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In the Light of Evolution Volume V: Cooperation and Conflict
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In the Light of Evolution Volume V: Cooperation and Conflict JOAN E. STRASSMANN, DAVID C. QUELLER, JOHN C. AVISE, and FRANCISCO J. AYALA, Editors THE NATIONAL ACADEMIES PRESS Washington, D.C. www.nap.edu
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THE NATIONAL ACADEMIES PRESS 500 Fifth Street., N.W. Washington, DC 20001 This volume is based on the Arthur M. Sackler Colloquium of the National Academy of Sci - ences, “Cooperation and Conflict,” held January 7-8, 2011, at the Arnold and Mabel Beckman Center of the National Academies of Sciences and Engineering in Irvine, California. The articles appearing in these pages were contributed by speakers at the colloquium and have been anonymously reviewed. Any opinions, findings, conclusions, or recommendations expressed in this volume are those of the authors and do not necessarily reflect the view of the National Academy of Sciences. In the light of evolution / John C. Avise and Francisco J. Ayala, editors. p. cm. Vol. I based on a colloquium of the National Academy of Sciences, held December 1–2, 2006, in Irvine, California. Includes bibliographical references. ISBN-13: 978-0-309-21836-8 ISBN-10: 0-309-21836-5 1. Evolution (Biology)—Congresses. I. Avise, John C, 1948–. II . Ayala, Francisco José, 1934– III . National Academy of Sciences (U.S.) QH359.I55 2007 576.8—dc22 2007032455 Additional copies of this book are available from the National Academies Press, 500 Fifth St., N.W., Lockbox 285, Washington, DC 10055; (800) 624-6242 or (202) 334-3313 (in the Washington metropolitan area); Internet, http://www.nap.edu. Cover image: Pictured is a Batik painting in woad, a plant-derived dye, on cotton. The two ants illustrate the phenomenon of cooperation, the focus of many of the chapters in this vol - ume. This collection of articles explores recent developments in the study of the evolution of cooperation among all organisms from the level of genes to that to societies—from bacteria to humans. Image courtesy of Robin Paris, www.robinparis.co.uk. Copyright 2011 by the National Academy of Sciences. All rights reserved. Printed in the United States of America
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The National Academy of Sciences is a private, nonprofit, self-perpetuating society of distinguished scholars engaged in scientific and engineering research, dedicated to the furtherance of science and technology and to their use for the general welfare. Upon the authority of the charter granted to it by the Congress in 1863, the Academy has a mandate that requires it to advise the federal govern- ment on scientific and technical matters. Dr. Ralph J. Cicerone is president of the National Academy of Sciences. The National Academy of Engineering was established in 1964, under the charter of the National Academy of Sciences, as a parallel organization of outstanding engineers. It is autonomous in its administration and in the selection of its mem - bers, sharing with the National Academy of Sciences the responsibility for advis - ing the federal government. The National Academy of Engineering also sponsors engineering programs aimed at meeting national needs, encourages education and research, and recognizes the superior achievements of engineers. Dr. Charles M. Vest is president of the National Academy of Engineering. The Institute of Medicine was established in 1970 by the National Academy of Sciences to secure the services of eminent members of appropriate professions in the examination of policy matters pertaining to the health of the public. The Insti - tute acts under the responsibility given to the National Academy of Sciences by its congressional charter to be an adviser to the federal government and, upon its own initiative, to identify issues of medical care, research, and education. Dr. Harvey V. Fineberg is president of the Institute of Medicine. The National Research Council was organized by the National Academy of Sciences in 1916 to associate the broad community of science and technology with the Academy’s purposes of furthering knowledge and advising the federal government. Functioning in accordance with general policies determined by the Academy, the Council has become the principal operating agency of both the National Academy of Sciences and the National Academy of Engineering in pro - viding services to the government, the public, and the scientific and engineering communities. The Council is administered jointly by both Academies and the Institute of Medicine. Dr. Ralph J. Cicerone and Dr. Charles M. Vest are chair and vice chair, respectively, of the National Research Council. www.national-academies.org
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Arthur M. Sackler, M.D. 1913–1987 Born in Brooklyn, New York, Arthur M. Sackler was educated in the arts, sciences, and humanities at New York University. These interests remained the focus of his life, as he became widely known as a scientist, art collector, and philanthropist, endowing institutions of learning and culture throughout the world. He felt that his fundamental role was as a doctor, a vocation he decided upon at the age of four. After completing his internship and service as house physician at Lincoln Hospital in New York City, he became a resident in psychiatry at Creedmoor State Hospital. There, in the 1940s, he started research that resulted in more than 150 papers in neuroendocri - nology, psychiatry, and experimental medicine. He considered his scien - tific research in the metabolic basis of schizophrenia his most significant contribution to science and served as editor of the Journal of Clinical and Experimental Psychobiology from 1950 to 1962. In 1960 he started publica- tion of Medical Tribune, a weekly medical newspaper that reached over one million readers in 20 countries. He established the Laboratories for Therapeutic Research in 1938, a facility in New York for basic research that he directed until 1983. As a generous benefactor to the causes of medicine and basic science, Arthur Sackler built and contributed to a wide range of scientific insti- tutions: the Sackler School of Medicine established in 1972 at Tel Aviv University, Tel Aviv, Israel; the Sackler Institute of Graduate Biomedical Science at New York University, founded in 1980; the Arthur M. Sackler Science Center dedicated in 1985 at Clark University, Worcester, Massachu- setts; and the Sackler School of Graduate Biomedical Sciences, established in 1980, and the Arthur M. Sackler Center for Health Communications, established in 1986, both at Tufts University, Boston, Massachusetts. His pre-eminence in the art world is already legendary. According to his wife Jillian, one of his favorite relaxations was to visit museums and art galleries and pick out great pieces others had overlooked. His interest in art is reflected in his philanthropy; he endowed galleries at the Metro- politan Museum of Art and Princeton University, a museum at Harvard vii
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University, and the Arthur M. Sackler Gallery of Asian Art in Washing- ton, D.C. True to his oft-stated determination to create bridges between peoples, he offered to build a teaching museum in China, which Jillian made possible after his death, and in 1993 opened the Arthur M. Sackler Museum of Art and Archaeology at Peking University in Beijing. In a world that often sees science and art as two separate cultures, Arthur Sackler saw them as inextricably related. In a speech given at the State University of New York at Stony Brook, Some reflections on the arts, sciences and humanities, a year before his death, he observed: ‘‘Communi- cation is, for me, the primum movens of all culture. In the arts . . . I find the emotional component most moving. In science, it is the intellectual content. Both are deeply interlinked in the humanities.’’ The Arthur M. Sackler Colloquia at the National Academy of Sciences pay tribute to this faith in communication as the prime mover of knowledge and culture. viii
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Contents Arthur M. Sackler Biography vii Preface to the In the Light of Evolution Series xiii Preface to In the Light of Evoluion V: Cooperation and Conflict xv PART I THE FUNDAMENTALS OF EVOLUTIONARY COOPERATION 1 1 Expanded Social Fitness and Hamilton’s Rule for Kin, Kith, and Kind 5 David C. Queller 2 Evolutionary Transitions in Bacterial Symbiosis 27 Joel L. Sachs, Ryan G. Skophammer, and John U. Regus 3 Kinship, Greenbeards, and Runaway Social Selection in the Evolution of Social Insect Cooperation 49 Peter Nonacs 4 Spatiotemporal Environmental Variation, Risk Aversion, and the Evolution of Cooperative Breeding as a Bet-Hedging Strategy 69 Dustin R. Rubenstein ix
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x / Contents PART II COOPERATION WRIT SMALL: MICROBES 87 5 Endemic Social Diversity Within Natural Kin Groups of a Cooperative Bacterium 91 Susanne A. Kraemer and Gregory J. Velicer 6 Evolution of Restraint in a Structured Rock–Paper–Scissors Community 117 Joshua R. Nahum, Brittany N. Harding, and Benjamin Kerr 7 Social Evolution in Multispecies Biofilms 137 Sara Mitri, João B. Xavier, and Kevin R. Foster PART III REAL SELFISH (AND COOPERATIVE) GENES 165 8 Molecular Evolutionary Analyses of Insect Societies 167 Brielle J. Fischman, S. Hollis Woodard, and Gene E. Robinson 9 Evolution of Cooperation and Control of Cheating in a Social Microbe 191 Joan E. Strassmann and David C. Queller 10 Selfish Genetic Elements, Genetic Conflict, and Evolutionary Innovation 213 John H. Werren PART IV SOCIALITY AND MEDICINE 235 11 The Evolution of Drug Resistance and the Curious Orthodoxy of Aggressive Chemotherapy 237 Andrew F. Read, Troy Day, and Silvie Huijben 12 Genomic Imprinting and the Evolutionary Psychology of Human Kinship 253 David Haig 13 Pathology from Evolutionary Conflict, with a Theory of X Chromosome Versus Autosome Conflict over Sexually Antagonistic Traits 275 Steven A. Frank and Bernard J. Crespi
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Contents / xi PART V ARE HUMANS DIFFERENT? 299 14 Cooperation and Competition in a Cliff-Dwelling People 303 Beverly I. Strassmann 15 Extent and Limits of Cooperation in Animals 325 Dorothy L. Cheney 16 Evolutionary Foundations of Human Prosocial Sentiments 343 Joan B. Silk and Bailey R. House 17 The Cultural Niche: Why Social Learning Is Essential for Human Adaptation 363 Robert Boyd, Peter J. Richerson, and Joseph Henrich References 383 Index 437
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Preface to the In the Light of Evolution Series B iodiversity—the genetic variety of life—is an exuberant product of the evolutionary past, a vast human-supportive resource (aesthetic, intellectual, and material) of the present, and a rich legacy to cher- ish and preserve for the future. Two urgent challenges, and opportunities, for 21st-century science are to gain deeper insights into the evolutionary processes that foster biotic diversity, and to translate that understanding into workable solutions for the regional and global crises that biodiver- sity currently faces. A grasp of evolutionary principles and processes is important in other societal arenas as well, such as education, medicine, sociology, and other applied fields including agriculture, pharmacology, and biotechnology. The ramifications of evolutionary thought also extend into learned realms traditionally reserved for philosophy and religion. In 1973, Theodosius Dobzhansky penned a short commentary entitled “Nothing in biology makes sense except in the light of evolution.” Most scientists agree that evolution provides the unifying framework for inter- preting biological phenomena that otherwise can often seem unrelated and perhaps unintelligible. Given the central position of evolutionary thought in biology, it is sadly ironic that evolutionary perspectives outside the sciences have often been neglected, misunderstood, or purposefully misrepresented. The central goal of the In the Light of Evolution (ILE) series is to pro- mote the evolutionary sciences through state-of-the-art colloquia—in the series of Arthur M. Sackler colloquia sponsored by the National Academy of Sciences—and their published proceedings. Each installment explores xiii
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xiv / Preface to the In the Light of Evolution Series evolutionary perspectives on a particular biological topic that is scientifi - cally intriguing but also has special relevance to contemporary societal issues or challenges. Individually and collectively, the ILE series aims to interpret phenomena in various areas of biology through the lens of evolution, address some of the most intellectually engaging as well as pragmatically important societal issues of our times, and foster a greater appreciation of evolutionary biology as a consolidating foundation for the life sciences. The organizers and founding editors of this effort (Avise and Ayala) are the academic grandson and son, respectively, of Theodosius Dobzhansky, to whose fond memory this ILE series is dedicated. May Dobzhansky’s words and insights continue to inspire rational scientific inquiry into nature’s marvelous operations. John C. Avise and Francisco J. Ayala Department of Ecology and Evolutionary Biology, University of California, Irvine (January 2007)
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Preface to In the Light of Evolution, Volume V: Cooperation and Conflict T his book is the outgrowth of the Arthur M. Sackler Colloquium “Cooperation and Conflict,” which was sponsored by the National Academy of Sciences on January 7–8, 2011, at the Academy’s Arnold and Mabel Beckman Center in Irvine, California. It is the fifth in a series of colloquia under the general title “In the Light of Evolution.” The first four books in this series were titled Adaptation and Complex Design (Avise and Ayala, 2007), Biodiversity and Extinction (Avise et al., 2008), Two Centuries of Darwin (Avise and Ayala, 2009a), and The Human Condition (Avise and Ayala, 2009b). The current volume explores recent developments in the study of cooperation and conflict, ranging from the level of the gene to societies and symbioses. Any student of history knows that we humans can be a vicious lot, but paradoxically we are also among nature’s great cooperators. Which of us, as an individual, can manufacture a cell phone or an airplane? Even our great conflicts—wars—are extremely cooperative endeavors on each side. Some of this cooperation is best understood culturally, but we are also products of evolution, with bodies, brains, and behaviors molded by natural selection. How cooperation evolves has been one of the big ques - tions in evolutionary biology, and how it pays or does not pay is a great intellectual puzzle. If nothing makes sense in biology except in the light of evolution (Dobzhansky, 1973), then for the first century after Darwin, cooperation and altruism did not make much sense. We could see that individual organisms sometimes helped others, even at a cost to their own fitness. It xv
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xvi / Preface to In the Light of Evolution, Volume V was clear that such behavior could benefit the group or the population, the species, or even other species and whole communities. However, it was not obvious how such effects would be heritable. All our mathematical models—the hard work of the modern synthesis—were about individuals with one allele out-reproducing those with an alternative. This process would favor individuals with higher reproduction but would not be expected to produce self-sacrifice. Yet, apparent cooperation was routinely attributed to the good of the group, species, or community. This situation changed in the first decade of Darwinism’s second century. William D. Hamilton (1964a,b) argued that cooperation was important in nature, and that social evolution could be understood in terms of direct gains to the actor’s own fitness or indirect benefits to the fitness of others who share the cooperation allele. There followed an intense period of exploring the indirect effects of cooperation and altruism, reinterpreting sexual selec - tion and many other phenomena in terms of individual advantage, and understanding frequency-dependent effects via game theory, efforts that continue to the present. The puzzle of cooperation was the dominant theme of research in the early years, whereas recent work has emphasized its importance and ubiquity. Far from being a rare trait shown by social insects and a few others, cooperation is both widespread taxonomically and essential to life. Major transitions in the hierarchy of life have often involved cooperation among lower-level units to the point where they evolve into higher-level organisms (Buss, 1987; Maynard Smith and Szathmary, 1995). Examples include the assembly of the eukaryotic cell with its symbiotic organelles, the evolution of multicellular organisms, and the organismal colonies of some social insects. Organisms are, at multiple levels, those units that have evolved to have, within their boundaries, extreme cooperation and minimal conflict (Queller and Strassmann, 2009; Strassmann and Queller, 2010). The depth of research on cooperation and conflict has increased greatly, most notably in the direction of small organisms. Microbes turn out to have highly developed cooperation (West et al., 2007a), and they, along with other model organisms, have proven instrumental in beginning to understand sociality at the genetic and molecular levels, the study of real selfish genes (Santorelli et al., 2008). The social evolution approach has given us new insights on diseases often caused by microbes (Foster, 2005). At the other end of the spectrum, we are getting a much better under- standing of the cooperation and conflict that matter most to our species (Alexander, 1979). Cooperation has been central to humanity’s spectacular success and will be central to our short-term and long-term fate.