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In the Light of Evolution

Volume IV: The Human Condition



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In the Light of Evolution Volume IV: The Human Condition

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In the Light of Evolution Volume IV: The Human Condition 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 sciences, “The human Condition,” held December 11-12, 2009, 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: 978-0-309-10405-0 (hardcover) isBn-10: 0-309-10405-X (hardcover) isBn-13: 978-0-309-66786-9 (pdf) isBn-10: 0-309-66786-0 (pdf) 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: Commonly referred to as the “priest-king,” this harappan Civilization sculpture (ca. 2000–1900 B.C.) was recovered from the indus valley Mohenjo-daro archaeological site in Pakistan. The statue symbolizes the human condition as a representation of bodily form and function, and is an example of culture and the arts. image copyright J. M. Kenoyer, courtesy of the Department of Archaeology and Museums, Government of Pakistan/harappa.com. Copyright 2010 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 IV: The Human Condition xv PART I HUMAN PHYLOGENETIC HISTORY AND THE PALEONTOLOGICAL RECORD 1 1 reconstructing human evolution: Achievements, Challenges, and opportunities 5 Bernard Wood 2 Terrestrial Apes and Phylogenetic Trees 27 Juan Luis Arsuaga 3 Phylogenomic evidence of Adaptive evolution in the Ancestry of humans 47 Morris Goodman and Kirstin N. Sterner ix

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x / Contents 4 human Adaptations to Diet, subsistence, and ecoregion Are Due to subtle shifts in Allele Frequency 63 Angela M. Hancock, David B. Witonsky, Edvard Ehler, Gorka Alkorta- Aranburu, Cynthia Beall, Amha Gebremedhin, Rem Sukernik, Gerd Utermann, Jonathan Pritchard, Graham Coop, and Anna Di Rienzo 5 Working Toward a synthesis of Archaeological, linguistic, and Genetic Data for inferring African Population history 81 Laura B. Scheinfeldt, Sameer Soi, and Sarah A. Tishkoff PART II STRUCTURE AND FUNCTION OF THE HUMAN GENOME 101 6 Uniquely human evolution of sialic Acid Genetics and Biology 105 Ajit Varki 7 Bioenergetics, the origins of Complexity, and the Ascent of Man 127 Douglas C. Wallace 8 Genome-wide Patterns of Population structure and Admixture Among hispanic/latino Populations 147 Katarzyna Bryc, Christopher Velez, Tatiana Karafet, Andres Moreno-Estrada, Andy Reynolds, Adam Auton, Michael Hammer, Carlos D. Bustamante, and Harry Ostrer 9 human skin Pigmentation as an Adaptation to Uv radiation 167 Nina G. Jablonski and George Chaplin 10 Footprints of nonsentient Design inside the human Genome 185 John C. Avise PART III CULTURAL EVOLUTION AND THE UNIQUENESS OF BEING HUMAN 205 11 how Grandmother effects Plus individual variation in Frailty shape Fertility and Mortality: Guidance from human-Chimpanzee Comparisons 211 Kristen Hawkes

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Contents / xi 12 Gene-Culture Coevolution in the Age of Genomics 231 Peter J. Richerson, Robert Boyd, and Joseph Henrich 13 The Cognitive niche: Coevolution of intelligence, sociality, and language 257 Steven Pinker 14 A role for relaxed selection in the evolution of the language Capacity 275 Terrence W. Deacon 15 Adaptive specializations, social exchange, and the evolution of human intelligence 293 Leda Cosmides, H. Clark Barrett, and John Tooby 16 The Difference of Being human: Morality 319 Francisco J. Ayala references 341 index 393

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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 Dobzhan- sky, to whose fond memory this ILE series is dedicated. May Dobzhan- sky’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 IV: The Human Condition T he year 2009 marked the 200th anniversary of Charles Darwin’s birth and the 150th anniversary of his most influential publica- tion (Darwin, 1859). Darwin transformed the biological sciences in much the same way that nicolaus Copernicus, Galileo Galilei, and isaac newton had transformed the physical sciences—by demonstrat - ing that the universe operates according to natural laws that fall within the purview of rational scientific inquiry. in 1543, Copernicus published De revolutionibus orbium celestium (“on the revolutions of the Celestial spheres”) which challenged conventional wisdom that the earth was the center of Creation, and instead promoted the idea that natural laws govern the motion of physical objects in the universe. More than three centuries later, in The Origin of Species, Darwin developed the equally revolutionary concept that a natural but nonrandom process—natural selection—can yield biological adaptations that otherwise exude the superficial aura of direct craftsmanship by an intelligent agent. This book is the outgrowth of the Arthur M. sackler Colloquium “The human Condition,” which was sponsored by the national Academy of sciences on December 11–12, 2009, at the Academy’s Arnold and Mabel Beckman Center in irvine, California. it is the fourth in a series of colloquia under the umbrella title “in the light of evolution.” The first book in this series was titled In the Light of Evolution, Volume I: Adaptation and Complex Design (Avise and Ayala, 2007). The second was In the Light of Evolution, Volume II: Biodiversity and Extinction (Avise et al., 2008). The third book—In the Light of Evolution, Volume III: Two Centuries of Darwin (Avise and Ayala, xv

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xvi  /  Preface  to  In the Light of Evolution, Volume IV 2009)—presented the proceedings of a Sackler Colloquium that kicked off the bicentennial celebration of Darwin’s birth and the sesquicentennial of   The Origin of Species. The current book registers the proceedings of a Sack- ler Symposium that was timed to help close the bicentennial celebration by addressing the modern Darwinian legacy as it relates to human evolution and the human condition. Thus, the papers in this collection are devoted to “anthropogeny” (Varki et al., 2008): understanding the evolutionary origins of humans and their biological and cultural traits. Actually, Darwin barely mentioned Homo sapiens in the Origin of Spe- cies, coyly stating only that “much light will be thrown on the origin of man and his history.” More than a decade later, however, Darwin addressed human evolution at considerable length in The  Descent  of  Man  and Selection in Relation to Sex (1871a) wherein can be found many thought- ful passages, such as, “Man may be excused for feeling some pride at hav - ing risen, though not through his own exertions, to the very summit of the organic scale; and the fact of his having thus risen, instead of having been aboriginally placed there, may give him hopes for a still higher destiny in the distant future.” Of course, much has been learned about humanity’s evolutionary origins and biological conditions since Darwin’s time, not least from the evidence of paleontology, comparative vertebrate biology, and genomics. In the chapters of this book, leading evolutionary biologists and philosophers of science reflect upon and commemorate the Darwinian Revolution as it relates to the human condition at levels ranging from the molecular to the theological. Chapters in these proceedings are organized into three parts: (I) Human Phylogenetic History and the Paleontological Record, (II) Structure and Function of the Human Genome; and (III) Cul- tural Evolution and the Uniqueness of Being Human. The diverse topics addressed in these chapters give some indication of the vast breadth and depth of modern scientific research on Darwinian evolution of the human state.