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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Genome-wide meta-analysis identifies new susceptibility loci for migraine</title><author><name sortKey="Anttila, Verneri" sort="Anttila, Verneri" uniqKey="Anttila V" first="Verneri" last="Anttila">Verneri Anttila</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Cambridge, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A2">Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A3">Analytical and Translational Genetics Unit, Department of Medicine, Massachusetts General Hospital and Harvard Medical School, Boston, MA, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A4">Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA</nlm:aff></affiliation></author><author><name sortKey="Winsvold, Bendik S" sort="Winsvold, Bendik S" uniqKey="Winsvold B" first="Bendik S." last="Winsvold">Bendik S. Winsvold</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Cambridge, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A5">Department of Neurology, Oslo University Hospital and University of Oslo, Oslo, Norway</nlm:aff></affiliation></author><author><name sortKey="Gormley, Padhraig" sort="Gormley, Padhraig" uniqKey="Gormley P" first="Padhraig" last="Gormley">Padhraig Gormley</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Kurth, Tobias" sort="Kurth, Tobias" uniqKey="Kurth T" first="Tobias" last="Kurth">Tobias Kurth</name><affiliation><nlm:aff id="A6">INSERM Unit 708 – Neuroepidemiology, F-33000 Bordeaux, France</nlm:aff></affiliation><affiliation><nlm:aff id="A7">University of Bordeaux, F-33000 Bordeaux, France</nlm:aff></affiliation><affiliation><nlm:aff id="A8">Division of Preventive Medicine, Department of Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, USA</nlm:aff></affiliation></author><author><name sortKey="Bettella, Francesco" sort="Bettella, Francesco" uniqKey="Bettella F" first="Francesco" last="Bettella">Francesco Bettella</name><affiliation><nlm:aff id="A9">deCODE genetics, Reykjavik, Iceland</nlm:aff></affiliation></author><author><name sortKey="Mcmahon, George" sort="Mcmahon, George" uniqKey="Mcmahon G" first="George" last="Mcmahon">George Mcmahon</name><affiliation><nlm:aff id="A10">MRC Centre for Causal Analyses in Translational Epidemiology, School of Social and Community Medicine, University of Bristol, Bristol, UK</nlm:aff></affiliation></author><author><name sortKey="Kallela, Mikko" sort="Kallela, Mikko" uniqKey="Kallela M" first="Mikko" last="Kallela">Mikko Kallela</name><affiliation><nlm:aff id="A11">Department of Neurology, Helsinki University Central Hospital, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Malik, Rainer" sort="Malik, Rainer" uniqKey="Malik R" first="Rainer" last="Malik">Rainer Malik</name><affiliation><nlm:aff id="A12">Institute for Stroke and Dementia Research, Klinikum der Universität München, Ludwig-Maximilians-Universität, Munich, Germany</nlm:aff></affiliation></author><author><name sortKey="De Vries, Boukje" sort="De Vries, Boukje" uniqKey="De Vries B" first="Boukje" last="De Vries">Boukje De Vries</name><affiliation><nlm:aff id="A13">Department of Human Genetics, Leiden University Medical Centre, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Terwindt, Gisela" sort="Terwindt, Gisela" uniqKey="Terwindt G" first="Gisela" last="Terwindt">Gisela Terwindt</name><affiliation><nlm:aff id="A14">Department of Neurology, Leiden University Medical Centre, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Medland, Sarah E" sort="Medland, Sarah E" uniqKey="Medland S" first="Sarah E." last="Medland">Sarah E. Medland</name><affiliation><nlm:aff id="A15">Queensland Institute of Medical Research, Brisbane, Queensland, Australia</nlm:aff></affiliation></author><author><name sortKey="Todt, Unda" sort="Todt, Unda" uniqKey="Todt U" first="Unda" last="Todt">Unda Todt</name><affiliation><nlm:aff id="A16">Institute of Human Genetics, University of Ulm, Ulm, Germany</nlm:aff></affiliation></author><author><name sortKey="Mcardle, Wendy L" sort="Mcardle, Wendy L" uniqKey="Mcardle W" first="Wendy L." last="Mcardle">Wendy L. Mcardle</name><affiliation><nlm:aff id="A10">MRC Centre for Causal Analyses in Translational Epidemiology, School of Social and Community Medicine, University of Bristol, Bristol, UK</nlm:aff></affiliation></author><author><name sortKey="Quaye, Lydia" sort="Quaye, Lydia" uniqKey="Quaye L" first="Lydia" last="Quaye">Lydia Quaye</name><affiliation><nlm:aff id="A17">Department of Twin Research and Genetic Epidemiology, King’s College London, London, UK</nlm:aff></affiliation></author><author><name sortKey="Koiranen, Markku" sort="Koiranen, Markku" uniqKey="Koiranen M" first="Markku" last="Koiranen">Markku Koiranen</name><affiliation><nlm:aff id="A18">Institute of Health Sciences, University of Oulu, Oulu, Finland</nlm:aff></affiliation></author><author><name sortKey="Ikram, M Arfan" sort="Ikram, M Arfan" uniqKey="Ikram M" first="M. Arfan" last="Ikram">M. Arfan Ikram</name><affiliation><nlm:aff id="A19">Genetic Epidemiology Unit, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, the Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A20">Department of Radiology Erasmus University Medical Centre, Rotterdam, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A21">Department of Neurology Erasmus University Medical Centre, Rotterdam, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Lehtim Ki, Terho" sort="Lehtim Ki, Terho" uniqKey="Lehtim Ki T" first="Terho" last="Lehtim Ki">Terho Lehtim Ki</name><affiliation><nlm:aff id="A22">Department of Clinical Chemistry, Fimlab Laboratories and University of Tampere School of Medicine, Tampere, Finland</nlm:aff></affiliation></author><author><name sortKey="Stam, Anine H" sort="Stam, Anine H" uniqKey="Stam A" first="Anine H." last="Stam">Anine H. Stam</name><affiliation><nlm:aff id="A14">Department of Neurology, Leiden University Medical Centre, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Ligthart, Lannie" sort="Ligthart, Lannie" uniqKey="Ligthart L" first="Lannie" last="Ligthart">Lannie Ligthart</name><affiliation><nlm:aff id="A23">Department of Biological Psychology, VU University, Amsterdam, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A24">EMGO+ Institute for Health and Care Research, VU University Medical Center, Amsterdam, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Wedenoja, Juho" sort="Wedenoja, Juho" uniqKey="Wedenoja J" first="Juho" last="Wedenoja">Juho Wedenoja</name><affiliation><nlm:aff id="A25">Department of Public Health, Hjelt Institute, University of Helsinki, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Dunham, Ian" sort="Dunham, Ian" uniqKey="Dunham I" first="Ian" last="Dunham">Ian Dunham</name><affiliation><nlm:aff id="A26">European Bioinformatics Insitute, Wellcome Trust Genome Campus, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Neale, Benjamin M" sort="Neale, Benjamin M" uniqKey="Neale B" first="Benjamin M." last="Neale">Benjamin M. Neale</name><affiliation><nlm:aff id="A3">Analytical and Translational Genetics Unit, Department of Medicine, Massachusetts General Hospital and Harvard Medical School, Boston, MA, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A4">Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA</nlm:aff></affiliation></author><author><name sortKey="Palta, Priit" sort="Palta, Priit" uniqKey="Palta P" first="Priit" last="Palta">Priit Palta</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Cambridge, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A2">Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Hamalainen, Eija" sort="Hamalainen, Eija" uniqKey="Hamalainen E" first="Eija" last="Hamalainen">Eija Hamalainen</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Cambridge, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A2">Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Schurks, Markus" sort="Schurks, Markus" uniqKey="Schurks M" first="Markus" last="Schürks">Markus Schürks</name><affiliation><nlm:aff id="A27">Department of Neurology, University Hospital Essen, Essen, Germany</nlm:aff></affiliation></author><author><name sortKey="Rose, Lynda M" sort="Rose, Lynda M" uniqKey="Rose L" first="Lynda M" last="Rose">Lynda M. Rose</name><affiliation><nlm:aff id="A8">Division of Preventive Medicine, Department of Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, USA</nlm:aff></affiliation></author><author><name sortKey="Buring, Julie E" sort="Buring, Julie E" uniqKey="Buring J" first="Julie E." last="Buring">Julie E. Buring</name><affiliation><nlm:aff id="A8">Division of Preventive Medicine, Department of Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, USA</nlm:aff></affiliation></author><author><name sortKey="Ridker, Paul M" sort="Ridker, Paul M" uniqKey="Ridker P" first="Paul M." last="Ridker">Paul M. Ridker</name><affiliation><nlm:aff id="A8">Division of Preventive Medicine, Department of Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A28">Harvard Medical School, Boston, MA 02215, USA</nlm:aff></affiliation></author><author><name sortKey="Steinberg, Stacy" sort="Steinberg, Stacy" uniqKey="Steinberg S" first="Stacy" last="Steinberg">Stacy Steinberg</name><affiliation><nlm:aff id="A9">deCODE genetics, Reykjavik, Iceland</nlm:aff></affiliation></author><author><name sortKey="Stefansson, Hreinn" sort="Stefansson, Hreinn" uniqKey="Stefansson H" first="Hreinn" last="Stefansson">Hreinn Stefansson</name><affiliation><nlm:aff id="A9">deCODE genetics, Reykjavik, Iceland</nlm:aff></affiliation></author><author><name sortKey="Jakobsson, Finnbogi" sort="Jakobsson, Finnbogi" uniqKey="Jakobsson F" first="Finnbogi" last="Jakobsson">Finnbogi Jakobsson</name><affiliation><nlm:aff id="A69">Department of Neurology, Landspitali University Hospital, Reykjavik, Iceland</nlm:aff></affiliation></author><author><name sortKey="Lawlor, Debbie A" sort="Lawlor, Debbie A" uniqKey="Lawlor D" first="Debbie A." last="Lawlor">Debbie A. Lawlor</name><affiliation><nlm:aff id="A10">MRC Centre for Causal Analyses in Translational Epidemiology, School of Social and Community Medicine, University of Bristol, Bristol, UK</nlm:aff></affiliation></author><author><name sortKey="Evans, David M" sort="Evans, David M" uniqKey="Evans D" first="David M." last="Evans">David M. Evans</name><affiliation><nlm:aff id="A10">MRC Centre for Causal Analyses in Translational Epidemiology, School of Social and Community Medicine, University of Bristol, Bristol, UK</nlm:aff></affiliation></author><author><name sortKey="Ring, Susan M" sort="Ring, Susan M" uniqKey="Ring S" first="Susan M." last="Ring">Susan M. Ring</name><affiliation><nlm:aff id="A10">MRC Centre for Causal Analyses in Translational Epidemiology, School of Social and Community Medicine, University of Bristol, Bristol, UK</nlm:aff></affiliation></author><author><name sortKey="F Rkkil, Markus" sort="F Rkkil, Markus" uniqKey="F Rkkil M" first="Markus" last="F Rkkil">Markus F Rkkil</name><affiliation><nlm:aff id="A11">Department of Neurology, Helsinki University Central Hospital, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Artto, Ville" sort="Artto, Ville" uniqKey="Artto V" first="Ville" last="Artto">Ville Artto</name><affiliation><nlm:aff id="A11">Department of Neurology, Helsinki University Central Hospital, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Kaunisto, Mari A" sort="Kaunisto, Mari A" uniqKey="Kaunisto M" first="Mari A" last="Kaunisto">Mari A. Kaunisto</name><affiliation><nlm:aff id="A2">Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A29">Folkhälsan Research Center, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Freilinger, Tobias" sort="Freilinger, Tobias" uniqKey="Freilinger T" first="Tobias" last="Freilinger">Tobias Freilinger</name><affiliation><nlm:aff id="A12">Institute for Stroke and Dementia Research, Klinikum der Universität München, Ludwig-Maximilians-Universität, Munich, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="A30">Department of Neurology, Klinikum der Universität München, Munich, Germany</nlm:aff></affiliation></author><author><name sortKey="Schoenen, Jean" sort="Schoenen, Jean" uniqKey="Schoenen J" first="Jean" last="Schoenen">Jean Schoenen</name><affiliation><nlm:aff id="A31">Headache Research Unit, Department of Neurology and Groupe Interdisciplinaire de Génoprotéomique Appliquée (GIGA)-Neurosciences, Liège University, Liège, Belgium</nlm:aff></affiliation></author><author><name sortKey="Frants, Rune R" sort="Frants, Rune R" uniqKey="Frants R" first="Rune R." last="Frants">Rune R. Frants</name><affiliation><nlm:aff id="A13">Department of Human Genetics, Leiden University Medical Centre, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Pelzer, Nadine" sort="Pelzer, Nadine" uniqKey="Pelzer N" first="Nadine" last="Pelzer">Nadine Pelzer</name><affiliation><nlm:aff id="A14">Department of Neurology, Leiden University Medical Centre, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Weller, Claudia M" sort="Weller, Claudia M" uniqKey="Weller C" first="Claudia M." last="Weller">Claudia M. Weller</name><affiliation><nlm:aff id="A13">Department of Human Genetics, Leiden University Medical Centre, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Zielman, Ronald" sort="Zielman, Ronald" uniqKey="Zielman R" first="Ronald" last="Zielman">Ronald Zielman</name><affiliation><nlm:aff id="A14">Department of Neurology, Leiden University Medical Centre, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Heath, Andrew C" sort="Heath, Andrew C" uniqKey="Heath A" first="Andrew C." last="Heath">Andrew C. Heath</name><affiliation><nlm:aff id="A32">Department of Psychiatry, Washington University School of Medicine, St. Louis, Missouri, USA</nlm:aff></affiliation></author><author><name sortKey="Madden, Pamela A F" sort="Madden, Pamela A F" uniqKey="Madden P" first="Pamela A. F." last="Madden">Pamela A. F. Madden</name><affiliation><nlm:aff id="A32">Department of Psychiatry, Washington University School of Medicine, St. Louis, Missouri, USA</nlm:aff></affiliation></author><author><name sortKey="Montgomery, Grant W" sort="Montgomery, Grant W" uniqKey="Montgomery G" first="Grant W." last="Montgomery">Grant W. Montgomery</name><affiliation><nlm:aff id="A15">Queensland Institute of Medical Research, Brisbane, Queensland, Australia</nlm:aff></affiliation></author><author><name sortKey="Martin, Nicholas G" sort="Martin, Nicholas G" uniqKey="Martin N" first="Nicholas G." last="Martin">Nicholas G. Martin</name><affiliation><nlm:aff id="A15">Queensland Institute of Medical Research, Brisbane, Queensland, Australia</nlm:aff></affiliation></author><author><name sortKey="Borck, Guntram" sort="Borck, Guntram" uniqKey="Borck G" first="Guntram" last="Borck">Guntram Borck</name><affiliation><nlm:aff id="A16">Institute of Human Genetics, University of Ulm, Ulm, Germany</nlm:aff></affiliation></author><author><name sortKey="Gobel, Hartmut" sort="Gobel, Hartmut" uniqKey="Gobel H" first="Hartmut" last="Göbel">Hartmut Göbel</name><affiliation><nlm:aff id="A33">Kiel Pain and Headache Center, Kiel, Germany</nlm:aff></affiliation></author><author><name sortKey="Heinze, Axel" sort="Heinze, Axel" uniqKey="Heinze A" first="Axel" last="Heinze">Axel Heinze</name><affiliation><nlm:aff id="A33">Kiel Pain and Headache Center, Kiel, Germany</nlm:aff></affiliation></author><author><name sortKey="Heinze Kuhn, Katja" sort="Heinze Kuhn, Katja" uniqKey="Heinze Kuhn K" first="Katja" last="Heinze-Kuhn">Katja Heinze-Kuhn</name><affiliation><nlm:aff id="A33">Kiel Pain and Headache Center, Kiel, Germany</nlm:aff></affiliation></author><author><name sortKey="Williams, Frances M K" sort="Williams, Frances M K" uniqKey="Williams F" first="Frances M. K." last="Williams">Frances M. K. Williams</name><affiliation><nlm:aff id="A17">Department of Twin Research and Genetic Epidemiology, King’s College London, London, UK</nlm:aff></affiliation></author><author><name sortKey="Hartikainen, Anna Liisa" sort="Hartikainen, Anna Liisa" uniqKey="Hartikainen A" first="Anna-Liisa" last="Hartikainen">Anna-Liisa Hartikainen</name><affiliation><nlm:aff id="A34">Department of Clinical Sciences/Obstetrics and Gynecology, University Hospital of Oulu, Oulu, Finland</nlm:aff></affiliation></author><author><name sortKey="Pouta, Anneli" sort="Pouta, Anneli" uniqKey="Pouta A" first="Anneli" last="Pouta">Anneli Pouta</name><affiliation><nlm:aff id="A18">Institute of Health Sciences, University of Oulu, Oulu, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A34">Department of Clinical Sciences/Obstetrics and Gynecology, University Hospital of Oulu, Oulu, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A35">Department of Children, Young People and Families, National Institute for Health and Welfare, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Van Den Ende, Joyce" sort="Van Den Ende, Joyce" uniqKey="Van Den Ende J" first="Joyce" last="Van Den Ende">Joyce Van Den Ende</name><affiliation><nlm:aff id="A19">Genetic Epidemiology Unit, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Uitterlinden, Andre G" sort="Uitterlinden, Andre G" uniqKey="Uitterlinden A" first="Andre G." last="Uitterlinden">Andre G. Uitterlinden</name><affiliation><nlm:aff id="A36">Department of Internal Medicine, Erasmus Medical Center, Rotterdam, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Hofman, Albert" sort="Hofman, Albert" uniqKey="Hofman A" first="Albert" last="Hofman">Albert Hofman</name><affiliation><nlm:aff id="A37">Genetic Epidemiology Unit, Department of Clinical Genetics, Erasmus University Medical Centre, Rotterdam, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Amin, Najaf" sort="Amin, Najaf" uniqKey="Amin N" first="Najaf" last="Amin">Najaf Amin</name><affiliation><nlm:aff id="A19">Genetic Epidemiology Unit, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Hottenga, Jouke Jan" sort="Hottenga, Jouke Jan" uniqKey="Hottenga J" first="Jouke-Jan" last="Hottenga">Jouke-Jan Hottenga</name><affiliation><nlm:aff id="A23">Department of Biological Psychology, VU University, Amsterdam, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Vink, Jacqueline M" sort="Vink, Jacqueline M" uniqKey="Vink J" first="Jacqueline M." last="Vink">Jacqueline M. Vink</name><affiliation><nlm:aff id="A23">Department of Biological Psychology, VU University, Amsterdam, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Heikkil, Kauko" sort="Heikkil, Kauko" uniqKey="Heikkil K" first="Kauko" last="Heikkil">Kauko Heikkil</name><affiliation><nlm:aff id="A25">Department of Public Health, Hjelt Institute, University of Helsinki, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Alexander, Michael" sort="Alexander, Michael" uniqKey="Alexander M" first="Michael" last="Alexander">Michael Alexander</name><affiliation><nlm:aff id="A38">Department of Genomics, Life & Brain Center, University of Bonn, Bonn, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="A39">Institute of Human Genetics, University of Bonn, Bonn, Germany</nlm:aff></affiliation></author><author><name sortKey="Muller Myhsok, Bertram" sort="Muller Myhsok, Bertram" uniqKey="Muller Myhsok B" first="Bertram" last="Muller-Myhsok">Bertram Muller-Myhsok</name><affiliation><nlm:aff id="A40">Max Planck Institute of Psychiatry, Munich, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="A68">Munich Cluster for Systems Neurology (SyNergy), Munich, Germany</nlm:aff></affiliation></author><author><name sortKey="Schreiber, Stefan" sort="Schreiber, Stefan" uniqKey="Schreiber S" first="Stefan" last="Schreiber">Stefan Schreiber</name><affiliation><nlm:aff id="A41">Department of Clinical Molecular Biology, Christian Albrechts University, Kiel, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="A42">Department of Internal Medicine I, Christian Albrechts University, Kiel, Germany</nlm:aff></affiliation></author><author><name sortKey="Meitinger, Thomas" sort="Meitinger, Thomas" uniqKey="Meitinger T" first="Thomas" last="Meitinger">Thomas Meitinger</name><affiliation><nlm:aff id="A43">Institute of Human Genetics, Helmholtz Center Munich, Neuherberg, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="A44">Institute of Human Genetics, Klinikum Rechts der Isar, Technische Universität München, Munich, Germany</nlm:aff></affiliation></author><author><name sortKey="Wichmann, Heinz Erich" sort="Wichmann, Heinz Erich" uniqKey="Wichmann H" first="Heinz Erich" last="Wichmann">Heinz Erich Wichmann</name><affiliation><nlm:aff id="A45">Institut für Medizinische Informationsverarbeitung, Biometrie und Epidemiologie, Ludwig-Maximilians-Universität München, Munich, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="A46">Institute of Epidemiology I, HelmholtzCenter Munich, Neuherberg, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="A47">Klinikum Großhadern, Ludwig-Maximilians-Universität München, Munich, Germany</nlm:aff></affiliation></author><author><name sortKey="Aromaa, Arpo" sort="Aromaa, Arpo" uniqKey="Aromaa A" first="Arpo" last="Aromaa">Arpo Aromaa</name><affiliation><nlm:aff id="A48">National Institute for Health and Welfare, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Eriksson, Johan G" sort="Eriksson, Johan G" uniqKey="Eriksson J" first="Johan G." last="Eriksson">Johan G. Eriksson</name><affiliation><nlm:aff id="A29">Folkhälsan Research Center, Helsinki, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A48">National Institute for Health and Welfare, Helsinki, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A49">Department of General Practice, Helsinki University Central Hospital, Helsinki, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A50">Vaasa Central Hospital, Vaasa, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A51">Department of General Practice and Primary Health Care, University of Helsinki, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Traynor, Bryan" sort="Traynor, Bryan" uniqKey="Traynor B" first="Bryan" last="Traynor">Bryan Traynor</name><affiliation><nlm:aff id="A52">Laboratory of Neurogenetics, National Institute on Aging, National Institutes of Health, Bethesda, MD, USA</nlm:aff></affiliation></author><author><name sortKey="Trabzuni, Daniah" sort="Trabzuni, Daniah" uniqKey="Trabzuni D" first="Daniah" last="Trabzuni">Daniah Trabzuni</name><affiliation><nlm:aff id="A53">Department of Molecular Neuroscience, Institute of Neurology, University College London, London, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A54">Department of Genetics, King Faisal Specialist Hospital and Research Centre, Riyadh, Saudi Arabia</nlm:aff></affiliation></author><author><name sortKey="Rossin, Elizabeth" sort="Rossin, Elizabeth" uniqKey="Rossin E" first="Elizabeth" last="Rossin">Elizabeth Rossin</name><affiliation><nlm:aff id="A3">Analytical and Translational Genetics Unit, Department of Medicine, Massachusetts General Hospital and Harvard Medical School, Boston, MA, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A4">Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A55">Center for Human Genetic Research, Massachusetts General Hospital, Boston, MA, USA</nlm:aff></affiliation></author><author><name sortKey="Lage, Kasper" sort="Lage, Kasper" uniqKey="Lage K" first="Kasper" last="Lage">Kasper Lage</name><affiliation><nlm:aff id="A3">Analytical and Translational Genetics Unit, Department of Medicine, Massachusetts General Hospital and Harvard Medical School, Boston, MA, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A4">Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A56">Pediatric Surgical Research Laboratories, Massachusetts General Hospital for Children, Massachusetts General Hospital, Boston, MA, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A57">Center for Biological Sequence Analysis, Department of Systems Biology, Technical University of Denmark, Lyngby, Denmark</nlm:aff></affiliation><affiliation><nlm:aff id="A58">Center for Protein Research, University of Copenhagen, Copenhagen, Denmark</nlm:aff></affiliation></author><author><name sortKey="Jacobs, Suzanne B R" sort="Jacobs, Suzanne B R" uniqKey="Jacobs S" first="Suzanne B. R." last="Jacobs">Suzanne B. R. Jacobs</name><affiliation><nlm:aff id="A4">Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA</nlm:aff></affiliation></author><author><name sortKey="Gibbs, J Raphael" sort="Gibbs, J Raphael" uniqKey="Gibbs J" first="J. Raphael" last="Gibbs">J. Raphael Gibbs</name><affiliation><nlm:aff id="A52">Laboratory of Neurogenetics, National Institute on Aging, National Institutes of Health, Bethesda, MD, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A53">Department of Molecular Neuroscience, Institute of Neurology, University College London, London, UK</nlm:aff></affiliation></author><author><name sortKey="Birney, Ewan" sort="Birney, Ewan" uniqKey="Birney E" first="Ewan" last="Birney">Ewan Birney</name><affiliation><nlm:aff id="A26">European Bioinformatics Insitute, Wellcome Trust Genome Campus, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Kaprio, Jaakko" sort="Kaprio, Jaakko" uniqKey="Kaprio J" first="Jaakko" last="Kaprio">Jaakko Kaprio</name><affiliation><nlm:aff id="A2">Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A59">Department of Mental Health and Alcohol Research, National Institute for Health and Welfare, Helsinki, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A25">Department of Public Health, Hjelt Institute, University of Helsinki, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Penninx, Brenda W" sort="Penninx, Brenda W" uniqKey="Penninx B" first="Brenda W." last="Penninx">Brenda W. Penninx</name><affiliation><nlm:aff id="A62">Department of Psychiatry, VU University Medical Center, Amsterdam, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A24">EMGO+ Institute for Health and Care Research, VU University Medical Center, Amsterdam, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A61">Department of Psychiatry, University Medical Center Groningen, Groningen, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A60">Department of Psychiatry, Leiden University Medical Center, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Boomsma, Dorret I" sort="Boomsma, Dorret I" uniqKey="Boomsma D" first="Dorret I." last="Boomsma">Dorret I. Boomsma</name><affiliation><nlm:aff id="A23">Department of Biological Psychology, VU University, Amsterdam, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Van Duijn, Cornelia" sort="Van Duijn, Cornelia" uniqKey="Van Duijn C" first="Cornelia" last="Van Duijn">Cornelia Van Duijn</name><affiliation><nlm:aff id="A19">Genetic Epidemiology Unit, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Raitakari, Olli" sort="Raitakari, Olli" uniqKey="Raitakari O" first="Olli" last="Raitakari">Olli Raitakari</name><affiliation><nlm:aff id="A63">Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku University Hospital, Turku, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A64">Department of Clinical Physiology and Nuclear Medicine, Turku University Hospital, Turku, Finland</nlm:aff></affiliation></author><author><name sortKey="Jarvelin, Marjo Riitta" sort="Jarvelin, Marjo Riitta" uniqKey="Jarvelin M" first="Marjo-Riitta" last="Jarvelin">Marjo-Riitta Jarvelin</name><affiliation><nlm:aff id="A65">Department of Epidemiology and Biostatistics, School of Public Health, MRC-HPA Centre for Environment and Health, Faculty of Medicine, Imperial College, London, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A18">Institute of Health Sciences, University of Oulu, Oulu, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A66">Biocenter Oulu, University of Oulu, Oulu, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A35">Department of Children, Young People and Families, National Institute for Health and Welfare, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Zwart, John Anker" sort="Zwart, John Anker" uniqKey="Zwart J" first="John-Anker" last="Zwart">John-Anker Zwart</name><affiliation><nlm:aff id="A5">Department of Neurology, Oslo University Hospital and University of Oslo, Oslo, Norway</nlm:aff></affiliation></author><author><name sortKey="Cherkas, Lynn" sort="Cherkas, Lynn" uniqKey="Cherkas L" first="Lynn" last="Cherkas">Lynn Cherkas</name><affiliation><nlm:aff id="A17">Department of Twin Research and Genetic Epidemiology, King’s College London, London, UK</nlm:aff></affiliation></author><author><name sortKey="Strachan, David P" sort="Strachan, David P" uniqKey="Strachan D" first="David P." last="Strachan">David P. Strachan</name><affiliation><nlm:aff id="A67">Division of Population Health Sciences and Education, St George’s, University of London, London, UK</nlm:aff></affiliation></author><author><name sortKey="Kubisch, Christian" sort="Kubisch, Christian" uniqKey="Kubisch C" first="Christian" last="Kubisch">Christian Kubisch</name><affiliation><nlm:aff id="A16">Institute of Human Genetics, University of Ulm, Ulm, Germany</nlm:aff></affiliation></author><author><name sortKey="Ferrari, Michel D" sort="Ferrari, Michel D" uniqKey="Ferrari M" first="Michel D." last="Ferrari">Michel D. Ferrari</name><affiliation><nlm:aff id="A14">Department of Neurology, Leiden University Medical Centre, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Van Den Maagdenberg, Arn M J M" sort="Van Den Maagdenberg, Arn M J M" uniqKey="Van Den Maagdenberg A" first="Arn M. J. M." last="Van Den Maagdenberg">Arn M. J. M. Van Den Maagdenberg</name><affiliation><nlm:aff id="A13">Department of Human Genetics, Leiden University Medical Centre, Leiden, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A14">Department of Neurology, Leiden University Medical Centre, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Dichgans, Martin" sort="Dichgans, Martin" uniqKey="Dichgans M" first="Martin" last="Dichgans">Martin Dichgans</name><affiliation><nlm:aff id="A12">Institute for Stroke and Dementia Research, Klinikum der Universität München, Ludwig-Maximilians-Universität, Munich, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="A68">Munich Cluster for Systems Neurology (SyNergy), Munich, Germany</nlm:aff></affiliation></author><author><name sortKey="Wessman, Maija" sort="Wessman, Maija" uniqKey="Wessman M" first="Maija" last="Wessman">Maija Wessman</name><affiliation><nlm:aff id="A2">Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A29">Folkhälsan Research Center, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Smith, George Davey" sort="Smith, George Davey" uniqKey="Smith G" first="George Davey" last="Smith">George Davey Smith</name><affiliation><nlm:aff id="A10">MRC Centre for Causal Analyses in Translational Epidemiology, School of Social and Community Medicine, University of Bristol, Bristol, UK</nlm:aff></affiliation></author><author><name sortKey="Stefansson, Kari" sort="Stefansson, Kari" uniqKey="Stefansson K" first="Kari" last="Stefansson">Kari Stefansson</name><affiliation><nlm:aff id="A9">deCODE genetics, Reykjavik, Iceland</nlm:aff></affiliation><affiliation><nlm:aff id="A70">School of Medicine, University of Iceland, Reykjavik, Iceland</nlm:aff></affiliation></author><author><name sortKey="Daly, Mark J" sort="Daly, Mark J" uniqKey="Daly M" first="Mark J." last="Daly">Mark J. Daly</name><affiliation><nlm:aff id="A3">Analytical and Translational Genetics Unit, Department of Medicine, Massachusetts General Hospital and Harvard Medical School, Boston, MA, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A4">Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA</nlm:aff></affiliation></author><author><name sortKey="Nyholt, Dale R" sort="Nyholt, Dale R" uniqKey="Nyholt D" first="Dale R." last="Nyholt">Dale R. Nyholt</name><affiliation><nlm:aff id="A15">Queensland Institute of Medical Research, Brisbane, Queensland, Australia</nlm:aff></affiliation></author><author><name sortKey="Chasman, Daniel" sort="Chasman, Daniel" uniqKey="Chasman D" first="Daniel" last="Chasman">Daniel Chasman</name><affiliation><nlm:aff id="A8">Division of Preventive Medicine, Department of Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A28">Harvard Medical School, Boston, MA 02215, USA</nlm:aff></affiliation></author><author><name sortKey="Palotie, Aarno" sort="Palotie, Aarno" uniqKey="Palotie A" first="Aarno" last="Palotie">Aarno Palotie</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Cambridge, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A2">Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A4">Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA</nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">23793025</idno><idno type="pmc">4041123</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4041123</idno><idno type="RBID">PMC:4041123</idno><idno type="doi">10.1038/ng.2676</idno><date when="2013">2013</date><idno type="wicri:Area/Pmc/Corpus">002E27</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">002E27</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Genome-wide meta-analysis identifies new susceptibility loci for migraine</title><author><name sortKey="Anttila, Verneri" sort="Anttila, Verneri" uniqKey="Anttila V" first="Verneri" last="Anttila">Verneri Anttila</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Cambridge, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A2">Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A3">Analytical and Translational Genetics Unit, Department of Medicine, Massachusetts General Hospital and Harvard Medical School, Boston, MA, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A4">Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA</nlm:aff></affiliation></author><author><name sortKey="Winsvold, Bendik S" sort="Winsvold, Bendik S" uniqKey="Winsvold B" first="Bendik S." last="Winsvold">Bendik S. Winsvold</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Cambridge, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A5">Department of Neurology, Oslo University Hospital and University of Oslo, Oslo, Norway</nlm:aff></affiliation></author><author><name sortKey="Gormley, Padhraig" sort="Gormley, Padhraig" uniqKey="Gormley P" first="Padhraig" last="Gormley">Padhraig Gormley</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Kurth, Tobias" sort="Kurth, Tobias" uniqKey="Kurth T" first="Tobias" last="Kurth">Tobias Kurth</name><affiliation><nlm:aff id="A6">INSERM Unit 708 – Neuroepidemiology, F-33000 Bordeaux, France</nlm:aff></affiliation><affiliation><nlm:aff id="A7">University of Bordeaux, F-33000 Bordeaux, France</nlm:aff></affiliation><affiliation><nlm:aff id="A8">Division of Preventive Medicine, Department of Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, USA</nlm:aff></affiliation></author><author><name sortKey="Bettella, Francesco" sort="Bettella, Francesco" uniqKey="Bettella F" first="Francesco" last="Bettella">Francesco Bettella</name><affiliation><nlm:aff id="A9">deCODE genetics, Reykjavik, Iceland</nlm:aff></affiliation></author><author><name sortKey="Mcmahon, George" sort="Mcmahon, George" uniqKey="Mcmahon G" first="George" last="Mcmahon">George Mcmahon</name><affiliation><nlm:aff id="A10">MRC Centre for Causal Analyses in Translational Epidemiology, School of Social and Community Medicine, University of Bristol, Bristol, UK</nlm:aff></affiliation></author><author><name sortKey="Kallela, Mikko" sort="Kallela, Mikko" uniqKey="Kallela M" first="Mikko" last="Kallela">Mikko Kallela</name><affiliation><nlm:aff id="A11">Department of Neurology, Helsinki University Central Hospital, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Malik, Rainer" sort="Malik, Rainer" uniqKey="Malik R" first="Rainer" last="Malik">Rainer Malik</name><affiliation><nlm:aff id="A12">Institute for Stroke and Dementia Research, Klinikum der Universität München, Ludwig-Maximilians-Universität, Munich, Germany</nlm:aff></affiliation></author><author><name sortKey="De Vries, Boukje" sort="De Vries, Boukje" uniqKey="De Vries B" first="Boukje" last="De Vries">Boukje De Vries</name><affiliation><nlm:aff id="A13">Department of Human Genetics, Leiden University Medical Centre, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Terwindt, Gisela" sort="Terwindt, Gisela" uniqKey="Terwindt G" first="Gisela" last="Terwindt">Gisela Terwindt</name><affiliation><nlm:aff id="A14">Department of Neurology, Leiden University Medical Centre, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Medland, Sarah E" sort="Medland, Sarah E" uniqKey="Medland S" first="Sarah E." last="Medland">Sarah E. Medland</name><affiliation><nlm:aff id="A15">Queensland Institute of Medical Research, Brisbane, Queensland, Australia</nlm:aff></affiliation></author><author><name sortKey="Todt, Unda" sort="Todt, Unda" uniqKey="Todt U" first="Unda" last="Todt">Unda Todt</name><affiliation><nlm:aff id="A16">Institute of Human Genetics, University of Ulm, Ulm, Germany</nlm:aff></affiliation></author><author><name sortKey="Mcardle, Wendy L" sort="Mcardle, Wendy L" uniqKey="Mcardle W" first="Wendy L." last="Mcardle">Wendy L. Mcardle</name><affiliation><nlm:aff id="A10">MRC Centre for Causal Analyses in Translational Epidemiology, School of Social and Community Medicine, University of Bristol, Bristol, UK</nlm:aff></affiliation></author><author><name sortKey="Quaye, Lydia" sort="Quaye, Lydia" uniqKey="Quaye L" first="Lydia" last="Quaye">Lydia Quaye</name><affiliation><nlm:aff id="A17">Department of Twin Research and Genetic Epidemiology, King’s College London, London, UK</nlm:aff></affiliation></author><author><name sortKey="Koiranen, Markku" sort="Koiranen, Markku" uniqKey="Koiranen M" first="Markku" last="Koiranen">Markku Koiranen</name><affiliation><nlm:aff id="A18">Institute of Health Sciences, University of Oulu, Oulu, Finland</nlm:aff></affiliation></author><author><name sortKey="Ikram, M Arfan" sort="Ikram, M Arfan" uniqKey="Ikram M" first="M. Arfan" last="Ikram">M. Arfan Ikram</name><affiliation><nlm:aff id="A19">Genetic Epidemiology Unit, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, the Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A20">Department of Radiology Erasmus University Medical Centre, Rotterdam, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A21">Department of Neurology Erasmus University Medical Centre, Rotterdam, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Lehtim Ki, Terho" sort="Lehtim Ki, Terho" uniqKey="Lehtim Ki T" first="Terho" last="Lehtim Ki">Terho Lehtim Ki</name><affiliation><nlm:aff id="A22">Department of Clinical Chemistry, Fimlab Laboratories and University of Tampere School of Medicine, Tampere, Finland</nlm:aff></affiliation></author><author><name sortKey="Stam, Anine H" sort="Stam, Anine H" uniqKey="Stam A" first="Anine H." last="Stam">Anine H. Stam</name><affiliation><nlm:aff id="A14">Department of Neurology, Leiden University Medical Centre, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Ligthart, Lannie" sort="Ligthart, Lannie" uniqKey="Ligthart L" first="Lannie" last="Ligthart">Lannie Ligthart</name><affiliation><nlm:aff id="A23">Department of Biological Psychology, VU University, Amsterdam, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A24">EMGO+ Institute for Health and Care Research, VU University Medical Center, Amsterdam, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Wedenoja, Juho" sort="Wedenoja, Juho" uniqKey="Wedenoja J" first="Juho" last="Wedenoja">Juho Wedenoja</name><affiliation><nlm:aff id="A25">Department of Public Health, Hjelt Institute, University of Helsinki, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Dunham, Ian" sort="Dunham, Ian" uniqKey="Dunham I" first="Ian" last="Dunham">Ian Dunham</name><affiliation><nlm:aff id="A26">European Bioinformatics Insitute, Wellcome Trust Genome Campus, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Neale, Benjamin M" sort="Neale, Benjamin M" uniqKey="Neale B" first="Benjamin M." last="Neale">Benjamin M. Neale</name><affiliation><nlm:aff id="A3">Analytical and Translational Genetics Unit, Department of Medicine, Massachusetts General Hospital and Harvard Medical School, Boston, MA, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A4">Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA</nlm:aff></affiliation></author><author><name sortKey="Palta, Priit" sort="Palta, Priit" uniqKey="Palta P" first="Priit" last="Palta">Priit Palta</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Cambridge, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A2">Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Hamalainen, Eija" sort="Hamalainen, Eija" uniqKey="Hamalainen E" first="Eija" last="Hamalainen">Eija Hamalainen</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Cambridge, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A2">Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Schurks, Markus" sort="Schurks, Markus" uniqKey="Schurks M" first="Markus" last="Schürks">Markus Schürks</name><affiliation><nlm:aff id="A27">Department of Neurology, University Hospital Essen, Essen, Germany</nlm:aff></affiliation></author><author><name sortKey="Rose, Lynda M" sort="Rose, Lynda M" uniqKey="Rose L" first="Lynda M" last="Rose">Lynda M. Rose</name><affiliation><nlm:aff id="A8">Division of Preventive Medicine, Department of Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, USA</nlm:aff></affiliation></author><author><name sortKey="Buring, Julie E" sort="Buring, Julie E" uniqKey="Buring J" first="Julie E." last="Buring">Julie E. Buring</name><affiliation><nlm:aff id="A8">Division of Preventive Medicine, Department of Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, USA</nlm:aff></affiliation></author><author><name sortKey="Ridker, Paul M" sort="Ridker, Paul M" uniqKey="Ridker P" first="Paul M." last="Ridker">Paul M. Ridker</name><affiliation><nlm:aff id="A8">Division of Preventive Medicine, Department of Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A28">Harvard Medical School, Boston, MA 02215, USA</nlm:aff></affiliation></author><author><name sortKey="Steinberg, Stacy" sort="Steinberg, Stacy" uniqKey="Steinberg S" first="Stacy" last="Steinberg">Stacy Steinberg</name><affiliation><nlm:aff id="A9">deCODE genetics, Reykjavik, Iceland</nlm:aff></affiliation></author><author><name sortKey="Stefansson, Hreinn" sort="Stefansson, Hreinn" uniqKey="Stefansson H" first="Hreinn" last="Stefansson">Hreinn Stefansson</name><affiliation><nlm:aff id="A9">deCODE genetics, Reykjavik, Iceland</nlm:aff></affiliation></author><author><name sortKey="Jakobsson, Finnbogi" sort="Jakobsson, Finnbogi" uniqKey="Jakobsson F" first="Finnbogi" last="Jakobsson">Finnbogi Jakobsson</name><affiliation><nlm:aff id="A69">Department of Neurology, Landspitali University Hospital, Reykjavik, Iceland</nlm:aff></affiliation></author><author><name sortKey="Lawlor, Debbie A" sort="Lawlor, Debbie A" uniqKey="Lawlor D" first="Debbie A." last="Lawlor">Debbie A. Lawlor</name><affiliation><nlm:aff id="A10">MRC Centre for Causal Analyses in Translational Epidemiology, School of Social and Community Medicine, University of Bristol, Bristol, UK</nlm:aff></affiliation></author><author><name sortKey="Evans, David M" sort="Evans, David M" uniqKey="Evans D" first="David M." last="Evans">David M. Evans</name><affiliation><nlm:aff id="A10">MRC Centre for Causal Analyses in Translational Epidemiology, School of Social and Community Medicine, University of Bristol, Bristol, UK</nlm:aff></affiliation></author><author><name sortKey="Ring, Susan M" sort="Ring, Susan M" uniqKey="Ring S" first="Susan M." last="Ring">Susan M. Ring</name><affiliation><nlm:aff id="A10">MRC Centre for Causal Analyses in Translational Epidemiology, School of Social and Community Medicine, University of Bristol, Bristol, UK</nlm:aff></affiliation></author><author><name sortKey="F Rkkil, Markus" sort="F Rkkil, Markus" uniqKey="F Rkkil M" first="Markus" last="F Rkkil">Markus F Rkkil</name><affiliation><nlm:aff id="A11">Department of Neurology, Helsinki University Central Hospital, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Artto, Ville" sort="Artto, Ville" uniqKey="Artto V" first="Ville" last="Artto">Ville Artto</name><affiliation><nlm:aff id="A11">Department of Neurology, Helsinki University Central Hospital, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Kaunisto, Mari A" sort="Kaunisto, Mari A" uniqKey="Kaunisto M" first="Mari A" last="Kaunisto">Mari A. Kaunisto</name><affiliation><nlm:aff id="A2">Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A29">Folkhälsan Research Center, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Freilinger, Tobias" sort="Freilinger, Tobias" uniqKey="Freilinger T" first="Tobias" last="Freilinger">Tobias Freilinger</name><affiliation><nlm:aff id="A12">Institute for Stroke and Dementia Research, Klinikum der Universität München, Ludwig-Maximilians-Universität, Munich, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="A30">Department of Neurology, Klinikum der Universität München, Munich, Germany</nlm:aff></affiliation></author><author><name sortKey="Schoenen, Jean" sort="Schoenen, Jean" uniqKey="Schoenen J" first="Jean" last="Schoenen">Jean Schoenen</name><affiliation><nlm:aff id="A31">Headache Research Unit, Department of Neurology and Groupe Interdisciplinaire de Génoprotéomique Appliquée (GIGA)-Neurosciences, Liège University, Liège, Belgium</nlm:aff></affiliation></author><author><name sortKey="Frants, Rune R" sort="Frants, Rune R" uniqKey="Frants R" first="Rune R." last="Frants">Rune R. Frants</name><affiliation><nlm:aff id="A13">Department of Human Genetics, Leiden University Medical Centre, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Pelzer, Nadine" sort="Pelzer, Nadine" uniqKey="Pelzer N" first="Nadine" last="Pelzer">Nadine Pelzer</name><affiliation><nlm:aff id="A14">Department of Neurology, Leiden University Medical Centre, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Weller, Claudia M" sort="Weller, Claudia M" uniqKey="Weller C" first="Claudia M." last="Weller">Claudia M. Weller</name><affiliation><nlm:aff id="A13">Department of Human Genetics, Leiden University Medical Centre, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Zielman, Ronald" sort="Zielman, Ronald" uniqKey="Zielman R" first="Ronald" last="Zielman">Ronald Zielman</name><affiliation><nlm:aff id="A14">Department of Neurology, Leiden University Medical Centre, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Heath, Andrew C" sort="Heath, Andrew C" uniqKey="Heath A" first="Andrew C." last="Heath">Andrew C. Heath</name><affiliation><nlm:aff id="A32">Department of Psychiatry, Washington University School of Medicine, St. Louis, Missouri, USA</nlm:aff></affiliation></author><author><name sortKey="Madden, Pamela A F" sort="Madden, Pamela A F" uniqKey="Madden P" first="Pamela A. F." last="Madden">Pamela A. F. Madden</name><affiliation><nlm:aff id="A32">Department of Psychiatry, Washington University School of Medicine, St. Louis, Missouri, USA</nlm:aff></affiliation></author><author><name sortKey="Montgomery, Grant W" sort="Montgomery, Grant W" uniqKey="Montgomery G" first="Grant W." last="Montgomery">Grant W. Montgomery</name><affiliation><nlm:aff id="A15">Queensland Institute of Medical Research, Brisbane, Queensland, Australia</nlm:aff></affiliation></author><author><name sortKey="Martin, Nicholas G" sort="Martin, Nicholas G" uniqKey="Martin N" first="Nicholas G." last="Martin">Nicholas G. Martin</name><affiliation><nlm:aff id="A15">Queensland Institute of Medical Research, Brisbane, Queensland, Australia</nlm:aff></affiliation></author><author><name sortKey="Borck, Guntram" sort="Borck, Guntram" uniqKey="Borck G" first="Guntram" last="Borck">Guntram Borck</name><affiliation><nlm:aff id="A16">Institute of Human Genetics, University of Ulm, Ulm, Germany</nlm:aff></affiliation></author><author><name sortKey="Gobel, Hartmut" sort="Gobel, Hartmut" uniqKey="Gobel H" first="Hartmut" last="Göbel">Hartmut Göbel</name><affiliation><nlm:aff id="A33">Kiel Pain and Headache Center, Kiel, Germany</nlm:aff></affiliation></author><author><name sortKey="Heinze, Axel" sort="Heinze, Axel" uniqKey="Heinze A" first="Axel" last="Heinze">Axel Heinze</name><affiliation><nlm:aff id="A33">Kiel Pain and Headache Center, Kiel, Germany</nlm:aff></affiliation></author><author><name sortKey="Heinze Kuhn, Katja" sort="Heinze Kuhn, Katja" uniqKey="Heinze Kuhn K" first="Katja" last="Heinze-Kuhn">Katja Heinze-Kuhn</name><affiliation><nlm:aff id="A33">Kiel Pain and Headache Center, Kiel, Germany</nlm:aff></affiliation></author><author><name sortKey="Williams, Frances M K" sort="Williams, Frances M K" uniqKey="Williams F" first="Frances M. K." last="Williams">Frances M. K. Williams</name><affiliation><nlm:aff id="A17">Department of Twin Research and Genetic Epidemiology, King’s College London, London, UK</nlm:aff></affiliation></author><author><name sortKey="Hartikainen, Anna Liisa" sort="Hartikainen, Anna Liisa" uniqKey="Hartikainen A" first="Anna-Liisa" last="Hartikainen">Anna-Liisa Hartikainen</name><affiliation><nlm:aff id="A34">Department of Clinical Sciences/Obstetrics and Gynecology, University Hospital of Oulu, Oulu, Finland</nlm:aff></affiliation></author><author><name sortKey="Pouta, Anneli" sort="Pouta, Anneli" uniqKey="Pouta A" first="Anneli" last="Pouta">Anneli Pouta</name><affiliation><nlm:aff id="A18">Institute of Health Sciences, University of Oulu, Oulu, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A34">Department of Clinical Sciences/Obstetrics and Gynecology, University Hospital of Oulu, Oulu, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A35">Department of Children, Young People and Families, National Institute for Health and Welfare, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Van Den Ende, Joyce" sort="Van Den Ende, Joyce" uniqKey="Van Den Ende J" first="Joyce" last="Van Den Ende">Joyce Van Den Ende</name><affiliation><nlm:aff id="A19">Genetic Epidemiology Unit, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Uitterlinden, Andre G" sort="Uitterlinden, Andre G" uniqKey="Uitterlinden A" first="Andre G." last="Uitterlinden">Andre G. Uitterlinden</name><affiliation><nlm:aff id="A36">Department of Internal Medicine, Erasmus Medical Center, Rotterdam, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Hofman, Albert" sort="Hofman, Albert" uniqKey="Hofman A" first="Albert" last="Hofman">Albert Hofman</name><affiliation><nlm:aff id="A37">Genetic Epidemiology Unit, Department of Clinical Genetics, Erasmus University Medical Centre, Rotterdam, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Amin, Najaf" sort="Amin, Najaf" uniqKey="Amin N" first="Najaf" last="Amin">Najaf Amin</name><affiliation><nlm:aff id="A19">Genetic Epidemiology Unit, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Hottenga, Jouke Jan" sort="Hottenga, Jouke Jan" uniqKey="Hottenga J" first="Jouke-Jan" last="Hottenga">Jouke-Jan Hottenga</name><affiliation><nlm:aff id="A23">Department of Biological Psychology, VU University, Amsterdam, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Vink, Jacqueline M" sort="Vink, Jacqueline M" uniqKey="Vink J" first="Jacqueline M." last="Vink">Jacqueline M. Vink</name><affiliation><nlm:aff id="A23">Department of Biological Psychology, VU University, Amsterdam, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Heikkil, Kauko" sort="Heikkil, Kauko" uniqKey="Heikkil K" first="Kauko" last="Heikkil">Kauko Heikkil</name><affiliation><nlm:aff id="A25">Department of Public Health, Hjelt Institute, University of Helsinki, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Alexander, Michael" sort="Alexander, Michael" uniqKey="Alexander M" first="Michael" last="Alexander">Michael Alexander</name><affiliation><nlm:aff id="A38">Department of Genomics, Life & Brain Center, University of Bonn, Bonn, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="A39">Institute of Human Genetics, University of Bonn, Bonn, Germany</nlm:aff></affiliation></author><author><name sortKey="Muller Myhsok, Bertram" sort="Muller Myhsok, Bertram" uniqKey="Muller Myhsok B" first="Bertram" last="Muller-Myhsok">Bertram Muller-Myhsok</name><affiliation><nlm:aff id="A40">Max Planck Institute of Psychiatry, Munich, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="A68">Munich Cluster for Systems Neurology (SyNergy), Munich, Germany</nlm:aff></affiliation></author><author><name sortKey="Schreiber, Stefan" sort="Schreiber, Stefan" uniqKey="Schreiber S" first="Stefan" last="Schreiber">Stefan Schreiber</name><affiliation><nlm:aff id="A41">Department of Clinical Molecular Biology, Christian Albrechts University, Kiel, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="A42">Department of Internal Medicine I, Christian Albrechts University, Kiel, Germany</nlm:aff></affiliation></author><author><name sortKey="Meitinger, Thomas" sort="Meitinger, Thomas" uniqKey="Meitinger T" first="Thomas" last="Meitinger">Thomas Meitinger</name><affiliation><nlm:aff id="A43">Institute of Human Genetics, Helmholtz Center Munich, Neuherberg, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="A44">Institute of Human Genetics, Klinikum Rechts der Isar, Technische Universität München, Munich, Germany</nlm:aff></affiliation></author><author><name sortKey="Wichmann, Heinz Erich" sort="Wichmann, Heinz Erich" uniqKey="Wichmann H" first="Heinz Erich" last="Wichmann">Heinz Erich Wichmann</name><affiliation><nlm:aff id="A45">Institut für Medizinische Informationsverarbeitung, Biometrie und Epidemiologie, Ludwig-Maximilians-Universität München, Munich, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="A46">Institute of Epidemiology I, HelmholtzCenter Munich, Neuherberg, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="A47">Klinikum Großhadern, Ludwig-Maximilians-Universität München, Munich, Germany</nlm:aff></affiliation></author><author><name sortKey="Aromaa, Arpo" sort="Aromaa, Arpo" uniqKey="Aromaa A" first="Arpo" last="Aromaa">Arpo Aromaa</name><affiliation><nlm:aff id="A48">National Institute for Health and Welfare, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Eriksson, Johan G" sort="Eriksson, Johan G" uniqKey="Eriksson J" first="Johan G." last="Eriksson">Johan G. Eriksson</name><affiliation><nlm:aff id="A29">Folkhälsan Research Center, Helsinki, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A48">National Institute for Health and Welfare, Helsinki, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A49">Department of General Practice, Helsinki University Central Hospital, Helsinki, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A50">Vaasa Central Hospital, Vaasa, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A51">Department of General Practice and Primary Health Care, University of Helsinki, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Traynor, Bryan" sort="Traynor, Bryan" uniqKey="Traynor B" first="Bryan" last="Traynor">Bryan Traynor</name><affiliation><nlm:aff id="A52">Laboratory of Neurogenetics, National Institute on Aging, National Institutes of Health, Bethesda, MD, USA</nlm:aff></affiliation></author><author><name sortKey="Trabzuni, Daniah" sort="Trabzuni, Daniah" uniqKey="Trabzuni D" first="Daniah" last="Trabzuni">Daniah Trabzuni</name><affiliation><nlm:aff id="A53">Department of Molecular Neuroscience, Institute of Neurology, University College London, London, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A54">Department of Genetics, King Faisal Specialist Hospital and Research Centre, Riyadh, Saudi Arabia</nlm:aff></affiliation></author><author><name sortKey="Rossin, Elizabeth" sort="Rossin, Elizabeth" uniqKey="Rossin E" first="Elizabeth" last="Rossin">Elizabeth Rossin</name><affiliation><nlm:aff id="A3">Analytical and Translational Genetics Unit, Department of Medicine, Massachusetts General Hospital and Harvard Medical School, Boston, MA, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A4">Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A55">Center for Human Genetic Research, Massachusetts General Hospital, Boston, MA, USA</nlm:aff></affiliation></author><author><name sortKey="Lage, Kasper" sort="Lage, Kasper" uniqKey="Lage K" first="Kasper" last="Lage">Kasper Lage</name><affiliation><nlm:aff id="A3">Analytical and Translational Genetics Unit, Department of Medicine, Massachusetts General Hospital and Harvard Medical School, Boston, MA, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A4">Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A56">Pediatric Surgical Research Laboratories, Massachusetts General Hospital for Children, Massachusetts General Hospital, Boston, MA, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A57">Center for Biological Sequence Analysis, Department of Systems Biology, Technical University of Denmark, Lyngby, Denmark</nlm:aff></affiliation><affiliation><nlm:aff id="A58">Center for Protein Research, University of Copenhagen, Copenhagen, Denmark</nlm:aff></affiliation></author><author><name sortKey="Jacobs, Suzanne B R" sort="Jacobs, Suzanne B R" uniqKey="Jacobs S" first="Suzanne B. R." last="Jacobs">Suzanne B. R. Jacobs</name><affiliation><nlm:aff id="A4">Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA</nlm:aff></affiliation></author><author><name sortKey="Gibbs, J Raphael" sort="Gibbs, J Raphael" uniqKey="Gibbs J" first="J. Raphael" last="Gibbs">J. Raphael Gibbs</name><affiliation><nlm:aff id="A52">Laboratory of Neurogenetics, National Institute on Aging, National Institutes of Health, Bethesda, MD, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A53">Department of Molecular Neuroscience, Institute of Neurology, University College London, London, UK</nlm:aff></affiliation></author><author><name sortKey="Birney, Ewan" sort="Birney, Ewan" uniqKey="Birney E" first="Ewan" last="Birney">Ewan Birney</name><affiliation><nlm:aff id="A26">European Bioinformatics Insitute, Wellcome Trust Genome Campus, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Kaprio, Jaakko" sort="Kaprio, Jaakko" uniqKey="Kaprio J" first="Jaakko" last="Kaprio">Jaakko Kaprio</name><affiliation><nlm:aff id="A2">Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A59">Department of Mental Health and Alcohol Research, National Institute for Health and Welfare, Helsinki, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A25">Department of Public Health, Hjelt Institute, University of Helsinki, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Penninx, Brenda W" sort="Penninx, Brenda W" uniqKey="Penninx B" first="Brenda W." last="Penninx">Brenda W. Penninx</name><affiliation><nlm:aff id="A62">Department of Psychiatry, VU University Medical Center, Amsterdam, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A24">EMGO+ Institute for Health and Care Research, VU University Medical Center, Amsterdam, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A61">Department of Psychiatry, University Medical Center Groningen, Groningen, The Netherlands</nlm:aff></affiliation><affiliation><nlm:aff id="A60">Department of Psychiatry, Leiden University Medical Center, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Boomsma, Dorret I" sort="Boomsma, Dorret I" uniqKey="Boomsma D" first="Dorret I." last="Boomsma">Dorret I. Boomsma</name><affiliation><nlm:aff id="A23">Department of Biological Psychology, VU University, Amsterdam, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Van Duijn, Cornelia" sort="Van Duijn, Cornelia" uniqKey="Van Duijn C" first="Cornelia" last="Van Duijn">Cornelia Van Duijn</name><affiliation><nlm:aff id="A19">Genetic Epidemiology Unit, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Raitakari, Olli" sort="Raitakari, Olli" uniqKey="Raitakari O" first="Olli" last="Raitakari">Olli Raitakari</name><affiliation><nlm:aff id="A63">Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku University Hospital, Turku, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A64">Department of Clinical Physiology and Nuclear Medicine, Turku University Hospital, Turku, Finland</nlm:aff></affiliation></author><author><name sortKey="Jarvelin, Marjo Riitta" sort="Jarvelin, Marjo Riitta" uniqKey="Jarvelin M" first="Marjo-Riitta" last="Jarvelin">Marjo-Riitta Jarvelin</name><affiliation><nlm:aff id="A65">Department of Epidemiology and Biostatistics, School of Public Health, MRC-HPA Centre for Environment and Health, Faculty of Medicine, Imperial College, London, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A18">Institute of Health Sciences, University of Oulu, Oulu, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A66">Biocenter Oulu, University of Oulu, Oulu, Finland</nlm:aff></affiliation><affiliation><nlm:aff id="A35">Department of Children, Young People and Families, National Institute for Health and Welfare, Helsinki, Finland</nlm:aff></affiliation></author><author><name sortKey="Zwart, John Anker" sort="Zwart, John Anker" uniqKey="Zwart J" first="John-Anker" last="Zwart">John-Anker Zwart</name><affiliation><nlm:aff id="A5">Department of Neurology, Oslo University Hospital and University of Oslo, Oslo, Norway</nlm:aff></affiliation></author><author><name sortKey="Cherkas, Lynn" sort="Cherkas, Lynn" uniqKey="Cherkas L" first="Lynn" last="Cherkas">Lynn Cherkas</name><affiliation><nlm:aff id="A17">Department of Twin Research and Genetic Epidemiology, King’s College London, London, UK</nlm:aff></affiliation></author><author><name sortKey="Strachan, David P" sort="Strachan, David P" uniqKey="Strachan D" first="David P." last="Strachan">David P. Strachan</name><affiliation><nlm:aff id="A67">Division of Population Health Sciences and Education, St George’s, University of London, London, UK</nlm:aff></affiliation></author><author><name sortKey="Kubisch, Christian" sort="Kubisch, Christian" uniqKey="Kubisch C" first="Christian" last="Kubisch">Christian Kubisch</name><affiliation><nlm:aff id="A16">Institute of Human Genetics, University of Ulm, Ulm, Germany</nlm:aff></affiliation></author><author><name sortKey="Ferrari, Michel D" sort="Ferrari, Michel D" uniqKey="Ferrari M" first="Michel D." last="Ferrari">Michel D. Ferrari</name><affiliation><nlm:aff id="A14">Department of Neurology, Leiden University Medical Centre, Leiden, The Netherlands</nlm:aff></affiliation></author><author><name sortKey="Van Den Maagdenberg, Arn M J M" sort="Van Den Maagdenberg, Arn M J M" uniqKey="Van Den Maagdenberg A" first="Arn M. J. M." last="Van Den Maagdenberg">Arn M. J. M. 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contrib-type="author"><name><surname>Smith</surname><given-names>George Davey</given-names></name><xref ref-type="aff" rid="A10">10</xref></contrib><contrib contrib-type="author"><name><surname>Stefansson</surname><given-names>Kari</given-names></name><xref ref-type="aff" rid="A9">9</xref><xref ref-type="aff" rid="A70">Xxx2</xref></contrib><contrib contrib-type="author"><name><surname>Daly</surname><given-names>Mark J.</given-names></name><xref ref-type="aff" rid="A3">3</xref><xref ref-type="aff" rid="A4">4</xref></contrib><contrib contrib-type="author"><name><surname>Nyholt</surname><given-names>Dale R.</given-names></name><xref ref-type="aff" rid="A15">15</xref></contrib><contrib contrib-type="author"><name><surname>Chasman</surname><given-names>Daniel</given-names></name><xref ref-type="aff" rid="A8">8</xref><xref ref-type="aff" rid="A28">28</xref></contrib><contrib contrib-type="author"><name><surname>Palotie</surname><given-names>Aarno</given-names></name><xref ref-type="aff" rid="A1">1</xref><xref ref-type="aff" rid="A2">2</xref><xref ref-type="aff" rid="A4">4</xref></contrib><on-behalf-of>for the International Headache Genetics Consortium</on-behalf-of><xref ref-type="author-notes" rid="FN1">55</xref></contrib-group><aff id="A1"><label>1</label>Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Cambridge, UK</aff><aff id="A2"><label>2</label>Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland</aff><aff id="A3"><label>3</label>Analytical and Translational Genetics Unit, Department of Medicine, Massachusetts General Hospital and Harvard Medical School, Boston, MA, USA</aff><aff id="A4"><label>4</label>Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA</aff><aff id="A5"><label>5</label>Department of Neurology, Oslo University Hospital and University of Oslo, Oslo, Norway</aff><aff id="A6"><label>6</label>INSERM Unit 708 – Neuroepidemiology, F-33000 Bordeaux, France</aff><aff id="A7"><label>7</label>University of Bordeaux, F-33000 Bordeaux, France</aff><aff id="A8"><label>8</label>Division of Preventive Medicine, Department of Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, USA</aff><aff id="A9"><label>9</label>deCODE genetics, Reykjavik, Iceland</aff><aff id="A10"><label>10</label>MRC Centre for Causal Analyses in Translational Epidemiology, School of Social and Community Medicine, University of Bristol, Bristol, UK</aff><aff id="A11"><label>11</label>Department of Neurology, Helsinki University Central Hospital, Helsinki, Finland</aff><aff id="A12"><label>12</label>Institute for Stroke and Dementia Research, Klinikum der Universität München, Ludwig-Maximilians-Universität, Munich, Germany</aff><aff id="A13"><label>13</label>Department of Human Genetics, Leiden University Medical Centre, Leiden, The Netherlands</aff><aff id="A14"><label>14</label>Department of Neurology, Leiden University Medical Centre, Leiden, The Netherlands</aff><aff id="A15"><label>15</label>Queensland Institute of Medical Research, Brisbane, Queensland, Australia</aff><aff id="A16"><label>16</label>Institute of Human Genetics, University of Ulm, Ulm, Germany</aff><aff id="A17"><label>17</label>Department of Twin Research and Genetic Epidemiology, King’s College London, London, UK</aff><aff id="A18"><label>18</label>Institute of Health Sciences, University of Oulu, Oulu, Finland</aff><aff id="A19"><label>19</label>Genetic Epidemiology Unit, Department of Epidemiology, Erasmus University Medical Center, Rotterdam, the Netherlands</aff><aff id="A20"><label>20</label>Department of Radiology Erasmus University Medical Centre, Rotterdam, The Netherlands</aff><aff id="A21"><label>21</label>Department of Neurology Erasmus University Medical Centre, Rotterdam, The Netherlands</aff><aff id="A22"><label>22</label>Department of Clinical Chemistry, Fimlab Laboratories and University of Tampere School of Medicine, Tampere, Finland</aff><aff id="A23"><label>23</label>Department of Biological Psychology, VU University, Amsterdam, The Netherlands</aff><aff id="A24"><label>24</label>EMGO+ Institute for Health and Care Research, VU University Medical Center, Amsterdam, The Netherlands</aff><aff id="A25"><label>25</label>Department of Public Health, Hjelt Institute, University of Helsinki, Helsinki, Finland</aff><aff id="A26"><label>26</label>European Bioinformatics Insitute, Wellcome Trust Genome Campus, Cambridge, UK</aff><aff id="A27"><label>27</label>Department of Neurology, University Hospital Essen, Essen, Germany</aff><aff id="A28"><label>28</label>Harvard Medical School, Boston, MA 02215, USA</aff><aff id="A29"><label>29</label>Folkhälsan Research Center, Helsinki, Finland</aff><aff id="A30"><label>30</label>Department of Neurology, Klinikum der Universität München, Munich, Germany</aff><aff id="A31"><label>31</label>Headache Research Unit, Department of Neurology and Groupe Interdisciplinaire de Génoprotéomique Appliquée (GIGA)-Neurosciences, Liège University, Liège, Belgium</aff><aff id="A32"><label>32</label>Department of Psychiatry, Washington University School of Medicine, St. Louis, Missouri, USA</aff><aff id="A33"><label>33</label>Kiel Pain and Headache Center, Kiel, Germany</aff><aff id="A34"><label>34</label>Department of Clinical Sciences/Obstetrics and Gynecology, University Hospital of Oulu, Oulu, Finland</aff><aff id="A35"><label>35</label>Department of Children, Young People and Families, National Institute for Health and Welfare, Helsinki, Finland</aff><aff id="A36"><label>36</label>Department of Internal Medicine, Erasmus Medical Center, Rotterdam, The Netherlands</aff><aff id="A37"><label>37</label>Genetic Epidemiology Unit, Department of Clinical Genetics, Erasmus University Medical Centre, Rotterdam, The Netherlands</aff><aff id="A38"><label>38</label>Department of Genomics, Life & Brain Center, University of Bonn, Bonn, Germany</aff><aff id="A39"><label>39</label>Institute of Human Genetics, University of Bonn, Bonn, Germany</aff><aff id="A40"><label>40</label>Max Planck Institute of Psychiatry, Munich, Germany</aff><aff id="A41"><label>41</label>Department of Clinical Molecular Biology, Christian Albrechts University, Kiel, Germany</aff><aff id="A42"><label>42</label>Department of Internal Medicine I, Christian Albrechts University, Kiel, Germany</aff><aff id="A43"><label>43</label>Institute of Human Genetics, Helmholtz Center Munich, Neuherberg, Germany</aff><aff id="A44"><label>44</label>Institute of Human Genetics, Klinikum Rechts der Isar, Technische Universität München, Munich, Germany</aff><aff id="A45"><label>45</label>Institut für Medizinische Informationsverarbeitung, Biometrie und Epidemiologie, Ludwig-Maximilians-Universität München, Munich, Germany</aff><aff id="A46"><label>46</label>Institute of Epidemiology I, HelmholtzCenter Munich, Neuherberg, Germany</aff><aff id="A47"><label>47</label>Klinikum Großhadern, Ludwig-Maximilians-Universität München, Munich, Germany</aff><aff id="A48"><label>48</label>National Institute for Health and Welfare, Helsinki, Finland</aff><aff id="A49"><label>49</label>Department of General Practice, Helsinki University Central Hospital, Helsinki, Finland</aff><aff id="A50"><label>50</label>Vaasa Central Hospital, Vaasa, Finland</aff><aff id="A51"><label>51</label>Department of General Practice and Primary Health Care, University of Helsinki, Helsinki, Finland</aff><aff id="A52"><label>52</label>Laboratory of Neurogenetics, National Institute on Aging, National Institutes of Health, Bethesda, MD, USA</aff><aff id="A53"><label>53</label>Department of Molecular Neuroscience, Institute of Neurology, University College London, London, UK</aff><aff id="A54"><label>54</label>Department of Genetics, King Faisal Specialist Hospital and Research Centre, Riyadh, Saudi Arabia</aff><aff id="A55"><label>56</label>Center for Human Genetic Research, Massachusetts General Hospital, Boston, MA, USA</aff><aff id="A56"><label>57</label>Pediatric Surgical Research Laboratories, Massachusetts General Hospital for Children, Massachusetts General Hospital, Boston, MA, USA</aff><aff id="A57"><label>58</label>Center for Biological Sequence Analysis, Department of Systems Biology, Technical University of Denmark, Lyngby, Denmark</aff><aff id="A58"><label>59</label>Center for Protein Research, University of Copenhagen, Copenhagen, Denmark</aff><aff id="A59"><label>60</label>Department of Mental Health and Alcohol Research, National Institute for Health and Welfare, Helsinki, Finland</aff><aff id="A60"><label>61</label>Department of Psychiatry, Leiden University Medical Center, Leiden, The Netherlands</aff><aff id="A61"><label>62</label>Department of Psychiatry, University Medical Center Groningen, Groningen, The Netherlands</aff><aff id="A62"><label>63</label>Department of Psychiatry, VU University Medical Center, Amsterdam, The Netherlands</aff><aff id="A63"><label>64</label>Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku University Hospital, Turku, Finland</aff><aff id="A64"><label>65</label>Department of Clinical Physiology and Nuclear Medicine, Turku University Hospital, Turku, Finland</aff><aff id="A65"><label>66</label>Department of Epidemiology and Biostatistics, School of Public Health, MRC-HPA Centre for Environment and Health, Faculty of Medicine, Imperial College, London, UK</aff><aff id="A66"><label>67</label>Biocenter Oulu, University of Oulu, Oulu, Finland</aff><aff id="A67"><label>68</label>Division of Population Health Sciences and Education, St George’s, University of London, London, UK</aff><aff id="A68"><label>69</label>Munich Cluster for Systems Neurology (SyNergy), Munich, Germany</aff><aff id="A69"><label>Xxx</label>Department of Neurology, Landspitali University Hospital, Reykjavik, Iceland</aff><aff id="A70"><label>Xxx2</label>School of Medicine, University of Iceland, Reykjavik, Iceland</aff><author-notes><corresp id="CR1">Corresponding authors: Aarno Palotie (<email>ap8@sanger.ac.uk</email>), Verneri Anttila (<email>anttila@atgu.mgh.harvard.edu</email>)</corresp><fn id="FN1"><label>55</label><p id="P1">Details appear in the <xref ref-type="supplementary-material" rid="SD1">Supplementary Note</xref>.</p></fn><fn id="FN4"><p id="P2"><bold>Author Contributions</bold></p><p id="P3">Jointly supervised research: A.Palotie, D.C., D.R.N., M.Daly, K.S., G.D.S., M.W., M.Dichgans, A.M., M.Ferrari, C.K., T.K., D.S., L.C., J.A.Z., M.R.J., C.D., D.I.B., J.K., L.Q., G.T.</p><p id="P4">Conceived and designed the study: A.Palotie, D.C., D.R.N., V.Anttila, B.S.W., M.Daly, M.W., M.Dichgans, A.M., C.K., L.C., J.A.Z., M.R.J., C.D., D.I.B., J.K., T.K., M.Kallela, R.M., B.V., G.T., L.Q., M.A.I., L.L., E.H., M.S., H.S., K.S., F.J., T.F., B.M.M.</p><p id="P5">Performed the experiments: V.Anttila, B.S.W., A.Palotie, D.R.N., G.D.S., M.W., M.Dichgans, A.M., C.K., J.A.Z., M.R.J., O.R., C.D., D.I.B., J.K., E.B., M.Kallela, B.V., G.T., E.H., T.F., R.R.F., N.G.M., A.G.U., T.M., J.G.E.</p><p id="P6">Performed statistical analysis: V.Anttila, B.S.W., P.G., D.C., D.R.N., M.Daly, D.S., E.B., J.R.G., S.B.R.J., T.K., F.B., G.M., R.M., B.V., L.Q., M.A.I., L.L., I.D., P.P., M.S., S.Steinberg, T.F., B.M.M.</p><p id="P7">Analysed the data: V.Anttila, B.S.W., P.G., A.Palotie, D.C., D.R.N., L.C., J.R.G., S.B.R.J., K.L., T.K., F.B., G.M., R.M., B.V., S.E.M., L.Q., M.A.I., L.L., J.W., P.P., M.S., S.Steinberg, H.S., T.F., N.A., B.M.M., D.T.</p><p id="P8">Contributed reagents/materials/analysis tools: D.C., D.R.N., M.Daly, A.Palotie, G.D.S., M.W., M.Dichgans, A.M., M.Ferrari, C.K., D.S., L.C., J.A.Z., M.R.J., O.R., C.D., D.I.B., B.W.P., J.K., E.B., J.R.G., K.L., E.R., V.Anttila, B.S.W., P.G., T.K., F.B., G.M., M.Kallela, R.M., B.V., G.T., U.T., W.L.M., L.Q., M.Koiranen, M.A.I., T.L., A.H.S., L.L., I.D., B.N., M.S., L.M.R., J.E.B., P.M.R., S.Steinberg, H.S., F.J., D.A.L., D.M.E., S.M.R., M.Färkkilä, V.Artto, M.A.K., T.F., J.S., R.R.F., N.P., C.M.W., R.Z., A.C.H., P.A.F.M., G.W.M., N.G.M., G.B., H.G., A.Heinze, K.H.K., F.M.K.W., A.L.H., A.Pouta, J.E., A.G.U., A.Hofman, J.J.H., J.M.V., K.H., M.A., B.M.M., S.Schreiber, T.M., E.W., A.A., J.G.E., B.T., D.T.</p><p id="P9">Wrote the paper: V.Anttila, B.S.W., A.Palotie, D.C., D.R.N., A.M., C.K.</p><p id="P10">All authors contributed to the final version of the manuscript.</p></fn></author-notes><pub-date pub-type="nihms-submitted"><day>27</day><month>5</month><year>2014</year></pub-date><pub-date pub-type="epub"><day>23</day><month>6</month><year>2013</year></pub-date><pub-date pub-type="ppub"><month>8</month><year>2013</year></pub-date><pub-date pub-type="pmc-release"><day>02</day><month>6</month><year>2014</year></pub-date><volume>45</volume><issue>8</issue><fpage>912</fpage><lpage>917</lpage><pmc-comment>elocation-id from pubmed: 10.1038/ng.2676</pmc-comment>
<permissions><license><license-p>Users may view, print, copy, download and text and data- mine the content in such documents, for the purposes of academic research, subject always to the full Conditions of use:
<uri xlink:type="simple" xlink:href="http://www.nature.com/authors/editorial_policies/license.html#terms">http://www.nature.com/authors/editorial_policies/license.html#terms</uri></license-p></license></permissions></article-meta></front><body><p id="P11">Migraine is the most common brain disorder, affecting approximately 14% of the adult population, but its molecular mechanisms are poorly understood. We report the results of a meta-analysis across 29 genome-wide association studies, including a total of 23 285 migraine cases and 95 425 population-matched controls. We identified 12 loci associated with migraine susceptibility (<italic>P</italic> < 5 × 10<sup>−8</sup>). Five loci are new (near <italic>AJAP1</italic> on 1p36, near <italic>TSPAN2</italic> on 1p13, within <italic>FHL5</italic> on 6q16, within <italic>c7orf10</italic> on 7p14, and near <italic>MMP16</italic> on 8q21). Three of these loci were identified in disease subgroup analyses. Brain tissue eQTL analysis suggests potential functional candidate genes at four loci: <italic>APOA1BP, TBC1D7, FUT9, STAT6,</italic> and <italic>ATP5B</italic>.</p><p id="P12">Recently, significant progress has been made in the identification of common genetic variants associated with migraine susceptibility through genome-wide association (GWA) studies of clinic-based migraine with aura (MA) patients<sup><xref rid="R1" ref-type="bibr">1</xref></sup>, migraineurs from the general population<sup><xref rid="R2" ref-type="bibr">2</xref>,<xref rid="R3" ref-type="bibr">3</xref></sup>, and clinic-based migraine without aura (MO) patients<sup><xref rid="R4" ref-type="bibr">4</xref></sup>. To further elucidate the genetic susceptibility of migraine, we performed a meta-analysis of 23 285 individuals with migraine from 29 clinic- and population-based studies (<xref ref-type="fig" rid="F1">Fig. 1</xref>, <xref ref-type="supplementary-material" rid="SD1">Supplementary Fig. 1, Supplementary Note</xref>). These include 5 175 cases from five clinic-based patient collections, matched to 13 972 population-based controls (<xref ref-type="supplementary-material" rid="SD1">Supplementary Table 1</xref>), as well as 18 110 cases from 14 population-based studies and 81 453 migraine-free or control individuals from the same studies (<xref ref-type="supplementary-material" rid="SD1">Supplementary Table 2</xref>). Results from GWA studies of the five clinic-based collections<sup><xref rid="R1" ref-type="bibr">1</xref>,<xref rid="R4" ref-type="bibr">4</xref></sup> and four of the population-based collections<sup><xref rid="R2" ref-type="bibr">2</xref>,<xref rid="R3" ref-type="bibr">3</xref></sup> have been previously reported (<xref ref-type="supplementary-material" rid="SD1">Supplementary Fig. 2</xref>).</p><p id="P13">In addition to the primary meta-analysis using all available genotype data, three subgroup analyses were performed in those cohorts where sufficient additional clinical information was available (<xref ref-type="supplementary-material" rid="SD1">Supplementary Table 3</xref>). The first two subgroups consisted of migraine cases fulfilling the International Headache Society diagnostic criteria<sup><xref rid="R5" ref-type="bibr">5</xref></sup> for either MA or MO. The third subgroup included only the clinic-based samples, under the hypothesis that they represent a group of migraineurs more enriched for severe migraines than cases identified from the general population.</p><p id="P14">Results from the primary meta-analysis and the three subgroups identify 142 single nucleotide polymorphisms (SNPs), at a total of 12 loci, to be significantly associated with migraine susceptibility (<xref ref-type="table" rid="T1">Table 1</xref>, <xref ref-type="supplementary-material" rid="SD1">Supplementary Fig. 3,4</xref>). Eight of those loci contain SNPs that lie within a known transcript. In addition, 1 168 SNPs at 134 loci (<xref ref-type="supplementary-material" rid="SD1">Supplementary Table 4</xref>) showed suggestive association to migraine, again combining the primary analysis and the three subgroup analyses. The single most significant <italic>P</italic> value overall was observed for rs11172113 in the primary analysis (<italic>P</italic> value 2.69 × 10<sup>−19</sup>; <xref ref-type="fig" rid="F2">Fig. 2a</xref>, <xref ref-type="table" rid="T1">Table 1</xref>) at the <italic>LRP1</italic> locus on 12q13.</p><p id="P15">Five of the 12 genome-wide significant loci are new (near <italic>AJAP1</italic>, near <italic>TSPAN2, FHL5, c7orf10,</italic> and near <italic>MMP16</italic>), while seven confirm previously reported migraine loci (<italic>PRDM16<sup><xref rid="R2" ref-type="bibr">2</xref></sup>, MEF2D<sup><xref rid="R4" ref-type="bibr">4</xref></sup>, TRPM8<sup><xref rid="R1" ref-type="bibr">1</xref>,<xref rid="R2" ref-type="bibr">2</xref></sup></italic>, near <italic>TGFBR2<sup><xref rid="R4" ref-type="bibr">4</xref></sup>, PHACTR1<sup><xref rid="R4" ref-type="bibr">4</xref></sup>, ASTN2<sup><xref rid="R4" ref-type="bibr">4</xref></sup></italic>, and <italic>LRP1<sup><xref rid="R2" ref-type="bibr">2</xref></sup></italic>). All seven previously reported loci seen in this study remained significant (all p < 6.25 × 10<sup>−3</sup>, correcting for eight previously reported loci) in analyses that exclude samples used in previous reports<sup><xref rid="R1" ref-type="bibr">1</xref>,<xref rid="R2" ref-type="bibr">2</xref>,<xref rid="R4" ref-type="bibr">4</xref></sup> (<xref ref-type="supplementary-material" rid="SD1">Supplementary Fig. 5 and Supplementary Table 5</xref>). Among the newly identified loci, two contain SNPs with significant association that are located within known transcripts. On 6q16, <italic>FHL5</italic> encodes a transcription factor that regulates cAMP responsive elements <italic>CREM</italic> and <italic>CREB</italic><sup><xref rid="R6" ref-type="bibr">6</xref></sup>, which play a role in synaptic plasticity<sup><xref rid="R7" ref-type="bibr">7</xref></sup> and memory formation<sup><xref rid="R8" ref-type="bibr">8</xref></sup>. The locus also overlaps KIAA0776, which encodes a hypothetical protein also known as UFL1. On 7p14, mutations in <italic>c7orf10</italic> have been found in phenotypically mild or even clinically asymptomatic forms of glutaric aciduria type III<sup><xref rid="R9" ref-type="bibr">9</xref></sup>, a rare metabolic abnormality leading to persistent excretion of glutaric acid.</p><p id="P16">The novel loci on 1p36, 1p13, and 8q21 are located outside known transcripts. On 1p36, rs10915437 is located approximately 500 kb telomeric from <italic>AJAP1</italic> and approximately 300kb centromeric from a gene cluster encoding the apoptosis-related proteins DFFB and TP73 as well as centrosomal protein CEP104. <italic>AJAP1</italic> is expressed in brain (<xref ref-type="supplementary-material" rid="SD1">Supplementary Fig. 6</xref>) and has been associated with tumor invasion and regulation of metalloproteinase activity<sup><xref rid="R10" ref-type="bibr">10</xref></sup>. On 1p13, rs12134493 is located 87 kb 5′ of <italic>TSPAN2</italic>, a member of the tetraspanin family, encoding a cell surface protein that mediates signal transduction events involved in the regulation of cell development, activation, growth and motility<sup><xref rid="R11" ref-type="bibr">11</xref></sup>. <italic>TSPAN2</italic> has further been shown to act as a regulator of metalloproteinase activity<sup><xref rid="R11" ref-type="bibr">11</xref></sup>. On 8q21, rs10504861 is located 200 kb telomeric from matrix metalloproteinase <italic>MMP16</italic>. Members of the metalloproteinase family are widely expressed in human tissues and are involved in the breakdown of extracellular matrix in normal physiological processes. Notably, the protein encoded by <italic>MMP16</italic> (MT-MMP2) cleaves LRP1<sup><xref rid="R12" ref-type="bibr">12</xref></sup>, encoded by a previously reported migraine gene<sup><xref rid="R2" ref-type="bibr">2</xref></sup>. In addition, <italic>MMP16</italic> has recently been shown to be involved in basal NgR1 (Nogo-66 receptor) shedding in cortical neurons, thereby increasing axonal and synaptic plasticity<sup><xref rid="R13" ref-type="bibr">13</xref></sup>.</p><p id="P17">Four of the twelve loci (near <italic>AJAP1</italic>, near <italic>TGFBR2, PHACTR1</italic>, near <italic>MMP16</italic>), including two of the novel associations, were identified exclusively in the subgroup analyses (<xref ref-type="table" rid="T1">Table 1</xref> and <xref ref-type="supplementary-material" rid="SD1">Supplementary Table 4</xref>). Two of the loci (rs9349379 in <italic>PHACTR1</italic> on 6p21, and rs10504861 near <italic>MMP16</italic> on 8q21) reached genome-wide significance only in MO, while no SNPs reached genome-wide significance in MA (<xref ref-type="fig" rid="F2">Fig. 2c, d</xref>). The lowest <italic>P</italic> value in MA was with SNP rs7015657 (p = 7.88 × 10<sup>−8</sup>), which is located approximately 582 kb 3′ from <italic>GFRA2</italic>, a member of the glial cell line-derived neurotrophic factor family.</p><p id="P18">A similar subgroup analysis was performed in only those samples that originate from specialized migraine clinics. Two loci with suggestive association in the primary analysis, rs6790925 (near <italic>TGFBR2</italic>) and rs6478241 (<italic>ASTN2</italic>), reached genome-wide significance in the clinic-based subgroup (<xref ref-type="table" rid="T1">Table 1</xref> and <xref ref-type="supplementary-material" rid="SD1">Supplementary Tables 6,7</xref>). All of the 12 genome-wide significant loci associated with migraine had larger estimated effect sizes in the clinic-based subgroup compared to the primary (All) analysis (<xref ref-type="supplementary-material" rid="SD1">Supplementary Table 6, 7</xref>). A two-tailed binomial test shows the chance of observing larger effects at all 12 loci is significantly different from that expected by chance (<italic>P</italic> = 4.88 × 10<sup>−4</sup>). Among all reported loci (<italic>P</italic> <1 × 10<sup>−5</sup>), only the clinic-based group showed a number of associated SNPs with higher effect sizes (OR > 1.2) at low frequency (MAF < 0.05; <xref ref-type="supplementary-material" rid="SD1">Supplementary Fig. 7</xref>). Thus, clinic-based migraine samples may represent a promising subgroup to help prioritise loci in the search for low frequency variants with moderate effects. Overall, among the 146 loci identified, twice as many have causative minor alleles over protective ones (with the ratio increasing towards the lower minor allele frequencies).</p><p id="P19">To explore the biological context for the identified loci we examined the properties of the most proximal genes to the 12 genome-wide significant top SNPs (<xref ref-type="table" rid="T1">Table 1</xref>). In expression data from 55 269 samples profiled using the Affymetrix HG-U133 Plus 2.0 microarray (including 1 990 brain and 384 endothelial samples), 11 of the 12 genes nearest to the identified loci (all except <italic>FHL5</italic>) were at least moderately (>20% of samples of the tissue showing a normalized log<sub>2</sub> expression value greater than 6; see Methods) expressed in disease-relevant brain regions (<xref ref-type="supplementary-material" rid="SD1">Supplementary Fig. 6</xref>). In contrast, only <italic>TGFBR2</italic> and <italic>MEF2D</italic> show moderate or greater expression in the endothelial samples. Possibly reflecting known co-morbidity between migraine and cardiovascular disease<sup><xref rid="R14" ref-type="bibr">14</xref></sup>, two of the 12 most proximal genes (<italic>TGFBR2</italic> and <italic>PHACTR1</italic>) have also been associated with cardiovascular traits: <italic>TGFBR2</italic> mutations have been reported to cause monogenic Marfan’s syndrome<sup><xref rid="R15" ref-type="bibr">15</xref></sup> and to be involved in abdominal aortic aneurysms<sup><xref rid="R16" ref-type="bibr">16</xref></sup>, while <italic>PHACTR1</italic> is associated with early onset myocardial infarction<sup><xref rid="R17" ref-type="bibr">17</xref></sup>. TSPAN2<sup><xref rid="R18" ref-type="bibr">18</xref></sup>, MEF2D<sup><xref rid="R19" ref-type="bibr">19</xref></sup>, TRPM8<sup><xref rid="R20" ref-type="bibr">20</xref></sup>, TGFBR2<sup><xref rid="R21" ref-type="bibr">21</xref></sup>, PHACTR1<sup><xref rid="R22" ref-type="bibr">22</xref></sup>, MMP16<sup><xref rid="R23" ref-type="bibr">23</xref></sup>, ASTN2<sup><xref rid="R24" ref-type="bibr">24</xref></sup>, and LRP1<sup><xref rid="R25" ref-type="bibr">25</xref></sup> have been suggested to have functions in synaptic formation or regulation; <italic>PRDM16</italic> has been linked to oxidative stress response, and <italic>AJAP1</italic> in maintaining tissue borders (<xref ref-type="supplementary-material" rid="SD1">Supplementary Fig. 8</xref>).</p><p id="P20">To identify possible non-proximity based genes underlying these associations, we examined eQTL data among 394 samples of brain tissue from the North American Brain Expression Consortium and the UK Brain Expression Consortium. Among the 12 regions with significant association to migraine, four were found to contain significant eQTLs (see Methods) among the SNP-probe pairs within the tested brain samples (four in the frontal cortex, one in the cerebellum; <xref ref-type="table" rid="T2">Table 2</xref>). On chromosome 1, rs12136718 is an eQTL for <italic>APOA1BP</italic>, rather than the gene closest to it, <italic>MEF2D. APOA1BP</italic> is widely expressed and is potentially linked with cholesterol efflux from cells<sup><xref rid="R26" ref-type="bibr">26</xref></sup>. On chromosome 6, rs35128104 is an eQTL for <italic>FUT9</italic>, encoding α1,3-fucosyl transferase IX. The FUT9 enzyme synthesizes the Lewis X (Le<sup>x</sup>) carbohydrate structure, which has been implicated in neurite outgrowth in several types of brain neuronal cells<sup><xref rid="R27" ref-type="bibr">27</xref>-<xref rid="R29" ref-type="bibr">29</xref></sup>. At the chromosome 12 locus, two different eQTLs in brain tissue were found within the peak, for <italic>STAT6</italic> (rs4559) and for <italic>ATP5B</italic> (rs113953523), the former at a very robust <italic>P</italic>-value (2.16 × 10<sup>−22</sup>). The <italic>STAT</italic> genes are known for transducing activation signals to transcription factors in macrophages<sup><xref rid="R30" ref-type="bibr">30</xref></sup>, and STAT6 phosphorylation has recently been shown to sense oxidative stress in astrocytes resulting in prostaglandin release<sup><xref rid="R31" ref-type="bibr">31</xref></sup>. A second eQTL gene at the same locus, <italic>ATP5B</italic>, is the β subunit of the mitochondrial ATP synthase, but a potential specific role in neuronal or vascular cells is not known. Finally, on chromosome 6, rs9349379 is an eQTL in cerebellar tissue for <italic>TBC1D7</italic>, that potentially down regulates the tuberous sclerosis gene, TSC1, through a positive regulation of the mTOR-signalling pathway<sup><xref rid="R32" ref-type="bibr">32</xref></sup>. In the central nervous system, <italic>TSC1/2</italic> signalling contributes to neural connectivity via its multi-faceted roles<sup><xref rid="R33" ref-type="bibr">33</xref></sup>. Based on the available data it is not possible to decide whether a gene identified by the eQTL analysis or the gene closest to the strongest positional association is the most relevant gene contributing to migraine pathogenesis.</p><p id="P21">In a hypersensitivity site analysis, associated SNPs in the migraine loci were found to occur significantly more often in DNAse I hypersensitivity sites in a number of tissues (<xref ref-type="supplementary-material" rid="SD1">Supplementary Fig. 9</xref>). This suggests that the loci associated with migraine are enriched for actively transcribed regions, supporting a regulatory role for the variants. Both neuronal and vascular tissue types carry an enriched set of sites within the detected loci. In addition, querying the RegulomeDB (see URLs) showed that several of the associated SNPs were found to overlap directly with known transcription factor binding motifs (<xref ref-type="supplementary-material" rid="SD1">Supplementary Table 8</xref>).</p><p id="P22">The number of significantly associated loci is still modest to form a broad understanding of the disease susceptibility, and any proposed functional hypothesis from the identified loci must thus be taken with caution. Some functional hypotheses could be inferred from the results of this study, as the majority of the identified loci harbour genes that can be linked to neuronal function.</p><p id="P23">The eQTL analysis further supported that regulatory effects in brain tissue may underlie several of the association signals. However, the lack of replication across brain regions suggests that our ability to use the eQTL data to pinpoint the functionally most significant gene within the locus is limited.</p><p id="P24">The observed difference between the number of significant loci in the MO and MA groups (6 vs 0, respectively; <xref ref-type="fig" rid="F2">Fig. 2c, d</xref>) despite reasonably similar sample sizes was somewhat unexpected. MA has been shown to have a considerably higher heritability estimate and sibling recurrence risk than MO (3.8 vs 1.9), and has thus been considered to be the more heritable of the two common migraine types<sup><xref rid="R34" ref-type="bibr">34</xref></sup>. One possible explanation could be that the genetic susceptibility to MA is mediated more by rare variants with larger effect sizes, although this remains speculative. Another explanation for the difference may be a higher degree of heterogeneity among the MA cases (due to genetically distinct subgroups, for example). No common variants specifically predisposing to aura were identified by this study (the lowest observed P-value was p = 7.88 × 10<sup>−8</sup>).</p><p id="P25">In summary, we conducted a large migraine meta-analysis and identified 12 loci associated with migraine susceptibility, including five loci not previously associated with migraine, as well as 134 additional suggestive loci. An eQTL analysis of brain tissue highlighted a further five genes potentially implicated in migraine susceptibility. Two of the 12 loci were observed only in the clinic-based sample group, possibly suggesting more specificity to severe migraine headache, and two only in the MO group. Seven previously reported loci for migraine susceptibility were replicated in independent samples in this study. The difference in the number of identified loci and the strength of association suggest that the genetic background of MA is considerably less influenced by common variants than that of MO, contrary to previous expectations. Finally, while pathway analysis of the 146 loci showed no concentration in any particular pathway or tissue, eight of the 12 identified loci are located in or immediately outside genes with known function in synaptic or neuronal regulation and several of them exert regulatory control on one another.</p><sec sec-type="methods" specific-use="web-only" id="S1"><title>Methods</title><sec id="S2"><title>Overall study design</title><p id="P26">For this meta-analysis, we used SNP marker data from 23 285 cases and 95 425 controls of European descent from 29 studies, including five clinic-based studies compared to population-matched control samples with unknown migraine status, as well as 14 entirely population-based cohorts. Four of the population-based cohorts (the B58C, NFBC, Young Finns and FinnTwin; see <xref ref-type="supplementary-material" rid="SD1">Supplementary Note</xref> for further details) were birth cohorts. The datasets for the meta-analysis included previously genotyped genome-wide association (GWA) study data from migraine-specific studies by the International Headache Genetics Consortium (see URLs) studies and the Women’s Genome Health Study, as well as a number of pre-existing population-based GWA studies (for complete list of references, see <xref ref-type="supplementary-material" rid="SD1">Supplementary Table 2</xref>). Local research ethics committees approved the individual studies, and informed consent was obtained from all participants when necessary (see the <xref ref-type="supplementary-material" rid="SD1">Supplementary Materials</xref> for full details of ethics and consent procedures for each study). Additional details on sample recruitment and phenotypes and summary details for each collection are given in the <xref ref-type="supplementary-material" rid="SD1">Supplementary Note</xref> and in <xref ref-type="supplementary-material" rid="SD1">Supplementary Tables 1 and 2</xref>. Genome-wide SNP genotyping was performed independently in each cohort with the use of various standard genotyping technologies, and imputed for each study with reference to HapMap release 21 or 22 CEU phased genotypes<sup><xref rid="R35" ref-type="bibr">35</xref></sup>.</p></sec><sec id="S3"><title>Study phenotypes</title><p id="P27">The primary phenotype analysed was migraine of any type, regardless of source. This was followed by a subgroup analysis consisting of 1) analysing only the clinical samples, 2) only samples satisfying criteria for MA, and 3) for MO. Population-based samples were not analysed genome-wide as a subgroup, due to forming 78% of cases and 85% of controls in the main analysis, but associations were calculated for the significant SNPs for comparative purposes. In the clinical cohorts, headache specialist has assigned a migraine diagnosis based on direct or telephone interview or through the use of an extensive questionnaire. For the population studies, migraine status for individuals in a study sample has been determined by a questionnaire (see <xref ref-type="supplementary-material" rid="SD1">Supplementary Note</xref>).</p></sec><sec id="S4"><title>Statistical analysis of GWA study data</title><p id="P28">Each study contributed summary statistic data from an association analysis performed using a frequentist additive model based on an expected allelic dosage model for SNP markers, adjusting for gender (using either SNPTEST or ProbABEL [see URLs]). SNPs were filtered on per-study level based on inclusion criteria of MAF>0.1% and imputation quality measures of I<sub>A</sub> > 0.6 (IMPUTE 2<sup><xref rid="R36" ref-type="bibr">36</xref></sup>) or r<sup>2</sup> > 0.3 (MACH<sup><xref rid="R37" ref-type="bibr">37</xref></sup>). Four of the included studies contain novel genotyping (HUNT) or imputation (the Finnish, German and Dutch MA studies and HUNT). In the meta-analysis, combined association data for ~2.3 million imputed and genotyped autosomal SNPs were analysed in a fixed-effects model using GWAMA. At this stage, SNPs with a heterogeneity coefficient <italic>I<sup>2</sup></italic> exceeding 75% or presence in only four or fewer studies were filtered out. In the meta-analysis, there was little evidence for population stratification at the study level (each genomic inflation factor λ≤ 1.1), though moderate inflation was observed at the meta-analysis level (λ = 1.15; <xref ref-type="supplementary-material" rid="SD1">Supplementary Fig. 4</xref>). For estimating genome-wide significance, we used the commonly accepted threshold of 5 × 10<sup>−8</sup> for primary loci <sup><xref rid="R38" ref-type="bibr">38</xref></sup>, and 1 × 10<sup>−5</sup> for secondary loci, in accordance with the reporting threshold for the GWAS catalog<sup><xref rid="R39" ref-type="bibr">39</xref></sup>. At secondary loci, to limit spurious associations, at least two SNPs were required to pass the significance threshold within a 50kb window. We also estimated the robustness of this threshold using the false discovery rate method of Benjamini & Hochberg<sup><xref rid="R40" ref-type="bibr">40</xref></sup>, showing that the <italic>P</italic> value threshold corresponding to FDR<0.05 was 2.33 × 10<sup>−5</sup>. The quantile-quantile plot (<xref ref-type="supplementary-material" rid="SD1">Supplementary Fig. 4</xref>) of the meta-analysis <italic>P</italic> values showed a marked excess of association signals well beyond those expected by chance below the suggestive reporting threshold. The significant loci were visualized using the LocusZoom interface<sup><xref rid="R41" ref-type="bibr">41</xref></sup>. For the heterogeneity analyses for migraine type, due to shared controls in some of the sets, the available study samples were divided into as equally-sized groups in terms of effective study size as possible, and then the data was analysed using the gender heterogeneity analysis method <sup><xref rid="R42" ref-type="bibr">42</xref></sup> (–sex option) of GWAMA<sup><xref rid="R43" ref-type="bibr">43</xref></sup>, with a dummy variable coding for MA and MO instead of gender. For the heterogeneity analyses for gender, the same method was used to compare <italic>P</italic> values from males-only and females-only analyses.</p></sec><sec id="S5"><title>Pathway analyses</title><p id="P29"><bold>MAGENTA</bold> - The MAGENTA software<sup><xref rid="R44" ref-type="bibr">44</xref></sup> was used to conduct an analysis to evaluate whether <italic>P</italic> values for association with migraine are enriched to particular biological networks, using pathway lists from GO, PANTHER, INGENUITY, KEGG, REACTOME and BIOCARTA. In the gene set enrichment analysis, <italic>P</italic> values were estimated via 10,000 permutations of genes evaluated at 75<sup>th</sup> FDR percentile (due to assumption of high polygenicity), manually corrected to account for FDR across all pathway sets. <bold>DAPPLE</bold> - Using a refined database of high-confidence protein-protein interactions (InWeb<sup><xref rid="R45" ref-type="bibr">45</xref>,<xref rid="R46" ref-type="bibr">46</xref></sup>) we used DAPPLE <sup><xref rid="R47" ref-type="bibr">47</xref></sup> to assess the amount of physical interactions connecting the genes within 50 kb of the 146 reported migraine loci, as well as an analysis of only the 16 proteins from the 12 genome-wide significant loci. Both direct and indirect (through 1st order common interaction partners) were measured and compared to a random expectation over 10,000 permutations, and the resulting network was plotted. <bold>GRAIL</bold> - The GRAIL web interface <sup><xref rid="R48" ref-type="bibr">48</xref></sup> was used to explore similarities in published PubMed articles (August 2012 freeze), using data from HapMap release 22 CEU and gene size correction set to on. From the GRAIL results, only genes with significant (<italic>P</italic> value<0.05) are shown, and the list of similar genes was capped at genes within the top 200 highest ranks.</p></sec><sec id="S6"><title>Overlap with DNAse I hypersensitivity sites</title><p id="P30">The positions of SNPs from migraine-associated loci were overlapped with DNase 1 ‘hotspot’ regions from the ENCODE project that mark generalized chromatin accessibility mapped for each of 125 diverse cell lines and tissues<sup><xref rid="R49" ref-type="bibr">49</xref></sup>. To assess the significance of overlap for the set of SNPs as a whole, 100 background sets of SNPs were chosen from the genome so that each migraine associated SNP was matched in each set by a SNP within the same decile for minor allele frequency, distance to the nearest transcription start site and GC content of the 100 base region surrounding the SNP. The background SNP sets were overlapped with the DNase 1 hotspots, and the enrichment for overlap with the migraine associated SNP set expressed as the Z score relative to the distribution of background SNP set overlaps on a per cell line basis. In addition migraine associated SNPs were analysed for other overlap with ENCODE data including transcription factor motifs using RegulomeDB (<ext-link ext-link-type="uri" xlink:href="http://regulome.stanford.edu">http://regulome.stanford.edu</ext-link>)<sup><xref rid="R50" ref-type="bibr">50</xref></sup>.</p></sec><sec id="S7"><title>Tissue-based gene expression analysis</title><p id="P31">For the tissue analysis, a microarray-based analysis of gene expression was performed on a dataset of 55 269 samples in the Gene Expression Omnibus (GEO) database that were measured on the Affymetrix U133 Plus 2.0 Array. Each sample in the raw expression data was first linearly transformed using a modified invariant set normalization method<sup><xref rid="R51" ref-type="bibr">51</xref></sup> on a set of eighty control genes with stable expression on U133 Plus 2.0. The expression data was log2 transformed to stabilize the variance and expression distribution. Finally, the data were quantile-normalized<sup><xref rid="R52" ref-type="bibr">52</xref></sup> to match the expression distribution of each sample. Expression values for genes with multiple probe sets were calculated by taking the median value of all probe sets for that gene. Following normalization, a log2 expression value of 4 is considered baseline and log2 expression values greater than 6 are considered expressed. Sample annotations were curated based on GEO descriptions provided by depositors. To account for variation in the number of samples representing each tissue in the dataset, expression of a gene was plotted to show the fraction of samples of a tissue that exceeds a log2 expression value of 6, with higher fractions indicating more ubiquitous expression in the tissue in question.</p></sec><sec id="S8"><title>eQTL analysis</title><p id="P32">Based on the meta-analysis results for association with migraine, 146 regions of interest were queried against the expression quantitative trait loci (eQTL) results from the North American Brain Expression and UK Brain Expression Consortium studies (GEO # GSE36192, dbGaP # phs000249). These eQTL results are based on Cerebellum and Frontal Cortex tissue mRNA expression levels from 394 human subjects (see <xref ref-type="supplementary-material" rid="SD1">Supplementary methods</xref>). Of the 146 associated regions 134 were represented in the eQTL analysis. Within these regions 831 mRNA expression traits and 222 668 <italic>cis</italic> SNP/expression trait pairs were considered in Cerebellum and 864 mRNA expression traits and 230 660 <italic>cis</italic> SNP/expression trait pairs were considered in within the Frontal Cortex. 45 SNPs within the migraine associated loci were found to have significant correlation (evaluated at FDR-corrected threshold; 0.0001668 for frontal cortex and 0.0002187 for cerebellum) with the expression of 12 mRNA transcripts in both the Cerebellum and the Frontal Cortex. A more detailed methods description can be found in <xref ref-type="supplementary-material" rid="SD1">Supplementary Materials</xref>. The extent of linkage disequilibrium between SNPs associated with migraine and the SNPs in the tested eQTL SNP-probe pairs was evaluated using SNAP (see <xref ref-type="fn" rid="FN3">URLs</xref>).</p></sec></sec><sec sec-type="supplementary-material" id="SM"><title>Supplementary Material</title><supplementary-material content-type="local-data" id="SD1"><label>1</label><media xlink:href="NIHMS53496-supplement-1.pdf" orientation="portrait" xlink:type="simple" id="d36e2012" position="anchor"></media></supplementary-material></sec></body><back><ack id="S9"><title>Acknowledgments</title><p>Study-specific acknowledgments appear as a <xref ref-type="supplementary-material" rid="SD1">Supplementary Note</xref>. We wish to thank Alison Coffey, S. Hunt, R. Gwillian, P. Whittaker, S. Potter and A. Tashakkori-Ghanbarian, for their invaluable help with this study, and to collectively thank everyone who has contributed to the collection, genotyping and analysis of the individual cohorts, as well as all the study participants.</p></ack><fn-group><fn id="FN2"><p id="P33"><bold>Competing Financial Interests</bold></p><p id="P34">The authors declare no competing financial interests</p></fn><fn id="FN3"><p id="P35"><bold>URLs.</bold></p><p id="P36">DAPPLE - <ext-link ext-link-type="uri" xlink:href="http://www.broadinstitute.org/mpg/dapple">www.broadinstitute.org/mpg/dapple</ext-link></p><p id="P37">GRAIL - <ext-link ext-link-type="uri" xlink:href="http://www.broadinstitute.org/mpg/grail">www.broadinstitute.org/mpg/grail</ext-link></p><p id="P38">GWAMA - <ext-link ext-link-type="uri" xlink:href="http://www.well.ox.ac.uk/gwama">www.well.ox.ac.uk/gwama</ext-link></p><p id="P39">IMPUTE2 - <ext-link ext-link-type="uri" xlink:href="http://mathgen.stats.ox.ac.uk/impute">mathgen.stats.ox.ac.uk/impute</ext-link></p><p id="P40">International Headache Genetics Consortium- <ext-link ext-link-type="uri" xlink:href="http://www.headachegenetics.org">www.headachegenetics.org</ext-link></p><p id="P41">MAGENTA - <ext-link ext-link-type="uri" xlink:href="http://www.broadinstitute.org/mpg/magenta">www.broadinstitute.org/mpg/magenta</ext-link></p><p id="P42">ProbAbel - <ext-link ext-link-type="uri" xlink:href="http://www.genabel.org/packages/ProbABEL">http://www.genabel.org/packages/ProbABEL</ext-link></p><p id="P43">RegulomeDB - <ext-link ext-link-type="uri" xlink:href="http://regulome.stanford.edu">http://regulome.stanford.edu</ext-link></p><p id="P44">SNAP - <ext-link ext-link-type="uri" xlink:href="http://www.broadinstitute.org/mpg/snap">http://www.broadinstitute.org/mpg/snap</ext-link></p><p id="P45">SNPTEST - <ext-link ext-link-type="uri" 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Participation in each analysis depended on the availability of the data in question and the recruitment method.</p></caption><graphic xlink:href="emss-53496-f0001"></graphic></fig><fig id="F2" orientation="portrait" position="float"><label>Figure 2</label><caption><title>Manhattan plot of the results of the meta-analysis</title><p>Results of the meta-analysis of all migraine cases, of any migraine subtype or recruiting method, versus all available controls, adjusting for gender. Red dots indicate SNPs with significant (< 5 × 10<sup>−8</sup>) <italic>P</italic> values.</p><p>A) All migraine (23 285 cases, 95 425 controls)</p><p>B) Clinic-based studies only (5 175 cases, 13 972 controls)</p><p>C) Migraine with aura (5 118 cases, 74 239 controls)</p><p>D) Migraine without aura (7 107 cases, 69 427 controls)</p></caption><graphic xlink:href="emss-53496-f0002"></graphic><graphic xlink:href="emss-53496-f0003"></graphic><graphic xlink:href="emss-53496-f0004"></graphic><graphic xlink:href="emss-53496-f0005"></graphic></fig><table-wrap id="T1" position="float" orientation="landscape"><label>Table 1</label><caption><title>Results of the association analyses</title></caption><table frame="box" rules="all"><thead><tr><th align="center" valign="middle" rowspan="1" colspan="1">SNP</th><th align="center" valign="middle" rowspan="1" colspan="1">Chr</th><th align="center" valign="middle" rowspan="1" colspan="1">Position</th><th align="center" valign="middle" rowspan="1" colspan="1">Location</th><th align="center" valign="middle" rowspan="1" colspan="1">Gene</th><th align="center" valign="middle" rowspan="1" colspan="1">Minor allele</th><th align="center" valign="middle" rowspan="1" colspan="1">MAF</th><th align="center" valign="middle" rowspan="1" colspan="1">P-value</th><th align="center" valign="middle" rowspan="1" colspan="1">OR (95% CI)</th><th align="center" valign="middle" rowspan="1" colspan="1">q/p-value</th><th align="center" valign="middle" rowspan="1" colspan="1"><italic>I</italic><sup>2</sup></th><th align="center" valign="middle" rowspan="1" colspan="1">Group or subgroup with the lowest p-value</th><th align="center" valign="middle" rowspan="1" colspan="1">Additional significance in</th></tr></thead><tbody><tr><td align="center" valign="middle" rowspan="1" colspan="1">rs2651899</td><td align="center" valign="middle" rowspan="1" colspan="1">1</td><td align="center" valign="middle" rowspan="1" colspan="1">3,073,572</td><td align="center" valign="middle" rowspan="1" colspan="1">Genic</td><td align="center" valign="middle" rowspan="1" colspan="1"><italic>PRDM16</italic><xref ref-type="table-fn" rid="TFN1">*</xref></td><td align="center" valign="middle" rowspan="1" colspan="1">C</td><td align="center" valign="middle" rowspan="1" colspan="1">0.41</td><td align="center" valign="middle" rowspan="1" colspan="1">3.28 × 10<sup>−14</sup></td><td align="center" valign="middle" rowspan="1" colspan="1">1.09 (1.07-1.12)</td><td align="center" valign="middle" rowspan="1" colspan="1">0.214</td><td align="center" valign="middle" rowspan="1" colspan="1">20%</td><td align="center" valign="middle" rowspan="1" colspan="1">All samples</td><td align="center" valign="middle" rowspan="1" colspan="1"></td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">rs10915437</td><td align="center" valign="middle" rowspan="1" colspan="1">1</td><td align="center" valign="middle" rowspan="1" colspan="1">4,082,866</td><td align="center" valign="middle" rowspan="1" colspan="1">Intergenic</td><td align="center" valign="middle" rowspan="1" colspan="1"><italic>near AJAP1</italic></td><td align="center" valign="middle" rowspan="1" colspan="1">G</td><td align="center" valign="middle" rowspan="1" colspan="1">0.36</td><td align="center" valign="middle" rowspan="1" colspan="1">2.81 × 10<sup>−8</sup></td><td align="center" valign="middle" rowspan="1" colspan="1">0.86 (0.82-0.91)</td><td align="center" valign="middle" rowspan="1" colspan="1">0.108</td><td align="center" valign="middle" rowspan="1" colspan="1">47%</td><td align="center" valign="middle" rowspan="1" colspan="1">Clinics only</td><td align="center" valign="middle" rowspan="1" colspan="1"></td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">rs12134493</td><td align="center" valign="middle" rowspan="1" colspan="1">1</td><td align="center" valign="middle" rowspan="1" colspan="1">115,479,469</td><td align="center" valign="middle" rowspan="1" colspan="1">Intergenic</td><td align="center" valign="middle" rowspan="1" colspan="1"><italic>near TSPAN2</italic></td><td align="center" valign="middle" rowspan="1" colspan="1">A</td><td align="center" valign="middle" rowspan="1" colspan="1">0.46</td><td align="center" valign="middle" rowspan="1" colspan="1">6.71 × 10<sup>−14</sup></td><td align="center" valign="middle" rowspan="1" colspan="1">1.14 (1.10-1.18)</td><td align="center" valign="middle" rowspan="1" colspan="1">0.480</td><td align="center" valign="middle" rowspan="1" colspan="1">14%</td><td align="center" valign="middle" rowspan="1" colspan="1">All samples</td><td align="center" valign="middle" rowspan="1" colspan="1"></td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">rs2274316</td><td align="center" valign="middle" rowspan="1" colspan="1">1</td><td align="center" valign="middle" rowspan="1" colspan="1">154,712,866</td><td align="center" valign="middle" rowspan="1" colspan="1">Genic</td><td align="center" valign="middle" rowspan="1" colspan="1"><italic>MEF2D</italic><xref ref-type="table-fn" rid="TFN1">*</xref></td><td align="center" valign="middle" rowspan="1" colspan="1">C</td><td align="center" valign="middle" rowspan="1" colspan="1">0.37</td><td align="center" valign="middle" rowspan="1" colspan="1">3.14 × 10<sup>−8</sup></td><td align="center" valign="middle" rowspan="1" colspan="1">1.07 (1.04-1.09)</td><td align="center" valign="middle" rowspan="1" colspan="1">0.021</td><td align="center" valign="middle" rowspan="1" colspan="1">45%</td><td align="center" valign="middle" rowspan="1" colspan="1">All samples</td><td align="center" valign="middle" rowspan="1" colspan="1"></td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">rs7577262</td><td align="center" valign="middle" rowspan="1" colspan="1">2</td><td align="center" valign="middle" rowspan="1" colspan="1">234,483,608</td><td align="center" valign="middle" rowspan="1" colspan="1">Genic</td><td align="center" valign="middle" rowspan="1" colspan="1"><italic>TRPM8</italic><xref ref-type="table-fn" rid="TFN1">*</xref></td><td align="center" valign="middle" rowspan="1" colspan="1">A</td><td align="center" valign="middle" rowspan="1" colspan="1">0.10</td><td align="center" valign="middle" rowspan="1" colspan="1">3.27 × 10<sup>−13</sup></td><td align="center" valign="middle" rowspan="1" colspan="1">0.87 (0.84-0.90)</td><td align="center" valign="middle" rowspan="1" colspan="1">0.070</td><td align="center" valign="middle" rowspan="1" colspan="1">33%</td><td align="center" valign="middle" rowspan="1" colspan="1">All samples</td><td align="center" valign="middle" rowspan="1" colspan="1">Clinics only, MO only</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">rs6790925</td><td align="center" valign="middle" rowspan="1" colspan="1">3</td><td align="center" valign="middle" rowspan="1" colspan="1">30,455,089</td><td align="center" valign="middle" rowspan="1" colspan="1">Intergenic</td><td align="center" valign="middle" rowspan="1" colspan="1"><italic>near TGFBR2</italic><xref ref-type="table-fn" rid="TFN1">*</xref></td><td align="center" valign="middle" rowspan="1" colspan="1">T</td><td align="center" valign="middle" rowspan="1" colspan="1">0.38</td><td align="center" valign="middle" rowspan="1" colspan="1">2.16 × 10<sup>−8</sup></td><td align="center" valign="middle" rowspan="1" colspan="1">1.15 (1.10-1.21)</td><td align="center" valign="middle" rowspan="1" colspan="1">0.780</td><td align="center" valign="middle" rowspan="1" colspan="1">0%</td><td align="center" valign="middle" rowspan="1" colspan="1">Clinics only</td><td align="center" valign="middle" rowspan="1" colspan="1"></td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">rs9349379</td><td align="center" valign="middle" rowspan="1" colspan="1">6</td><td align="center" valign="middle" rowspan="1" colspan="1">13,011,943</td><td align="center" valign="middle" rowspan="1" colspan="1">Genic</td><td align="center" valign="middle" rowspan="1" colspan="1"><italic>PHACTR1</italic><xref ref-type="table-fn" rid="TFN1">*</xref></td><td align="center" valign="middle" rowspan="1" colspan="1">G</td><td align="center" valign="middle" rowspan="1" colspan="1">0.40</td><td align="center" valign="middle" rowspan="1" colspan="1">2.81 × 10<sup>−10</sup></td><td align="center" valign="middle" rowspan="1" colspan="1">0.86 (0.82-0.90)</td><td align="center" valign="middle" rowspan="1" colspan="1">0.443</td><td align="center" valign="middle" rowspan="1" colspan="1">0%</td><td align="center" valign="middle" rowspan="1" colspan="1">MO only</td><td align="center" valign="middle" rowspan="1" colspan="1">All samples</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">rs13208321</td><td align="center" valign="middle" rowspan="1" colspan="1">6</td><td align="center" valign="middle" rowspan="1" colspan="1">96,967,075</td><td align="center" valign="middle" rowspan="1" colspan="1">Genic</td><td align="center" valign="middle" rowspan="1" colspan="1"><italic>FHL5</italic></td><td align="center" valign="middle" rowspan="1" colspan="1">A</td><td align="center" valign="middle" rowspan="1" colspan="1">0.22</td><td align="center" valign="middle" rowspan="1" colspan="1">2.15 × 10<sup>−12</sup></td><td align="center" valign="middle" rowspan="1" colspan="1">1.18 (1.13-1.24)</td><td align="center" valign="middle" rowspan="1" colspan="1">0.168</td><td align="center" valign="middle" rowspan="1" colspan="1">0%</td><td align="center" valign="middle" rowspan="1" colspan="1">All samples</td><td align="center" valign="middle" rowspan="1" colspan="1">MO only</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">rs4379368</td><td align="center" valign="middle" rowspan="1" colspan="1">7</td><td align="center" valign="middle" rowspan="1" colspan="1">40,432,725</td><td align="center" valign="middle" rowspan="1" colspan="1">Genic</td><td align="center" valign="middle" rowspan="1" colspan="1"><italic>c7orf10</italic></td><td align="center" valign="middle" rowspan="1" colspan="1">T</td><td align="center" valign="middle" rowspan="1" colspan="1">0.12</td><td align="center" valign="middle" rowspan="1" colspan="1">1.46 × 10<sup>−9</sup></td><td align="center" valign="middle" rowspan="1" colspan="1">1.11 (1.08-1.15)</td><td align="center" valign="middle" rowspan="1" colspan="1">0.441</td><td align="center" valign="middle" rowspan="1" colspan="1">2%</td><td align="center" valign="middle" rowspan="1" colspan="1">All samples</td><td align="center" valign="middle" rowspan="1" colspan="1">MO only</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">rs10504861</td><td align="center" valign="middle" rowspan="1" colspan="1">8</td><td align="center" valign="middle" rowspan="1" colspan="1">89,617,048</td><td align="center" valign="middle" rowspan="1" colspan="1">Intergenic</td><td align="center" valign="middle" rowspan="1" colspan="1"><italic>near MMP16</italic></td><td align="center" valign="middle" rowspan="1" colspan="1">T</td><td align="center" valign="middle" rowspan="1" colspan="1">0.16</td><td align="center" valign="middle" rowspan="1" colspan="1">1.32 × 10<sup>−8</sup></td><td align="center" valign="middle" rowspan="1" colspan="1">0.86 (0.81-0.90)</td><td align="center" valign="middle" rowspan="1" colspan="1">0.755</td><td align="center" valign="middle" rowspan="1" colspan="1">0%</td><td align="center" valign="middle" rowspan="1" colspan="1">MO only</td><td align="center" valign="middle" rowspan="1" colspan="1"></td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">rs6478241</td><td align="center" valign="middle" rowspan="1" colspan="1">9</td><td align="center" valign="middle" rowspan="1" colspan="1">118,292,450</td><td align="center" valign="middle" rowspan="1" colspan="1">Genic</td><td align="center" valign="middle" rowspan="1" colspan="1"><italic>ASTN2</italic><xref ref-type="table-fn" rid="TFN1">*</xref></td><td align="center" valign="middle" rowspan="1" colspan="1">A</td><td align="center" valign="middle" rowspan="1" colspan="1">0.38</td><td align="center" valign="middle" rowspan="1" colspan="1">1.04 × 10<sup>−9</sup></td><td align="center" valign="middle" rowspan="1" colspan="1">1.16 (1.11-1.22)</td><td align="center" valign="middle" rowspan="1" colspan="1">0.646</td><td align="center" valign="middle" rowspan="1" colspan="1">0%</td><td align="center" valign="middle" rowspan="1" colspan="1">Clinics only</td><td align="center" valign="middle" rowspan="1" colspan="1">All samples</td></tr><tr><td align="center" valign="middle" rowspan="1" colspan="1">rs11172113</td><td align="center" valign="middle" rowspan="1" colspan="1">12</td><td align="center" valign="middle" rowspan="1" colspan="1">55,813,550</td><td align="center" valign="middle" rowspan="1" colspan="1">Genic</td><td align="center" valign="middle" rowspan="1" colspan="1"><italic>LRP1</italic><xref ref-type="table-fn" rid="TFN1">*</xref></td><td align="center" valign="middle" rowspan="1" colspan="1">C</td><td align="center" valign="middle" rowspan="1" colspan="1">0.43</td><td align="center" valign="middle" rowspan="1" colspan="1">2.69 × 10<sup>−19</sup></td><td align="center" valign="middle" rowspan="1" colspan="1">0.90 (0.88-0.92)</td><td align="center" valign="middle" rowspan="1" colspan="1">0.188</td><td align="center" valign="middle" rowspan="1" colspan="1">22%</td><td align="center" valign="middle" rowspan="1" colspan="1">All samples</td><td align="center" valign="middle" rowspan="1" colspan="1">MO only</td></tr></tbody></table><table-wrap-foot><p>When multiple subgroups show significant association, <italic>P</italic> values and odds ratios (OR) are shown for the analysis with the lowest <italic>P</italic> value. ORs reported for the minor allele. Chromosomal positions are based on NCBI build 36. In the Location and Gene columns, SNPs located within the gene transcript (‘Genic’) list that gene, while for intergenic SNPs (‘Intergenic’) the gene reported for that locus is listed.</p><fn id="TFN1"><label>*</label><p id="P46">Previously identified migraine locus. MAF – minor allele frequency. q/p value - <italic>P</italic> value from Cochran’s heterogeneity statistic. <italic>I</italic><sup>2</sup> – heterogeneity index. MO – migraine without aura.</p></fn></table-wrap-foot></table-wrap><table-wrap id="T2" position="float" orientation="landscape"><label>Table 2</label><caption><title>Significant eQTLs observed in 394 samples of two different brain tissue at loci significantly associated with migraine</title></caption><table frame="void" rules="groups"><thead><tr><th align="left" valign="bottom" rowspan="1" colspan="1"></th><th align="left" valign="bottom" rowspan="1" colspan="1"></th><th align="left" valign="bottom" rowspan="1" colspan="1"></th><th align="left" valign="bottom" rowspan="1" colspan="1"></th><th align="left" valign="bottom" rowspan="1" colspan="1"></th><th colspan="4" align="center" valign="bottom" rowspan="1">Frontal cortex</th><th align="left" valign="bottom" rowspan="1" colspan="1"></th></tr><tr><th align="left" valign="bottom" rowspan="1" colspan="1">Chr</th><th align="left" valign="bottom" rowspan="1" colspan="1">Position (bp)</th><th align="left" valign="bottom" rowspan="1" colspan="1">Locus</th><th align="left" valign="bottom" rowspan="1" colspan="1">Probe</th><th align="left" valign="bottom" rowspan="1" colspan="1">Gene name</th><th align="left" valign="bottom" rowspan="1" colspan="1">SNPs with sign.<break></break>eQTL with<break></break>probe</th><th align="left" valign="bottom" rowspan="1" colspan="1">SNP with<break></break>lowest eQTL<break></break>p-value</th><th align="left" valign="bottom" rowspan="1" colspan="1">Lowest eQTL<break></break>p-value</th><th align="left" valign="bottom" rowspan="1" colspan="1">Highest LD<break></break>with reported<break></break>SNP</th><th align="left" valign="bottom" rowspan="1" colspan="1">SNPs with sign.<break></break>eQTL with<break></break>probe</th></tr></thead><tbody><tr><td align="left" valign="middle" rowspan="1" colspan="1">1</td><td align="left" valign="middle" rowspan="1" colspan="1">154,830,352</td><td align="left" valign="middle" rowspan="1" colspan="1"><italic>MEF2D</italic></td><td align="left" valign="middle" rowspan="1" colspan="1">ILMN_1729533</td><td align="left" valign="middle" rowspan="1" colspan="1"><italic>APOA1BP</italic></td><td align="right" valign="middle" rowspan="1" colspan="1"><bold>1</bold></td><td align="left" valign="middle" rowspan="1" colspan="1"><bold>rs12136718</bold></td><td align="left" valign="middle" rowspan="1" colspan="1"><bold>2.18 × 10<sup>−5</sup></bold></td><td align="right" valign="middle" rowspan="1" colspan="1"><bold>0.38</bold></td><td align="right" valign="middle" rowspan="1" colspan="1">0</td></tr><tr><td align="left" valign="middle" rowspan="1" colspan="1">6</td><td align="left" valign="middle" rowspan="1" colspan="1">13,413,333</td><td align="left" valign="middle" rowspan="1" colspan="1"><italic>PHACTR1</italic></td><td align="left" valign="middle" rowspan="1" colspan="1">ILMN_1661622</td><td align="left" valign="middle" rowspan="1" colspan="1"><italic>TBC1D7</italic></td><td align="right" valign="middle" rowspan="1" colspan="1">0</td><td align="center" valign="middle" rowspan="1" colspan="1">-</td><td align="center" valign="middle" rowspan="1" colspan="1">-</td><td align="left" valign="middle" rowspan="1" colspan="1">-</td><td align="right" valign="middle" rowspan="1" colspan="1"><bold>1</bold></td></tr><tr><td align="left" valign="middle" rowspan="1" colspan="1">6</td><td align="left" valign="middle" rowspan="1" colspan="1">96,769,539</td><td align="left" valign="middle" rowspan="1" colspan="1"><italic>FHL5</italic></td><td align="left" valign="middle" rowspan="1" colspan="1">ILMN_1878007</td><td align="left" valign="middle" rowspan="1" colspan="1"><italic>FUT9</italic></td><td align="right" valign="middle" rowspan="1" colspan="1"><bold>2</bold></td><td align="left" valign="middle" rowspan="1" colspan="1"><bold>rs35128104</bold></td><td align="left" valign="middle" rowspan="1" colspan="1"><bold>5.96 × 10<sup>−5</sup></bold></td><td align="right" valign="middle" rowspan="1" colspan="1"><bold>0.74</bold></td><td align="right" valign="middle" rowspan="1" colspan="1">0</td></tr><tr><td align="left" valign="middle" rowspan="1" colspan="1">12</td><td align="left" valign="middle" rowspan="1" colspan="1">55,775,568</td><td align="left" valign="middle" rowspan="1" colspan="1"><italic>LRP1</italic></td><td align="left" valign="middle" rowspan="1" colspan="1">ILMN_1763198</td><td align="left" valign="middle" rowspan="1" colspan="1"><italic>STAT6</italic></td><td align="right" valign="middle" rowspan="1" colspan="1"><bold>40</bold></td><td align="left" valign="middle" rowspan="1" colspan="1"><bold>rs4559</bold></td><td align="left" valign="middle" rowspan="1" colspan="1"><bold>2.16 × 10<sup>−22</sup></bold></td><td align="right" valign="middle" rowspan="1" colspan="1"><bold>1</bold></td><td align="left" valign="middle" rowspan="1" colspan="1">-</td></tr><tr><td align="left" valign="middle" rowspan="1" colspan="1">12</td><td align="left" valign="middle" rowspan="1" colspan="1">55,318,337</td><td align="left" valign="middle" rowspan="1" colspan="1"><italic>LRP1</italic></td><td align="left" valign="middle" rowspan="1" colspan="1">ILMN_1772132</td><td align="left" valign="middle" rowspan="1" colspan="1"><italic>ATP5B</italic></td><td align="right" valign="middle" rowspan="1" colspan="1"><bold>1</bold></td><td align="left" valign="middle" rowspan="1" colspan="1"><bold>rs113953523</bold></td><td align="left" valign="middle" rowspan="1" colspan="1"><bold>1.62 × 10<sup>−4</sup></bold></td><td align="right" valign="middle" rowspan="1" colspan="1"><bold>0.39</bold></td><td align="right" valign="middle" rowspan="1" colspan="1">0</td></tr></tbody></table><table-wrap-foot><p>-Chr – chromosome. All basepair positions are given for build 36/hg18. ‘Highest LD…’ column indicates the maximum extent of LD observed between a significant eQTL SNP-probe pair (see Methods), and a SNP passing the reporting threshold (p< × 10<sup>−5</sup>) for association with migraine. Due to multiple overlapping signals, the HLA region has been excluded from the analysis.</p></table-wrap-foot></table-wrap></floats-group></pmc></record>Pour manipuler ce document sous Unix (Dilib)
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