A Companion to the Philosophy of Biology Blackwell Companions to Philosophy This outstanding student reference series offers a comprehensive and authoritative survey of philosophy as a whole. Written by today’s leading philosophers, each volume provides lucid and engaging coverage of the key fi gures, terms, topics, and problems of the fi eld. Taken together, the volumes provide the ideal basis for course use, representing an unparalleled work of reference for students and specialists alike. Already published in the series: 23. A Companion to Early Modern Philosophy Edited by Steven Nadler 1. The Blackwell Companion to Philosophy, 24. A Companion to Philosophy in the Middle Second Edition Ages Edited by Nicholas Bunnin and Eric Tsui-James Edited by Jorge J. E. Gracia and Timothy B. Noone 2. A Companion to Ethics Edited by Peter Singer 25. A Companion to African-American Philosophy 3. A Companion to Aesthetics Edited by Tommy L. Lott and John P. Pittman Edited by David Cooper 26. A Companion to Applied Ethics 4. A Companion to Epistemology Edited by R. G. Frey and Edited by Jonathan Dancy and Ernest Sosa Christopher Heath Wellman 5. A Companion to Contemporary Political 27. A Companion to the Philosophy of Education Philosophy (2 Volume Set), Second Edition Edited by Randall Curren Edited by Robert E. Goodin and Philip Pettit 28. A Companion to African Philosophy 6. A Companion to Philosophy of Mind Edited by Kwasi Wiredu Edited by Samuel Guttenplan 29. A Companion to Heidegger 7. A Companion to Metaphysics Edited by Hubert L. Dreyfus and Edited by Jaegwon Kim and Ernest Sosa Mark A. Wrathall 8. A Companion to Philosophy of Law and Legal 30. A Companion to Rationalism Theory Edited by Alan Nelson Edited by Dennis Patterson 31. A Companion to Ancient Philosophy 9. A Companion to Philosophy of Religion Edited by Mary Louise Gill and Pierre Pellegrin Edited by Philip L. Quinn and Charles Taliaferro 32. A Companion to Pragmatism 10. A Companion to the Philosophy of Language Edited by John R. Shook and Joseph Margolis Edited by Bob Hale and Crispin Wright 33. A Companion to Nietzsche 11. A Companion to World Philosophies Edited by Keith Ansell Pearson Edited by Eliot Deutsch and Ron Bontekoe 34. A Companion to Socrates 12. A Companion to Continental Philosophy Edited by Sara Ahbel-Rappe and Edited by Simon Critchley and William Schroeder Rachana Kamtekar 13. A Companion to Feminist Philosophy 35. A Companion to Phenomenology and Edited by Alison M. Jaggar and Existentialism Iris Marion Young Edited by Hubert L. Dreyfus and 14. A Companion to Cognitive Science Mark A. Wrathall Edited by William Bechtel and George Graham 36. A Companion to Kant 15. A Companion to Bioethics Edited by Graham Bird Edited by Helga Kuhse and Peter Singer 37. A Companion to Plato 16. A Companion to the Philosophers Edited by Hugh H. Benson Edited by Robert L. Arrington 38. A Companion to Descartes 17. A Companion to Business Ethics Edited by Janet Broughton and John Carriero Edited by Robert E. Frederick 39. A Companion to the Philosophy of Biology 18. A Companion to the Philosophy of Science Edited by Sahotra Sarkar and Anya Plutynski Edited by W. H. Newton-Smith 19. A Companion to Environmental Philosophy Edited by Dale Jamieson 20. A Companion to Analytic Philosophy Forthcoming Edited by A. P. Martinich and David Sosa 21. A Companion to Genethics 40. A Companion to Hume Edited by Justine Burley and John Harris Edited by Elizabeth S. Radcliffe 22. A Companion to Philosophical Logic 41. A Companion to Aristotle Edited by Dale Jacquette Edited by Georgios Anagnostopoulos A Companion to the Philosophy of Biology Edited by Sahotra Sarkar and Anya Plutynski © 2008 by Blackwell Publishing Ltd except for editorial material and organization © 2008 by Sahotra Sarkar and Anya Plutynski BLACKWELL PUBLISHING 350 Main Street, Malden, MA 02148-5020, USA 9600 Garsington Road, Oxford OX4 2DQ, UK 550 Swanston Street, Carlton, Victoria 3053, Australia The right of Sahotra Sarkar and Anya Plutynski to be identifi ed as the authors of the editorial material in this work has been asserted in accordance with the UK Copyright, Designs, and Patents Act 1988. 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, electronic, mechanical, photocopying, recording or otherwise, except as permitted by the UK Copyright, Designs, and Patents Act 1988, without the prior permission of the publisher. Designations used by companies to distinguish their products are often claimed as trademarks. All brand names and product names used in this book are trade names, service marks, trademarks, or registered trademarks of their respective owners. The publisher is not associated with any product or vendor mentioned in this book. This publication is designed to provide accurate and authoritative information in regard to the subject matter covered. It is sold on the understanding that the publisher is not engaged in rendering professional services. If professional advice or other expert assistance is required, the services of a competent professional should be sought. First published 2008 by Blackwell Publishing Ltd 1 2008 Library of Congress Cataloging-in-Publication Data A companion to the philosophy of biology / edited by Sahotra Sarkar and Anya Plutynski. p. cm. – (Blackwell companions to philosophy ; 39) Includes bibliographical references and index. ISBN 978-1-4051-2572-7 (hardcover : alk. paper) 1. Biology–Philosophy. I. Sarkar, Sahotra. II. Plutynski, Anya. QH331.C8423 2006 570.1–dc22 2007024735 A catalogue record for this title is available from the British Library. Set in 10 on 12.5 pt Photina by SNP Best-set Typesetter Ltd., Hong Kong Printed and bound in Singapore by Utopia Press Pte Ltd The publisher’s policy is to use permanent paper from mills that operate a sustainable forestry policy, and which has been manufactured from pulp processed using acid-free and elementary chlorine-free practices. Furthermore, the publisher ensures that the text paper and cover board used have met acceptable environmental accreditation standards. For further information on Blackwell Publishing, visit our website at www.blackwellpublishing.com Contents List of Figures viii List of Tables x Notes on Contributors xi Acknowledgments xvii Introduction xviii Sahotra Sarkar and Anya Plutynski Part I Molecular Biology and Genetics 1 1 Gene Concepts 3 Hans-Jörg Rheinberger and Staffan Müller-Wille 2 Biological Information 22 Stefan Artmann 3 Heredity and Heritability 40 Richard C. Lewontin 4 Genomics, Proteomics, and Beyond 58 Sahotra Sarkar Part II Evolution 75 5 Darwinism and Neo-Darwinism 77 James G. Lennox 6 Systematics and Taxonomy 99 Marc Ereshefsky 7 Population Genetics 119 Christopher Stephens 8 The Units and Levels of Selection 138 Samir Okasha v contents 9 Molecular Evolution 157 Michael R. Dietrich 10 Speciation and Macroevolution 169 Anya Plutynski 11 Adaptationism 186 Peter Godfrey-Smith and Jon F. Wilkins Part III Developmental Biology 203 12 Phenotypic Plasticity and Reaction Norms 205 Jonathan M. Kaplan 13 Explaining the Ontogeny of Form: Philosophical Issues 223 Alan C. Love 14 Development and Evolution 248 Ron Amundson Part IV Medicine 269 15 Self and Nonself 271 Moira Howes 16 Health and Disease 287 Dominic Murphy Part V Ecology 299 17 Population Ecology 301 Mark Colyvan 18 Complexity, Diversity, and Stability 321 James Justus 19 Ecosystems 351 Kent A. Peacock 20 Biodiversity: Its Meaning and Value 368 Bryan G. Norton Part VI Mind and Behavior 391 21 Ethology, Sociobiology, and Evolutionary Psychology 393 Paul E. Griffi ths 22 Cooperation 415 J. McKenzie Alexander 23 Language and Evolution 431 Derek Bickerton vi contents Part VII Experimentation, Theory, and Themes 453 24 What is Life? 455 Mark A. Bedau 25 Experimentation 472 Marcel Weber 26 Laws and Theories 489 Marc Lange 27 Models 506 Jay Odenbaugh 28 Function and Teleology 525 Justin Garson 29 Reductionism in Biology 550 Alexander Rosenberg Index 568 vii Figures Figure 2.1 Schematic diagram of a general communication system. 24 Figure 3.1 Norms of reaction of eye size as a function of temperature for three different genotypes of Drosophila melanogaster. 42 Figure 3.2 Growth of clones of Achillea millefolium at three different elevations. 43 Figure 3.3 Four different models of the determination of phenotype. 45 Figure 3.4 Hypothetical norms of reaction for two genotypes and the phenotypic distribution resulting from variation in the environment and genetic variation. 55 Figure 6.1 Phylogenetic trees. 105 Figure 6.2 Speciation by cladogenesis and anagenesis. 108 Figure 6.3 A phylogenetic tree of lizards, snakes, crocodiles, and birds. 109 Figure 6.4 Two cladograms of the same taxa. 111 Figure 12.1 Phenotypic plasticity and reaction norms. 206 Figure 12.2 Developmental sensitivity versus developmental conversion. 210 Figure 12.3 A developmental reaction norm visualization of buffering. 216 Figure 12.4 Developmental stability versus phenotypic plasticity. 217 Figure 13.1 Different mechanisms of morphogenesis. 228 Figure 15.1 A two signal model of T helper lymphocyte activation. 277 Figure 15.2 Idiotypic interactions between antibodies. 279 Figure 21.1 The “hydraulic model” of instinctual motivation. 395 Figure 21.2 The Prisoner’s Dilemma. 401 Figure 21.3 Two versions of the Wason card selection task, one an abstract problem and the other a problem concerning social exchange. 403 Figure 22.1 The Prisoner’s Dilemma payoffs. 416 Figure 22.2 Reciprocity changes the Prisoner’s Dilemma into an Assurance Game. 421 Figure 22.3 The payoff matrix for cooperative behavior generated through byproduct mutualism. 425 Figure 22.4 The spatial prisoner’s dilemma illustrating the evolution of stable cooperative regions. 427 viii fi gures Figure 22.5 The spatial prisoner’s dilemma illustrating the evolution of stable cooperative regions.