wellington, Wellington, New Zealand.
Department of Theoretical Biology, University of Vienna, Vienna, Austria.
Department of Biology, Indiana University, Bloomington, IN 47405-7107, USA.
Cohn Institute for the History of Philosophy of Science and Ideas, Tel Aviv University, Tel Aviv, Israel.
Mansfield College, University of Oxford, Oxford, UK.


Scientific activities take place within the structured sets of ideas and assumptions that define a field and its practices. The conceptual framework of evolutionary biology emerged with the Modern Synthesis in the early twentieth century and has since expanded into a highly successful research program to explore the processes of diversification and adaptation. Nonetheless, the ability of that framework satisfactorily to accommodate the rapid advances in developmental biology, genomics and ecology has been questioned. We review some of these arguments, focusing on literatures (evo-devo, developmental plasticity, inclusive inheritance and niche construction) whose implications for evolution can be interpreted in two ways—one that preserves the internal structure of contemporary evolutionary theory and one that points towards an alternative conceptual framework. The latter, which we label the 'extended evolutionary synthesis' (EES), retains the fundaments of evolutionary theory, but differs in its emphasis on the role of constructive processes in development and evolution, and reciprocal portrayals of causation. In the EES, developmental processes, operating through developmental bias, inclusive inheritance and niche construction, share responsibility for the direction and rate of evolution, the origin of character variation and organism-environment complementarity. We spell out the structure, core assumptions and novel predictions of the EES, and show how it can be deployed to stimulate and advance research in those fields that study or use evolutionary biology.


developmental plasticity; evolutionary developmental biology; extended evolutionary synthesis; inclusive inheritance; niche construction; reciprocal causation

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