Abstract

Schizophrenia is a devastating psychiatric illness with high heritability. Brain structure and function differ, on average, between people with schizophrenia and healthy individuals. As common genetic associations are emerging for both schizophrenia and brain imaging phenotypes, we can now use genome-wide data to investigate genetic overlap. Here we integrated results from common variant studies of schizophrenia (33,636 cases, 43,008 controls) and volumes of several (mainly subcortical) brain structures (11,840 subjects). We did not find evidence of genetic overlap between schizophrenia risk and subcortical volume measures either at the level of common variant genetic architecture or for single genetic markers. These results provide a proof of concept (albeit based on a limited set of structural brain measures) and define a roadmap for future studies investigating the genetic covariance between structural or functional brain phenotypes and risk for psychiatric disorders.

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Acknowledgements

PGC. The authors are grateful to the many family members who participated in the studies that recruited these samples, to the many clinicians who assisted in their recruitment, and to our team members, without whom this study would have been impossible. Core funding for the PGC is from the US National Institute of Mental Health (NIMH; U01 MH094421). Statistical analyses were carried out on the Genetic Cluster Computer (http://www.geneticcluster.org/) hosted by SURFsara and financially supported by the Netherlands Scientific Organization (NWO 480-05-003), along with a supplement from the Dutch Brain Foundation and the VU University Amsterdam. The GRAS data collection was supported by the Max Planck Society, the Max-Planck-Förderstiftung and the DFG Center for Nanoscale Microscopy & Molecular Physiology of the Brain (CNMPB), Göttingen, Germany. The Boston CIDAR project was supported by the NIMH (P50 MH080272, R.W.M.; U01 MH081928, L.J.S.; R01 MH092380, T.L.P.) and the Massachusetts General Hospital Executive Committee on Research (T.L.P.). P.H.L. is supported by NIMH K99 MH101367. ISC Portugal: C.N.P. and M.T.P. have been supported by NIMH grants MH085548, MH085542, MH071681, MH061884, MH58693 and MH52618 and NCRR grant RR026075. C.N.P., M.T.P. and A.H.F. have been supported by grants from the Department of Veterans Affairs Merit Review Program. The Danish Aarhus study was supported by grants from Lundbeck Foundation, Danish Strategic Research Council, Aarhus University, and Stanley Research Foundation. Work in Cardiff was supported by UK Medical Research Council (MRC) Centre (G0800509) and MRC Programme (G0801418) grants, the European Community's Seventh Framework Programme (HEALTH-F2-2010-241909, Project EU-GEI) the European Union Seventh Framework Programme (FP7/2007-2013) under grant agreement 279227, a fellowship to J.W. from the MRC/Welsh Assembly Government and the Margaret Temple Award from the British Medical Association. We thank Novartis for their input in obtaining CLOZUK samples, staff at The Doctor's Laboratory (L. Levett and A. Levett) for help with sample acquisition and data linkage, and staff in Cardiff (K. Mantripragada and L. Hopkins) for sample management. CLOZUK and some other samples were genotyped at the Broad Institute or by the Wellcome Trust Case Control Consortium (WTCCC) and Wellcome Trust Case Control Consortium 2 (WTCCC2) (WT 083948/Z/07/Z). We acknowledge use of the British 1958 Birth Cohort DNA (MRC: G0000934) and the Wellcome Trust (068545/Z/0 and 076113/C/04/Z), the UK Blood Services Common Controls (UKBS-CC collection), funded by the Wellcome Trust (076113/C/04/Z) and by a National Institute for Health Research programme grant to National Health Service Blood and Transplant (RP-PG-0310-1002). Virginia Commonwealth University investigators were supported by NIMH grants R01 MH083094, R01 MH041953, and R01 MH068881 and WTCCC2 grant WTCCC-084710. Recruitment of families in Bulgaria was funded by the Janssen Research Foundation, Beerse, Belgium. We thank the staff in the Neuroscience Biomarkers Genomic Lab led by R. Favis at Janssen for sample processing and the staff at Illumina for genotyping Janssen DNA samples. We also thank A. Santos, N. Bottrel, M.-A. Franc and W. Cafferty of Janssen Research & Development) for operational support. Dutch samples were funded by the Netherlands Organization for Health Research and Development (ZonMw) in the Mental Health program and by NIMH R01 MH078075. Danish samples were funded by the Danish Council for Strategic Research (Journ.nr. 09-067048), the Danish National Advanced Technology Foundation (Journ.nr. 001-2009-2), the Lundbeck Foundation (Journ.nr. R24-A3243), and the EU 7th Framework Programme. The Wellcome Trust supported this study as part of the WTCCC2 project. E. Bramon holds a MRC New Investigator Award and a MRC Centenary Award. The TOP Study was supported by the Research Council of Norway (213837, 217776, 223273), South-East Norway Health Authority (2013-123) and K.G. Jebsen Foundation. This work was supported by the Donald and Barbara Zucker Foundation, the North Shore – Long Island Jewish Health System Foundation and grants from the Stanley Foundation (A.K.M.), NARSAD (A.K.M.), NIMH (MH065580 to T. Lencz; MH001760 to A.K.M.), and NIMH RC2 MH089964 and R01 MH084098. Finnish samples were funded by SynSys (EU FP7-242167), Sigrid Juselius Foundation, the Academy of Finland (grant 251704), and the Sohlberg Foundation. The Swedish Research Council (grants 2006-4472, 2009-5269, 2009-3413) and the County Councils of Västerbotten and Norrbotten, Sweden, supported the collection of the Umeå samples. The Betula Study, from which the Umeå controls were recruited, is supported by grants from the Swedish Research Council (grants 345-2003-3883, 315-2004-6977) and the Bank of Sweden Tercentenary Foundation, the Swedish Council for Planning and Coordination of Research, the Swedish Council for Research in the Humanities and Social Sciences and the Swedish Council for Social Research. We acknowledge support from NIMH K01 MH085812 (M.C.K.) and NIMH R01 MH100141 (M.C.K.). Estonian Genome Center at the University of Tartu (EGCUT) work was supported by Targeted Financing from the Estonian Ministry of Science and Education (SF0180142s08), US National Institutes of Health (NIH) grant R01 DK075787, the Development Fund of the University of Tartu (grant SP1GVARENG), the European Regional Development Fund to the Centre of Excellence in Genomics (EXCEGEN; grant 3.2.0304.11-0312) and FP7 grant 313010. M. Macek was supported by CZ.2.16/3.1.00/24022OPPK, NT/13770-4 and 00064203 FN Motol. Funding from the Singapore National Medical Research Council (NMRC/TCR/003/2008) and the Singapore Biomedical Research Council. We acknowledge the support of the Singapore Agency for Science, Technology and Research (A*STAR). Genotyping of the Swedish Hubin sample was performed by the SNP&SEQ Technology Platform in Uppsala, which is supported by Uppsala University, Uppsala University Hospital, Science for Life Laboratory and the Swedish Research Council (contracts 80576801 and 70374401). The Swedish Hubin sample was supported by Swedish Research Council (I.A., E.G.J.) and Stockholm County Council and the Karolinska Insititutet (E.G.J.). B.J.M., V.J.C., R.J.S., S.V.C., F.A.H., A.V.J., C.M.L., P.T.M., C.P. and U.S. were supported by the Australian Schizophrenia Research Bank, which is supported by an enabling grant from the National Health and Medical Research Council (386500), the Pratt Foundation, Ramsay Health Care, the Viertel Charitable Foundation, the Schizophrenia Research Institute and the NSW Department of Health. C.P. is supported by a Senior Principal Research Fellowship from the National Health and Medical Research Council (Australia). The Perth sample collection was funded by Australian National Health and Medical Research Council project grants and the Australian Schizophrenia Research Bank. The Bonn/Mannheim sample was genotyped in a study that was supported by the German Federal Ministry of Education and Research (BMBF) through the Integrated Genome Research Network MooDS (Systematic Investigation of the Molecular Causes of Major Mood Disorders and Schizophrenia; grant 01GS08144 to M.M.N. and S.C., grant 01GS08147 to M.R.), under the National Genome Research Network plus (NGFNplus), and the Integrated Network IntegraMent (Integrated Understanding of Causes and Mechanisms in Mental Disorders), under e:Med Programme (GlaxoSmithKline control sample; B.M.-M.) This work has been funded by the Bavarian Ministry of Commerce and by the BMBF in the framework of the National Genome Research Network, Förderkennzeichen 01GS0481 and the Bavarian Ministry of Commerce. M.M.N. is a member of the German Research Foundation (DFG)-funded Excellence Cluster ImmunoSensation. M.M.N. also received support from the Alfried Krupp von Bohlen und Halbach-Stiftung. M.R. was supported by the 7th Framework Programme of the European Union (ADAMS project, HEALTH-F4-2009-242257; CRESTAR project, HEALTH-2011-1.1-2) grant 279227. J. Knight holds the Joanne Murphy Professor in Behavioural Science. The Stanley Center for Psychiatric Research at the Broad Institute acknowledges funding from the Stanley Medical Research Institute. Support for the Sweden Schizophrenia Study (principal investigators P.F.S., C.M.H. and P. Sklar) was provided by the NIMH (R01 MH077139 and R01 MH095034), the Stanley Center for Psychiatric Research, the Sylvan Herman Foundation, the Friedman Brain Institute at the Mount Sinai School of Medicine, the Karolinska Institutet, Karolinska University Hospital, the Swedish Research Council, the Swedish County Council and the Söderström Königska Foundation. We acknowledge use of DNA from the UK Blood Services Collection of Common Controls (UKBS collection), funded by the Wellcome Trust grant 076113/CI04/Z, by Juvenile Diabetes Research Foundation grant WT0618S8, and by the National Institute of Health Research of England. The Multicenter Genetics Studies of Schizophrenia and Molecular Genetics of Schizophrenia studies study were supported by NIMH grant R01 MH062276 (to D.F.L., C.L., M.J.O. and D.B.W.), grant R01 MH068922 (to P.V.G.), grant R01 MH068921 (to A.E.P.) and grant R01 MH068881 (to B.P.R.). D.F.L. was supported by the Walter E. Nichols, M.D., Professorship in the School of Medicine, the Eleanor Nichols Endowment, the Walter F. & Rachael L. Nichols Endowment and the William and Mary McIvor Endowment, Stanford University. This study was supported by NIH R01 grants (MH67257 to N.G.B., MH59588 to B.J.M., MH59571 to P.V.G., MH59565 to R.F., MH59587 to F.A., MH60870 to W.F.B., MH59566 to D.W.B., MH59586 to J.M.S., MH61675 to D.F.L., MH60879 to C.R.C. and MH81800 to P.V.G.), NIH U01 grants (MH46276 to C.R.C., MH46289 to C. Kaufmann, MH46318 to M.T. Tsuang, MH79469 to P.V.G. and MH79470 to D.F.L.), the Genetic Association Information Network (GAIN) and The Paul Michael Donovan Charitable Foundation. Genotyping was carried out by the Center for Genotyping and Analysis at the Broad Institute of Harvard and MIT (S. Gabriel and D.B. Mirel), supported by grant U54 RR020278 from the National Center for Research Resources. D.R.W. and R.E.S. thank the staff of the Lieber Institute and the Clinical Brain Disorders Branch of the Intramural Research Program of the NIMH for their assistance in data collection and management. We acknowledge the Irish contribution to the International Schizophrenia Consortium (ISC) study, the WTCCC2 schizophrenia study and WTCCC2 controls from the 1958BC and UKNBS, the Science Foundation Ireland (08/IN.1/B1916). We acknowledge use of the Trinity Biobank sample from the Irish Blood Transfusion Service and the Trinity Centre for High Performance Computing. Funding for this study was provided by the WTCCC2 project (085475/B/08/Z and 085475/Z/08/Z), the Wellcome Trust (072894/Z/03/Z, 090532/Z/09/Z and 075491/Z/04/B), NIMH grants (MH 41953 and MH083094) and British 1958 Birth Cohort DNA collection funded by the MRC (grant G0000934) and the Wellcome Trust (grant 068545/Z/02). Collection of the UK National Blood Service controls was funded by the Wellcome Trust. We acknowledge Hong Kong Research Grants Council project grants GRF 774707M, 777511M, 776412M and 776513M.ENIGMA. ENIGMA was supported in part by a consortium grant (U54 EB020403 to P.M.T.) from the NIH institutes contributing to the Big Data to Knowledge (BD2K) initiative, including the NIBIB and NCI. ADNI and ADNI2GO: Data collection and sharing for this project was funded by the Alzheimer's Disease Neuroimaging Initiative (ADNI) (NIH grant U01 AG024904) and US Department of Defense (award W81XWH-12-2-0012). ADNI is funded by the US National Institute on Aging, the National Institute of Biomedical Imaging and Bioengineering, and through generous contributions from the following: Alzheimer's Association; Alzheimer's Drug Discovery Foundation; BioClinica, Inc.; Biogen Idec Inc.; Bristol-Myers Squibb Company; Eisai Inc.; Elan Pharmaceuticals, Inc.; Eli Lilly and Company; F. Hoffmann-La Roche Ltd and its affiliated company Genentech, Inc.; GE Healthcare; Innogenetics, N.V.; IXICO Ltd.; Janssen Alzheimer Immunotherapy Research & Development, LLC.; Johnson & Johnson Pharmaceutical Research & Development LLC.; Medpace, Inc.; Merck & Co., Inc.; Meso Scale Diagnostics, LLC.; NeuroRx Research; Novartis Pharmaceuticals Corporation; Pfizer Inc.; Piramal Imaging; Servier; Synarc Inc.; and Takeda Pharmaceutical Company. The Canadian Institutes of Health Research is providing funds to support ADNI clinical sites in Canada. Private sector contributions are facilitated by the Foundation for the National Institutes of Health. The grantee organization is the Northern California Institute for Research and Education, and the study is coordinated by the Alzheimer's Disease Cooperative Study at the University of California, San Diego. ADNI data are disseminated by the Laboratory of Neuro Imaging at the University of Southern California. Betula: this sample collection was supported by a Wallenberg Scholar grant from the Knut and Alice Wallenberg Foundation and a grant from Torsten and Ragnar Söderbergs Foundation to L.N., and a grant from HelseVest RHF (grant 911554) to S.L.H. Bipolar Family Study: The Bipolar Family Study wishes to thank the Scottish Mental Health Research Network for research assistant support; the Brain Research Imaging Centre Edinburgh, a center in the Scottish Funding Council Scottish Imaging Network–A Platform for Scientific Excellence (SINAPSE) Collaboration, for image acquisition; and the Wellcome Trust Clinical Research Facility for genotyping. Genotyping was supported by a National Alliance for Research on Schizophrenia and Depression (NARSAD) Independent Investigator Award (to A.M.M.), and data collection was supported by the Health Foundation Clinician Scientist Fellowship. BIG: This work makes use of the BIG (Brain Imaging Genetics) database, first established in Nijmegen, the Netherlands, in 2007. This resource is now part of the Cognomics Initiative (http://www.cognomics.nl/), a joint initiative by researchers of the Donders Centre for Cognitive Neuroimaging, the Human Genetics and Cognitive Neuroscience departments of the Radboud University Medical Centre and the Max Planck Institute for Psycholinguistics in Nijmegen. The Cognomics Initiative is supported by the participating departments and centers and by external grants from the Biobanking and Biomolecular Resources Research Infrastructure (Netherlands) (BBMRI-NL), the Hersenstichting Nederland and the Netherlands Organization for Scientific Research (NWO). We wish to thank all persons who kindly participated in the BIG research. The research leading to these results also receives funding from the European Community's Seventh Framework Programme (FP7/2007– 2013) under grant agreements 602450 (IMAGEMEND) and 602805 (Aggressotype) and from ERC-2010-AdG 268800-NEUROSCHEMA. B.F. is supported by a Vici grant from the NWO (grant 016.130.669). Brain Genomics Superstruct Project (GSP): Data were provided [in part] by the Brain GSP of Harvard University and the Massachusetts General Hospital, with support from the Center for Brain Science Neuroinformatics Research Group, the Athinoula A. Martinos Center for Biomedical Imaging and the Center for Human Genetic Research. Twenty individual investigators at Harvard and Massachusetts General Hospital generously contributed data to GSP. GIG: The GIG (Genomic Imaging Göttingen) sample was established at the Center for Translational Research in Systems Neuroscience and Psychiatry at Göttingen University. We thank M. Keil, E. Diekhof, T. Melcher and I. Henseler for assistance in MRI data acquisition, and E. Binder and H. Mohr for help with genotyping. We are grateful to all persons who kindly participated in the GIG study. IMAGEN: IMAGEN was supported by the European Union-funded FP6 Integrated Project IMAGEN (Reinforcement-Related Behaviour in Normal Brain Function and Psychopathology) (LSHM-CT- 2007-037286), the FP7 projects IMAGEMEND (602450) and MATRICS (603016), and the Innovative Medicine Initiative Project EU-AIMS (115300-2), the Medical Research Council programme grant “Developmental pathways into adolescent substance abuse” (93558), as well as the NIHR Biomedical Research Center “Mental Health”. Further support was provided by the Swedish Research Council (FORMAS) and the BMBF (eMED SysAlc 01ZX1311A; Forschungsnetz AERIAL; 1EV0711). MooDS: The establishment of the MooDS sample was funded by the BMBF through the Integrated Genome Research Network MooDS (grant 01GS08144 to M.M.N. and S.C., grant 01GS08147 to M.R. and A.M.-L. and grant 01GS08148 to A. Heinz), under the auspices of the National Genome Research Network plus (NGFNplus), and through the Integrated Network IntegraMent, under the auspices of the e:Med Programme (grant 01ZX1314A to M.M.N., grant 01ZX1314C to H. Walter, grant 01ZX1314G to M.R.). MPIP: The MPIP Munich Morphometry Sample comprises images acquired as part of the Munich Antidepressant Response Signature Study and the Recurrent Unipolar Depression (RUD) Case-Control study performed at the MPIP, and control subject data acquired at the Ludwig Maximilians University, Munich, Department of Psychiatry. We wish to acknowledge A. Olynyik and radiographers R. Schirmer, E. Schreiter, R. Borschke and I. Eidner for image acquisition and data preparation. We thank D.P. Auer for local study management in the initial phase of the RUD study. We are grateful to GlaxoSmithKline for providing the genotypes of the RUD case-control sample. We thank the staff of the Center of Applied Genotyping for generating the genotypes of the MARS cohort. The study is supported by a grant of the Exzellenz-Stiftung of the Max Planck Society. This work has also been funded by the BMBF in the framework of the National Genome Research Network (NGFN), FKZ 01GS0481. NCNG: Sample collection was supported by grants from the Bergen Research Foundation and the University of Bergen, the Dr Einar Martens Fund, the K.G. Jebsen Foundation and the Research Council of Norway, to S.L.H., V.M.S. and T.E. NESDA: Funding was obtained from the Netherlands Organization for Scientific Research (Geestkracht program grant 10-000-1002), the Center for Medical Systems Biology (CSMB, NWO Genomics), Biobanking and Biomolecular Resources Research Infrastructure (BBMRI-NL), VU University's Institutes for Health and Care Research (EMGO+) and Neuroscience Campus Amsterdam, University Medical Center Groningen, Leiden University Medical Center and NIH (R01D0042157-01A, MH081802, Grand Opportunity grants 1RC2 MH089951 and 1RC2 MH089995). Part of the genotyping and analyses were funded by the Genetic Association Information Network (GAIN) of the Foundation for the National Institutes of Health. Computing was supported by BiG Grid, the Dutch e-Science Grid, which is financially supported by NWO. NeuroIMAGE: NeuroIMAGE was supported by NIH grant R01MH62873 (to S.V. Faraone), NWO large investment grant 1750102007010 (to J. Buitelaar) and grants from Radboud University Nijmegen Medical Center, University Medical Center Groningen and Accare, and VU University Amsterdam. The research leading to these results also receives funding from the European Community's Seventh Framework Programme (FP7/2007– 2013) under grant agreements 602450 (IMAGEMEND), 278948 (TACTICS) and 602805 (Aggressotype). NTR-Adults and Brainscale: We would like to thank all twin participants from the Netherlands Twin Register. The NTR-Adult and Brainscale studies were supported by the NWO (MW904-61-193 (E.J.C.d.G. and D.I.B.), MaGW-nr 400-07-080 (D.v.t.E.), MagW 480-04-004 (D.B), (51.02.060 (H.H.), 668.772 (D.B. and H.H.); NWO/SPI 56-464-14192 (D.B.), the European Research Council (ERC-230374) (D.B.), High Potential Grant Utrecht University (H.H.), NWO Brain and Cognition 433-09-220 (H.H.) and the Neuroscience Campus Amsterdam. Older Australian Twins Study (OATS): We would like to acknowledge and thank the OATS participants, their supporters and respective research teams. OATS is supported by Australian National Health and Medical Research Council (NHMRC)/Australian Research Council Strategic Award 401162 and NHMRC project grant 1045325 to P.S.S. and colleagues. OATS was facilitated through access to the Australian Twin Registry, a national research resource supported by NHMRC enabling grant 310667, administered by the University of Melbourne. DNA was extracted by Genetic Repositories Australia, an Enabling Facility supported by NHMRC grant 401184. OATS genotyping was partly funded by a Commonwealth Scientific and Industrial Research Organisation Flagship Collaboration Fund grant. H.B. is supported by the Australian Government funded Dementia Collaborative Research Centre, UNSW. N.J.A. was supported by NHMRC project grant 525453 and K.A.M. is supported by an Alzheimer's Australia Dementia Research Foundation postdoctoral fellowship and NHMRC capacity building grant 568940. QTIM: D.P.H., N.J., C.R.K.C. and P.M.T. are supported, in part, by NIH grants R01 NS080655, R01AG040060, R01 EB008432, R01 MH097268, U01 AG024904, R01 MH085667, R01 MH089722, P41 EB015922 and R01 MH094343. R.K.W. is supported by National Science Foundation (BCS-1229450). J.L.S. was supported by the NIMH (K99MH102357) and Autism Speaks. S.E.M. and G.I.d.Z. are supported by Future Fellowships (FT110100548, FT0991634) from the Australian Research Council, and G.W.M. is supported by an NHMRC fellowship (619667). The QTIM study is supported by grants from the NIH (R01 HD050735) and the NHMRC (389875, 486682, 1009064). We thank the twins and siblings for their participation, M. Grace and A. Eldridge for twin recruitment, A. Al Najjar and other radiographers for scanning, K. McAloney and D. Park for research support, and A. Henders and staff for DNA sample processing and preparation. SHIP: The Study of Health in Pomerania (SHIP) is supported by the BMBF (grants 01ZZ9603, 01ZZ0103 and 01ZZ0403) and the DFG (GR 1912/5-1). Genome-wide data and MRI scans were supported by a joint grant from Siemens Healthcare, Erlangen, Germany, and the Federal State of Mecklenburg–West Pomerania. SHIP-TREND-0: This cohort is part of the Community Medicine Research (CMR) net of the University of Greifswald, which is funded by the BMBF and the German Ministry of Cultural Affairs, as well as by the Social Ministry of the Federal State of Mecklenburg–West Pomerania. CMR encompasses several research projects that share data from SHIP. MRI scans were supported by a joint grant from Siemens Healthcare, Erlangen, Germany, and the Federal State of Mecklenburg–West Pomerania. The SHIP authors are grateful to M. Stanke for the opportunity to use his server cluster for SNP imputation as well as to H. Prokisch and T. Meitinger (HelmholtzZentrum München) for genotyping the SHIP-TREND cohort, which was supported by the BMBF (grant 03ZIK012). We thank all staff members and participants of the SHIP studies, as well as all of the genotyping staff for generating the SHIP SNP data set. D.J. is supported by a scholarship from the Gerhard-Domagk Programme of the University Medicine Greifswald. Sydney Memory and Ageing Study (Sydney MAS): We would like to thank the Sydney MAS participants, their supporters and respective research teams. Sydney MAS was supported by NHMRC program grants 350833 and 568969 to P.S.S., H.B. and G. Andrews. DNA was extracted by Genetic Repositories Australia, an Enabling Facility supported by NHMRC grant 401184. H.B. is supported by the Australian Government funded Dementia Collaborative Research Centre, UNSW. N.J.A. was supported by NHMRC project grant 525453 and K.A.M. is supported by an Alzheimer's Australia Dementia Research Foundation postdoctoral fellowship. Both S. Reppermund and K.A.M. are supported by NHMRC capacity building grant 568940. Data used in preparing this article were obtained from the ADNI database (http://adni.loni.usc.edu/). Many investigators in ADNI contributed to the design and implementation of ADNI and/or provided data but did not participate in analysis or writing of this report. A complete listing of ADNI investigators can be found at http://adni.loni.usc.edu/wp-content/uploads/how_to_apply/ADNI_Acknowledgement_List.pdf.

Author information

Author notes

    • Barbara Franke
    • , Jason L Stein
    • , Stephan Ripke
    • , Stephan Ripke
    • , Derrek P Hibar
    • , Jason L Stein
    • , Miguel E Renteria
    • , Alejandro Arias-Vasquez
    •  & Sylvane Desrivières

    These authors contributed equally to this work.

    • Benjamin M Neale
    • , Patrick F Sullivan
    • , Michael C O'Donovan
    • , Nicholas G Martin
    • , Margaret J Wright
    • , Gunter Schumann
    • , Barbara Franke
    • , Paul M Thompson
    • , Sarah E Medland
    • , Michael C O'Donovan
    • , Paul M Thompson
    • , Benjamin M Neale
    • , Sarah E Medland
    •  & Patrick F Sullivan

    These authors jointly directed this work.

Affiliations

  1. Department of Human Genetics, Radboud University Medical Center, Nijmegen, the Netherlands.

    • Barbara Franke
    • , Kimm J E van Hulzen
    • , Alejandro Arias-Vasquez
    • , Janita Bralten
    • , Martine Hoogman
    • , Marieke Klein
    • , Marjolein M J van Donkelaar
    • , Marina M H Hakobjan
    • , Angelien J G A M Heister
    • , Remco R R Makkinje
    • , Marlies A M Naber
    • , Saskia S L van der Marel
    • , Jeanette C Mostert
    • , Han G Brunner
    •  & Hans van Bokhoven
  2. Department of Psychiatry, Radboud University Medical Center, Nijmegen, the Netherlands.

    • Barbara Franke
    •  & Alejandro Arias-Vasquez
  3. Donders Institute for Brain, Cognition and Behaviour, Raboud University, Nijmegen, the Netherlands.

    • Barbara Franke
    • , Kimm J E van Hulzen
    • , Alejandro Arias-Vasquez
    • , Janita Bralten
    • , Martine Hoogman
    • , Marieke Klein
    • , Marjolein M J van Donkelaar
    • , Marcel P Zwiers
    • , Marina M H Hakobjan
    • , Angelien J G A M Heister
    • , Remco R R Makkinje
    • , Marlies A M Naber
    • , Saskia S L van der Marel
    • , Jeanette C Mostert
    • , Han G Brunner
    • , Jan K Buitelaar
    • , Hans van Bokhoven
    • , Guillén Fernández
    • , Simon E Fisher
    •  & Clyde Francks
  4. Imaging Genetics Center, Mark and Mary Stevens Neuroimaging & Informatics Institute, Keck School of Medicine of the University of Southern California, Marina del Rey, California, USA.

    • Jason L Stein
    • , Derrek P Hibar
    •  & Paul M Thompson
  5. Neurogenetics Program, Department of Neurology, UCLA School of Medicine, Los Angeles, California, USA.

    • Jason L Stein
  6. Analytic and Translational Genetics Unit, Massachusetts General Hospital, Boston, Massachusetts, USA.

    • Stephan Ripke
    • , Verneri Anttila
    • , Benjamin M Neale
    • , Kai-How Farh
    • , Brendan Bulik-Sullivan
    • , Hailiang Huang
    • , Menachem Fromer
    • , Jacqueline I Goldstein
    • , Mark J Daly
    •  & Raymond K Walters
  7. Stanley Center for Psychiatric Research, Broad Institute of MIT and Harvard, Cambridge, Massachusetts, USA.

    • Stephan Ripke
    • , Verneri Anttila
    • , Jordan W Smoller
    • , Phil Lee
    • , Benjamin M Neale
    • , Brendan Bulik-Sullivan
    • , Richard A Belliveau Jr
    • , Sarah E Bergen
    • , Elizabeth Bevilacqua
    • , Kimberley D Chambert
    • , Menachem Fromer
    • , Giulio Genovese
    • , Colm O'Dushlaine
    • , Edward M Scolnick
    • , Steven A McCarroll
    • , Jennifer L Moran
    • , Aarno Palotie
    • , Tracey L Petryshen
    •  & Mark J Daly
  8. Department of Psychiatry and Psychotherapy, Charité Universitätsmedizin Berlin, Campus Charité Mitte, Berlin, Germany.

    • Stephan Ripke
    • , Susanne Erk
    • , Andreas Heinz
    • , Sebastian Mohnke
    • , Nina Romanczuk-Seiferth
    •  & Henrik Walter
  9. Department of Cognitive Neuroscience, Radboud University Medical Center, Nijmegen, the Netherlands.

    • Alejandro Arias-Vasquez
    • , Janita Bralten
    • , Jan K Buitelaar
    •  & Guillén Fernández
  10. Psychiatric and Neurodevelopmental Genetics Unit, Massachusetts General Hospital, Boston, Massachusetts, USA.

    • Jordan W Smoller
    • , Phil Lee
    • , Benjamin M Neale
    • , Menachem Fromer
    •  & Aarno Palotie
  11. Department of Psychiatry, Harvard Medical School, Boston, Massachusetts, USA.

    • Jordan W Smoller
    • , Phil Lee
    • , Lynn E DeLisi
    • , Robert W McCarley
    • , Raquelle I Mesholam-Gately
    • , Larry J Seidman
    •  & Tracey L Petryshen
  12. FMRIB Centre, University of Oxford, Oxford, UK.

    • Thomas E Nichols
  13. Department of Statistics & Warwick Manufacturing Group, University of Warwick, Coventry, UK.

    • Thomas E Nichols
  14. Departments of Psychiatry & Human Genetics, Virginia Commonwealth University, Richmond, Virginia, USA.

    • Michael C Neale
  15. Division of Psychiatry, Royal Edinburgh Hospital, Centre for Cognitive Ageing and Cognitive Epidemiology, University of Edinburgh, Edinburgh, UK.

    • Andrew M McIntosh
    • , Martina Papmeyer
    • , Emma Sprooten
    • , Stephen M Lawrie
    •  & Jessika E Sussmann
  16. Intramural Research Program, National Institutes of Health, US Department of Health & Human Services, Bethesda, Maryland, USA.

    • Francis J McMahon
    •  & Yin Yao
  17. Central Institute of Mental Health, Medical Faculty Mannheim, University Heidelberg, Mannheim, Germany.

    • Andreas Meyer-Lindenberg
    • , Emanuel Schwarz
    •  & Oliver Grimm
  18. Department of Biomedicine, Aarhus University, Aarhus, Denmark.

    • Manuel Mattheisen
  19. The Lundbeck Foundation Initiative for Integrative Psychiatric Research, iPSYCH, Aarhus and Copenhagen, Denmark.

    • Esben Agerbo
    • , Ditte Demontis
    • , Thomas Hansen
    • , Manuel Mattheisen
    • , Ole Mors
    • , Line Olsen
    • , Henrik B Rasmussen
    • , Anders D Børglum
    • , Preben B Mortensen
    •  & Thomas Werge
  20. Center for integrated Sequencing, iSEQ, Aarhus University, Aarhus, Denmark.

    • Manuel Mattheisen
  21. NORMENT - KG Jebsen Centre, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.

    • Ole A Andreassen
    • , Andrew A Brown
    • , Lavinia Athanasiu
    • , Cecilie B Hartberg
    •  & Unn Haukvik
  22. Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.

    • Ole A Andreassen
    •  & Ingrid Melle
  23. Center for Translational Research in Systems Neuroscience and Psychiatry, Department of Psychiatry and Psychotherapy, University Medical Center, Goettingen, Germany.

    • Oliver Gruber
    • , Bernd Kraemer
    •  & Maria Keil
  24. Centre for Healthy Brain Ageing, School of Psychiatry, University of New South Wales (UNSW), Sydney, New South Wales, Australia.

    • Perminder S Sachdev
    • , Karen A Mather
    • , Anbupalam Thalamuthu
    • , Nicola J Armstrong
    • , Amelia A Assareh
    • , Henry Brodaty
    • , Simone Reppermund
    •  & Wei Wen
  25. Neuropsychiatric Institute, Prince of Wales Hospital, Sydney, New South Wales, Australia.

    • Perminder S Sachdev
  26. Department of Psychiatry, University Hospital Marqués de Valdecilla, School of Medicine, University of Cantabria-IDIVAL, Santander, Spain.

    • Roberto Roiz-Santiañez
  27. Cibersam (Centro Investigación Biomédica en Red Salud Mental), Madrid, Spain.

    • Roberto Roiz-Santiañez
    •  & Rocio Perez-Iglesias
  28. Center for Neuroimaging, Radiology and Imaging Sciences, Indiana University School of Medicine, Indianapolis, Indiana, USA.

    • Andrew J Saykin
    • , Sungeun Kim
    • , Kwangsik Nho
    • , Shannon L Risacher
    •  & Li Shen
  29. Indiana Alzheimer Disease Center, Indiana University School of Medicine, Indianapolis, Indiana, USA.

    • Andrew J Saykin
    • , Sungeun Kim
    • , Kwangsik Nho
    • , Shannon L Risacher
    • , Li Shen
    •  & Tatiana M Foroud
  30. Medical and Molecular Genetics, Indiana University School of Medicine, Indianapolis, Indiana, USA.

    • Andrew J Saykin
    •  & Tatiana M Foroud
  31. Department of Child and Adolescent Psychiatry, Faculty of Medicine and University Hospital, Technische Universität Dresden, Dresden, Germany.

    • Stefan Ehrlich
    • , Johanna Hass
    •  & Esther Walton
  32. Georgia State University, Atlanta, Georgia, USA.

    • Jessica A Turner
  33. Mind Research Network, Albuquerque, New Mexico, USA.

    • Jessica A Turner
  34. QIMR Berghofer Medical Research Institute, Brisbane, Queensland, Australia.

    • Yvonne Y W Ho
    • , Nicholas G Martin
    • , Margaret J Wright
    • , Miguel E Renteria
    • , Gabriel Cuellar-Partida
    • , Lachlan T Strike
    • , Narelle K Hansell
    • , Grant W Montgomery
    •  & Sarah E Medland
  35. School of Psychology, University of Queensland, Brisbane, Queensland, Australia.

    • Margaret J Wright
    • , Lachlan T Strike
    •  & Greig I de Zubicaray
  36. MRC Centre for Neuropsychiatric Genetics and Genomics, Institute of Psychological Medicine and Clinical Neurosciences, School of Medicine, Cardiff University, Cardiff, UK.

    • James T R Walters
    • , Peter A Holmans
    • , Noa Carrera
    • , Nick Craddock
    • , Valentina Escott-Price
    • , Lyudmila Georgieva
    • , Marian L Hamshere
    • , David Kavanagh
    • , Sophie E Legge
    • , Andrew J Pocklington
    • , Alexander L Richards
    • , Douglas M Ruderfer
    • , Nigel M Williams
    • , George Kirov
    • , Michael J Owen
    •  & Michael C O'Donovan
  37. National Centre for Mental Health, Cardiff University, Cardiff, UK.

    • Peter A Holmans
    • , Nick Craddock
    • , Alexander L Richards
    • , Michael J Owen
    •  & Michael C O'Donovan
  38. Medical and Population Genetics Program, Broad Institute of MIT and Harvard, Cambridge, Massachusetts, USA.

    • Benjamin M Neale
    • , Hailiang Huang
    • , Tune H Pers
    • , Jacqueline I Goldstein
    • , Joel N Hirschhorn
    • , Alkes Price
    • , Eli A Stahl
    • , Tõnu Esko
    • , Mark J Daly
    •  & Raymond K Walters
  39. Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden.

    • Sarah E Bergen
    • , Anna K Kähler
    • , Patrik K E Magnusson
    • , Christina M Hultman
    •  & Patrick F Sullivan
  40. Department of Genetics, University of North Carolina, Chapel Hill, North Carolina, USA.

    • James J Crowley
    • , Martilias S Farrell
    • , Paola Giusti-Rodríguez
    • , Yunjung Kim
    • , Jin P Szatkiewicz
    • , Stephanie Williams
    •  & Patrick F Sullivan
  41. Department of Psychiatry, University of North Carolina, Chapel Hill, North Carolina, USA.

    • Diana O Perkins
    •  & Patrick F Sullivan
  42. Neuropsychiatric Genetics Research Group, Department of Psychiatry, Trinity College Dublin, Dublin, Ireland.

    • Aiden Corvin
    • , Paul Cormican
    • , Gary Donohoe
    • , Derek W Morris
    • , Michael Gill
    • , Emma J Rose
    •  & Margaret Needham
  43. Eli Lilly and Company Limited, Erl Wood Manor, Sunninghill Road, Windlesham, Surrey, UK.

    • David A Collier
  44. Social, Genetic and Developmental Psychiatry Centre, Institute of Psychiatry, King's College London, London, UK.

    • David A Collier
  45. NORMENT, KG Jebsen Centre for Psychosis Research, Institute of Clinical Medicine, University of Oslo, Oslo, Norway.

    • Ingrid Agartz
    • , Srdjan Djurovic
    • , Morten Mattingsdal
    •  & Erik G Jönsson
  46. Department of Psychiatry, Diakonhjemmet Hospital, Oslo, Norway.

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

    • Ingrid Agartz
    • , Erik Söderman
    •  & Erik G Jönsson
  48. State Mental Hospital, Haar, Germany.

    • Margot Albus
  49. Department of Psychiatry and Behavioral Sciences, Stanford University, Stanford, California, USA.

    • Madeline Alexander
    • , Claudine Laurent
    •  & Douglas F Levinson
  50. Department of Psychiatry and Behavioral Sciences, Emory University, Atlanta, Georgia, USA.

    • Farooq Amin
  51. Department of Psychiatry and Behavioral Sciences, Atlanta Veterans Affairs Medical Center, Atlanta, Georgia, USA.

    • Farooq Amin
  52. Virginia Institute for Psychiatric and Behavioral Genetics, Virginia Commonwealth University, Richmond, Virginia, USA.

    • Silviu A Bacanu
    • , Tim B Bigdeli
    • , Mark A Reimers
    • , Bradley T Webb
    • , Aaron R Wolen
    • , Brandon K Wormley
    • , Kenneth S Kendler
    •  & Brien P Riley
  53. Clinical Neuroscience, Max Planck Institute of Experimental Medicine, Göttingen, Germany.

    • Martin Begemann
    • , Christian Hammer
    • , Sergi Papiol
    •  & Hannelore Ehrenreich
  54. Department of Medical Genetics, University of Pécs, Pécs, Hungary.

    • Judit Bene
    •  & Bela Melegh
  55. Szentagothai Research Center, University of Pécs, Pécs, Hungary.

    • Judit Bene
    •  & Bela Melegh
  56. Department of Psychiatry, University of Iowa Carver College of Medicine, Iowa City, Iowa, USA.

    • Donald W Black
  57. University Medical Center Groningen, Department of Psychiatry, University of Groningen, the Netherlands.

    • Richard Bruggeman
  58. School of Nursing, Louisiana State University Health Sciences Center, New Orleans, Louisiana, USA.

    • Nancy G Buccola
  59. Center for Brain Science, Harvard University, Cambridge, Massachusetts, USA.

    • Randy L Buckner
  60. Department of Psychiatry, Massachusetts General Hospital, Boston, Massachusetts, USA.

    • Randy L Buckner
    • , Joshua L Roffman
    • , Avram J Holmes
    •  & Randy L Gollub
  61. Athinoula A Martinos Center, Massachusetts General Hospital, Boston, Massachusetts, USA.

    • Randy L Buckner
    •  & Joshua L Roffman
  62. Department of Psychiatry, University of California at San Francisco, San Francisco, California, USA.

    • William F Byerley
  63. University Medical Center Utrecht, Department of Psychiatry, Rudolf Magnus Institute of Neuroscience, Utrecht, the Netherlands.

    • Wiepke Cahn
    • , René S Kahn
    • , Eric Strengman
    •  & Roel A Ophoff
  64. Department of Psychiatry, Icahn School of Medicine at Mount Sinai, New York, New York, USA.

    • Guiqing Cai
    • , Kenneth L Davis
    • , Elodie Drapeau
    • , Joseph I Friedman
    • , Vahram Haroutunian
    • , Elena Parkhomenko
    • , Abraham Reichenberg
    • , Jeremy M Silverman
    •  & Joseph D Buxbaum
  65. Department of Human Genetics, Icahn School of Medicine at Mount Sinai, New York, New York, USA.

    • Guiqing Cai
    •  & Joseph D Buxbaum
  66. Schizophrenia Research Institute, Sydney, New South Wales, Australia.

    • Murray J Cairns
    • , Vaughan J Carr
    • , Stanley V Catts
    • , Frans A Henskens
    • , Carmel M Loughland
    • , Patricia T Michie
    • , Christos Pantelis
    • , Ulrich Schall
    • , Rodney J Scott
    • , Paul A Tooney
    • , Jing Qin Wu
    •  & Assen V Jablensky
  67. Priority Centre for Translational Neuroscience and Mental Health, University of Newcastle, Newcastle, New South Wales, Australia.

    • Murray J Cairns
    • , Brian J Kelly
    • , Carmel M Loughland
    •  & Ulrich Schall
  68. School of Biomedical Sciences and Pharmacy, University of Newcastle, Callaghan, New South Wales, Australia.

    • Murray J Cairns
    • , Rodney J Scott
    • , Paul A Tooney
    •  & Jing Qin Wu
  69. Centre Hospitalier du Rouvray and INSERM U1079 Faculty of Medicine, Rouen, France.

    • Dominique Campion
  70. Department of Human Genetics, David Geffen School of Medicine, University of California, Los Angeles, California, USA.

    • Rita M Cantor
    •  & Roel A Ophoff
  71. School of Psychiatry, University of New South Wales, Sydney, New South Wales, Australia.

    • Vaughan J Carr
  72. Royal Brisbane and Women's Hospital, University of Queensland, Brisbane, Queensland, Australia.

    • Stanley V Catts
  73. Institute of Psychology, Chinese Academy of Science, Beijing, China.

    • Raymond C K Chan
  74. State Key Laboratory for Brain and Cognitive Sciences, Li Ka Shing Faculty of Medicine, The University of Hong Kong, Hong Kong, China.

    • Eric Y H Chen
    • , Miaoxin Li
    •  & Pak C Sham
  75. Department of Psychiatry, Li Ka Shing Faculty of Medicine, The University of Hong Kong, Hong Kong, China.

    • Eric Y H Chen
    • , Ronald Y L Chen
    • , Miaoxin Li
    • , Hon-Cheong So
    • , Emily H M Wong
    •  & Pak C Sham
  76. Department of Computer Science, University of North Carolina, Chapel Hill, North Carolina, USA.

    • Wei Cheng
  77. Castle Peak Hospital, Hong Kong, China.

    • Eric F C Cheung
  78. Institute of Mental Health, Singapore, Singapore.

    • Siow Ann Chong
    • , Jimmy Lee Chee Keong
    • , Kang Sim
    •  & Mythily Subramaniam
  79. Department of Psychiatry, Washington University, St. Louis, Missouri, USA.

    • C Robert Cloninger
    •  & Dragan M Svrakic
  80. Department of Child and Adolescent Psychiatry, Assistance Publique Hospitaux de Paris, Pierre and Marie Curie Faculty of Medicine and Institute for Intelligent Systems and Robotics, Paris, France.

    • David Cohen
  81. Blue Note Biosciences, Princeton, New Jersey, USA.

    • Nadine Cohen
  82. Department of Psychiatry, University Hospital Marqués de Valdecilla, School of Medicine, University of Cantabria-IDIVAL, Santander, Spain.

    • Benedicto Crespo-Facorro
  83. Centro Investigacion Biomedica en Red Salud Mental, Madrid, Spain.

    • Benedicto Crespo-Facorro
  84. Department of Psychological Medicine, Queen Mary University of London, London, UK.

    • David Curtis
  85. Molecular Psychiatry Laboratory, Division of Psychiatry, University College London, London, UK.

    • David Curtis
    • , Jonathan Pimm
    • , Hugh Gurling
    •  & Andrew McQuillin
  86. Sheba Medical Center, Tel Hashomer, Israel.

    • Michael Davidson
    •  & Mark Weiser
  87. Institute of Human Genetics, University of Bonn, Bonn, Germany.

    • Franziska Degenhardt
    • , Stefan L Herms
    • , Per Hoffmann
    • , Andrea Hofman
    • , Sven Cichon
    •  & Markus M Nöthen
  88. Department of Genomics, Life and Brain Center, Bonn, Germany.

    • Franziska Degenhardt
    • , Stefan L Herms
    • , Per Hoffmann
    • , Andrea Hofman
    • , Sven Cichon
    •  & Markus M Nöthen
  89. Applied Molecular Genomics Unit, VIB Department of Molecular Genetics, University of Antwerp, Antwerp, Belgium.

    • Jurgen Del Favero
  90. First Department of Psychiatry, University of Athens Medical School, Athens, Greece.

    • Dimitris Dikeos
    •  & George N Papadimitriou
  91. Department of Psychiatry, University College Cork, County Cork, Ireland.

    • Timothy Dinan
  92. Department of Medical Genetics, Oslo University Hospital, Oslo, Norway.

    • Srdjan Djurovic
  93. Cognitive Genetics and Therapy Group, School of Psychology and Discipline of Biochemistry, National University of Ireland Galway, County Galway, Ireland.

    • Gary Donohoe
    •  & Derek W Morris
  94. Department of Psychiatry and Behavioral Sciences, NorthShore University HealthSystem, Evanston, Illinois, USA.

    • Jubao Duan
    • , Alan R Sanders
    •  & Pablo V Gejman
  95. Department of Psychiatry and Behavioral Neuroscience, University of Chicago, Chicago, Illinois, USA.

    • Jubao Duan
  96. Department of Non-Communicable Disease Epidemiology, London School of Hygiene and Tropical Medicine, London, UK.

    • Frank Dudbridge
  97. Department of Psychiatry, University of Regensburg, Regensburg, Germany.

    • Peter Eichhammer
  98. Folkhälsan Research Center, Helsinki, Finland, Biomedicum Helsinki, Helsinki, Finland.

    • Johan Eriksson
  99. National Institute for Health and Welfare, Helsinki, Finland.

    • Johan Eriksson
    •  & Veikko Salomaa
  100. Department of General Practice, Helsinki University Central Hospital, University of Helsinki, Helsinki, Finland.

    • Johan Eriksson
  101. Translational Technologies and Bioinformatics, Pharma Research and Early Development, F. Hoffman-La Roche, Basel, Switzerland.

    • Laurent Essioux
  102. Department of Medicine, McGill University, Montreal, Quebec, Canada.

    • Ayman H Fanous
  103. Department of Psychiatry, Georgetown University School of Medicine, Washington, DC, USA.

    • Ayman H Fanous
  104. Department of Psychiatry, Virginia Commonwealth University School of Medicine, Richmond, Virginia, USA.

    • Ayman H Fanous
  105. Department of Psychiatry, Keck School of Medicine of the University of Southern California, Los Angeles, California, USA.

    • Ayman H Fanous
  106. Department of Genetic Epidemiology in Psychiatry, Central Institute of Mental Health, Medical Faculty Mannheim, University of Heidelberg, Heidelberg, Mannheim, Germany.

    • Josef Frank
    • , Thomas G Schulze
    • , Stephanie H Witt
    •  & Marcella Rietschel
  107. Department of Genetics, University of Groningen, University Medical Centre Groningen, Groningen, the Netherlands.

    • Lude Franke
    •  & Juha Karjalainen
  108. Department of Psychiatry, University of Colorado Denver, Aurora, Colorado, USA.

    • Robert Freedman
    •  & Ann Olincy
  109. Center for Neurobehavioral Genetics, Semel Institute for Neuroscience and Human Behavior, University of California, Los Angeles, California, USA.

    • Nelson B Freimer
    •  & Roel A Ophoff
  110. Departments of Psychiatry and Human Genetics, University of Chicago, Chicago, Illinois, USA.

    • Elliot S Gershon
  111. Department of Psychiatry, University of Halle, Halle, Germany.

    • Ina Giegling
    • , Annette M Hartmann
    • , Bettina Konte
    •  & Dan Rujescu
  112. Department of Psychiatry, University of Munich, Munich, Germany.

    • Ina Giegling
    •  & Dan Rujescu
  113. Departments of Psychiatry and Human and Molecular Genetics, INSERM, Institut de Myologie, Hôpital de la Pitiè-Salpêtrière, Paris, France.

    • Stephanie Godard
  114. Neuroscience Therapeutic Area, Janssen Research and Development, Raritan, New Jersey, USA.

    • Srihari Gopal
    • , Dai Wang
    •  & Qingqin S Li
  115. Queensland Brain Institute, The University of Queensland, Brisbane, Queensland, Australia.

    • Jacob Gratten
    • , S Hong Lee
    • , Naomi R Wray
    •  & Peter M Visscher
  116. Academic Medical Centre University of Amsterdam, Department of Psychiatry, Amsterdam, the Netherlands.

    • Lieuwe de Haan
    •  & Carin J Meijer
  117. Illumina, La Jolla, California, California, USA.

    • Mark Hansen
  118. JJ Peters Veterans Affairs Medical Center, Bronx, New York, New York, USA.

    • Vahram Haroutunian
  119. Friedman Brain Institute, Icahn School of Medicine at Mount Sinai, New York, New York, USA.

    • Vahram Haroutunian
    • , Joseph D Buxbaum
    •  & Pamela Sklar
  120. School of Electrical Engineering and Computer Science, University of Newcastle, Newcastle, New South Wales, Australia.

    • Frans A Henskens
  121. Priority Research Centre for Health Behaviour, University of Newcastle, Newcastle, New South Wales, Australia.

    • Frans A Henskens
  122. Division of Medical Genetics, Department of Biomedicine, University of Basel, Basel, Switzerland.

    • Stefan L Herms
    • , Per Hoffmann
    •  & Sven Cichon
  123. Section of Neonatal Screening and Hormones, Department of Clinical Biochemistry, Immunology and Genetics, Statens Serum Institut, Copenhagen, Denmark.

    • Mads V Hollegaard
    •  & David M Hougaard
  124. Department of Psychiatry, Fujita Health University School of Medicine, Toyoake, Aichi, Japan.

    • Masashi Ikeda
    •  & Nakao Iwata
  125. Regional Centre for Clinical Research in Psychosis, Department of Psychiatry, Stavanger University Hospital, Stavanger, Norway.

    • Inge Joa
  126. Rheumatology Research Group, Vall d'Hebron Research Institute, Barcelona, Spain.

    • Antonio Julià
    •  & Sara Marsal
  127. Centre for Medical Research, The University of Western Australia, Perth, Western Australia, Australia.

    • Luba Kalaydjieva
  128. The Perkins Institute for Medical Research, The University of Western Australia, Perth, Western Australia, Australia.

    • Luba Kalaydjieva
  129. Department of Medical Genetics, Medical University, Sofia, Bulgaria.

    • Sena Karachanak-Yankova
  130. Department of Psychology, University of Colorado Boulder, Boulder, Colorado, USA.

    • Matthew C Keller
  131. Campbell Family Mental Health Research Institute, Centre for Addiction and Mental Health, Toronto, Ontario, Canada.

    • James L Kennedy
    • , Clement C Zai
    •  & Jo Knight
  132. Department of Psychiatry, University of Toronto, Toronto, Ontario, Canada.

    • James L Kennedy
    • , Clement C Zai
    •  & Jo Knight
  133. Institute of Medical Science, University of Toronto, Toronto, Ontario, Canada.

    • James L Kennedy
    •  & Jo Knight
  134. Institute of Molecular Genetics, Russian Academy of Sciences, Moscow, Russia.

    • Andrey Khrunin
    • , Svetlana Limborska
    •  & Petr Slominsky
  135. Latvian Biomedical Research and Study Centre, Riga, Latvia.

    • Janis Klovins
    •  & Liene Nikitina-Zake
  136. Department of Psychiatry and Zilkha Neurogenetics Institute, Keck School of Medicine at University of Southern California, Los Angeles, California, USA.

    • James A Knowles
    • , Michele T Pato
    •  & Carlos N Pato
  137. Faculty of Medicine, Vilnius University, Vilnius, Lithuania.

    • Vaidutis Kucinskas
    •  & Zita Ausrele Kucinskiene
  138. Department of Biology and Medical Genetics, 2nd Faculty of Medicine and University Hospital Motol, Prague, Czech Republic.

    • Hana Kuzelova-Ptackova
  139. Department of Child and Adolescent Psychiatry, Pierre and Marie Curie Faculty of Medicine, Paris, France.

    • Claudine Laurent
  140. Duke–National University Singapore Graduate Medical School, Singapore, Singapore.

    • Jimmy Lee Chee Keong
  141. Department of Psychiatry, Hadassah-Hebrew University Medical Center, Jerusalem, Israel.

    • Bernard Lerer
  142. Centre for Genomic Sciences, The University of Hong Kong, Hong Kong, China.

    • Miaoxin Li
    •  & Pak C Sham
  143. Mental Health Centre and Psychiatric Laboratory, West China Hospital, Sichuan University, Chengdu, Sichuan, China.

    • Tao Li
  144. Department of Biostatistics, Johns Hopkins University Bloomberg School of Public Health, Baltimore, Maryland, USA.

    • Kung-Yee Liang
  145. Department of Psychiatry, Columbia University, New York, New York, USA.

    • Jeffrey Lieberman
    •  & T Scott Stroup
  146. Department of Mental Health and Substance Abuse Services, National Institute for Health and Welfare, Helsinki, Finland.

    • Jouko Lönnqvist
    •  & Jaana Suvisaari
  147. Department of Genetics and Pathology, International Hereditary Cancer Center, Pomeranian Medical University in Szczecin, Szczecin, Poland.

    • Jan Lubinski
  148. Department of Biology and Medical Genetics, 2nd Faculty of Medicine and University Hospital Motol, Prague, Czech Republic.

    • Milan Macek Jr
  149. Department of Mental Health, Bloomberg School of Public Health, Johns Hopkins University, Baltimore, Maryland, USA.

    • Brion S Maher
  150. Department of Psychiatry, University of Bonn, Bonn, Germany.

    • Wolfgang Maier
  151. Centre National de la Recherche Scientifique, Laboratoire de Génétique Moléculaire de la Neurotransmission et des Processus Neurodégénératifs, Hôpital de la Pitié Salpêtrière, Paris, France.

    • Jacques Mallet
  152. Research Unit, Sørlandet Hospital, Kristiansand, Norway.

    • Morten Mattingsdal
  153. Department of Psychiatry, National University of Ireland Galway, County Galway, Ireland.

    • Colm McDonald
  154. Department of Genetic Epidemiology in Psychiatry, Central Institute of Mental Health, Medical Faculty Mannheim, University of Heidelberg, Heidelberg, Mannheim, Germany.

    • Sandra Meier
    •  & Jana Strohmaier
  155. Estonian Genome Center, University of Tartu, Tartu, Estonia.

    • Andres Metspalu
    • , Lili Milani
    •  & Mari Nelis
  156. School of Psychology, University of Newcastle, Newcastle, New South Wales, Australia.

    • Patricia T Michie
  157. First Psychiatric Clinic, Medical University, Sofia, Bulgaria.

    • Vihra Milanova
  158. Eli Lilly and Company Limited, Windlesham, Surrey, UK.

    • Younes Mokrab
  159. Max Planck Institute of Psychiatry, Munich, Germany.

    • Bertram Müller-Myhsok
  160. Institute of Translational Medicine, University of Liverpool, Liverpool, UK.

    • Bertram Müller-Myhsok
  161. Munich Cluster for Systems Neurology (SyNergy), Munich, Germany.

    • Bertram Müller-Myhsok
  162. Department of Psychiatry, Royal College of Surgeons in Ireland, Dublin, Ireland.

    • Kieran C Murphy
  163. King's College London, London, UK.

    • Robin M Murray
    •  & John Powell
  164. Maastricht University Medical Centre, South Limburg Mental Health Research and Teaching Network, EURON, Maastricht, the Netherlands.

    • Inez Myin-Germeys
    •  & Jim Van Os
  165. Department of Psychiatry and Psychotherapy, Jena University Hospital, Jena, Germany.

    • Igor Nenadic
  166. Queensland Centre for Mental Health Research, University of Queensland, Brisbane, Queensland, Australia.

    • Deborah A Nertney
    •  & Bryan J Mowry
  167. Department of Psychiatry and Behavioral Sciences, Johns Hopkins University School of Medicine, Baltimore, Maryland, USA.

    • Gerald Nestadt
    •  & Ann E Pulver
  168. Department of Psychiatry, Trinity College Dublin, Dublin, Ireland.

    • Kristin K Nicodemus
  169. Eli Lilly and Company, Lilly Corporate Center, Indianapolis, Indiana, USA.

    • Laura Nisenbaum
  170. Department of Clinical Sciences, Psychiatry, Umeå University, Umeå, Sweden.

    • Annelie Nordin
    •  & Rolf Adolfsson
  171. DETECT Early Intervention Service for Psychosis, Blackrock, County Dublin, Ireland.

    • Eadbhard O'Callaghan
  172. Centre for Public Health, Institute of Clinical Sciences, Queen's University Belfast, Belfast, UK.

    • F Anthony O'Neill
  173. Lawrence Berkeley National Laboratory, University of California at Berkeley, Berkeley, California, USA.

    • Sang-Yun Oh
  174. Institute of Psychiatry, King's College London, London, UK.

    • Jim Van Os
    •  & Rocio Perez-Iglesias
  175. Melbourne Neuropsychiatry Centre, University of Melbourne & Melbourne Health, Melbourne, Victoria, Australia.

    • Christos Pantelis
  176. Public Health Genomics Unit, National Institute for Health and Welfare, Helsinki, Finland.

    • Tiina Paunio
    •  & Olli Pietiläinen
  177. Department of Psychiatry, University of Helsinki, Helsinki, Finland.

    • Tiina Paunio
  178. Institute for Molecular Medicine Finland, FIMM, University of Helsinki, Helsinki, Finland.

    • Olli Pietiläinen
  179. Division of Psychiatric Genomics, Department of Psychiatry, Icahn School of Medicine at Mount Sinai, New York, New York, USA.

    • Shaun M Purcell
    • , Panos Roussos
    •  & Pamela Sklar
  180. Department of Psychiatry, University of Oxford, Oxford, UK.

    • Digby Quested
    •  & Simon Lovestone
  181. Institute for Multiscale Biology, Icahn School of Medicine at Mount Sinai, New York, New York, USA.

    • Panos Roussos
    •  & Pamela Sklar
  182. Department of Psychiatry and Behavioral Neuroscience, University of Chicago, Chicago, Illinois, USA.

    • Alan R Sanders
    •  & Pablo V Gejman
  183. PharmaTherapeutics Clinical Research, Pfizer Worldwide Research and Development, Cambridge, Massachusetts, USA.

    • Christian R Schubert
    •  & Jens R Wendland
  184. Department of Psychiatry and Psychotherapy, University of Gottingen, Gottingen, Germany.

    • Thomas G Schulze
    •  & Knut Schnell
  185. Psychiatry and Psychotherapy Clinic, University of Erlangen, Erlangen, Germany.

    • Sibylle G Schwab
  186. Hunter New England Health Service, Newcastle, New South Wales, Australia.

    • Rodney J Scott
  187. Division of Cancer Epidemiology and Genetics, National Cancer Institute, Bethesda, Maryland, USA.

    • Jianxin Shi
  188. Research and Development, Bronx Veterans Affairs Medical Center, New York, New York, USA.

    • Jeremy M Silverman
  189. Wellcome Trust Centre for Human Genetics, Oxford, UK.

    • Chris C A Spencer
  190. Department of Clinical Neurology, Medical University of Vienna, Vienna, Austria.

    • Elisabeth Stogmann
    •  & Fritz Zimprich
  191. Lieber Institute for Brain Development, Baltimore, Maryland, USA.

    • Richard E Straub
    • , Daniel R Weinberger
    • , Qiang Chen
    • , Aaron L Goldman
    •  & Venkata S Mattay
  192. Department of Medical Genetics, University Medical Centre Utrecht, Utrecht, the Netherlands.

    • Eric Strengman
  193. Berkshire Healthcare NHS Foundation Trust, Bracknell, UK.

    • Srinivas Thirumalai
  194. Department of Medical Genetics, Medical University, Sofia, Bulgaria.

    • Draga Toncheva
  195. Priority Research Centre for Translational Neuroscience and Mental Health, University of Newcastle, Newcastle, New South Wales, Australia.

    • Paul A Tooney
  196. Department of Psychiatry, University of Oulu, Oulu, Finland.

    • Juha Veijola
  197. University Hospital of Oulu, Oulu, Finland.

    • Juha Veijola
  198. Molecular and Cellular Therapeutics, Royal College of Surgeons in Ireland, Dublin, Ireland.

    • John Waddington
  199. Health Research Board, Dublin, Ireland.

    • Dermot Walsh
  200. Mental Health Centre and Psychiatric Laboratory, West China Hospital, Sichuan University, Chendu, Sichuan, China.

    • Qiang Wang
  201. School of Psychiatry and Clinical Neurosciences, The University of Western Australia, Perth, Western Australia, Australia.

    • Dieter B Wildenauer
  202. Computational Sciences CoE, Pfizer Worldwide Research and Development, Cambridge, Massachusetts, USA.

    • Hualin Simon Xi
  203. Human Genetics, Genome Institute of Singapore, A*STAR, Singapore, Singapore.

    • Xuebin Zheng
    •  & Jianjun Liu
  204. Division of Psychiatry, University of Edinburgh, Edinburgh, UK.

    • Douglas H R Blackwood
  205. University College London, London, UK.

    • Elvira Bramon
  206. Department of Neuroscience, Icahn School of Medicine at Mount Sinai, New York, New York, USA.

    • Joseph D Buxbaum
  207. Institute of Neuroscience and Medicine (INM-1), Research Center Juelich, Juelich, Germany.

    • Sven Cichon
    •  & Thomas W Mühleisen
  208. Department of Genetics, The Hebrew University of Jerusalem, Jerusalem, Israel.

    • Ariel Darvasi
  209. Neuroscience Discovery and Translational Area, Pharma Research and Early Development, F. Hoffman-La Roche, Basel, Switzerland.

    • Enrico Domenici
  210. School of Psychiatry and Clinical Neurosciences, The University of Western Australia, Perth, Western Australia, Australia.

    • Assen V Jablensky
  211. Centre for Clinical Research in Neuropsychiatry, School of Psychiatry and Clinical Neurosciences, The University of Western Australia, Medical Research Foundation Building, Perth, Western Australia, Australia.

    • Assen V Jablensky
  212. The Perkins Institute for Medical Research, The University of Western Australia, Perth, Western Australia, Australia.

    • Assen V Jablensky
  213. The Zucker Hillside Hospital, Glen Oaks, New York, USA.

    • Todd Lencz
    •  & Anil K Malhotra
  214. The Feinstein Institute for Medical Research, Manhasset, New York, USA.

    • Todd Lencz
    •  & Anil K Malhotra
  215. The Hofstra North Shore–Long Island Jewish School of Medicine, Hempstead, New York, USA.

    • Todd Lencz
    •  & Anil K Malhotra
  216. Saw Swee Hock School of Public Health, National University of Singapore, Singapore, Singapore.

    • Jianjun Liu
  217. Queensland Brain Institute, The University of Queensland, Brisbane, Queensland, Australia.

    • Bryan J Mowry
  218. Department of Functional Genomics, Center for Neurogenomics and Cognitive Research, Neuroscience Campus Amsterdam, VU University, Amsterdam, the Netherlands.

    • Danielle Posthuma
  219. Department of Complex Trait Genetics, Neuroscience Campus Amsterdam, VU University Medical Center Amsterdam, Amsterdam, the Netherlands.

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

    • Danielle Posthuma
  221. University of Aberdeen, Institute of Medical Sciences, Aberdeen, UK.

    • David St Clair
  222. Departments of Psychiatry, Neurology, Neuroscience and Institute of Genetic Medicine, Johns Hopkins School of Medicine, Baltimore, Maryland, USA.

    • Daniel R Weinberger
  223. MRC-SGDP Centre, Institute of Psychiatry, King's College London, London, UK.

    • Sylvane Desrivières
    • , Tianye Jia
    • , Christine Macare
    •  & Gunter Schumann
  224. Imaging Genetics Center, Institute for Neuroimaging & Informatics, Keck School of Medicine of the University of Southern California, Los Angeles, USA.

    • Neda Jahanshad
    •  & Christopher R K Ching
  225. Institut Pasteur, Paris, France.

    • Roberto Toro
  226. German Center for Neurodegenerative Diseases (DZNE) Rostock/Greifswald, Greifswald, Germany.

    • Katharina Wittfeld
    •  & Wolfgang Hoffmann
  227. Department of Psychiatry, University Medicine Greifswald, Greifswald, Germany.

    • Katharina Wittfeld
    • , Deborah Janowitz
    •  & Hans J Grabe
  228. Brain Center Rudolf Magnus, Department of Psychiatry, UMC Utrecht, Utrecht, the Netherlands.

    • Lucija Abramovic
    • , Marc M Bohlken
    • , Marco P Boks
    • , Kristel R van Eijk
    • , Neeltje E M van Haren
    • , Rachel M Brouwer
    •  & Hilleke E Hulshoff Pol
  229. Umeå Centre for Functional Brain Imaging (UFBI), Umeå University, Umeå, Sweden.

    • Micael Andersson
    • , Alireza Salami
    • , Lars G Nilsson
    •  & Lars Nyberg
  230. Brain Research Imaging Centre, University of Edinburgh, Edinburgh, UK.

    • Benjamin S Aribisala
    • , Natalie A Royle
    • , Mark E Bastin
    •  & Joanna M Wardlaw
  231. Department of Computer Science, Lagos State University, Lagos, Nigeria.

    • Benjamin S Aribisala
  232. Scottish Imaging Network, A Platform for Scientific Excellence (SINAPSE) Collaboration, Department of Neuroimaging Sciences, University of Edinburgh, Edinburgh, UK.

    • Benjamin S Aribisala
    • , Natalie A Royle
    • , Mark E Bastin
    •  & Joanna M Wardlaw
  233. Hospital for Sick Children, University of Toronto, Toronto, Ontario, Canada.

    • Manon Bernard
    • , Jean Shin
    •  & Zdenka Pausova
  234. Cerebral Imaging Centre, Douglas Mental Health University Institute, Montreal, Quebec, Canada.

    • M Mallar Chakravarty
  235. Department of Psychiatry and Biomedical Engineering, McGill University, Montreal, Quebec, Canada.

    • M Mallar Chakravarty
  236. Biological Psychology, Neuroscience Campus Amsterdam, VU University & VU Medical Center, Amsterdam, the Netherlands.

    • Anouk den Braber
    • , Iryna O Fedko
    • , Jouke-Jan Hottenga
    • , Dennis van 't Ent
    • , Dorret I Boomsma
    •  & Eco J C de Geus
  237. NORMENT – KG Jebsen Centre for Psychosis Research, Department of Clinical Science, University of Bergen, Bergen, Norway.

    • Sudheer Giddaluru
    • , Vidar M Steen
    •  & Stephanie LeHellard
  238. Dr Einar Martens Research Group for Biological Psychiatry, Center for Medical Genetics and Molecular Medicine, Haukeland University Hospital, Bergen, Norway.

    • Sudheer Giddaluru
    • , Vidar M Steen
    •  & Stephanie LeHellard
  239. Language and Genetics Department, Max Planck Institute for Psycholinguistics, Nijmegen, the Netherlands.

    • Tulio Guadalupe
  240. International Max Planck Research School for Language Sciences, Nijmegen, the Netherlands.

    • Tulio Guadalupe
  241. National Institute of Mental Health Intramural Research Program, Bethesda, Maryland, USA.

    • Girma Woldehawariat
    • , Allison C Nugent
    • , Xinmin Liu
    • , Wayne C Drevets
    •  & Dara M Cannon
  242. Center for Neurobehavioral Genetics, University of California, Los Angeles, California, USA.

    • Loes M Olde Loohuis
  243. Centre for Cognitive Ageing and Cognitive Epidemiology, Psychology, University of Edinburgh, Edinburgh, UK.

    • Michelle Luciano
    • , David C M Liewald
    • , Lorna M Lopez
    • , Natalie A Royle
    • , Mark E Bastin
    • , Ian J Deary
    •  & Joanna M Wardlaw
  244. Department of Psychiatry, Neuroscience Campus Amsterdam, VU University Medical Center, Amsterdam, the Netherlands.

    • Yuri Milaneschi
    • , Lianne Schmaal
    • , Dick J Veltman
    •  & Brenda W J H Penninx
  245. Department of Medical and Molecular Genetics, King's College London, London, UK.

    • Adaikalavan Ramasamy
    • , Mina Ryten
    •  & Michael E Weale
  246. Reta Lila Weston Institute and Department of Molecular Neuroscience, UCL Institute of Neurology, London, UK.

    • Adaikalavan Ramasamy
    • , Daniah Trabzuni
    • , Raphael Gibbs
    • , Sebastian Guelfi
    • , Dena G Hernandez
    • , John Hardy
    •  & Mina Ryten
  247. Max Planck Institute of Psychiatry, Munich, Germany.

    • Philipp G Sämann
    • , David Höhn
    • , Benno Pütz
    • , Christiane Wolf
    • , Michael Czisch
    •  & Florian Holsboer
  248. Multimodal Imaging Laboratory, Department of Neurosciences, University of California, San Diego, California, USA.

    • Andrew J Schork
  249. Department of Cognitive Sciences, University of California, San Diego, California, USA.

    • Andrew J Schork
  250. Institute for Community Medicine, University Medicine Greifswald, Greifswald, Germany.

    • Alexander Teumer
    • , Wolfgang Hoffmann
    •  & Henry Völzke
  251. NORMENT – KG Jebsen Centre, Division of Mental Health and Addiction, Oslo University Hospital, Oslo, Norway.

    • Lars T Westlye
    •  & Thomas Espeseth
  252. NORMENT – KG Jebsen Centre, Department of Psychology, University of Oslo, Oslo, Norway.

    • Lars T Westlye
    • , Ivar Reinvang
    •  & Thomas Espeseth
  253. Molecular and Cellular Therapeutics, The Royal College of Surgeons, Dublin, 2, Ireland.

    • Christopher D Whelan
    • , Saud Alhusaini
    • , Gianpiero L Cavalleri
    •  & Norman Delanty
  254. The Oxford Center for Functional MRI of the Brain, Nuffield Department of Clinical Neurosciences, Oxford University, Oxford, UK.

    • Anderson M Winkler
  255. Department of Neurology and Neurosurgery, Montreal Neurological Institute, McGill University, Montreal, Quebec, Canada.

    • Saud Alhusaini
  256. UCL Institute of Neurology, University College London, and Epilepsy Society, London, UK.

    • Dalia Kasperaviciute
    • , Mar Matarin
    •  & Sanjay M Sisodiya
  257. Department of Medicine, Imperial College London, London, UK.

    • Dalia Kasperaviciute
  258. Department of Psychiatry, Yale University, New Haven, Connecticut, USA.

    • David R McKay
    •  & David C Glahn
  259. Olin Neuropsychiatric Research Center, Hartford, Connecticut, USA.

    • David R McKay
    •  & David C Glahn
  260. Department of Genetics, King Faisal Specialist Hospital and Research Centre, Riyadh, Saudi Arabia.

    • Daniah Trabzuni
  261. Texas Biomedical Research Institute, San Antonio, Texas, USA.

    • Laura Almasy
    • , Melanie A Carless
    • , Joanne E Curran
    • , Ravi Duggirala
    • , Thomas D Dyer
    • , Harald H H Göring
    • , Jack W Kent Jr
    •  & John Blangero
  262. National Ageing Research Institute, Royal Melbourne Hospital, Melbourne, Victoria, Australia.

    • David Ames
  263. Academic Unit for Psychiatry of Old Age, University of Melbourne, Melbourne, Victoria, Australia.

    • David Ames
  264. Laboratory of Neurogenetics, National Institute on Aging, National Institutes of Health, Bethesda, Maryland, USA.

    • Sampath Arepalli
    • , Allissa Dillman
    • , Raphael Gibbs
    • , Dena G Hernandez
    • , Michael A Nalls
    • , Bryan Traynor
    • , Mark R Cookson
    •  & Andrew Singleton
  265. Centre for Clinical Brain Sciences, University of Edinburgh, Edinburgh, UK.

    • Mark E Bastin
    • , Maria C Valdés Hernández
    •  & Joanna M Wardlaw
  266. NI Vavilov Institute of General Genetics, Russian Academy of Sciences, Moscow, Russia.

    • Kazima B Bulayeva
  267. Clinical Research Branch, National Institute on Aging, Baltimore, Maryland, USA.

    • Luigi Ferrucci
  268. University of Texas Health Science Center, San Antonio, USA.

    • Peter T Fox
    •  & Rene L Olvera
  269. Biofunctional Imaging, Immunology Frontier Research Center, Osaka University, Osaka, Japan.

    • Masaki Fukunaga
  270. Harvard Medical School, Cambridge, Massachusetts, USA.

    • Robert C Green
  271. Division of Genetics, Department of Medicine, Brigham and Women's Hospital, Boston, Massachusetts, USA.

    • Robert C Green
  272. Department of Psychiatry, University of Groningen, University Medical Center Groningen, Groningen, the Netherlands.

    • Catharina A Hartman
    •  & Pieter J Hoekstra
  273. Institute of Diagnostic Radiology and Neuroradiology, University Medicine Greifswald, Greifswald, Germany.

    • Katrin Hegenscheid
    •  & Norbert Hosten
  274. Department of Psychology, VU University Amsterdam, Amsterdam, the Netherlands.

    • Dirk J Heslenfeld
  275. Interfaculty Institute for Genetics and Functional Genomics, University Medicine Greifswald, Greifswald, Germany.

    • Georg Homuth
  276. Radiology, Mayo Clinic, Rochester, Minnesota, USA.

    • Clifford R Jack Jr
  277. FMRIB Centre, University of Oxford, Oxford, UK.

    • Mark Jenkinson
  278. NICHD Brain and Tissue Bank for Developmental Disorders, University of Maryland Medical School, Baltimore, Maryland, USA.

    • Robert Johnson
    •  & Ronald H Zielke
  279. School of Psychology, University of Sussex, Brighton, UK.

    • Ryota Kanai
  280. Institute of Cognitive Neuroscience, University College London, London, UK.

    • Ryota Kanai
  281. Department of Psychiatry, University of Maryland, Catonsville, Maryland, USA.

    • Peter Kochunov
  282. Neuroscience Research Australia, Sydney, New South Wales, Australia.

    • John B Kwok
    •  & Peter R Schofield
  283. School of Medical Sciences, University of New South Wales, Sydney, New South Wales, Australia.

    • John B Kwok
    •  & Peter R Schofield
  284. Columbia University Medical Center, New York, USA.

    • Xinmin Liu
  285. Lymphocyte Cell Biology Unit, Laboratory of Immunology, National Institute on Aging, National Institutes of Health, Baltimore, Maryland, USA.

    • Dan L Longo
  286. Centre for Advanced Imaging, University of Queensland, Brisbane, Queensland, Australia.

    • Katie L McMahon
  287. Ludwig-Maximilians-Universität, Munich, Germany.

    • Eva Meisenzahl
  288. Department of Psychiatry, Osaka University Graduate School of Medicine, Osaka, Japan.

    • Kazutaka Ohi
  289. Department of Neurology, University of Calgary, Calgary, Canada.

    • G Bruce Pike
  290. Department of Clinical Neuroscience, University of Calgary, Calgary, Canada.

    • G Bruce Pike
  291. Psychiatry and Human Behavior, University of California, Irvine, California, USA.

    • Steven G Potkin
  292. Beth Israel Deaconess Medical Center, Boston, Massachusetts, USA.

    • Glenn D Rosen
  293. Department of Neurology, Harvard Medical School, Boston, Massachusetts, USA.

    • Glenn D Rosen
  294. Department of Neuropathology, MRC Sudden Death Brain Bank Project, University of Edinburgh, Edinburgh, UK.

    • Colin Smith
  295. Laboratory of Neuro Imaging, Institute for Neuroimaging and Informatics, Keck School of Medicine of the University of Southern California, Los Angeles, California, USA.

    • Arthur W Toga
  296. Brain Resource Center, Johns Hopkins University, Baltimore, Maryland, USA.

    • Juan Troncoso
  297. The Scripps Research Institute, Jupiter, Florida, USA.

    • Marcel van der Brug
  298. Leiden University Medical Center, Leiden, the Netherlands.

    • Nic J A van der Wee
  299. Neuroimaging Centre, University of Groningen, University Medical Center Groningen, Groningen, the Netherlands.

    • Marie-Jose van Tol
  300. Department of Psychiatry, Carver College of Medicine, University of Iowa, Iowa City, lowa, USA.

    • Thomas H Wassink
  301. Department of Neurobiology, Care Sciences and Society, Karolinska Institutet, Stockholm, Sweden.

    • Eric Westman
  302. Research Resources Branch, National Institute on Aging, National Institutes of Health, Bethesda, Maryland, USA.

    • Alan Zonderman
  303. Faculty of Life Sciences, University of Manchester, Manchester, UK.

    • David G Ashbrook
    •  & Reinmar Hager
  304. Center for Integrative and Translational Genomics, University of Tennessee Health Science Center, Memphis, Tennessee, USA.

    • Lu Lu
    •  & Robert W Williams
  305. Department of Anatomy and Neurobiology, University of Tennessee Health Science Center, Memphis, Tennessee, USA.

    • Lu Lu
    •  & Robert W Williams
  306. The Mind Research Network & LBERI, Albuquerque, New Mexico, USA.

    • Vince D Calhoun
  307. Department of ECE, University of New Mexico, Albuquerque, New Mexico, USA.

    • Vince D Calhoun
  308. Center for Translational Imaging and Personalized Medicine, University of California, San Diego, California, USA.

    • Anders M Dale
  309. Departments of Neurosciences, Radiology, Psychiatry, and Cognitive Science, University of California, San Diego, California, USA.

    • Anders M Dale
  310. Avera Institute for Human Genetics, Sioux Falls, South Dakota, USA.

    • Gareth E Davies
  311. Neurology Division, Beaumont Hospital, Dublin, Ireland.

    • Norman Delanty
  312. Department of Neurology, Hopital Erasme, Universite Libre de Bruxelles, Brussels, Belgium.

    • Chantal Depondt
    •  & Massimo Pandolfo
  313. Janssen Research & Development, Johnson & Johnson, Raritan, New Jersey, USA.

    • Wayne C Drevets
  314. Department of Psychiatry, University of Iowa, Iowa City. lowa, USA.

    • Beng-Choon Ho
  315. Institute of Clinical Chemistry and Laboratory Medicine, University Medicine Greifswald, Greifswald, Germany.

    • Matthias Nauck
  316. Center for Imaging of Neurodegenerative Disease, San Francisco Veterans Affairs Medical Center, University of California, San Francisco, USA.

    • Michael W Weiner
  317. Department of Child Psychiatry, Erasmus University Medical Centre, Rotterdam, the Netherlands.

    • Tonya White
  318. Department of Radiology, Erasmus University Medical Centre, Rotterdam, the Netherlands.

    • Tonya White
    • , Hieab H H Adams
    •  & M Arfan Ikram
  319. Clinical Neuroimaging Laboratory, College of Medicine, Nursing and Health Sciences, National University of Ireland Galway, Galway, Ireland.

    • Dara M Cannon
  320. Department of Psychiatry and Psychotherapy, HELIOS Hospital Stralsund, Stralsund, Germany.

    • Hans J Grabe
  321. Molecular Research Center for Children's Mental Development, United Graduate School of Child Development, Osaka University, Osaka, Japan.

    • Ryota Hashimoto
  322. Medical University of Lodz, Lodz, Poland.

    • Iwona Kloszewska
  323. Section of Gerontology and Geriatrics, Department of Medicine, University of Perugia, Perugia, Italy.

    • Patrizia Mecocci
  324. Rotman Research Institute, University of Toronto, Toronto, Ontario, Canada.

    • Tomas Paus
  325. Departments of Psychology and Psychiatry, University of Toronto, Ontario, Canada.

    • Tomas Paus
  326. Departments of Physiology and Nutritional Sciences, University of Toronto, Ontario, Canada.

    • Zdenka Pausova
  327. Department of Neuroimaging, Institute of Psychiatry, King's College London, London, UK.

    • Andy Simmons
  328. Biomedical Research Centre for Mental Health, King's College London, London, UK.

    • Andy Simmons
  329. Biomedical Research Unit for Dementia, King's College London, London, UK.

    • Andy Simmons
  330. Institute of Clinical Medicine, Neurology, University of Eastern Finland, Kuopio, Finland.

    • Hilkka Soininen
  331. Neurocentre Neurology, Kuopio University Hospital, Kuopio, Finland.

    • Hilkka Soininen
  332. Department of Epidemiology, Erasmus University Medical Centre, Rotterdam, the Netherlands.

    • Hieab H H Adams
    • , Sven J van der Lee
    • , Cornelia M van Duijn
    •  & M Arfan Ikram
  333. Laboratory of Epidemiology and Population Sciences, Intramural Research Program, National Institute on Aging, Bethesda, Maryland, USA.

    • Lenore J Launer
  334. Department of Neurology, Clinical Division of Neurogeriatrics, Medical University Graz, Graz, Austria.

    • Stephan Seiler
    •  & Reinhold Schmidt
  335. INSERM U897, University of Bordeaux, Bordeaux, France.

    • Ganesh Chauhan
  336. Department of Neurology, Boston University School of Medicine, Boston, Massachusetts, USA.

    • Claudia L Satizabal
    •  & Sudha Seshadri
  337. Framingham Heart Study, Framingham, Massachusetts, USA.

    • Claudia L Satizabal
    •  & Sudha Seshadri
  338. Department of Neurology, School of Medicine, University of Pittsburgh, Pittsburgh, Pennsylvania, USA.

    • James T Becker
  339. Department of Psychiatry, School of Medicine, University of Pittsburgh, Pittsburgh, Pennsylvania, USA.

    • James T Becker
  340. Department of Psychology, School of Medicine, University of Pittsburgh, Pittsburgh, Pennsylvania, USA.

    • James T Becker
  341. General Internal Medicine, Johns Hopkins School of Medicine, Baltimore, Maryland, USA.

    • Lisa Yanek
  342. Martinos Center for Biomedical Imaging, Massachusetts General Hospital, Charlestown, Massachusetts, USA.

    • Maritza Ebling
    • , Bruce Fischl
    • , Douglas Greve
    •  & Louis N Vinke
  343. Department of Radiology, Massachusetts General Hospital, Harvard Medical School, Boston, Massachusetts, USA.

    • Maritza Ebling
    • , Bruce Fischl
    • , Douglas Greve
    •  & Louis N Vinke
  344. Department of Neurology University of Washington, Seattle, Washington, USA.

    • W T Longstreth
  345. Institute of Molecular Biology and Biochemistry, Medical University Graz, Graz, Austria.

    • Helena Schmidt
  346. Department of Neurology, Johns Hopkins University School of Medicine, Baltimore, Maryland, USA.

    • Paul Nyquist
  347. Department of Biostatistics, Boston University School of Public Health, Boston, Massachuestts, USA.

    • Xue Luting
  348. UMR5296, Centre d'études scientifiques et techniques d'Aquitaine and University of Bordeaux, Bordeaux, France.

    • Bernard Mazoyer
  349. Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, Washington, USA.

    • Joshua C Bis
  350. Icelandic Heart Association, Kopavogur, University of Iceland, Faculty of Medicine, Reykjavik, Iceland.

    • Vilmundur Gudnason

Consortia

  1. Schizophrenia Working Group of the Psychiatric Genomics Consortium

    Psychosis Endophenotypes International Consortium

    Wellcome Trust Case Control Consortium 2

  2. Enigma Consortium

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Contributions

Study conception and supervision: B. Franke, B.M.N., J.L.S., M.C.O'D., P.F.S., P.M.T., S.E.M. Design of ENIGMA or PGC: A.A.-V., A.M.M., B. Franke, B.M.N., D.P.H., J.A.T., J.L.S., J.W.S., K.J.E.v.H., M.C.N., M.C.O'D., O.A.A., P.F.S., P.L., P.M.T., S.E.M., S. Ripke, T.E.N., V.A., Y.Y., Y.Y.W.H. Obtained funding: B. Franke, M.C.O'D., M.J.W., N.G.M., P.F.S., P.M.T. Provided samples: PGC2 Schizophrenia Working Group and ENIGMA2 Consortium. Conducted analyses: A.A.-V., B.M.N., D.P.H., J.L.S., K.J.E.v.H., M.C.N., P.L., S.E.M., S. Ripke, T.E.N., V.A., Y.Y.W.H. Writing group: A.A.-V., B. Franke, B.M.N., D.P.H., J.L.S., K.J.E.v.H., M.C.O'D., P.F.S., P.M.T., S.E.M., S. Ripke, V.A. All authors reviewed and approved the final version of this manuscript.

Competing interests

Some of the authors or collaborators were employees of the following pharmaceutical companies: Pfizer (C.R.S., J.R.W., H.S.X.), F. Hoffman-La Roche (E. Domenici, L.E.), Eli Lilly (D.A.C., Y. Mokrab, L. Nisenbaum) and Janssen (S. Gopal, D. Wang, Q.S.L., N. Cohen). None of these companies influenced the design of the study, the interpretation of the data or the amount of data reported, or financially profit by publication of the results, which are precompetitive.

Corresponding authors

Correspondence to Barbara Franke or Patrick F Sullivan.

Integrated supplementary information

Supplementary information

PDF files

  1. 1.

    Supplementary Text and Figures

    Supplementary Figures 1–10 and Supplementary Table 4

  2. 2.

    Supplementary Methods Checklist

Excel files

  1. 1.

    Supplementary Table 1: Characteristics of the PGC schizophrenia and ENIGMA data sets

    Tag refers to the PGC sample identifier. The numbers shown are before exclusion of overlap with ENIGMA.

  2. 2.

    Supplementary Table 2: Reciprocal look-up of genome-wide significant findings in the other data set

    Panels 1–8: Shown are LD-independent genome-wide significant SNP associations for schizophrenia from the final analysis (sorted by genomic position according to UCSC hg19/NCBI Build 37). Markers are ranked from 1 to 128 in order of significance. Insertion/deletion variants are given in the form “chrA_B_C” where A = chromosome, B = position, and C = insertion (I) or deletion (D). Column A12 has the SNP alleles, with the first allele (a1) the reference allele for the frequency and odds ratio columns. Frq = frequency of allele 1. Chr and Position denote the associated region surrounding the index SNP containing 1 or more SNPs in LD (r2 > 0.6) with the index SNP. OR = odds ratio for allele 1, CI = 95% confidence interval for OR. Effect = effect size for allele 1, CI = 95% confidence interval for the effect size and are provided in units of mm3 per effect allele. †Top finding from schizophrenia (rs115329265) was not available in ENIGMA and was replaced by a SNP in moderate LD (6:28305863R; r2 = 0.64). *X-chromosome-located SNPs were excluded because X-chromosomal information was not available in ENIGMA. **rs77149735 was not available in ENIGMA and could not be replaced by a SNP in moderate or high LD. Panel 9: The allele frequency (Freq) and effect size (Effect) are given with reference to allele 1 (A1). CI = 95% confidence interval for the effect size and is provided in units of mm3 per effect allele. OR = odds ratio for allele 1, CI = 95% confidence interval for OR.

  3. 3.

    Supplementary Table 3: P-values for the PGC SCZ and ENIGMA volumes and combined meta-analyses

    The combined meta-analysis results for each brain volume was ‘clumped’ with the 1000 Genomes V3 reference panel for Europeans using PLINK. P-value-informed LD clumping was done using 5 × 10-8 as significance threshold for index SNPs, 5 × 10-6 as secondary significance threshold for clumped SNPs, 0.8 as LD r2 threshold, and 1 MB as physical distance threshold.

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DOI

https://doi.org/10.1038/nn.4228

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