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Essays on Genetics and the Social Sciences Genoeconomics, Social-Science Genetics, Or Biological Economics 413 AYSU OKBAY Evidence from behavior-genetic studies of twins, adoptees and other pairs of relatives shows that virtually all human traits, including economic preferences and behaviors, are at least moderately heritable. With increasing availability of genetic data, it is now becoming feasible to identify specific genetic variants associated with complex outcomes, thus blurring the disciplinary boundaries between biological and Essays on Genetics and social sciences. Building on these developments, this thesis explores questions at the intersection of economics and biology and thus contributes to an emerging field of research commonly referred to as: - Essays on Genetics and the Social Sciences OKBAY AYSU genoeconomics, social-science genetics, or biological economics. the Social Sciences The research described here identifies specific genetic variants robustly associated with a suite of complex outcomes – ranging from educational attainment, to subjective well-being, neuroticism and depression – and shows how these findings can be used, in conjunction with quasi-experimental research designs from economics, to conduct rigorous and well-powered investigations of the interactions between genes and environment. It illustrates some hard-won lessons about the relative merits of various research strategies that have been proposed for efforts to discover genetic associations with complex traits. Furthermore, it provides a framework for quantifying tradeoffs between outcome heterogeneity and sample-size in gene-discovery efforts. The results reported in this thesis strongly suggest that there will be many settings in which genetic data will prove valuable to social sciences. In addition to broadening our knowledge about the biological mechanisms underlying behavioral outcomes, these advances will provide researchers from many disciplines with increasingly powerful tools that can be used to integrate genetic factors into a wide range of empirical models. The Erasmus Research Institute of Management (ERIM) is the Research School (Onderzoekschool) in the field of management of the Erasmus University Rotterdam. The founding participants of ERIM are the Rotterdam School of Management (RSM), and the Erasmus School of Economics (ESE). ERIM was founded in 1999 and is officially accredited by the Royal Netherlands Academy of Arts and Sciences (KNAW). The research undertaken by ERIM is focused on the management of the firm in its environment, its intra- and interfirm relations, and its business processes in their interdependent connections. The objective of ERIM is to carry out first rate research in management, and to offer an advanced doctoral programme in Research in Management. Within ERIM, over three hundred senior researchers and PhD candidates are active in the different research programmes. From a variety of academic backgrounds and expertises, the ERIM community is united in striving for excellence and working at the forefront of creating new business knowledge. ERIM PhD Series Research in Management Erasmus University Rotterdam (EUR) Erasmus Research Institute of Management Mandeville (T) Building Burgemeester Oudlaan 50 3062 PA Rotterdam, The Netherlands P.O. Box 1738 3000 DR Rotterdam, The Netherlands T +31 10 408 1182 E [email protected] W www.erim.eur.nl Essays on Genetics and the Social Sciences Essays on Genetics and the Social Sciences Essays over Genetica en de Sociale Wetenschappen THESIS to obtain the degree of Doctor from the Erasmus University Rotterdam by command of the rector magnificus Prof.dr. H.A.P. Pols and in accordance with the decision of the Doctorate Board. The public defense shall be held on Thursday, February 9, 2017, at 13.30 hours by Aysu Okbay born in Istanbul, Turkey. Doctoral Committee Doctoral dissertation supervisors: Prof.dr. A.R. Thurik Prof.dr. P.D. Koellinger Prof.dr. P.J.F. Groenen Other members: Prof.dr. M. Johannesson Prof.dr. D. Posthuma Prof.dr. A. Uitterlinden Erasmus Research Institute of Management – ERIM The joint research institute of the Rotterdam School of Management (RSM) and the Erasmus School of Economics (ESE) at the Erasmus University Rotterdam Internet: http://www.erim.eur.nl ERIM Electronic Series Portal: http://repub.eur.nl/ ERIM PhD Series in Research in Management, 413 ERIM reference number: EPS-2017-413-S&E ISBN 978-90-5892-477-3 © 2017, Aysu Okbay Design: PanArt, www.panart.nl This publication (cover and interior) is printed by Tuijtel on recycled paper, BalanceSilk® The ink used is produced from renewable resources and alcohol free fountain solution. Certifications for the paper and the printing production process: Recycle, EU Ecolabel, FSC®, ISO14001. More info: www.tuijtel.com All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means electronic or mechanical, including photocopying, recording, or by any information storage and retrieval system, without permission in writing from the author. Contents 1 Introduction and Conclusion 1 Motivation 3 Basic Genetic Concepts 5 Research Questions and Main Results 6 Declaration of Contribution 10 Implications and Discussion 12 Conclusion 16 2 On Improving the Credibility of Candidate Gene Studies: A Review of Candidate Gene Studies Published in Emotion 21 Introduction 23 Power and Credibility 24 Model Specification 29 Conclusion 32 3 Genetic Variants Associated with Subjective Well-being, Depressive Symptoms and Neuroticism Identified through Genome-wide Analyses 35 Introduction 37 Results 38 Discussion 55 Supplementary Methods 56 4 Genome-wide Association Study Identifies 74 Loci Associated with Educational Attainment 91 Introduction and Results 93 Supplementary Methods 110 5 Of Genes and Screens: Educational Reform, Ability, and Labor Market Screening 133 Introduction 135 The Swedish school reform 138 Theoretical Framework 139 Data 140 Empirical framework 146 Results 148 Conclusion 155 6 Meta-GWAS Accuracy and Power (MetaGAP) Calculator Shows that Hiding Heritability is Partially due to Imperfect Genetic Correlations across Studies 159 Introduction 161 Materials and Methods 164 Results 169 Discussion 181 Supplementary Methods 185 Appendix A: Supplementary Tables to Chapter 3 193 Appendix B: Supplementary Tables to Chapter 4 215 Appendix C: Supplementary Material to Chapter 5 235 Summary 249 Samenvatting (Summary in Dutch) 251 References 253 Acknowledgments 273 PhD Portfolio 277 About the Author 275 CHAPTER 1 Introductionintroduction and conclusion and Conclusion 2 INTRODUCTION AND CONCLUSION Abstract This thesis consists of six chapters that examine the genetic architecture of complex behav- ioral outcomes. Chapter 1 lists and substantiates the research questions and gives a summary overview of the key findings and their implications. Chapter 2 discusses methodological challenges with traditional candidate gene approaches, which until recently were the domi- nant paradigm for gene discovery in the social sciences. The chapter explains why well- known problems of low statistical power, population stratification and undisclosed multiple- hypothesis testing render most of the existing evidence from candidate gene studies unreli- able. Each potential problem is illustrated with reference to specific studies published by the journal Emotion. Recognizing these limitations, most modern gene discovery efforts are ge- nome-wide association studies (GWAS), in which millions of markers are tested for associ- ation with the outcome of interest in large samples and with stringent controls for population stratification. Chapters 3 and 4 report the results of two such studies, performed by meta- analyzing summary statistics from a large number of cohorts with combined sample sizes in the hundreds of thousands. The first study (Chapter 3) examines four genetically correlated phenotypes related to well-being (depressive symptoms, life satisfaction, neuroticism and positive affect) and identifies 14 genome-wide significant associations. The second study (Chapter 4) examines educational attainment and identifies 74 genome-wide significant as- sociations. Chapter 5 builds on these findings by exploiting the staggered rollout of an ex- pansion in comprehensive schooling in Sweden to test for gene-by-environment interactions, finding that the impact of the reform on educational outcomes and earnings was heteroge- neous by genotype for females: females with higher polygenic scores (derived using the results from Chapter 4) were more likely to obtain a high school degree, which is beyond the new minimum established by the reform. Chapter 6 develops a theoretical multi-study framework relating statistical power and predictive accuracy to cross-study heterogeneity. The chapter shows that in GWAS meta-analyses, including those in Chapters 3 and 4, cross- cohort genetic correlations lower than unity can have a substantial effect on statistical power, and predictive accuracy of polygenic scores derived from meta-analyses of summary statis- tics from heterogenous cohorts. The methodology developed in Chapter 6 is easily imple- mented using an online tool that researchers can use to quantify the tradeoff between gains from larger sample sizes and the potential losses in power from pooling measures that are too heterogeneous. M OTIVAT ION 3 Motivation Evidence from behavior-genetic studies of twins, adoptees and other pairs of relatives shows that virtually all human traits, including economic preferences and behaviors, are at least moderately heritable (Polderman et al., 2015; Sacerdote, 2010; Turkheimer,
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