Nature

, volume 604 , issue 7906 , pages 502-508

Mapping genomic loci implicates genes and synaptic biology in schizophrenia

Vassily Trubetskoy ¹

Antonio F. Pardiñas ²

TING QI ^{3, 4}

Georgia Panagiotaropoulou ¹

Swapnil Awasthi ¹

Tim B. Bigdeli ^{5, 6, 7}

Chen Chia-Yen ^{8, 9, 10}

Charlotte A. Dennison ²

Lynsey S. Hall ²

Max Lam ^{11, 12, 13}

Kyoko Watanabe ¹⁴

Oleksandr Frei ^{15, 16, 17}

Tian Ge ^{10, 11, 18}

Janet C Harwood ²

Sigurdur Magnusson ¹⁹

Alexander L. Richards ²

Julia Sidorenko ³

Yang Wu ³

Jian Zeng ³

JAKOB GROVE ^{20, 21, 22}

Minsoo Kim ²³

Zhiqiang Li ^{24, 25}

Georgios Voloudakis ²⁶

Wen Zhang ^{26, 27}

Mark Adams ²⁸

Ingrid Agartz ^{15, 29, 30}

Elizabeth G. Atkinson ^{9, 11}

Esben Agerbo ^{20, 31}

Mariam Al Eissa ³²

Margot Albus ³³

Madeline Alexander ³⁴

Behrooz Z. Alizadeh ^{35, 36}

Köksal Alptekin ^{37, 38}

Thomas D. Als ^{20, 21, 22}

Farooq Amin ³⁹

Volker Arolt ⁴⁰

Manuel Arrojo ⁴¹

Lavinia Athanasiu ^{15, 16}

Maria-Helena Azevedo ⁴²

Silviu A Bacanu ⁴³

Nicholas J. Bass ³²

Martin Begemann ⁴⁴

Richard A. Belliveau ¹¹

Judit Bene ⁴⁵

Beben Benyamin ^{46, 47, 48}

Sarah E. Bergen ⁴⁹

Giuseppe Blasi ⁵⁰

Julio Bobes ^{51, 52, 53}

Stefano Bonassi ⁵⁴

Alice Braun ¹

Rodrigo Affonseca Bressan ^{55, 56}

Evelyn J. Bromet ⁵⁷

Richard Bruggeman ^{35, 58}

Peter F. Buckley ⁵⁹

Randy L. Buckner ⁶⁰

Jonas Bybjerg-Grauholm ^{20, 61}

Wiepke Cahn ^{62, 63}

Murray J Cairns ^{64, 65, 66}

Monica E. Calkins ⁶⁷

Vaughan J Carr ^{68, 69, 70}

David Castle ^{71, 72}

STANLEY V. CATTS ^{73, 74}

Kimberley D Chambert ¹¹

Raymond C. K. Chan ^{75, 76}

Boris Chaumette ^{77, 78}

Cheng Wei ⁷⁹

Eric F.C. Cheung ⁸⁰

Siow Ann Chong ^{12, 81}

David Cohen ^{82, 83, 84}

Angèle Consoli ^{82, 83}

Quirino Cordeiro ⁸⁵

Javier Costas ⁸⁶

Charles Curtis ^{87, 88}

Michael Davidson ⁸⁹

KENNETH L. DAVIS ⁹⁰

Franziska Degenhardt ⁹¹

LYNN E. DELISI ^{18, 92}

Ditte Demontis ^{20, 21, 22}

Faith Dickerson ⁹³

Dimitris Dikeos ⁹⁴

Timothy Dinan ^{95, 96}

Srdjan Djurovic ^{97, 98}

Jubao Duan ^{99, 100}

Giuseppe Ducci ¹⁰¹

Frank Dudbridge ¹⁰²

Johan G. Eriksson ^{103, 104, 105}

Lourdes Fañanas ^{106, 107}

Stephen V. Faraone ¹⁰⁸

Alessia Fiorentino ³²

Andreas Forstner ^{91, 109}

Josef Frank ¹¹⁰

Nelson B. Freimer ^{111, 112}

Alessandra Frustaci ¹¹³

Ary Gadelha ^{55, 56}

Giulio Genovese ¹¹

Elliot S. Gershon ¹¹⁴

Marianna Giannitelli ^{82, 83}

Ina Giegling ¹¹⁵

Paola Giusti-Rodriguez ¹¹⁶

Stephanie Godard ¹¹⁷

Jacqueline I Goldstein ⁹

Javier González Peñas ^{107, 118}

Ana González-Pinto ^{107, 119}

Srihari Gopal ¹²⁰

Jacob Gratten ^{3, 121}

Michael F. Green ^{122, 123}

Tiffany A. Greenwood ¹²⁴

Olivier Guillin ^{125, 126, 127}

Sinan Gülöksüz ^{128, 129}

RAQUEL E. GUR ⁶⁷

RUBEN C. GUR ⁶⁷

Blanca Gutiérrez ¹³⁰

Eric Hahn ¹³¹

Hakon Hakonarson ¹³²

Vahram Haroutunian ^{90, 133, 134}

Annette M. Hartmann ¹¹⁵

Carol Harvey ^{71, 135}

Caroline Hayward ¹³⁶

Frans A Henskens ¹³⁷

Stefan Herms ¹³⁸

Per Hoffmann ¹³⁸

Daniel P. Howrigan ^{9, 139}

Masashi IKEDA ¹⁴⁰

Inge Joa ¹⁴¹

Antonio Juliá ¹⁴²

Anna K. Kähler ⁴⁹

Tony Kam Thong ¹⁴³

Yoichiro Kamatani ^{144, 145}

Sena Karachanak Yankova ^{146, 147}

Oussama Kebir ⁷⁷

Matthew C Keller ¹⁴⁸

Brian J. Kelly ¹³⁷

Andrey Khrunin ¹⁴⁹

Sung Wan Kim ¹⁵⁰

Janis KLOVINS ¹⁵¹

Nikolay Kondratiev ¹⁵²

Bettina Konte ¹¹⁵

Julia Kraft ^{1, 153}

Michiaki Kubo ¹⁵⁴

Vaidutis Kučinskas ¹⁵⁵

Zita Aušrelė Kučinskienė ¹⁵⁵

Agung Kusumawardhani ¹⁵⁶

Hana Kuzelova Ptackova ¹⁵⁷

Stefano Landi ¹⁵⁸

Laura C. Lazzeroni ^{159, 160}

Phil H Lee ^{11, 161}

Sophie E Legge ²

Douglas S. Lehrer ¹⁶²

Rebecca Lencer ⁴⁰

Bernard Lerer ¹⁶³

Miaoxin Li ¹⁶⁴

Jeffrey Lieberman ¹⁶⁵

Gregory A. Light ^{124, 166}

Svetlana Limborska ¹⁴⁹

Chih-Min Liu ^{167, 168}

Jouko Lönnqvist ^{169, 170}

Carmel M Loughland ¹⁷¹

Jan Lubinski ¹⁷²

Jurjen J. Luykx ^{128, 173, 174, 175}

Amy Lynham ²

Milan Macek ¹⁷⁶

Andrew MacKinnon ^{177, 178}

Patrik K.E. Magnusson ⁴⁹

Brion S Maher ¹⁷⁹

Wolfgang Maier ¹⁸⁰

Dolores Malaspina ^{90, 181}

Jacques Mallet ¹⁸²

STEPHEN R. MARDER ¹⁸³

Sara Marsal ¹⁴²

Alicia R. Martin ^{9, 11, 184}

Lourdes Martorell ¹⁸⁵

Manuel Mattheisen ^{21, 186, 187, 188}

ROBERT W. MCCARLEY ¹⁸⁹

Colm McDonald ¹⁹⁰

JOHN J. McGRATH ^{31, 191, 192}

Helena Medeiros ^{193, 194}

Sandra Meier ^{186, 195}

Bela Melegh ¹⁹⁶

Ingrid Melle ^{15, 16}

Raquelle I. Mesholam-Gately ^{18, 197}

Andres Metspalu ¹⁹⁸

Patricia T. Michie ¹⁹⁹

Lili Milani ¹⁹⁸

VIHRA MILANOVA ²⁰⁰

Marina Mitjans ⁴⁴

Espen Molden ^{201, 202}

Esther Molina ²⁰³

María Dolores Moltó ^{107, 204, 205}

Valeria Mondelli ^{88, 206}

Carmen Moreno ^{107, 118}

Christopher P. Morley ²⁰⁷

Gerard Muntané ^{185, 208}

Kieran C. Murphy ²⁰⁹

Inez Myin-Germeys ²¹⁰

Igor Nenadic ^{211, 212}

Gerald Nestadt ²¹³

Liene Ņikitina Zaķe ¹⁵¹

Cristiano Noto ^{55, 56}

Keith H. Nuechterlein ¹²²

Niamh Louise Obrien ³²

F. Anthony O??Neill ²¹⁴

Sang Yun Oh ^{215, 216}

Ann Olincy ²¹⁷

Vanessa Kiyomi Ota ^{56, 218}

Christos Pantelis ^{135, 219, 220, 221}

GEORGE N. PAPADIMITRIOU ⁹⁴

Mara Parellada ^{107, 118}

Tiina Paunio ^{222, 223}

Renata Pellegrino ¹³²

Sathish Periyasamy ^{191, 224}

Diana O. Perkins ²²⁵

Bruno Pfuhlmann ²²⁶

Pietiläinen Olli ^{11, 227, 228}

Jonathan Pimm ³²

David Porteous ²²⁹

Diego Quattrone ^{87, 88, 230}

Digby Quested ^{231, 232}

Allen D. Radant ^{233, 234}

Antonio Rampino ⁵⁰

Mark H. Rapaport ²³⁵

Anna Rautanen ¹⁴³

Abraham Reichenberg ⁹⁰

Cheryl Roe ²³⁶

Joshua L. Roffman ²³⁷

Julian Roth ²³⁸

Matthias Rothermundt ⁴⁰

Bart P. F. Rutten ¹²⁸

Safaa Saker Delye ²³⁹

Veikko Salomaa ²⁴⁰

Julio Sanjuán ^{107, 205, 241}

Marcos Leite Santoro ^{56, 218}

Adam Savitz ¹²⁰

Ulrich Schall ^{65, 242}

Rodney J Scott ^{64, 65, 243}

Larry J. Seidman ^{18, 197}

Sally Isabel Sharp ³²

Jianxin Shi ²⁴⁴

LARRY J. SIEVER ^{90, 245}

Engilbert Sigurdsson ^{246, 247}

Kang Sim ^{248, 249, 250}

Nora Skarabis ¹

Petr Slominsky ¹⁴⁹

Hon Cheong So ^{251, 252}

Janet L. Sobell ¹⁹³

Erik Söderman ³⁰

Helen J. Stain ^{253, 254}

Nils Eiel Steen ^{16, 255}

Agnes A Steixner Kumar ⁴⁴

Elisabeth Stögmann ²⁵⁶

William S. Stone ^{257, 258}

Richard E. Straub ²⁵⁹

Fabian Streit ¹¹⁰

Eric Strengman ²⁶⁰

T Scott Stroup ¹⁶⁵

Mythily Subramaniam ^{12, 81}

Catherine A Sugar ^{122, 261}

Jaana Suvisaari ²⁴⁰

Dragan M Svrakic ²⁶²

Neal R. Swerdlow ¹²⁴

Jin P. Szatkiewicz ¹¹⁶

Thi Minh Tam Ta ^{263, 264}

Atsushi Takahashi ²⁶⁵

Chikashi Terao ²⁶⁵

Florence Thibaut ^{266, 267}

Draga Toncheva ^{146, 268}

Paul A. Tooney ^{64, 65, 66}

Silvia Torretta ⁵⁰

Sarah Tosato ²⁶⁹

Gian Battista Tura ²⁷⁰

Bruce I. Turetsky ⁶⁷

Alp Üçok ²⁷¹

Arne Vaaler ^{272, 273}

Juha Veijola ^{274, 275}

John Waddington ²⁷⁶

Henrik Walter ¹

Anna Waterreus ^{277, 278}

Bradley T. Webb ⁴³

Mark Weiser ²⁷⁹

Nigel M. Williams ²

Stephanie H. Witt ¹¹⁰

Brandon K Wormley ⁴³

Jing Qin Wu ²⁸⁰

Zhida Xu ²⁸¹

Robert Yolken ²⁸²

Clement C. Zai ^{283, 284}

Wei Zhou ²⁵

Feng Zhu ^{285, 286}

Fritz Zimprich ²⁵⁶

Eşref Cem Atbaşoğlu ^{181, 287}

Muhammad Ayub ²⁸⁸

Christian Benner ²²⁸

Alessandro Bertolino ⁵⁰

DONALD W. BLACK ²⁸⁹

Nicholas J. Bray ²

Gerome Breen ⁸⁷

Nancy G. Buccola ²⁹⁰

WILLIAM F. BYERLEY ²⁹¹

Wei J. Chen ^{292, 293}

C Robert Cloninger ²⁶²

Benedicto Crespo-Facorro ^{294, 295}

Gary Donohoe ¹⁹⁰

Robert Freedman ²¹⁷

Cherrie Galletly ^{296, 297, 298}

Michael J Gandal ²³

Massimo Gennarelli ^{299, 300}

David M. Hougaard ^{20, 61}

Hai-Gwo Hwu ^{168, 301}

Assen V. Jablensky ²⁷⁸

Steven A. McCarroll ¹¹

Jennifer L. Moran ^{11, 237}

Ole Mors ^{20, 302}

Preben B. Mortensen ^{20, 31}

Bertram Müller-Myhsok ^{303, 304, 305}

Amanda L. Neil ³⁰⁶

Merete Nordentoft ^{20, 307}

MICHELE T. PATO ^{308, 309}

Tracey L Petryshen ¹⁶¹

Matti Pirinen ^{228, 310, 311}

Ann E. Pulver ²¹³

Thomas G. Schulze ^{188, 312, 313, 314}

Jeremy M. Silverman ^{90, 245}

Jordan W. Smoller ^{11, 161}

Eli A. Stahl ^{315, 316, 317}

Debby W. Tsuang ^{233, 234}

Elisabet Vilella ¹⁸⁵

Shi‐Heng Wang ³¹⁸

Shuhua Xu ^{319, 320, 321}

Nan Dai ³²²

Wenwen Qin ^{107, 118}

D B Wildenauer ^{40, 220, 221}

Feranindhya Agiananda ^{56, 218}

Nurmiati Amir ^{20, 21, 22}

Ronald Antoni ^{124, 166}

Tiana Arsianti ³²³

Asmarahadi Asmarahadi ⁹⁰

H Diatri ^{125, 126}

Prianto Djatmiko ³²⁴

Siti Khalimah ³²⁵

Irmia Kusumadewi ³²⁶

Profitasari Kusumaningrum ^{327, 328}

Petrin R Lukman ^{87, 88, 230}

Martina W Nasrun ³²⁹

N S Safyuni ⁴⁴

Prasetyawan Prasetyawan ^{2, 330}

G Semen ^{198, 316}

Kristiana Siste ^{7, 331, 332}

Heriani Tobing ^{333, 334}

Natalia Widiasih ²³⁸

Tjhin Wiguna ^{99, 100}

D. Wulandari ³²⁶

None Evalina ³³⁵

A J Hananto ¹⁵²

Joni H Ismoyo ³³⁶

T M Marini ⁴⁹

Supiyani Henuhili ^{11, 227}

Muhammad Reza ¹⁴⁰

Suzy Yusnadewi ^{30, 255}

ALEXEJ ABYZOV ^{62, 90}

Schahram Akbarian ^{283, 284}

Allison Ashley-Koch ^{333, 334}

Harm van Bakel ²

Michael Breen ^{337, 338}

Miguel Brown ⁷⁷

Julien Bryois ^{82, 83}

Becky Carlyle ^{339, 340}

Alex Charney ^{13, 341, 342}

Gerard Coetzee ¹⁵⁹

Gregory Crawford ¹²⁰

Stella Dracheva ^{343, 344}

Prashant Emani ^{13, 341, 342}

Peggy Farnham ³⁴⁵

Menachem Fromer ²⁸

Timur Galeev ³²

Mike Gandal ³⁴⁶

Mark Gerstein ^{277, 278}

Gina Giase ¹⁹⁰

Kiran Girdhar ^{191, 224}

Fernando Goes ⁸⁸

Kay Grennan ³⁴⁷

Mengting Gu ⁹¹

Gamze Gürsoy ³⁴⁸

Brittney Guerra ^{111, 349, 350}

Gabriel Hoffman ^{11, 228, 351}

THOMAS HYDE ^{308, 309}

Andrew Jaffe ^{25, 352}

Yan Jiang ⁴³

Shan) Jiang ¹¹⁰

Amira Kefi ^{353, 354}

Yunjung Kim ¹¹⁵

Robert Kitchen ²⁸⁷

Fides Lay ^{355, 356}

Li Mingfeng ^{357, 358, 359}

Donghoon Lee ³⁶⁰

Shuang Liu ³⁶¹

Chunyu Liu ^{24, 25}

Eugenio Mattei ¹⁹

Fabio Navarro ¹⁹

Xinghua Pan ^{362, 363}

Mette A. Peters ^{364, 365}

Dalila Pinto ³⁶⁶

Sirisha Pochareddy ²⁵⁹

Damon Polioudakis ^{367, 368, 369, 370}

Michael Purcaro ³⁷¹

Shaun Purcell ^{372, 373}

Henry Pratt ^{372, 373, 374}

Tim Reddy ²

Suhn Rhie ²

Panagiotis Roussos ^{26, 375}

Joel Rozowsky ^{87, 88, 376}

Stephan Sanders ^{377, 378}

Nenad Sestan ³

Anurag Sethi ^{3, 4, 379}

Xu Shi ³⁷⁸

Annie Shieh ^{15, 16}

Vivek Swarup ⁴³

Anna Szekely ²

Daifeng Wang ^{3, 191}

Jonathan Warrell ^{9, 139, 228}

Sherman Weissman ^{9, 11, 184}

Zhiping Weng ^{9, 11}

Kevin White ^{49, 116, 225}

Jennifer Wiseman ^{1, 9, 11}

Heather Witt ²

Hyejung Won ²

Hide authors affiliations Show authors affiliations: 379 affiliations

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

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

Institute for Molecular Bioscience, University of Queensland, Brisbane, Australia |

⁴

School of Life Sciences, Westlake University, Hangzhou, China |

⁵

Department of Psychiatry and the Behavioral Sciences, SUNY Downstate Medical Center, New York, USA |

⁶

Institute for Genomic Health, SUNY Downstate Medical Center, New York, USA |

⁷

Department of Psychiatry, Veterans Affairs New York Harbor Healthcare System, New York, USA |

⁸

Biogen, Cambridge, USA |

⁹

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

¹⁰

Psychiatric and Neurodevelopmental Genetics Unit, Center for Genomic Medicine, Massachusetts General Hospital, Boston, USA |

¹¹

Stanley Center for Psychiatric Research, Broad Institute of MIT and Harvard, Cambridge, USA |

¹²

Research Division, Institute of Mental Health, Singapore, Republic of Singapore |

¹³

Division of Psychiatry Research, Zucker Hillside Hospital, Glen Oaks, USA |

¹⁴

Department of Complex Trait Genetics, Center for Neurogenomics and Cognitive Research, Amsterdam Neuroscience, Vrije Universiteit Amsterdam, Amsterdam, the Netherlands

Vrije Universiteit Amsterdam

Amsterdam Neuroscience

¹⁵

NORMENT Centre, Division of Mental Health and Addiction, University of Oslo, Oslo, Norway |

¹⁶

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

¹⁷

Center for Bioinformatics, Department of Informatics, University of Oslo, Oslo, Norway |

¹⁸

Department of psychiatry, Harvard Medical School, Boston, USA |

¹⁹

deCODE genetics, Amgen, Reykjavik, Iceland |

²⁰

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

²¹

Department of Biomedicine and Centre for Integrative Sequencing (iSEQ), Aarhus University, Aarhus, Denmark |

²²

Center for Genomics and Personalized Medicine, Aarhus, Denmark |

²³

Department of Psychiatry, Semel Institute, David Geffen School of Medicine, University of California Los Angeles, Los Angeles, USA |

²⁴

Affiliated Hospital of Qingdao University and Biomedical Sciences Institute of Qingdao University (Qingdao Branch of SJTU Bio-X Institutes), Qingdao University, Qingdao, China

Shanghai Jiao Tong University

Qingdao University

²⁵

Bio-X Institutes, Key Laboratory for the Genetics of Developmental and Neuropsychiatric Disorders (Ministry of Education), Collaborative Innovation Center for Brain Science, Shanghai Jiao Tong University, Shanghai, China |

²⁶

Department of Psychiatry, Pamela Sklar Division of Psychiatric Genomics, Friedman Brain Institute, Department of Genetics and Genomic Science and Institute for Data Science and Genomic Technology, Icahn School of Medicine at Mount Sinai, New York, USA |

²⁷

Department of Genetics and Genomic Sciences and Institute for Genomics and Multiscale Biology, Icahn School of Medicine at Mount Sinai, New York, USA |

²⁸

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

²⁹

Department of Psychiatric Research, Diakonhjemmet Hospital, Oslo, Norway |

³⁰

Centre for Psychiatry Research, Department of Clinical Neuroscience, Karolinska Institutet and Stockholm Health Care Services, Stockholm Region, Stockholm, Sweden |

³¹

National Centre for Register-based Research, Aarhus University, Aarhus, Denmark |

³²

Molecular Psychiatry Laboratory, Division of Psychiatry, University College London, London, UK |

³³

Comedicum Lindwurmhof, Munich, Germany |

³⁴

Center for Depression, Anxiety and Stress Research, McLean Hospital, Belmont, USA |

³⁵

University Medical Center Groningen, University Center for Psychiatry, Rob Giel Research center, University of Groningen, Groningen, The Netherlands |

³⁶

Department of Epidemiology, University Medical Center Groningen, University of Groningen, Groningen, the Netherlands |

³⁷

Department of Psychiatry, Dokuz Eylül University School of Medicine, Izmir, Turkey |

³⁸

Department of Neuroscience, Dokuz Eylül University Graduate School of Health Sciences, Izmir, Turkey |

³⁹

Department of Psychiatry and Behavioral Sciences, Emory University, Atlanta, USA |

⁴⁰

Department of Psychiatry, University of Münster, Münster, Germany |

⁴¹

Servizo de Psiquiatría, Complexo Hospitalario Universitario de Santiago de Compostela, Servizo Galego de Saúde (SERGAS), Santiago de Compostela, Spain |

⁴²

Institute of Medical Psychology, Faculty of Medicine, University of Coimbra, Coimbra, Portugal |

⁴³

Virginia Institute for Psychiatric and Behavioral Genetics, Department of Psychiatry, Virginia Commonwealth University, Richmond, USA |

⁴⁴

Clinical Neuroscience, Max Planck Institute of Experimental Medicine, Göttingen, Germany |

⁴⁵

Department of Medical Genetics, Medical School, University of Pécs, Pécs, Hungary |

⁴⁶

Australian Centre for Precision Health, University of South Australia Cancer Research Institute, University of South Australia, Adelaide, Australia |

⁴⁷

UniSA Allied Health and Human Performance, University of South Australia, Adelaide, Australia |

⁴⁸

South Australian Health and Medical Research Institute, Adelaide, Australia |

⁴⁹

Department of medical epidemiology and biostatistics, Karolinska Institutet, Stockholm, Sweden |

⁵⁰

Department of Basic Medical Science, Neuroscience and Sense Organs, University of Bari ‘Aldo Moro’, Bari, Italy |

⁵¹

Área de Psiquiatría-Universidad de Oviedo, Hospital Universitario Central de Asturias (HUCA), Asturias, Spain |

⁵²

Instituto de Investigación Sanitaria Del Principado de Asturias (ISPA), Oviedo, Spain |

⁵³

Centro de Investigación Biomédica en Red de Salud Mental, Oviedo, Spain |

⁵⁴

Unit of Clinical and Molecular Epidemiology, IRCCS San Raffaele Roma and San Raffaele University, Rome, Italy

Istituto di Ricovero e Cura a Carattere Scientifico San Raffaele

San Raffaele University of Rome

⁵⁵

Department of Psychiatry, Universidade federal de São Paulo, São Paulo, Brazil |

⁵⁶

Laboratory of Integrative Neuroscience, Universidade Federal de São Paulo, São Paulo, Brazil |

⁵⁷

Department of Psychiatry and Behavioural Health, Stony Brook University, Stony Brook, USA |

⁵⁸

Department of clinical and Developmental Neuropsychology, University of Groningen, Groningen, the Netherlands |

⁵⁹

Health Science Center, University of Tennessee, Memphis, USA |

⁶⁰

Department of psychology, Harvard University, Cambridge, USA |

⁶¹

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

⁶²

University Medical Center Utrecht, Department of Psychiatry, Rudolf Magnus Institute of Neuroscience, Utrecht, The Netherlands |

⁶³

Altrecht, General Menthal Health Care, Utrecht, The Netherlands |

⁶⁴

School of Biomedical Sciences and Pharmacy, University of Newcastle, Newcastle, Australia |

⁶⁵

Hunter Medical Research Institute, Newcastle, Australia |

⁶⁶

Centre for Brain and Mental Health Research, University of Newcastle, Newcastle, Australia |

⁶⁷

Department of Psychiatry, University of Pennsylvania, Philadelphia, USA |

⁶⁸

School of psychiatry, University of New South Wales, Sydney, Australia |

⁶⁹

Department of Psychiatry, Monash University, Melbourne, Australia |

⁷⁰

Neuroscience Research Australia, Sydney, Australia |

⁷¹

Department of Psychiatry, University of Melbourne, Parkville, Australia |

⁷²

St Vincent’s Hospital, Melbourne, Australia |

⁷³

Brain and Mind Centre, the University of Sydney, Sydney, Australia |

⁷⁴

School of Medicine, University of Queensland, Herston, Australia |

⁷⁵

Institute of Psychology, Chinese Academy of Science, Beijing, China |

⁷⁶

Department of psychology, University of Chinese Academy of Sciences, Beijing, China |

⁷⁷

INSERM U1266, Institute of Psychiatry and Neuroscience of Paris, Université de Paris, GHU Paris Psychiatrie & Neurosciences, Paris, France |

⁷⁸

Department of psychiatry, McGill University, Montreal, Canada |

⁷⁹

Department of Computer Science, University of North Carolina, Chapel Hill, USA |

⁸⁰

Castle Peak Hospital, Hong Kong, China |

⁸¹

Saw Swee Hock School of Public Health, National University of Singapore, Singapore, Republic of Singapore |

⁸²

Faculté de Médecine Sorbonne Université, Groupe de Recherche Clinique no. 15 - Troubles Psychiatriques et Développement (PSYDEV), Department of Child and Adolescent Psychiatry, Hôpital Universitaire de la Pitié-Salpêtrière, Paris, France |

⁸³

Centre de Référence des Maladies Rares à Expression Psychiatrique, Department of Child and Adolescent Psychiatry, AP-HP Sorbonne Université, Hôpital Universitaire de la Pitié-Salpêtrière, Paris, France |

⁸⁴

Institut des Systèmes Intelligents et de Robotique (ISIR), CNRS UMR7222, Faculté des Sciences et Ingénierie, Sorbonne Université, Paris, France

Sorbonne University

Institute for Intelligent Systems and Robotics

⁸⁵

Department of Psychiatry, Irmandade da Santa Casa de Misericórdia de São Paulo, São Paulo, Brazil |

⁸⁶

Instituto de Investigación Sanitaria (IDIS) de Santiago de Compostela, Complexo Hospitalario Universitario de Santiago de Compostela (CHUS), Servizo Galego de Saúde (SERGAS), Santiago de Compostela, Spain |

⁸⁷

Social, Genetic and Developmental Psychiatry Centre, Institute of Psychiatry, Psychology and Neuroscience, King’S College London, London, UK |

⁸⁸

National Institute for Health Research (NIHR) Maudsley Biomedical Research Centre at South London and Maudsley NHS Foundation Trust and King’s College London, London, UK

King's College London

NIHR Maudsley Biomedical Research Centre

⁸⁹

University of Nicosia Medical School, Nicosia, Cyprus |

⁹⁰

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

⁹¹

Institute of Human genetics, University of Bonn, Bonn, Germany |

⁹²

Cambridge Health Alliance, Cambridge, USA |

⁹³

Sheppard Pratt Health System, Baltimore, USA |

⁹⁴

First Department of Psychiatry, Medical School, National and Kapodistrian University of Athens, Eginition Hospital, Athens, Greece |

⁹⁵

Department of Psychiatry and Neurobehavioural Sciences, University College Cork, Cork, Ireland |

⁹⁶

APC microbiome Ireland, University College Cork, Cork, Ireland |

⁹⁷

NORMENT Centre, Department of Clinical Science, University of Bergen, Bergen, Norway |

⁹⁸

Department of medical genetics, Oslo University Hospital, Oslo, Norway |

⁹⁹

Center for Psychiatric Genetics, NorthShore University HealthSystem, Evanston, USA |

¹⁰⁰

Department of Psychiatry and Behavioral Neurosciences, The University of Chicago, Chicago, USA |

¹⁰¹

Department of Mental Health, ASL Rome 1, Rome, Italy |

¹⁰²

Department of Health Sciences, University of Leicester, Leicester, UK |

¹⁰³

Department of General Practice and Primary Health Care, University of Helsinki and Helsinki University Hospital, Helsinki, Finland |

¹⁰⁴

Folkhälsan Research Center, Helsinki, FInland |

¹⁰⁵

Department of Obstetrics and Gynecology, Yong Loo Lin School of Medicine, National University of Singapore, Singapore, Republic of Singapore |

¹⁰⁶

Department of Evolutionary Biology, Ecology and Environmental Sciences, Faculty of Biology, University of Barcelona, Barcelona, Spain |

¹⁰⁷

Centro de Investigación Biomédica En Red en Salud Mental (CIBERSAM), Madrid, Spain |

¹⁰⁸

Departments of Psychiatry and Neuroscience and Physiology, SUNY Upstate Medical University, Syracuse, USA |

¹⁰⁹

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

¹¹⁰

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

Heidelberg University

University of Mannheim

Central Institute of Mental Health

¹¹¹

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

¹¹²

Department of Psychiatry and Biobehavioral Sciences, University of California Los Angeles, Los Angeles, USA |

¹¹³

Barnet, Enfield and Haringey Mental Health NHS Trust, St Ann’s Hospital, London, UK |

¹¹⁴

Departments of Psychiatry and Human Genetics, University of Chicago, Chicago, USA |

¹¹⁵

Department of Psychiatry and Psychotherapy, Medical University of Vienna, Vienna, Austria |

¹¹⁶

Department of Genetics, University of North Carolina, Chapel Hill, USA |

¹¹⁷

Departments of Psychiatry and Human and Molecular Genetics, INSERM, Institut de Myologie, Hôpital de la Pitiè-Salpêtrière, Paris, France |

¹¹⁸

Department of Child and Adolescent Psychiatry, Institute of Psychiatry and Mental Health, General Universitario Gregorio Marañón, School of Medicine, Universidad Complutense, Madrid, Spain

Universidad Complutense de Madrid

Hospital General Universitario Gregorio Marañón

¹¹⁹

BIOARABA Health Research Institute, OSI Araba, University Hospital, University of the Basque Country, Vitoria, Spain

University of the Basque Country

Araba University Hospital

¹²⁰

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

¹²¹

Mater Research Institute, University of Queensland, Brisbane, Australia |

¹²²

Department of Psychiatry and Biobehavioral Sciences, Geffen School of Medicine, University of California Los Angeles, Los Angeles, USA |

¹²³

VA Greater Los Angeles Healthcare System, Los Angeles, USA |

¹²⁴

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

¹²⁵

Inserm, Rouen, France |

¹²⁶

Centre Hospitalier du Rouvray, Rouen, France |

¹²⁷

UFR Santé, Université de Rouen Normandie, Rouen, France |

¹²⁸

Department of Psychiatry and Neuropsychology, School for Mental Health and Neuroscience, Maastricht University Medical Centre, Maastricht, the Netherlands |

¹²⁹

Department of Psychiatry, Yale School of Medicine, New Haven, USA |

¹³⁰

Department of Psychiatry, Faculty of Medicine and Biomedical Research Centre (CIBM), University of Granada, Granada, Spain |

¹³¹

Department of Psychiatry, Charité - Universitätsmedizin, Berlin, Germany |

¹³²

Children’s Hospital of Philadelphia, Leonard Madlyn Abramson Research Center, Philadelphia, USA |

¹³³

Department of Neuroscience, ICAHN School of Medicine at Mount Sinai, New York, USA |

¹³⁴

Mental Illness Research Clinical and Education Center (MIRECC), JJ Peters VA Medical Center, New York, USA |

¹³⁵

Northwestern Mental Health, Melbourne, Australia |

¹³⁶

MRC Human Genetics Unit, University of Edinburgh, Institute of Genetics and Cancer, Western General Hospital, Edinburgh, UK |

¹³⁷

School of Medicine and Public Health, University of Newcastle, Newcastle, AUSTRALIA |

¹³⁸

Division of Medical Genetics, Department of Biomedicine, University of Basel, Basel, Switzerland |

¹³⁹

Broad Institute of MIT and Harvard, Cambridge, USA |

¹⁴⁰

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

¹⁴¹

Regional Centre for Clinical Research in Psychosis, Department of Psychiatry, Stavanger University Hospital, Stavanger, Norway |

¹⁴²

Rheumatology Research Group, Vall d’Hebron Research Institute, Barcelona, Spain |

¹⁴³

Roche Pharma Research and Early Development, Pharmaceutical Sciences, Roche Innovation Center Basel, F. Hoffman-La Roche, Basel, Switzerland |

¹⁴⁴

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

¹⁴⁵

Laboratory for Statistical Analysis, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan |

¹⁴⁶

Department of Medical Genetics, Medical University, Sofia, Bulgaria |

¹⁴⁷

Department of Genetics, Faculty of Biology, Sofia University “St. Kliment Ohridski”, Sofia, Bulgaria |

¹⁴⁸

Institute for Behavioural Genetics, University of Colorado Boulder, Boulder, USA |

¹⁴⁹

Institute of Molecular Genetics of National Research Centre “Kurchatov Institute”, Moscow, Russia

Institute of Molecular Genetics of NRC «Kurchatov Institute»

National Research Centre "Kurchatov Institute"

¹⁵⁰

Department of Psychiatry, Chonnam National University Medical School, Gwangju, Korea |

¹⁵¹

Latvian Biomedical Research and Study centre, Riga, Latvia |

¹⁵²

Mental Health Research Center, Moscow, Russian Federation |

¹⁵³

Berlin School of Mind and Brain, Humboldt-Universität Zu Berlin, Berlin, Germany |

¹⁵⁴

RIKEN Center for Integrative Medical Sciences, Yokohama, Japan |

¹⁵⁵

Faculty of Medicine, Vilnius University, Vilnius, Lithuania |

¹⁵⁶

Psychiatry Department, University of Indonesia - Cipto Mangunkusumo National General Hospital, Jakarta, Indonesia |

¹⁵⁷

Department of Psychiatry, 1st Faculty of Medicine and General University Hospital, Prague, Czech Republic |

¹⁵⁸

Dipartimento di Biologia, Universita’ di Pisa, Pisa, Italy |

¹⁵⁹

Department of Psychiatry and Behavioral Sciences, Stanford University, Stanford, USA |

¹⁶⁰

Department of Biomedical Data Science, Stanford University, Stanford, USA |

¹⁶¹

Psychiatric and Neurodevelopmental Genetics Unit, Department of Psychiatry and Center for Genomic Medicine, Massachusetts General Hospital, Harvard Medical School, Boston, USA

Harvard University

Massachusetts General Hospital

¹⁶²

Department of Psychiatry, Wright State University, Dayton, USA

Wright State University

University of Dayton

¹⁶³

Department of Psychiatry, Hadassah-Hebrew University Medical Center, Jerusalem, Israel |

¹⁶⁴

Zhongshan School of Medicine and Key Laboratory of Tropical Diseases Control (SYSU), Sun Yat-sen University, Guangzhou, China |

¹⁶⁵

Department of Psychiatry, Columbia University, New York, USA |

¹⁶⁶

VISN 22, Mental Illness Research, Education and Clinical Center (MIRECC), VA San Diego Healthcare System, San Diego, USA |

¹⁶⁷

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

¹⁶⁸

Neurobiology and Cognitive Science Center, National Taiwan University, Taipei, Taiwan |

¹⁶⁹

Mental Health Unit, Department of Public Health Solutions, National Institute for Health and Welfare, Helsinki, Finland |

¹⁷⁰

Department of Psychiatry, University of Helsinki, Helsinki, Finland |

¹⁷¹

Hunter New England Health and University of Newcastle, Newcastle, Australia

University of Newcastle Australia

Hunter New England Local Health District

¹⁷²

Department of Genetics and Pathology, International Hereditary Cancer Center, Pomeranian Medical University in Szczecin, Szczecin, Poland |

¹⁷³

Department of Psychiatry, UMC Utrecht Brain Center, University Medical Centre Utrecht, Utrecht University, Utrecht, The Netherlands |

¹⁷⁴

Department of Translational Neuroscience, UMC Utrecht Brain Center, University Medical Center Utrecht, Utrecht University, Utrecht, the Netherlands |

¹⁷⁵

Second Opinion Outpatient Clinic, GGNet Mental Health, Warnsveld, the Netherlands |

¹⁷⁶

Department of Biology and Medical Genetics, 2nd Faculty of Medicine and University Hospital Motol, Prague, Czech Republic |

¹⁷⁷

Black Dog Institute, University of New South Wales, Randwick, Australia

University of New South Wales

Black Dog Institute

¹⁷⁸

Melbourne School of Population and Global Health, University of Melbourne, Melbourne, Australia |

¹⁷⁹

Department of Mental Health, Bloomberg School of Public Health, Johns Hopkins University, Baltimore, USA |

¹⁸⁰

Department for Neurodegenerative Diseases and Geriatric Psychiatry, University Hospital Bonn, Bonn, Germany |

¹⁸¹

Department of Genetics and Genomics, Icahn School of Medicine at Mount Sinai, New York, USA |

¹⁸²

Asfalia Biologics, iPEPS-ICM, Hôpital Universitaire de la Pitié-Salpêtrière, Paris, France |

¹⁸³

Semel Institute for Neurosciene, University of California Los Angeles, Los Angeles, USA |

¹⁸⁴

Department of medicine, Harvard Medical School, Boston, USA |

¹⁸⁵

Hospital Universitari Institut Pere Mata, IISPV, Universitat Rovira i Virgili, CIBERSAM, Reus, Spain

University of Rovira i Virgili

Centro de Investigación Biomédica en Red de Salud Mental

¹⁸⁶

Department of Psychiatry, Dalhousie University, Halifax, Canada |

¹⁸⁷

Department of Community Health and Epidemiology, Dalhousie University, Halifax, Canada |

¹⁸⁸

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

¹⁸⁹

VA Boston Health Care System, Brockton, USA |

¹⁹⁰

Centre for Neuroimaging, Cognition and Genomics (NICOG), National University of Ireland Galway, Galway, Ireland |

¹⁹¹

Queensland Brain Institute, University of Queensland, Brisbane, Australia |

¹⁹²

Queensland Centre for Mental Health Research, The Park Centre for Mental Health, Brisbane, Australia |

¹⁹³

Department of Psychiatry and the Behavioral Sciences, Keck School of Medicine, University of Southern California, Los Angeles, USA |

¹⁹⁴

College of Medicine, SUNY Downstate Health Sciences University, New York, USA |

¹⁹⁵

Department of Biomedicine, Aarhus University, Aarhus, Denmark |

¹⁹⁶

Department of Medical Genetics, University of Pécs, School of Medicine, Pécs, Hungary |

¹⁹⁷

Massachusetts Mental Health Center Public Psychiatry Division of the Beth Israel Deaconess Medical Center, Boston, USA |

¹⁹⁸

Estonian Genome Center, Institute of Genomics, University of Tartu, Tartu, Estonia |

¹⁹⁹

School of Psychology, University of Newcastle, Newcastle, Australia |

²⁰⁰

Psychiatric Clinic, Alexandrovska University Hospital, Sofia, Bulgaria |

²⁰¹

Department of Pharmacy, University of Oslo, Oslo, Norway |

²⁰²

Center for Psychopharmacology, Diakonhjemmet Hospital, Oslo, Norway |

²⁰³

Department of Nursing, Faculty of Health Sciences and Biomedical Research Centre (CIBM), University of Granada, Granada, Spain |

²⁰⁴

Department of Genetics, Faculty of Biological Sciences, Universidad de Valencia, Valencia, Spain |

²⁰⁵

Biomedical Research Institute INCLIVA, Valencia, Spain |

²⁰⁶

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

²⁰⁷

Departments of Public Health and Preventive Medicine, Family Medicine, and Psychiatry and Behavioral Sciences, State University of New York, Upstate Medical University, Syracuse, USA |

²⁰⁸

Institut de Biologia Evolutiva (UPF-CSIC), Departament de Ciències Experimentals i de la Salut, Universitat Pompeu Fabra, PRBB, Barcelona, Spain

Barcelona Biomedical Research Park

Institut de Biologia Evolutiva

²⁰⁹

Department of Psychiatry, Royal College of Surgeons in Ireland, Dublin, Ireland |

²¹⁰

Department for Neurosciences, Center for Contextual Psychiatry, KU Leuven, Leuven, Belgium |

²¹¹

Cognitive Neuropsychiatry Laboratory, Department of Psychiatry and Psychotherapy, Philipps Universität Marburg, Marburg, Germany |

²¹²

Department of Psychiatry and Psychotherapy, Jena University Hospital, Jena, Germany |

²¹³

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

²¹⁴

Centre for Public Health, Institute of Clinical Sciences, Queen’s University Belfast, Belfast, UK |

²¹⁵

Department of Statistics and Applied Probability, University of California at Santa Barbara, Santa Barbara, USA |

²¹⁶

Computational Research Division, Lawrence Berkeley National Laboratory, Berkeley, USA |

²¹⁷

Department of Psychiatry, University of Colorado Denver, Aurora, USA |

²¹⁸

Department of Morphology and Genetics, Laboratorio de Genetica, Universidade Federal de São Paulo, São Paulo, Brazil |

²¹⁹

Melbourne Neuropsychiatry Centre, University of Melbourne and Melbourne Health, Melbourne, Australia |

²²⁰

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

University of Melbourne

Florey Institute of Neuroscience and Mental Health

²²¹

Department of Psychiatry, Melbourne Medical School, University of Melbourne, Parkville, Australia |

²²²

Department of Public Health Solutions, Genomics and Biomarkers Unit, National Institute for Health and Welfare, Helsinki, Finland |

²²³

Department of Psychiatry and SleepWell Research Program, Faculty of Medicine, University of Helsinki and Helsinki University Central Hospital, Helsinki, Finland |

²²⁴

Queensland Centre for Mental Health Research, University of Queensland, Brisbane, Australia |

²²⁵

Department of Psychiatry, University of North Carolina, Chapel Hill, USA |

²²⁶

Clinic of Psychiatry and Psychotherapy, Weißer Hirsch, Dresden, Germany |

²²⁷

Department of Stem cell and Regenerative Biology, Harvard University, Cambridge, USA |

²²⁸

Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland

University of Helsinki

Institute for Molecular Medicine Finland

²²⁹

Centre for Genomic and Experimental Medicine, Institute of Genetics and Molecular Medicine, University of Edinburgh, Western General Hospital, Edinburgh, UK |

²³⁰

South London and Maudsley NHS Mental Health Foundation Trust, London, UK |

²³¹

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

²³²

Department of Psychiatry, University of Oxford, Oxford, UK |

²³³

Department of Psychiatry and Behavioral Sciences, University of Washington, Seattle, USA |

²³⁴

VA Puget Sound Health Care System, Seattle, USA |

²³⁵

Huntsman Mental Health Institute, Department of Psychiatry, University of Utah School of Medicine, Salt Lake City, USA |

²³⁶

SUNY Upstate Medical University, Syracuse, USA |

²³⁷

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

²³⁸

Department of Psychiatry, Psychosomatics and Psychotherapy, Julius-Maximilians-Universität Würzburg, Würzburg, Germany |

²³⁹

Généthon, Evry, France |

²⁴⁰

THL–Finnish Institute for Health and Welfare, Helsinki, Finland |

²⁴¹

Department of Psychiatry, School of Medicine, University of Valencia, Hospital Clínico Universitario de Valencia, Valencia, Spain |

²⁴²

Priority Centre for Brain and Mental Health Research, University of Newcastle, Mater Hospital, McAuley Centre, Newcastle, Australia |

²⁴³

Division of Molecular Medicine, NSW Health Pathology North, Newcastle, Australia |

²⁴⁴

Division of Cancer Epidemiology and Genetics, National Cancer Institute, Bethesda, USA |

²⁴⁵

James J. Peters VA Medical Center, Bronx, USA |

²⁴⁶

Faculty of medicine, University of Iceland, Reykjavik, Iceland |

²⁴⁷

Department of Psychiatry, Landspitali University Hospital, Reykjavik, Iceland |

²⁴⁸

West Region, Institute of Mental Health, Singapore, Singapore |

²⁴⁹

Yoo Loo Lin School of Medicine, National University of Singapore, Singapore, Singapore |

²⁵⁰

Lee Kong Chian School of Medicine, Nanyang Technological University, Singapore, Singapore |

²⁵¹

School of Biomedical Sciences, The Chinese University of Hong Kong, Hong Kong, China |

²⁵²

Department of Psychiatry, The Chinese University of Hong Kong, Hong Kong, China |

²⁵³

School of social and health sciences, Leeds trinity University, Leeds, UK |

²⁵⁴

TIPS - Network for Clinical Research in Psychosis, Stavanger University Hospital, Stavanger, Norway |

²⁵⁵

NORMENT Centre, Institute of Clinical Medicine, University of Oslo, Oslo, Norway |

²⁵⁶

Department of Neurology, Medical University of Vienna, Vienna, Austria |

²⁵⁷

Harvard Medical School Department of Psychiatry at Beth Israel Deaconess Medical Center, Boston, USA

Harvard University

Beth Israel Deaconess Medical Center

²⁵⁸

Massachusetts Mental Health Center, Boston, USA |

²⁵⁹

Lieber Institute for Brain Development, Baltimore, USA |

²⁶⁰

Department of Medical Genetics, University Medical Centre Utrecht, Utrecht, the Netherlands |

²⁶¹

Department of Biostatistics, Fielding School of Public Health, University of California Los Angeles, Los Angeles, USA |

²⁶²

Department of Psychiatry, Washington University, St Louis, USA |

²⁶³

Department of Psychiatry, Charité – Universitätsmedizin Berlin, Berlin, Germany |

²⁶⁴

Berlin Institute of Health (BiH), Berlin, Germany |

²⁶⁵

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

²⁶⁶

Université de Paris, Faculté de Médecine, Hôpital Cochin-Tarnier, Paris, France |

²⁶⁷

INSERM U1266, Institut de Psychiatrie et de Neurosciences, Paris, France |

²⁶⁸

Bulgarian Academy of Science, Sofia, Bulgaria |

²⁶⁹

Department of Neuroscience, Biomedicine and Movement Sciences, Section of Psychiatry, University of Verona, Verona, Italy |

²⁷⁰

Psychiatry Unit, IRCCS Istituto Centro San Giovanni di Dio Fatebenefratelli, Brescia, Italy |

²⁷¹

Department of Psychiatry, Faculty of Medicine, Istanbul University, Istanbul, Turkey |

²⁷²

Division of Mental Health, St. Olav’s Hospital, Trondheim University Hospital, Trondheim, Norway |

²⁷³

Department of Mental Health, Norwegian University of Science and Technology, Trondheim, Norway |

²⁷⁴

Department of Psychiatry, Research Unit of Clinical Neuroscience, University of Oulu, Oulu, Finland |

²⁷⁵

Medical Research Center Oulu, Oulu University Hospital and University of Oulu, Oulu, Finland |

²⁷⁶

Molecular and Cellular Therapeutics, Royal College of Surgeons in Ireland, Dublin, Ireland |

²⁷⁷

Neuropsychiatric Epidemiology Research Unit, School Of Population And Global Health, University Of Western Australia, Perth, Australia |

²⁷⁸

Centre for Clinical Research in Neuropsychiatry, University of Western Australia, Perth, Australia |

²⁷⁹

Sheba Medical Center, Tel HaShomer, Israel |

²⁸⁰

Baker Heart and Diabetes Institute, Melbourne, Australia |

²⁸¹

Department of Psychiatry, GGz Centraal, Utrecht, The Netherlands |

²⁸²

Stanley Neurovirology Laboratory, Johns Hopkins University School of Medicine, Baltimore, USA |

²⁸³

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

²⁸⁴

Department of Psychiatry, University Of Toronto, Toronto, Canada |

²⁸⁵

Department of Psychiatry, The First Affiliated Hospital of Xi’an Jiaotong University, Xi’an, China |

²⁸⁶

Center for Translational Medicine, the First Affiliated Hospital of Xi’an Jiaotong University, Xi’an, China |

²⁸⁷

Department of Psychiatry, School of Medicine, Ankara University, Ankara, Turkey |

²⁸⁸

Department of Psychiatry, Queens University Kingston, Kingston, Canada |

²⁸⁹

Department of Psychiatry, University of Iowa Carver College of Medicine, Iowa City, USA |

²⁹⁰

School of Nursing, Louisiana State University Health Sciences Center, New Orleans, USA |

²⁹¹

Department of Psychiatry, University of California San Francisco, San Francisco, USA |

²⁹²

Center for Neuropsychiatric Research, National Health Research Institutes, Zhunan Town, Taiwan |

²⁹³

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

²⁹⁴

University of Sevilla, CIBERSAM IBiS, Seville, Spain

Universidad de Sevilla

Centro de Investigación Biomédica en Red de Salud Mental

²⁹⁵

Hospital Universitario Virgen del Rocio, Department of Psychiatry, Universidad del Sevilla, Seville, Spain

Universidad de Sevilla

Virgen del Rocío University Hospital

²⁹⁶

Discipline of Psychiatry, Adelaide Medical School, University of Adelaide, Adelaide, Australia |

²⁹⁷

Ramsay Health Care (SA) Mental Health, Adelaide, Australia |

²⁹⁸

Northern Adelaide Local Health Network, Adelaide, Australia |

²⁹⁹

Department of Molecular and Translational medicine, University of Brescia, Brescia, Italy |

³⁰⁰

Genetic Unit, IRCCS Istituto Centro San Giovanni di Dio Fatebenefratelli, Brescia, Italy |

³⁰¹

Department of Psychiatry, College of Medicine and National Taiwan University Hospital, National Taiwan University, Taipei, Taiwan |

³⁰²

Psychosis Research Unit, Aarhus University Hospital, Aarhus, Denmark |

³⁰³

Max Planck Institute of Psychiatry, Munich, Germany |

³⁰⁴

Munich Cluster for Systems Neurology, Munich, Germany |

³⁰⁵

Department of Health Data Science, University of Liverpool, Liverpool, UK |

³⁰⁶

Menzies institute for Medical Research, University of Tasmania, Hobart, Australia |

³⁰⁷

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

³⁰⁸

Rutgers University, Robert Wood Johnson Medical School, New Brunswick, USA |

³⁰⁹

Rutgers University, New Jersey Medical School, Newark, USA |

³¹⁰

Department of Mathematics and statistics, University of Helsinki, Helsinki, Finland |

³¹¹

Department of Public health, University of Helsinki, Helsinki, Finland |

³¹²

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

³¹³

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

³¹⁴

Department of Psychiatry and Behavioral Sciences, The Johns Hopkins University, Baltimore, USA |

³¹⁵

Division of Psychiatric Genomics, Department of Psychiatry, Icahn School of Medicine at Mount Sinai, New York, USA |

³¹⁶

Program in Medical and Population Genetics, The Broad Institute of MIT and Harvard, Cambridge, USA |

³¹⁷

Regeneron Genetics Center, Orange, USA |

³¹⁸

College of Public Health, China Medical University, Taichung, Taiwan |

³¹⁹

State Key Laboratory of Genetic Engineering and Ministry of Education (MOE) Key Laboratory of Contemporary Anthropology, Collaborative Innovation Center of Genetics and Development, Human Phenome Institute, School of Life Sciences, Fudan University, Shanghai, China |

³²⁰

School of Life Science and Technology, ShanghaiTech University, Shanghai, China |

³²¹

Center for Excellence in Animal Evolution and Genetics, Chinese Academy of Sciences, Kunming, China |

³²²

Department of Clinical Sciences, Psychiatry, Umeå university, Umeå, Sweden |

³²³

Division of Psychiatry, Department of Mental Health Neuroscience, University College London, London, UK |

³²⁴

Department of Psychiatry, San Cecilio University Hospital, University of Granada, Granada, Spain |

³²⁵

Eli Lilly and Company, Windlesham, UK |

³²⁶

Neuropsychiatric Genetics Research Group, Department of Psychiatry, Trinity College Dublin, Dublin, Ireland |

³²⁷

UCL Genetics Institute, University College London, London, UK |

³²⁸

Centre for Psychiatry, Queen Mary University London, London, UK |

³²⁹

Department of Cellular, Computational and Integrative Biology, University of Trento, Trento, Italy |

³³⁰

Dementia Research Institute, Cardiff University, Cardiff, UK |

³³¹

Department of Psychiatry, Phoenix VA Healthcare System, Phoenix, USA |

³³²

Banner-University Medical Center, Phoenix, USA |

³³³

Department of Human Molecular Genetics of the Institute of Biochemistry and Genetics of the Ufa Federal Research Center of the Russian Academy of Sciences (IBG UFRC RAS), Ufa, Russia

Institute of Biochemistry and Genetics of the Ufa Federal Research Center of the Russian Academy of Sciences

Ufa Federal Research Center of the Russian Academy of Sciences

³³⁴

Federal State Educational Institution of Highest Education Bashkir State Medical University of Public Health Ministry of Russian Federation (BSMU), Ufa, Russia |

³³⁵

Psychiatric Genetic Epidemiology and Neurobiology Laboratory (PsychGENe lab), Department of Psychiatry and Behavioral Sciences, SUNY Upstate Medical University, Syracuse, USA |

³³⁶

Department of Psychiatry, Sungkyunkwan University School of Medicine, Samsung Medical Center, Seoul, Korea

Sungkyunkwan University

Samsung Medical Center

³³⁷

Department of Psychiatry and Zilkha Neurogenetics Institute, Keck School of Medicine at University of Southern California, Los Angeles, USA |

³³⁸

Department of Cell Biology, State University of New York, Downstate Health Sciences University, New York, USA |

³³⁹

Department of Psychosis, Institute of Mental Health, Singapore, Singapore |

³⁴⁰

Neuroscience and Mental Health, Lee Kong Chian School of Medicine, Nanyang Technological University, Singapore, Singapore |

³⁴¹

Institute of Behavioral Science, Feinstein Institutes for Medical Research, Manhasset, USA |

³⁴²

Department of Psychiatry, Zucker School of Medicine at Hofstra/Northwell, Hempstead, USA |

³⁴³

Human Genetics, Genome Institute of Singapore, A*STAR, Singapore, Singapore

Agency for Science, Technology and Research

Genome Institute of Singapore

³⁴⁴

Department of Medicine, Yong Loo Lin School of Medicine, National University of Singapore, Singapore, SINGAPORE |

³⁴⁵

Roche Pharma Research and Early Development, Roche Innovation Center Basel, F. Hoffman-La Roche, Basel, Switzerland |

³⁴⁶

Department of Preventative Medicine, Faculdade de Medicina FMUSP, University of São Paulo, São Paulo, Brazil |

³⁴⁷

Department of Psychiatry, University of Pittsburgh, Pittsburgh, USA |

³⁴⁸

Early Clinical Development, Pfizer Worldwide Research and Development, Groton, USA |

³⁴⁹

Center for Neurobehavioral Genetics, Semel Institute for Neuroscience and Human Behavior, University of California Los Angeles, Los Angeles, USA |

³⁵⁰

Department of Psychiatry, Erasmus University Medical Center, Rotterdam, the Netherlands |

³⁵¹

Analytic and Translational Genetics Unit, Department of Medicine, Department of Neurology and Department of Psychiatry, Massachusetts General Hospital, Boston, USA |

³⁵²

Shanghai Key Laboratory of Psychotic disorders, Shanghai Mental Health Center, Shanghai Jiao Tong University School of Medicine, Shanghai, China |

³⁵³

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

³⁵⁴

Institute of Neurosciences, Biomedical Research Center (CIBM), University of Granada, Granada, Spain |

³⁵⁵

Faculty of Science, Medicine and Health, University of Wollongong, Wollongong, Australia |

³⁵⁶

Illawarra Health and Medical Research Institute, Wollongong, Australia |

³⁵⁷

Centre for PanorOmic Sciences, LKS Faculty of Medicine, The University of Hong Kong, Hong Kong, China |

³⁵⁸

State Key Laboratory of Brain and Cognitive Sciences, LKS Faculty of Medicine, The University of Hong Kong, Hong Kong, China |

³⁵⁹

Department of Psychiatry, LKS Faculty of Medicine, The University of Hong Kong, Hong Kong, China |

³⁶⁰

Department of Biomedical and Neuromotor Sciences, University of Bologna, Bologna, ITALY |

³⁶¹

Institute of Medical Sciences, University of Aberdeen, Aberdeen, UK |

³⁶²

Center for Behavioral Genomics, Department of Psychiatry, University of California San Diego, La Jolla, USA |

³⁶³

Institute of Genomic Medicine, University of California San Diego, La Jolla, USA |

³⁶⁴

Department of Psychosis studies, Institute of Psychiatry, Psychology and Neuroscience, King’s College London, London, UK |

³⁶⁵

University Medical Center Utrecht, Department of Psychiatry, Utrecht, The Netherlands |

³⁶⁶

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

³⁶⁷

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

³⁶⁸

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

³⁶⁹

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

³⁷⁰

The Lundbeck Foundation Initiative for Integrative Psychiatric Research, iPSYCH, Copenhagen, Denmark |

³⁷¹

School of Psychiatry and Clinical Neurosciences, University of Western Australia, Perth, Australia |

³⁷²

Peking University Sixth Hospital, Peking University Institute of Mental Health, Beijing, China |

³⁷³

National Clinical Research Center for Mental Disorders, NHC Key Laboratory of Mental Health (Peking University) and Chinese Academy of Medical Sciences Research Unit, Beijing, China

University of Chinese Academy of Sciences

Peking University

Chinese Academy of Medical Sciences & Peking Union Medical College

³⁷⁴

PKU–IDG/McGovern Institute for Brain Research, Peking University, Beijing, China |

³⁷⁵

Mental Illness Research, Education, and Clinical Center (VISN 2 South), James J. Peters VA Medical Center, New York, USA |

³⁷⁶

Oxford Health NHS Foundation Trust, Oxford, UK |

³⁷⁷

Department of Clinical Genetics, Center for Neurogenomics and Cognitive Research, University Medical Center Amsterdam, Amsterdam, The Netherlands |

³⁷⁸

Department of Functional Genomics, Faculty of Exact Science, Center for Neurogenomics and Cognitive Research, VU University Amsterdam and VU Medical Center, Amsterdam, The Netherlands

Vrije Universiteit Medical Center

University of Amsterdam

³⁷⁹

Westlake Laboratory of Life Sciences and Biomedicine, Hangzhou, China |

Publication type: Journal Article

Publication date: 2022-04-08

Springer Nature

Nature

scimago Q1

wos Q1

SJR: 18.288

CiteScore: 78.1

Impact factor: 48.5

ISSN: 00280836, 14764687

DOI: 10.1038/s41586-022-04434-5

Copy DOI

PubMed ID: 35396580

Multidisciplinary

Abstract

Schizophrenia has a heritability of 60–80%1, much of which is attributable to common risk alleles. Here, in a two-stage genome-wide association study of up to 76,755 individuals with schizophrenia and 243,649 control individuals, we report common variant associations at 287 distinct genomic loci. Associations were concentrated in genes that are expressed in excitatory and inhibitory neurons of the central nervous system, but not in other tissues or cell types. Using fine-mapping and functional genomic data, we identify 120 genes (106 protein-coding) that are likely to underpin associations at some of these loci, including 16 genes with credible causal non-synonymous or untranslated region variation. We also implicate fundamental processes related to neuronal function, including synaptic organization, differentiation and transmission. Fine-mapped candidates were enriched for genes associated with rare disruptive coding variants in people with schizophrenia, including the glutamate receptor subunit GRIN2A and transcription factor SP4, and were also enriched for genes implicated by such variants in neurodevelopmental disorders. We identify biological processes relevant to schizophrenia pathophysiology; show convergence of common and rare variant associations in schizophrenia and neurodevelopmental disorders; and provide a resource of prioritized genes and variants to advance mechanistic studies. A genome-wide association study including over 76,000 individuals with schizophrenia and over 243,000 control individuals identifies common variant associations at 287 genomic loci, and further fine-mapping analyses highlight the importance of genes involved in synaptic processes.

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Trubetskoy V. et al. Mapping genomic loci implicates genes and synaptic biology in schizophrenia // Nature. 2022. Vol. 604. No. 7906. pp. 502-508.

GOST all authors (up to 50) Copy

Trubetskoy V. et al. Mapping genomic loci implicates genes and synaptic biology in schizophrenia // Nature. 2022. Vol. 604. No. 7906. pp. 502-508.

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RIS

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BibTex (up to 50 authors) Copy

@article{2022_Trubetskoy,

author = {Vassily Trubetskoy and Antonio F. Pardiñas and TING QI and Georgia Panagiotaropoulou and Swapnil Awasthi and Tim B. Bigdeli and Chen Chia-Yen and Charlotte A. Dennison and Lynsey S. Hall and Max Lam and Kyoko Watanabe and Oleksandr Frei and Tian Ge and Janet C Harwood and Sigurdur Magnusson and Alexander L. Richards and Julia Sidorenko and Yang Wu and Jian Zeng and JAKOB GROVE and Minsoo Kim and Zhiqiang Li and Georgios Voloudakis and Wen Zhang and Mark Adams and Ingrid Agartz and Elizabeth G. Atkinson and Esben Agerbo and Mariam Al Eissa and Margot Albus and Madeline Alexander and Behrooz Z. Alizadeh and Köksal Alptekin and Thomas D. Als and Farooq Amin and Volker Arolt and Manuel Arrojo and Lavinia Athanasiu and Maria-Helena Azevedo and Silviu A Bacanu and Nicholas J. Bass and Martin Begemann and Richard A. Belliveau and Judit Bene and Beben Benyamin and Sarah E. Bergen and Giuseppe Blasi and Julio Bobes and Stefano Bonassi and Alice Braun and others},

title = {Mapping genomic loci implicates genes and synaptic biology in schizophrenia},

journal = {Nature},

year = {2022},

volume = {604},

publisher = {Springer Nature},

month = {apr},

url = {https://www.nature.com/articles/s41586-022-04434-5},

number = {7906},

pages = {502--508},

doi = {10.1038/s41586-022-04434-5}

}

MLA

Cite this

MLA Copy

Trubetskoy, Vassily, et al. “Mapping genomic loci implicates genes and synaptic biology in schizophrenia.” Nature, vol. 604, no. 7906, Apr. 2022, pp. 502-508. https://www.nature.com/articles/s41586-022-04434-5.

Publisher

Springer Nature

Journal

Nature

scimago Q1

wos Q1

SJR

18.288

CiteScore

78.1

Impact factor

48.5

ISSN

00280836 (Print)

14764687 (Electronic)

Profiles