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Genetic influences on schizophrenia and subcortical brain volumes: large-scale proof of concept

Nature Neuroscience volume 19, pages 420–431 (2016)Cite this article

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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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Figure 1: Genetic predisposition score analyses examining the predictive capacity of ENIGMA brain volumetric results on schizophrenia case-control status using different P-value thresholds.
Figure 2: Evaluating the genome-wide overlap between genetic influences on schizophrenia and subcortical volumes.

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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

  1. Paul Nyquist: Full lists of members and affiliations appear above.

  2. Barbara Franke, Jason L Stein, Stephan Ripke, Stephan Ripke, Derrek P Hibar, Jason L Stein, Miguel E Renteria, Alejandro Arias-Vasquez, Sylvane Desrivières and Xue Luting: These authors contributed equally to this work.

  3. Benjamin M Neale, Patrick F Sullivan, Michael C O'Donovan, Bernard Mazoyer, 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 and Patrick F Sullivan: These authors jointly directed this work.

Authors and Affiliations

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

    Barbara Franke, Kimm J E van Hulzen, Alejandro Arias-Vasquez, 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, Kimm J E van Hulzen, Jeanette C Mostert, Han G Brunner, Hans van Bokhoven & Barbara Franke

  2. Department of Psychiatry, Radboud University Medical Center, Nijmegen, the Netherlands

    Barbara Franke, Alejandro Arias-Vasquez, Alejandro Arias-Vasquez & Barbara Franke

  3. Donders Institute for Brain, Cognition and Behaviour, Raboud University, Nijmegen, the Netherlands

    Barbara Franke, Kimm J E van Hulzen, Alejandro Arias-Vasquez, 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, Kimm J E van Hulzen, Jeanette C Mostert, Han G Brunner, Jan K Buitelaar, Hans van Bokhoven, Guillén Fernández, Simon E Fisher, Clyde Francks & Barbara Franke

  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, Derrek P Hibar, Jason L Stein, Paul M Thompson & Paul M Thompson

  5. Department of Neurology, Neurogenetics Program, UCLA School of Medicine, Los Angeles, California, USA

    Jason L Stein & Jason L Stein

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

    Stephan Ripke, Verneri Anttila, Stephan Ripke, Benjamin M Neale, Kai-How Farh, Brendan Bulik-Sullivan, Hailiang Huang, Menachem Fromer, Jacqueline I Goldstein, Mark J Daly, Raymond K Walters & Benjamin M Neale

  7. Stanley Center for Psychiatric Research, Broad Institute of MIT and Harvard, Cambridge, Massachusetts, USA

    Stephan Ripke, Verneri Anttila, Jordan W Smoller, Phil Lee, Stephan Ripke, Benjamin M Neale, Phil Lee, 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, Jordan W Smoller, Steven A McCarroll, Jennifer L Moran, Aarno Palotie, Tracey L Petryshen, Mark J Daly, Phil Lee, Jordan W Smoller & Benjamin M Neale

  8. Department of Psychiatry and Psychotherapy, Charité Universitätsmedizin Berlin, Campus Charité Mitte, Berlin, Germany

    Stephan Ripke, 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, 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, Phil Lee, Menachem Fromer, Jordan W Smoller, Aarno Palotie, Phil Lee, Jordan W Smoller & Benjamin M Neale

  11. Department of Psychiatry, Harvard Medical School, Boston, Massachusetts, USA

    Jordan W Smoller, Phil Lee, Phil Lee, Lynn E DeLisi, Robert W McCarley, Raquelle I Mesholam-Gately, Larry J Seidman, Jordan W Smoller, Tracey L Petryshen, Phil Lee & Jordan W Smoller

  12. FMRIB Centre, University of Oxford, Oxford, UK

    Thomas E Nichols & Thomas E Nichols

  13. Department of Statistics & Warwick Manufacturing Group, University of Warwick, Coventry, UK

    Thomas E Nichols & 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, Andrew M McIntosh, Martina Papmeyer, Emma Sprooten, Stephen M Lawrie, Jessika E Sussmann & Andrew M McIntosh

  16. US Department of Health & Human Services, Intramural Research Program, National Institutes of Health, Bethesda, Maryland, USA

    Francis J McMahon, Yin Yao & Francis J McMahon

  17. Central Institute of Mental Health, Medical Faculty Mannheim, University Heidelberg, Mannheim, Germany

    Andreas Meyer-Lindenberg, Emanuel Schwarz, Oliver Grimm & Andreas Meyer-Lindenberg

  18. Department of Biomedicine, Aarhus University, Aarhus, Denmark

    Manuel Mattheisen, Manuel Mattheisen, Manuel Mattheisen & 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 & Manuel Mattheisen

  20. Center for integrated Sequencing, iSEQ, Aarhus University, Aarhus, Denmark

    Manuel Mattheisen, Manuel Mattheisen & Manuel Mattheisen

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

    Ole A Andreassen, Ole A Andreassen, Andrew A Brown, Lavinia Athanasiu, Cecilie B Hartberg, Unn Haukvik & Ole A Andreassen

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

    Ole A Andreassen, Ingrid Melle, Ole A Andreassen, Ingrid Melle & Ole A Andreassen

  23. Department of Psychiatry and Psychotherapy, Center for Translational Research in Systems Neuroscience and Psychiatry, University Medical Center, Goettingen, Germany

    Oliver Gruber, Bernd Kraemer, Maria Keil & Oliver Gruber

  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, Karen A Mather, Amelia A Assareh, Henry Brodaty, Simone Reppermund, Anbupalam Thalamuthu, Wei Wen & Perminder S Sachdev

  25. Neuropsychiatric Institute, Prince of Wales Hospital, Sydney, New South Wales, Australia.,

    Perminder S Sachdev & Perminder S Sachdev

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

    Roberto Roiz-Santiañez & Roberto Roiz-Santiañez

  27. Cibersam (Centro Investigación Biomédica en Red Salud Mental), Madrid, Spain

    Roberto Roiz-Santiañez, 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 & Andrew J Saykin

  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 & Andrew J Saykin

  30. Medical and Molecular Genetics, Indiana University School of Medicine, Indianapolis, Indiana, USA

    Andrew J Saykin, Tatiana M Foroud & Andrew J Saykin

  31. Department of Child and Adolescent Psychiatry, Faculty of Medicine and University Hospital, Technische Universität Dresden, Dresden, Germany

    Stefan Ehrlich, Johanna Hass, Stefan Ehrlich & Esther Walton

  32. Georgia State University, Atlanta, Georgia, USA

    Jessica A Turner & Jessica A Turner

  33. Mind Research Network, Albuquerque, New Mexico, USA

    Jessica A Turner & 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, Nicholas G Martin, Margaret J Wright, Sarah E Medland & Sarah E Medland

  35. School of Psychology, University of Queensland, Brisbane, Queensland, Australia

    Margaret J Wright, Lachlan T Strike, Greig I de Zubicaray & Margaret J Wright

  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 & 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 & 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 & Benjamin M Neale

  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 & 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 & Patrick F Sullivan

  41. Department of Psychiatry, University of North Carolina, Chapel Hill, North Carolina, USA.,

    Diana O Perkins, Patrick F Sullivan & Patrick F Sullivan

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

    Aiden Corvin, Paul Cormican, Gary Donohoe, Derek W Morris, Michael Gill, Emma J Rose, Margaret Needham, Aiden Corvin, Derek W Morris & Gary Donohoe

  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, Srdjan Djurovic, Ingrid Agartz & Erik G Jönsson

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

    Ingrid Agartz & Ingrid Agartz

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

    Ingrid Agartz, Erik Söderman, Erik G Jönsson, Ingrid Agartz & 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. Department of Psychiatry, University Medical Center Groningen, 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 & Randy L Buckner

  60. Department of Psychiatry, Massachusetts General Hospital, Boston, Massachusetts, USA

    Randy L Buckner, Joshua L Roffman, Avram J Holmes, Randy L Buckner, Randy L Gollub & Joshua L Roffman

  61. Athinoula A Martinos Center, Massachusetts General Hospital, Boston, Massachusetts, USA

    Randy L Buckner, Joshua L Roffman, Randy L Buckner & Joshua L Roffman

  62. Department of Psychiatry, University of California at San Francisco, San Francisco, California, USA

    William F Byerley

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

    Wiepke Cahn, René S Kahn, Eric Strengman, Roel A Ophoff, Wiepke Cahn, René S Kahn & 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 & 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 & Benedicto Crespo-Facorro

  83. Centro Investigacion Biomedica en Red Salud Mental, Madrid, Spain

    Benedicto Crespo-Facorro & Benedicto Crespo-Facorro

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

    David Curtis

  85. Division of Psychiatry, Molecular Psychiatry Laboratory, 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, 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, Sven Cichon & Markus M Nöthen

  89. VIB Department of Molecular Genetics, Applied Molecular Genomics Unit, 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 & 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, Derek W Morris & Gary Donohoe

  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 & 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 & 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 & Dan Rujescu

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

    Ina Giegling, Dan Rujescu & 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. Department of Psychiatry, Academic Medical Centre University of Amsterdam, 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 & Sven Cichon

  123. Department of Clinical Biochemistry, Section of Neonatal Screening and Hormones, 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 & Masashi Ikeda

  125. Department of Psychiatry, Regional Centre for Clinical Research in Psychosis, 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 & 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 & Bertram Müller-Myhsok

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

    Bertram Müller-Myhsok & Bertram Müller-Myhsok

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

    Bertram Müller-Myhsok & 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 & Daniel R Weinberger

  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, 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 & 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. Department of Psychiatry, Brain Center Rudolf Magnus, 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. Department of Neuroimaging Sciences, Scottish Imaging Network, A Platform for Scientific Excellence (SINAPSE) Collaboration, 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. Department of Clinical Science, NORMENT – KG Jebsen Centre for Psychosis Research, 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. Department of Neurosciences, Multimodal Imaging Laboratory, 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. Division of Mental Health and Addiction, NORMENT – KG Jebsen Centre, Oslo University Hospital, Oslo, Norway

    Lars T Westlye & Thomas Espeseth

  252. Department of Psychology, NORMENT – KG Jebsen Centre, 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. Nuffield Department of Clinical Neurosciences, The Oxford Center for Functional MRI of the Brain, 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. Department of Medicine, Section of Gerontology and Geriatrics, 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. Department of Medicine, Cardiovascular Health Research Unit, University of Washington, Seattle, Washington, USA

    Joshua C Bis

  350. Icelandic Heart Association, Kopavogur, University of Iceland, Faculty of Medicine, Reykjavik, Iceland

    Vilmundur Gudnason

Authors
  1. Barbara Franke
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  2. Jason L Stein
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  3. Stephan Ripke
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  6. Kimm J E van Hulzen
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  7. Alejandro Arias-Vasquez
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  8. Jordan W Smoller
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  9. Thomas E Nichols
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  10. Michael C Neale
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  11. Andrew M McIntosh
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  12. Phil Lee
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  13. Francis J McMahon
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  14. Andreas Meyer-Lindenberg
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  15. Manuel Mattheisen
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  16. Ole A Andreassen
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  17. Oliver Gruber
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  18. Perminder S Sachdev
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  19. Roberto Roiz-Santiañez
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  20. Andrew J Saykin
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  21. Stefan Ehrlich
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  22. Karen A Mather
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  23. Jessica A Turner
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  24. Emanuel Schwarz
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  25. Anbupalam Thalamuthu
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  26. Yin Yao
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  27. Yvonne Y W Ho
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  28. Nicholas G Martin
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  29. Margaret J Wright
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  30. Michael C O'Donovan
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  31. Paul M Thompson
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  32. Benjamin M Neale
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  33. Sarah E Medland
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  34. Patrick F Sullivan
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Consortia

Schizophrenia Working Group of the Psychiatric Genomics Consortium

  • Stephan Ripke
  • , Benjamin M Neale
  • , Aiden Corvin
  • , James T R Walters
  • , Kai-How Farh
  • , Peter A Holmans
  • , Phil Lee
  • , Brendan Bulik-Sullivan
  • , David A Collier
  • , Hailiang Huang
  • , Tune H Pers
  • , Ingrid Agartz
  • , Esben Agerbo
  • , Margot Albus
  • , Madeline Alexander
  • , Farooq Amin
  • , Silviu A Bacanu
  • , Martin Begemann
  • , Richard A Belliveau Jr
  • , Judit Bene
  • , Sarah E Bergen
  • , Elizabeth Bevilacqua
  • , Tim B Bigdeli
  • , Donald W Black
  • , Richard Bruggeman
  • , Nancy G Buccola
  • , Randy L Buckner
  • , William F Byerley
  • , Wiepke Cahn
  • , Guiqing Cai
  • , Murray J Cairns
  • , Dominique Campion
  • , Rita M Cantor
  • , Vaughan J Carr
  • , Noa Carrera
  • , Stanley V Catts
  • , Kimberley D Chambert
  • , Raymond C K Chan
  • , Eric Y H Chen
  • , Ronald Y L Chen
  • , Wei Cheng
  • , Eric F C Cheung
  • , Siow Ann Chong
  • , C Robert Cloninger
  • , David Cohen
  • , Nadine Cohen
  • , Paul Cormican
  • , Nick Craddock
  • , Benedicto Crespo-Facorro
  • , James J Crowley
  • , David Curtis
  • , Michael Davidson
  • , Kenneth L Davis
  • , Franziska Degenhardt
  • , Jurgen Del Favero
  • , Lynn E DeLisi
  • , Ditte Demontis
  • , Dimitris Dikeos
  • , Timothy Dinan
  • , Srdjan Djurovic
  • , Gary Donohoe
  • , Elodie Drapeau
  • , Jubao Duan
  • , Frank Dudbridge
  • , Peter Eichhammer
  • , Johan Eriksson
  • , Valentina Escott-Price
  • , Laurent Essioux
  • , Ayman H Fanous
  • , Martilias S Farrell
  • , Josef Frank
  • , Lude Franke
  • , Robert Freedman
  • , Nelson B Freimer
  • , Joseph I Friedman
  • , Menachem Fromer
  • , Giulio Genovese
  • , Lyudmila Georgieva
  • , Elliot S Gershon
  • , Ina Giegling
  • , Paola Giusti-Rodríguez
  • , Stephanie Godard
  • , Jacqueline I Goldstein
  • , Srihari Gopal
  • , Jacob Gratten
  • , Lieuwe de Haan
  • , Christian Hammer
  • , Marian L Hamshere
  • , Mark Hansen
  • , Thomas Hansen
  • , Vahram Haroutunian
  • , Annette M Hartmann
  • , Frans A Henskens
  • , Stefan L Herms
  • , Joel N Hirschhorn
  • , Per Hoffmann
  • , Andrea Hofman
  • , Mads V Hollegaard
  • , David M Hougaard
  • , Masashi Ikeda
  • , Inge Joa
  • , Antonio Julià
  • , Anna K Kähler
  • , René S Kahn
  • , Luba Kalaydjieva
  • , Sena Karachanak-Yankova
  • , Juha Karjalainen
  • , David Kavanagh
  • , Matthew C Keller
  • , Brian J Kelly
  • , James L Kennedy
  • , Andrey Khrunin
  • , Yunjung Kim
  • , Janis Klovins
  • , James A Knowles
  • , Bettina Konte
  • , Vaidutis Kucinskas
  • , Zita Ausrele Kucinskiene
  • , Hana Kuzelova-Ptackova
  • , Claudine Laurent
  • , S Hong Lee
  • , Jimmy Lee Chee Keong
  • , Sophie E Legge
  • , Bernard Lerer
  • , Miaoxin Li
  • , Tao Li
  • , Kung-Yee Liang
  • , Jeffrey Lieberman
  • , Svetlana Limborska
  • , Jouko Lönnqvist
  • , Carmel M Loughland
  • , Jan Lubinski
  • , Milan Macek Jr
  • , Patrik K E Magnusson
  • , Brion S Maher
  • , Wolfgang Maier
  • , Jacques Mallet
  • , Sara Marsal
  • , Manuel Mattheisen
  • , Morten Mattingsdal
  • , Robert W McCarley
  • , Colm McDonald
  • , Andrew M McIntosh
  • , Sandra Meier
  • , Carin J Meijer
  • , Bela Melegh
  • , Ingrid Melle
  • , Raquelle I Mesholam-Gately
  • , Andres Metspalu
  • , Patricia T Michie
  • , Lili Milani
  • , Vihra Milanova
  • , Younes Mokrab
  • , Derek W Morris
  • , Ole Mors
  • , Bertram Müller-Myhsok
  • , Kieran C Murphy
  • , Robin M Murray
  • , Inez Myin-Germeys
  • , Mari Nelis
  • , Igor Nenadic
  • , Deborah A Nertney
  • , Gerald Nestadt
  • , Kristin K Nicodemus
  • , Liene Nikitina-Zake
  • , Laura Nisenbaum
  • , Annelie Nordin
  • , Eadbhard O'Callaghan
  • , Colm O'Dushlaine
  • , F Anthony O'Neill
  • , Sang-Yun Oh
  • , Ann Olincy
  • , Line Olsen
  • , Jim Van Os
  • , Christos Pantelis
  • , George N Papadimitriou
  • , Sergi Papiol
  • , Elena Parkhomenko
  • , Michele T Pato
  • , Tiina Paunio
  • , Psychosis Endophenotypes International Consortium
  • , Diana O Perkins
  • , Olli Pietiläinen
  • , Jonathan Pimm
  • , Andrew J Pocklington
  • , John Powell
  • , Alkes Price
  • , Ann E Pulver
  • , Shaun M Purcell
  • , Digby Quested
  • , Henrik B Rasmussen
  • , Abraham Reichenberg
  • , Mark A Reimers
  • , Alexander L Richards
  • , Joshua L Roffman
  • , Panos Roussos
  • , Douglas M Ruderfer
  • , Veikko Salomaa
  • , Alan R Sanders
  • , Ulrich Schall
  • , Christian R Schubert
  • , Thomas G Schulze
  • , Sibylle G Schwab
  • , Edward M Scolnick
  • , Rodney J Scott
  • , Larry J Seidman
  • , Jianxin Shi
  • , Jeremy M Silverman
  • , Kang Sim
  • , Petr Slominsky
  • , Jordan W Smoller
  • , Hon-Cheong So
  • , Erik Söderman
  • , Chris C A Spencer
  • , Eli A Stahl
  • , Elisabeth Stogmann
  • , Richard E Straub
  • , Eric Strengman
  • , Jana Strohmaier
  • , T Scott Stroup
  • , Mythily Subramaniam
  • , Jaana Suvisaari
  • , Dragan M Svrakic
  • , Jin P Szatkiewicz
  • , Srinivas Thirumalai
  • , Draga Toncheva
  • , Paul A Tooney
  • , Juha Veijola
  • , John Waddington
  • , Dermot Walsh
  • , Dai Wang
  • , Qiang Wang
  • , Bradley T Webb
  • , Mark Weiser
  • , Dieter B Wildenauer
  • , Nigel M Williams
  • , Stephanie Williams
  • , Stephanie H Witt
  • , Aaron R Wolen
  • , Emily H M Wong
  • , Brandon K Wormley
  • , Jing Qin Wu
  • , Hualin Simon Xi
  • , Clement C Zai
  • , Xuebin Zheng
  • , Fritz Zimprich
  • , Naomi R Wray
  • , Peter M Visscher
  • , Wellcome Trust Case Control Consortium 2
  • , Rolf Adolfsson
  • , Ole A Andreassen
  • , Douglas H R Blackwood
  • , Anders D Børglum
  • , Elvira Bramon
  • , Joseph D Buxbaum
  • , Sven Cichon
  • , Ariel Darvasi
  • , Enrico Domenici
  • , Hannelore Ehrenreich
  • , Tõnu Esko
  • , Pablo V Gejman
  • , Michael Gill
  • , Hugh Gurling
  • , Christina M Hultman
  • , Nakao Iwata
  • , Assen V Jablensky
  • , Erik G Jönsson
  • , Kenneth S Kendler
  • , George Kirov
  • , Jo Knight
  • , Todd Lencz
  • , Douglas F Levinson
  • , Qingqin S Li
  • , Jianjun Liu
  • , Anil K Malhotra
  • , Steven A McCarroll
  • , Andrew McQuillin
  • , Jennifer L Moran
  • , Preben B Mortensen
  • , Bryan J Mowry
  • , Markus M Nöthen
  • , Roel A Ophoff
  • , Michael J Owen
  • , Aarno Palotie
  • , Carlos N Pato
  • , Tracey L Petryshen
  • , Danielle Posthuma
  • , Marcella Rietschel
  • , Brien P Riley
  • , Dan Rujescu
  • , Pak C Sham
  • , Pamela Sklar
  • , David St Clair
  • , Daniel R Weinberger
  • , Jens R Wendland
  • , Thomas Werge
  • , Mark J Daly
  • , Patrick F Sullivan
  •  & Michael C O'Donovan

Enigma Consortium

  • Derrek P Hibar
  • , Jason L Stein
  • , Miguel E Renteria
  • , Alejandro Arias-Vasquez
  • , Sylvane Desrivières
  • , Neda Jahanshad
  • , Roberto Toro
  • , Katharina Wittfeld
  • , Lucija Abramovic
  • , Micael Andersson
  • , Benjamin S Aribisala
  • , Nicola J Armstrong
  • , Manon Bernard
  • , Marc M Bohlken
  • , Marco P Boks
  • , Janita Bralten
  • , Andrew A Brown
  • , M Mallar Chakravarty
  • , Qiang Chen
  • , Christopher R K Ching
  • , Gabriel Cuellar-Partida
  • , Anouk den Braber
  • , Sudheer Giddaluru
  • , Aaron L Goldman
  • , Oliver Grimm
  • , Tulio Guadalupe
  • , Johanna Hass
  • , Girma Woldehawariat
  • , Avram J Holmes
  • , Martine Hoogman
  • , Deborah Janowitz
  • , Tianye Jia
  • , Sungeun Kim
  • , Marieke Klein
  • , Bernd Kraemer
  • , Phil Lee
  • , Loes M Olde Loohuis
  • , Michelle Luciano
  • , Christine Macare
  • , Karen A Mather
  • , Manuel Mattheisen
  • , Yuri Milaneschi
  • , Kwangsik Nho
  • , Martina Papmeyer
  • , Adaikalavan Ramasamy
  • , Shannon L Risacher
  • , Roberto Roiz-Santiañez
  • , Emma J Rose
  • , Alireza Salami
  • , Philipp G Sämann
  • , Lianne Schmaal
  • , Andrew J Schork
  • , Jean Shin
  • , Lachlan T Strike
  • , Alexander Teumer
  • , Marjolein M J van Donkelaar
  • , Kristel R van Eijk
  • , Raymond K Walters
  • , Lars T Westlye
  • , Christopher D Whelan
  • , Anderson M Winkler
  • , Marcel P Zwiers
  • , Saud Alhusaini
  • , Lavinia Athanasiu
  • , Stefan Ehrlich
  • , Marina M H Hakobjan
  • , Cecilie B Hartberg
  • , Unn Haukvik
  • , Angelien J G A M Heister
  • , David Höhn
  • , Dalia Kasperaviciute
  • , David C M Liewald
  • , Lorna M Lopez
  • , Remco R R Makkinje
  • , Mar Matarin
  • , Marlies A M Naber
  • , David R McKay
  • , Margaret Needham
  • , Allison C Nugent
  • , Benno Pütz
  • , Natalie A Royle
  • , Li Shen
  • , Emma Sprooten
  • , Daniah Trabzuni
  • , Saskia S L van der Marel
  • , Kimm J E van Hulzen
  • , Esther Walton
  • , Christiane Wolf
  • , Laura Almasy
  • , David Ames
  • , Sampath Arepalli
  • , Amelia A Assareh
  • , Mark E Bastin
  • , Henry Brodaty
  • , Kazima B Bulayeva
  • , Melanie A Carless
  • , Sven Cichon
  • , Aiden Corvin
  • , Joanne E Curran
  • , Michael Czisch
  • , Greig I de Zubicaray
  • , Allissa Dillman
  • , Ravi Duggirala
  • , Thomas D Dyer
  • , Susanne Erk
  • , Iryna O Fedko
  • , Luigi Ferrucci
  • , Tatiana M Foroud
  • , Peter T Fox
  • , Masaki Fukunaga
  • , Raphael Gibbs
  • , Harald H H Göring
  • , Robert C Green
  • , Sebastian Guelfi
  • , Narelle K Hansell
  • , Catharina A Hartman
  • , Katrin Hegenscheid
  • , Andreas Heinz
  • , Dena G Hernandez
  • , Dirk J Heslenfeld
  • , Pieter J Hoekstra
  • , Florian Holsboer
  • , Georg Homuth
  • , Jouke-Jan Hottenga
  • , Masashi Ikeda
  • , Clifford R Jack Jr
  • , Mark Jenkinson
  • , Robert Johnson
  • , Ryota Kanai
  • , Maria Keil
  • , Jack W Kent Jr
  • , Peter Kochunov
  • , John B Kwok
  • , Stephen M Lawrie
  • , Xinmin Liu
  • , Dan L Longo
  • , Katie L McMahon
  • , Eva Meisenzahl
  • , Ingrid Melle
  • , Sebastian Mohnke
  • , Grant W Montgomery
  • , Jeanette C Mostert
  • , Thomas W Mühleisen
  • , Michael A Nalls
  • , Thomas E Nichols
  • , Lars G Nilsson
  • , Markus M Nöthen
  • , Kazutaka Ohi
  • , Rene L Olvera
  • , Rocio Perez-Iglesias
  • , G Bruce Pike
  • , Steven G Potkin
  • , Ivar Reinvang
  • , Simone Reppermund
  • , Marcella Rietschel
  • , Nina Romanczuk-Seiferth
  • , Glenn D Rosen
  • , Dan Rujescu
  • , Knut Schnell
  • , Peter R Schofield
  • , Colin Smith
  • , Vidar M Steen
  • , Jessika E Sussmann
  • , Anbupalam Thalamuthu
  • , Arthur W Toga
  • , Bryan Traynor
  • , Juan Troncoso
  • , Jessica A Turner
  • , Maria C Valdés Hernández
  • , Dennis van 't Ent
  • , Marcel van der Brug
  • , Nic J A van der Wee
  • , Marie-Jose van Tol
  • , Dick J Veltman
  • , Thomas H Wassink
  • , Eric Westman
  • , Ronald H Zielke
  • , Alan Zonderman
  • , David G Ashbrook
  • , Reinmar Hager
  • , Lu Lu
  • , Francis J McMahon
  • , Derek W Morris
  • , Robert W Williams
  • , Han G Brunner
  • , Randy L Buckner
  • , Jan K Buitelaar
  • , Wiepke Cahn
  • , Vince D Calhoun
  • , Gianpiero L Cavalleri
  • , Benedicto Crespo-Facorro
  • , Anders M Dale
  • , Gareth E Davies
  • , Norman Delanty
  • , Chantal Depondt
  • , Srdjan Djurovic
  • , Wayne C Drevets
  • , Thomas Espeseth
  • , Randy L Gollub
  • , Beng-Choon Ho
  • , Wolfgang Hoffmann
  • , Norbert Hosten
  • , René S Kahn
  • , Stephanie LeHellard
  • , Andreas Meyer-Lindenberg
  • , Bertram Müller-Myhsok
  • , Matthias Nauck
  • , Lars Nyberg
  • , Massimo Pandolfo
  • , Brenda W J H Penninx
  • , Joshua L Roffman
  • , Sanjay M Sisodiya
  • , Jordan W Smoller
  • , Hans van Bokhoven
  • , Neeltje E M van Haren
  • , Henry Völzke
  • , Henrik Walter
  • , Michael W Weiner
  • , Wei Wen
  • , Tonya White
  • , Ingrid Agartz
  • , Ole A Andreassen
  • , John Blangero
  • , Dorret I Boomsma
  • , Rachel M Brouwer
  • , Dara M Cannon
  • , Mark R Cookson
  • , Eco J C de Geus
  • , Ian J Deary
  • , Gary Donohoe
  • , Guillén Fernández
  • , Simon E Fisher
  • , Clyde Francks
  • , David C Glahn
  • , Hans J Grabe
  • , Oliver Gruber
  • , John Hardy
  • , Ryota Hashimoto
  • , Hilleke E Hulshoff Pol
  • , Erik G Jönsson
  • , Iwona Kloszewska
  • , Simon Lovestone
  • , Venkata S Mattay
  • , Patrizia Mecocci
  • , Colm McDonald
  • , Andrew M McIntosh
  • , Roel A Ophoff
  • , Tomas Paus
  • , Zdenka Pausova
  • , Mina Ryten
  • , Perminder S Sachdev
  • , Andrew J Saykin
  • , Andy Simmons
  • , Andrew Singleton
  • , Hilkka Soininen
  • , Joanna M Wardlaw
  • , Michael E Weale
  • , Daniel R Weinberger
  • , Hieab H H Adams
  • , Lenore J Launer
  • , Stephan Seiler
  • , Reinhold Schmidt
  • , Ganesh Chauhan
  • , Claudia L Satizabal
  • , James T Becker
  • , Lisa Yanek
  • , Sven J van der Lee
  • , Maritza Ebling
  • , Bruce Fischl
  • , W T Longstreth
  • , Douglas Greve
  • , Helena Schmidt
  • , Paul Nyquist
  • , Louis N Vinke
  • , Cornelia M van Duijn
  • , Xue Luting
  • , Bernard Mazoyer
  • , Joshua C Bis
  • , Vilmundur Gudnason
  • , Sudha Seshadri
  • , M Arfan Ikram
  • , Nicholas G Martin
  • , Margaret J Wright
  • , Gunter Schumann
  • , Barbara Franke
  • , Paul M Thompson
  •  & Sarah E Medland

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.

Corresponding authors

Correspondence to Barbara Franke or Patrick F Sullivan.

Ethics declarations

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.

Additional information

Psychosis Endophenotypes International Consortium, London, UK

Integrated supplementary information

Supplementary Figure 1 P-value-based meta-analysis between PGC schizophrenia (SCZ) and ENIGMA nucleus accumbens volume

PGC SCZ MA is shown in the top panel, the MA between PGC and ENIGMA is shown in the middle panel, and the ENIGMA MA is shown in the lower panel.

Supplementary Figure 2 P-value-based meta-analysis between PGC schizophrenia (SCZ) and ENIGMA amygdala volume.

PGC SCZ MA is shown in the top panel, the MA between PGC and ENIGMA is shown in the middle panel, and the ENIGMA MA is shown in the lower panel.

Supplementary Figure 3 P-value-based meta-analysis between PGC schizophrenia (SCZ) and ENIGMA caudate nucleus volume.

PGC SCZ MA is shown in the top panel, the MA between PGC and ENIGMA is shown in the middle panel, and the ENIGMA MA is shown in the lower panel.

Supplementary Figure 4 P-value-based meta-analysis between PGC schizophrenia (SCZ) and ENIGMA hippocampus volume.

PGC SCZ MA is shown in the top panel, the MA between PGC and ENIGMA is shown in the middle panel, and the ENIGMA MA is shown in the lower panel.

Supplementary Figure 5 P-value-based meta-analysis between PGC schizophrenia (SCZ) and ENIGMA ICV.

PGC SCZ MA is shown in the top panel, the MA between PGC and ENIGMA is shown in the middle panel, and the ENGIMA MA is shown in the lower panel.

Supplementary Figure 6 P-value-based meta-analysis between PGC schizophrenia (SCZ) and ENIGMA pallidum volume.

PGC SCZ MA is shown in the top panel, the MA between PGC and ENIGMA is shown in the middle panel, and the ENIGMA MA is shown in the lower panel.

Supplementary Figure 7 P-value-based meta-analysis between PGC schizophrenia (SCZ) and ENIGMA putamen volume.

PGC SCZ MA is shown in the top panel, the MA between PGC and ENIGMA is shown in the middle panel, and the ENGIMA MA is shown in the lower panel.

Supplementary Figure 8 P-value-based meta-analysis between PGC schizophrenia (SCZ) and ENIGMA thalamus volume.

PGC SCZ MA is shown in the top panel, the MA between PGC and ENIGMA is shown in the middle panel, and the ENIGMA MA is shown in the lower panel.

Supplementary Figure 9 Conjunction analysis: regional plots of genetic loci that influence both risk for schizophrenia and brain structure

We used a conjunction test to identify specific genetic variants influencing both risk for schizophrenia and changes in brain structure genome-wide. The conjunction test was run genome-wide using 7,510,842 SNPs found in both the ENIGMA2 and PGC2 studies with MAF ≥ 0.01. Conjunction results are then corrected for a downward bias in significance using the relaxed intersection union test. No SNP demonstrated genome-wide significant association to both schizophrenia and brain structure. Several sub-threshold loci did arise from this analysis that bear comment given the possibility that stronger significance may be achieved in future studies. (a) First, the conjunction between intracranial volume and schizophrenia was associated with a locus on chromosome 6 marked by rs381349, where the T allele increases ICV and decreases schizophrenia risk, in the expected direction for a schizophrenia risk factor. This locus is found intronic in the FOXO3 gene known to be involved in neural stem cell proliferation and renewal. (b) The conjunction between hippocampal volume and schizophrenia was associated with a previously mentioned locus on chromosome 2 marked by rs11693702, although in the opposite direction expected for a schizophrenia risk factor (A allele increases schizophrenia risk and increases hippocampal volume). (c,d) The conjunction between putamen volume and schizophrenia was associated with two interesting loci in the expected direction for schizophrenia risk factors marked by rs958246 (chromosome 11; A allele is associated with increased schizophrenia risk and increased putamen volume) and rs7231178 (chromosome 18; A allele is associated with increased schizophrenia risk and increased putamen volume). These SNPs are found in intronic regions of the DLG2 gene, which encodes a key component of the post-synaptic density, and the DCC gene, a netrin receptor involved in axon guidance.

Supplementary Figure 10 Comparison of SNP effect sizes

Effect sizes of individual SNPs having achieved replicated evidence for association with the phenotypes indicated. For brain structure phenotypes and schizophrenia, effect sizes for both, discovery and replication analyses are shown. For schizophrenia, only the five top SNPs from the 128 recently described genome-wide significant findings were included in the comparison. Height and educational attainment are shown as additional comparators. Effect sizes were measured in percent variance explained (for quantitative traits) or percent variance explained on the liability scale (for disease categories).

Supplementary information

Supplementary Text and Figures

Supplementary Figures 1–10 and Supplementary Table 4 (PDF 2300 kb)

Supplementary Methods Checklist (PDF 384 kb)

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. (XLSX 12 kb)

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. (XLSX 116 kb)

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. (XLSX 245 kb)

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Franke, B., Stein, J., Ripke, S. et al. Genetic influences on schizophrenia and subcortical brain volumes: large-scale proof of concept. Nat Neurosci 19, 420–431 (2016). https://doi.org/10.1038/nn.4228

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