Handbook of Behavior Genetics

Yong-Kyu KimEditor

Handbook of BehaviorGenetics

123

EditorYong-Kyu KimUniversity of GeorgiaAthens, GAUSAyongkyu@arches.uga.edu

ISBN 978-0-387-76726-0 e-ISBN 978-0-387-76727-7DOI 10.1007/978-0-387-76727-7

Library of Congress Control Number: 2008941695

c© Springer Science+Business Media, LLC 2009All rights reserved. This work may not be translated or copied in whole or in part without the written permission of thepublisher (Springer Science+Business Media, LLC, 233 Spring Street, New York, NY 10013, USA), except for briefexcerpts in connection with reviews or scholarly analysis. Use in connection with any form of information storage andretrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafterdeveloped is forbidden.The use in this publication of trade names, trademarks, service marks, and similar terms, even if they are not identifiedas such, is not to be taken as an expression of opinion as to whether or not they are subject to proprietary rights.While the advice and information in this book are believed to be true and accurate at the date of going to press, neitherthe authors nor the editors nor the publisher can accept any legal responsibility for any errors or omissions that maybe made. The publisher makes no warranty, express or implied, with respect to the material contained herein.

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Dedicated to my teachers

Preface

Behavior genetics is an interdisciplinary area combining the behavioral sciences and genetics.The study of behavior genetics has become increasingly important as we see growth spurtsin finding genes involved in complex behaviors following on advances in molecular genetictechniques. This domain has been growing rapidly since the 1970s and increasingly receivesattention from many different disciplines. It has now become a vast common ground for sci-entists from very diverse fields including psychology, psychiatry, neurology, endocrinology,biochemistry, neuroimaging, and genetics.

When I was invited to organize this book by Springer, I was preparing for a new course,Behavior Genetics, at the University of Georgia in fall, 2005. Only a few textbooks wereavailable at that time, but I could not find good references for graduate students and scien-tists. I thought that we needed to offer research guides to the studies of genetic and environ-mental influences on a variety of complex behaviors in humans and animals. I had little ideaabout the proper scope for such a book. I contacted senior colleagues of the Behavior Genet-ics Association and they gave me excellent advice. I initially invited contributors who werelargely members of the Behavior Genetics Association and the handbook was outlined with14 chapters. As the Handbook developed, it became clear that the first draft was not sufficientto cover all important domains in behavior genetics. In the second meeting with contributorsduring the BGA meeting in Hollywood, CA, we discussed expanding the handbook to otherrelated domains, such as evolutionary psychology, health behavior, and neurosciences. I invitedadditional contributors from other disciplines, and added chapters on the history of behaviorgenetics, quantitative methods and models, as well as more studies of animal models. Now thehandbook stands with 34 chapters and integrates many of the basic issues in behavior genetics.In each chapter, current research and issues on the selected topics are intensively reviewedand directions for future research on these topics are highlighted: new research designs, ana-lytic methods, and their implications are addressed. It is anticipated that the handbook willcontribute to our understanding of behavior genetics and future research endeavors in the 21stcentury.

Chapter 1 addresses a history of behavior genetics going back to some of Plato’s ideas anddiscusses the nature–nurture controversies on behavior in the modern era which sometimesbrought about uproar in our community. In Part I, we address designs and methods in behaviorgenetic research. Chapters 2 and 4 introduce statistical models and analyses, i.e., biometricalmodels and multivariate genetic analyses, which explain genetic and environmental causesof covariation between quantitative traits and comorbidity between disorders. In Chapter 3,quantitative trait locus (QTL) analysis is introduced and methods of linkage and associationmapping of continuous traits are discussed. Results of the QTL analyses in several quantitativetraits are presented throughout this volume. Chapter 5 addresses the importance of animalsas models of human behaviors – cognition, personality, and pathology are presented in thisvolume.

Part II addresses the genetics of cognition in humans and animals with nine selected top-ics. Chapter 6 discusses genetic and environmental influences on general intelligence using

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twin studies, followed by new twin research designs, analytic methods, findings, and theirimplications. In Chapter 7, behavioral genetic research on cognitive aging is reviewed: geneticand environmental contributions to age-related changes in cognitive abilities; contributions ofgenes and lifestyle variables to dementia, and to the terminal decline in cognitive functioning;and quantitative methods for investigating cognitive aging are presented. Chapter 8 addressesbehavioral genetic research on reading, and the genetic and environmental etiologies of read-ing ability and disability are discussed. Chapter 9 explores behavioral and molecular geneticstudies elucidating the role of the genome in the development and manifestation of disordersof speech and language. The human brain continues to show dynamic changes from child-hood into adulthood. Genetic and environmental influences in brain volumes are addressed inChapter 10. Using quantitative magnetic resonance imaging (MRI), brain structures in patientswith a clear genetic etiology are reviewed. Genetic approaches to the search for genes asso-ciated with brain volume are discussed. Cognitive abilities in animals as models of humanbehavior are presented in Chapters 11, 12, 13, and 14. Quantitative and molecular geneticapproaches to cognition research in rodents are presented in Chapter 11. Cognitive deficitsaffected by genetic manipulations and mouse models for human cognitive disabilities are dis-cussed. Specifically, Chapter 12 reviews human cognitive impairment associated with chromo-somal abnormalities, and mouse models of trisomy 21 are discussed addressing the relation-ships among genes, brain, and cognitive function. Drosophila (fruit fly) models of Alzheimer’sdisease are introduced in Chapter 13. Pathological roles of Aβ peptides in fly brains, memorydefects, and locomotor dysfunctions are discussed. Chapter 14 addresses Drosophila courtshipsongs which are utilized for intersexual selection and species recognition in nature. Quantita-tive and molecular genetic studies on the phylogenetic patterns of song evolution in differentspecies groups are reviewed.

In Part III, the genetics of personality in humans and animals is addressed with 10selected topics. Personality is influenced by both genes and environment during development.Chapter 15 explores genotype–environment correlation through a review of the behavioralgenetic literature on genetic and environmental influences on family relationships. It is veryimportant that behavioral genetic models that measure behaviors of interest reflect the contentof the domains. Chapter 16 reviews behavioral genetic methods and models for personalityresearch and theory, and addresses some methodological issues. Chapter 17 addresses the rolesof specific genes, i.e., DRD4 and 5-HTTLPR genes, contributing to the multifaceted dimen-sions of human personality, including altruism. Temperament, developing early in life andpossibly forming the basis for later personality and psychopathology, is explored in Chapter 18in which quantitative and molecular genetic findings, as well as endophenotypic approaches,are discussed. Sexual orientation is a controversial issue in our communities. A growing bodyof evidence suggests that familial and genetic factors affect human sexual orientation. Quan-titative and molecular genetic studies on sexual orientation are reviewed in Chapter 19. Threechapters introduce animal models of personality and aggression. Chapter 20 explores personal-ity differences in rats widely used in laboratories and discusses anatomical and neurochemicalanalyses in this endeavor. Behavioral and genetic research on offensive aggression in mice isreviewed and comparative genetic studies of aggression across species are addressed in Chap-ter 21. Chapter 22 discusses aggressive behavior in fruit flies from the ecological, genetic,neurological, and evolutionary perspectives. Approximately 10% of the population are left-handers. The history, determination, and etiology of handedness are addressed in Chapter 23.Chapter 24 introduces exercise behavior as a new discipline in behavior genetics. A large pro-portion of adults in the world do not regularly engage in exercise, although benefits of exerciseare well documented. Genetic determinants of variability in exercise behavior are discussed.

In Part IV the genetics of psychopathology is represented with nine selected topics. Somepsychiatric disorders like ADHD are only diagnosed by questionnaires or psychiatric inter-views, rather than by clinical tests, and consequently the genetic studies of the disorderscan vary as a function of applied assessment methods and informants. Chapter 25 addressessuch behavioral measure issues concerning ADHD. Depression and anxiety have their origins

Preface ix

in childhood and arise from genetic and shared environmental effects. Epidemiological andbehavior genetic research on childhood depression and anxiety are discussed in Chapter 26.Autism is familial and, thus, relatives of probands with autism are at high risk for depres-sion, anxiety, and personality attributes. Chapter 27 reviews current findings in the geneticepidemiology of autism and its etiological issues concerning the definition of autism pheno-types are discussed. Two chapters address substance abuse behaviors, that is, smoking, drugs,and drinking. Smoking behaviors aggregate in families and in peer networks due to geneticdispositions and common environmental influences. Chapter 28 reviews behavioral geneticresearch on smoking behavior and nicotine dependence, using Finnish sample studies, and itscomorbidities with other substance use, depression, and schizophrenia are discussed. Behav-ioral and molecular genetic research on the use and abuse of both alcohol and drugs is reviewedin Chapter 29. Substance abuse and substance use disorder co-occur with conduct disorder andantisocial behavior. Chapter 30 gives results of a meta-analysis of twin and adoption studiesexamining genetic and environmental influences on conduct disorder and antisocial behavior.Association and linkage studies for genes influencing antisocial behavior are discussed. Chap-ter 31 explores the behavioral and molecular genetic approaches to the origins of two majorpsychoses: schizophrenia and bipolar mood disorder. The concept of endophenotypes, whichare measured intermediate traits or states between genotypes (genetic liability) and pheno-types (disorders), is discussed. Chapter 32 discusses indepth longitudinal ”high-risk” studiesthat intend to identify endophenotypes in the first-degree relatives of schizophrenic probandsand to offer putative behavioral predictors of future schizophrenia spectrum disorders. Mousemodels of cognitive dysfunctions in schizophrenia are explored in Chapter 33 where the roleof dopamine in attention and working memory is discussed. Finally, in Chapter 34, futuredirections for behavior genetics are addressed.

It is not surprising that, at the final publication date of a book like the Handbook of BehaviorGenetics, research has moved on. In 2008 we saw the publication of genome wide associationstudies for Bipolar disorder (Ferreira et al., 2008), for five dimensions of personality (Ter-racciano et al., 2008), ADHD (Neale et al., 2008) and major depressive disorder (Sullivanet al., 2008). Many more GWA studies of complex behavioral and psychiatric phenotypes areexpected in the next few years. The landscape of behavior genetics has changed remarkablyin a relatively short space of time. The field continues to progress from comparatively smallstudies to consortia-based efforts that target the inherited components of complex diseases andbehaviors and which typically involve thousands of participants (Orr & Chanock, 2008).

References

Ferreira, M. A., O’Donovan, M. C., Meng, Y. A., Jones, I. R., Ruderfer, D. M., Jones, L., et al. (2008).Collaborative genome-wide association analysis supports a role for ANK3 and CACNA1C in bipolardisorder. Nature Genetics, 40, 1056–1058.

Neale, B. M., Lasky-Su, J., Anney, R., Franke, B., Zhou, K., Maller, J. B., et al. (2008). Genome-wideassociation scan of attention deficit hyperactivity disorder. American Journal of Medical Genetics BNeuropsychiatric Genetics, 147B, 1337–1344.

Orr, N., & Chanock, S. (2008). Common genetic variation and human disease. Advances in Genetics, 62, 1–32.Sullivan, P. F., de Geus, E. J. C., Willemsen, G., James, M. R., Smit, J. H., Zandbelt, T., et al. (2008).

Genome-wide association for major depressive disorder: a possible role for the presynaptic protein piccolo.Molecular Psychiatry, 1–17.

Terracciano, A., Sanna, S., Uda, M., Deiana, B., Usala, G., Busonero, F., et al. (2008). Genome-wide associationscan for five major depressions of personality. Molecuar Psychiatry (in press).

Acknowledgments

I gratefully acknowledge John DeFries, L. Erlenmeyer-Kimling, Irving Gottesman, JenaeNeiderhiser, Nancy Pedersen, and Anita Thapar who provided useful suggestions for shapingthe contents of this handbook. I owe a great debt to my colleagues whose contributions createthe substance of this exciting book. They have been very patient, and they have promptlyresponded and revised their chapters in light of reviewers’ comments and suggestions. I hopethat they are as pleased with the final result as I am. I also acknowledge Janice Stern, mySpringer Editor, for her patience on the long delay of the handbook, and I thank her staff fortheir help in publishing this book.

I am especially grateful to the numerous colleagues who critically reviewed and improvedthe earlier versions of the chapters with very valuable comments and suggestions. These peopleinclude W. Anderson, J. Ando, R. Asarnow, H. Bleaut, D. Blizzard, D. Boomsma, J. Bridle,D. Bruck, T. Button, A. Caspi, S. Cherny, K. Christensen, J. Crabbe, W. Crusio, K. Deater-Deckard, S. de Boer, C. DeCarli, J. DeFries, D. Dick, L. DiLalla, M. Dunne, L. Ehrman,T. Eley, L. Erlenmeyer-Kimling, W. Etges, J.-F. Ferveur, G. Fisch, B. Fish, J. Flint, S. Gammie,X. Ge, M. Geyer, N. Gillespie, J. Gleason, D. Gooding, I. Gottesman, T. Gould, P. A. Gowaty,E. Grigorenko, N. Harlaar, J. Harris, N. Henderson, J. Hewitt, A. Holmes, J. Horn, G. Jackson,J.-M. Jallon, K. Jang, J. Kaprio, W. Kates, J. Kuntsi, C. Kyriacou, R. Landgraf, J.-O. Larsson,L. Leve, J. Loehlin, M. Luciano, H. Maes, T. Markow, L. Matzel, S. Maxson, R. McCrae,S. McGuire, S. Medland, K. Morley, P. Mullineaux, J. Neiderhiser, R. Nelson, Y. Oguma,D. Overstreet, D. Patterson, S. Petrill, A. Pike, R. Pillard, R. Plomin, M. Pogue-Geile,G. Riedel, G. Rieger, F. Rijsdijk, J. Ringo, P. Roubertoux, M. Rutter, S. Smith, M. Stallings,T. Suzuki, G. Swan, P. Szatmari, M. Thomis, R. Todd, T. Verson, E. Viding, J. Vink, I. Weiner,H. Welzl, J. Wilson, D. Wolfer, and J. Young.

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Contents

Introduction

1 History of Behavior Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3John C. Loehlin

Part I Quantitative Methods and Models

2 Biometrical Models in Behavioral Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15Michael C. Neale

3 QTL Methodology in Behavior Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35Stacey S. Cherny

4 Multivariate Genetic Analysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47Danielle Posthuma

5 Models of Human Behavior: Talking to the Animals . . . . . . . . . . . . . . . . . . . . . . . . . 61Gene S. Fisch

Part II Genetics of Cognition

6 Twin Studies of General Mental Ability . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81Nancy L. Segal and Wendy Johnson

7 Behavioral Genetic Investigations of Cognitive Aging . . . . . . . . . . . . . . . . . . . . . . . . 101Deborah Finkel and Chandra A. Reynolds

8 The Genetics and Environments of Reading: A Behavioral Genetic Perspective . 113Sara A. Hart and Stephen A. Petrill

9 Behavior-Genetic and Molecular Studies of Disorders of Speechand Language: An Overview . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125Elena L. Grigorenko

10 Human Brain Volume: What’s in the Genes? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137Jiska S. Peper, Marcel P. Zwiers, Dorret I. Boomsma, Rene S. Kahn, and HillekeE. Hulshoff Pol

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11 Cognition in Rodents . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159Christopher Janus, Michael J. Galsworthy, David P. Wolfer, and Hans Welzl

12 Neurogenetic Analysis and Cognitive Functions in Trisomy 21 . . . . . . . . . . . . . . . . 175Pierre L. Roubertoux and Michele Carlier

13 Evolution of Complex Acoustic Signals in Drosophila Species . . . . . . . . . . . . . . . . . 187Anneli Hoikkala and Dominique Mazzi

14 Drosophila Model of Alzheimer’s Amyloidosis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197Koichi Iijima, Kanae Iijima-Ando, and Yi Zhong

Part III Genetics of Personality

15 Genotype–Environment Correlation and Family Relationships . . . . . . . . . . . . . . . 209Jennifer A. Ulbricht and Jenae M. Neiderhiser

16 Personality . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223Kerry L. Jang and Shinji Yamagata

17 Molecular Genetics of Personality: How Our Genes can Bring Us to a BetterUnderstanding of Why We Act the Way We Do . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 239Richard P. Ebstein and Salomon Israel

18 The Genetics of Childhood Temperament . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 251Jeffrey R. Gagne, Matthew K. Vendlinski, and H. Hill Goldsmith

19 Genetic and Environmental Influences on Sexual Orientation . . . . . . . . . . . . . . . . . 269Khytam Dawood, J. Michael Bailey, and Nicholas G. Martin

20 Some Guidelines for Defining Personality Differences in Rats . . . . . . . . . . . . . . . . . 281Peter Driscoll, Alberto Fernandez-Teruel, Maria G. Corda, Osvaldo Giorgi,and Thierry Steimer

21 The Genetics of Offensive Aggression in Mice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301Stephen C. Maxson

22 Sexual Selection and Aggressive Behavior in Drosophila . . . . . . . . . . . . . . . . . . . . . 317Yong-Kyu Kim

23 Handedness: A Behavioral Laterality Manifestation . . . . . . . . . . . . . . . . . . . . . . . . . 331Ira B. Perelle and Lee Ehrman

24 Genetics of Exercise Behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 343Janine H. Stubbe and Eco J.C. de Geus

Part IV Genetics of Psychopathology

25 Genetics of ADHD, Hyperactivity, and Attention Problems . . . . . . . . . . . . . . . . . . . 361Eske M. Derks, James J. Hudziak, and Dorret I. Boomsma

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26 Depression and Anxiety in Childhood and Adolescence: DevelopmentalPathways, Genes and Environment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 379Frances Rice and Anita Thapar

27 Genetics of Autism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 397Sarah Curran and Patrick Bolton

28 Genetics of Smoking Behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 411Richard J. Rose, Ulla Broms, Tellervo Korhonen, Danielle M. Dick, and JaakkoKaprio

29 The Genetics of Substance Use and Substance Use Disorders . . . . . . . . . . . . . . . . . 433Danielle M. Dick, Carol Prescott, and Matt McGue

30 Genetic Analysis of Conduct Disorder and Antisocial Behavior . . . . . . . . . . . . . . . 455Soo Hyun Rhee and Irwin D. Waldman

31 Schizophrenia and Affective Psychotic Disorders – Inputsfrom a Genetic Perspective . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 473Daniel R. Hanson

32 Genetic Risks in Schizophrenia: Cross-National Prospective LongitudinalHigh-Risk Studies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 487Judith G. Auerbach, L. Erlenmeyer-Kimling, Barbara Fish, Sydney L. Hans,Loring J. Ingraham, Joseph Marcus, Thomas F. McNeil, and Erland Schubert

33 Attention and Working Memory: Animal Models for CognitiveSymptoms of Schizophrenia – Studies on D2-Like ReceptorKnockout Mice . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 501Claudia Schmauss

Conclusion

34 Future Directions for Behavior Genetics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 515Yong-Kyu Kim

Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 519

Contributors

Judith G. Auerbach Department of Psychology, Ben-Gurion University, Beer-Sheva,Israel; Center for Advanced Studies, Norwegian Academy of Science and Letters, Oslo,Norway,Email: judy@bgu.ac.il

J. Michael Bailey Department of Psychology, Northwestern University, Evanston, IL 60637,USA,Email: jm-bailey@northwestern.edu

Patrick F. Bolton Department of Psychologist Medicine, Institute of Psychiatry, London SE58AF, UK,Email: p.bolton@iop.kcl.ac.uk

Dorret I. Boomsma Department of Biological Psychology, Vrije Universiteit, Amsterdam1081 BT, The Netherlands,Email: DI.Boomsma@psy.vu.nl

Ulla Broms Department of Public Health, University of Helsinki, Helsinki 00014, Finland;Department of Mental Health and Alcohol Research, National Public Health Institute,Helsinki, Finland,Email: ulla.broms@helsinki.fi

Michele Carlier Institut Universitaire de France and Laboratoire Psychologie Cognitive,UMR 6146, CNRS Aix-Marseille 1, Marseille Cedex 20, France,Email: Michele.Carlier@univ-provence.fr

Stacey S. Cherny Department of Psychiatry, Genome Research Centre, and The State Key,Laboratory of Brain and Cognitive Sciences, University of Hong Kong, Pokfulam, HongKong,Email: cherny@hku.hk

Maria G. Corda Department of Toxicology, University of Cagliari, Cagliari 09124, Italy,Email: mgcorda@unica.it

Sarah Curran Department of Psychologist Medicine, Institute of Psychiatry, London SE58AF, UK,Email: s.curran@iop.kcl.ac.uk

Khytam Dawood Department of Psychology and Center for Developmental and Health Genet-ics, Pennsylvania State University, University Park, PA 16802, USA,Email: Khytam@psu.edu

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xviii Contributors

Eco J.C. de Geus Department of Biological Psychology, Vrije Universiteit, Amsterdam 1081BT, The Netherlands,Email: JCN.de.Geus@psy.vu.nl

Eske M. Derks Department of Biological Psychology, Vrije Universiteit, Amsterdam 1081BT, The Netherlands; University Medical Center Utrecht, Divisie hersenen, Heidelberglaan100, 3584 CX Utrecht, The Netherlands,Email: E.M.Derks@umcutrecht.nl

Danielle M. Dick Virginia Institute of Psychiatric and Behavioral Genetics, VirginiaCommonwealth University, Richmond, VA 23298, USA,Email: ddick@vcu.edu

Peter Driscoll Institute for Animal Science, ETHZ, Schwerzenbach 8603, Switzerland,Email: peter-driscoll@ethz.ch

Richard P. Ebstein Department of Psychology, Scheinfeld Center of Genetic Studies for theSocial Sciences, Hebrew University of Jerusalem; Sarah Herzog Memorial Hospital, GivatShaul, Jerusalem 91905, Israel,Email: ebstein@mscc.huji.ac.il

Lee Ehrman School of Natural and Social Sciences, Purchase College, State University ofNew York, Purchase, NY 10577, USA,Email: lee.ehrman@purchase.edu

L. Erlenmeyer-Kimling Departments of Medical Genetics, New York State PsychiatricInstitute, New York, NY 10032, USA; Departments of Psychiatry and of Genetics andDevelopment, Columbia University, New York, NY, USA,Email: le4@columbia.edu

Alberto Fernandez-Teruel Medical Psychology Unit, Department of Psychiatry and ForensicMedicine, University of Barcelona, Bellaterra 08193, Spain,Email: albert.fernandez.teruel@uab.es

Deborah Finkel Department of Psychology, School of Social Sciences, Indiana UniversitySoutheast, New Albany, IN 47150, USA,Email: dfinkel@ius.edu

Gene S. Fisch Bluestone Clinical Research Center, NYU Colleges of Dentistry & Nursing,Yeshiva University, New York, NY 10010, USA,Email: gene.fisch@nyu.edu

Barbara Fish Department of Psychiatry and Behavioral Sciences, University of California,Los Angeles, CA 90095, USA,Email: bfish@ucla.edu

Jeffrey R. Gagne Department of Psychology, University of Wisconsin, Madison, WI 53706,USA,Email: jgagne@wisc.edu

Contributors xix

Michael J. Galsworthy Division of Neuroanatomy and Behavior, Institute of Anatomy,University of Zurich, Zurich CH-8057, Switzerland,Email: mike galsworthy@yahoo.co.uk

Osvaldo Giorgi Department of Toxicology, University of Cagliari, Cagliari 09124, Italy,Email: giorgi@unica.it

H. Hill Goldsmith Department of Psychology, University of Wisconsin, Madison, WI 53706,USA,Email: hhgoldsm@wisc.edu

Elena L. Grigorenko Child Study Center, Department of Psychology, Department ofEpidemiology & Public Health, Yale University, New Haven, CT 06519-1124, USA,Email: elena.grigorenko@yale.edu

Sydney L. Hans School of Social Service Administration, University of Chicago, Chicago,IL 60637, USA,Email: s-hans@uchicago.edu

Daniel R. Hanson Departments of Psychiatry & Psychology, University of Minnesota,Minneapolis, MN 55454, USA,Email: drhanson@umn.edu

Sara A. Hart Department of Human Development and Family Science, The Ohio StateUniversity, Columbus, OH 43210, USA,Email: hart.327@osu.edu

Anneli Hoikkala Department of Biological and Environmental Science, University ofJyvaskyla, Jyvaskyla, Finland,Email: anneli.a.hoikkala@jyu.fi

James J. Hudziak Department of Psychiatry and Medicine (Division of Human Genetics),Center for Children, Youth and Families, University of Vermont, Burlington, VT, USA,Email: James.Hudziak@uvm.edu

Hilleke E. Hulshoff Pol Department of Psychiatry, Rudolf Magnus Institute of Neuroscience,University Medical Center, Utrecht, The Netherlands,Email: h.e.hulshoff@umcutrecht.nl

Koichi Iijima Laboratory of Neurodegenerative Diseases and Gene Discovery; FarberInstitute for Neurosciences; Department of Biochemistry and Molecular Biology, ThomasJefferson University, Philadelphia PA19107, USA,Email: Koichi.Iijima@jefferson.edu

Kanae Iijima-Ando Laboratory of Neurogenetics and Pathobiology; Farber Institute forNeurosciences; Department of Biochemistry and Molecular Biology, Thomas JeffersonUniversity, Philadelphia PA 19107, USA,Email: Kanae.Iijima-Ando@jefferson.edu

Loring J. Ingraham Department of Psychology, The George Washington University,Washington, DC 20037, USA,Email: ingraham@gwu.edu

xx Contributors

Salomon Israel Department of Psychology, Scheinfeld Center of Genetic Studies for theSocial Sciences, Hebrew University of Jerusalem, Jerusalem 91905, Israel,Email: salomon.israel@mail.huji.ac.il

Kerry L. Jang Department of Psychiatry, University of British Columbia, Vancouver, BritishColumbia, Canada, V6T 2A1.Email: kjang@interchange.ubc.ca

Christopher Janus Mayo Clinic Jacksonville, Department of Neuroscience, University ofFlorida, Jacksonville, FL 32224, USA,Email: Janus.Christopher@mayo.edu

Wendy Johnson Centre for Cognitive Ageing and Cognitive Epidemiology and Departmentof Psychology, University of Edinburgh, Edinburgh, UK, and Department of Psychology,University of Minnesota, Minneapolis, MN 55455, USA,Email: john4350@umn.edu

Rene S. Kahn Department of Psychiatry, Rudolf Magnus Institute of Neuroscience, UniversityMedical Center, Utrecht, The Netherlands,Email: r.kahn@umcutrecht.nl

Jaakko Kaprio Department of Public Health, University of Helsinki, Helsinki 14, Finland;Department of Mental Health and Alcohol Research, National Public Health Institute,Helsinki, Finland,Email: jaakko.kaprio@helsinki.fi

Yong-Kyu Kim Department of Genetics, University of Georgia, Athens, GA 30602, USA,Email: yongkyu@uga.edu

Tellervo Korhonen Department of Public Health, University of Helsinki, Helsinki 14,Finland; Department of Mental Health and Alcohol Research, National Public HealthInstitute, Helsinki, Finland,Email: tellervo.korhonen@helsinki.fi

John C. Loehlin Department of Psychology, University of Texas at Austin, Austin, TX 78712,USA,Email: loehlin@psy.utexas.edu

Joseph Marcus Department of Psychiatry, University of Chicago, Chicago, IL 60637, USA,Email: jmarcusmd@cox.net

Nicholas G. Martin Department of Genetic Epidemiology, Queensland Institute of MedicalResearch, Brisbane, Queensland, Australia,Email: Nick.Martin@qimr.edu.au

Stephen C. Maxson Department of Psychology, University of Connecticut, Storrs, CT 06269-1020, USA,Email: Stephen.Maxson@Uconn.Edu

Contributors xxi

Dominique Mazzi Institute of Plant Sciences, Swiss Federal Institute of Technology, Zurich,Switzerland,Email: dominique.mazzi@ipw.argl.ethz.ch

Matthew McGue Department of Psychology, University of Minnesota, Minneapolis, MN55455, USA; Department of Epidemiology, Southern Denmark University, Denmark,Email: mcgue001@tc.umn.edu

Thomas F. McNeil Department of Psychiatric Epidemiology, University Hospital, LundUniversity, Israel,Email: Thomas.McNeil@med.lu.se

Michael C. Neale Departments of Psychiatry and Human Genetics, Virginia Institute forPsychiatric and Behavioral Genetics, Virginia Commonwealth University, Richmond, VA23298, USA,Email: neale@vcu.edu

Jenae M. Neiderhiser Department of Psychiatry and Behavioral Sciences, Center for FamilyResearch, The Pennsylvania State University, University Park, PA 16802, USA,Email: jenaemn@psu.edu

Jiska S. Peper Department of Psychiatry, Rudolf Magnus Institute of Neuroscience,University Medical Center, Utrecht, The Netherlands,Email: j.s.peper@umcutrecht.nl

Ira B. Perelle Department of Psychology, Mercy College, Dobbs Ferry, NY 10522, USA,Email: iperelle@mercy.edu

Stephen A. Petrill Department of Human Development and Family Science, Ohio StateUniversity, Columbus, OH 43210, USA,Email: spetrill@ehe.osu.edu

Danielle Posthuma Department of Biological Psychology, Section Medical Genomics, andSection Functional Genomics, Vrije Universiteit and Vrije Universiteit Medical Center,Amsterdam 1081BT, The Netherlands,Email: danielle@psy.vu.nl

Carol Prescott Department of Psychology, University of Southern California, Los Angeles,CA 90089, USA,Email: cprescot@usc.edu

Chandra A. Reynolds Department of Psychology, University of California, Riverside, CA92521, USA,Email: chandra.reynolds@ucr.edu

Soo Hyun Rhee Institute for Behavioral Genetics, University of Colorado, Boulder, CO80309, USA,Email: Soo.Rhee@colorado.edu

Frances Rice Child & Adolescent Psychiatry Section, Department of Psychological Medicine,School of Medicine, Cardiff University, Cardiff CF14 4XN, UK,Email: ricef@cardiff.ac.uk

xxii Contributors

Richard J. Rose Department of Psychological and Brain Sciences, Indiana University,Bloomington, IN 47405, USA; Department of Public Health, University of Helsinki, Finland,Email: rose@indiana.edu

Pierre L. Roubertoux INSERM UMR910 Genetique Medicale et Genomique Fonctionnelle,Universite d’Aix-Marseille 2, Faculte de Medecine, 27 Bvd Jean Moulin, 13385 MarseilleCedex 05, France,Email: pierre.roubertoux@univmed.fr

Claudia Schmauss Department of Psychiatry and Molecular Therapeutics, ColumbiaUniversity and New York State Psychiatric Institute, New York, NY 10032, USA,Email: cs581@columbia.edu

Erland Schubert Department of Psychiatric Epidemiology, University Hospital, LundUniversity, Israel,Email: erland.schbert@psychepi.lu.se

Nancy L. Segal Department of Psychology, California State University, Fullerton, CA 92834,USA,Email: nsegal@fullerton.edu

Thierry Steimer Clinical Psychopharmacology Unit, University Hospital of Geneva, 1225Chene-Bourg, Switzerland,Email: thierry.steimer@hcuge.ch

Janine H. Stubbe Department of Biological Psychology, Vrije Universiteit, Amsterdam 1081BT, The Netherlands,Email: janine.stubbe@tno.nl

Anita Thapar Child & Adolescent Psychiatry Section, Department of PsychologicalMedicine, Cardiff University, Cardiff CF14 4XN, UK,Email: thapar@Cardiff.ac.uk

Jennifer A. Ulbricht Center for Family Research; Department of Psychology, GeorgeWashington University, Washington, DC 20037, USA,Email: jau5b@gwu.edu

Matthew K. Vendlinski Department of Psychology, University of Wisconsin, Madison, WI53706, USA,Email: vendlinski@wise.edu

Irwin D. Waldman Department of Psychology, Emory University, Atlanta, GA 30322, USA,Email: psyiw@emory.edu

Hans Welzl Division of Neuroanatomy and Behavior, Institute of Anatomy, University ofZurich, Zurich CH-8057, Switzerland,Email: welzl@anatom.uzh.ch

David P. Wolfer Division of Neuroanatomy and Behavior, Institute of Anatomy, University ofZurich, Zurich CH-8057, Switzerland,Email: dpwolfer@anatom.uzh.ch

Contributors xxiii

Shinji Yamagata Faculty of Letters, Keio University, Tokyo, 108-8345, Japan,Email: yamagata@bayes.cu-tokyo.ad.jp

Yi Zhong Cold Spring Harbor Laboratory, Cold Spring Harbor, NY 11724, USA,Email: zhongyi@cshl.edu

Marcel P. Zwiers F.C. Donders Center for Cognitive Neuroscience, Nijmegen, TheNetherlands; Psychiatry Department, Radboud University, Nijmegen Medical Center,Nijmegen 6525 HE, The Netherlands,Email: m.zwiers@fcdonders.ru.nl

Introduction

Chapter 1

History of Behavior Genetics

John C. Loehlin

Hermann Ebbinghaus (1908) said of psychology that it had along past, but only a short history. The same may be said ofbehavior genetics. One cannot specify an exact date at whichbehavior genetics came to be regarded as a distinct scientificdiscipline, but for convenience let us say 1960, the publica-tion date of Fuller and Thompson’s textbook of that title.

This chapter considers both the long past and someaspects of the short history of behavior genetics. We beginwith the long past: the recognition since antiquity that behav-ioral traits are in part inherited, and the controversy concern-ing the extent to which this is so, a discussion often goingunder the label of the nature–nurture controversy.

The Long Past of Behavior Genetics

From Ancient Times to the Renaissance

Ancient Times

Where does the long past start? Perhaps with the domes-tication of dogs for behavioral as well as physical traits,a process which probably took place at least 15,000 yearsago (Savolainen, Zhang, Luo, Lundeberg, & Leitner, 2002) –although one must suppose that in its early days this wasmore an evolution of a subgroup of wolves to fit a nichearound human habitation than a process deliberately under-taken by man (Morey, 1994). In any case, about 5000 yearsago in Egypt and the Near East, it appears that deliber-ate animal breeding was well established (Brewer, Clark, &Phillips, 2001); several distinctive varieties of cattle and dogsare portrayed in ancient Egyptian art.

J.C. Loehlin (B)Department of Psychology, The University of Texas at Austin, Austin,TX 78712, USA

Greeks, Romans, Hebrews

By classical times, 3000–1500 years ago, many varieties ofdogs with distinctive physical and behavioral characteristicswere recognized. More than 50 breeds are named in sur-viving Greek and Roman documents, falling into such cate-gories as scent- and sight hounds, shepherd dogs, guard dogs,war dogs, and pets (Brewer et al., 2001).

The ancient Greeks held that humans inherited quali-ties, including behavioral ones, from their ancestors. Thusin Book IV of Homer’s Odyssey, Menelaus greets two youngvisiting strangers, “Ye are of the line of men that are scep-tred kings . . . for no churls could beget sons like you”(Homer, trans. 1909, p. 49). And later (p. 53), to one of them,“Thou has said all that a wise man might say or do, yea, andan elder than thou; – for from such a sire too thou art sprung,wherefore thou dost even speak wisely.” A similar notionwas expressed in the Hebrew scriptures: “I am the heir ofwise men, and spring from ancient kings” (Isaiah 19:11, NewEnglish Bible).

A few hundred years later, the Greek philosopher Platoin Book V of the Republic – his prescription for an idealstate – took both inheritance and instruction into accountin the development of the “Guardians,” the ruling elite. Hebegins with the question, “How can marriages be made mostbeneficial?” He discusses the breeding of hunting dogs andbirds, noting that “Although they are all of a good sort, arenot some better than others?” “True.” “And do you breedfrom them all indifferently, or do you take care to breed fromthe best only?” “From the best” (Plato, trans. 1901, p. 149).From there Plato goes on to generalize to the class of elitehumans in his ideal state – to the desirability of matching thebest with the best, and rearing their offspring with specialattention.

Plato recognizes that good ancestry is not infallibly pre-dictive and recommends applying, at least in early youth,a universal education to the citizens of his state; demoting,when inferior, offspring of the elite class of guardians andelevating into the ranks of the guardians offspring of thelower classes who show merit.

Y.-K. Kim (ed.), Handbook of Behavior Genetics, 3DOI 10.1007/978-0-387-76727-7 1, c© Springer Science+Business Media, LLC 2009

4 J.C. Loehlin

We need not debate the pros and cons of Plato’s partic-ular social proposals; people have been arguing about themever since his day. We only need observe that well over 2000years ago the interplay of nature and nurture – and its socialimplications – was being discussed.

Middle Ages

What of the contrary view, the notion that all men areborn equal? A major impetus to such an idea came fromthe medieval Catholic Church (Pearson, 1995). All men aresons of God, and therefore of equal value in His sight. Or,from another perspective, as the fourteenth-century Englishproverb had it, “When Adam delved and Eve span/Who wasthen a gentleman?”

The Renaissance

Ideas concerning the inheritance of behavior were present inShakespeare’s day. The Countess of Rossilon in All’s WellThat Ends Well says, about a wise daughter of a wise father,“Her dispositions she inherits” (Act I:i). The nature–nurturecontroversy itself appears to have got its label from Pros-pero’s remark in The Tempest about his subhuman creature,Caliban, “A devil, a born devil, on whose nature nurture willnever stick” (Act IV:i).

The Nature–Nurture Controversy in theModern Era

Although ideas about the roles of nature and nurture inhuman and animal behaviors have been with us for thousandsof years, the modern form of the controversy traces backfairly directly to the seventeenth-century philosopher JohnLocke and the nineteenth-century naturalist Charles Darwin.

John Locke

Locke may be considered to be the chief ideological fatherof the nurture side of the controversy. In An Essay Concern-ing Human Understanding (Locke, 1690/1975), he invokedthe metaphor of the mind as a blank sheet of paper uponwhich knowledge is written by the hand of experience. In theopening paragraph of his book Some Thoughts ConcerningEducation, he said, “I think I may say, that of all the Menwe meet with, nine Parts of ten are what they are, good orevil, useful or not, by their Education” (Locke, 1693/1913,Sect. 1). Locke’s political view that all men are by natureequal and independent, and that society is a mutual contract

entered into for the common good, had an immense influencevia Jefferson, Voltaire, Rousseau, and the other theorists ofthe American and French revolutions.

Indeed, one may view many of the events of the nature–nurture controversy since Locke’s day as a series of chal-lenges to the prevailing Lockean position, with those steepedin that tradition rising indignantly to battle what they per-ceived to be threats to inalienable human rights of liberty andequality.

Locke himself, however, was not nearly as alien to hered-itarian concepts as some of his followers have been. Herejected the concept of inborn ideas, but not of all innate char-acteristics. In a marginal note on a pamphlet by one ThomasBurnet, Locke wrote “I think noe body but this Author whoever read my book [An Essay Concerning Human Under-standing] could doubt that I spoke only of innate Ideas . . .

and not of innate powers . . . ” (see Porter, 1887). Elsewherein Some Thoughts Concerning Education Locke wrote,

Some Men by the unalterable Frame of their Constitutions arestout, others timorous, some confident, others modest, tractable,or obstinate, curious or careless, quick or slow. There are notmore Differences in Men’s Faces, or in the outward Lineamentsof their Bodies, than there are in the Makes and Tempers of theirMinds. (1693/1913, Sect. 101)

John Stuart Mill

Many of Locke’s successors in the English liberal traditioncame out more strongly than Locke did on the side of nurture.John Stuart Mill wrote in his Autobiography (1873, p. 192),

I have long felt that the prevailing tendency to regard all themarked distinctions of human character as innate, and in themain indelible, and to ignore the irresistible proofs that by far thegreater part of these differences, whether between individuals,races, or sexes, are such as not only might but naturally wouldbe produced by differences in circumstances, is one of the chiefhindrances to the rational treatment of great social questions, andone of the greatest stumbling blocks to human improvement.

Charles Darwin

During roughly the same period as Mill, Charles Darwingave the nature side of the controversy its modern formby placing behavior, including human behavior, solidly inthe framework of biological evolution. In addition to hismajor treatise The Origin of Species (1859), Darwin in suchworks as The Descent of Man (1871) and The Expressionof the Emotions in Man and Animals (1872) made it clearthat human behavior shared ancestry with that of other ani-mal forms, and was subject to the same evolutionary pro-cess of hereditary variation followed by natural selection ofthe variants that proved most successful in their particularenvironments.

1 History of Behavior Genetics 5

In The Descent of Man (1871, pp. 110–111) Darwinwrote,

So in regard to mental qualities, their transmission is manifestin our dogs, horses, and other domestic animals. Besides specialtastes and habits, general intelligence, courage, bad and goodtemper, etc. are certainly transmitted. With man we see simi-lar facts in almost every family; and we now know through theadmirable labours of Mr. Galton that genius, which implies awonderfully complex combination of high faculties, tends to beinherited; and on the other hand, it is too certain that insanity anddeteriorated mental powers likewise run in the same families.

Francis Galton

Darwin’s younger cousin Francis Galton agreed with Darwinand disagreed with Mill. In his book Inquiries into HumanFaculty (1883, p. 241) he concluded,

There is no escape from the conclusion that nature prevails enor-mously over nurture when the differences of nurture do notexceed what is commonly to be found among persons of thesame rank of society and in the same country.

Galton is not saying that environment never matters. How-ever, he is saying that the ordinary differences we observeamong people in the same general social context are mostlydue to heredity.

Galton was a central, crystallizing figure in behaviorgenetics’ “long past.” His emphasis on the measurement ofindividual differences and their statistical treatment becamea core theme in the development of the field. His studiesof “hereditary genius” and “the comparative worth of differ-ent races” (Galton, 1869) foreshadowed recent controversiesabout IQ. He proposed the study of twins as a way of gettingat the relative effect of nature and nurture. And his promotionof eugenics – that is, the encouragement of the more usefulmembers of society to have more children and the less usefulto have fewer (as in Plato’s scheme for an ideal state) – hasgenerated on occasion a good deal of heat. Here is a recentexample (Graves, 2001, p. 100): “Galton’s scientific accom-plishments are sufficient for some still to consider him anintellectual hero. Whereas for others (this author included)he was an intellectual mediocrity, a sham, and a villain.”

The Twentieth Century

Vigorous disagreements on the relative impact of nature andnurture on behavior continued into the twentieth century. Onthe whole, twentieth-century psychology was heavily envi-ronmentalistic, emphasizing the crucial role of learning inshaping behavior. The high-water mark of this tradition wasthe famous claim of John B. Watson (1925, p. 82):

Give me a dozen healthy infants, well-formed, and my own spec-ified world to bring them up in and I’ll guarantee to take any

one at random and train him to become any type of specialistI might select – doctor, lawyer, artist, merchant-chief, and yes,even beggar-man and thief, regardless of his talents, penchants,tendencies, abilities, vocations, and race of his ancestors.

The year 1928 saw the publication of the Twenty-SeventhYearbook of the National Society for the Study of Educa-tion. It was entitled Nature and Nurture, and it containedthe reports of two adoption studies of IQ. One, by BarbaraBurks, emphasized the effects of nature. The other, by Free-man, Holzinger, and Mitchell, came down on the side ofnurture. The nature–nurture controversy continued, but stu-dents of the effects of heredity and environment on behaviorwere gathering data. When enough had been gathered for atextbook to be written, the short history of behavior geneticscould begin.

The Short History of Behavior Genetics

Most of the short history of behavior genetics, as it appliesto the study of both humans and other animal species, willnot be discussed in this chapter. It is a tale of steady sci-entific progress on a variety of fronts, despite occasionalcontroversies, confusions, and setbacks, and it is a tale toldin the other chapters of this handbook. The reader who wantsa quick sense of the scope of scientific progress in the fieldof behavior genetics during the last 40-odd years, and theprospects opening up in it today, can achieve this by scanningthrough the chapter introductions and summaries, and theeditor’s final chapter. The reader who aspires to a more solidgrasp of this short history will need, of course, to proceedmore systematically through the book, as well as followingup some of its many references.

The remainder of this chapter addresses two other aspectsof behavior genetics’ short history. First, we look briefly atsome institutional features of the field: its principal schol-arly and scientific organization, the Behavior Genetics Asso-ciation; the discipline’s key journal, Behavior Genetics; andsome major centers of behavior genetics research. Followingthis, we look at the social context of behavior genetics, atinstances in which the scientific and scholarly pursuits of thefield have become entangled with public political and socialconcerns. These instances include a series of controversiesconcerning the genetic or environmental bases of differencesin psychological characteristics between groups defined byrace, sex, or social class. Controversies about group differ-ences have roots in behavior genetics’ long past and havepersisted into its short history. They are far from central inthe activities of most working behavior geneticists, but theyrepresent an important part of the public face of the field.

6 J.C. Loehlin

The Institutional History of Behavior Genetics

The Behavior Genetics Association

After some informal discussions in the late 1960s, and thecirculation of a mailing to a list of persons who had recentlypublished in the area of behavior genetics, an organizationalmeeting took place at Urbana, Illinois, in March 1970. R.H. Osborne, then editor of the journal Social Biology, waschosen to act as president pro tem, and five committees wereappointed to lay the groundwork for a Behavior GeneticsAssociation (or Society – there was some argument abouta suitable name). In April 1971, the fledgling organizationheld its first formal meeting, at Storrs, Connecticut. In addi-tion to scientific sessions, a draft constitution was discussedto be submitted to the initial membership via mail ballot forapproval. Nominations and an election followed, and at thetime of the second annual meeting at Boulder, Colorado,in April 1972, the Behavior Genetics Association (BGA)was officially underway, and its first set of officers tookoffice: Theodosius Dobzhansky was president, John Fullerwas president-elect, R. H. Osborne served as past president,the secretary was Elving Anderson, the treasurer was JohnLoehlin, and the two executive committee members-at-largewere Seymour Kessler and L. Erlenmeyer-Kimling.

The association proved viable. Table 1.1 shows the suc-cessive presidents of the BGA and the location of its annualmeetings. Note that a special extra international meeting washeld in Jerusalem in 1981, and that thereafter the regularannual BGA meeting was periodically held in countries out-side the USA: in England (twice), the Netherlands (twice),France, Australia, Spain, Canada (twice), and Sweden.

Over time, the association grew in size. Forty-four personsresponded to the initial mailing indicating interest in such anassociation. There were 69 paid-up members at the time ofthe first annual meeting at Storrs. By the time of the 34thannual meeting in Aix-en-Provence, France, in 2004, theBGA had 270 regular and 109 associate members (the latterchiefly graduate students). Approximately two-thirds werefrom North America and one-third from other continents.

The Journal Behavior Genetics

In 1970, a decade after Fuller and Thompson’s textbook, thescientific journal Behavior Genetics began with Vol. 1, No. 1.Its founding editors were Steven G. Vandenberg and John C.DeFries. They stated their hopes for the new journal in aneditorial (p. 1):

Research in behavior genetics continues to be undertaken atan accelerating rate. Nevertheless, no single journal has existedheretofore which was dedicated primarily to the publication ofpapers in this important area. Since manuscripts in behavior

Table 1.1 BGA Presidents and Annual Meetings

Year President Site of meeting

1971 R. H. Osborne [pro tem] Storrs CT1972 Th. Dobzhansky Boulder CO1973 John L. Fuller Chapel Hill NC1974 Gerald E. McClearn Minneapolis MN1975 J. P. Scott Austin TX1976 Irving I. Gottesman Boulder CO1977 W. R. Thompson Louisville KY1978 Lee Ehrman Davis CA1979 V. Elving Anderson Middletown CT1980 John C. Loehlin Chicago IL1981 Norman D. Henderson Purchase NY/Jerusalem1982 John C. DeFries Ft Collins CO1983 David W. Fulker London, England1984 Steven G. Vandenberg Bloomington IN1985 Sandra Scarr State College PA1986 Ronald S. Wilson Honolulu HI1987 Peter A. Parsons Minneapolis MN1988 Leonard L. Heston Nijmegen, Netherlands1989 Robert Plomin Charlottesville VA1990 Carol B. Lynch Aussois, France1991 Lindon J. Eaves St. Louis MO1992 David A. Blizard Boulder CO1993 Thomas J. Bouchard, Jr. Sydney, Australia1994 Glayde Whitney Barcelona, Spain1995 James Wilson Richmond VA1996 Nicholas G. Martin Pittsburgh PA1997 Nicholas G. Martin Toronto, Canada1998 Norman D. Henderson Stockholm, Sweden1999 Richard Rose Vancouver, Canada2000 John Hewitt Burlington VT2001 Matt McGue Cambridge, England2002 Nancy Pedersen Keystone CO2003 Andrew Heath Chicago IL2004 Michele Carlier Aix-en-Provence, France2005 H. Hill Goldsmith Hollywood CA2006 Laura Baker Storrs CT2007 Pierre Roubertoux Amsterdam, Netherlands

Source: BGA web site (June 27, 2007); http://www.bga.org

genetics have thus been published in widely scattered journals, aclear identification with this discipline has been lacking. It is ourhope that BEHAVIOR GENETICS will fulfill this need.

The journal has largely lived up to their hopes. It neverstood completely alone – for example, at the time there wasan existing journal focused on twin research, Acta GeneticaeMedicae et Gemellologiae, which published many behav-iorally oriented papers. The journal Social Biology – whoseeditor, R. H. Osborne, played an important role in found-ing the Behavior Genetics Association – initially served asthe official organ of the BGA. (Behavior Genetics assumedthat role in 1974.) Other journals have since emerged –for example, the recent journals Genes, Brains, and Behav-ior and Twin Research. Many important papers in behaviorgenetics continue to be published in journals in the neighbor-ing behavioral and biological sciences. Nevertheless, Behav-ior Genetics, as the official organ of the Behavior GeneticsAssociation, remains a major defining force in the field.

1 History of Behavior Genetics 7

It is instructive to compare Vol. 1 (1970) of BehaviorGenetics with Vol. 35 (2005). The journal became a gooddeal bigger: from three issues in Vol. 1 (Nos. 3 and 4 werebound together) to six in Vol. 35 from 274 to 854 pages(and nearly twice the number of words per page becauseof larger pages). In Vol. 1, there were 24 papers, an edito-rial, and 2 “short communications.” In Vol. 35 there were66 papers, plus 142 abstracts from the Behavior GeneticsAssociation meeting, and various BGA minutes, announce-ments, etc. Behavior Genetics continues to publish both sub-stantively and methodologically oriented papers, featuringvarious animal species, but the mix changed from Vol. 1to Vol. 35. In Vol. 1 there were 7 papers (27%) focused onhuman behavior, 16 papers (62%) involving rodents, mostlyinbred mice, 1 paper on another species (Drosophila), and2 papers primarily methodological (statistical) in character.In Vol. 35, there was an increased proportion of substantivepapers involving humans, 28 (42%); proportionately fewerinvolving rodents, 14 (21%); an increase in those involvingother animal species, 9 (15%) – mostly Drosophila, but oneon rainbow trout. For many of the remaining 22% of papers,the species might be described as the computer: These weremethodological papers, many involving a heavy dose of com-puter model-fitting or simulation.

Major Behavior Genetics Centers

Preeminent among academic centers for teaching andresearch in behavior genetics has been the Institute forBehavioral Genetics (IBG) at the University of Colorado atBoulder. Among the notable behavior geneticists who haveserved on its faculty are Gregory Carey, John DeFries, DavidFulker, John Hewitt, Carol Lynch, Gerald McClearn, RobertPlomin, Steven Vandenberg, and James Wilson. It has alsoserved as home for the journal Behavior Genetics, except for1978–1985 when Jan Bruell edited the journal at the Uni-versity of Texas and 2000–2002 when Norman Hendersonedited it at Oberlin College. The IBG has also hosted sev-eral BGA annual meetings and a number of summer traininginstitutes on behavior genetics methods.

Next in line as a center of behavior genetics activity wouldprobably be the University of Minnesota, whose faculty hasincluded important behavior geneticists like Elving Ander-son, Thomas Bouchard, Irving Gottesman, Leonard Heston,Gardner Lindzey, David Lykken, Matthew McGue, Shel-don Reed, Sandra Scarr, and Auke Tellegen. A third cen-ter, at least in the early days, was the University of Texasat Austin, with Jan Bruell, Joseph Horn, Gardner Lindzey,John Loehlin, Delbert Thiessen, and Lee Willerman. A cur-rent major behavior genetics center is at the Virginia Com-monwealth University; its faculty includes Lindon Eaves,Kenneth Kendler, Hermine Maes, and Michael Neale. Other

important U.S. centers include Washington University inSt. Louis (Robert Cloninger, Andrew Heath, & John Rice)and Penn State (David Blizard, Gerald McClearn, & GeorgeVogler). Outside the USA, Kings College, London, hasrecruited an eminent group of behavior genetics researchers,including Peter McGuffin, Robert Plomin, and Michael Rut-ter. The Vrije Universiteit in Amsterdam also has a substan-tial behavior genetics contingent, including Dorret Boomsmaand Danielle Postuma. Stable international coalitions arebecoming increasingly common, greatly facilitated by theInternet. Notable examples include collaborations betweengroups at Indiana University and the University of Helsinki,Penn State and the Karolinska Institute in Stockholm, andseveral U.S. groups with the Queensland Institute for Medi-cal Research in Australia.

Beside the institutions mentioned above, dozens of otheruniversities and research institutes, including many outsidethe USA, have developed and maintained strong programsin human or animal behavior genetics on the strength of oneor two distinguished researchers on their faculties. Almosthalf the presidents of the BGA, for example, would representthis category. The hosting of an annual BGA meeting (seeTable 1.1) also tends to reflect a strong local program.

Public Controversies – Group Differences

The possibility that there might be genetic differences inpsychological traits between groups defined by race, sex, orsocial class has led to a good deal of public uproar and nota little confusion. It has provided an inflammatory intersec-tion between the scientific discipline of behavior geneticsand Western attitudes of equality stemming from religious,political, and philosophical roots. Racist, sexist, and class-ist ideas (as references to such group differences are some-times called) tend to drive traditional Lockean ideologists upthe wall, so that clear thinking has not always prevailed inthis area.

A few general points should be noted. First, the mainbusiness of behavior geneticists has always been individualdifferences, not group differences, so that for the day-to-dayresearch of most behavior geneticists, questions about groupdifferences are at best an unwelcome distraction. Second, asLewontin (1970) made clear, a demonstration that individ-ual differences are due to genes does not imply that groupdifferences are genetic. He used the analogy of geneticallyvaried seeds raised in a greenhouse in two pots under iden-tical regimens, except that one pot lacked a crucial tracenutrient present for the other. The heights of the plantsare subsequently measured. The variation of height withineach pot, except for random measurement errors, is entirelygenetic, since the plants within each pot vary genetically,

8 J.C. Loehlin

but are treated exactly the same. The average difference inplant height between the two pots is entirely environmen-tal, because it stems from the presence or absence of thecritical nutrient. Clearly, this example implies that groupdifferences may be different in their genetic and environ-mental origins from individual differences. However, it issometimes forgotten that may does not imply are. Thereremains the empirical question for any particular trait andany particular group difference in any particular population:To what relative extent are genetic and environmental dif-ferences between the groups in fact involved? There alsoremains the social question: How much (if at all) does thismatter?

The empirical question is not necessarily an easy one toanswer. For one thing, it may well have different answers fordifferent traits and different groups (Loehlin, 2000). If onewere to demonstrate that profiles of cognitive ability differfor genetic reasons between Asian Americans and EuropeanAmericans, it would not imply that a difference in averageintellectual performance between European Americans andAfrican Americans has a genetic origin. To make mattersworse, the social excitement and media hoopla surroundingthe issue of group differences has discouraged most behav-ior geneticists from addressing such matters empirically. It isnot as though informative research designs do not exist. Onelisting of promising areas of research on racial-ethnic abilitydifferences listed ten possible approaches, ranging from stud-ies of race mixtures and cross-racial adoptions to piggy-backstudies on educational or nutritional programs which werebeing undertaken for other reasons (Loehlin, Lindzey, &Spuhler, 1975, pp. 251–254).

Jensen

Less than a decade into behavior genetics’ short history,the educational psychologist Arthur Jensen published along article in the Harvard Educational Review entitled“How much can we boost IQ and scholastic achievement?”(Jensen, 1969). Jensen noted the fact that compensatoryeducation programs had not lived up to their advance billingand concluded that this might partly reflect the geneticcontribution to IQ, which he estimated at a fairly high80%. Almost in passing, he noted the possibility that thepersistent IQ gap between U.S. blacks and whites mightin part be genetic in origin. He did not say that this hadbeen demonstrated to be the case, but suggested that thematter should be looked into empirically. Jensen’s article,particularly the suggestion that there might be a geneticcontribution to black–white IQ differences, created animmediate furor. There were numerous published critiques,not all judicious and carefully thought out. And this was notjust a genteel academic debate – tires were slashed and public

meetings disrupted. A graphic account of the goings-on maybe found in Pearson (1991). The controversy about possibleracial differences in mental abilities has continued to thepresent – the interested reader may wish to consult RaceDifferences in Intelligence (Loehlin et al., 1975), Race, IQand Jensen (Flynn, 1980), The Black–White Test Score Gap(Jencks & Phillips, 1998), and The New Know-Nothings(Hunt, 1999). Rushton and Jensen (2005) provide a recentreview emphasizing the genes: “Thirty years of research onrace differences in cognitive ability,” which, along with anumber of critiques from various points of view, fills an issueof Psychology, Public Policy, and Law [Vol. 11(2), 2005].

The Bell Curve

Twenty-five years after Jensen’s article, a similar uproararose, this time due to the publication of a book by thepsychologist Richard Herrnstein and the sociologist CharlesMurray entitled The Bell Curve (Herrnstein & Murray, 1994).Although much of the furor focused on race differences incognitive skills, the authors did not in fact devote a great dealof attention to this topic and took a fairly mild position onit. After emphasizing via a version of Lewontin’s metaphorthat a genetic basis for individual differences does not implya genetic basis for group differences, they said of U.S. ethnicdifferences in average IQ (p. 312):

They may well include some (as yet unknown) genetic compo-nent, but nothing suggests that they are entirely genetic. And,most important, it matters little whether the genes are involvedat all.

Their argument in support of the second sentence was thatfor an appropriate treatment of an individual it is his or herown IQ that is relevant (if IQ is relevant at all), not the aver-age IQs of some group to which the individual may belong.One might add, however, that for long-term social policy, thefact that an average group difference has its source in genesor in the environment can sometimes matter, because it canaffect the choice of a remedy to alter that difference – eugen-ics versus Head Start, for example.

Herrnstein on Social Class and IQ

The Bell Curve did not represent Herrnstein’s first engage-ment with group differences and public controversy. In anarticle in The Atlantic (Herrnstein, 1971) and in a subsequentbook, I.Q. in the Meritocracy (1973), Herrnstein elaboratedon an idea by Cyril Burt (1961) that social class and occu-pational differences in IQ will be partly genetic in a soci-ety that features social mobility. If IQ is partly genetic, andhigher IQ individuals tend to move up in social and occupa-tional status, while lower IQ individuals tend to move down,then IQ differences between social classes and occupational

1 History of Behavior Genetics 9

groups will come to be partly genetic. This is not a heredi-tary aristocracy – far from it – it is a dynamic phenomenonthat depends on continued mobility up and down the socialscale. An important question is, How much? Some evidencesuggests that about 40% of IQ differences in occupation andincome in Western societies are associated with genetic dif-ferences (Rowe, Vesterdal, & Rodgers, 1998; Tambs, Sundet,Magnus, & Berg, 1989). Phenotypically, there are substan-tial average differences in IQ between different occupationalgroups. For example, in the U.S. standardization sample forthe 1981 revision of the Wechsler Adult Intelligence Scale,there was a 22-point difference between the average IQs ofpersons in professional and technical occupations and per-sons who were unskilled laborers (Reynolds, Chastain, Kauf-man, & McLean, 1987). And yet there was nearly as muchvariation in IQ within these two occupational groups (stan-dard deviations of 14.4 and 15.2) as in the U.S. populationas a whole (standard deviation of 15.1). It is an interestingparadox that there may be real and significant differences inaverage IQ between different groups, yet individuals vary sowidely within them that an individual’s group membership isof almost no value for predicting his or her IQ.

The Glayde Whitney Affair

In his 1995 presidential address to the Behavior GeneticsAssociation, Glayde Whitney, whose distinguished researchcareer had mostly focused on taste sensitivity in mice, turnedto humans and elected to address the topic of black–whitedifferences in the frequency of criminal behavior. He pointedout the large discrepancies on the phenotypic level, such asa ninefold difference in murder rates between blacks andwhites in the USA. Compared to a dozen other industrializedcountries, the USA had the highest overall murder rate. How-ever, based only on its white population, it ranked third fromthe bottom, with a lower murder rate than such countriesas Switzerland, Denmark, Finland, and Sweden. Whitneyargued that behavior geneticists should be willing to exploreboth genetic and environmental hypotheses about such dif-ferences; he also argued that the current intellectual climatein the USA made such discussion virtually impossible – andhe made some critical remarks about the contribution of thepolitical Left to this situation (Whitney, 1995).

Whitney’s address was perhaps not a model of tact: forexample, in addition to his comments about the Left, henoted that Richmond, Virginia, the city in which he wasspeaking as a guest, was the second-worst large city in theUSA with respect to its murder rate. Nor did he addressthe question of how behavior geneticists were to go aboutdeciding to what extent the group differences in criminalitywere genetic or environmental. Subsequent events within theBehavior Genetics Association proved, however, that he was

clearly right about the difficulty of public discussion of suchquestions. An announcement was issued the next day by theBGA Executive Committee to the effect that Whitney wasnot acting as the official spokesman of the association, thatpresentations at BGA meetings should be strictly scientific,and that “members are not encouraged to express their per-sonal political and moral views” (Heath, 1995, p. 590). Aspecial December meeting of the BGA Executive Committeewas scheduled to consider removing Whitney from the BGABoard of Directors, of which he was automatically a memberas past president (e-mail announcement to the BGA member-ship, October 12, 1995). President-elect Pierre Roubertouxand Wim Crusio, a member-at-large of the Executive Com-mittee, resigned from the association because it was unwill-ing to adopt sufficiently strong sanctions against Whitney.The incoming president-elect, Nicholas Martin, took overfor Roubertoux as president, and later served his own term,accounting for his double appearance in Table 1.1, in 1996and 1997 (Heath, 1996).

Lawrence Summers and Sex Differences

On January 14, 2005, Harvard President Lawrence H. Sum-mers informally addressed a conference on “Diversifying theScience and Engineering Workforce” which was consideringthe reasons for a shortage of women at the highest levels inthe scientific professions (Summers, 2005). With the avowedintention of provoking discussion, Summers proposed threehypotheses for his audience’s consideration: (a) Many tal-ented women prefer devoting some of their time to childrenand families rather than undertaking the 80-hour work-weeksrequired for reaching the top levels in elite research organi-zations; (b) there may be biological differences between thesexes, such as a greater variance for males on many traits,producing an excess of males at the extremes; and (c) subtleand not-so-subtle patterns of discrimination may exist thatlead the present elite in these fields, mostly males, to chooseothers like them to join them. Summers thought it likely thatall three of these factors contributed, and he guessed thatthey might rank in importance in the order given. Summersis an economist by training, not a behavior geneticist, but hecited some behavior genetic evidence against an overwhelm-ing role of socialization in producing behavioral differences,and suggested that the effects in hypotheses (a) and (b) mighthave in part a biological basis. Summers’ remarks aroused afirestorm in the press and in feminist circles, which in turnprovoked assorted indignant rejoinders. It is not necessary topursue these in detail here – a quick survey on the Internetwill yield an ample sampling of widely varying views aboutSummers’ remarks – views expressed with widely varyingdegrees of heat and light. Pinker (2002, Chap. 18) providesa readable survey of the considerable evidence that at least

10 J.C. Loehlin

some male–female psychological differences have a biolog-ical component – although, presumably, few are exclusivelyso, and many questions remain open empirically.

The Future?

One take-home lesson from the various controversies con-cerning group differences is that the nature–nurture contro-versy is not dead, even though it has been declared moribundon many occasions in recent decades. Although behaviorgeneticists have had an appreciable impact on public think-ing about individual differences, the question of the relativegenetic and environmental contributions to group differenceshas been both more socially explosive and much less suc-cessfully addressed empirically.

What does the future hold? This will depend, in part, onfuture behavior genetics research on these topics – some ofit, perhaps, carried out by readers of this book. One maybe fairly confident that nature–nurture controversies will notvanish completely anytime soon. However, one may hopethat as knowledge expands, the cloud of misunderstandingson which these controversies feed will gradually shrink, andthat one day we may have an agreed-upon body of facts onwhich to base social policy.

Conclusion

Yes, behavior genetics has had a long past, which extendsinto the nature–nurture controversies of the present day. Ithas also had a short but solid history of substantive accom-plishment and institutional establishment. The date at whichthe short history will make the long past seem quaint andobsolete in the eyes of the general educated public remainsto be determined. Readers of this book will help determine it.

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Part IQuantitative Methods and Models

Chapter 2

Biometrical Models in Behavioral Genetics

Michael C. Neale

Introduction

The main goal of this chapter is to describe the researchdesigns and statistical methods that are in popular use inbehavioral genetics (BG). We begin with a brief overview ofthe historical background to BG in general and twin studiesin particular. Next, we describe some elementary statisticsrequired for understanding biometrical modeling. Then fol-lows a statistical model for genetic variation, as articulated byFisher in his classic 1918 paper, in which additive and dom-inance genetic variance terms are defined. The coefficientsof resemblance between relatives derived from this modelare then implemented in structural equation models for theanalysis of data from twins and other relatives. Overall theintent is to provide a general and extensible infrastructure forthe modeling of genetically informative data.

Historical Background

Behavior genetics is the synthesis of two domains: behavior,which is defined as the actions or reactions of an object ororganism, and genetics, which is the science of heredity andvariation. The primary focus of contemporary BG is variationin behavior, while broader psychological constructs such asinternal mental states and cognition are frequently included.Individual differences in this activity are readily observed invirtually all forms of animal life and may also be seen incertain plant species, such as Dionaea muscipula (the VenusFly Trap). The ability to predict behavior in other organisms,be they of the same or a different species – would seem tohave substantial survival value. Today, tremendous invest-ment is made by both medical and military agencies in order

M.C. Neale (B)Departments of Psychiatry and Human Genetics, Virginia Institute forPsychiatric and Behavioral Genetics, Virginia CommonwealthUniversity, Richmond, VA 23298-0126, USAe-mail: neale@vou.edu

to understand the origins of behavioral differences, and withgood reason. Many of the most pressing health problems inmodern cultures have behavioral components: obesity, car-diovascular disease, cancer, drug abuse and psychopathologyare obvious examples. It is also the case that human con-flicts, be it a marital dispute, a street fight, or a world war,are primarily behavioral. Thus much of human suffering hasbehavioral origins. One aim of BG is to identify potentialways to alleviate this distress by correctly identifying bothgenetic and environmental sources of individual differencesin behavior and susceptibility to environmental insults.

Behavioral genetics as a field was perhaps first establishedby the exceptional 18th century cousins, Charles Darwin andFrancis Galton. The former, in Chapter 8 of On the Ori-gin of Species (Darwin, 1859) discusses instincts in animalsas diverse as dogs, birds, insects, and notes individual dif-ferences in behavior within species. In his later work, theDescent of Man (Darwin, 1871), he wrote:

If no organic being excepting man had possessed any mentalpower, or if his powers had been of a wholly different naturefrom those of the lower animals, then we should never have beenable to convince ourselves that our high faculties had been grad-ually developed. But it can be shown that there is no fundamentaldifference of this kind.

It is well known that different breeds of dog have differ-ent average temperaments. Typically, Labradors are affec-tionate, Border Collies are intelligent and Bull Terriers areaggressive. Even the most ardent critics of behavioral genet-ics do not seem to quibble with this or any other behavioraldifferences that are observed in either domesticated speciesor those in the wild (Mann, 1994). That selection experi-ments can produce reliable behavioral differences betweenstrains of rats and mice is well established for numeroustraits, including mazesolving ability (Tryon, 1941), activity(Defries, Gervais, & Thomas, 1978), brain weight (Fuller& Herman, 1974), and alcohol preference (Li, Lumeng, &Doolittle, 1993). Selective breeding experiments are essen-tially univariate in design; those with high or low scores onthe single trait of interest are used to populate the next gen-eration. However, it is commonly observed that changes in

Y.-K. Kim (ed.), Handbook of Behavior Genetics, 15DOI 10.1007/978-0-387-76727-7 2, c© Springer Science+Business Media, LLC 2009