Article | Published:

Multiancestry genome-wide association study of 520,000 subjects identifies 32 loci associated with stroke and stroke subtypes

Nature Geneticsvolume 50pages524537 (2018) | Download Citation

Abstract

Stroke has multiple etiologies, but the underlying genes and pathways are largely unknown. We conducted a multiancestry genome-wide-association meta-analysis in 521,612 individuals (67,162 cases and 454,450 controls) and discovered 22 new stroke risk loci, bringing the total to 32. We further found shared genetic variation with related vascular traits, including blood pressure, cardiac traits, and venous thromboembolism, at individual loci (n = 18), and using genetic risk scores and linkage-disequilibrium-score regression. Several loci exhibited distinct association and pleiotropy patterns for etiological stroke subtypes. Eleven new susceptibility loci indicate mechanisms not previously implicated in stroke pathophysiology, with prioritization of risk variants and genes accomplished through bioinformatics analyses using extensive functional datasets. Stroke risk loci were significantly enriched in drug targets for antithrombotic therapy.

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Acknowledgements

A full list of Acknowledgements appears in the Supplementary Note.

Author information

Author notes

  1. A list of members and affiliations appears in the Supplementary Note.

  2. A list of members and affiliations appears at the end of the paper.

  3. These authors contributed equally: Rainer Malik, Ganesh Chauhan, Matthew Traylor, Muralidharan Sargurupremraj and Yukinori Okada.

  4. These authors jointly supervised this work: Kari Stefansson, Bradford B. Worrall, Steven J. Kittner, Sudha Seshadri, Myriam Fornage, Hugh S. Markus, Joanna M. M. Howson, Yoichiro Kamatani, Stephanie Debette and Martin Dichgans.

Affiliations

  1. Institute for Stroke and Dementia Research (ISD), University Hospital, LMU Munich, Munich, Germany

    • Rainer Malik
    • , Steffen Tiedt
    • , Martin Dichgans
    • , Rainer Malik
    • , Steffen Tiedt
    •  & Martin Dichgans
  2. Centre for Brain Research, Indian Institute of Science, Bangalore, India

    • Ganesh Chauhan
    •  & Ganesh Chauhan
  3. INSERM U1219 Bordeaux Population Health Research Center, University of Bordeaux, France

    • Matthew Traylor
    • , Loes Rutten-Jacobs
    • , Hugh S. Markus
    • , Matthew Traylor
    • , Loes Rutten-Jacobs
    •  & Hugh S. Markus
  4. Stroke Research Group, Division of Clinical Neurosciences, University of Cambridge, Cambridge, UK

    • Muralidharan Sargurupremraj
    • , Aniket Mishra
    • , Stephanie Debette
    • , Muralidharan Sargurupremraj
    • , Aniket Mishra
    • , Sara Kaffashian
    •  & Stephanie Debette
  5. Department of Neurology, Institute for Neurodegenerative Disease, Bordeaux University Hospital, Bordeaux, France

    • Muralidharan Sargurupremraj
    • , Aniket Mishra
    • , Christophe Tzourio
    • , Stephanie Debette
    • , Muralidharan Sargurupremraj
    • , Aniket Mishra
    • , Christophe Tzourio
    • , Sara Kaffashian
    •  & Stephanie Debette
  6. Laboratory for Statistical Analysis, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan

    • Yukinori Okada
    • , Masahiro Kanai
    • , Yoichiro Kamatani
    • , Yukinori Okada
    • , Masahiro Kanai
    •  & Yoichiro Kamatani
  7. Department of Statistical Genetics, Osaka University Graduate School of Medicine, Osaka, Japan

    • Yukinori Okada
    • , Masahiro Kanai
    • , Saori Sakaue
    • , Yukinori Okada
    • , Masahiro Kanai
    •  & Saori Sakaue
  8. Laboratory of Statistical Immunology, Immunology Frontier Research Center (WPI-IFReC), Osaka University, Suita, Japan

    • Yukinori Okada
    •  & Yukinori Okada
  9. Department of Neurology, Massachusetts General Hospital (MGH), Harvard Medical School, Boston, MA, USA

    • Anne-Katrin Giese
    • , Natalia S. Rost
    • , Anne-Katrin Giese
    •  & Natalia S. Rost
  10. Laboratory of Experimental Cardiology, Department of Cardiology, Division of Heart and Lungs, University Medical Center Utrecht, University of Utrecht, Utrecht, the Netherlands

    • Sander W. van der Laan
    •  & Sander W. van der Laan
  11. deCODE genetics/AMGEN Inc., Reykjavik, Iceland

    • Solveig Gretarsdottir
    • , Gudmar Thorleifsson
    • , Unnur Thorsteinsdottir
    • , Kari Stefansson
    • , Solveig Gretarsdottir
    • , Gudmar Thorleifsson
    • , Unnur Thorsteinsdottir
    •  & Kari Stefansson
  12. Center for Genomic Medicine, MGH, Boston, MA, USA

    • Christopher D. Anderson
    • , Jonathan Rosand
    • , Christopher D. Anderson
    •  & Jonathan Rosand
  13. J. Philip Kistler Stroke Research Center, Department of Neurology, MGH, Boston, MA, USA

    • Christopher D. Anderson
    • , Hakan Ay
    • , Natalia S. Rost
    • , Jonathan Rosand
    • , Christopher D. Anderson
    • , Hakan Ay
    • , Natalia S. Rost
    •  & Jonathan Rosand
  14. Program in Medical and Population Genetics, Broad Institute, Cambridge, MA, USA

    • Christopher D. Anderson
    • , Jonathan Rosand
    • , Christopher D. Anderson
    •  & Jonathan Rosand
  15. Population Health Research Institute, McMaster University, Hamilton, Ontario, Canada

    • Michael Chong
    • , Martin J. O’Donnell
    • , Salim Yusuf
    • , Guillaume Pare
    • , Michael Chong
    • , Martin J. O’Donnell
    • , Salim Yusuf
    •  & Guillaume Pare
  16. Department of Epidemiology, Erasmus University Medical Center, Rotterdam, the Netherlands

    • Hieab H. H. Adams
    • , M. Arfan Ikram
    • , Hieab H. H. Adams
    • , M. Arfan Ikram
    • , Najaf Amin
    •  & Sven J. van der Lee
  17. Department of Radiology and Nuclear Medicine, Erasmus University Medical Center, Rotterdam, the Netherlands

    • Hieab H. H. Adams
    •  & Hieab H. H. Adams
  18. Department of Medicine and Clinical Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan

    • Tetsuro Ago
    •  & Tetsuro Ago
  19. Department of Clinical Sciences, Lund University, Malmö, Sweden

    • Peter Almgren
    • , Olle Melander
    • , Peter Almgren
    • , Gunnar Engström
    •  & Olle Melander
  20. INSERM, Institut Pasteur de Lille, LabEx DISTALZ-UMR1167, Risk Factors and Molecular Determinants of Aging-Related Diseases, Université Lille, Lille, France

    • Philippe Amouyel
    •  & Philippe Amouyel
  21. Centre Hospitalier Université Lille, Epidemiology and Public Health Department, Lille, France

    • Philippe Amouyel
    •  & Philippe Amouyel
  22. AA Martinos Center for Biomedical Imaging, Department of Radiology, MGH, Harvard Medical School, Boston, MA, USA

    • Hakan Ay
    •  & Hakan Ay
  23. Cardiovascular Health Research Unit, Departments of Biostatistics and Medicine, University of Washington, Seattle, WA, USA

    • Traci M. Bartz
    •  & Traci M. Bartz
  24. Division of Neurology, Faculty of Medicine, Brain Research Center, University of British Columbia, Vancouver, British Columbia, Canada

    • Oscar R. Benavente
    •  & Oscar R. Benavente
  25. School of Life Science, University of Lincoln, Lincoln, UK

    • Steve Bevan
    •  & Steve Bevan
  26. Department of Cerebrovascular Diseases, Fondazione IRCCS Istituto Neurologico ‘Carlo Besta’, Milan, Italy

    • Giorgio B. Boncoraglio
    •  & Giorgio B. Boncoraglio
  27. Department of Neurology, Mayo Clinic Rochester, Rochester, MN, USA

    • Robert D. Brown Jr
    •  & Robert D. Brown Jr
  28. MRC/BHF Cardiovascular Epidemiology Unit, Department of Public Health and Primary Care, University of Cambridge, Cambridge, UK

    • Adam S. Butterworth
    • , John Danesh
    • , Wei-Yu Lin
    • , Joanna M. M. Howson
    • , Adam S. Butterworth
    • , John Danesh
    • , Wei-Yu Lin
    •  & Joanna M. M. Howson
  29. National Institute for Health Research Blood and Transplant Research Unit in Donor Health and Genomics, University of Cambridge, Cambridge, UK

    • Adam S. Butterworth
    •  & Adam S. Butterworth
  30. Neurovascular Research Laboratory, Vall d’Hebron Institut of Research, Neurology and Medicine Departments-Universitat Autònoma de Barcelona, Vall d’Hebrón Hospital, Barcelona, Spain

    • Caty Carrera
    • , Israel Fernandez-Cadenas
    • , Caty Carrera
    • , Iolanda Riba-Llena
    •  & Israel Fernandez-Cadenas
  31. Stroke Pharmacogenomics and Genetics, Fundacio Docència i Recerca MutuaTerrassa, Terrassa, Spain

    • Caty Carrera
    • , Israel Fernandez-Cadenas
    • , Caty Carrera
    • , Natalia Cullell
    • , Elena Muiño
    • , Nuria P. Torres-Aguila
    •  & Israel Fernandez-Cadenas
  32. Children’s Research Institute, Children’s National Medical Center, Washington, DC, USA

    • Cara L. Carty
    •  & Cara L. Carty
  33. Center for Translational Science, George Washington University, Washington, DC, USA

    • Cara L. Carty
    •  & Cara L. Carty
  34. Division of Preventive Medicine, Brigham and Women’s Hospital, Boston, MA, USA

    • Daniel I. Chasman
    • , Paul M. Ridker
    • , Daniel I. Chasman
    • , Paul M. Ridker
    •  & Julie E. Buring
  35. Harvard Medical School, Boston, MA, USA

    • Daniel I. Chasman
    • , Paul M. Ridker
    • , Daniel I. Chasman
    • , Paul M. Ridker
    •  & Julie E. Buring
  36. Center for Public Health Genomics, Department of Public Health Sciences, University of Virginia, Charlottesville, VA, USA

    • Wei-Min Chen
    • , Ani Manichaikul
    • , Stephen S. Rich
    • , Wei-Min Chen
    • , Ani Manichaikul
    •  & Stephen S. Rich
  37. Department of Neurology, University of Maryland School of Medicine and Baltimore VAMC, Baltimore, MD, USA

    • John W. Cole
    • , Steven J. Kittner
    • , John W. Cole
    •  & Steven J. Kittner
  38. Departments of Medicine, Pediatrics and Population Health Science, University of Mississippi Medical Center, Jackson, MS, USA

    • Adolfo Correa
    •  & Adolfo Correa
  39. Institute of Cardiovascular Research, Royal Holloway University of London, London, UK, and Ashford and St Peters Hospital, Surrey, UK

    • Ioana Cotlarciuc
    • , Pankaj Sharma
    • , Ioana Cotlarciuc
    •  & Pankaj Sharma
  40. Department of Psychiatry, Hope Center Program on Protein Aggregation and Neurodegeneration (HPAN), Washington University School of Medicine, St. Louis, MO, USA

    • Carlos Cruchaga
    •  & Carlos Cruchaga
  41. Department of Developmental Biology, Washington University School of Medicine, St. Louis, MO, USA

    • Carlos Cruchaga
    •  & Carlos Cruchaga
  42. NIHR Blood and Transplant Research Unit in Donor Health and Genomics, Department of Public Health and Primary Care, University of Cambridge, Cambridge, UK

    • John Danesh
    •  & John Danesh
  43. Wellcome Trust Sanger Institute, Hinxton, Cambridge, UK

    • John Danesh
    •  & John Danesh
  44. British Heart Foundation, Cambridge Centre of Excellence, Department of Medicine, University of Cambridge, Cambridge, UK

    • John Danesh
    •  & John Danesh
  45. Department of Medical Genetics, University Medical Center Utrecht, Utrecht, the Netherlands

    • Paul I. W. de Bakker
    • , Sara L. Pulit
    • , Paul I. W. de Bakker
    •  & Sara L. Pulit
  46. Department of Epidemiology, Julius Center for Health Sciences and Primary Care, University Medical Center Utrecht, Utrecht, the Netherlands

    • Paul I. W. de Bakker
    •  & Paul I. W. de Bakker
  47. Boston University School of Public Health, Boston, MA, USA

    • Anita L. DeStefano
    • , Qiong Yang
    • , Anita L. DeStefano
    • , Qiong Yang
    • , Alexa S. Beiser
    •  & Chloe Sarnowski
  48. Framingham Heart Study, Framingham, MA, USA

    • Anita L. DeStefano
    • , Andrew D. Johnson
    • , Sudha Seshadri
    • , Anita L. DeStefano
    • , Hugo S. Aparicio
    • , Alexa S. Beiser
    • , Seung Hoan Choi
    • , Ayesha Chowhan
    • , Vasileios Lioutas
    • , Jose R. Romero
    • , Chloe Sarnowski
    • , Claudia L. Satizabal
    •  & Sudha Seshadri
  49. Department of Immunology, Genetics and Pathology and Science for Life Laboratory, Uppsala University, Uppsala, Sweden

    • Marcel den Hoed
    •  & Marcel den Hoed
  50. Department of Genetics, University of North Carolina, Chapel Hill, NC, USA

    • Qing Duan
    •  & Qing Duan
  51. Department of Neurology and Stroke Center, Basel University Hospital, Basel, Switzerland

    • Stefan T. Engelter
    •  & Stefan T. Engelter
  52. Neurorehabilitation Unit, University of Basel and University Center for Medicine of Aging and Rehabilitation Basel, Felix Platter Hospital, Basel, Switzerland

    • Stefan T. Engelter
    •  & Stefan T. Engelter
  53. Department of Neurology, Yale University School of Medicine, New Haven, CT, USA

    • Guido J. Falcone
    •  & Guido J. Falcone
  54. Program in Medical and Population Genetics, Broad Institute of Harvard and MIT, Cambridge, MA, USA

    • Guido J. Falcone
    • , Ingrid E. Christophersen
    • , Carolina Roselli
    • , Steven A. Lubitz
    • , Patrick T. Ellinor
    •  & Guido J. Falcone
  55. Department of Neurology, Johns Hopkins University School of Medicine, Baltimore, MD, USA

    • Rebecca F. Gottesman
    •  & Rebecca F. Gottesman
  56. Neuroscience Institute, SF Medical Center, Trenton, NJ, USA

    • Raji P. Grewal
    • , Raji P. Grewal
    •  & Leema Reddy Peddareddygari
  57. Icelandic Heart Association Research Institute, Kopavogur, Iceland

    • Vilmundur Gudnason
    •  & Vilmundur Gudnason
  58. Faculty of Medicine, University of Iceland, Reykjavik, Iceland

    • Vilmundur Gudnason
    •  & Vilmundur Gudnason
  59. Department of Medical Sciences, Molecular Epidemiology and Science for Life Laboratory, Uppsala University, Uppsala, Sweden

    • Stefan Gustafsson
    •  & Stefan Gustafsson
  60. Division of Public Health Sciences, Fred Hutchinson Cancer Research Center, Seattle, WA, USA

    • Jeffrey Haessler
    • , Charles Kooperberg
    • , Jeffrey Haessler
    •  & Charles Kooperberg
  61. Laboratory of Epidemiology and Population Science, National Institute on Aging, National Institutes of Health, Bethesda, MD, USA

    • Tamara B. Harris
    •  & Tamara B. Harris
  62. Department of Neurology, Leeds General Infirmary, Leeds Teaching Hospitals NHS Trust, Leeds, UK

    • Ahamad Hassan
    •  & Ahamad Hassan
  63. National Institute for Health and Welfare, Helsinki, Finland

    • Aki S. Havulinna
    • , Veikko Salomaa
    • , Aki S. Havulinna
    • , Veikko Salomaa
    • , Pekka Jousilahti
    •  & Markus Perola
  64. FIMM–Institute for Molecular Medicine Finland, Helsinki, Finland

    • Aki S. Havulinna
    •  & Aki S. Havulinna
  65. Department of Epidemiology, University of Washington, Seattle, WA, USA

    • Susan R. Heckbert
    • , Alexander P. Reiner
    • , Nicholas L. Smith
    • , W. T. Longstreth Jr
    • , Susan R. Heckbert
    • , Alexander P. Reiner
    •  & W. T. Longstreth Jr
  66. Public Health Stream, Hunter Medical Research Institute, New Lambton, New South Wales, Australia

    • Elizabeth G. Holliday
    •  & Elizabeth G. Holliday
  67. Faculty of Health and Medicine, University of Newcastle, Newcastle, New South Wales, Australia

    • Elizabeth G. Holliday
    •  & Elizabeth G. Holliday
  68. School of Public Health, University of Alabama at Birmingham, Birmingham, AL, USA

    • George Howard
    •  & George Howard
  69. Department of Biostatistical Sciences, Wake Forest School of Medicine, Winston-Salem, NC, USA

    • Fang-Chi Hsu
    •  & Fang-Chi Hsu
  70. Aflac Cancer and Blood Disorder Center, Department of Pediatrics, Emory University School of Medicine, Atlanta, GA, USA

    • Hyacinth I. Hyacinth
    •  & Hyacinth I. Hyacinth
  71. Department of Medicine, Division of Cardiovascular Medicine, Stanford University School of Medicine, Stanford, CA, USA

    • Erik Ingelsson
    •  & Erik Ingelsson
  72. Department of Medical Sciences, Molecular Epidemiology and Science for Life Laboratory, Uppsala University, Uppsala, Sweden

    • Erik Ingelsson
    •  & Erik Ingelsson
  73. Epidemiology, School of Public Health, University of Alabama at Birmingham, Birmingham, AL, USA

    • Marguerite R. Irvin
    •  & Marguerite R. Irvin
  74. Brown Foundation Institute of Molecular Medicine, University of Texas Health Science Center at Houston, Houston, TX, USA

    • Xueqiu Jian
    • , Myriam Fornage
    • , Xueqiu Jian
    •  & Myriam Fornage
  75. Neurovascular Research Group (NEUVAS), Neurology Department, Institut Hospital del Mar d’Investigació Mèdica, Universitat Autònoma de Barcelona, Barcelona, Spain

    • Jordi Jiménez-Conde
    •  & Jordi Jiménez-Conde
  76. Department of Pharmacotherapy and Translational Research and Center for Pharmacogenomics, University of Florida, College of Pharmacy, Gainesville, FL, USA

    • Julie A. Johnson
    •  & Julie A. Johnson
  77. Division of Cardiovascular Medicine, College of Medicine, University of Florida, Gainesville, FL, USA

    • Julie A. Johnson
    •  & Julie A. Johnson
  78. Department of Cardiology, Leiden University Medical Center, Leiden, the Netherlands

    • J. Wouter Jukema
    •  & J. Wouter Jukema
  79. Program in Bioinformatics and Integrative Genomics, Harvard Medical School, Boston, MA, USA

    • Masahiro Kanai
    •  & Masahiro Kanai
  80. Department of Biology, East Carolina University, Greenville, NC, USA

    • Keith L. Keene
    •  & Keith L. Keene
  81. Center for Health Disparities, East Carolina University, Greenville, NC, USA

    • Keith L. Keene
    •  & Keith L. Keene
  82. University of Cincinnati College of Medicine, Cincinnati, OH, USA

    • Brett M. Kissela
    • , Dawn O. Kleindorfer
    • , Daniel Woo
    • , Brett M. Kissela
    • , Dawn O. Kleindorfer
    •  & Daniel Woo
  83. RIKEN Center for Integrative Medical Sciences, Yokohama, Japan

    • Michiaki Kubo
    •  & Michiaki Kubo
  84. Department of Medicine, University of Colorado Denver, Anschutz Medical Campus, Aurora, CO, USA

    • Leslie A. Lange
    •  & Leslie A. Lange
  85. Center for Public Health Genomics and Department of Biostatistical Sciences, Wake Forest School of Medicine, Winston-Salem, NC, USA

    • Carl D. Langefeld
    •  & Carl D. Langefeld
  86. MRC Epidemiology Unit, University of Cambridge School of Clinical Medicine, Institute of Metabolic Science, Cambridge, UK

    • Claudia Langenberg
    • , Nicholas J. Wareham
    • , Claudia Langenberg
    •  & Nicholas J. Wareham
  87. Intramural Research Program, National Institute on Aging, National Institutes of Health, Bethesda, MD, USA

    • Lenore J. Launer
    •  & Lenore J. Launer
  88. Department of Neurology, Radiology, and Biomedical Engineering, Washington University School of Medicine, St. Louis, MO, USA

    • Jin-Moo Lee
    •  & Jin-Moo Lee
  89. Department of Neurosciences, Experimental Neurology, KU Leuven–University of Leuven, Leuven, Belgium

    • Robin Lemmens
    •  & Robin Lemmens
  90. VIB Center for Brain & Disease Research, University Hospitals Leuven, Department of Neurology, Leuven, Belgium

    • Robin Lemmens
    •  & Robin Lemmens
  91. INSERM U 1171, CHU Lille, Université Lille, Lille, France

    • Didier Leys
    •  & Didier Leys
  92. Department of Medical and Molecular Genetics, King’s College London, London, UK

    • Cathryn M. Lewis
    •  & Cathryn M. Lewis
  93. SGDP Centre, Institute of Psychiatry, Psychology & Neuroscience, King’s College London, London, UK

    • Cathryn M. Lewis
    •  & Cathryn M. Lewis
  94. Northern Institute for Cancer Research, Newcastle University, Newcastle, UK

    • Wei-Yu Lin
    •  & Wei-Yu Lin
  95. Department of Clinical Sciences Lund, Neurology, Lund University, Lund, Sweden

    • Arne G. Lindgren
    • , Arne G. Lindgren
    • , Hossein Delavaran
    • , Andreea Ilinca
    •  & Bo Norrving
  96. Department of Neurology and Rehabilitation Medicine, Skåne University Hospital, Lund, Sweden

    • Arne G. Lindgren
    • , Arne G. Lindgren
    • , Hossein Delavaran
    • , Andreea Ilinca
    •  & Bo Norrving
  97. Bioinformatics Core Facility, University of Gothenburg, Gothenburg, Sweden

    • Erik Lorentzen
    •  & Erik Lorentzen
  98. Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden

    • Patrik K. Magnusson
    • , Patrik K. Magnusson
    •  & Nancy L. Pedersen
  99. University of Technology Sydney, Faculty of Health, Ultimo, New South Wales, Australia

    • Jane Maguire
    •  & Jane Maguire
  100. Department of Medicine, University of Maryland School of Medicine, Baltimore, MD, USA

    • Patrick F. McArdle
    • , Braxton D. Mitchell
    • , Patrick F. McArdle
    •  & Braxton D. Mitchell
  101. Department of Neurology, Mayo Clinic, Jacksonville, FL, USA

    • James F. Meschia
    •  & James F. Meschia
  102. Geriatrics Research and Education Clinical Center, Baltimore Veterans Administration Medical Center, Baltimore, MD, USA

    • Braxton D. Mitchell
    •  & Braxton D. Mitchell
  103. Division of Geriatrics, School of Medicine, University of Mississippi Medical Center, Jackson, MS, USA

    • Thomas H. Mosley
    •  & Thomas H. Mosley
  104. Memory Impairment and Neurodegenerative Dementia Center, University of Mississippi Medical Center, Jackson, MS, USA

    • Thomas H. Mosley
    •  & Thomas H. Mosley
  105. Laboratory of Neurogenetics, National Institute on Aging, National Institutes of Health, Bethesda, MD, USA

    • Michael A. Nalls
    •  & Michael A. Nalls
  106. Data Tecnica International, Glen Echo, MD, USA

    • Michael A. Nalls
    •  & Michael A. Nalls
  107. Department of Epidemiology and Public Health, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan

    • Toshiharu Ninomiya
    • , Jun Hata
    • , Toshiharu Ninomiya
    •  & Jun Hata
  108. Clinical Research Facility, Department of Medicine, NUI Galway, Galway, Ireland

    • Martin J. O’Donnell
    •  & Martin J. O’Donnell
  109. Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, WA, USA

    • Bruce M. Psaty
    • , Kerri L. Wiggins
    • , Joshua C. Bis
    • , Bruce M. Psaty
    •  & Kerri L. Wiggins
  110. Department of Epidemiology, University of Washington, Seattle, WA, USA

    • Bruce M. Psaty
    •  & Bruce M. Psaty
  111. Department of Health Services, University of Washington, Seattle, WA, USA

    • Bruce M. Psaty
    •  & Bruce M. Psaty
  112. Kaiser Permanente Washington Health Research Institute, Seattle, WA, USA

    • Bruce M. Psaty
    •  & Bruce M. Psaty
  113. Brain Center Rudolf Magnus, Department of Neurology, University Medical Center Utrecht, Utrecht, the Netherlands

    • Sara L. Pulit
    •  & Sara L. Pulit
  114. Usher Institute of Population Health Sciences and Informatics, University of Edinburgh, Edinburgh, UK

    • Kristiina Rannikmäe
    • , Cathie L. M. Sudlow
    • , Kristiina Rannikmäe
    •  & Cathie L. M. Sudlow
  115. Centre for Clinical Brain Sciences, University of Edinburgh, Edinburgh, UK

    • Kristiina Rannikmäe
    • , Cathie L. M. Sudlow
    • , Kristiina Rannikmäe
    •  & Cathie L. M. Sudlow
  116. Fred Hutchinson Cancer Research Center, University of Washington, Seattle, WA, USA

    • Alexander P. Reiner
    •  & Alexander P. Reiner
  117. Department of Medicine, Brigham and Women’s Hospital, Boston, MA, USA

    • Kathryn M. Rexrode
    •  & Kathryn M. Rexrode
  118. Department of Biostatistics, University of Washington, Seattle, WA, USA

    • Kenneth Rice
    • , Kenneth Rice
    • , Cathy C. Laurie
    •  & Quenna Wong
  119. Nuffield Department of Clinical Neurosciences, University of Oxford, Oxford, UK

    • Peter M. Rothwell
    •  & Peter M. Rothwell
  120. Institute for Translational Genomics and Population Sciences, Los Angeles Biomedical Research Institute at Harbor–UCLA Medical Center, Torrance, CA, USA

    • Jerome I. Rotter
    • , Kent D. Taylor
    • , Jerome I. Rotter
    •  & Kent D. Taylor
  121. Division of Genomic Outcomes, Department of Pediatrics, Harbor–UCLA Medical Center, Torrance, CA, USA

    • Jerome I. Rotter
    • , Kent D. Taylor
    • , Jerome I. Rotter
    •  & Kent D. Taylor
  122. Department of Neurology, Miller School of Medicine, University of Miami, Miami, FL, USA

    • Tatjana Rundek
    • , Ralph L. Sacco
    • , Tatjana Rundek
    •  & Ralph L. Sacco
  123. Department of Allergy and Rheumatology, Graduate School of Medicine, University of Tokyo, Tokyo, Japan

    • Saori Sakaue
    •  & Saori Sakaue
  124. Center for Public Health Genomics, University of Virginia, Charlottesville, VA, USA

    • Michele M. Sale
    • , Mete Civelek
    •  & Michele M. Sale
  125. Department of Pediatrics, College of Medicine, University of Oklahoma Health Sciences Center, Oklahoma City, OK, USA

    • Bishwa R. Sapkota
    • , Dharambir K. Sanghera
    • , Bishwa R. Sapkota
    •  & Dharambir K. Sanghera
  126. Department of Neurology, Medical University of Graz, Graz, Austria

    • Reinhold Schmidt
    •  & Reinhold Schmidt
  127. Institute for Community Medicine, SHIP-KEF, University Medicine Greifswald, Greifswald, Germany

    • Carsten O. Schmidt
    •  & Carsten O. Schmidt
  128. Department of Neurology, University Medicine Greifswald, Greifswald, Germany

    • Ulf Schminke
    •  & Ulf Schminke
  129. Department of Neurology, Jagiellonian University, Krakow, Poland

    • Agnieszka Slowik
    • , Agnieszka Slowik
    •  & Joanna Pera
  130. Department of Neurology, Justus Liebig University, Giessen, Germany

    • Christian Tanislav
    •  & Christian Tanislav
  131. Department of Clinical Neurosciences/Neurology, Institute of Neuroscience and Physiology, Sahlgrenska Academy at University of Gothenburg, Gothenburg, Sweden

    • Turgut Tatlisumak
    •  & Turgut Tatlisumak
  132. Sahlgrenska University Hospital, Gothenburg, Sweden

    • Turgut Tatlisumak
    •  & Turgut Tatlisumak
  133. Stroke Division, Florey Institute of Neuroscience and Mental Health, University of Melbourne, Heidelberg, Victoria, Australia

    • Vincent N. S. Thijs
    •  & Vincent N. S. Thijs
  134. Austin Health, Department of Neurology, Heidelberg, Victoria, Australia

    • Vincent N. S. Thijs
    •  & Vincent N. S. Thijs
  135. Department of Internal Medicine, Section Gerontology and Geriatrics, Leiden University Medical Center, Leiden, the Netherlands

    • Stella Trompet
    •  & Stella Trompet
  136. INSERM, U1219, Bordeaux, France

    • Christophe Tzourio
    •  & Christophe Tzourio
  137. Department of Public Health, Bordeaux University Hospital, Bordeaux, France

    • Christophe Tzourio
    •  & Christophe Tzourio
  138. Genetic Epidemiology Unit, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, the Netherlands

    • Cornelia M. van Duijn
    •  & Cornelia M. van Duijn
  139. Center for Medical Systems Biology, Leiden, the Netherlands

    • Cornelia M. van Duijn
    •  & Cornelia M. van Duijn
  140. School of Medicine, Dentistry and Nursing at the University of Glasgow, Glasgow, UK

    • Matthew Walters
    •  & Matthew Walters
  141. Department of Epidemiology and Population Health, Albert Einstein College of Medicine, New York, NY, USA

    • Sylvia Wassertheil-Smoller
    •  & Sylvia Wassertheil-Smoller
  142. Department of Physiology and Biophysics, University of Mississippi Medical Center, Jackson, MS, USA

    • James G. Wilson
    •  & James G. Wilson
  143. Department of Human Genetics, McGill University, Montreal, Quebec, Canada

    • Tomi Pastinen
  144. Department of Pathophysiology, Institute of Biomedicine and Translation Medicine, University of Tartu, Tartu, Estonia

    • Arno Ruusalepp
  145. Department of Cardiac Surgery, Tartu University Hospital, Tartu, Estonia

    • Arno Ruusalepp
  146. Clinical Gene Networks AB, Stockholm, Sweden

    • Arno Ruusalepp
  147. Department of Genetics and Genomic Sciences, Icahn Institute for Genomics and Multiscale Biology Icahn School of Medicine at Mount Sinai, New York, NY, USA

    • Eric E. Schadt
    • , Simon Koplev
    •  & Johan L. M. Björkegren
  148. Department of Pathophysiology, Institute of Biomedicine and Translation Medicine, Biomeedikum, University of Tartu, Tartu, Estonia

    • Johan L. M. Björkegren
  149. Integrated Cardio Metabolic Centre, Department of Medicine, Karolinska Institutet, Karolinska Universitetssjukhuset, Huddinge, Sweden

    • Johan L. M. Björkegren
  150. Clinical Gene Networks AB, Stockholm, Sweden

    • Johan L. M. Björkegren
  151. UPMC Univ. Paris 06, INSERM, UMR_S 1166, Team Genomics & Pathophysiology of Cardiovascular Diseases, Sorbonne Universités, Paris, France

    • Veronica Codoni
    •  & David A. Trégouët
  152. ICAN Institute for Cardiometabolism and Nutrition, Paris, France

    • Veronica Codoni
    •  & David A. Trégouët
  153. Department of Biomedical Engineering, University of Virginia, Charlottesville, VA, USA

    • Mete Civelek
  154. Group Health Research Institute, Group Health Cooperative, Seattle, WA, USA

    • Nicholas L. Smith
  155. Seattle Epidemiologic Research and Information Center, VA Office of Research and Development, Seattle, WA, USA

    • Nicholas L. Smith
  156. Cardiovascular Research Center, MGH, Boston, MA, USA

    • Ingrid E. Christophersen
    • , Steven A. Lubitz
    •  & Patrick T. Ellinor
  157. Department of Medical Research, Bærum Hospital, Vestre Viken Hospital Trust, Gjettum, Norway

    • Ingrid E. Christophersen
  158. Saw Swee Hock School of Public Health, National University of Singapore and National University Health System, Singapore, Singapore

    • E. Shyong Tai
  159. National Heart and Lung Institute, Imperial College London, London, UK

    • Jaspal S. Kooner
  160. Department of Gene Diagnostics and Therapeutics, Research Institute, National Center for Global Health and Medicine, Tokyo, Japan

    • Norihiro Kato
    •  & Fumihiko Takeuchi
  161. Department of Epidemiology, Tulane University School of Public Health and Tropical Medicine, New Orleans, LA, USA

    • Jiang He
  162. Department of Cardiology, University Medical Center Groningen, University of Groningen, Groningen, the Netherlands

    • Pim van der Harst
  163. MRC-PHE Centre for Environment and Health, School of Public Health, Department of Epidemiology and Biostatistics and the NIHR Imperial Biomedical Research Centre, Imperial College London, London, UK

    • Paul Elliott
  164. Department of Epidemiology and Biostatistics, Imperial College London, London, UK

    • John C. Chambers
  165. Department of Cardiology, Ealing Hospital NHS Trust, Southall, UK

    • John C. Chambers
  166. National Heart, Lung and Blood Research Institute, Division of Intramural Research, Population Sciences Branch, Framingham, MA, USA

    • Andrew D. Johnson
  167. Department of Pharmaceutical Sciences, College of Pharmacy, University of Oklahoma Health Sciences Center, Oklahoma City, OK, USA

    • Dharambir K. Sanghera
    •  & Dharambir K. Sanghera
  168. Oklahoma Center for Neuroscience, Oklahoma City, OK, USA

    • Dharambir K. Sanghera
    •  & Dharambir K. Sanghera
  169. Department of Pathology and Genetics, Institute of Biomedicine, Sahlgrenska Academy at University of Gothenburg, Gothenburg, Sweden

    • Christina Jern
    •  & Christina Jern
  170. Department of Neurology, Helsinki University Hospital, Helsinki, Finland

    • Daniel Strbian
    • , Nicolas Martinez-Majander
    • , Gerli Sibolt
    • , Liisa Tomppo
    •  & Daniel Strbian
  171. Clinical Neurosciences, Neurology, University of Helsinki, Helsinki, Finland

    • Daniel Strbian
    •  & Daniel Strbian
  172. Department of Neurology, University of Washington, Seattle, WA, USA

    • W. T. Longstreth Jr
    •  & W. T. Longstreth Jr
  173. Albrecht Kossel Institute, University Clinic of Rostock, Rostock, Germany

    • Arndt Rolfs
    •  & Arndt Rolfs
  174. Clinical Trial Service Unit and Epidemiological Studies Unit, Nuffield Department of Population Health, University of Oxford, Oxford, UK

    • Jemma C. Hopewell
    •  & Jemma C. Hopewell
  175. Department of Genetics, Perelman School of Medicine, University of Pennsylvania, Philadelphia, PA, USA

    • Danish Saleheen
    •  & Danish Saleheen
  176. Faculty of Medicine, University of Iceland, Reykjavik, Iceland

    • Kari Stefansson
    •  & Kari Stefansson
  177. Departments of Neurology and Public Health Sciences, University of Virginia School of Medicine, Charlottesville, VA, USA

    • Bradford B. Worrall
    •  & Bradford B. Worrall
  178. Glenn Biggs Institute for Alzheimer’s and Neurodegenerative Diseases, University of Texas Health Sciences Center, San Antonio, San Antonio, TX, USA

    • Sudha Seshadri
    •  & Sudha Seshadri
  179. Human Genetics Center, University of Texas Health Science Center at Houston, Houston, TX, USA

    • Myriam Fornage
    •  & Myriam Fornage
  180. Center for Genomic Medicine, Kyoto University Graduate School of Medicine, Kyoto, Japan

    • Yoichiro Kamatani
    •  & Yoichiro Kamatani
  181. Munich Cluster for Systems Neurology (SyNergy), Munich, Germany

    • Martin Dichgans
    •  & Martin Dichgans
  182. German Center for Neurodegenerative Diseases (DZNE), Munich, Germany

    • Martin Dichgans
    •  & Martin Dichgans
  183. Boston University School of Medicine, Boston, MA, USA

    • Hugo S. Aparicio
    • , Ayesha Chowhan
    • , Jose R. Romero
    •  & Claudia L. Satizabal
  184. University of Kentucky College of Public Health, Lexington, KY, USA

    • Donna K. Arnett
  185. University of Newcastle and Hunter Medical Research Institute, New Lambton, New South Wales, Australia

    • John Attia
  186. INSERM, U1061, Université Montpellier, Montpellier, France

    • Claudine Berr
  187. Centre for Research in Environmental Epidemiology, Barcelona, Spain

    • Mariana Bustamante
  188. Department of Neurology, Università degli Studi di Perugia, Umbria, Italy

    • Valeria Caso
  189. Department of Medicine, University of Maryland School of Medicine, Baltimore, MD, USA

    • Yu-Ching Cheng
    •  & Huichun Xu
  190. Broad Institute, Cambridge, MA, USA

    • Seung Hoan Choi
  191. Bordeaux Population Health Research Center, INSERM, UMR 1219, Université Bordeaux, Bordeaux, France

    • Jean-François Dartigues
  192. Department of Neurology, Memory Clinic, Bordeaux University Hospital, Bordeaux, France

    • Jean-François Dartigues
  193. Neurovascular Research Laboratory. Vall d’Hebron Institut of Research, Neurology and Medicine Departments-Universitat Autònoma de Barcelona. Vall d’Hebrón Hospital, Barcelona, Spain

    • Pilar Delgado
  194. Department of Internal Medicine B, University Medicine Greifswald, Greifswald, Germany

    • Marcus Dörr
  195. DZHK, Greifswald, Germany

    • Marcus Dörr
  196. Robertson Center for Biostatistics, University of Glasgow, Glasgow, UK

    • Ian Ford
  197. Hero DMC Heart Institute, Dayanand Medical College & Hospital, Ludhiana, India

    • Wander S. Gurpreet
    •  & Ralhan Sarju
  198. Atherosclerosis Research Unit, Department of Medicine Solna, Karolinska Institutet, Stockholm, Sweden

    • Anders Hamsten
  199. Karolinska Institutet, Stockholm, Sweden

    • Anders Hamsten
  200. Division of Emergency Medicine, and Department of Neurology, Washington University School of Medicine, St. Louis, MO, USA

    • Laura Heitsch
  201. Tohoku Medical Megabank Organization, Sendai, Japan

    • Atsushi Hozawa
    • , Naoko Minegishi
    • , Soichi Ogishima
    •  & Takako Takai
  202. Department of Psychiatry, Washington University School of Medicine, St. Louis, MO, USA

    • Laura Ibanez
  203. Department of Public Health and Caring Sciences/Geriatrics, Uppsala University, Uppsala, Sweden

    • Martin Ingelsson
  204. Epidemiology and Prevention Group, Center for Public Health Sciences, National Cancer Center, Tokyo, Japan

    • Motoki Iwasaki
    • , Norie Sawada
    • , Shoichiro Tsugane
    •  & Taiki Yamaji
  205. Department of Internal Medicine and the Center for Clinical and Translational Science, Ohio State University, Columbus, OH, USA

    • Rebecca D. Jackson
  206. Institute of Neuroscience and Physiology, Sahlgrenska Academy at University of Gothenburg, Goteborg, Sweden

    • Katarina Jood
  207. Department of Basic and Clinical Neurosciences, King’s College London, London, UK

    • Lalit Kalra
  208. Department of Health Care Administration and Management, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan

    • Masahiro Kamouchi
  209. Department of Medicine and Clinical Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan

    • Takanari Kitazono
  210. Departments of Neurology & Radiology, Landspitali National University Hospital, Reykjavik, Iceland

    • Olafur Kjartansson
  211. Department of Neurology, Heidelberg University Hospital, Heidelberg, Germany

    • Manja Kloss
  212. Department of Neurology, Erasmus University Medical Center, Rotterdam, the Netherlands

    • Peter J. Koudstaal
  213. Hospital Universitari Mutua Terrassa, Terrassa (Barcelona), Spain

    • Jerzy Krupinski
  214. Montefiore Medical Center, Albert Einstein College of Medicine, New York, NY, USA

    • Daniel L. Labovitz
  215. John Hunter Hospital, Hunter Medical Research Institute and University of Newcastle, Newcastle, New South Wales, Australia

    • Christopher R. Levi
  216. Centre for Prevention of Stroke and Dementia, Nuffield Department of Clinical Neurosciences, University of Oxford, Oxford, UK

    • Linxin Li
  217. Department of Medical Sciences, Uppsala University, Uppsala, Sweden

    • Lars Lind
  218. Genetic and Genomic Epidemiology Unit, Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, UK

    • Cecilia M. Lindgren
  219. Wellcome Trust Centre for Human Genetics, Oxford, UK

    • Cecilia M. Lindgren
  220. Beth Israel Deaconess Medical Center, Boston, MA, USA

    • Vasileios Lioutas
  221. Wake Forest School of Medicine, Wake Forest, NC, USA

    • Yong Mei Liu
  222. Department of Neurology, University of Pittsburgh, Pittsburgh, PA, USA

    • Oscar L. Lopez
  223. BioBank Japan, Laboratory of Clinical Sequencing, Department of Computational Biology and Medical Sciences, Graduate School of Frontier Sciences, University of Tokyo, Tokyo, Japan

    • Hirata Makoto
    •  & Koichi Matsuda
  224. Neurovascular Research Laboratory, Vall d’Hebron Institut of Research, Neurology and Medicine Departments-Universitat Autònoma de Barcelona, Vall d’Hebrón Hospital, Barcelona, Spain

    • Joan Montaner
  225. Department of Biostatistics, University of Liverpool, Liverpool, UK

    • Andrew P. Morris
  226. Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, UK

    • Andrew P. Morris
  227. Institute of Genetic Epidemiology, Helmholtz Zentrum München–German Research Center for Environmental Health, Neuherberg, Germany

    • Martina Müller-Nurasyid
    •  & Konstantin Strauch
  228. Department of Medicine I, Ludwig-Maximilians-Universität, Munich, Germany

    • Martina Müller-Nurasyid
  229. DZHK (German Centre for Cardiovascular Research), partner site Munich Heart Alliance, Munich, Germany

    • Martina Müller-Nurasyid
  230. Department of Cerebrovascular Diseases, Fondazione IRCCS Istituto Neurologico ‘Carlo Besta’, Milan, Italy

    • Eugenio A. Parati
  231. MEB, Karolinska Institutet, Stockholm, Sweden

    • Nancy L. Pedersen
  232. Estonian Genome Center, University of Tartu, Tartu, Estonia

    • Markus Perola
  233. Department of Clinical and Experimental Sciences, Neurology Clinic, University of Brescia, Brescia, Italy

    • Alessandro Pezzini
  234. Translational Genomics Unit, Department of Oncology, IRCCS Istituto di Ricerche Farmacologiche Mario Negri, Milan, Italy

    • Silvana Pileggi
  235. Department of Genetics, Microbiology and Statistics, University of Barcelona, Barcelona, Spain

    • Raquel Rabionet
  236. Psychiatric Genetics Unit, Group of Psychiatry, Mental Health and Addictions, Vall d’Hebron Research Institute (VHIR), Universitat Autònoma de Barcelona, Biomedical Network Research Centre on Mental Health (CIBERSAM), Barcelona, Spain

    • Marta Ribasés
  237. Department of Neurology, IMIM–Hospital del Mar, and Universitat Autònoma de Barcelona, Barceloina, Spain

    • Jaume Roquer
  238. IMIM (Hospital del Mar Medical Research Institute), Barcelona, Spain

    • Jaume Roquer
    •  & Carolina Soriano-Tárraga
  239. National Institute for Health Research Comprehensive Biomedical Research Centre, Guy’s & St. Thomas’ NHS Foundation Trust and King’s College London, London, UK

    • Anthony G. Rudd
    •  & Charles D. A. Wolfe
  240. Division of Health and Social Care Research, King’s College London, London, UK

    • Anthony G. Rudd
    •  & Charles D. A. Wolfe
  241. FIMM–Institute for Molecular Medicine Finland, Helsinki, Finland

    • Antti-Pekka Sarin
  242. THL–National Institute for Health and Welfare, Helsinki, Finland

    • Antti-Pekka Sarin
  243. Iwate Tohoku Medical Megabank Organization, Iwate Medical University, Iwate, Japan

    • Makoto Sasaki
    • , Mamoru Satoh
    • , Kenji Sobue
    •  & Kozo Tanno
  244. BHF Glasgow Cardiovascular Research Centre, Faculty of Medicine, Glasgow, UK

    • Naveed Sattar
  245. deCODE Genetics/Amgen, Inc., Reykjavik, Iceland

    • Ásgeir Sigurdsson
  246. Icelandic Heart Association, Reykjavik, Iceland

    • Albert Smith
  247. Institute of Biomedicine, Sahlgrenska Academy at University of Gothenburg, Goteborg, Sweden

    • Tara Stanne
  248. Department of Epidemiology, University of Maryland School of Medicine, Baltimore, MD, USA

    • O. Colin Stine
  249. Institute of Cardiovascular and Medical Sciences, Faculty of Medicine, University of Glasgow, Glasgow, UK

    • David J. Stott
  250. IBE, Faculty of Medicine, LMU Munich, Munich, Germany

    • Konstantin Strauch
  251. Division of Epidemiology and Prevention, Aichi Cancer Center Research Institute, Nagoya, Japan

    • Hideo Tanaka
    •  & Kenji Wakai
  252. Department of Epidemiology, Nagoya University Graduate School of Medicine, Nagoya, Japan

    • Hideo Tanaka
  253. Institute for Community Medicine, SHIP-KEF, University Medicine Greifswald, Greifswald, Germany

    • Alexander Teumer
    •  & Henry Völzke
  254. Department of Neurology, Caen University Hospital, Caen, France

    • Emmanuel Touze
  255. University of Caen Normandy, Caen, France

    • Emmanuel Touze
  256. Department of Internal Medicine, Erasmus University Medical Center, Rotterdam, the Netherlands

    • Andre G. Uitterlinden
  257. Landspitali University Hospital, Reykjavik, Iceland

    • Einar M. Valdimarsson
  258. Survey Research Center, University of Michigan, Ann Arbor, MI, USA

    • David Weir
  259. Department of Neurology, University of Virginia, Charlottesville, VA, USA

    • Stephen R. Williams

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