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Prelims.fm Page ii Friday, November 10, 2006 9:45 AM Handbook of the Philosophy of Science General Editors Dov M. Gabbay Paul Thagard John Woods Cover figure taken from: Darwin’s Origin of Species, 1859 AMSTERDAM · BOSTON · HEIDELBERG · LONDON · NEW YORK · OXFORD PARIS · SAN DIEGO · SAN FRANCISCO · SINGAPORE · SYDNEY · TOKYO North-Holland is an imprint of Elsevier Prelims.fm Page iii Friday, November 10, 2006 9:45 AM Philosophy of Biology Edited by Mohan Matthen University of Toronto, Canada and Christopher Stephens University of British Columbia, Canada AMSTERDAM · BOSTON · HEIDELBERG · LONDON · NEW YORK · OXFORD PARIS · SAN DIEGO · SAN FRANCISCO · SINGAPORE · SYDNEY · TOKYO North-Holland is an imprint of Elsevier Prelims.fm Page iv Friday, November 10, 2006 9:45 AM North-Holland is an imprint of Elsevier Radarweg 29, PO Box 211, 1000 AE Amsterdam, The Netherlands The Boulevard, Langford Lane, Kidlington, Oxford OX5 1GB, UK First edition 2007 Copyright © 2007 Elsevier B.V. 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 without the prior written permission of the publisher Permissions may be sought directly from Elsevier’s Science & Technology Rights Department in Oxford, UK: phone (+44) (0) 1865 843830; fax (+44) (0) 1865 853333; email: [email protected]. Alternatively you can submit your request online by visiting the Elsevier web site at http://elsevier.com/locate/permissions, and selecting Obtaining permission to use Elsevier material Notice No responsibility is assumed by the publisher for any injury and/or damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or operation of any methods, products, instructions or ideas contained in the material herein. Because of rapid advances in the medical sciences, in particular, independent verification of diagnoses and drug dosages should be made Library of Congress Cataloging-in-Publication Data A catalog record for this book is available from the Library of Congress British Library Cataloguing in Publication Data A catalogue record for this book is available from the British Library ISBN-13: 978-0-444-51543-8 ISBN-10: 0-444-51543-7 For information on all North-Holland publications visit our website at books.elsevier.com Printed and bound in The Netherlands 07 08 09 10 11 10 9 8 7 6 5 4 3 2 1 GENERAL PREFACE Dov Gabbay, Paul Thagard, and John Woods Whenever science operates at the cutting edge of what is known, it invariably runsintophilosophicalissuesaboutthenatureofknowledgeandreality. Scientific controversies raise such questions as the relation of theory and experiment, the nature of explanation, and the extent to which science can approximate to the truth. Within particular sciences, special concerns arise about what exists and how it can be known, for example in physics about the nature of space and time, and in psychology about the nature of consciousness. Hence the philosophy of science is an essential part of the scientific investigation of the world. In recent decades, philosophy of science has become an increasingly central part of philosophy in general. Although there are still philosophers who think that theories of knowledge and reality can be developed by pure reflection, much current philosophical work finds it necessary and valuable to take into account relevant scientific findings. For example, the philosophy of mind is now closely tied to empirical psychology, and political theory often intersects with economics. Thus philosophy of science provides a valuable bridge between philosophical and scientific inquiry. More and more, the philosophy of science concerns itself not just with general issuesaboutthenatureandvalidityofscience,butespeciallywithparticularissues that arise in specific sciences. Accordingly, we have organized this Handbook into many volumes reflecting the full range of current research in the philosophy of science. We invited volume editors who are fully involved in the specific sciences, and are delighted that they have solicited contributions by scientifically-informed philosophers and (in a few cases) philosophically-informed scientists. The result is the most comprehensive review ever provided of the philosophy of science. Here are the volumes in the Handbook: Philosophy of Science: Focal Issues, edited by Theo Kuipers. Philosophy of Physics, edited by John Earman and Jeremy Butterfield. PhilosophyofBiology,editedbyMohanMatthenandChristopherStephens. Philosophy of Mathematics, edited by Andrew Irvine. Philosophy of Logic, edited by Dale Jacquette. Philosophy of Chemistry and Pharmacology, edited by Andrea Woody and Robin Hendry. vi DovGabbay,PaulThagard,andJohnWoods PhilosophyofStatistics,editedbyPrasantaS.BandyopadhyayandMalcolm Forster. Philosophy of Information, edited by Pieter Adriaans and Johan van Benthem. Philosophy of Technological Sciences, edited by Anthonie Meijers. Philosophy of Complex Systems, edited by Cliff Hooker and John Collier. Philosophy of Earth Systems Science, edited by Bryson Brown and Kent Peacock. Philosophy of Psychology and Cognitive Science, edited by Paul Thagard. Philosophy of Economics, edited by Uskali Ma¨ki. PhilosophyofLinguistics,editedbyMartinStokhofandJeroenGroenendijk. Philosophy of Anthropology and Sociology, edited by Stephen Turner and Mark Risjord. Philosophy of Medicine, edited by Fred Gifford. Detailsaboutthecontentsandpublishingscheduleofthevolumescanbefound at http://www.johnwoods.ca/HPS/. Asgeneraleditors,weareextremelygratefultothevolumeeditorsforarranging such a distinguished array of contributors and for managing their contributions. Production of these volumes has been a huge enterprise, and our warmest thanks go to Jane Spurr and Carol Woods for putting them together. Thanks also to Andy Deelen and Arjen Sevenster at Elsevier for their support and direction. CONTENTS General Preface v Dov Gabbay, Paul Thagard, and John Woods Preface xi Mohan Matthen and Christopher Stephens List of Contributors xv I. Biography Charles Darwin 1 Michael Ruse Sir Ronald Aylmer Fisher 37 Robert A. Skipper, Jr. Haldane and the Emergence of Modern Evolutionary Theory 49 Sahotra Sarkar Sewall Wright 87 James F. Crow Motoo Kimura 101 James F. Crow II. Evolution Natural Selection 111 Christopher Stephens Neutralism 129 Anya Plutynski Levels of Selection 141 Robert A. Wilson What is Evolvability? 163 Kim Sterelny viii Contents Development: Three Grades of Ontogenetic Involvement 179 D. M. Walsh Evolution and Normativity 201 Michael Bradie Evolutionary Ethics 219 Catherine Wilson III. Genetics Genetic Analysis 249 Raphael Falk The Development of Population Genetics 309 Margaret Morrison Maximisation Principles in Evolutionary Biology 335 A. W. F. Edwards Reductionism in Biology 349 Alex Rosenberg Traits, Genes, and Coding 369 Michael Wheeler IV. Taxonomy Species, Taxonomy,and Systematics 403 Marc Ereshefsky Homology and Homoplasy 429 Brian K. Hall Biological Conceptions of Race 455 Robin O. Andreasen V. Special Topics Formalisations of Evolutionary Biology 485 Paul Thompson Functions 525 Tim Lewens Biological Approaches to Mental Representation 549 Karen Neander Contents ix Innateness 567 André Ariew Artificial Life 585 Mark A. Bedau Index 605 PREFACE Philosophy of Biology has taken flight in the last quarter of the twentieth cen- tury. In very large part, of course, this is because biology itself has made huge strides in this period of time. Not only has it taken large strides, but it has done so in a way that it has made it a player in many of the largest public issues of the day: DNA, cloning, evolution, evolutionary psychology. It is not an exaggeration either to say that Darwin, cloning, and DNA now occupy as much space in academic and intellectual conversations as Shakespeare or the Bible. This hashadadramaticeffectonphilosophers. Whatkindofprocessisevolution? Are morals and meaning reducible to biology? What is the significance of genetic in- heritance? Ofthemolecularsubstrateoflife? Questionslikethesebubbleupfrom common discourse, from popular culture, from discussions in coffee shops. And philosophers have been willing and well-equipped to take them up. As recently as 1975, philosophy of biology was not generally recognized as a subfield, though texts by Michael Ruse and David Hull had appeared. Many of those whom we recognize today as the most prominent philosophers of biology were engaged in other things: Elliott Sober was writing about simplicity, Daniel C. Dennett about Rylean theories of mind, Philip Kitcher about mathematics. Virtually no University Department thought: “What we really need right now is a philosopher of biology.” Philosophical discussions of biological issues were only just beginning to go beyond the rather mechanical business of localizing debates emanating from general philosophy of science. The pioneers were in place, of course. Logical empiricists like Carl Hempel and Ernest Nagel had discussed tele- ology. David Hull and Nicolas Jardine had done some startlingly original work on biologicalsystematics. ArthurBurks, thevisionaryfromMichigan, wasprovoking interest in genetic algorithms. Marjorie Grene and Michael Ruse blazed trails in discussions of evolution (not to mention Karl Popper in his peculiarly opinion- ated way). There was desultory interest in philosophical issues surrounding the reduction of Mendelian genetics to molecular biology, though here the general as- sumptionwasthattheissueswerereallynodifferentfromthoseinthereductionof thermodynamics to statistical mechanics. Some of the philosophically interesting workinthisperiodtookplaceinhistoricaldiscussions—intheworkofsuchAris- totelian scholars as David Balme and Geoffrey Lloyd on taxonomy, for example, andinT.S.Hall’sIdeasofLifeand Matter. Andthen,ofcourse,therewerethose outside philosophy, writing in ways that were recognizably philosophical: Richard Dawkins, E. O. Wilson, Richard Lewontin, Stephen Gould, Michael Ghiselin, and