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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.
