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Gene expression imputation across multiple brain regions provides insights into schizophrenia risk

Huckins, Laura M.; Dobbyn, Amanda; Ruderfer, Douglas M.; Hoffman, Gabriel; Wang, Weiqing; Pardiñas, Antonio; Rajagopal, Veera M.; Als, Thomas D.; Nguyen, Hoang; Girdhar, Kiran; Boocock, James; Roussos, Panos; Fromer, Menachem; Kramer, Robin; Domencini, Enrico; Gamazon, Eric; Purcell, Shaun; CommonMind Consortium; The Schizophrenia Working Group of the Psychiatric Genomics Consortium; iPSYCH-GEMS Schizophrenia Working Group; Demontis, Ditte; Børglum, Anders D.; Walters, James T. R.; O'Donovan, Michael C.; Sullivan, Patrick; Owen, Michael J.; Devlin, Bernie; Sieberts, Solveig K.; Cox, Nancy J.; Im, Hae Kyung; Sklar, Pamela; Stahl, Eli A.. (2019-03-25)

 
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URL:
https://doi.org/10.1038/s41588-019-0364-4

Huckins, Laura M.
Dobbyn, Amanda
Ruderfer, Douglas M.
Hoffman, Gabriel
Wang, Weiqing
Pardiñas, Antonio
Rajagopal, Veera M.
Als, Thomas D.
Nguyen, Hoang
Girdhar, Kiran
Boocock, James
Roussos, Panos
Fromer, Menachem
Kramer, Robin
Domencini, Enrico
Gamazon, Eric
Purcell, Shaun
CommonMind Consortium
The Schizophrenia Working Group of the Psychiatric Genomics Consortium
iPSYCH-GEMS Schizophrenia Working Group
Demontis, Ditte
Børglum, Anders D.
Walters, James T. R.
O'Donovan, Michael C.
Sullivan, Patrick
Owen, Michael J.
Devlin, Bernie
Sieberts, Solveig K.
Cox, Nancy J.
Im, Hae Kyung
Sklar, Pamela
Stahl, Eli A..
Springer Nature
25.03.2019

Huckins, L.M., Dobbyn, A., Ruderfer, D.M. et al. Gene expression imputation across multiple brain regions provides insights into schizophrenia risk. Nat Genet 51, 659–674 (2019). https://doi.org/10.1038/s41588-019-0364-4

https://rightsstatements.org/vocab/InC/1.0/
© 2020 Springer Nature Limited. This is a post-peer-review, pre-copyedit version of an article published in Nature Genetics. The final authenticated version is available online at: http://dx.doi.org/10.1038/s41588-019-0364-4
https://rightsstatements.org/vocab/InC/1.0/
doi:https://doi.org/10.1038/s41588-019-0364-4
Näytä kaikki kuvailutiedot
Julkaisun pysyvä osoite on
https://urn.fi/URN:NBN:fi-fe2020102085760
Tiivistelmä

Abstract

Transcriptomic imputation approaches combine eQTL reference panels with large-scale genotype data in order to test associations between disease and gene expression. These genic associations could elucidate signals in complex genome-wide association study (GWAS) loci and may disentangle the role of different tissues in disease development. We used the largest eQTL reference panel for the dorso-lateral prefrontal cortex (DLPFC) to create a set of gene expression predictors and demonstrate their utility. We applied DLPFC and 12 GTEx-brain predictors to 40,299 schizophrenia cases and 65,264 matched controls for a large transcriptomic imputation study of schizophrenia. We identified 413 genic associations across 13 brain regions. Stepwise conditioning identified 67 non-MHC genes, of which 14 did not fall within previous GWAS loci. We identified 36 significantly enriched pathways, including hexosaminidase-A deficiency, and multiple porphyric disorder pathways. We investigated developmental expression patterns among the 67 non-MHC genes and identified specific groups of pre- and postnatal expression.

 

Working Group

The Schizophrenia Working Group of the Psychiatric Genomics Consortium

Stephan Ripke37,38

Benjamin M. Neale37,38,39,40

Aiden Corvin41

James T. R. Walters6

Kai-How Farh37

Peter A. Holmans6,42

Phil Lee37,38,40

Brendan Bulik-Sullivan37,38

David A. Collier43,44

Hailiang Huang37,39

Tune H. Pers39,45,46

Ingrid Agartz47,48,49

Esben Agerbo8,32,33

Margot Albus50

Madeline Alexander51

Farooq Amin52,53

Silviu A. Bacanu54

Martin Begemann55

Richard A. Belliveau Jr38

Judit Bene56,57

Sarah E. Bergen38,58

Elizabeth Bevilacqua38

Tim B. Bigdeli54

Donald W. Black59

Richard Bruggeman60

Nancy G. Buccola61

Randy L. Buckner62,63,64

William Byerley65

Wiepke Cahn66

Guiqing Cai2,3

Dominique Campion67

Rita M. Cantor10

Vaughan J. Carr68,69

Noa Carrera6

Stanley V. Catts68,70

Kimberly D. Chambert38

Raymond C. K. Chan71

Ronald Y. L. Chen72

Eric Y. H. Chen72,73

Wei Cheng15

Eric F. C. Cheung74

Siow Ann Chong75

C. Robert Cloninger76

David Cohen77

Nadine Cohen78

Paul Cormican41

Nick Craddock6,42

James J. Crowley79

David Curtis80,81

Michael Davidson82

Kenneth L. Davis3

Franziska Degenhardt83,84

Jurgen Del Favero85

Ditte Demontis7,8,9

Dimitris Dikeos86

Timothy Dinan87

Srdjan Djurovic49,88

Gary Donohoe41,89

Elodie Drapeau3

Jubao Duan90,91

Frank Dudbridge92

Naser Durmishi93

Peter Eichhammer94

Johan Eriksson95,96,97

Valentina Escott-Price6

Laurent Essioux98

Ayman H. Fanous99,100,101,102

Martilias S. Farrell79

Josef Frank103

Lude Franke104

Robert Freedman105

Nelson B. Freimer106

Marion Friedl107

Joseph I. Friedman3

Menachem Fromer1,37,38,40

Giulio Genovese38

Lyudmila Georgieva6

Ina Giegling107,108

Paola Giusti-Rodríguez79

Stephanie Godard109

Jacqueline I. Goldstein37,39

Vera Golimbet110

Srihari Gopal111

Jacob Gratten112

Lieuwe de Haan113

Christian Hammer55

Marian L. Hamshere6

Mark Hansen114

Thomas Hansen8,30

Vahram Haroutunian3,25,23

Annette M. Hartmann107

Frans A. Henskens68,115,116

Stefan Herms83,84,117

Joel N. Hirschhorn39,46,118

Per Hoffmann83,84,117

Andrea Hofman83,84

Mads V. Hollegaard36

David M. Hougaard36

Masashi Ikeda119

Inge Joa120

Antonio Julia121

Rene S. Kahn66

Luba Kalaydjieva122,123

Sena Karachanak-Yankova124

Juha Karjalainen104

David Kavanagh6

Matthew C. Keller125

James L. Kennedy126,127,128

Andrey Khrunin129

Yunjung Kim79

Janis Klovins130

James A. Knowles131

Bettina Konte107

Vaidutis Kucinskas132

Zita Ausrele Kucinskiene132

Hana Kuzelova-Ptackova133

Anna K. Kahler58

Claudine Laurent51,134

Jimmy Lee Chee Keong75,135

S. Hong Lee112

Sophie E. Legge6

Bernard Lerer136

Miaoxin Li72,73,137

Tao Li138

Kung-Yee Liang139

Jeffrey Lieberman140

Svetlana Limborska129

Carmel M. Loughland68,141

Jan Lubinski142

Jouko Lonnqvist143

Milan Macek Jr133

Patrik K. E. Magnusson58

Brion S. Maher144

Wolfgang Maier145

Jacques Mallet146

Sara Marsal121

Manuel Mattheisen7,8,9,147

Morten Mattingsdal49,148

Robert W. McCarley149,150

Colm McDonald151

Andrew M. McIntosh152,153

Sandra Meier103

Carin J. Meijer113

Bela Melegh56,57

Ingrid Melle49,154

Raquelle I. Mesholam-Gately149,155

Andres Metspalu156

Patricia T. Michie68,157

Lili Milani156

Vihra Milanova158

Younes Mokrab43

Derek W. Morris41,89

Ole Mors8,9,159

Kieran C. Murphy160

Robin M. Murray161

Inez Myin-Germeys162

Bertram Muller-Myhsok163,164,165

Mari Nelis156

Igor Nenadic166

Deborah A. Nertney167

Gerald Nestadt168

Kristin K. Nicodemus169

Liene Nikitina-Zake130

Laura Nisenbaum170

Annelie Nordin171

Eadbhard O’Callaghan172

Colm O’Dushlaine38

F. Anthony O’Neill173

Sang-Yun Oh174

Ann Olincy126

Line Olsen8,64

Jim Van Os162,175

Christos Pantelis68,176

George N. Papadimitriou86

Sergi Papiol55

Elena Parkhomenko3

Michele T. Pato131

Tiina Paunio177,178

Milica Pejovic-Milovancevic179

Diana O. Perkins180

Olli Pietiläinen178,181

Jonathan Pimm81

Andrew J. Pocklington6

John Powell161

Alkes Price39,182

Ann E. Pulver168

Shaun M. Purcell1

Digby Quested183

Henrik B. Rasmussen30,43

Abraham Reichenberg3

Mark A. Reimers184

Alexander L. Richards6

Joshua L. Roffman62,64

Panos Roussos1,4

Douglas M. Ruderfer1,5,6

Veikko Salomaa97

Alan R. Sanders90,91

Ulrich Schall68,141

Christian R. Schubert185

Thomas G. Schulze103,186

Sibylle G. Schwab187

Edward M. Scolnick38

Rodney J. Scott68,188,189

Larry J. Seidman144,155

Jianxin Shi190

Engilbert Sigurdsson191

Teimuraz Silagadze192

Jeremy M. Silverman3,193

Kang Sim75

Petr Slominsky129

Jordan W. Smoller38,40

Hon-Cheong So72

Chris C. A. Spencer194

Eli A. Stahl1,2,3,4

Hreinn Stefansson195

Stacy Steinberg195

Elisabeth Stogmann196

Richard E. Straub197

Eric Strengman66,198

Jana Strohmaier103

T. Scott Stroup140

Mythily Subramaniam75

Jaana Suvisaari143

Dragan M. Svrakic76

Jin P. Szatkiewicz79

Erik Soderman47

Srinivas Thirumalai199

Draga Toncheva124

Sarah Tosato200

Juha Veijola201,202

John Waddington203

Dermot Walsh204

Dai Wang111

Qiang Wang138

Bradley T. Webb54

Mark Weiser82

Dieter B. Wildenauer205

Nigel M. Williams6

Stephanie Williams79

Stephanie H. Witt103

Aaron R. Wolen184

Emily H. M. Wong72

Brandon K. Wormley54

Hualin Simon Xi206

Clement C. Zai126,127

Xuebin Zheng207

Fritz Zimprich196

Naomi R. Wray112

Kari Stefansson195

Peter M. Visscher112

Rolf Adolfsson171

Ole A. Andreassen49,154

Douglas H. R. Blackwood153

Elvira Bramon208

Joseph D. Buxbaum2,3,24,25

Anders D. Børglum7,8,9,159

Sven Cichon83,84,117,209

Ariel Darvasi210

Enrico Domenici211

Hannelore Ehrenreich55

Tonu Esko39,46,118,156

Pablo V. Gejman90,91

Michael Gill41

Hugh Gurling81

Christina M. Hultman58

Nakao Iwata119

Assen V. Jablensky68,123,205,212

Erik G. Jonsson47,49

Kenneth S. Kendler213

George Kirov6

Jo Knight125,127,128

Todd Lencz214,215,216

Douglas F. Levinson51

Qingqin S. Li111

Jianjun Liu207,217

Anil K. Malhotra214,215,216

Steven A. McCarroll38,118

Andrew McQuillin81

Jennifer L. Moran38

Preben B. Mortensen8,32,33

Bryan J. Mowry112,218

Markus M. Nothen83,84

Roel A. Ophoff10,66,105

Michael J. Owen6,42

Aarno Palotie38,40,181

Carlos N. Pato131

Tracey L. Petryshen38,149,219

Danielle Posthuma220,221,222

Marcella Rietschel103

Brien P. Riley213

Dan Rujescu107,108

Pak C. Sham72,73,137

Pamela Sklar1,2,3,4,25

David St Clair223

Daniel R. Weinberger197,224

Jens R. Wendland185

Thomas Werge8,30,225

Mark J. Daly37,38,39

Patrick F. Sullivan58,79,180

Michael C. O’Donovan6,42

20Integrated Technology Research Laboratories, Pharmaceutical Research Division, Takeda Pharmaceutical Company Limited, Fujisawa, Japan.

21Neuropsychiatry Section, Department of Psychiatry, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, USA.

22Neuropsychiatric Signaling Program, Department of Psychiatry, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, USA.

23Psychiatry, JJ Peters Virginia Medical Center, Bronx, NY, USA.

24Department of Neuroscience, Icahn School of Medicine at Mount Sinai, New York, New York, NY, USA.

25Friedman Brain Institute, Icahn School of Medicine at Mount Sinai, New York, NY, USA.

26Seaver Autism Center for Research and Treatment, Icahn School of Medicine at Mount Sinai, New York, NY, USA.

27CNS Drug Discovery Unit, Pharmaceutical Research Division, Takeda Pharmaceutical Company Limited, Fujisawa, Japan.

28F. Hoffman-La Roche Ltd, Basel, Switzerland.

29Bioinformatics Research Centre, Aarhus University, Aarhus, Denmark.

30Institute of Biological Psychiatry, MHC Sct. Hans, Mental Health Services Copenhagen, Roskilde, Denmark.

31Department of Clinical Medicine, University of Copenhagen, Copenhagen, Denmark.

32National Centre for Register-Based Research, Aarhus University, Aarhus, Denmark.

33Centre for Integrated Registerbased Research, Aarhus University, Aarhus, Denmark.

34Psychosis Research Unit, Aarhus University Hospital, Risskov, Denmark.

35Mental Health Services in the Capital Region of Denmark, Mental Health Center Copenhagen, University of Copenhagen, Copenhagen, Denmark.

36Center for Neonatal Screening, Department for Congenital Disorders, Statens Serum Institut, Copenhagen, Denmark.

37Analytic and Translational Genetics Unit, Massachusetts General Hospital, Boston, MA, USA.

38Stanley Center for Psychiatric Research, Broad Institute of MIT and Harvard, Cambridge, MA, USA.

39Medical and Population Genetics Program, Broad Institute of MIT and Harvard, Cambridge, MA, USA.

40Psychiatric and Neurodevelopmental Genetics Unit, Massachusetts General Hospital, Boston, MA, USA.

41Neuropsychiatric Genetics Research Group, Department of Psychiatry, Trinity College Dublin, Dublin, Ireland.

42NationalCentre for Mental Health, Cardiff University, Cardiff, UK.

43Eli Lilly and Company Limited, Erl Wood Manor, Sunninghill Road, Windlesham, Surrey, UK.

44Social, Genetic and Developmental Psychiatry Centre, Institute of Psychiatry, King’s College London, London, UK.

45Center for BiologicalSequence Analysis, Department of Systems Biology, Technical University of Denmark, Kongens Lyngby, Denmark.

46Division of Endocrinology and Center for Basic and Translational Obesity Research, Boston Children’s Hospital, Boston, MA, USA.

47Department of Clinical Neuroscience, Psychiatry Section, Karolinska Institutet, Stockholm, Sweden.

48Department of Psychiatry, Diakonhjemmet Hospital, Oslo, Norway.

49NORMENT, KG Jebsen Centre for Psychosis Research, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.

50State Mental Hospital, Haar, Germany.

51Department of Psychiatry and Behavioral Sciences, Stanford University, Stanford, CA, USA.

52Department of Psychiatry and Behavioral Sciences, Atlanta Veterans Affairs Medical Center, Atlanta, GA, USA.

53Department of Psychiatry and Behavioral Sciences, Emory University, Atlanta, GA, USA.

54Virginia Institute for Psychiatric and Behavioral Genetics, Department of Psychiatry, Virginia Commonwealth University, Richmond, VA, USA.

55Clinical Neuroscience, Max Planck Institute of Experimental Medicine, Gottingen, Germany.

56Department of Medical Genetics, University of Pécs, Pécs, Hungary.

57Szentagothai Research Center, University of Pécs, Pécs, Hungary.

58Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden.

59Department of Psychiatry, University of Iowa Carver College of Medicine, Iowa City, IA, USA.

60University Medical Center Groningen, Department of Psychiatry, University of Groningen, Groningen, the Netherlands.

61School of Nursing, Louisiana State University Health Sciences Center, New Orleans, LA, USA.

62Athinoula A. Martinos Center, Massachusetts General Hospital, Boston, MA, USA.

63Center for Brain Science, Harvard University, Cambridge, MA, USA.

64Department of Psychiatry, Massachusetts General Hospital, Boston, MA, USA.

65Department of Psychiatry, University of California at San Francisco, San Francisco, CA, USA.

66University Medical Center Utrecht, Department of Psychiatry, Rudolf Magnus Institute of Neuroscience, Utrecht, the Netherlands.

67Centre Hospitalier du Rouvray and INSERM U1079 Faculty of Medicine, Rouen, France.

68Schizophrenia Research Institute, Sydney, New South Wales, Australia.

69School of Psychiatry, University of New South Wales, Sydney, New South Wales, Australia.

70Royal Brisbane and Women’s Hospital, University of Queensland, Brisbane, Queensland, Australia.

71Institute of Psychology, Chinese Academy of Science, Beijing, China.

72Department of Psychiatry, Li Ka Shing Faculty of Medicine, The University of Hong Kong, Hong Kong, China.

73State Key Laboratory for Brain and Cognitive Sciences, Li Ka Shing Faculty of Medicine, The University of Hong Kong, Hong Kong, China.

74Castle Peak Hospital, Hong Kong, China.

75Institute of Mental Health, Singapore, Singapore.

76Department of Psychiatry, Washington University, St. Louis, MO, USA.

77Department of Child and Adolescent Psychiatry, Assistance Publique Hopitaux de Paris, Pierre and Marie Curie Faculty of Medicine and Institute for Intelligent Systems and Robotics, Paris, France.

78Blue Note Biosciences, Princeton, NJ, USA.

79Department of Genetics, University of North Carolina, Chapel Hill, NC, USA.

80Department of Psychological Medicine, Queen Mary University of London, London, UK.

81Molecular Psychiatry Laboratory, Division of Psychiatry, University College London, London, UK.

82Sheba Medical Center, Tel Hashomer, Israel.

83Department of Genomics, Life and Brain Center, Bonn, Germany.

84Institute of Human Genetics, University of Bonn, Bonn, Germany.

85AppliedMolecular Genomics Unit, VIB Department of Molecular Genetics, University of Antwerp, Antwerp, Belgium.

86First Department of Psychiatry, University of Athens Medical School, Athens, Greece.

87Department of Psychiatry, University College Cork, Co, Cork, Ireland.

88Department of Medical Genetics, Oslo University Hospital, Oslo, Norway.

89Cognitive Genetics and Therapy Group, School of Psychology and Discipline of Biochemistry, National University of Ireland Galway, Co, Galway, Ireland.

90Department of Psychiatry and Behavioral Neuroscience, University of Chicago, Chicago, IL, USA.

91Department of Psychiatry and Behavioral Sciences, North Shore University Health System, Evanston, IL, USA.

92Department of Non-Communicable Disease Epidemiology, London School of Hygiene and Tropical Medicine, London, UK.

93Department of Child and Adolescent Psychiatry, University Clinic of Psychiatry, Skopje, Republic of Macedonia.

94Department of Psychiatry, University of Regensburg, Regensburg, Germany.

95Department of General Practice, Helsinki University Central Hospital, University of Helsinki, Helsinki, Finland.

96Folkhälsan Research Center, Helsinki, Finland, Biomedicum Helsinki, Helsinki, Finland.

97National Institute for Health and Welfare, Helsinki, Finland.

98Translational Technologies and Bioinformatics, Pharma Research and Early Development, F. Hoffman-La Roche, Basel, Switzerland.

99Department of Psychiatry, Georgetown University School of Medicine, Washington, DC, USA.

100Department of Psychiatry, Keck School of Medicine of the University of Southern California, Los Angeles, CA, USA.

101Department of Psychiatry, Virginia Commonwealth University School of Medicine, Richmond, VA, USA.

102Mental Health Service Line, Washington VA Medical Center, Washington, DC, USA.

103Department of Genetic Epidemiology in Psychiatry, Central Institute of Mental Health, Medical Faculty Mannheim, University of Heidelberg, Heidelberg, Mannheim, Germany.

104Department of Genetics, University of Groningen, University Medical Centre Groningen, Groningen, the Netherlands.

105Department of Psychiatry, University of Colorado Denver, Aurora, CO, USA.

106Center for Neurobehavioral Genetics, Semel Institute for Neuroscience and Human Behavior, University of California Los Angeles, Los Angeles, CA, USA.

107Department of Psychiatry, University of Halle, Halle, Germany.

108Department of Psychiatry, University of Munich, Munich, Germany.

109Departments of Psychiatry and Human and Molecular Genetics, INSERM, Institut de Myologie, Hôpital de la Pitiè-Salpêtrière, Paris, France.

110Mental Health Research Centre, Russian Academy of Medical Sciences, Moscow, Russia.

111Neuroscience Therapeutic Area, Janssen Research and Development, Raritan, NJ, USA.

112Queensland Brain Institute, The University of Queensland, Brisbane, Queensland, Australia.

113Academic Medical Centre University of Amsterdam, Department of Psychiatry, Amsterdam, the Netherlands.

114Illumina, La Jolla, CA, USA.

115Priority Research Centre for Health Behaviour, University of Newcastle, Newcastle, New South Wales, Australia.

116School of Electrical Engineering and Computer Science, University of Newcastle, Newcastle, New South Wales, Australia.

117Division of Medical Genetics, Department of Biomedicine, University of Basel, Basel, Switzerland.

118Department of Genetics, Harvard Medical School, Boston, MA, USA.

119Department of Psychiatry, Fujita Health University School of Medicine, Toyoake, Japan.

120Regional Centre for Clinical Researchin Psychosis, Department of Psychiatry, Stavanger University Hospital, Stavanger, Norway.

121Rheumatology Research Group, Vall d’Hebron Research Institute, Barcelona, Spain.

122Centre for Medical Research, The University of Western Australia, Perth, Western Australia, Australia.

123The Perkins Institute for Medical Research, The University of Western Australia, Perth, Western Australia, Australia.

124Department of Medical Genetics, Medical University, Sofia, Bulgaria.

125Department of Psychology, University of Colorado Boulder, Boulder, CO, USA.

126Campbell Family Mental Health Research Institute, Centre for Addiction and Mental Health, Toronto, Ontario, Canada.

127Department of Psychiatry, University of Toronto, Toronto, Ontario, Canada.

128Institute of Medical Science, University of Toronto, Toronto, Ontario, Canada.

129Institute of Molecular Genetics, Russian Academy of Sciences, Moscow, Russia.

130Latvian Biomedical Research and Study Centre, Riga, Latvia.

131Department of Psychiatry and Zilkha Neurogenetics Institute, Keck School of Medicine at University of Southern California, Los Angeles, CA, USA.

132Faculty of Medicine, Vilnius University, Vilnius, Lithuania.

133Department of Biology and Medical Genetics, 2nd Faculty of Medicine and University Hospital Motol, Prague, Czech Republic.

134Department of Child and Adolescent Psychiatry, Pierre and Marie Curie Faculty of Medicine, Paris, France.

135Duke-NUS Graduate Medical School, Singapore, Singapore.

136Department of Psychiatry, Hadassah-Hebrew University Medical Center, Jerusalem, Israel.

137Centre for Genomic Sciences, The University of Hong Kong, Hong Kong, China.

138Mental Health Centre and Psychiatric Laboratory, West China Hospital, Sichuan University, Chengdu, Sichuan, China.

139Department of Biostatistics, Johns Hopkins University Bloomberg School of Public Health, Baltimore, MD, USA.

140Department of Psychiatry, Columbia University, New York, New York, NY, USA.

141Priority Centre for Translational Neuroscience and Mental Health, University of Newcastle, Newcastle, New South Wales, Australia.

142Department of Genetics and Pathology, International Hereditary Cancer Center, Pomeranian Medical University in Szczecin, Szczecin, Poland.

143Department of Mental Health and Substance Abuse Services, National Institute for Health and Welfare, Helsinki, Finland.

144Department of Mental Health, Bloomberg School of Public Health, Johns Hopkins University, Baltimore, MD, USA.

145Department of Psychiatry, University of Bonn, Bonn, Germany.

146Centre National de la Recherche Scientifique, Laboratoire de Génétique Moléculaire de la Neurotransmission et des Processus Neurodénégératifs, Hôpital de la Pitiè-Salpêtrière, Paris, France.

147Department of Genomics Mathematics, University of Bonn, Bonn, Germany.

148Research Unit, Sørlandet Hospital, Kristiansand, Norway.

149Department of Psychiatry, Harvard Medical School, Boston, MA, USA.

150VA Boston Health Care System, Brockton, MA, USA.

151Department of Psychiatry, National University of Ireland Galway, Co, Galway, Ireland.

152Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, UK.

153Division of Psychiatry, University of Edinburgh, Edinburgh, UK.

154Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.

155Massachusetts Mental Health Center Public Psychiatry Division of the Beth Israel Deaconess Medical Center, Boston, MA, USA.

156Estonian Genome Center, University of Tartu, Tartu, Estonia.

157School of Psychology, University of Newcastle, Newcastle, New South Wales, Australia.

158First Psychiatric Clinic, Medical University, Sofia, Bulgaria.

159Department P, Aarhus University Hospital, Risskov, Denmark.

160Department of Psychiatry, Royal College of Surgeons in Ireland, Dublin, Ireland.

161King’s College London, London, UK.

162Maastricht University Medical Centre, South Limburg Mental Health Research and TeachingNetwork, EURON, Maastricht, the Netherlands.

163Institute of Translational Medicine, University of Liverpool, Liverpool, UK.

164Max Planck Institute of Psychiatry, Munich, Germany.

165Munich Cluster for Systems Neurology (SyNergy), Munich, Germany.

166Department of Psychiatry and Psychotherapy, Jena University Hospital, Jena, Germany.

167Department of Psychiatry, Queensland Brain Institute and Queensland Centre for Mental Health Research, University of Queensland, Brisbane, Queensland, Australia.

168Department of Psychiatry and Behavioral Sciences, Johns Hopkins University School of Medicine, Baltimore, MD, USA.

169Department of Psychiatry, Trinity College Dublin, Dublin, Ireland.

170Eli Lilly and Company, Lilly Corporate Center, Indianapolis, IN, USA.

171Department of Clinical Sciences, Psychiatry, Umeå University, Umeå, Sweden.

172DETECT Early Intervention Service for Psychosis, Blackrock, Co, Dublin, Ireland.

173Centre for Public Health, Institute of Clinical Sciences, Queen’s University Belfast, Belfast, UK.

174Lawrence Berkeley National Laboratory, University of California at Berkeley, Berkeley, CA, USA.

175Institute of Psychiatry, King’s College London, London, UK.

176Melbourne Neuropsychiatry Centre, University of Melbourne & Melbourne Health, Melbourne, Victoria, Australia.

177Department of Psychiatry, University of Helsinki, Helsinki, Finland.

178Public Health Genomics Unit, National Institute for Health and Welfare, Helsinki, Finland.

179Medical Faculty, University of Belgrade, Belgrade, Serbia.

180Department of Psychiatry, University of North Carolina, Chapel Hill, NC, USA.

181Institute for Molecular Medicine Finland, FIMM, University of Helsinki, Helsinki, Finland.

182Department of Epidemiology, Harvard School of Public Health, Boston, MA, USA.

183Department of Psychiatry, University of Oxford, Oxford, UK.

184Virginia Institute for Psychiatric and Behavioral Genetics, Virginia Commonwealth University, Richmond, VA, USA.

185Pharma Therapeutics Clinical Research, Pfizer Worldwide Research and Development, Cambridge, MA, USA.

186Department of Psychiatry and Psychotherapy, University of Gottingen, Göttingen, Germany.

187Psychiatry and Psychotherapy Clinic, University of Erlangen, Erlangen, Germany.

188Hunter New England Health Service, Newcastle, New South Wales, Australia.

189School of Biomedical Sciences, University of Newcastle, Newcastle, New South Wales, Australia.

190Division of Cancer Epidemiology and Genetics, National Cancer Institute, Bethesda, MD, USA.

191University of Iceland, Landspitali, National University Hospital, Reykjavik, Iceland.

192Department of Psychiatry and Drug Addiction, Tbilisi State Medical University (TSMU), Tbilisi, Georgia.

193Research and Development, Bronx Veterans Affairs Medical Center, New York, NY, USA.

194WellcomeTrust Centre for Human Genetics, Oxford, UK.

195deCODE Genetics, Reykjavik, Iceland.

196Department of Clinical Neurology, Medical University of Vienna, Wien, Austria.

197Lieber Institute for Brain Development, Baltimore, MD, USA.

198Department of Medical Genetics, University Medical Centre Utrecht, Utrecht, the Netherlands.

199Berkshire Healthcare NHS Foundation Trust, Bracknell, UK.

200Section of Psychiatry, University of Verona, Verona, Italy.

201Department of Psychiatry, University of Oulu, Oulu, Finland.

202University Hospital of Oulu, Oulu, Finland.

203Molecular and Cellular Therapeutics, Royal College of Surgeons in Ireland, Dublin, Ireland.

204Health Research Board, Dublin, Ireland.

205School of Psychiatry and Clinical Neurosciences, The University of Western Australia, Perth, Western Australia, Australia.

206Computational Sciences CoE, Pfizer Worldwide Research and Development, Cambridge, MA, USA.

207Human Genetics, Genome Institute of Singapore, A*STAR, Singapore, Singapore.

208University College London, London, UK.

209Institute of Neuroscience and Medicine (INM-1), Research Center Juelich, Juelich, Germany.

210Department of Genetics, The Hebrew University of Jerusalem, Jerusalem, Israel.

211NeuroscienceDiscovery and Translational Area, Pharma Research and Early Development, F. Hoffman-La Roche, Basel, Switzerland.

212Centre for Clinical Research in Neuropsychiatry, School of Psychiatry and Clinical Neurosciences, The University of Western Australia, Medical Research Foundation Building, Perth, Western Australia, Australia.

213Virginia Institute for Psychiatric and Behavioral Genetics, Departments of Psychiatry and Human and Molecular Genetics, Virginia Commonwealth University, Richmond, VA, USA.

214The Feinstein Institute for Medical Research, Manhasset, NY, USA.

215The Hofstra NS-LIJ School of Medicine, Hempstead, NY, USA.

216The Zucker Hillside Hospital, Glen Oaks, NY, USA.

217Saw Swee Hock School of Public Health, National University of Singapore, Singapore, Singapore.

218Queensland Centre for Mental Health Research, University of Queensland, Brisbane, Queensland, Australia.

219Center for HumanGenetic Research and Department of Psychiatry, Massachusetts General Hospital, Boston, MA, USA.

220Department of Child and Adolescent Psychiatry, Erasmus University Medical Centre, Rotterdam, the Netherlands.

221Department of Complex Trait Genetics, Neuroscience Campus Amsterdam, VU University Medical Center Amsterdam, Amsterdam, the Netherlands.

222Department of Functional Genomics, Center for Neurogenomics and Cognitive Research, Neuroscience Campus Amsterdam, VU University, Amsterdam, the Netherlands.

223University of Aberdeen, Institute of Medical Sciences, Aberdeen, UK.

224Departments of Psychiatry, Neurology, Neuroscience and Institute of Genetic Medicine, Johns Hopkins School of Medicine, Baltimore, MD, USA.

225Department of Clinical Medicine, University of Copenhagen, Copenhagen, Denmark.

 
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