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The Changing Role of the Embryo in Evolutionary Thought P1: JZZ/KAB P2: IKB/JZN QC: KOD/JZN T1: KOD 0521806992agg.xml CB793B/Amundson 0 521 80699 2 April 24, 2005 17:42 The Changing Role of the Embryo in Evolutionary Thought In this book, Ron Amundson examines 200 years of scientific views on the evolution–development relationship from the perspective of evolutionary devel- opmental biology (evo–devo). This new perspective challenges several popular views about the history of evolutionary thought by claiming that many earlier authors made history come out right for the Evolutionary Synthesis. The book starts with a revised history of nineteenth-century evolutionary thought. It then investigates how development became irrelevant to evolution with the Evolutionary Synthesis. It concludes with an examination of the contrasts that persist between mainstream evolutionary theory and evo–devo. This book will appeal to students and professionals in the philosophy of science, and the philosophy and history of biology. Ron Amundson is Professor of Philosophy, University of Hawaii at Hilo. i P1: JZZ/KAB P2: IKB/JZN QC: KOD/JZN T1: KOD 0521806992agg.xml CB793B/Amundson 0 521 80699 2 April 24, 2005 17:42 ii P1: JZZ/KAB P2: IKB/JZN QC: KOD/JZN T1: KOD 0521806992agg.xml CB793B/Amundson 0 521 80699 2 April 24, 2005 17:42 cambridge studies in philosophy and biology General Editor Michael Ruse Florida State University Advisory Board Michael Donoghue Yale University Jean Gayon University of Paris Jonathan Hodge University of Leeds Jane Maienschein Arizona State University Jes´us Moster´ın Instituto de Filosof´ıa (Spanish Research Council) Elliott Sober University of Wisconsin Alfred I. Tauber The Immune Self: Theory or Metaphor? Elliott Sober From a Biological Point of View Robert Brandon Concepts and Methods in Evolutionary Biology Peter Godfrey-Smith Complexity and the Function of Mind in Nature William A. Rottschaefer The Biology and Psychology of Moral Agency Sahotra Sarkar Genetics and Reductionism Jean Gayon Darwinism’s Struggle for Survival Jane Maienschein and Michael Ruse (eds.) Biology and the Foundation of Ethics Jack Wilson Biological Individuality Richard Creath and Jane Maienschein (eds.) Biology and Epistemology Alexander Rosenberg Darwinism in Philosophy, Social Science, and Policy Peter Beurton, Raphael Falk, and Hans-J¨org Rheinberger (eds.) The Concept of the Gene in Development and Evolution David Hull Science and Selection James G. Lennox Aristotle’s Philosophy of Biology Marc Ereshefsky The Poverty of the Linnaean Hierarchy Kim Sterelny The Evolution of Agency and Other Essays William S. Cooper The Evolution of Reason Peter McLaughlin What Functions Explain Steven Hecht Orzack and Elliott Sober (eds.) Adaptationism and Optimality Bryan G. Norton Searching for Sustainability Sandra D. Mitchell Biological Complexity and Integrative Pluralism Joseph LaPorte Natural Kinds and Conceptual Change Greg Cooper The Science of the Struggle for Existence Jason Scott Robert Embryology, Epigenesis, and Evolution William F. Harms Information & Meaning in Evolutionary Processes iii P1: JZZ/KAB P2: IKB/JZN QC: KOD/JZN T1: KOD 0521806992agg.xml CB793B/Amundson 0 521 80699 2 April 24, 2005 17:42 iv P1: JZZ/KAB P2: IKB/JZN QC: KOD/JZN T1: KOD 0521806992agg.xml CB793B/Amundson 0 521 80699 2 April 24, 2005 17:42 The Changing Role of the Embryo in Evolutionary Thought Roots of Evo–Devo RON AMUNDSON University of Hawaii at Hilo v Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, São Paulo Cambridge University Press The Edinburgh Building, Cambridge , UK Published in the United States of America by Cambridge University Press, New York www.cambridge.org Information on this title: www.cambridge.org/9780521806992 © Ron Amundson 2005 This book is in copyright. Subject to statutory exception and to the provision of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press. First published in print format 2005 - ---- eBook (NetLibrary) - --- eBook (NetLibrary) - ---- hardback - --- hardback Cambridge University Press has no responsibility for the persistence or accuracy of s for external or third-party internet websites referred to in this book, and does not guarantee that any content on such websites is, or will remain, accurate or appropriate. P1: JZZ/KAB P2: IKB/JZN QC: KOD/JZN T1: KOD 0521806992agg.xml CB793B/Amundson 0 521 80699 2 April 24, 2005 17:42 To my father, Art Amundson. He taught me curiosity, and the pleasure of taking things apart to see how they work. vii P1: JZZ/KAB P2: IKB/JZN QC: KOD/JZN T1: KOD 0521806992agg.xml CB793B/Amundson 0 521 80699 2 April 24, 2005 17:42 viii P1: JZZ/KAB P2: IKB/JZN QC: KOD/JZN T1: KOD 0521806992agg.xml CB793B/Amundson 0 521 80699 2 April 24, 2005 17:42 Contents Acknowledgments page xiii 1 Introduction 1.1 Evo–Devo as New and Old Science1 1.2 Evo–Devo and the Windfall of the 1990s4 1.3 How I Came to Write this Book9 1.4 Historical Format 11 1.5 Epistemological Concepts in Historical Context 14 1.5.1 Inductivist Caution 14 1.5.2 Idealism 16 1.5.3 Two Essentialisms 18 1.6 Explanatory Relativity 20 1.7 Historical Conventions 22 1.8 Historical Precis´ 23 part i. darwin’s century: beyond the essentialism story 2 Systematics and the Birth of the Natural System 2.1 Introduction 31 2.2 The Discovery of Species Fixism 34 2.3 Linnaeus and His Contemporaries 39 2.4 French Systems: Jussieu and Cuvier 41 2.5 British Systems and the Growth of Taxonomic Realism 45 2.6 Review of Species Fixism, Essentialism, and Real Groups 50 3 The Origins of Morphology, the Science of Form 3.1 Morphology and Natural Theology 53 ix P1: JZZ/KAB P2: IKB/JZN QC: KOD/JZN T1: KOD 0521806992agg.xml CB793B/Amundson 0 521 80699 2 April 24, 2005 17:42 x Contents 3.2 Form as a Topic of Study 55 3.2.1 Goethe 55 3.2.2 The Great Cuvier-Geoffroy Debate 56 3.2.3 Von Baer and Development 58 3.2.4 The Study of Form Summarized 61 3.3 Natural Theologians on Unity of Type 62 3.3.1 William Paley 63 3.3.2 William Buckland 64 3.3.3 Charles Bell 64 3.3.4 William Whewell 65 3.3.5 Peter Mark Roget 66 3.4 The Structural Turn 67 3.4.1 Martin Barry 68 3.4.2 William Carpenter 70 3.4.3 Rudolph Leuckart 72 3.5 What is Natural Theology? 73 4 Owen and Darwin, The Archetype and the Ancestor 4.1 Introduction 76 4.2 Typology Defined: Kinds of Types 78 4.3 Owen Builds the Archetype 82 4.4 Owen on Species Origins 88 4.5 Anti-Adaptationism 93 4.6 Darwin’s use of Morphological Types 96 4.7 Misunderstanding Darwin on Owen 99 4.8 Darwin on Unity of Type 102 4.9 A Structuralist Evolutionary Theory? 103 4.10 How Darwin Differed 104 5Evolutionary Morphology: The First Generation of Evolutionists 5.1 The Program of Evolutionary Morphology 107 5.2 Evolutionary Morphology as Non-Darwinian and as Darwinian 108 5.3 The Biogenetic Law 112 5.4 Early Origins in Phylogeny and Ontogeny 114 5.5 Explaining Form 118 5.6 The Struggles of Evolutionary Morphology 121 5.7 The Conflict between Adaptation and Structure 125 6 Interlude 6.1 Two Narratives of the History of Evolutionary Biology 130 6.2 One Theory or Two? 130 P1: JZZ/KAB P2: IKB/JZN QC: KOD/JZN T1: KOD 0521806992agg.xml CB793B/Amundson 0 521 80699 2 April 24, 2005 17:42 Contents xi 6.3 Grounds for Species Fixism 131 6.4 Darwin’s other Primary Achievement: The Tree of Life 132 6.5 The Significance of Gappiness 134 6.6 And Forward 136 part ii. neo-darwin’s century: explaining the absence and the reappearance of development in evolutionary thought 7 The Invention of Heredity 7.1 Truisms of Heredity 139 7.2 Epigenetic Origins of Heredity 140 7.3 Epigenetic Heredity During the Nineteenth Century 143 7.3.1 Martin Barry 143 7.3.2 Charles Darwin 144 7.3.3 August Weismann 144 7.4 The Cleavage between Heredity and Development 148 7.5 Reinforcing the Dichotomy: Rewriting Weismann and Johannsen 152 7.6 Broad and Narrow Heredity 155 8 Basics of the Evolutionary Synthesis 8.1 A Long Story Made Short 159 8.2 The Struggles of Natural Selection 160 8.3 Problems in Characterizing the Evolutionary Synthesis 161 8.4 The Evolutionary Synthesis Characterized 163 8.5 By-Products of the Core of Synthesis Thought 166 8.5.1 Systematics 166 8.5.2 Phylogeny 167 8.5.3 Mechanisms 167 9 Structuralist Reactions to the Synthesis 9.1 Experimental Embryology and the Synthesis 169 9.2 The Program of Experimental Embryology 170 9.3 The Embryological Critique of the Synthesis 175 9.3.1 Critique 1: The Causal Completeness Principle 175 9.3.2 Critique 2: The Developmental Paradox 177 9.3.3 Critique 3: Fundamental versus Superficial Characters 180 9.3.4 Cytoplasmic Inheritance versus Darwinian Extrapolation 185 P1: JZZ/KAB P2: IKB/JZN QC: KOD/JZN T1: KOD 0521806992agg.xml CB793B/Amundson 0 521 80699 2 April 24, 2005 17:42 xii Contents 9.4 Points of Contact among Developmental and Genetic Biologists, and Synthesis Evolutionists 189 9.4.1 Sewall Wright 190 9.4.2 Oxford Morphology 191 9.4.3 Waddington and Schmalhausen 193 9.4.4 Richard Goldschmidt 195 9.5 Historical Reflection: Explanatory Goals 196 9.5.1 Form-Theoretic Evolutionary Theory 196 9.5.2 Change-Theoretic Evolutionary Theory 197 10 The Synthesis Matures 10.1 The Darwin Centennial Celebration 198 10.2 Uses of Dichotomies 201 10.3 Proximate versus Ultimate: Context 203 10.4 Population Thinking versus Typological Thinking: Context 204 10.5 Ernst Mayr as a Structuralist? 209 10.6 The Enlarged Quiver of Dichotomies 211 11 Recent Debates and the Continuing Tension 11.1 Diversity versus Commonality: Starting with Genes 213 11.2 The Four Dichotomies Defend the Synthesis 218 11.2.1
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