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Genomics yields biological and phenotypic insights into bipolar disorder

Abstract

Bipolar disorder is a leading contributor to the global burden of disease1. Despite high heritability (60–80%), the majority of the underlying genetic determinants remain unknown2. We analysed data from participants of European, East Asian, African American and Latino ancestries (n = 158,036 cases with bipolar disorder, 2.8 million controls), combining clinical, community and self-reported samples. We identified 298 genome-wide significant loci in the multi-ancestry meta-analysis, a fourfold increase over previous findings3, and identified an ancestry-specific association in the East Asian cohort. Integrating results from fine-mapping and other variant-to-gene mapping approaches identified 36 credible genes in the aetiology of bipolar disorder. Genes prioritized through fine-mapping were enriched for ultra-rare damaging missense and protein-truncating variations in cases with bipolar disorder4, highlighting convergence of common and rare variant signals. We report differences in the genetic architecture of bipolar disorder depending on the source of patient ascertainment and on bipolar disorder subtype (type I or type II). Several analyses implicate specific cell types in the pathophysiology of bipolar disorder, including GABAergic interneurons and medium spiny neurons. Together, these analyses provide additional insights into the genetic architecture and biological underpinnings of bipolar disorder.

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Fig. 1: Genetic correlation and bivariate MiXeR estimates for the genetic overlap of BD ascertainment and subtypes.

Fig. 2: Genetic correlations (with standard errors) between BD and other psychiatric disorders.

Fig. 3: Phenotypic variance in BD in EUR cohorts explained by PRSs derived from the multi-ancestry and EUR meta-analyses (with and without self-reported data).

Fig. 4: Supercluster-level SNP-h2 enrichment for BD.

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Data availability

Genome-wide association summary statistics for these analyses are available at https://www.med.unc.edu/pgc/download-results/. The full GWAS summary statistics for the 23andMe datasets will be made available through 23andMe to qualified researchers under an agreement with 23andMe that protects the privacy of the 23andMe participants. Please visit https://research.23andme.com/collaborate/#dataset-access for more information and to apply to access the data. After applying with 23andMe, the full summary statistics including all analysed SNPs and samples in the GWAS meta-analyses will be accessible to the approved researchers. Genotype data are available for a subset of cohorts, including dbGAP accession numbers and/or restrictions, as described in the ‘Cohort descriptions’ section of the supplementary materials.

Code availability

No custom code was developed for this study. All software and tools used for the analyses presented are publicly available and referenced within the respective sections in the Methods of the article.

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Acknowledgements

We thank the participants who donated their time, life experiences and DNA to this research, the clinical and scientific teams that worked with them, and the investigators who comprise the PGC. The PGC has received major funding from the US National Institute of Mental Health (PGC4: R01 MH124839, PGC3: U01 MH109528, PGC2: U01 MH094421 and PGC1: U01 MH085520). Statistical analyses were carried out on the NL Genetic Cluster Computer (http://www.geneticcluster.org) hosted by SURFsara. The content is solely the responsibility of the authors and does not necessarily represent the official views of the US National Institutes of Health. Individual and cohort-specific funding acknowledgements are detailed in the Supplementary Information.

Author information

Author notes

These authors contributed equally: Maria Koromina, Tracey van der Veen, Toni Boltz, Friederike S. David, Jessica Mei Kay Yang, Keng-Han Lin, Xin Wang, Jonathan R. I. Coleman, Brittany L. Mitchell, Caroline C. McGrouther, Aaditya V. Rangan, Penelope A. Lind, Elise Koch, Arvid Harder, Nadine Parker, Jaroslav Bendl

These authors jointly supervised this work: Andrew McQuillin, Andreas J. Forstner, Niamh Mullins, Arianna Di Florio, Roel A. Ophoff, Ole A. Andreassen

Lists of members and their affiliations appear in the Supplementary Information

Authors and Affiliations

Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway

Kevin S. O’Connell, Elise Koch, Nadine Parker, Oleksandr Frei, Alexey Shadrin, Olav B. Smeland, Markos Tesfaye, Martin Tesli, Ingrid Melle & Ole A. Andreassen

Center for Precision Psychiatry, University of Oslo, Oslo, Norway

Kevin S. O’Connell, Elise Koch, Nadine Parker, Ole Kristian Drange, Alexey Shadrin, Olav B. Smeland, Markos Tesfaye, Martin Tesli & Ole A. Andreassen

Department of Psychiatry, Icahn School of Medicine at Mount Sinai, New York, NY, USA

Maria Koromina, Jaroslav Bendl, Tereza Clarence, John F. Fullard, Jessica S. Johnson, Dolores Malaspina, Laura G. Sloofman, René S. Kahn, Panos Roussos, Eli A. Stahl & Niamh Mullins

Charles Bronfman Institute for Personalized Medicine, Icahn School of Medicine at Mount Sinai, New York, NY, USA

Maria Koromina & Niamh Mullins

Department of Genetics and Genomic Sciences, Icahn School of Medicine at Mount Sinai, New York, NY, USA

Maria Koromina, Jaroslav Bendl, Tereza Clarence, John F. Fullard, Jessica S. Johnson, Dolores Malaspina, Laura G. Sloofman, Panos Roussos, Eli A. Stahl & Niamh Mullins

Division of Psychiatry, University College London, London, UK

Tracey van der Veen, Nicholas Bass, Andrew McQuillin & Tracey van der Veen

Department of Human Genetics, David Geffen School of Medicine, University of California Los Angeles, Los Angeles, CA, USA

Toni Boltz & Roel A. Ophoff

Institute of Human Genetics, University of Bonn, School of Medicine and University Hospital Bonn, Bonn, Germany

Friederike S. David, Eva C. Beins, Isabelle Claus, Franziska Degenhardt, Stefan Herms, Per Hoffmann, Eva C. Schulte, Lisa Sindermann, Sven Cichon, Markus M. Nöthen & Andreas J. Forstner

Department of Psychiatry and Psychotherapy, University of Marburg, Marburg, Germany

Friederike S. David, Frederike Stein & Tilo Kircher

Centre for Neuropsychiatric Genetics and Genomics, Division of Psychological Medicine and Clinical Neurosciences, Cardiff University, Cardiff, UK

Jessica Mei Kay Yang, Valentina Escott-Price, Liz Forty, Peter A. Holmans, Antonio F. Pardiñas, James T. R. Walters, Nicholas Craddock, Ian Jones, George Kirov, Michael C. O’Donovan, Michael J. Owen & Arianna Di Florio

23andMe Inc., Sunnyvale, CA, USA

Keng-Han Lin & Xin Wang

Social, Genetic and Developmental Psychiatry Centre, King’s College London, London, UK

Jonathan R. I. Coleman, Gerome Breen & Panagiotis Ferentinos

NIHR Maudsley BRC, King’s College London, London, UK

Jonathan R. I. Coleman & Gerome Breen

Mental Health and Neuroscience, QIMR Berghofer Medical Research Institute, Brisbane, Queensland, Australia

Brittany L. Mitchell, Penelope A. Lind, Jackson G. Thorp & Sarah E. Medland

School of Biomedical Sciences and Faculty of Medicine, The University of Queensland, Brisbane, Queensland, Australia

Brittany L. Mitchell & Penelope A. Lind

New York University, New York, NY, USA

Caroline C. McGrouther & Aaditya V. Rangan

Flatiron Institute, New York, NY, USA

Aaditya V. Rangan

School of Biomedical Sciences, Queensland University of Technology, Brisbane, Queensland, Australia

Penelope A. Lind

Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden

Arvid Harder, Dennis Hellgren, Mikael Landén & Patrick F. Sullivan

Center for Disease Neurogenomics, Icahn School of Medicine at Mount Sinai, New York, NY, USA

Jaroslav Bendl, Tereza Clarence, John F. Fullard & Panos Roussos

Friedman Brain Institute, Icahn School of Medicine at Mount Sinai, New York, NY, USA

Jaroslav Bendl, Tereza Clarence, John F. Fullard & Panos Roussos

Institute of Psychiatric Phenomics and Genomics (IPPG), LMU University Hospital, LMU Munich, Munich, Germany

Kristina Adorjan, Monika Budde, Urs Heilbronner, Janos L. Kalman, Sergi Papiol, Eva C. Schulte, Fanny Senner & Thomas G. Schulze

Department of Psychiatry and Psychotherapy, University Hospital, LMU Munich, Munich, Germany

Kristina Adorjan, Janos L. Kalman, Sergi Papiol & Fanny Senner

University Hospital of Psychiatry and Psychotherapy, University of Bern, Bern, Switzerland

Kristina Adorjan

iPSYCH, The Lundbeck Foundation Initiative for Integrative Psychiatric Research, Aarhus, Denmark

Esben Agerbo, Thomas D. Als, Carsten Bøcker Pedersen, Jonas Bybjerg-Grauholm, Jakob Grove, Manuel Mattheisen, Anders D. Børglum, David M. Hougaard, Ole Mors, Preben Bo Mortensen, Merete Nordentoft & Thomas Werge

National Centre for Register-Based Research, Aarhus University, Aarhus, Denmark

Esben Agerbo & Carsten Bøcker Pedersen

Centre for Integrated Register-based Research, Aarhus University, Aarhus, Denmark

Esben Agerbo & Carsten Bøcker Pedersen

Department of Neuroscience, Istituto Di Ricerche Farmacologiche Mario Negri IRCCS, Milano, Italy

Diego Albani

Instituto de Salud Carlos III, Biomedical Network Research Centre on Mental Health (CIBERSAM), Madrid, Spain

Silvia Alemany, Rosa Bosch, Judit Cabana-Domínguez, Sergi Papiol, Maria Soler Artigas, Josep Antoni Ramos-Quiroga & Marta Ribasés

Department of Psychiatry, Hospital Universitari Vall d’Hebron, Barcelona, Spain

Silvia Alemany, Judit Cabana-Domínguez, Maria Soler Artigas, Josep Antoni Ramos-Quiroga & Marta Ribasés

Psychiatric Genetics Unit, Group of Psychiatry Mental Health and Addictions, Vall d’Hebron Research Institut (VHIR), Universitat Autònoma de Barcelona, Barcelona, Spain

Silvia Alemany, Judit Cabana-Domínguez, Maria Soler Artigas, Josep Antoni Ramos-Quiroga & Marta Ribasés

Department of Psychiatry and Behavioral Neuroscience, University of Chicago, Chicago, IL, USA

Ney Alliey-Rodriguez & Elliot S. Gershon

Northwestern University, Chicago, IL, USA

Ney Alliey-Rodriguez

iSEQ, Center for Integrative Sequencing, Aarhus University, Aarhus, Denmark

Thomas D. Als & Manuel Mattheisen

Department of Biomedicine-Human Genetics, Aarhus University, Aarhus, Denmark

Thomas D. Als & Manuel Mattheisen

Department of Neurology, Klinikum rechts der Isar, School of Medicine, Technical University of Munich, Munich, Germany

Till F. M. Andlauer

National and Kapodistrian University of Athens, 2nd Department of Psychiatry, Attikon General Hospital, Athens, Greece

Anastasia Antoniou, Konstantinos Dafnas & Panagiotis Ferentinos

PsychGen Centre for Genetic Epidemiology and Mental Health, Norwegian Institute of Public Health, Oslo, Norway

Helga Ask, Elizabeth C. Corfield & Alexandra Havdahl

PROMENTA Research Centre, Department of Psychology, University of Oslo, Oslo, Norway

Helga Ask & Alexandra Havdahl

Department of Psychiatry and Psychotherapy, University Hospital Carl Gustav Carus, Technische Universität Dresden, Dresden, Germany

Michael Bauer & Andrea Pfennig

Department of Psychiatry and Behavioral Sciences, SUNY Downstate Health Sciences University, Brooklyn, NY, USA

Tim B. Bigdeli & Chris Chatzinakos

VA NY Harbor Healthcare System, Brooklyn, NY, USA

Tim B. Bigdeli

Institute for Genomics in Health, SUNY Downstate Health Sciences University, Brooklyn, NY, USA

Tim B. Bigdeli & Chris Chatzinakos

Department of Epidemiology and Biostatistics, School of Public Health, SUNY Downstate Health Sciences University, Brooklyn, NY, USA

Tim B. Bigdeli

Psychiatry, Brain Center UMC Utrecht, Utrecht, The Netherlands

Marco P. Boks, Annabel Vreeker & René S. Kahn

Research and Communication Unit for Musculoskeletal Health, Division of Clinical Neuroscience, Oslo University Hospital, Ullevål, Oslo, Norway

Sigrid Børte

Institute of Clinical Medicine, University of Oslo, Oslo, Norway

Sigrid Børte, Oleksandr Frei & John-Anker Zwart

HUNT Center for Molecular and Clinical Epidemiology, Department of Public Health and Nursing, Faculty of Medicine and Health Sciences, Norwegian University of Science and Technology, Trondheim, Norway

Sigrid Børte, Bendik S. Winsvold & John-Anker Zwart

Programa SJD MIND Escoles, Hospital Sant Joan de Déu, Institut de Recerca Sant Joan de Déu, Esplugues de Llobregat, Spain

Rosa Bosch & Miquel Casas

Department of Psychiatry, Psychosomatic Medicine and Psychotherapy, University Hospital Frankfurt, Frankfurt am Main, Germany

Murielle Brum, Nathalie Brunkhorst-Kanaan, Thorsten M. Kranz & Andreas Reif

K. G. Jebsen Center for Genetic Epidemiology, Department of Public Health and Nursing, Faculty of Medicine and Health Sciences, Norwegian University of Science and Technology, Trondheim, Norway

Ben M. Brumpton & Kristian Hveem

Center for Neonatal Screening, Department for Congenital Disorders, Statens Serum Institut, Copenhagen, Denmark

Jonas Bybjerg-Grauholm & David M. Hougaard

Psychiatry, University of California San Francisco, San Francisco, CA, USA

William Byerley

School of Biomedical Sciences and Pharmacy, The University of Newcastle, Callaghan, New South Wales, Australia

Murray J. Cairns

Precision Medicine Research Program, Hunter Medical Research Institute, New Lambton, New South Wales, Australia

Murray J. Cairns

Section of Psychiatry, Department of Medical Sciences and Public Health, University of Cagliari, Cagliari, Italy

Bernardo Carpiniello & Mirko Manchia

Department of Psychiatry and Forensic Medicine, Universitat Autònoma de Barcelona, Barcelona, Spain

Miquel Casas & Josep Antoni Ramos-Quiroga

Fundació Privada d’Investigació Sant Pau (FISP), Barcelona, Spain

Miquel Casas

Department of Psychiatry, Mood Disorders Program, McGill University Health Center, Montreal, Québec, Canada

Pablo Cervantes & Cristiana Cruceanu

Department of Psychiatry, National Taiwan University Hospital, Taipei, Taiwan

Hsi-Chung Chen & Po-Hsiu Kuo

Department of Psychiatry, College of Medicine, National Taiwan University, Taipei, Taiwan

Hsi-Chung Chen

Division of Psychiatry, University of Edinburgh, Edinburgh, UK

Toni-Kim Clarke

Department of Quantitative Health Sciences Research, Mayo Clinic, Rochester, MN, USA

Brandon Coombes & Joanna M. Biernacka

Nic Waals Institute, Lovisenberg Diaconal Hospital, Oslo, Norway

Elizabeth C. Corfield & Alexandra Havdahl

Department of Genetics and Bioinformatics, Norwegian Institute of Public Health, Oslo, Norway

Elizabeth C. Corfield

Department of Physiology and Pharmacology, Karolinska Institutet, Stockholm, Sweden

Cristiana Cruceanu

Department of Psychiatry, Universidad Autonoma de Nuevo Leon, Monterrey, Mexico

Alfredo Cuellar-Barboza

Department of Psychiatry and Psychology, Mayo Clinic, Rochester, MN, USA

Alfredo Cuellar-Barboza, Joanna M. Biernacka & Mark Frye

Department of Psychiatry, Laboratory of Psychiatric Genetics, Poznan University of Medical Sciences, Poznan, Poland

Piotr M. Czerski

Center for Multimodal Imaging and Genetics, Departments of Neurosciences, Radiology, and Psychiatry, University of California, San Diego, CA, USA

Anders M. Dale

Division of Psychiatry and Psychotherapeutic Medicine, Medical University of Graz, Graz, Austria

Nina Dalkner, Frederike T. Fellendorf, Susanne Bengesser & Eva Z. Reininghaus

Department of Child and Adolescent Psychiatry, Psychosomatics and Psychotherapy, University Hospital Essen, University of Duisburg-Essen, Duisburg, Germany

Franziska Degenhardt

Department of Psychiatry and Behavioral Sciences, Johns Hopkins University School of Medicine, Baltimore, MD, USA

J. Raymond DePaulo, James B. Potash, Peter P. Zandi, Fernando S. Goes & Thomas G. Schulze

Department of Medical Genetics, Oslo University Hospital Ullevål, Oslo, Norway

Srdjan Djurovic

Department of Clinical Science, University of Bergen, Bergen, Norway

Srdjan Djurovic & Markos Tesfaye

Department of Psychiatry, Sørlandet Hospital, Kristiansand, Norway

Ole Kristian Drange

Department of Psychiatry, University of Arizona College of Medicine-Phoenix, Phoenix, AZ, USA

Ayman H. Fanous

Carl T. Hayden Veterans Affairs Medical Center, Phoenix, AZ, USA

Ayman H. Fanous

Banner-University Medical Center, Phoenix, AZ, USA

Ayman H. Fanous

Academic Psychiatry, Newcastle University, Newcastle upon Tyne, UK

I. Nicol Ferrier

Department of Genetic Epidemiology in Psychiatry, Central Institute of Mental Health, Medical Faculty Mannheim, Heidelberg University, Mannheim, Germany

Josef Frank, Lea Sirignano, Stephanie H. Witt, Lea Zillich, Marcella Rietschel, Thomas G. Schulze & Fabian Streit

Center for Neurobehavioral Genetics, Semel Institute for Neuroscience and Human Behavior, Los Angeles, CA, USA

Nelson B. Freimer & Roel A. Ophoff

Department of Psychiatry and Biobehavioral Science, Semel Institute, David Geffen School of Medicine, University of California, Los Angeles, Los Angeles, CA, USA

Nelson B. Freimer & Roel A. Ophoff

Department of Psychiatry, Dalhousie University, Halifax, Nova Scotia, Canada

Julie Garnham, Claire O’Donovan, Claire Slaney, Evangelia Eirini Tsermpini & Martin Alda

Department of Psychological Sciences, University of Missouri, Columbia, MO, USA

Ian R. Gizer

Genetics and Computational Biology, QIMR Berghofer Medical Research Institute, Brisbane, Queensland, Australia

Scott D. Gordon & Nicholas G. Martin

Psychological Medicine, University of Worcester, Worcester, UK

Katherine Gordon-Smith, Amy Perry & Lisa A. Jones

Department of Psychiatry, University of California San Diego, La Jolla, CA, USA

Tiffany A. Greenwood, Adam X. Maihofer, Paul D. Shilling, John R. Kelsoe & Caroline M. Nievergelt

Department of Biomedicine and the iSEQ Center, Aarhus University, Aarhus, Denmark

Jakob Grove & Anders D. Børglum

Center for Genomics and Personalized Medicine, CGPM, Aarhus, Denmark

Jakob Grove & Anders D. Børglum

Bioinformatics Research Centre, Aarhus University, Aarhus, Denmark

Jakob Grove

Mental Health Department, University Regional Hospital, Biomedicine Institute (IBIMA), Málaga, Spain

José Guzman-Parra & Fermin Mayoral

Department of Neuropsychiatry, Seoul National University Bundang Hospital, Seongnam, Republic of Korea

Tae Hyon Ha, Euitae Kim, Jaeyoung Kim, Hye Youn Park & Woojae Myung

Department of Neuropsychiatry, Seoul National University College of Medicine, Seoul, Republic of Korea

Tae Hyon Ha, Euitae Kim, Hye Youn Park & Woojae Myung

Institute for Translational Psychiatry, University of Münster, Münster, Germany

Tim Hahn & Udo Dannlowski

Faculty of Medicine, Department of Psychiatry, School of Health Sciences, University of Iceland, Reykjavik, Iceland

Magnus Haraldsson & Engilbert Sigurdsson

Landspitali University Hospital, Reykjavik, Iceland

Magnus Haraldsson & Engilbert Sigurdsson

Department of Psychology, Eberhard Karls Universität Tübingen, Tubingen, Germany

Martin Hautzinger

Department of Biomedicine, University of Basel, Basel, Switzerland

Stefan Herms, Per Hoffmann, Thomas W. Mühleisen & Sven Cichon

Institute of Medical Genetics and Pathology, University Hospital Basel, Basel, Switzerland

Stefan Herms, Per Hoffmann & Sven Cichon

Brain and Mind Centre, The University of Sydney, Sydney, New South Wales, Australia

Ian B. Hickie

Department of Psychiatry, Taipei City Psychiatric Center, Taipei City Hospital, Taipei, Taiwan

Ming-Chyi Huang

Department of Psychiatry, Nagoya University Graduate School of Medicine, Nagoya, Japan

Masashi Ikeda

Department of Psychiatry, Fujita Health University School of Medicine, Toyoake, Japan

Masashi Ikeda, Nakao Iwata & Takeo Saito

Université Paris Est Créteil, INSERM, IMRB, Translational Neuropsychiatry, Créteil, France

Stéphane Jamain & Marion Leboyer

Department of Psychiatry, UNC Chapel Hill School of Medicine, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA

Jessica S. Johnson

Institute of Neuroscience and Physiology, University of Gothenburg, Gothenburg, Sweden

Lina Jonsson & Mikael Landén

Laboratory of Complex Trait Genomics, Department of Computational Biology and Medical Sciences, Graduate School of Frontier Sciences, The University of Tokyo, Tokyo, Japan

Yoichiro Kamatani

Laboratory for Statistical and Translational Genetics, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan

Yoichiro Kamatani & Chikashi Terao

Campbell Family Mental Health Research Institute, Centre for Addiction and Mental Health, Toronto, Ontario, Canada

James L. Kennedy & John B. Vincent

Neurogenetics Section, Centre for Addiction and Mental Health, Toronto, Ontario, Canada

James L. Kennedy

Department of Psychiatry, University of Toronto, Toronto, Ontario, Canada

James L. Kennedy

Institute of Medical Sciences, University of Toronto, Toronto, Ontario, Canada

James L. Kennedy

Department of Brain and Cognitive Sciences, Seoul National University College of Natural Sciences, Seoul, Republic of Korea

Euitae Kim

Samsung Advanced Institute for Health Sciences and Technology (SAIHST), Sungkyunkwan University, Samsung Medical Center, Seoul, Republic of Korea

Jaeyoung Kim & Hong-Hee Won

Department of Psychiatry and Neurobehavioral Science, University College Cork, Cork, Ireland

Sarah Kittel-Schneider

Department of Psychiatry, Psychosomatics and Psychotherapy, Center of Mental Health, University Hospital Würzburg, Würzburg, Germany

Sarah Kittel-Schneider, Manuel Mattheisen & Micha Gawlik

Human Genetics Institute of New Jersey, Rutgers University, Piscataway, NJ, USA

James A. Knowles

ISGlobal, Barcelona, Spain

Manolis Kogevinas

Estonian Genome Centre, Institute of Genomics, University of Tartu, Tartu, Estonia

Kristi Krebs & Lili Milani

Department of Psychiatry, Erasmus MC, University Medical Center Rotterdam, Rotterdam, The Netherlands

Steven A. Kushner

Translational Psychiatry, Department of Molecular Medicine and Surgery, Karolinska Institutet, Stockholm, Sweden

Catharina Lavebratt, Martin Lundberg, Vincent Millischer, Lena Backlund & Martin Schalling

Center for Molecular Medicine, Karolinska University Hospital, Stockholm, Sweden

Catharina Lavebratt, Martin Lundberg, Vincent Millischer, Lena Backlund & Martin Schalling

Psychiatry, North East London NHS Foundation Trust, Ilford, UK

Jacob Lawrence

Clinic for Psychiatry and Psychotherapy, University Hospital Cologne, Cologne, Germany

Markus Leber

Department of Psychiatry, Korea University College of Medicine, Seoul, Republic of Korea

Heon-Jeong Lee

Analytic and Translational Genetics Unit, Massachusetts General Hospital, Boston, MA, USA

Calwing Liao, Robert Ye, Benjamin M. Neale & Stephan Ripke

Stanley Center for Psychiatric Research, Broad Institute, Cambridge, MA, USA

Calwing Liao, Robert Ye, Benjamin M. Neale, Stephan Ripke & Jordan W. Smoller

Department of Translational Research in Psychiatry, Max Planck Institute of Psychiatry, Munich, Germany

Susanne Lucae & Bertram Müller-Myhsok

Division of Psychiatry, Centre for Clinical Brain Sciences, The University of Edinburgh, Edinburgh, UK

Donald J. MacIntyre & Andrew M. McIntosh

Department of Psychiatry and Psychotherapy, University of Bonn, School of Medicine and University Hospital Bonn, Bonn, Germany

Wolfgang Maier & Eva C. Schulte

Research/Psychiatry, Veterans Affairs San Diego Healthcare System, San Diego, CA, USA

Adam X. Maihofer & Caroline M. Nievergelt

Unit of Clinical Psychiatry, University Hospital Agency of Cagliari, Cagliari, Italy

Mirko Manchia

National and Kapodistrian University of Athens, Medical School, Clinical Biochemistry Laboratory, Attikon General Hospital, Athens, Greece

Eirini Maratou & Paraskevi Moutsatsou

Department of Clinical Neuroscience, Karolinska Institutet, Stockholm, Sweden

Lina Martinsson & Tomas Olsson

Centre for Psychiatry Research, SLSO Region Stockholm, Stockholm, Sweden

Lina Martinsson

Department of Clinical Neuroscience, Centre for Psychiatry Research, Karolinska Institutet, Stockholm, Sweden

Manuel Mattheisen

Human and Systems Genetics Working Group, Department of Genetics, Stellenbosch University, Stellenbosch, South Africa

Nathaniel W. McGregor

Department of Psychiatry, University of Michigan, Ann Arbor, MI, USA

Melvin G. McInnis

Genetic Cancer Susceptibility Group, International Agency for Research on Cancer, Lyon, France

James D. McKay

Institute for Genomic Health, SUNY Downstate Medical Center College of Medicine, Brooklyn, NY, USA

Helena Medeiros

Department of Psychiatry and Psychotherapy, Central Institute of Mental Health, Medical Faculty Mannheim, University of Heidelberg, Mannheim, Germany

Andreas Meyer-Lindenberg & Fabian Streit

German Centre for Mental Health (DZPG), partner site Mannheim-Heidelberg-Ulm, Mannheim, Germany

Andreas Meyer-Lindenberg

Department of Psychiatry and Psychotherapy, Clinical Division of General Psychiatry, Medical University of Vienna, Vienna, Austria

Vincent Millischer

Comprehensive Center for Clinical Neurosciences and Mental Health, Medical University of Vienna, Vienna, Austria

Vincent Millischer

Centre for Neuroimaging and Cognitive Genomics (NICOG), School of Biological and Chemical Sciences, University of Galway, Galway, Ireland

Derek W. Morris

Institute of Neuroscience and Medicine (INM-1), Research Centre Jülich, Jülich, Germany

Thomas W. Mühleisen, Sven Cichon & Andreas J. Forstner

Population Health, QIMR Berghofer Medical Research Institute, Brisbane, Queensland, Australia

Catherine M. Olsen & David C. Whiteman

Department of Psychiatry and Psychotherapy, Charité-Universitätsmedizin, Berlin, Germany

Georgia Panagiotaropoulou & Stephan Ripke

Department of Biomedical Sciences, University of Cagliari, Cagliari, Italy

Claudia Pisanu & Alessio Squassina

Oxford Health NHS Foundation Trust, Warneford Hospital, Oxford, UK

Digby Quested

Department of Psychiatry, University of Oxford, Warneford Hospital, Oxford, UK

Digby Quested

Department of Psychiatry and Behavioral Sciences, Emory University School of Medicine, Atlanta, GA, USA

Mark H. Rapaport

Outpatient Clinic for Bipolar Disorder, Altrecht, Utrecht, The Netherlands

Eline J. Regeer

Department of Psychiatry, Washington University in Saint Louis, Saint Louis, MO, USA

John P. Rice

Department of Biochemistry and Molecular Biology II, Faculty of Pharmacy, University of Granada, Granada, Spain

Margarita Rivera

Institute of Neurosciences ‘Federico Olóriz’, Biomedical Research Center (CIBM), University of Granada, Granada, Spain

Margarita Rivera

Instituto de Investigación Biosanitaria ibs.GRANADA, Granada, Spain

Margarita Rivera

KG Jebsen Centre for Neurodevelopmental disorders, University of Oslo, Oslo, Norway

Alexey Shadrin

Faculty of Medicine, University of Queensland, Brisbane, Queensland, Australia

Dan Siskind & Dan Siskind

Division of Psychiatry, Centre for Clinical Brain Sciences, University of Edinburgh, Edinburgh, UK

Daniel J. Smith

Psychiatry and the Behavioral Sciences, University of Southern California, Los Angeles, CA, USA

Janet L. Sobell

Department of Genetics, Microbiology, and Statistics, Faculty of Biology, Universitat de Barcelona, Barcelona, Spain

Maria Soler Artigas & Marta Ribasés

SAMRC Unit on Risk and Resilience in Mental Disorders, Dept of Psychiatry and Neuroscience Institute, University of Cape Town, Cape Town, South Africa

Dan J. Stein

Virginia Institute for Psychiatric and Behavioral Genetics, Virginia Commonwealth University, Richmond, VA, USA

Mei-Hsin Su

Human Genetics Branch, Intramural Research Program, National Institute of Mental Health, Bethesda, MD, USA

Heejong Sung & Francis J. McMahon

Department of Environmental Epidemiology, Nofer Institute of Occupational Medicine, Lodz, Poland

Beata Świątkowska

Department of Mental Disorders, Norwegian Institute of Public Health, Oslo, Norway

Martin Tesli

deCODE Genetics/Amgen, Reykjavik, Iceland

Thorgeir E. Thorgeirsson, Hreinn Stefansson & Kari Stefansson

Neuroscience Research Australia, Sydney, New South Wales, Australia

Claudio Toma, Janice M. Fullerton, Melissa J. Green, Peter R. Schofield, Cynthia S. Weickert & Thomas W. Weickert

Discipline of Psychiatry and Mental Health, School of Clinical Medicine, Faculty of Medicine and Health, University of New South Wales, Sydney, New South Wales, Australia

Claudio Toma, Vaughan J. Carr, Melissa J. Green, Philip B. Mitchell, Cynthia S. Weickert & Thomas W. Weickert

Centro de Biología Molecular Severo Ochoa, Universidad Autónoma de Madrid and CSIC, Madrid, Spain

Claudio Toma

Department of Psychiatry, Harvard Medical School, Boston, MA, USA

Leonardo Tondo

School of Biomedical Science and Pharmacy, University of Newcastle, Newcastle, New South Wales, Australia

Paul A. Tooney

Department of Psychiatry, Taipei Veterans General Hospital, Taipei, Taiwan

Shih-Jen Tsai

Division of Psychiatry, National Yang Ming Chiao Tung University, Taipei, Taiwan

Shih-Jen Tsai

Department of Psychiatry and Human Behavior, School of Medicine, University of California, Irvine, CA, USA

Marquis P. Vawter

Psychiatry, Psychiatrisches Zentrum Nordbaden, Wiesloch, Germany

Helmut Vedder

Department of Child and Adolescent Psychiatry/Psychology, Erasmus MC Sophia Children Hospital, Erasmus University, Rotterdam, The Netherlands

Annabel Vreeker

Department of Psychology Education and Child Studies, Erasmus School of Social and Behavioral Sciences, Erasmus University Rotterdam, Rotterdam, The Netherlands

Annabel Vreeker

Department of Research, Innovation and Education, Division of Clinical Neuroscience, Oslo University Hospital, Oslo, Norway

Bendik S. Winsvold & John-Anker Zwart

Department of Neurology, Oslo University Hospital, Oslo, Norway

Bendik S. Winsvold

Samsung Genome Institute, Samsung Medical Center, Sungkyunkwan University School of Medicine, Seoul, Republic of Korea

Hong-Hee Won

Department of Psychological Medicine, Institute of Psychiatry, Psychology and Neuroscience, King’s College London, London, UK

Allan H. Young

South London and Maudsley NHS Foundation Trust, Bethlem Royal Hospital, Kent, UK

Allan H. Young

Department of Clinical Sciences, Psychiatry, Umeå University Medical Faculty, Umeå, Sweden

Rolf Adolfsson

National Institute of Mental Health, Klecany, Czech Republic

Martin Alda

Institute of Environmental Medicine, Karolinska Institutet, Stockholm, Sweden

Lars Alfredsson

Department of Psychiatry, University of Münster, Münster, Germany

Bernhard T. Baune

Department of Psychiatry, Melbourne Medical School, The University of Melbourne, Melbourne, Victoria, Australia

Bernhard T. Baune

The Florey Institute of Neuroscience and Mental Health, The University of Melbourne, Parkville, Victoria, Australia

Bernhard T. Baune & Christos Pantelis

Université Paris Cité, INSERM, Optimisation Thérapeutique en Neuropsychopharmacologie, UMRS-1144, Paris, France

Frank Bellivier & Bruno Etain

APHP Nord, DMU Neurosciences, GHU Saint Louis-Lariboisière-Fernand Widal, Département de Psychiatrie et de Médecine Addictologique, Paris, France

Frank Bellivier & Bruno Etain

Psychiatry, University of Pennsylvania, Philadelphia, PA, USA

Wade H. Berrettini

Center for Statistical Genetics and Department of Biostatistics, University of Michigan, Ann Arbor, MI, USA

Michael Boehnke & Laura J. Scott

University of Queensland, Brisbane, Queensland, Australia

Stanley Catts

Neuropsychiatric Genetics Research Group, Department of Psychiatry and Trinity Translational Medicine Institute, Trinity College Dublin, Dublin, Ireland

Aiden Corvin

National and Kapodistrian University of Athens, 1st Department of Psychiatry, Eginition Hospital, Athens, Greece

Dimitris Dikeos

School of Biomedical Sciences, Faculty of Medicine and Health, University of New South Wales, Sydney, New South Wales, Australia

Janice M. Fullerton & Peter R. Schofield

Department of Human Genetics, University of Chicago, Chicago, IL, USA

Elliot S. Gershon

Biometric Psychiatric Genetics Research Unit, Alexandru Obregia Clinical Psychiatric Hospital, Bucharest, Romania

Maria Grigoroiu-Serbanescu

Department of Psychiatric Genetics, Poznan University of Medical Sciences, Poznan, Poland

Joanna Hauser & Joanna M. Pawlak

School of Medicine and Public Health, University of Newcastle, Newcastle, New South Wales, Australia

Frans A. Henskens

Department of Medical Biochemistry and Biophysics, Karolinska Institutet, Stockholm, Sweden

Jens Hjerling-Leffler

HUNT Research Center, Department of Public Health and Nursing, Faculty of Medicine and Health Sciences, Norwegian University of Science and Technology, Trondheim, Norway

Kristian Hveem

Department of Public Health and Institute of Epidemiology and Preventive Medicine, College of Public Health, National Taiwan University, Taipei, Taiwan

Po-Hsiu Kuo

Neuroscience Therapeutic Area, Janssen Research and Development, Titusville, NJ, USA

Qingqin S. Li

JRD Data Science, Janssen Research and Development, Titusville, NJ, USA

Qingqin S. Li

Cancer Epidemiology and Prevention, M. Sklodowska-Curie National Research Institute of Oncology, Warsaw, Poland

Jolanta Lissowska

SA MRC Unit on Risk and Resilience in Mental Disorders, Department of Psychiatry, Stellenbosch University, Stellenbosch, South Africa

Christine Lochner

University of Newcastle, Newcastle, New South Wales, Australia

Carmel Loughland

Department of Psychiatry, Amsterdam University Medical Center, Amsterdam, The Netherlands

Jurjen J. Luykx

Department of Psychiatry and Neuropsychology, School for Mental Health and Neuroscience, Maastricht University Medical Center, Maastricht, The Netherlands

Jurjen J. Luykx

School of Psychology, The University of Queensland, Brisbane, Queensland, Australia

Nicholas G. Martin & Sarah E. Medland

Department of Psychiatry, University of Florida, Gainesville, FL, USA

Carol A. Mathews

Research Institute, Lindner Center of HOPE, Mason, OH, USA

Susan L. McElroy

School of Psychology and Counselling, Queensland University of Technology, Brisbane, Queensland, Australia

Sarah E. Medland

Division of Mental Health and Addiction, University of Oslo, Institute of Clinical Medicine, Oslo, Norway

Ingrid Melle

Department of Mental Health, Faculty of Medicine and Health Sciences, Norwegian University of Science and Technology (NTNU), Trondheim, Norway

Gunnar Morken

Psychiatry, St Olavs University Hospital, Trondheim, Norway

Gunnar Morken

Psychosis Research Unit, Aarhus University Hospital-Psychiatry, Risskov, Denmark

Ole Mors

NCRR and CIRRAU, Aarhus BSS, Aarhus University, Aarhus, Denmark

Preben Bo Mortensen

Munich Cluster for Systems Neurology (SyNergy), Munich, Germany

Bertram Müller-Myhsok

University of Liverpool, Liverpool, UK

Bertram Müller-Myhsok

HudsonAlpha Institute for Biotechnology, Huntsville, AL, USA

Richard M. Myers

Medical and Population Genetics, Broad Institute, Cambridge, MA, USA

Benjamin M. Neale & Eli A. Stahl

Mental Health Services in the Capital Region of Denmark, Mental Health Center Copenhagen, University of Copenhagen, Copenhagen, Denmark

Merete Nordentoft

Psychiatry, Indiana University School of Medicine, Indianapolis, IN, USA

John I. Nurnberger

Division of Psychiatry, Haukeland Universitetssjukehus, Bergen, Norway

Ketil J. Oedegaard

Faculty of Medicine and Dentistry, University of Bergen, Bergen, Norway

Ketil J. Oedegaard

Center for Molecular Medicine, Stockholm, Sweden

Tomas Olsson

Human Genetics and Computational Biomedicine, Pfizer Global Research and Development, Groton, CT, USA

Sara A. Paciga

Melbourne Neuropsychiatry Centre, Department of Psychiatry, The University of Melbourne, Melbourne, Victoria, Australia

Christos Pantelis

Monash Institute of Pharmaceutical Sciences (MIPS), Monash University, Parkville, Victoria, Australia

Christos Pantelis

Rutgers Health, Rutgers University, Piscataway, NJ, USA

Carlos N. Pato & Michele T. Pato

University of Patras, School of Health Sciences, Department of Pharmacy, Laboratory of Pharmacogenomics and Individualized Therapy, Patras, Greece

George P. Patrinos

Department of Genetics and Genomics, College of Medicine and Health Sciences, United Arab Emirates University, Al-Ain, United Arab Emirates

George P. Patrinos

Zayed Center for Health Sciences, United Arab Emirates University, Al-Ain, United Arab Emirates

George P. Patrinos

Department of Pathology, Faculty of Medicine and Health Sciences, Clinical Bioinformatics Unit, Erasmus University Medical Center Rotterdam, Rotterdam, The Netherlands

George P. Patrinos

Department of Neurology and Neurosurgery, Faculty of Medicine, McGill University, Montreal, Québec, Canada

Guy A. Rouleau

Montreal Neurological Institute and Hospital, McGill University, Montréal, Québec, Canada

Guy A. Rouleau

Center for Precision Medicine and Translational Therapeutics, James J. Peters VA Medical Center, Bronx, NY, USA

Panos Roussos

Centre for Brain and Mental Health Research, The University of Newcastle, Newcastle, New South Wales, Australia

Ulrich Schall

Hunter Medical Research Institute, New Lambtion Heights, New South Wales, Australia

Ulrich Schall

Department of Psychiatry and Psychotherapy, University Medical Center Göttingen, Göttingen, Germany

Thomas G. Schulze

Department of Psychiatry and Behavioral Sciences, SUNY Upstate Medical University, Syracuse, NY, USA

Thomas G. Schulze

The School of Biomedical Sciences and Pharmacy, Faculty of Medicine, Health and Wellbeing, University of Newcastle, Newcastle, New South Wales, Australia

Rodney J. Scott

Cancer Detection and Therapies Program, Hunter Medical Research Institute, University of Newcastle, Newcastle, New South Wales, Australia

Rodney J. Scott

Department of Medicine and Surgery, Kore University of Enna, Enna, Italy

Alessandro Serretti

Department of Biomedical and Neuromotor Sciences, University of Bologna, Bologna, Italy

Alessandro Serretti

Oasi Research Institute-IRCCS, Troina, Italy

Alessandro Serretti

Department of Psychiatry, Massachusetts General Hospital, Boston, MA, USA

Jordan W. Smoller

Psychiatric and Neurodevelopmental Genetics Unit (PNGU), Massachusetts General Hospital, Boston, MA, USA

Jordan W. Smoller

Faculty of Medicine, University of Iceland, Reykjavik, Iceland

Kari Stefansson

Department of Psychiatry, Hospital Namsos, Namsos, Norway

Eystein Stordal

Department of Neuroscience, Norges Teknisk Naturvitenskapelige Universitet Fakultet for naturvitenskap og teknologi, Trondheim, Norway

Eystein Stordal

Hector Institute for Artificial Intelligence in Psychiatry, Central Institute of Mental Health, Medical Faculty Mannheim, Heidelberg University, Mannheim, Germany

Fabian Streit

Department of Genetics, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA

Patrick F. Sullivan

Department of Psychiatry, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA

Patrick F. Sullivan & Arianna Di Florio

Department of Psychiatry, McGill University, Montreal, Québec, Canada

Gustavo Turecki

Department of Psychiatry, Sankt Olavs Hospital Universitetssykehuset i Trondheim, Trondheim, Norway

Arne E. Vaaler

Clinical Institute of Neuroscience, Hospital Clinic, University of Barcelona, IDIBAPS, CIBERSAM, Barcelona, Spain

Eduard Vieta

Department of Psychology, Emory University, Atlanta, GA, USA

Irwin D. Waldman

Department of Neuroscience, SUNY Upstate Medical University, Syracuse, NY, USA

Cynthia S. Weickert & Thomas W. Weickert

Institute of Biological Psychiatry, Mental Health Services, Copenhagen University Hospital, Copenhagen, Denmark

Thomas Werge

Department of Clinical Medicine, University of Copenhagen, Copenhagen, Denmark

Thomas Werge

Center for GeoGenetics, GLOBE Institute, University of Copenhagen, Copenhagen, Denmark

Thomas Werge

Biochemistry and Molecular Biology, Indiana University School of Medicine, Indianapolis, IN, USA

Howard J. Edenberg

Department of Medical and Molecular Genetics, Indiana University School of Medicine, Indianapolis, IN, USA

Howard J. Edenberg

Centre for Human Genetics, University of Marburg, Marburg, Germany

Andreas J. Forstner

Genoplan RnD Division, Genoplan Korea, Seoul, Republic of Korea

Byung-Chul Lee, Ji-Woong Kim, Young Kee Lee, Joon Ho Kang, Myeong Jae Cheon & Dong Jun Kim

Genoplan RnD Division, Genoplan Japan, Fukuoka, Japan

Byung-Chul Lee, Ji-Woong Kim, Young Kee Lee, Joon Ho Kang, Myeong Jae Cheon & Dong Jun Kim

Clinical Epidemiology Research Center (CERC), VA Connecticut Healthcare System, West Haven, CT, USA

Mihaela Aslan

Bruce W. Carter Miami Veterans Affairs (VA) Medical Center, Miami, FL, USA

Philip D. Harvey

Office of Research and Development, Veterans Health Administration, Washington, DC, USA

Grant D. Huang

Authors

Kevin S. O’Connell

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