OFFSPRING

HUMAN FERTILITY BEHAVIOR IN BIODEMOGRAPHIC PERSPECTIVE

Panel for the Workshop on the Biodemography of Fertility and Family Behavior

Kenneth W. Wachter and Rodolfo A. Bulatao, Editors

Committee on Population

Division of Behavioral and Social Sciences and Education

NATIONAL RESEARCH COUNCIL
OF THE NATIONAL ACADEMIES

THE NATIONAL ACADEMIES PRESS
Washington, D.C.
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Offspring: Human Fertility Behavior in Biodemographic Perspective OFFSPRING HUMAN FERTILITY BEHAVIOR IN BIODEMOGRAPHIC PERSPECTIVE Panel for the Workshop on the Biodemography of Fertility and Family Behavior Kenneth W. Wachter and Rodolfo A. Bulatao, Editors Committee on Population Division of Behavioral and Social Sciences and Education NATIONAL RESEARCH COUNCIL OF THE NATIONAL ACADEMIES THE NATIONAL ACADEMIES PRESS Washington, D.C. www.nap.edu

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Offspring: Human Fertility Behavior in Biodemographic Perspective THE NATIONAL ACADEMIES PRESS 500 Fifth Street, N.W. Washington, DC 20001 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 study was supported by Award No. NO1-OD-4-2139, TO #71 to the National Academies from the National Institute for Child Health and Human Development. Additional funding was provided by the Andrew W. Mellon Foundation. Any opinions, findings, conclusions, or recommendations expressed in this publication are those of the author(s) and do not necessarily reflect the views of the organizations or agencies that provided support for the project. Library of Congress Cataloging-in-Publication Data Workshop on the Biodemography of Fertility and Family Behavior. Offspring : human fertility behavior in biodemographic perspective : proceedings of a workshop / Kenneth W. Wachter, Rodolfo A. Bulatao, editors. p. cm. “Panel for the Workshop on the Biodemography of Fertility and Family Behavior.” Includes bibliographical references and index. ISBN 0-309-08718-X 1. Fertility, Human—Congresses. 2. Demography—Congresses. I. Title: Human fertility behavior in biodemographic perspective. II. Wachter, Kenneth W. III. Bulatao, Rodolfo A., 1944- IV. Title. QP251.W665 2003 304.6’32—dc21 2003000077 Additional copies of this report are available from the National Academies Press, 500 Fifth Street, N.W., Lockbox 285, Washington, DC 20055; (800) 624-6242 or (202) 334-3313 (in the Washington metropolitan area); Internet, http://www.nap.edu. Copyright 2003 by the National Academy of Sciences. All rights reserved. Printed in the United States of America. Suggested citation: National Research Council. (2003). Offspring: Human Fertility Behavior in Biodemographic Perspective. Panel for the Workshop on the Biodemography of Fertility and Family Behavior. Kenneth W. Wachter and Rodolfo A. Bulatao, eds. Committeee on Population, Division of Behavioral and Social Sciences and Education. Washington, DC: The National Academies Press.

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Offspring: Human Fertility Behavior in Biodemographic Perspective THE NATIONAL ACADEMIES Advisers to the Nation on Science, Engineering, and 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. Wm. 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. 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 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. Wm. A. Wulf are chair and vice chair, respectively, of the National Research Council. www.national-academies.org

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Offspring: Human Fertility Behavior in Biodemographic Perspective PANEL FOR THE WORKSHOP ON THE BIODEMOGRAPHY OF FERTILITY AND FAMILY BEHAVIOR KENNETH W. WACHTER (Chair), Department of Demography, University of California, Berkeley JOHN N. HOBCRAFT, Population Investigation Committee, London School of Economics JEROME KAGAN, Department of Psychology, Harvard University HILLARD S. KAPLAN, Department of Anthropology, University of New Mexico HANS-PETER KOHLER, Research Group on Social Dynamics and Fertility, Max Planck Institute for Demographic Research, Rostock, Germany DAVID LAM, Population Studies Center, University of Michigan, Ann Arbor JANE MENKEN, Institute of Behavioral Sciences, University of Colorado, Boulder GERALD P. SCHATTEN, Department of Obstetrics, Gynecology and Reproductive Sciences and Department of Cell Biology and Physiology, University of Pittsburgh School of Medicine RODOLFO A. BULATAO, Study Director ANA-MARIA IGNAT, Senior Project Assistant

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Offspring: Human Fertility Behavior in Biodemographic Perspective COMMITTEE ON POPULATION (June 2000) JANE MENKEN (Chair), Institute of Behavioral Sciences, University of Colorado, Boulder ELLEN BRENNAN-GALVIN, Woodrow Wilson Center for International Scholars, Washington, DC JANET CURRIE, Department of Economics, University of California, Los Angeles JOHN N. HOBCRAFT, Population Investigation Committee, London School of Economics CHARLES B. KEELY, Institute for the Study of International Migration, Georgetown University DAVID I. KERTZER, Department of Anthropology, Brown University DAVID LAM, Population Studies Center, University of Michigan, Ann Arbor CYNTHIA B. LLOYD, Population Council, New York W. HENRY MOSLEY, Department of Population and Family Health Sciences, Johns Hopkins University ALBERTO PALLONI, Center for Demography and Ecology, University of Wisconsin, Madison JAMES W. VAUPEL, Max Planck Institute for Demographic Research, Rostock, Germany KENNETH W. WACHTER, Department of Demography, University of California, Berkeley LINDA J. WAITE, Population Research Center, University of Chicago BARNEY COHEN, Director

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Offspring: Human Fertility Behavior in Biodemographic Perspective Preface Having children is a biological imperative for the survival of a group, and a wanted child can be a biological coup for a couple, a supreme biological moment—of possible peril and triumph—for a woman. Yet demographic interest in fertility has, for decades, been driven less by such concerns than by the threat of rapid population growth and the issue, for couples and women, of unplanned fertility. This volume seeks to encourage more balance and depth in the treatment of fertility in population studies. It suggests that many fertility behaviors that concern demographers may follow biodemographic templates, are influenced by genetic endowment, are triggered through hormonal pathways, and have been shaped in specific directions in the course of human evolution. Since the middle of the 20th century, the contrast between small families in developed countries and large families and burgeoning populations in developing countries has fueled concern about deepening poverty, global inequality, and escalating environmental burdens. These concerns have not disappeared, as global population continues to expand. Yet much of the expansion is now effectively a hangover from the high-fertility decades, and many countries, both developed and developing, have entered an era of subreplacement fertility. The confluence of the largely successful (though still incomplete) worldwide effort to tame high fertility and the emergence of genetic approaches to understanding human behavior provide a stimulus to review the focus of demographic work on fertility, to enlarge its concerns with biological and evolutionary questions. The Committee on Population took on this task with the understanding that researchers in the area were pursuing a variety

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Offspring: Human Fertility Behavior in Biodemographic Perspective of independent approaches that required a common, unifying focus. Some work in the area, such as studies of twins and adoptees, has also generated controversy, despite being pursued aggressively. With advice from the Committee on Population, the Board on Life Sciences, and the Institute of Medicine’s Board on Neuroscience and Behavioral Health, the National Research Council (NRC) appointed a panel to organize a workshop on the topic. This volume is the result. The papers contained in this volume were presented at the Workshop on the Biodemography of Fertility and Family Behavior, held at the National Academies in Washington, D.C., in June 2002, building on a preliminary meeting in February 2002 at the Beckman Center in Irvine, California. The workshop in a sense complemented an earlier workshop in April 1996 on biodemographic aspects of longevity (published as Between Zeus and the Salmon: The Biodemography of Longevity). Both workshops brought together demographers, evolutionary theorists, geneticists, and biologists to consider questions at the interface between the social sciences and the life sciences. The papers were subsequently reviewed in draft form by individuals chosen for their diverse perspectives and technical expertise, in accordance with procedures approved by the NRC’s Report Review Committee. The purpose of this independent review was to provide candid and critical comments to assist the institution in making the published volume as sound as possible and to ensure that the volume meets institutional standards for objectivity, evidence, and responsiveness to the study charge. We thank the following individuals for contributing to the review: Nicholas G. Blurton Jones (University of California at Los Angeles), Sue Carter (University of Illinois at Chicago), Peter T. Ellison (Harvard University), John Haaga (Population Reference Bureau), Jennifer Harris (National Institute on Aging), Kristin Hawkes (University of Utah), Jerome Kagan (Harvard University), John J. Lepri (University of North Carolina at Greensboro), Kimber McKay (University of Montana), Monique Borgerhoff Mulder (University of California, Davis), Gerald P. Schatten (University of Pittsburgh School of Medicine), Barbara Smuts (University of Michigan, Ann Arbor), Kim Wallen (Emory University), and Maxine Weinstein (Georgetown University). The review of the entire volume was overseen by Michael Murphy of the London School of Economics. Although these individuals provided constructive comments and suggestions, responsibility for the content of this volume rests entirely with the authors and the institution. Work on this project was stimulated and encouraged by Christine Bachrach of the National Institute for Child Health and Human Development and was supported under a contract with the institute. We are grateful for her continued attention to the core issues that the volume addresses and

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Offspring: Human Fertility Behavior in Biodemographic Perspective to raising many questions that helped shape and enrich the volume. Additional funding was provided by the Andrew W. Mellon Foundation. We are especially grateful to Kenneth Wachter, who chaired the panel that organized the workshop, providing the spark and vital judgments about appropriate participants. The other members of the panel that organized the workshop each made important contributions in helping delineate relevant topics, identify participants, and critique their contributions, and in some cases writing papers themselves. We also wish to thank Randy Bulatao who, as study director, enriched the workshop with broad perspectives and guided the endeavor through many challenges. Ana-Maria Ignat provided active staff support. Barbara Bodling O’Hare skillfully edited the report and Yvonne Wise guided the manuscript through the publication process. Jane Menken, Chair Committee on Population

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Offspring: Human Fertility Behavior in Biodemographic Perspective Contents 1   Biodemography of Fertility and Family Formation Kenneth W. Wachter   1 2   Genetic Influences on Fertility: Strengths and Limitations of Quantitative Inferences Michael L. Rutter   18 3   Education, Fertility, and Heritability: Explaining a Paradox Hans-Peter Kohler and Joseph L. Rodgers   46 4   The Neural Basis of Pair Bonding in a Monogamous Species: A Model for Understanding the Biological Basis of Human Behavior Larry J. Young   91 5   Hormonal Mediation of Physiological and Behavioral Processes That Influence Fertility Judy L. Cameron   104 6   Intraspecific Variability in Fertility and Offspring Survival in a Nonhuman Primate: Behavioral Control of Ecological and Social Sources Jeanne Altmann and Susan C. Alberts   140

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Offspring: Human Fertility Behavior in Biodemographic Perspective 7   An Evolutionary and Ecological Analysis of Human Fertility, Mating Patterns, and Parental Investment Hillard S. Kaplan and Jane B. Lancaster   170 8   Sexually Antagonistic Coevolution: Theory, Evidence, and Implications for Patterns of Human Mating and Fertility Steven W. Gangestad   224 9   Pubertal Maturation, Andrenarche, and the Onset of Reproduction in Human Males Benjamin Campbell   260 10   Energetics, Sociality, and Human Reproduction: Life History Theory in Real Life Carol M. Worthman   289 11   Evolutionary Biology and Rational Choice in Models of Fertility David Lam   322 12   Reflections on Demographic, Evolutionary, and Genetic Approaches to the Study of Human Reproductive Behavior John N. Hobcraft   339     Contributors and Other Workshop Participants   358     Index   365

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Offspring: Human Fertility Behavior in Biodemographic Perspective Tables and Figures TABLES 3-1   Number of Twins by Birth Cohort,   60 3-2   Completed Education for Females and Males by Birth Cohort,   61 3-3   Years of Primary and Secondary Schooling,   63 3-4   Number of Children Born Until End of 1998,   63 3-5   Females—Bivariate Ordered Probit Estimation for Number of Children,   71 3-6   Males—Bivariate Ordered Probit Estimation for Number of Children,   80 6-1   Per Annum Vital Rates for Wild-Foraging Amboseli Baboon Females (1971-1999) by One-Year Age Classes for Analysis in Projection Matrix Models,   145 6-2   Per Annum Vital Rates for Wild-Foraging Amboseli Baboon Males (1971-1999) by One-Year Age Classes for Analysis in Projection Matrix Models,   146 6-3   Impact of Foraging Environment on Various Life History Components,   154 6-4   Impact of Ecological and Social Factors on Various Life History Components,   155 7-1   Regressions for Percent of Families That Are Polygynous Among Foragers,   187 7-2   The Wives of Despots,   194 10-1   Humans’ Expectable Environments of Rearing,   313

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Offspring: Human Fertility Behavior in Biodemographic Perspective FIGURES 3-1   Bivariate behavioral genetics model for education and fertility, females,   66 3-2   Bivariate behavioral genetics model for education and fertility, males,   67 3-3   Cohort and education effects on within-MZ correlation,   76 5-1   Schematic diagram of the hypothalamic-pituitary-gonadal axis,   107 5-2   Diagrammatic representation of changes in plasma levels of estradiol, progesterone, LH, FSH, and portal levels of GnRH over the human menstrual cycle,   109 5-3   Examples of the pulsatile pattern of LH secretion in a woman during the late follicular phase (A) and midluteal phase (B) of the menstrual cycle,   110 5-4   Schematic diagram of factors that mediate the effects of psychological stress on the activity of the hypothalamic-pituitary-gonadal axis,   128 6-1   Duration of several life history stages as a function of body size in anthropoid primates,   141 6-2   Proportion of elasticity in the matrix model attributable to immature survival, fertility, and adult survival,   149 6-3   Projected population growth (or relative fitness) for different levels of first-year survivorship holding all other survival and fertility values to those in Table 6-1,   151 6-4   Amboseli study groups that contributed data to the present analyses,   152 6-5   High density results in lower reproductive rate (longer interbirth intervals following a surviving offspring),   157 6-6   Age of reproductive maturity and reproductive rate (measured as in Figure 6-5) are functions of social status,   159 6-7   Daughters’ social status as predicted by that of their mothers,   160 6-8   Overview of behaviors affecting fertility and offspring survival by fully wild-foraging baboons in Amboseli over three decades,   161 7-1   Production as a function of the capital stock,   176 7-2   Offspring viability isoclines (indifference curves) as a function of male and female inputs,   177

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Offspring: Human Fertility Behavior in Biodemographic Perspective 7-3   Net food production and survival: Human foragers and chimpanzees,   181 7-4   Frequency distribution of percentage of marriages that are polygynous among foragers,   184 7-5   Father’s probability of ceasing to live with his child before the age of 6,   204 7-6   Living arrangements of I.D. Mkize High School students in Guguletu, South Africa,   206 9-1   Temporal relationship of adrenarche and puberty in human males,   263 9-2   Causal relationship of adrenarche and pubertal maturation,   267 9-3   Schematic representation of the impact of gonadal and adrenal hormones on reproductive strategies,   270 9-4   Conceptual model of biological dimension of adolescent sexual behavior,   273 10-1   Scheduling in timing of reproductive events in diverse human populations,   292 10-2   Secular trend in timing of puberty in terms of life history,   299 10-3   Representative endocrine axes, pathways, and interrelations, with particular regard to resource partitioning,   301 11-1   Number of live births by years of schooling for married women, ages 45 to 54, in Brazil 1984, and for black South Africans, 1995-1998,   329 11-2   Employment rates by years of schooling for married women, ages 35 to 44, in Brazil, 1984, and for black South Africans, 1995-1998,   329 11-3   Natural logarithm of monthly wages by years of schooling (relative to women with zero schooling) for married women, ages 35 to 44, with positive wages, in Brazil, 1984, and for black South Africans, 1995-1998,   330 11-4   Mean years of schooling of 15-year-olds by mother’s education, in Brazil, 1984, and for black South Africans, 1995-1998,   330

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