Between Zeus and the Salmon

The Biodemography of Longevity

Kenneth W. Wachter and Caleb E. Finch, Editors

Committee on Population

Commission on Behavioral and Social Sciences and Education

National Research Council

Washington, D.C.

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Between Zeus and the Salmon The Biodemography of Longevity Kenneth W. Wachter and Caleb E. Finch, Editors Committee on Population Commission on Behavioral and Social Sciences and Education National Research Council NATIONAL ACADEMY PRESS Washington, D.C. 1997

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NATIONAL ACADEMY PRESS 2101 Constitution Avenue, NW Washington, D.C. 20418 NOTICE: The project that is the subject of this report was approved by the Governing Board of the National Research Council, whose members are drawn from the councils of the National Academy of Sciences, the National Academy of Engineering, and the Institute of Medicine. The members of the committee responsible for the report were chosen for their special competences and with regard for appropriate balance. This report has been reviewed by a group other than the authors according to procedures approved by a Report Review Committee consisting of members of the National Academy of Sciences, the National Academy of Engineering, and the Institute of Medicine. 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 government on scientific and technical matters. Dr. Bruce M. Alberts 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 members. sharing with the National Academy of Sciences the responsibility for advising 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. William A. Wulf 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 Institute 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. Kenneth I. Shine 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 providing 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. Bruce M. Alberts and Dr. William A. Wulf are chairman and vice chairman, respectively, of the National Research Council. This study was supported by Contract No. N01-OD-4-2139 between the National Academy of Sciences and the National Institute on Aging. Any opinions, findings. conclusions, or recommendations expressed in this publication are those of the author(s) and do not necessarily reflect the view of the organizations or agencies that provided support for this project. Library of Congress Catalog Card No. 97-33767 International Standard Book Number 0-309-05787-6 This book is available for sale from the National Academy Press , 2101 Constitution Avenue. N.W. Box 285. Washington. D.C. 20055. Call 800-624-6242 or 202-334-3313 (in the Washington Metropolitan Area). Order electronically via Internet at Copyright 1997 by the National Academy of Sciences. All rights reserved. Printed in the United States of America.

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COMMITTEE ON POPULATION RONALD D. LEE (Chair), Department of Demography and Economics, University of California, Berkeley CAROLINE H. BLEDSOE, Department of Anthropology, Northwestern University JOSÉ LUIS BOBADILLA,* Inter-American Development Bank, Washington, D.C. JOHN BONGAARTS, The Population Council, New York JOHN B. CASTERLINE, The Population Council, New York LINDA G. MARTIN, RAND, Santa Monica, California JANE MENKEN, University of Pennsylvania ROBERT A. MOFFITT, Department of Economics, Johns Hopkins University MARK R. MONTGOMERY, The Population Council, New York W. HENRY MOSLEY, Department of Population Dynamics, Johns Hopkins University School of Hygiene and Public Health ALBERTO PALLONI, Department of Sociology, University of Wisconsin, Madison JAMES P. SMITH, RAND, Santa Monica, California BETH J. SOLDO, Department of Demography, Georgetown University MARTA TIENDA,** Population Research Center, University of Chicago AMY O. TSUI,** Carolina Population Center, University of North Carolina, Chapel Hill JOHN G. HAAGA, Director BARNEY COHEN, Program Officer TRISH DeFRISCO, Senior Project Assistant KAREN A. FOOTE, Program Officer (to July 1996) JOEL A. ROSENQUIST, Senior Project Assistant *   deceased October 1996 **   through October 1996

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CONTRIBUTORS STEVEN N. AUSTAD, University of Idaho JAMES R. CAREY, University of California, Davis CALEB E. FINCH, University of Southern California CATHERINE GRUENFELDER, University of California, Davis THOMAS E. JOHNSON, University of Colorado HILLARD KAPLAN, University of New Mexico RONALD D. LEE, University of California, Berkeley LINDA PARTRIDGE, University College London MICHAEL R. ROSE, University of California, Irvine DAVID R. SHOOK, University of Colorado SHRIPAD TULJAPURKAR, Mountain View Research, California, and Stanford University JAMES W. VAUPEL, Max Planck Institute for Demographic Research, Rostock, Germany KENNETH W. WACHTER, University of California, Berkeley ROBERT B. WALLACE, University of Iowa JOHN R. WILMOTH, University of California, Berkeley

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Preface The Committee on Population was established in 1983 by the National Research Council, under the charter of the National Academy of Sciences, to bring the knowledge and methods of the population sciences to bear on major issues of science and public policy. Much of the committee's work has concerned the demography and health of aging populations. Together with the Committee on National Statistics and the Institute of Medicine, the Committee on Population sponsored a Workshop on Forecasting Survival, Health, and Disability in 1992. Also in 1992, the committee organized a Workshop on the Demography of Aging covering a range of topics, from household and family demography, to work and retirement, intergenerational transfers, and health (Demography of Aging, 1994). A 1994 workshop examined the reasons for continued racial and ethnic differences in health and survival at older ages (Racial and Ethnic Differences in the Health of Older Americans, 1997). It was clear from all these efforts that future work on the progress of life expectancy at older ages would require genuine interdisciplinary communication between (and within) the social and life sciences. This concern led the committee, with funding from the National Institute on Aging (NIA), to organize a series of planning and discussion meetings, culminating in a workshop in Washington in April 1996, at which demographers, evolutionary theorists, genetic epidemiologists, anthropologists, and biologists from many different scientific taxa could meet and share contributions to the understanding of human longevity. These were exciting occasions, as I believe this volume, consisting of revised versions of some papers originally presented at the April workshop, attests.

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The call for interdisciplinary communication is often just a polite but inconsequential line in conclusions or book reviews, or a grandiose promise in conference proposals. Making it happen productively requires a good deal of preparation and commitment on the part of scholars, all with competing demands on their time and attention. The committee was especially fortunate to get advice and help from thoughtful and busy individuals. We would like to thank Richard Suzman of NIA, who saw the need, shared his ideas, and infected all of us with his enthusiasm. Kenneth Weiss of Pennsylvania State University was especially helpful; he was uniquely qualified to tell us what would interest scientists working in a whole range of fields, and what they knew that would interest the rest. We thank Richard Hodes of NIA for his interest and support. Harold Morowitz of George Mason University, Richard Sprott, Richard Havlik, and Evan Hadley (all of NIA), and Eric Fischer of the NRC Board on Biology gave us sound advice in the design stages and good ideas in the discussions. Elizabeth Corder of Duke University and the Stockholm Gerontology Research Center, Marcus Feldman of Stanford University, Trudy Mackay of North Carolina State University, Jennifer Madans of the National Center on Health Statistics, Randolph Nesse of the University of Michigan, Alan Rogers of the University of Utah, Burton Singer of Princeton University, and Robert Willis of the University of Michigan were generous with their ideas at the workshops and afterward. Our greatest debt is to Kenneth Wachter of the University of California, Berkeley, and Caleb Finch of the University of Southern California, who devoted time and energy to planning and chairing the workshop, guiding authors in their revisions, and editing this volume. The success of the project depended on their ability to listen and read carefully and critically, and discern what each speaker and author had to contribute to those working in different areas. Thanks also are due to Beth Soldo, who reviewed the manuscript on behalf of the committee. Colene Walden gave a helpful copy editing of the report and drafted the Glossary. Christine McShane guided the manuscript through the publication process. Karen Foote was diligent and thorough in handling the organization of the workshop and the review process. Trish DeFrisco handled all the administrative tasks for the workshop and manuscript production with her customary efficiency and good grace. Trang Ta helped with manuscript preparation and citation checking in the final stages. John Haaga, the committee staff director, wrote the proposals, supervised the process between meetings, and ably replaced Karen as project officer when she moved to the University of Illinois. Most of all, of course, we appreciate the contributions of authors and workshop participants. We trust that this volume will inspire others to join in their valuable work. RONALD D. LEE CHAIR, COMMITTEE ON POPULATION

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Contents     INTRODUCTION         1 Between Zeus and the Salmon: Introduction Kenneth W. Wachter   1     THE EMPIRICAL DEMOGRAPHY OF SURVIVAL         2 Trajectories of Mortality at Advanced Ages James W. Vaupel   17     3 In Search of Limits John R. Wilmoth   38     EVOLUTIONARY THEORY AND SENESCENCE         4 The Evolution of Senescence Shripad Tuljapurkar   65     5 Evolutionary Biology and Age-Related Mortality Linda Partridge   78     6 Toward an Evolutionary Demography Michael R. Rose   96     7 Identification and Mapping of Genes Determining Longevity Thomas E. Johnson and David R. Shook   108     THE ELDERLY IN NATURE         8 Population Biology of the Elderly James R. Carey and Catherine Gruenfelder   127     9 Postreproductive Survival Steven N. Austad   161

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    10 The Evolution of the Human Life Course Hillard Kaplan   175     11 Intergenerational Relations and the Elderly Ronald D. Lee   212     DATA FOR THE FUTURE         12 The Potential of Population Surveys for Genetic Studies Robert B. Wallace   234     CONCLUSION         13 Comparative Perspectives on Plasticity in Human Aging and Life Spans Caleb E. Finch   245     GLOSSARY   269     INDEX   275

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Between Zeus and the Salmon The Biodemography of Longevity

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