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A THEORY OF CONCEPTUAL ADVANCE : EXPLAINING CONCEPTUAL CHANGE IN EVOLUTIONARY, MOLECULAR, AND EVOLUTIONARY DEVELOPMENTAL BIOLOGY by Ingo Brigandt M.Sc. in Mathematics, University of Konstanz, 1999 M.A. in Philosophy, University of Pittsburgh, 2005 Submitted to the Graduate Faculty of the Faculty of Arts and Sciences in partial fulfillment of the requirements for the degree of Doctor of Philosophy University of Pittsburgh 2006 UNIVERSITY OF PITTSBURGH FACULTY OF ARTS AND SCIENCES This dissertation was presented by Ingo Brigandt It was defended on August 2, 2006 and approved by Robert B. Brandom Distinguished Service Professor of Philosophy, University of Pittsburgh James G. Lennox Professor of History and Philosophy of Science, University of Pittsburgh Sandra D. Mitchell Professor of History and Philosophy of Science, University of Pittsburgh G¨unter P. Wagner Alison Richard Professor of Ecology and Evolutionary Biology, Yale University Dissertation Directors: Anil Gupta Distinguished Professor of Philosophy, University of Pittsburgh Paul E. Griffiths ARC Federation Fellow, Professor of Philosophy, University of Queensland ii Copyright c by Ingo Brigandt 2006 iii A THEORY OF CONCEPTUAL ADVANCE : EXPLAINING CONCEPTUAL CHANGE IN EVOLUTIONARY, MOLECULAR, AND EVOLUTIONARY DEVELOPMENTAL BIOLOGY Ingo Brigandt, PhD University of Pittsburgh, 2006 Abstract The theory of concepts advanced in the dissertation aims at accounting for a) how a concept makes successful practice possible, and b) how a scientific concept can be subject to rational change in the course of history. Traditional accounts in the philosophy of science have usually studied concepts in terms only of their reference; their concern is to establish a stability of reference in order to address the incommensurability problem. My discussion, in contrast, suggests that each scientific concept consists of three components of content: 1) reference, 2) inferential role, and 3) the epistemic goal pursued with the concept’s use. I argue that in the course of history a concept can change in any of these three components, and that change in one component — including change of reference — can be accounted for as being rational relative to other components, in particular a concept’s epistemic goal. This semantic framework is applied to two cases from the history of biology: the homology concept as used in 19th and 20th century biology, and the gene concept as used in different parts of the 20th century. The homology case study argues that the advent of Darwinian evolutionary theory, despite introducing a new definition of homology, did not bring about a new homology concept (distinct from the pre-Darwinian concept) in the 19th century. Nowadays, however, distinct homology concepts are used in systematics/evolutionary biology, in evolutionary developmental biology, and in molecular biology. The emergence of these different homology concepts is explained as occurring in a rational fashion. The gene case study argues that conceptual progress occurred iv with the transition from the classical to the molecular gene concept, despite a change in reference. In the last two decades, change occurred internal to the molecular gene concept, so that nowadays this concept’s usage and reference varies from context to context. I argue that this situation emerged rationally and that the current variation in usage and reference is conducive to biological practice. The dissertation uses ideas and methodological tools from the philosophy of mind and language, the philosophy of science, the history of science, and the psychology of concepts. v TABLE OF CONTENTS PREFACE .............................................. x 1.0 INTRODUCTION ..................................... 1 2.0 CONCEPTUAL CHANGE AS A PHILOSOPHICAL PROBLEM ....... 13 2.1 Referential Approaches to Concepts and Conceptual Change ............ 14 2.1.1 Limitations of Referential Accounts in the Philosophy of Science ..... 18 2.2 Bringing Concepts Back In : Explaining and Assessing Conceptual Change .... 24 2.2.1 Kitcher’s Theory of Conceptual Change ................... 24 2.2.2 Unsolved Issues in Kitcher’s Account ..................... 27 2.2.3 Critique of Kitcher’s Referential Framework of Conceptual Change .... 38 2.2.4 Summary of the Chapter ............................ 43 3.0 AN ACCOUNT OF CONCEPTS IN TERMS OF SEVERAL SEMANTIC COMPONENTS ............ 45 3.1 Inferential Role and Epistemic Goal As Components of Content .......... 48 3.1.1 Inferential Role Semantics ........................... 51 3.1.2 Formal Semantics and Semantic Atomism .................. 55 3.1.3 Motivations for Inferential Role Semantics .................. 61 3.1.4 Material Inference ............................... 68 3.2 Criticisms of Inferential Role Semantics ........................ 73 3.2.1 Compositionality of Concepts ......................... 73 3.2.2 Inferential Role and Reference ......................... 80 3.3 Concept Individuation .................................. 89 3.3.1 Troubles With Meaning Monism ....................... 91 3.3.2 Toward a Meaning Pluralism ......................... 101 3.3.3 Individuating Scientific Concepts ....................... 110 3.4 Summary of the Framework of Concepts ........................ 117 vi 4.0 THE HOMOLOGY CONCEPT BEFORE 1950 ................... 125 4.0.1 Homology: The Very Idea ........................... 127 4.0.2 Creationism, Teleosemantics, and Other Non-Biological Theories ..... 131 4.1 Homology before 1859 : The Emergence of a Crucial Conceptual Practice ..... 135 4.1.1 The Positional Criterion ............................ 136 4.1.2 Serial Homology ................................ 145 4.1.3 The Embryological Criterion .......................... 148 4.1.4 Richard Owen: Naming and Homology .................... 154 4.2 A Semantic Characterization of the Homology Concept Used Before 1859 ..... 162 4.3 Homology from 1859 until 1950 : An Instance of Conceptual Stability ....... 172 4.3.1 A Novel Homology Concept? ......................... 174 4.3.2 Evolutionary Morphology in the 19th Century ................ 180 4.3.3 Evolutionary Morphology in the 20th Century ................ 191 4.4 A Semantic Characterization of the Homology Concept Used Before 1950 ..... 196 5.0 HOMOLOGY IN CONTEMPORARY BIOLOGY : THE PHYLOGENETIC, THE DEVELOPMENTAL, AND THE MOLECULAR HOMOLOGY CONCEPT ...................... 211 5.1 Homology in Evolutionary Biology and Systematics : The Study and Explanation of Differences Between Species ............................. 213 5.1.1 Evolutionary Biology: Transformational Approach to Homology ...... 214 5.1.2 Systematic Biology: Taxic Approach to Homology ............. 216 5.1.3 A Semantic Account of the Phylogenetic Homology Concept ........ 221 5.2 Homology in Evolutionary Developmental Biology : The Search for an Account of Morphological Unity ................................... 227 5.2.1 Irreducible Levels of Morphological Organization .............. 228 5.2.2 Developmental Approaches to Homology ................... 235 5.2.3 A Semantic Account of the Developmental Homology Concept ....... 244 5.2.4 Implications for Theories of Reference .................... 252 5.3 Homology in Molecular Biology : The Practical Success of an Operational Concept .......................................... 259 5.3.1 The Phylogenetic Concept in Molecular Evolution and Phylogeny ..... 260 5.3.2 A Semantic Account of the Molecular Homology Concept ......... 265 5.4 Summary of the Discussion of Homology ....................... 270 vii 6.0 THE CHANGE OF THE GENE CONCEPT .................... 280 6.1 The Classical Gene Concept .............................. 282 6.1.1 From the Notion of Unit-Characters to the Genotype Concept ....... 285 6.1.2 The Chromosome Theory and the Morgan Group .............. 296 6.1.3 The Content of the Classical Gene Concept and Its Limited Explanatory Potential ..................................... 307 6.1.4 The Classical Gene Concept in (Contemporary) Population Genetics ... 315 6.2 The Classical Molecular Gene Concept ......................... 319 6.2.1 Genetics on the Way Toward a Molecular Gene Concept .......... 319 6.2.2 The Content and Explanatory Potential of the Classical Molecular Gene Concept ..................................... 329 6.2.3 Conceptual Progress Without Reduction and Stable Reference ....... 339 6.3 The Contemporary Molecular Gene Concept ..................... 351 6.3.1 Recent Empirical Findings and the Fragmentation of a Concept ...... 353 6.3.2 A Semantic Analysis of the Contemporary Gene Concept ......... 362 6.4 Summary of the Discussion of the Gene Concept ................... 375 7.0 CONCLUSION ....................................... 380 7.1 Philosophical Insights Yielded by the Application of the Framework of Concepts to the Case Studies ................................... 389 7.2 Summary and Implications of the Case Studies: Constraints on Any Theory of Concepts ......................................... 402 APPENDIX A. CONCEPTS IN PSYCHOLOGY .................... 406 APPENDIX B. CRITIQUES OF NEO-DARWINISM AND THE EMERGENCE OF EVOLUTIONARY DEVELOPMENTAL BIOLOGY ............. 419 B.1 Developmental Constraints and Evolvability ................. 423 B.2 The Explanation of the Origin of Body Plans and Evolutionary Novelties 430 B.3 Morphology or Developmental Genetics? ................... 438 BIBLIOGRAPHY ......................................... 441 viii LIST OF FIGURES 1 Homologies of the Mammalian Forelimb ......................... 128 2 Table from Geoffroy Saint-Hilaire’s Philosophie
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