CHALLENGING THE MODERN SYNTHESIS CHALLENGING THE MODERN SYNTHESIS Adaptation, Development, and Inheritance Edited by Philippe Huneman and Denis M. Walsh 1 1 Oxford University Press is a department of the University of Oxford. It furthers the University’s objective of excellence in research, scholarship, and education by publishing worldwide. Oxford is a registered trade mark of Oxford University Press in the UK and certain other countries. Published in the United States of America by Oxford University Press 198 Madison Avenue, New York, NY 10016, United States of America. © Oxford University Press 2017 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, without the prior permission in writing of Oxford University Press, or as expressly permitted by law, by license, or under terms agreed with the appropriate reproduction rights organization. Inquiries concerning reproduction outside the scope of the above should be sent to the Rights Department, Oxford University Press, at the address above. You must not circulate this work in any other form and you must impose this same condition on any acquirer. CIP data is on file at the Library of Congress ISBN 978– 0– 19– 937717– 6 9 8 7 6 5 4 2 1 Printed by Sheridan Books, Inc., United States of America CONTENTS Contributors vii Introduction: Challenging the Modern Synthesis—D enis M. Walsh and Philippe Huneman 1 PART I: Adaptation and Selection 1. Natural Selection, Adaptation, and the Recovery of Development— David J. Depew 37 2. Why Would We Call for a New Evolutionary Synthesis? The Variation Issue and the Explanatory Alternatives—P hilippe Huneman 68 3. Genetic Assimilation and the Paradox of Blind Variation— Arnaud Pocheville and Étienne Danchin 111 4. Evolutionary Theory Evolving—P atrick Bateson 137 PART II: Development 5. Evo- Devo and the Structure(s) of Evolutionary Theory: A Different Kind of Challenge—A lan C. Love 159 vi • Contents 6. Toward a Nonidealist Evolutionary Synthesis— Stuart A. Newman 188 7. Evolvability and Its Evolvability— Alessandro Minelli 211 8. “Chance Caught on the Wing”: Metaphysical Commitment or Methodological Artifact?—D enis M. Walsh 239 PART III: Inheritance 9. Limited Extended Inheritance— Francesca Merlin 263 10. Heredity and Evolutionary Theory—T obias Uller and Heikki Helanterä 280 11. Serial Homology as a Challenge to Evolutionary Theory: The Repeated Parts of Organisms from Idealistic Morphology to Evo- Devo—S téphane Schmitt 317 Index 349 CONTRIBUTORS Patrick Bateson is emeritus professor of ethology at the University of Cambridge. Much of his scientific career has been concerned with bridging the gap between the studies of behavior and those of underlying mechanisms, focusing on the process of behavioral imprinting in birds. Another aspect of his work was on the development of play and the induction of alternative pathways in cats depending on their early experience. Subsequently he has published extensively on develop- ment and evolution, including the modern study of epigenetics. Étienne Danchin is director of research at the French CNRS at the Evolution & Diversité Biologique (EDB) laboratoire in Toulouse gathering 60 academics, plus about 50 PhD and postdoc students. He coleads the Laboratoire of Excellence TULIP that regroups 5 laboratories totaling 400 staff. He is an expert in behav- ioral ecology, on which he wrote a textbook for Oxford University Press. His fields of interest range from social information use in habitat and mate choices to the evolution of coloniality in birds, and nongenetic inheritance (epigenetic and cultural), as well as information flows as determinants of ecological and evolu- tionary dynamics. He advocates the necessity to generalize the modern synthesis of evolution into an inclusive evolutionary synthesis incorporating all dimen- sions of nongenetic inheritance. David J. Depew is professor emeritus of rhetoric of inquiry at the University of Iowa. His work is in the history, philosophy, and rhetoric of evolutionary biology, with special interest in the history and conceptual foundations of the modern evolutionary synthesis. He is the coauthor, with Bruce H. Weber, of Darwinism Evolving: Systems Dynamics and the Genealogy of Natural Selection (1995); with the late Marjorie Grene of Philosophy of Biology: An Episodic History (2005); and most recently with John P. Jackson of an extended study of interactions between anthropologists and evolutionary biologists in 20th- century America. Heikki Helanterä is an evolutionary biologist with his main focus in social evo- lution. His research tackles the causes and consequences of variation in social viii • Contributors traits at both individual and group levels, and employs a wide range of methods from theoretical and conceptual work to genomic, population genetic, compar- ative, and behavioral studies. He has published over 50 papers and is currently a Kone Foundation Research Fellow at the University of Helsinki, Finland. Philippe Huneman is a research director at the Institut d’Histoire et de Philosophie des Sciences et des Techniques (CNRS/U niversité Paris I Sorbonne). A philosopher of biology, he works and publishes papers on issues related to evolutionary biology, on the concept of organism and its relations to Kant’s metaphysics, and on kinds of explanation in biology and ecology (with a focus on structural/t opological explanations). He has edited several books, including Understanding Purpose: Essays on Kant’s Philosophy of Biology (2007), Functions: Selection and Mechanisms (2013); From Groups to Individuals (2013, with F. Bouchard); and Handbook of Evolutionary Thinking in the Sciences (2015, with T. Heams, G. Lecointre, M. Silberstein). He is coeditor of the book series History, Philosophy, and Theory in Life Sciences. Alan C. Love is Associate Professor of Philosophy at the University of Minnesota and Director of the Minnesota Center for Philosophy of Science. His areas of specialization are philosophy of biology and philosophy of sci- ence. Alan’s research focuses on a variety of conceptual issues in evolution- ary and developmental biology with an emphasis on philosophical questions about conceptual change, explanatory pluralism, the structure of evolutionary theory, reductionism, the nature of historical science, and interdisciplinary epistemology. Francesca Merlin is researcher in philosophy of biology at CNRS (IHPST, UMR 8590, Paris, France). She obtained a PhD in philosophy at Université Paris 1— Panthéon Sorbonne in 2009. Her research focuses on central concepts in biol- ogy such as chance and probability, inheritance and epigenetics, in particular in the context of evolutionary theory. She currently works on the extension of bio- logical inheritance in the light of the fact that organisms inherit much more than DNA. She is a recipient of the 2010 Young Researcher Prize from the Société de Philosophie des Sciences in France. She published Mutations et aléas: Le hasard dans la théorie de l’évolution (2013). She has also coedited, with Thierry Hoquet, Précis de philosophie de la biologie (2014). Alessandro Minelli was full professor of zoology at the University of Padua, Italy, until his retirement in 2011; he has been vice president (1997– 1999) of the European Society for Evolutionary Biology. For several years his research focus was biological systematics— the subject of his book Biological Systematics: The State of the Art (1993)— but since the mid- 1990s his research interests have turned toward evolutionary developmental biology, the subject of his monographs The Contributors • ix Development of Animal Form (2003) and Perspectives in Animal Phylogeny and Evolution (2009). He is currently serving as specialty editor- in- chief for evolu- tionary developmental biology of Frontiers in Ecology and Evolutionary Biology. Stuart A. Newman is professor of cell biology and anatomy at New York Medical College, where he directs a research program in developmental biology. He has contributed to several scientific fields, including cell differentiation and pattern formation, the theory of biochemical networks, protein folding and assembly, and the mechanisms of morphological evolution. He has also written on the social and cultural dimensions of developmental biology. He is coauthor (with Gabor Forgacs) of Biological Physics of the Developing Embryo (2005), and coedi- tor (with Gerd B. Müller) of Origination of Organismal Form: Beyond the Gene in Developmental and Evolutionary Biology (2003) and (with Karl J. Niklas) of Multicellularity: Origins and Evolution (2016). He is editor- in- chief of Biological Theory, the journal of the KLI (Klosterneuburg, Austria). Arnaud Pocheville is a theoretical biologist and a philosopher of biology, cur- rently a research fellow at the University of Sydney, working on the project ‘Causal Foundations of Biological Information’ headed by Paul E. Griffiths. He holds a PhD in Life Sciences from the École Normale Supérieure, Paris. He advo- cates that we need new concepts and mathematical approaches to handle cases of time-scales and levels entanglements in biology. He has worked in particular on the evo/devo entanglement in relation to niche construction and non-genetic inheritance. Stéphane Schmitt is a research director at the French Centre National de la Recherche Scientifique (Paris). He works on the history of the life sciences, espe- cially in the 18th and 19th centuries. He has published many books and papers on the history of anatomy, embryology, and the sciences of evolution, and is the main editor of Buffon’s Œuvres complètes (2007– , 9 volumes published to date). Tobias Uller is an evolutionary biologist. His current work focuses on the rela- tions between development, heredity, and evolution by studying the evolutionary causes and consequences of developmental plasticity and parental effects. He has published over 130 papers on a broad range of topics in ecology and evolution and currently holds a Wallenberg Academy Fellowship and a Senior Lectureship at Lund University, Sweden. Denis M. Walsh is a professor in the Department of Philosophy, the Institute for the History and Philosophy of Science and Technology, and the Department of Ecology and Evolutionary Biology at the University of Toronto. He is author of Organisms, Agency and Evolution (2015).
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