Genome-wide association study identifies five new schizophrenia loci (original) (raw)

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Acknowledgements

We thank the study participants and the research staff at the many study sites. Over 40 US National Institutes of Health grants and similar numbers of government grants from other countries, along with substantial private and foundation support, enabled this work. We greatly appreciate the sustained efforts of T. Lehner (National Institute of Mental Health) on behalf of the Schizophrenia Psychiatric Genome-Wide Association Study (GWAS) Consortium (PGC). Detailed acknowledgments, including grant support, are listed in the Supplementary Note.

Author information

Author notes

  1. Maurice Jay, Virginia K Lasseter and Leena Peltonen: Deceased.

Authors and Affiliations

  1. Center for Human Genetic Research, Massachusetts General Hospital, Boston, Massachusetts, USA
    Stephan Ripke, Pamela Sklar, Lizzy Rossin, Douglas M Ruderfer, Sarah E Bergen & Mark J Daly
  2. Department of Psychiatry and Behavioral Sciences, NorthShore University HealthSystem, Evanston, Illinois, USA
    Alan R Sanders, Jubao Duan & Pablo V Gejman
  3. Department of Psychiatry and Behavioral Sciences, University of Chicago, Chicago, Illinois, USA
    Alan R Sanders, Jubao Duan & Pablo V Gejman
  4. Virginia Institute for Psychiatric and Behavioral Genetics, Virginia Commonwealth University School of Medicine, Richmond, Virginia, USA
    Kenneth S Kendler, Brion S Maher, Robert Ribble, Brien P Riley & Brandon Wormley
  5. Department of Psychiatry, Virginia Commonwealth University School of Medicine, Richmond, Virginia, USA
    Kenneth S Kendler, Ayman Fanous, Brion S Maher, Robert Ribble, Brien P Riley & Brandon Wormley
  6. Department of Human and Molecular Genetics, Virginia Commonwealth University School of Medicine, Richmond, Virginia, USA
    Kenneth S Kendler, Brion S Maher, Robert Ribble, Brien P Riley & Brandon Wormley
  7. Department of Psychiatry and Behavioral Sciences, Stanford University, Stanford, California, USA
    Douglas F Levinson & Madeline Alexander
  8. Department of Psychiatry, Mount Sinai School of Medicine, New York, New York, USA
    Pamela Sklar & Jeremy M Silverman
  9. Medical Research Council (MRC) Centre for Neuropsychiatric Genetics and Genomics, School of Medicine, Cardiff University, Cardiff, UK
    Peter A Holmans, Nicholas Craddock, Sarah Dwyer, Lyudmila Georgieva, Marian Hamshere, George K Kirov, Valentina Moskvina, Ivan Nikolov, Nadine Norton, Michael J Owen, Hywel J Williams, Nigel M Williams, Stan Zammit & Michael C O'Donovan
  10. Department of Psychological Medicine and Neurology, School of Medicine, Cardiff University, Cardiff, UK
    Peter A Holmans, Nicholas Craddock, Sarah Dwyer, Lyudmila Georgieva, Marian Hamshere, George K Kirov, Valentina Moskvina, Ivan Nikolov, Nadine Norton, Michael J Owen, Hywel J Williams, Nigel M Williams, Stan Zammit & Michael C O'Donovan
  11. Department of Biostatistics, University of North Carolina, Chapel Hill, North Carolina, USA
    Dan-Yu Lin
  12. Department of Medical Genetics, University Medical Center Utrecht, Utrecht, The Netherlands
    Roel A Ophoff & Eric Strengman
  13. Rudolf Magnus Institute of Neuroscience, University Medical Center Utrecht, Utrecht, The Netherlands
    Roel A Ophoff & Eric Strengman
  14. University of California at Los Angeles (UCLA) Center for Neurobehavioral Genetics, University of California at Los Angeles, Los Angeles, California, USA
    Roel A Ophoff, Rita M Cantor & Nelson B Freimer
  15. Department of Human Genetics, University of California at Los Angeles, Los Angeles, California, USA
    Roel A Ophoff & Rita M Cantor
  16. Psychiatry Section, Institute of Clinical Medicine, University of Oslo, Oslo, Norway
    Ole A Andreassen, Ingrid Agartz, Srdjan Djurovic, Morten Mattingsdal & Ingrid Melle
  17. Department of Psychiatry, Oslo University Hospital, Oslo, Norway
    Ole A Andreassen & Ingrid Melle
  18. Broad Institute, Cambridge, Massachusetts, USA
    Edward Scolnick, Shaun Purcell & Benjamin M Neale
  19. Department of Genomics, Life and Brain Center, University of Bonn, Bonn, Germany
    Sven Cichon, Manuel Mattheisen & Markus M Nöthen
  20. Institute of Neuroscience and Medicine (INM-1), Research Center Juelich, Juelich, Germany
    Sven Cichon
  21. Institute of Human Genetics, University of Bonn, Bonn, Germany
    Sven Cichon, Manuel Mattheisen & Markus M Nöthen
  22. Institute of Medical Sciences, University of Aberdeen, Foresterhill, Aberdeen, UK
    David St. Clair
  23. Neuropsychiatric Genetics Research Group, Trinity College Dublin, Dublin, Ireland
    Aiden Corvin, Paul Cormican, Gary Donohoe, Michael Gill, Elaine Kenny, Derek W Morris, Colm T O'Dushlaine & Emma M Quinn
  24. Research Department of Mental Health Sciences, Molecular Psychiatry Laboratory, University College London Medical School, Windeyer Institute of Medical Sciences, London, UK
    Hugh Gurling, Nicholas Bass, Khalid Choudhury, Susmita Datta, Robert Krasucki, Jacob Lawrence, Andrew McQuillin, Jonathan Pimm & Vinay Puri
  25. Institute of Biological Psychiatry, Mental Health Center (MHC) Sct. Hans, Copenhagen University Hospital, Roskilde, Denmark
    Thomas Werge, Linh Duong, Thomas Hansen, Andrés Ingason, Klaus D Jakobsen, Line Olsen, Henrik B Rasmussen & Johan H Thygesen
  26. Department of Psychiatry, Molecular and Clinical Neurobiology, Ludwig-Maximilians-University, Munich, Germany
    Dan Rujescu, Marion Friedl, Ina Giegling, Annette M Hartmann, Heike Konnerth & Bettina Konte
  27. Division of Psychiatry, University of Edinburgh, Royal Edinburgh Hospital, Edinburgh, UK
    Douglas H R Blackwood, Alan W Maclean, Pat Malloy, Kevin A McGhee & Andrew McIntosh
  28. Keck School of Medicine, University of Southern California, Los Angeles, California, USA
    Carlos N Pato & Michele T Pato
  29. Department of Psychiatry, Division of Research, The Zucker Hillside Hospital Division of the North Shore-Long Island Jewish Health System, Glen Oaks, New York, USA
    Anil K Malhotra & Todd Lencz
  30. Center for Psychiatric Neuroscience, The Feinstein Institute for Medical Research, Manhasset, New York, USA
    Anil K Malhotra & Todd Lencz
  31. Department of Psychiatry and Behavioral Science, Albert Einstein College of Medicine of Yeshiva University, New York, New York, USA
    Anil K Malhotra & Todd Lencz
  32. Department of Non-Communicable Disease Epidemiology, London School of Hygiene and Tropical Medicine, London, UK
    Frank Dudbridge
  33. Queensland Statistical Genetics Laboratory, Queensland Institute of Medical Research, Brisbane, Queensland, Australia
    Peter M Visscher
  34. Department of Functional Genomics, Vrije Universiteit (VU), Center for Neurogenomics and Cognitive Research (CNCR), Amsterdam, The Netherlands
    Danielle Posthuma
  35. Department of Medical Genomics, VU Medical Centre, Amsterdam, The Netherlands
    Danielle Posthuma
  36. Washington Veteran's Affairs Medical Center, Washington, DC, USA
    Ayman Fanous & Richard L Amdur
  37. Department of Psychiatry, Georgetown University School of Medicine, Washington, DC, USA
    Ayman Fanous & Richard L Amdur
  38. deCODE Genetics, Reykjavik, Iceland
    Hreinn Stefansson, Stacy Steinberg & Kari Stefansson
  39. Queensland Brain Institute, University of Queensland, Brisbane, Queensland, Australia
    Bryan J Mowry, John J McGrath & Deborah A Nertney
  40. Queensland Centre for Mental Health Research, University of Queensland, Brisbane, Queensland, Australia
    Bryan J Mowry, John J McGrath & Deborah A Nertney
  41. Mental Health Research Center, Russian Academy of Medical Sciences, Moscow, Russia
    Vera Golimbet
  42. University Psychiatric Centre, Catholic University Leuven, Kortenberg, Belgium
    Marc De Hert & Ruud van Winkel
  43. Department of Clinical Neuroscience, Human Brain Informatics (HUBIN) Project, Karolinska Institutet and Hospital, Stockholm, Sweden
    Erik G Jönsson & Lars Terenius
  44. Department of Psychiatry and Psychotherapy, Semmelweis University, Budapest, Hungary
    István Bitter & János M Réthelyi
  45. Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland
    Olli P H Pietiläinen & Leena Peltonen
  46. Department of Medical Genetics, University of Helsinki, Helsinki, Finland
    Olli P H Pietiläinen & Leena Peltonen
  47. Social, Genetic and Developmental Psychiatry Centre, Institute of Psychiatry, King's College, London, UK
    David A Collier
  48. Section of Psychiatry and Clinical Psychology, University of Verona, Verona, Italy
    Sarah Tosato & Mirella Ruggeri
  49. Department of Research, Diakonhjemmet Hospital, Oslo, Norway
    Ingrid Agartz
  50. State Mental Hospital, Haar, Germany
    Margot Albus
  51. Department of Psychiatry and Behavioral Sciences, Emory University, Atlanta, Georgia, USA
    Farooq Amin
  52. Department of Psychiatry and Behavioral Sciences, Atlanta Veterans Affairs Medical Center, Atlanta, Georgia, USA
    Farooq Amin
  53. Department of Psychiatry, University of Iowa Carver College of Medicine, Iowa City, Iowa, USA
    Donald W Black
  54. Institute of Human Genetics, University of Aarhus, Aarhus, Denmark
    Anders D Børglum & Ditte Demontis
  55. Centre for Psychiatric Research, Aarhus University Hospital, Risskov, Denmark
    Anders D Børglum & Ole Mors
  56. University of Queensland Diamantina Institute, Princess Alexandra Hospital, University of Queensland, Brisbane, Queensland, Australia
    Matthew A Brown & Patrick A Danoy
  57. Department of Psychiatry, University Medical Center Groningen, University of Groningen, Groningen, The Netherlands
    Richard Bruggeman & Durk Wiersma
  58. School of Nursing, Louisiana State University Health Sciences Center, New Orleans, Louisiana, USA
    Nancy G Buccola
  59. Department of Psychiatry, University of California at San Francisco, San Francisco, California, USA
    William F Byerley
  60. NCIRE (Northern California Institute for Research and Education), San Francisco, California, USA
    William F Byerley
  61. Department of Psychiatry, Rudolf Magnus Institute of Neuroscience, University Medical Center Utrecht, Utrecht, The Netherlands
    Wiepke Cahn & René S Kahn
  62. School of Psychiatry, University of New South Wales and Schizophrenia Research Institute, Sydney, New South Wales, Australia
    Vaughan J Carr
  63. Department of Psychiatry, University of Queensland, Royal Brisbane Hospital, Brisbane, Australia
    Stanley V Catts
  64. Department of Psychiatry, Washington University, St. Louis, Missouri, USA
    C Robert Cloninger
  65. Department of Psychiatry, Academic Medical Centre, University of Amsterdam, Amsterdam, The Netherlands
    Lieuwe de Haan & Don H Linszen
  66. Department of Psychiatry, University of Athens Medical School, Athens, Greece
    Dimitris Dikeos & George Papadimitriou
  67. Department of Medical Genetics, Oslo University Hospital, Oslo, Norway
    Srdjan Djurovic
  68. Wellcome Trust Centre for Human Genetics, Oxford, UK
    Peter Donnelly, Chris C A Spencer & Amy Strange
  69. Department of Statistics, University of Oxford, Oxford, UK
    Peter Donnelly
  70. Mental Health Center Copenhagen, Copenhagen University Hospital, Copenhagen, Denmark
    Anders Fink-Jensen
  71. Department of Psychiatry, University of Colorado Denver, Aurora, Colorado, USA
    Robert Freedman & Ann Olincy
  72. Center for Clinical Intervention and Neuropsychiatric Schizophrenia Research, Mental Health Center Glostrup, Copenhagen University Hospital, Glostrup, Denmark
    Birte Glenthøj
  73. INSERM, Institut de Myologie, Hôpital de la Pitié-Salpêtrière, Paris, France
    Stephanie Godard
  74. Illumina, Inc., La Jolla, California, USA
    Mark Hansen
  75. School of Electrical Engineering and Computing Science, University of Newcastle, Newcastle, New South Wales, Australia
    Frans A Henskens
  76. Department of Clinical Chemistry and Immunology, Section of Neonatal Screening and Hormones, The State Serum Institute, Copenhagen, Denmark
    David M Hougaard
  77. Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden
    Christina M Hultman, Paul Lichtenstein & Patrick F Sullivan
  78. Centre for Clinical Research in Neuropsychiatry, School of Psychiatry and Clinical Neurosciences, The University of Western Australia, Perth, Western Australia, Australia
    Assen V Jablensky
  79. Department of Child and Adolescent Psychiatry, Pierre and Marie Curie Faculty of Medicine, Paris, France
    Maurice Jay & Claudine Laurent
  80. Department of Clinical Pharmacology, Bispebjerg University Hospital, Copenhagen, Denmark
    Gesche Jürgens
  81. Department of Psychology, University of Colorado, Boulder, Boulder, Colorado, USA
    Matthew C Keller
  82. Department of Psychiatry and Psychology, School of Mental Health and Neuroscience, European Graduate School of Neuroscience (EURON), South Limburg Mental Health Research and Teaching Network (SEARCH), Maastricht University Medical Centre, Maastricht, The Netherlands
    Gunter Kenis & Ruud van Winkel
  83. Department of Genetics, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, USA
    Yunjung Kim & Patrick F Sullivan
  84. Maastricht University Medical Centre, South Limburg Mental Health Research and Teaching Network, Maastricht, The Netherlands
    Lydia Krabbendam, Inez Myin-Germeys & Jim van Os
  85. Department of Psychiatry and Behavioral Sciences, Johns Hopkins University School of Medicine, Baltimore, Maryland, USA
    Virginia K Lasseter, Gerald Nestadt & Ann E Pulver
  86. Department of Psychiatry, Hadassah-Hebrew University Medical Center, Jerusalem, Israel
    F Bernard Lerer
  87. Department of Biostatistics, Johns Hopkins University Bloomberg School of Public Health, Baltimore, Maryland, USA
    Kung-Yee Liang
  88. Department of Psychiatry, Columbia University, New York, New York, USA
    Jeffrey A Lieberman & T Scott Stroup
  89. Department of Mental Health and Substance Abuse Services, National Institute for Health and Welfare, Helsinki, Finland
    Jouko Lönnqvist & Jaana Suvisaari
  90. Schizophrenia Research Institute, Sydney and Centre for Brain and Mental Health Research, University of Newcastle, Newcastle, New South Wales, Australia
    Carmel M Loughland
  91. Department of Psychiatry, University of Bonn, Bonn, Germany
    Wolfgang Maier & Marcella Rietschel
  92. Laboratoire de Génétique Moléculaire de la Neurotransmission et des Processus Neurodégénératifs, Centre National de la Recherche Scientifique, Hôpital de la Pitié Salpêtrière, Paris, France
    Jacques Mallet
  93. Institute of Medical Biometry, Informatics and Epidemiology (IMBIE), University of Bonn, Bonn, Germany
    Manuel Mattheisen
  94. Department of Research, Sørlandet Hospital, Kristiansand, Norway
    Morten Mattingsdal
  95. Queensland Centre for Mental Health Research, The Park Centre for Mental Health, Wacol, Queensland, Australia
    Duncan E McLean
  96. Functional NeuroImaging Laboratory, School of Psychology, University of Newcastle, Sydney, New South Wales, Australia
    Patricia T Michie
  97. Schizophrenia Research Institute, Sydney, New South Wales, Australia
    Patricia T Michie & Ulrich Schall
  98. Department of Psychiatry, First Psychiatric Clinic, Alexander University Hospital, Sofia, Bulgaria
    Vihra Milanova
  99. National Centre for Register-Based Research, University of Aarhus, Aarhus, Denmark
    Preben B Mortensen
  100. Department of Psychiatry, University of Toronto, Toronto, Canada
    Pierandrea Muglia
  101. NeuroSearch A/S, Ballerup, Denmark
    Pierandrea Muglia
  102. Unit for Psychiatric Research, Aalborg Psychiatric Hospital, Aalborg, Denmark
    Jimmi Nielsen
  103. Psychiatric Centre Copenhagen, Copenhagen University Hospital, Copenhagen, Denmark
    Merete Nordentoft
  104. Department of Psychiatry, Queens University, Belfast, Ireland
    F Anthony O'Neill
  105. ARoS Applied Biotechnology A/S, Skejby, Denmark
    Torben F Ørntoft
  106. Department of Molecular Medicine, Aarhus University Hospital, Skejby, Denmark
    Torben F Ørntoft
  107. Department of Psychiatry, Melbourne Neuropsychiatry Centre, The University of Melbourne and Melbourne Health, Melbourne, Victoria, Australia
    Christos Pantelis
  108. Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, Cambridge, UK
    Leena Peltonen
  109. Department of Psychiatry, National University Hospital, Reykjavik, Iceland
    Hannes Petursson & Engilbert Sigurdsson
  110. Strathclyde Institute of Pharmacy and Biomedical Sciences, The John Arbuthnott Building, University of Strathclyde, Glasgow, UK
    Ben Pickard
  111. Department of Psychiatry, University of Oxford, Warneford Hospital, Headington, Oxford, UK
    Digby Quested
  112. Department of Genetic Epidemiology in Psychiatry, Central Institute of Mental Health, University of Heidelberg, Mannheim, Germany
    Marcella Rietschel & Thomas G Schulze
  113. Priority Centre for Brain and Mental Health Research, University of Newcastle, Sydney, New South Wales, Australia
    Ulrich Schall
  114. Department of Psychiatry and Psychotherapy, Georg-August-University, Göttingen, Germany
    Thomas G Schulze
  115. School of Psychiatry and Clinical Neurosciences, University of Western Australia, Perth, Western Australia, Australia
    Sibylle G Schwab & Dieter B Wildenauer
  116. Department of Psychiatry, University of Erlangen-Nuremberg, Erlangen, Germany
    Sibylle G Schwab
  117. Centre for Medical Research, Western Australian Institute for Medical Research, University of Western Australia, Perth, Western Australia, Australia
    Sibylle G Schwab
  118. Centre for Information Based Medicine, University of Newcastle, Hunter Medical Research Institute, Newcastle and Schizophrenia Research Institute, Sydney, New South Wales, Australia
    Rodney J Scott
  119. Division of Cancer Epidemiology and Genetics, National Cancer Institute, Bethesda, Maryland, USA
    Jianxin Shi
  120. Department of Psychiatry, University of Iceland, Reykjavik, Iceland
    Engilbert Sigurdsson
  121. Department of Psychiatry, Veterans Affairs Medical Center, New York, New York, USA
    Jeremy M Silverman
  122. West Berkshire National Health Service (NHS) Trust, Reading, UK
    Srinivasa Thirumalai
  123. Mental Health Center Frederiksberg, Copenhagen University Hospital, Copenhagen, Denmark
    Sally Timm
  124. Department of Medical Genetics, University Hospital Maichin Dom, Sofia, Bulgaria
    Draga Toncheva
  125. Department of Pharmacy, Virginia Commonwealth University, Richmond, Virginia, USA
    Edwin van den Oord
  126. Department of Neurology, Rudolf Magnus Institute of Neuroscience, Universitair Medisch Centrum (UMC) Utrecht, Utrecht, The Netherlands
    Jan Veldink
  127. The Health Research Board, Dublin, Ireland
    Dermot Walsh
  128. Mental Health Center Amager, Copenhagen University Hospital, Copenhagen, Denmark
    August G Wang
  129. Centre for Clinical Research in Neuropsychiatry, Graylands Hospital, Mt Claremont, Western Australia, Australia
    Dieter B Wildenauer
  130. Department of Psychiatry, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, USA
    Patrick F Sullivan
  131. Department of Epidemiology, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina, USA
    Patrick F Sullivan

Consortia

The Schizophrenia Psychiatric Genome-Wide Association Study (GWAS) Consortium

Contributions

The Schizophrenia Psychiatric Genome-Wide Association Study (GWAS) Consortium (PGC): overall coordination: P.V.G. Coordination of statistical analyses: M.J.D. Coordination of phenotypic analyses: K.S.K. Statistical analyses: S.R., M.J.D., P.A.H., D.-Y.L., S.P., F.D., B.M.N., L.R., P.M.V., D.P., D.M.R. Manuscript preparation: P.V.G. (primary), M.J.D. (primary), A.R.S. (primary), S.R. (primary), M.C.O. (primary), K.S.K., D.F.L., P.S., P.A.H., P.F.S. (primary), D.-Y.L., J.D., R.A.O., O.A.A., E. Scolnick. Phenotypic analyses: K.S.K., A.F., A.C., R.L.A. Stage 1 GWAS sample 1–Cardiff, UK: M.C.O., N.C., P.A.H., M. Hamshere, H.J.W., V. Moskvina, S. Dwyer, L.G., S.Z., M.J.O. Stage 1 GWAS sample 2–Clinical Antipsychotic Trials of Intervention Effectiveness (CATIE): P.F.S., D.-Y.L., E.v.d.O., Y.K., T.S.S., J.A.L. Stage 1 GWAS sample 3–International Schizophrenia Consortium (ISC)–Aberdeen: D.St.C. Stage 1 GWAS sample 4–ISC–Cardiff: G.K.K., M.C.O., P.A.H., L.G., I.N., H.J.W., D.T., V. Milanova, M.J.O. Stage 1 GWAS sample 5–ISC–Dublin: D.W.M., C.T.O., E.K., E.M.Q., M.G., A.C. Stage 1 GWAS sample 6–ISC–Edinburgh: D.H.R.B., K.A.M., B.P., P. Malloy, A.W.M., A. McIntosh. Stage 1 GWAS sample 7–ISC–London: A. McQuillin, K.C., S. Datta, J.P., S. Thirumalai, V.P., R.K., J. Lawrence, D.Q., N.B., H.G. Stage 1 GWAS sample 8–ISC–Portugal: M.T.P., C.N.P., A.F. Stage 1 GWAS sample 9–ISC–SW1–Sweden, stage 1 GWAS sample 10–ISC–SW2–Sweden, stage 2 replication follow-up sample 16–SW3–Sweden, stage 2 replication follow-up sample 17–SW4–Sweden: C.M.H., P.L., S.E.B., S.P., E. Scolnick, P.S., P.F.S. Stage 1 GWAS sample 11–Molecular Genetics of Schizophrenia (MGS): J. Shi, D.F.L., J.D., A.R.S., M.C.K., B.J.M., A.O., F.A., C.R.C., J.M.S., N.G.B., W.FB., D.W.B., K.S.K., R.F., P.V.G. Stage 1 GWAS sample 12–Schizophrenia Genetics Consortium (SGENE)–Bonn: S.C., M. Rietschel, M.M.N., W.M., T.G.S., M. Mattheisen. Stage 1 GWAS sample 13–SGENE–Copenhagen, stage 2 replication follow-up sample 5–SGENE–Copenhagen: T.H., A.I., K.D.J., L.D., G.J., H.B.R., B.G., J.N., S. Timm, L.O., A.G.W., A.F.-J., J.H.T., T.W. Stage 1 GWAS sample 14–SGENE–Munich, stage 2 replication follow-up sample 12–SGENE–Munich, stage 2 replication follow-up sample 13–SGENE–Munich: I.G., A.M.H., H.K., M.F., B.K., P. Muglia, D.R. Stage 1 GWAS sample 15–SGENE–Thematic Organized Psychoses Research 3 (TOP3): S. Djurovic, M. Mattingsdal, I.A., I.M., O.A.A. Stage 1 GWAS sample 16–SGENE–UCLA: R.A.O., R.M.C., N.B.F., R.S.K., D.H.L., J.v.O., D. Wiersma, R.B., W.C., L.d.H., L.K., I.M.-G., E. Strengman. Stage 1 GWAS sample 1–Zucker Hillside A.K.M., T.L.Stage 2 replication follow-up sample –multicenter pedigree P.A.H., B.P.R., A.E.P., M.J.O., D.B.W., P.V.G., B.J.M., C.L., K.S.K., G.N., N.M.W., S.G.S., A.R.S., M. Hansen, D.A.N., J.M., B.W., V.K.L., M.C.O., J.D., M. Albus, M. Alexander, S.G., R.R., K.-Y.L., N.N., W.M., G.P., D. Walsh, M.J., F.A.O., F.B.L., D. Dikeos, J.M.S., D.F.L.Stage 2 replication follow-up sample –SGEN–Aarhus A.D.B., D. Demontis, P.B.M., D.M.H., T.FØ., O.M.Stage 2 replication follow-up sample –SGEN–Aarhus O.M., M.N., A.D.B.Stage 2 replication follow-up sample –SGEN–Belgium R.v.W., G.K., M.D.H., J.V.Stage 2 replication follow-up sample –SGEN–Iceland H.S., S.S., E. Sigurdsson, H.P., K.S.Stage 2 replication follow-up sample –SGEN–England D.A.C.Stage 2 replication follow-up sample –SGEN–Helsinki, stage 2 replication follow-up sample 1–SGEN–Kuusamo L.P., O.P.H.P., J. Suvisaari, J. önnqvist.Stage 2 replication follow-up sample –SGEN–Hungary I.B., J.M.R.Stage 2 replication follow-up sample 1–SGEN–Italy M. Ruggeri, S. Tosato.Stage 2 replication follow-up sample 1–SGEN–Russia V.G.Stage 2 replication follow-up sample 1–SGEN–Sweden E.G.J., I.A., L.T.Stage 2 replication follow-up sample 1–University of Queensland B.J.M., M.A.B., P.A.D., J.J.M., D.E.M.Stage 2 replication follow-up sample 1–Australian Schizophrenia Research Bank 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., U.S.Stage 2 replication follow-up sample 1–Irish Schizophrenia Genomics Consortium (ISGC) A.C., D.W.M., P.C., B.S.M., C.T.O., G.D., F.A.O., M.G., K.S.K., B.P.R., ISGC (see the Acknowledgments in theSupplementary Not for additional contributors not listed above).Stage 2 replication follow-up sample 1–Wellcome Trust Case Control Consortium 2 (WTCCC2) P.D. (Chair of Management Committee; Data and Analysis Group), C.C.A.S. (Data and Analysis Group; Publications Committee), A.S. (Data and Analysis Group), WTCCC2 (see Acknowledgments in theSupplementary Not for additional contributors not listed above). All authors contributed to the current version of the paper

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Correspondence toPablo V Gejman.

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Competing interests

Eli Lilly funded portions of the genotyping for CATIE and TOP. P.F.S. received research funding from Eli Lilly in connection with CATIE. T.S.S. received research funding from Eli Lilly and consulting fees from Janssen Pharmaceutica, GlaxoSmithKline and Bristol-Myers Squibb. J.A.L. received research funding from AstraZeneca Pharmaceuticals, Bristol-Myers Squibb, GlaxoSmithKline, Janssen Pharmaceutica and Pfizer and consulting and educational fees from AstraZeneca Pharmaceuticals, Bristol-Myers Squibb, Eli Lilly, Forest Pharmaceuticals, GlaxoSmithKline, Janssen Pharmaceutica, Novartis, Pfizer and Solvay. D.St.C. received research funding from GlaxoSmithKline and Generation Scotland, Genetics Health Initiative. F.A. received funds from Pfizer, Organon and the Foundation for the National Institutes of Health. D.W.B. has received research support from Shire and Forest, has been on the speakers' bureau for Pfizer and has received consulting honoraria from Forest and Jazz. T.W. has received consulting and lecture fees from H. Lundbeck A/S. O.A.A. has received Speaker's honorarium from AstraZeneca, Janssen, Bristol-Myers Squibb and GlaxoSmithKline. I.M. has received a Speaker's honorarium from Janssen and AstraZeneca. A.K.M. has received consulting fees or honoraria from Eli Lilly & Company, Janssen Pharmaceutica, Merck, Bristol-Meyers Squibb, Pfizer, PGxHealth (a division of Clinical Data, Inc.), Roche Diagnostics and Vanda Pharmaceuticals and has received research support from Eli Lilly & Company. T.L. has received consulting fees or honoraria from Merck, Eli Lilly & Company, Golden Helix, Inc., InforMed Insights and PGxHealth (a division of Clinical Data, Inc.). I.B. has been an advisory board member, consultant and lecturer for AstraZeneca, Bristol-Myers Squibb, Eli Lilly, EGIS, Janssen, H. Lundbeck A/S, Novartis, Pfizer, Richter and Schering-Plough and received a grant for an investigator-initiated study from H. Lundbeck A/S. J.J.M. has received consulting and speaker's fees from Johnson & Johnson, Schering-Plough and Eli Lilly. C.P. has received grant support from Janssen-Cilag, Eli Lilly, Hospira (Mayne) and AstraZeneca, provided consultancy to Janssen-Cilag, Eli Lilly, Hospira (Mayne), AstraZeneca, Pfizer and Schering-Plough and has undertaken investigator-initiated studies supported by Eli Lilly, Hospira, Janssen Cilag and AstraZeneca. The Denmark-Aarhus group (The GEMS Stud with principal investigators A.D.B., O.M. and P.B.M.) received research funding from H. Lundbeck A/S. E.G.J. has served as an unpaid consultant for Eli Lilly.

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The Schizophrenia Psychiatric Genome-Wide Association Study (GWAS) Consortium. Genome-wide association study identifies five new schizophrenia loci.Nat Genet 43, 969–976 (2011). https://doi.org/10.1038/ng.940

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