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Common genetic variants influence human subcortical brain structures

Nature volume 520, pages 224–229 (2015)Cite this article

Subjects

Abstract

The highly complex structure of the human brain is strongly shaped by genetic influences1. Subcortical brain regions form circuits with cortical areas to coordinate movement2, learning, memory3 and motivation4, and altered circuits can lead to abnormal behaviour and disease2. To investigate how common genetic variants affect the structure of these brain regions, here we conduct genome-wide association studies of the volumes of seven subcortical regions and the intracranial volume derived from magnetic resonance images of 30,717 individuals from 50 cohorts. We identify five novel genetic variants influencing the volumes of the putamen and caudate nucleus. We also find stronger evidence for three loci with previously established influences on hippocampal volume5 and intracranial volume6. These variants show specific volumetric effects on brain structures rather than global effects across structures. The strongest effects were found for the putamen, where a novel intergenic locus with replicable influence on volume (rs945270; P = 1.08 × 10−33; 0.52% variance explained) showed evidence of altering the expression of the KTN1 gene in both brain and blood tissue. Variants influencing putamen volume clustered near developmental genes that regulate apoptosis, axon guidance and vesicle transport. Identification of these genetic variants provides insight into the causes of variability in human brain development, and may help to determine mechanisms of neuropsychiatric dysfunction.

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Figure 1: Common genetic variants associated with subcortical volumes and the ICV.
Figure 2: Effect of rs945270 on KTN1 expression and putamen shape.

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Acknowledgements

Funding sources for contributing sites and acknowledgments of contributing consortia authors can be found in Supplementary Note 3.

Author information

Author notes

  1. Derrek P. Hibar, Jason L. Stein, Miguel E. Renteria, Alejandro Arias-Vasquez and Sylvane Desrivières: These authors contributed equally to this work.

  2. Nicholas G. Martin, Margaret J. Wright, Gunter Schumann, Barbara Franke, Paul M. Thompson and Sarah E. Medland: These authors jointly supervised this work.

Authors and Affiliations

  1. Imaging Genetics Center, Institute for Neuroimaging & Informatics, Keck School of Medicine of the University of Southern California, Los Angeles, 90292, California, USA

    Derrek P. Hibar, Jason L. Stein, Neda Jahanshad, Christopher R. K. Ching, Christopher D. Whelan & Paul M. Thompson

  2. Department of Neurology, Neurogenetics Program, UCLA School of Medicine, Los Angeles, 90095, California, USA

    Jason L. Stein

  3. QIMR Berghofer Medical Research Institute, Brisbane 4006, Australia.,

    Miguel E. Renteria, Gabriel Cuellar-Partida, Lachlan T. Strike, Narelle K. Hansell, Grant W. Montgomery, Nicholas G. Martin, Margaret J. Wright & Sarah E. Medland

  4. Department of Human Genetics, Radboud university medical center, Nijmegen 6500 HB, The Netherlands.,

    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

  5. Department of Psychiatry, Radboud university medical center, Nijmegen 6500 HB, The Netherlands.,

    Alejandro Arias-Vasquez & Barbara Franke

  6. Department of Cognitive Neuroscience, Radboud university medical center, Nijmegen 6500 HB, The Netherlands.,

    Alejandro Arias-Vasquez, Janita Bralten, Jan K. Buitelaar & Guillén Fernández

  7. Donders Institute for Brain, Cognition and Behaviour, Radboud University, Nijmegen 6500 GL, The Netherlands.,

    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

  8. MRC-SGDP Centre, Institute of Psychiatry, Psychology and Neuroscience, King’s College London, London SE5 8AF, UK.,

    Sylvane Desrivières, Tianye Jia, Christine Macare & Gunter Schumann

  9. Laboratory of Human Genetics and Cognitive Functions, Institut Pasteur, Paris 75015, France.,

    Roberto Toro

  10. Centre Nationale de Recherche Scientifique (CNRS) Unité de Recherche Associée (URA) 2182 Genes, Synapses and Cognition, Institut Pasteur, Paris 75015, France.,

    Roberto Toro

  11. Université Paris Diderot, Sorbonne Paris Cité, Paris 75015, France.,

    Roberto Toro

  12. German Center for Neurodegenerative Diseases (DZNE) Rostock/Greifswald, Greifswald 17487, Germany.,

    Katharina Wittfeld & Wolfgang Hoffmann

  13. Department of Psychiatry, University Medicine Greifswald, Greifswald 17489, Germany.,

    Katharina Wittfeld, Deborah Janowitz & Hans J. Grabe

  14. Department of Psychiatry, Brain Center Rudolf Magnus, University Medical Center Utrecht, Utrecht, 3584 CX, The Netherlands.,

    Lucija Abramovic, Marc M. Bohlken, Marco P. Boks, Kristel R. van Eijk, Wiepke Cahn, René S. Kahn, Neeltje E. M. van Haren, Rachel M. Brouwer, Hilleke E. Hulshoff Pol & Roel A. Ophoff

  15. Umeå Centre for Functional Brain Imaging (UFBI), Umeå University, Umeå 901 87, Sweden.,

    Micael Andersson, Alireza Salami, Lars G. Nilsson & Lars Nyberg

  16. Brain Research Imaging Centre, University of Edinburgh, Edinburgh EH4 2XU, UK.,

    Benjamin S. Aribisala, Natalie A. Royle, Mark E. Bastin & Joanna M. Wardlaw

  17. Department of Computer Science, Lagos State University, Lagos, Nigeria

    Benjamin S. Aribisala

  18. Department of Neuroimaging Sciences, Scottish Imaging Network, A Platform for Scientific Excellence (SINAPSE) Collaboration, University of Edinburgh, Edinburgh EH4 2XU, UK.,

    Benjamin S. Aribisala, Natalie A. Royle, Mark E. Bastin & Joanna M. Wardlaw

  19. Centre for Healthy Brain Ageing, School of Psychiatry, University of New South Wales, Sydney 2052, Australia.,

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

  20. School of Mathematics and Statistics, University of Sydney, Sydney 2006, Australia.,

    Nicola J. Armstrong

  21. The Hospital for Sick Children, University of Toronto, Toronto M5G 1X8, Canada.,

    Manon Bernard, Jean Shin & Zdenka Pausova

  22. NORMENT - KG Jebsen Centre, Institute of Clinical Medicine, University of Oslo, Oslo N-0316, Norway.,

    Andrew A. Brown, Lavinia Athanasiu, Cecilie B. Hartberg, Unn K. Haukvik, Ingrid Melle, Srdjan Djurovic, Ingrid Agartz, Ole A. Andreassen & Erik G. Jönsson

  23. Division of Mental Health and Addiction, NORMENT - KG Jebsen Centre, Oslo University Hospital, Oslo 0424, Norway.,

    Andrew A. Brown, Lars T. Westlye, Lavinia Athanasiu, Ingrid Melle, Thomas Espeseth & Ole A. Andreassen

  24. Cerebral Imaging Centre, Douglas Mental Health University Institute, Montreal H4H 1R3, Canada.,

    M. Mallar Chakravarty

  25. Department of Psychiatry and Biomedical Engineering, McGill University, Montreal H3A 2B4, Canada.,

    M. Mallar Chakravarty

  26. Lieber Institute for Brain Development, Baltimore, 21205, Maryland, USA

    Qiang Chen, Aaron L. Goldman, Venkata S. Mattay & Daniel R. Weinberger

  27. Interdepartmental Neuroscience Graduate Program, UCLA School of Medicine, Los Angeles, 90095, California, USA

    Christopher R. K. Ching

  28. Biological Psychology, Neuroscience Campus Amsterdam & EMGO Institute for Health and Care Research, VU University & VU Medical Center, Amsterdam 1081 BT, The Netherlands.,

    Anouk den Braber, Iryna O. Fedko, Jouke-Jan Hottenga, Dennis van 't Ent, Dorret I. Boomsma & Eco J. C. de Geus

  29. Department of Clinical Science, NORMENT - KG Jebsen Centre for Psychosis Research, University of Bergen, 5021 Bergen, Norway.,

    Sudheer Giddaluru, Vidar M. Steen & Stephanie Le Hellard

  30. Dr. Einar Martens Research Group for Biological Psychiatry, Center for Medical Genetics and Molecular Medicine, Haukeland University Hospital, Bergen 5021, Norway.,

    Sudheer Giddaluru, Vidar M. Steen & Stephanie Le Hellard

  31. Central Institute of Mental Health, Medical Faculty Mannheim, University Heidelberg, Mannheim 68159, Germany.,

    Oliver Grimm, Marcella Rietschel & Andreas Meyer-Lindenberg

  32. Language and Genetics Department, Max Planck Institute for Psycholinguistics, Nijmegen 6525 XD, The Netherlands.,

    Tulio Guadalupe, Simon E. Fisher & Clyde Francks

  33. International Max Planck Research School for Language Sciences, Nijmegen 6525 XD, The Netherlands.,

    Tulio Guadalupe

  34. Department of Child and Adolescent Psychiatry, Faculty of Medicine of the TU Dresden, Dresden 01307 Germany.,

    Johanna Hass, Stefan Ehrlich & Esther Walton

  35. Human Genetics Branch and Experimental Therapeutics and Pathophysiology Branch, National Institute of Mental Health Intramural Research Program, Bethesda, 20892, Maryland, USA

    Girma Woldehawariat, Allison C. Nugent, Xinmin Liu, Francis J. McMahon, Wayne C. Drevets & Dara M. Cannon

  36. Department of Psychology, Yale University, New Haven, 06511, Connecticut, USA

    Avram J. Holmes

  37. Department of Psychiatry, Massachusetts General Hospital, Boston, 02115, Massachusetts, USA

    Avram J. Holmes, Phil H. Lee, Stefan Ehrlich, Randy L. Buckner, Randy L. Gollub, Joshua L. Roffman & Jordan W. Smoller

  38. Center for Neuroimaging, Radiology and Imaging Sciences, Indiana University School of Medicine, Indianapolis, 46202, Indiana, USA

    Sungeun Kim, Kwangsik Nho, Shannon L. Risacher, Li Shen & Andrew J. Saykin

  39. Center for Computational Biology and Bioinformatics, Indiana University School of Medicine, Indianapolis, 46202, Indiana, USA

    Sungeun Kim, Kwangsik Nho & Li Shen

  40. Indiana Alzheimer Disease Center, Indiana University School of Medicine, Indianapolis, 46202, Indiana, USA

    Sungeun Kim, Kwangsik Nho, Shannon L. Risacher, Li Shen, Tatiana M. Foroud & Andrew J. Saykin

  41. Department of Psychiatry and Psychotherapy, Center for Translational Research in Systems Neuroscience and Psychiatry, University Medical Center, 37075, Goettingen, Germany

    Bernd Kraemer, Maria Keil & Oliver Gruber

  42. Psychiatric and Neurodevelopmental Genetics Unit, Center for Human Genetic Research, Massachusetts General Hospital, Boston, 02115, Massachusetts, USA

    Phil H. Lee & Jordan W. Smoller

  43. Stanley Center for Psychiatric Research, Broad Institute of MIT and Harvard, Boston, 02141, Massachusetts, USA

    Phil H. Lee & Jordan W. Smoller

  44. Department of Psychiatry, Harvard Medical School, Boston, 02115, Massachusetts, USA

    Phil H. Lee

  45. Center for Neurobehavioral Genetics, University of California, Los Angeles, 90095, California, USA

    Loes M. Olde Loohuis & Roel A. Ophoff

  46. Centre for Cognitive Ageing and Cognitive Epidemiology, Psychology, University of Edinburgh, Edinburgh EH8 9JZ, UK.,

    Michelle Luciano, David C. M. Liewald, Lorna M. Lopez, Natalie A. Royle, Mark E. Bastin, Ian J. Deary, Andrew M. McIntosh & Joanna M. Wardlaw

  47. Department of Biomedicine, Aarhus University, Aarhus DK-8000, Denmark.,

    Manuel Mattheisen

  48. The Lundbeck Foundation Initiative for Integrative Psychiatric Research, iPSYCH, Aarhus and Copenhagen DK-8000, Denmark.,

    Manuel Mattheisen

  49. Center for integrated Sequencing, iSEQ, Aarhus University, Aarhus DK-8000, Denmark.,

    Manuel Mattheisen

  50. Department of Psychiatry, Neuroscience Campus Amsterdam, VU University Medical Center/GGZ inGeest, Amsterdam 1081 HL, The Netherlands.,

    Yuri Milaneschi, Lianne Schmaal, Dick J. Veltman & Brenda W. J. H. Penninx

  51. Division of Psychiatry, Royal Edinburgh Hospital, University of Edinburgh, Edinburgh EH10 5HF, UK.,

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

  52. Department of Medical and Molecular Genetics, King’s College London, London SE1 9RT, UK.,

    Adaikalavan Ramasamy, Mina Ryten & Michael E. Weale

  53. Reta Lila Weston Institute and Department of Molecular Neuroscience, UCL Institute of Neurology, London WC1N 3BG, UK.,

    Adaikalavan Ramasamy, Daniah Trabzuni, J. Raphael Gibbs, Sebastian Guelfi, Dena G. Hernandez, John Hardy & Mina Ryten

  54. Department of Psychiatry, University Hospital Marqués de Valdecilla, School of Medicine, University of Cantabria-IDIVAL, Santander 39008, Spain.,

    Roberto Roiz-Santiañez & Benedicto Crespo-Facorro

  55. Cibersam (Centro Investigación Biomédica en Red Salud Mental), Madrid 28029, Spain.,

    Roberto Roiz-Santiañez, Rocio Perez-Iglesias & Benedicto Crespo-Facorro

  56. Neuropsychiatric Genetics Research Group and Department of Psychiatry, Trinity College Institute of Psychiatry, Trinity College Dublin, Dublin 2, Ireland.,

    Emma J. Rose, Margaret Needham, Aiden Corvin, Derek W. Morris & Gary Donohoe

  57. Department of Psychiatry, Center for Translational Research on Adversity, Neurodevelopment and Substance Abuse (C-TRANS), University of Maryland School of Medicine, Baltimore, 21045, Maryland, USA

    Emma J. Rose

  58. Aging Research Center, Karolinska Institutet and Stockholm University, 11330 Stockholm, Sweden.,

    Alireza Salami

  59. Max Planck Institute of Psychiatry, Munich 80804, Germany.,

    Philipp G. Sämann, David Hoehn, Benno Pütz, Christiane Wolf, Michael Czisch, Florian Holsboer, Bertram Müller-Myhsok & Oliver Gruber

  60. Department of Neurosciences, Multimodal Imaging Laboratory, University of California, San Diego, 92093, California, USA

    Andrew J. Schork

  61. Department of Cognitive Sciences, University of California, San Diego, 92161, California, USA

    Andrew J. Schork

  62. School of Psychology, University of Queensland, Brisbane 4072, Australia.,

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

  63. Centre for Advanced Imaging, University of Queensland, Brisbane 4072, Australia.,

    Lachlan T. Strike & Katie L. McMahon

  64. Institute for Community Medicine, University Medicine Greifswald, Greifswald D-17475, Germany.,

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  65. Analytic and Translational Genetics Unit, Massachusetts General Hospital, Boston, 02114, Massachusetts, USA

    Raymond K. Walters

  66. Medical and Population Genetics Program, Broad Institute of Harvard and MIT, Boston, 02142, Massachusetts, USA

    Raymond K. Walters

  67. Department of Psychology, University of Oslo, Oslo 0373, Norway.,

    Lars T. Westlye, Ivar Reinvang & Thomas Espeseth

  68. Nuffield Department of Clinical Neurosciences, The Oxford Centre for Functional MRI of the Brain, Oxford University, Oxford OX3 9DU, UK.,

    Anderson M. Winkler

  69. Department of Psychiatry, Yale School of Medicine, New Haven, 06511, Connecticut, USA

    Anderson M. Winkler, D. Reese McKay, Emma Sprooten & David C. Glahn

  70. Department of Neurology and Neurosurgery, Montreal Neurological Institute, McGill University, Montreal H3A 2B4, Canada.,

    Saud Alhusaini

  71. Molecular and Cellular Therapeutics, The Royal College of Surgeons, Dublin 2, Ireland.,

    Norman Delanty

  72. The Athinoula A.Martinos Center for Biomedical Imaging, Massachusetts General Hospital, Charlestown, 02129, Massachusetts, USA

    Stefan Ehrlich, Randy L. Gollub, Maritza Ebling, Bruce Fischl, Douglas Greve & Louis N. Vinke

  73. Department of Psychiatric Research and Development, Diakonhjemmet Hospital, Oslo 0319, Norway.,

    Cecilie B. Hartberg & Ingrid Agartz

  74. UCL Institute of Neurology, London, United Kingdom and Epilepsy Society, London WC1N 3BG, UK.,

    Dalia Kasperaviciute & Sanjay M. Sisodiya

  75. Department of Medicine, Imperial College London, London W12 0NN, UK.,

    Dalia Kasperaviciute

  76. Department of Clinical and Experimental Epilepsy, UCL Institute of Neurology, London WC1N 3BG, UK.,

    Mar Matarin

  77. Olin Neuropsychiatric Research Center, Institute of Living, Hartford Hospital, Hartford, 06106, Connecticut, USA

    D. Reese McKay, Emma Sprooten & David C. Glahn

  78. Department of Genetics, King Faisal Specialist Hospital and Research Centre, Riyadh 11211, Saudi Arabia.,

    Daniah Trabzuni

  79. Texas Biomedical Research Institute, San Antonio, 78245, 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

  80. University of Texas Health Science Center, San Antonio, 78229, Texas, USA

    Laura Almasy, Thomas D. Dyer, Peter T. Fox & John Blangero

  81. National Ageing Research Institute, Royal Melbourne Hospital, Melbourne 3052, Australia.,

    David Ames

  82. Academic Unit for Psychiatry of Old Age, University of Melbourne, Melbourne 3101, Australia.,

    David Ames, J. Raphael Gibbs & Dena G. Hernandez

  83. Laboratory of Neurogenetics, National Institute on Aging, National Institutes of Health, Bethesda, 20892, Maryland, USA

    Sampath Arepalli, Allissa Dillman, Michael A. Nalls, Bryan J. Traynor, Mark R. Cookson & Andrew Singleton

  84. Centre for Clinical Brain Sciences, University of Edinburgh, Edinburgh EH4 2XU, UK.,

    Mark E. Bastin, Maria C. Valdés Hernández & Joanna M. Wardlaw

  85. N.I. Vavilov Institute of General Genetics, Russian Academy of Sciences, Moscow 119333, Russia.,

    Kazima B. Bulayeva

  86. Division of Medical Genetics, Department of Biomedicine, University of Basel, Basel 4055, Switzerland.,

    Sven Cichon

  87. Institute of Human Genetics, University of Bonn, Bonn, D-53127, Germany.,

    Sven Cichon, Thomas W. Mühleisen & Markus M. Nöthen

  88. Institute of Neuroscience and Medicine (INM-1), Research Centre Jülich, Jülich, D-52425, Germany.,

    Sven Cichon & Thomas W. Mühleisen

  89. Department of Genomics, Life & Brain Center, University of Bonn, Bonn D-53127, Germany.,

    Sven Cichon, Andreas Heinz, Thomas W. Mühleisen & Markus M. Nöthen

  90. Department of Psychiatry and Psychotherapy, Charité Universitätsmedizin Berlin, CCM, Berlin 10117, Germany.,

    Susanne Erk, Sebastian Mohnke, Nina Romanczuk-Seiferth, Henrik Walter & Andrew J. Saykin

  91. Clinical Research Branch, National Institute on Aging, Baltimore, 20892, Maryland, USA

    Luigi Ferrucci

  92. Department of Medical and Molecular Genetics, Indiana University School of Medicine, Indianapolis, Indiana 46202, USA.,

    Tatiana M. Foroud & Rene L. Olvera

  93. South Texas Veterans Health Care System, San Antonio, 78229, Texas, USA

    Peter T. Fox

  94. Biofunctional Imaging, Immunology Frontier Research Center, Osaka University, Osaka 565-0871, Japan.,

    Masaki Fukunaga

  95. Division of Genetics, Department of Medicine, Brigham and Women’s Hospital, Boston, 02115, Massachusetts, USA

    Robert C. Green

  96. Harvard Medical School, Boston, 02115, Massachusetts, USA

    Robert C. Green, Randy L. Gollub & Jordan W. Smoller

  97. Department of Psychiatry, University of Groningen, University Medical Center Groningen, 9713 GZ Groningen, The Netherlands.,

    Catharina A. Hartman & Pieter J. Hoekstra

  98. Institute of Diagnostic Radiology and Neuroradiology, University Medicine Greifswald, Greifswald 17475, Germany.,

    Katrin Hegenscheid & Norbert Hosten

  99. Departments of Cognitive and Clinical Neuropsychology, VU University Amsterdam, 1081 BT Amsterdam, The Netherlands,

    Dirk J. Heslenfeld

  100. Interfaculty Institute for Genetics and Functional Genomics, University Medicine Greifswald, Greifswald 17489, Germany.,

    Georg Homuth

  101. Department of Psychiatry, Fujita Health University School of Medicine, Toyoake 470-1192, Japan.,

    Masashi Ikeda

  102. Radiology, Mayo Clinic, Rochester, 55905, Minnesota, USA

    Clifford R. Jack Jr

  103. FMRIB Centre, University of Oxford, Oxford OX3 9DU, UK.,

    Mark Jenkinson & Thomas E. Nichols

  104. NICHD Brain and Tissue Bank for Developmental Disorders, University of Maryland Medical School, Baltimore, 21201, Maryland, USA

    Robert Johnson & Ronald H. Zielke

  105. School of Psychology, University of Sussex, Brighton BN1 9QH, UK.,

    Ryota Kanai

  106. Institute of Cognitive Neuroscience, University College London, London WC1N 3AR, UK.,

    Ryota Kanai

  107. Department of Psychiatry, Maryland Psychiatric Research Center, University of Maryland, Baltimore, 21201, Maryland, USA

    Peter Kochunov

  108. Neuroscience Research Australia, Sydney 2031, Australia.,

    John B. Kwok & Peter R. Schofield

  109. School of Medical Sciences, UNSW, Sydney 2052, Australia.,

    John B. Kwok & Peter R. Schofield

  110. Department of Pathology and Cell Biology, Columbia University Medical Center, 10032, New York, USA

    Xinmin Liu

  111. Lymphocyte Cell Biology Unit, Laboratory of Genetics, National Institute on Aging, National Institutes of Health, Baltimore, 21224, Maryland, USA

    Dan L. Longo

  112. Department of Psychiatry, Ludwig-Maximilians-Universität, Munich 80336, Germany.,

    Eva Meisenzahl & Dan Rujescu

  113. Department of Statistics & WMG, University of Warwick, Coventry CV4 7AL, UK.,

    Thomas E. Nichols

  114. Department of Psychiatry, Osaka University Graduate School of Medicine, Osaka 565-0871, Japan.,

    Kazutaka Ohi

  115. Institute of Psychiatry, King’s College London, London SE5 8AF, UK.,

    Rocio Perez-Iglesias

  116. Department of Neurology, University of Calgary, Calgary T2N 2T9, Canada.,

    G. Bruce Pike

  117. Department of Clinical Neuroscience, University of Calgary, Calgary T2N 2T9, Canada.,

    G. Bruce Pike

  118. Psychiatry and Human Behavior, University of California, Irvine, 92617, California, USA

    Steven G. Potkin

  119. Department of Neurology, Beth Israel Deaconess Medical Center, Boston, 02215, Massachusetts, USA

    Glenn D. Rosen

  120. Harvard Medical School, Boston, 02115, Massachusetts, USA

    Glenn D. Rosen

  121. Department of General Psychiatry, Heidelberg University Hospital, Heidelberg 69115, Germany.,

    Knut Schnell

  122. Department of Neuropathology, MRC Sudden Death Brain Bank Project, University of Edinburgh, Edinburgh EH8 9AG, UK.,

    Colin Smith

  123. Laboratory of Neuro Imaging, Institute for Neuroimaging and Informatics, Keck School of Medicine of the University of Southern California, Los Angeles, 90033, California, USA

    Arthur W. Toga

  124. Department of Pathology, Johns Hopkins University, Baltimore, 21287, Maryland, USA

    Juan Troncoso

  125. Psychology Department and Neuroscience Institute, Georgia State University, Atlanta, 30302, Georgia, USA

    Jessica A. Turner

  126. Genentech, South San Francisco, 94080, California, USA

    Marcel van der Brug

  127. Psychiatry and Leiden Institute for Brain and Cognition, Leiden University Medical Center, Leiden 2333 ZA, The Netherlands.,

    Nic J. A. van der Wee

  128. Neuroimaging Centre, University of Groningen, University Medical Center Groningen, Groningen 9713 AW, The Netherlands.,

    Marie-Jose van Tol

  129. Department of Psychiatry, Carver College of Medicine, University of Iowa, Iowa City, 52242, Iowa, USA

    Thomas H. Wassink

  130. Department of Neurobiology, Care Sciences and Society, Karolinska Institutet, Stockholm SE-141 83, Sweden.,

    Eric Westman

  131. Behavioral Epidemiology Section, National Institute on Aging Intramural Research Program, Baltimore, 20892, Maryland, USA

    Alan B. Zonderman

  132. Faculty of Life Sciences, University of Manchester, Manchester M13 9PT, UK.,

    David G. Ashbrook & Reinmar Hager

  133. Center for Integrative and Translational Genomics, University of Tennessee Health Science Center, Memphis, 38163, Tennessee, USA

    Lu Lu & Robert W. Williams

  134. Department of Genetics, Genomics, and Informatics, University of Tennessee Health Science Center, Memphis, 38163, Tennessee, USA

    Lu Lu & Robert W. Williams

  135. Jiangsu Province Key Laboratory for Inflammation and Molecular Drug Target, Medical College of Nantong University, Nantong, 226001, China

    Lu Lu

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

    Derek W. Morris & Gary Donohoe

  137. Department of Clinical Genetics, Maastricht University Medical Center, Maastricht 6200 MD, The Netherlands.,

    Han G. Brunner

  138. Department of Psychology, Center for Brain Science, Harvard University, Boston, 02138, Massachusetts, USA

    Randy L. Buckner

  139. Karakter Child and Adolescent Psychiatry, Radboud university medical center, Nijmegen 6500 HB, The Netherlands.,

    Jan K. Buitelaar

  140. The Mind Research Network & LBERI, Albuquerque, 87106, New Mexico, USA

    Vince D. Calhoun

  141. Department of ECE, University of New Mexico, Albuquerque, 87131, New Mexico, USA

    Vince D. Calhoun

  142. Center for Translational Imaging and Personalized Medicine, University of California, San Diego, 92093, California, USA

    Anders M. Dale

  143. Departments of Neurosciences, Radiology, Psychiatry, and Cognitive Science, University of California, San Diego, 92093, California, USA

    Anders M. Dale

  144. Avera Institute for Human Genetics, Sioux Falls, 57108, South Dakota, USA

    Gareth E. Davies

  145. Neurology Division, Beaumont Hospital, Dublin 9, Ireland.,

    Norman Delanty

  146. Department of Neurology, Hopital Erasme, Universite Libre de Bruxelles, Brussels 1070, Belgium.,

    Chantal Depondt & Massimo Pandolfo

  147. Department of Medical Genetics, Oslo University Hospital, Oslo 0450, Norway.,

    Srdjan Djurovic

  148. Janssen Research & Development, Johnson & Johnson, Titusville, 08560, New Jersey, USA

    Wayne C. Drevets

  149. Department of Psychiatry, University of Iowa, Iowa City, 52242, Iowa, USA

    Beng-Choon Ho

  150. Munich Cluster for Systems Neurology (SyNergy), Munich 81377, Germany.,

    Bertram Müller-Myhsok

  151. University of Liverpool, Institute of Translational Medicine, Liverpool L69 3BX, UK.,

    Bertram Müller-Myhsok

  152. Institute of Clinical Chemistry and Laboratory Medicine, University Medicine Greifswald, Greifswald 17475, Germany.,

    Matthias Nauck

  153. Center for Imaging of Neurodegenerative Disease, San Francisco VA Medical Center, University of California, San Francisco, 94121, California, USA

    Michael W. Weiner

  154. Department of Child and Adolescent Psychiatry, Erasmus University Medical Centre, Rotterdam 3000 CB, The Netherlands.,

    Tonya White

  155. Department of Radiology, Erasmus University Medical Centre, Rotterdam 3015 CN, The Netherlands.,

    Tonya White, Hieab H. H. Adams & M. Arfan Ikram

  156. Department of Clinical Neuroscience, Psychiatry Section, Karolinska Institutet, Stockholm SE-171 76, Sweden.,

    Ingrid Agartz & Erik G. Jönsson

  157. Clinical Neuroimaging Laboratory, College of Medicine, Nursing and Health Sciences, National University of Ireland Galway, Galway, Ireland

    Dara M. Cannon & Colm McDonald

  158. Department of Psychiatry and Psychotherapy, HELIOS Hospital Stralsund 18435, Germany.,

    Hans J. Grabe

  159. Molecular Research Center for Children’s Mental Development, United Graduate School of Child Development, Osaka University, Osaka 565-0871, Japan.,

    Ryota Hashimoto

  160. Medical University of Lodz, Lodz 90-419, Poland.,

    Iwona Kloszewska

  161. Department of Psychiatry, University of Oxford, Oxford OX3 7JX, UK.,

    Simon Lovestone

  162. NIHR Dementia Biomedical Research Unit, King’s College London, London SE5 8AF, UK.,

    Simon Lovestone

  163. Department of Neurology, Johns Hopkins University School of Medicine, Baltimore, 21205, Maryland, USA

    Venkata S. Mattay

  164. Department of Medicine, Section of Gerontology and Geriatrics, University of Perugia, Perugia 06156, Italy.,

    Patrizia Mecocci

  165. Rotman Research Institute, University of Toronto, Toronto M6A 2E1, Canada.,

    Tomas Paus

  166. Departments of Psychology and Psychiatry, University of Toronto, Toronto M5T 1R8, Canada.,

    Tomas Paus

  167. Departments of Physiology and Nutritional Sciences, University of Toronto, Toronto M5S 3E2, Canada.,

    Zdenka Pausova

  168. Neuropsychiatric Institute, Prince of Wales Hospital, Sydney 2031, Australia.,

    Perminder S. Sachdev

  169. Department of Neuroimaging, Institute of Psychiatry, King’s College London, London SE5 8AF, UK.,

    Andy Simmons

  170. Biomedical Research Centre for Mental Health, King’s College London, London SE5 8AF, UK.,

    Andy Simmons

  171. Biomedical Research Unit for Dementia, King’s College London, London SE5 8AF, UK.,

    Andy Simmons

  172. Institute of Clinical Medicine, Neurology, University of Eastern Finland, Kuopio FI-70211, Finland.,

    Hilkka Soininen

  173. Neurocentre Neurology, Kuopio University Hospital, Kuopio FI-70211, Finland.,

    Hilkka Soininen

  174. Departments of Psychiatry, Neurology, Neuroscience and the Institute of Genetic Medicine, Johns Hopkins University School of Medicine, Baltimore, 21205, Maryland, USA

    Daniel R. Weinberger

  175. Department of Epidemiology, Erasmus University Medical Centre, Rotterdam 3015 CN, The Netherlands.,

    Hieab H. H. Adams, Sven J. van der Lee, Cornelia M. van Duijn & M. Arfan Ikram

  176. Laboratory of Epidemiology and Population Sciences, Intramural Research Program, National Institute on Aging, Bethesda, 20892, Maryland, USA

    Lenore J. Launer

  177. Department of Neurology, Clinical Division of Neurogeriatrics, Medical University Graz, Graz 8010, Austria.,

    Stephan Seiler & Reinhold Schmidt

  178. INSERM U897, University of Bordeaux, Bordeaux 33076, France.,

    Ganesh Chauhan

  179. Department of Neurology, Boston University School of Medicine, Boston, 02118, Massachusetts, USA

    Claudia L. Satizabal & Sudha Seshadri

  180. Framingham Heart Study, Framingham, 01702, Massachusetts, USA

    Claudia L. Satizabal

  181. Department of Neurology, School of Medicine, University of Pittsburgh, Pittsburgh, 15260, Pennsylvania, USA

    James T. Becker & Sudha Seshadri

  182. Department of Psychiatry, School of Medicine, University of Pittsburgh, Pittsburgh, 15260, Pennsylvania, USA

    James T. Becker

  183. Department of Psychology, Dietrich School of Arts and Sciences, University of Pittsburgh, Pittsburgh, 15260, Pennsylvania, USA

    James T. Becker

  184. General Internal Medicine, Johns Hopkins School of Medicine, Baltimore, 21205, Maryland, USA

    Lisa Yanek

  185. Department of Radiology, Massachusetts General Hospital, Harvard Medical School, Boston, 02114, Massachusetts, USA

    Maritza Ebling, Bruce Fischl, Douglas Greve & Louis N. Vinke

  186. Computer Science and AI Lab, Massachusetts Institute of Technology, Boston, 02141, Massachusetts, USA

    Bruce Fischl

  187. Department of Neurology University of Washington, Seattle, 98195, Washington, USA

    W. T. Longstreth Jr

  188. Institute of Molecular Biology and Biochemistry, Medical University Graz, 8010 Graz, Austria.,

    Helena Schmidt

  189. Department of Neurology, Johns Hopkins University School of Medicine, Baltimore, 21205, Maryland, USA

    Paul Nyquist

  190. Department of Biostatistics, Boston University School of Public Health, Boston, 02118, Massachusetts, USA

    Luting Xue

  191. Groupe d’Imagerie Neurofonctionnelle, UMR5296 CNRS, CEA and University of Bordeaux, Bordeaux 33076, France.,

    Bernard Mazoyer

  192. Department of Medicine, Cardiovascular Health Research Unit, University of Washington, Seattle, 98101, Washington, USA

    Joshua C. Bis

  193. Icelandic Heart Association, University of Iceland, Faculty of Medicine, Reykjavik 101, Iceland.,

    Vilmundur Gudnason

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