Meta-analysis identifies 29 additional ulcerative colitis risk loci, increasing the number of confirmed associations to 47
Identifieur interne : 002E05 ( Pmc/Corpus ); précédent : 002E04; suivant : 002E06Meta-analysis identifies 29 additional ulcerative colitis risk loci, increasing the number of confirmed associations to 47
Auteurs : Carl A. Anderson ; Gabrielle Boucher ; Charlie W. Lees ; Andre Franke ; Mauro D Mato ; Kent D. Taylor ; James C. Lee ; Philippe Goyette ; Marcin Imielinski ; Anna Latiano ; Caroline Lagacé ; Regan Scott ; Leila Amininejad ; Suzannah Bumpstead ; Leonard Baidoo ; Robert N. Baldassano ; Murray Barclay ; Theodore M. Bayless ; Stephan Brand ; Carsten Büning ; Jean-Frédéric Colombel ; Lee A. Denson ; Martine De Vos ; Marla Dubinsky ; Cathryn Edwards ; David Ellinghaus ; Rudolf S. N. Fehrmann ; James A. B. Floyd ; Tim Florin ; Denis Franchimont ; Lude Franke ; Michel Georges ; Jürgen Glas ; Nicole L. Glazer ; Stephen L. Guthery ; Talin Haritunians ; Nicholas K. Hayward ; Jean-Pierre Hugot ; Gilles Jobin ; Debby Laukens ; Ian Lawrance ; Marc Lémann ; Arie Levine ; Cecile Libioulle ; Edouard Louis ; Dermot P. Mcgovern ; Monica Milla ; Grant W. Montgomery ; Katherine I. Morley ; Craig Mowat ; Aylwin Ng ; William Newman ; Roel A. Ophoff ; Laura Papi ; Orazio Palmieri ; Laurent Peyrin-Biroulet ; Julián Panés ; Anne Phillips ; Natalie J. Prescott ; Deborah D. Proctor ; Rebecca Roberts ; Richard Russell ; Paul Rutgeerts ; Jeremy Sanderson ; Miquel Sans ; Philip Schumm ; Frank Seibold ; Yashoda Sharma ; Lisa Simms ; Mark Seielstad ; A. Hillary Steinhart ; Stephan R. Targan ; Leonard H. Van Den Berg ; Morten Vatn ; Hein Verspaget ; Thomas Walters ; Cisca Wijmenga ; David C. Wilson ; Harm-Jan Westra ; Ramnik J. Xavier ; Zhen Z. Zhao ; Cyriel Y. Ponsioen ; Vibeke Andersen ; Leif Torkvist ; Maria Gazouli ; Nicholas P. Anagnou ; Tom H. Karlsen ; Limas Kupcinskas ; Jurgita Sventoraityte ; John C. Mansfield ; Subra Kugathasan ; Mark S. Silverberg ; Jonas Halfvarson ; Jerome I. Rotter ; Christopher G. Mathew ; Anne M. Griffiths ; Richard Gearry ; Tariq Ahmad ; Steven R. Brant ; Mathias Chamaillard ; Jack Satsangi ; Judy H. Cho ; Stefan Schreiber ; Mark J. Daly ; Jeffrey C. Barrett ; Miles Parkes ; Vito Annese ; Hakon Hakonarson ; Graham Radford-Smith ; Richard H. Duerr ; Séverine Vermeire ; Rinse K. Weersma ; John D. RiouxSource :
- Nature genetics [ 1061-4036 ] ; 2011.
Abstract
Genome-wide association studies (GWAS) and candidate gene studies in ulcerative colitis (UC) have identified 18 susceptibility loci. We conducted a meta-analysis of 6 UC GWAS, comprising 6,687 cases and 19,718 controls, and followed-up the top association signals in 9,628 cases and 12,917 controls. We identified 29 additional risk loci (P<5×10-8), increasing the number of UC associated loci to 47. After annotating associated regions using GRAIL, eQTL data and correlations with non-synonymous SNPs, we identified many candidate genes providing potentially important insights into disease pathogenesis, including
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DOI: 10.1038/ng.764
PubMed: 21297633
PubMed Central: 3084597
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<record><TEI><teiHeader><fileDesc><titleStmt><title xml:lang="en">Meta-analysis identifies 29 additional ulcerative colitis risk loci, increasing the number of confirmed associations to 47</title><author><name sortKey="Anderson, Carl A" sort="Anderson, Carl A" uniqKey="Anderson C" first="Carl A." last="Anderson">Carl A. Anderson</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Boucher, Gabrielle" sort="Boucher, Gabrielle" uniqKey="Boucher G" first="Gabrielle" last="Boucher">Gabrielle Boucher</name><affiliation><nlm:aff id="A2">Université de Montréal, Medicine, Montréal, Québec, Canada</nlm:aff></affiliation><affiliation><nlm:aff id="A3">Montreal Heart Institute, Research Center, Montréal, Québec, Canada</nlm:aff></affiliation></author><author><name sortKey="Lees, Charlie W" sort="Lees, Charlie W" uniqKey="Lees C" first="Charlie W." last="Lees">Charlie W. Lees</name><affiliation><nlm:aff id="A4">University of Edinburgh, Western General Hospital, Gastrointestinal Unit, Molecular Medicine Centre, Edinburgh, UK</nlm:aff></affiliation></author><author><name sortKey="Franke, Andre" sort="Franke, Andre" uniqKey="Franke A" first="Andre" last="Franke">Andre Franke</name><affiliation><nlm:aff id="A5">Christian-Albrechts-University Kiel, Institute of Clinical Molecular Biology, Kiel, Germany</nlm:aff></affiliation></author><author><name sortKey="D Mato, Mauro" sort="D Mato, Mauro" uniqKey="D Mato M" first="Mauro" last="D Mato">Mauro D Mato</name><affiliation><nlm:aff id="A6">Karolinska Institute, Department of Biosciences and Nutrition, Stockholm, Sweden</nlm:aff></affiliation></author><author><name sortKey="Taylor, Kent D" sort="Taylor, Kent D" uniqKey="Taylor K" first="Kent D." last="Taylor">Kent D. Taylor</name><affiliation><nlm:aff id="A7">Cedars-Sinai Medical Center, Inflammatory Bowel and Immunobiology Research Institute, Los Angeles, California, USA</nlm:aff></affiliation></author><author><name sortKey="Lee, James C" sort="Lee, James C" uniqKey="Lee J" first="James C." last="Lee">James C. Lee</name><affiliation><nlm:aff id="A8">Addenbrooke’s Hospital, University of Cambridge, Gastroenterology Research Unit, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Goyette, Philippe" sort="Goyette, Philippe" uniqKey="Goyette P" first="Philippe" last="Goyette">Philippe Goyette</name><affiliation><nlm:aff id="A2">Université de Montréal, Medicine, Montréal, Québec, Canada</nlm:aff></affiliation><affiliation><nlm:aff id="A3">Montreal Heart Institute, Research Center, Montréal, Québec, Canada</nlm:aff></affiliation></author><author><name sortKey="Imielinski, Marcin" sort="Imielinski, Marcin" uniqKey="Imielinski M" first="Marcin" last="Imielinski">Marcin Imielinski</name><affiliation><nlm:aff id="A9">The Children’s Hospital of Philadelphia, Center for Applied Genomics, Philadelphia, Pennsylvania, USA</nlm:aff></affiliation></author><author><name sortKey="Latiano, Anna" sort="Latiano, Anna" uniqKey="Latiano A" first="Anna" last="Latiano">Anna Latiano</name><affiliation><nlm:aff id="A10">IRCCS-CSS Hospital, Unit of Gastroenterology, San Giovanni Rotondo, Italy</nlm:aff></affiliation></author><author><name sortKey="Lagace, Caroline" sort="Lagace, Caroline" uniqKey="Lagace C" first="Caroline" last="Lagacé">Caroline Lagacé</name><affiliation><nlm:aff id="A2">Université de Montréal, Medicine, Montréal, Québec, Canada</nlm:aff></affiliation><affiliation><nlm:aff id="A3">Montreal Heart Institute, Research Center, Montréal, Québec, Canada</nlm:aff></affiliation></author><author><name sortKey="Scott, Regan" sort="Scott, Regan" uniqKey="Scott R" first="Regan" last="Scott">Regan Scott</name><affiliation><nlm:aff id="A11">University of Pittsburgh School of Medicine, Division of Gastroenterology, Hepatology and Nutrition, Department of Medicine, Pittsburgh, Pennsylvania, USA</nlm:aff></affiliation></author><author><name sortKey="Amininejad, Leila" sort="Amininejad, Leila" uniqKey="Amininejad L" first="Leila" last="Amininejad">Leila Amininejad</name><affiliation><nlm:aff id="A12">Erasmus Hospital, Free University of Brussels, Department of Gastroenterology, Brussels, Belgium</nlm:aff></affiliation></author><author><name sortKey="Bumpstead, Suzannah" sort="Bumpstead, Suzannah" uniqKey="Bumpstead S" first="Suzannah" last="Bumpstead">Suzannah Bumpstead</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Baidoo, Leonard" sort="Baidoo, Leonard" uniqKey="Baidoo L" first="Leonard" last="Baidoo">Leonard Baidoo</name><affiliation><nlm:aff id="A11">University of Pittsburgh School of Medicine, Division of Gastroenterology, Hepatology and Nutrition, Department of Medicine, Pittsburgh, Pennsylvania, USA</nlm:aff></affiliation></author><author><name sortKey="Baldassano, Robert N" sort="Baldassano, Robert N" uniqKey="Baldassano R" first="Robert N." last="Baldassano">Robert N. Baldassano</name><affiliation><nlm:aff id="A13">The Children’s Hospital of Philadelphia, Department of Pediatrics, Center for Pediatric Inflammatory Bowel Disease, Philadelphia, Pennsylvania, USA</nlm:aff></affiliation></author><author><name sortKey="Barclay, Murray" sort="Barclay, Murray" uniqKey="Barclay M" first="Murray" last="Barclay">Murray Barclay</name><affiliation><nlm:aff id="A14">University of Otago, Department of Medicine, Christchurch, New Zealand</nlm:aff></affiliation></author><author><name sortKey="Bayless, Theodore M" sort="Bayless, Theodore M" uniqKey="Bayless T" first="Theodore M." last="Bayless">Theodore M. Bayless</name><affiliation><nlm:aff id="A15">Johns Hopkins University School of Medicine, Meyeroff Inflammatory Bowel Disease Center, Dept. of Medicine, Baltimore, Maryland, USA</nlm:aff></affiliation></author><author><name sortKey="Brand, Stephan" sort="Brand, Stephan" uniqKey="Brand S" first="Stephan" last="Brand">Stephan Brand</name><affiliation><nlm:aff id="A16">University Hospital Munich, Department of Medicine II, Munich, Germany</nlm:aff></affiliation></author><author><name sortKey="Buning, Carsten" sort="Buning, Carsten" uniqKey="Buning C" first="Carsten" last="Büning">Carsten Büning</name><affiliation><nlm:aff id="A17">Universitätsmedizin Berlin, Department of Gastroenterology, Charité, Campus Mitte, Berlin, Germany</nlm:aff></affiliation></author><author><name sortKey="Colombel, Jean Frederic" sort="Colombel, Jean Frederic" uniqKey="Colombel J" first="Jean-Frédéric" last="Colombel">Jean-Frédéric Colombel</name><affiliation><nlm:aff id="A18">Université de Lille Department of Hepato-Gastroenterology, Lille, France</nlm:aff></affiliation></author><author><name sortKey="Denson, Lee A" sort="Denson, Lee A" uniqKey="Denson L" first="Lee A." last="Denson">Lee A. Denson</name><affiliation><nlm:aff id="A19">Cincinnati Children’s Hospital Medical Center, Pediatric Gastroenterology, Cincinnati, Ohio, USA</nlm:aff></affiliation></author><author><name sortKey="De Vos, Martine" sort="De Vos, Martine" uniqKey="De Vos M" first="Martine" last="De Vos">Martine De Vos</name><affiliation><nlm:aff id="A20">Ghent University Hospital, Department of Hepatology and Gastroenterology, Ghent, Belgium</nlm:aff></affiliation></author><author><name sortKey="Dubinsky, Marla" sort="Dubinsky, Marla" uniqKey="Dubinsky M" first="Marla" last="Dubinsky">Marla Dubinsky</name><affiliation><nlm:aff id="A21">Cedars-Sinai Medical Center, Department of Pediatrics, Los Angeles, California, USA</nlm:aff></affiliation></author><author><name sortKey="Edwards, Cathryn" sort="Edwards, Cathryn" uniqKey="Edwards C" first="Cathryn" last="Edwards">Cathryn Edwards</name><affiliation><nlm:aff id="A22">Torbay Hospital, Department of Gastroenterology, Torbay, Devon, UK</nlm:aff></affiliation></author><author><name sortKey="Ellinghaus, David" sort="Ellinghaus, David" uniqKey="Ellinghaus D" first="David" last="Ellinghaus">David Ellinghaus</name><affiliation><nlm:aff id="A5">Christian-Albrechts-University Kiel, Institute of Clinical Molecular Biology, Kiel, Germany</nlm:aff></affiliation></author><author><name sortKey="Fehrmann, Rudolf S N" sort="Fehrmann, Rudolf S N" uniqKey="Fehrmann R" first="Rudolf S. N." last="Fehrmann">Rudolf S. N. Fehrmann</name><affiliation><nlm:aff id="A23">University Medical Center Groningen, Department of Genetics, Groningen, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Floyd, James A B" sort="Floyd, James A B" uniqKey="Floyd J" first="James A. B." last="Floyd">James A. B. Floyd</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Florin, Tim" sort="Florin, Tim" uniqKey="Florin T" first="Tim" last="Florin">Tim Florin</name><affiliation><nlm:aff id="A24">Mater Health Services, Department of Gastroenterology, Brisbane, Australia</nlm:aff></affiliation></author><author><name sortKey="Franchimont, Denis" sort="Franchimont, Denis" uniqKey="Franchimont D" first="Denis" last="Franchimont">Denis Franchimont</name><affiliation><nlm:aff id="A25">Erasmus Hospital, Free University of Brussels, Department of Gastroenterology, Brussels, Belgium</nlm:aff></affiliation></author><author><name sortKey="Franke, Lude" sort="Franke, Lude" uniqKey="Franke L" first="Lude" last="Franke">Lude Franke</name><affiliation><nlm:aff id="A23">University Medical Center Groningen, Department of Genetics, Groningen, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Georges, Michel" sort="Georges, Michel" uniqKey="Georges M" first="Michel" last="Georges">Michel Georges</name><affiliation><nlm:aff id="A26">University of Liège, Department of Genetics, Faculty of Veterinary Medicine, Liège, Belgium</nlm:aff></affiliation></author><author><name sortKey="Glas, Jurgen" sort="Glas, Jurgen" uniqKey="Glas J" first="Jürgen" last="Glas">Jürgen Glas</name><affiliation><nlm:aff id="A16">University Hospital Munich, Department of Medicine II, Munich, Germany</nlm:aff></affiliation></author><author><name sortKey="Glazer, Nicole L" sort="Glazer, Nicole L" uniqKey="Glazer N" first="Nicole L." last="Glazer">Nicole L. Glazer</name><affiliation><nlm:aff id="A27">University of Washington, Cardiovascular Health Research Unit, Department of Internal Medicine, Seattle, Washington, USA</nlm:aff></affiliation></author><author><name sortKey="Guthery, Stephen L" sort="Guthery, Stephen L" uniqKey="Guthery S" first="Stephen L." last="Guthery">Stephen L. Guthery</name><affiliation><nlm:aff id="A28">University of Utah School of Medicine, Department of Pediatrics, Salt Lake City, Utah, USA</nlm:aff></affiliation></author><author><name sortKey="Haritunians, Talin" sort="Haritunians, Talin" uniqKey="Haritunians T" first="Talin" last="Haritunians">Talin Haritunians</name><affiliation><nlm:aff id="A29">Cedars-Sinai Medical Center, Medical Genetics Institute, Los Angeles, California, USA</nlm:aff></affiliation></author><author><name sortKey="Hayward, Nicholas K" sort="Hayward, Nicholas K" uniqKey="Hayward N" first="Nicholas K." last="Hayward">Nicholas K. Hayward</name><affiliation><nlm:aff id="A30">Queensland Institute of Medical Research, Genetic Epidemiology, Brisbane, Australia</nlm:aff></affiliation></author><author><name sortKey="Hugot, Jean Pierre" sort="Hugot, Jean Pierre" uniqKey="Hugot J" first="Jean-Pierre" last="Hugot">Jean-Pierre Hugot</name><affiliation><nlm:aff id="A31">Université Paris Diderot & INSERM & Hopital Robert Debre APHP, Gastroenterology, Paris, France</nlm:aff></affiliation></author><author><name sortKey="Jobin, Gilles" sort="Jobin, Gilles" uniqKey="Jobin G" first="Gilles" last="Jobin">Gilles Jobin</name><affiliation><nlm:aff id="A2">Université de Montréal, Medicine, Montréal, Québec, Canada</nlm:aff></affiliation><affiliation><nlm:aff id="A32">Hôpital Maisonneuve-Rosemont, Dept of Gastroenterology, Montréal, Québec, Canada</nlm:aff></affiliation></author><author><name sortKey="Laukens, Debby" sort="Laukens, Debby" uniqKey="Laukens D" first="Debby" last="Laukens">Debby Laukens</name><affiliation><nlm:aff id="A20">Ghent University Hospital, Department of Hepatology and Gastroenterology, Ghent, Belgium</nlm:aff></affiliation></author><author><name sortKey="Lawrance, Ian" sort="Lawrance, Ian" uniqKey="Lawrance I" first="Ian" last="Lawrance">Ian Lawrance</name><affiliation><nlm:aff id="A33">The University of Western Australia, School of Medicine and Pharmacology, Fremantle, Australia</nlm:aff></affiliation></author><author><name sortKey="Lemann, Marc" sort="Lemann, Marc" uniqKey="Lemann M" first="Marc" last="Lémann">Marc Lémann</name><affiliation><nlm:aff id="A34">Université Paris Diderot, GETAID group, Paris, France</nlm:aff></affiliation></author><author><name sortKey="Levine, Arie" sort="Levine, Arie" uniqKey="Levine A" first="Arie" last="Levine">Arie Levine</name><affiliation><nlm:aff id="A35">Tel Aviv University, Pediatric Gastroenterology Unit, Wolfson Medical Center and Sackler School of Medicine, Tel Aviv, Israel</nlm:aff></affiliation></author><author><name sortKey="Libioulle, Cecile" sort="Libioulle, Cecile" uniqKey="Libioulle C" first="Cecile" last="Libioulle">Cecile Libioulle</name><affiliation><nlm:aff id="A36">Centre Hospitalier Universitaire Université de Liège, Division of Gastroenterology, Liège, Belgium</nlm:aff></affiliation></author><author><name sortKey="Louis, Edouard" sort="Louis, Edouard" uniqKey="Louis E" first="Edouard" last="Louis">Edouard Louis</name><affiliation><nlm:aff id="A36">Centre Hospitalier Universitaire Université de Liège, Division of Gastroenterology, Liège, Belgium</nlm:aff></affiliation></author><author><name sortKey="Mcgovern, Dermot P" sort="Mcgovern, Dermot P" uniqKey="Mcgovern D" first="Dermot P." last="Mcgovern">Dermot P. Mcgovern</name><affiliation><nlm:aff id="A7">Cedars-Sinai Medical Center, Inflammatory Bowel and Immunobiology Research Institute, Los Angeles, California, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A29">Cedars-Sinai Medical Center, Medical Genetics Institute, Los Angeles, California, USA</nlm:aff></affiliation></author><author><name sortKey="Milla, Monica" sort="Milla, Monica" uniqKey="Milla M" first="Monica" last="Milla">Monica Milla</name><affiliation><nlm:aff id="A37">AOU Careggi, Unit of Gastroenterology SOD2, Florence, Italy</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="A30">Queensland Institute of Medical Research, Genetic Epidemiology, Brisbane, Australia</nlm:aff></affiliation></author><author><name sortKey="Morley, Katherine I" sort="Morley, Katherine I" uniqKey="Morley K" first="Katherine I." last="Morley">Katherine I. Morley</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Mowat, Craig" sort="Mowat, Craig" uniqKey="Mowat C" first="Craig" last="Mowat">Craig Mowat</name><affiliation><nlm:aff id="A38">Ninewells Hospital and Medical School, Dept of Medicine, Dundee, UK</nlm:aff></affiliation></author><author><name sortKey="Ng, Aylwin" sort="Ng, Aylwin" uniqKey="Ng A" first="Aylwin" last="Ng">Aylwin Ng</name><affiliation><nlm:aff id="A39">Massachusetts General Hospital, Harvard Medical School, Gastroenterology Unit Boston, Massachusetts, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A40">Center for Computational and Integrative Biology, Massachusetts General Hospital, Boston, Massachusetts, USA</nlm:aff></affiliation></author><author><name sortKey="Newman, William" sort="Newman, William" uniqKey="Newman W" first="William" last="Newman">William Newman</name><affiliation><nlm:aff id="A41">University of Manchester Department of Medical Genetics, Manchester, UK</nlm:aff></affiliation></author><author><name sortKey="Ophoff, Roel A" sort="Ophoff, Roel A" uniqKey="Ophoff R" first="Roel A" last="Ophoff">Roel A. Ophoff</name><affiliation><nlm:aff id="A42">University Medical Center Utrecht, Department of Medical Genetics, Utrecht, Netherlands</nlm:aff></affiliation></author><author><name sortKey="Papi, Laura" sort="Papi, Laura" uniqKey="Papi L" first="Laura" last="Papi">Laura Papi</name><affiliation><nlm:aff id="A43">University of Florence, Institute of Human Genetics, Florence, Italy</nlm:aff></affiliation></author><author><name sortKey="Palmieri, Orazio" sort="Palmieri, Orazio" uniqKey="Palmieri O" first="Orazio" last="Palmieri">Orazio Palmieri</name><affiliation><nlm:aff id="A10">IRCCS-CSS Hospital, Unit of Gastroenterology, San Giovanni Rotondo, Italy</nlm:aff></affiliation></author><author><name sortKey="Peyrin Biroulet, Laurent" sort="Peyrin Biroulet, Laurent" uniqKey="Peyrin Biroulet L" first="Laurent" last="Peyrin-Biroulet">Laurent Peyrin-Biroulet</name><affiliation><nlm:aff id="A44">University Hospital of Nancy, Department of Hepato-Gastroenterology, Vandoeuvre-lès-Nancy, France</nlm:aff></affiliation></author><author><name sortKey="Panes, Julian" sort="Panes, Julian" uniqKey="Panes J" first="Julián" last="Panés">Julián Panés</name><affiliation><nlm:aff id="A45">Hospital Clínic de Barcelona, IDIBAPS, CIBERehd, Department of Gastroenterology, Barcelona, Spain</nlm:aff></affiliation></author><author><name sortKey="Phillips, Anne" sort="Phillips, Anne" uniqKey="Phillips A" first="Anne" last="Phillips">Anne Phillips</name><affiliation><nlm:aff id="A38">Ninewells Hospital and Medical School, Dept of Medicine, Dundee, UK</nlm:aff></affiliation></author><author><name sortKey="Prescott, Natalie J" sort="Prescott, Natalie J" uniqKey="Prescott N" first="Natalie J." last="Prescott">Natalie J. Prescott</name><affiliation><nlm:aff id="A46">King’s College London School of Medicine, Guy’s Hospital, Department of Medical and Molecular Genetics, London, UK</nlm:aff></affiliation></author><author><name sortKey="Proctor, Deborah D" sort="Proctor, Deborah D" uniqKey="Proctor D" first="Deborah D." last="Proctor">Deborah D. Proctor</name><affiliation><nlm:aff id="A47">Yale University, Section of Digestive Diseases, Department of Medicine, New Haven, Connecticut, USA</nlm:aff></affiliation></author><author><name sortKey="Roberts, Rebecca" sort="Roberts, Rebecca" uniqKey="Roberts R" first="Rebecca" last="Roberts">Rebecca Roberts</name><affiliation><nlm:aff id="A14">University of Otago, Department of Medicine, Christchurch, New Zealand</nlm:aff></affiliation></author><author><name sortKey="Russell, Richard" sort="Russell, Richard" uniqKey="Russell R" first="Richard" last="Russell">Richard Russell</name><affiliation><nlm:aff id="A48">Royal Hospital for Sick Children, Paediatric Gastroenterology and Nutrition, Glasgow, UK</nlm:aff></affiliation></author><author><name sortKey="Rutgeerts, Paul" sort="Rutgeerts, Paul" uniqKey="Rutgeerts P" first="Paul" last="Rutgeerts">Paul Rutgeerts</name><affiliation><nlm:aff id="A49">University Hospital Gasthuisberg, Division of Gastroenterology, Leuven, Belgium</nlm:aff></affiliation></author><author><name sortKey="Sanderson, Jeremy" sort="Sanderson, Jeremy" uniqKey="Sanderson J" first="Jeremy" last="Sanderson">Jeremy Sanderson</name><affiliation><nlm:aff id="A50">Guy’s & St Thomas’ NHS Foundation Trust, St Thomas’ Hospital, Dept Gastroenterology, London, UK</nlm:aff></affiliation></author><author><name sortKey="Sans, Miquel" sort="Sans, Miquel" uniqKey="Sans M" first="Miquel" last="Sans">Miquel Sans</name><affiliation><nlm:aff id="A51">Hospital Clínic de Barcelona, IDIBAPS, CIBERehd, Department of Gastroenterology, Barcelona, Spain</nlm:aff></affiliation></author><author><name sortKey="Schumm, Philip" sort="Schumm, Philip" uniqKey="Schumm P" first="Philip" last="Schumm">Philip Schumm</name><affiliation><nlm:aff id="A52">University of Chicago, Department of Health Studies, Chicago, Illinois, USA</nlm:aff></affiliation></author><author><name sortKey="Seibold, Frank" sort="Seibold, Frank" uniqKey="Seibold F" first="Frank" last="Seibold">Frank Seibold</name><affiliation><nlm:aff id="A53">University of Bern, Division of Gastroenterology, Inselspital, Bern, Switzerland</nlm:aff></affiliation></author><author><name sortKey="Sharma, Yashoda" sort="Sharma, Yashoda" uniqKey="Sharma Y" first="Yashoda" last="Sharma">Yashoda Sharma</name><affiliation><nlm:aff id="A47">Yale University, Section of Digestive Diseases, Department of Medicine, New Haven, Connecticut, USA</nlm:aff></affiliation></author><author><name sortKey="Simms, Lisa" sort="Simms, Lisa" uniqKey="Simms L" first="Lisa" last="Simms">Lisa Simms</name><affiliation><nlm:aff id="A30">Queensland Institute of Medical Research, Genetic Epidemiology, Brisbane, Australia</nlm:aff></affiliation></author><author><name sortKey="Seielstad, Mark" sort="Seielstad, Mark" uniqKey="Seielstad M" first="Mark" last="Seielstad">Mark Seielstad</name><affiliation><nlm:aff id="A54">Genome Institute of Singapore, Human Genetics, Singapore</nlm:aff></affiliation><affiliation><nlm:aff id="A55">Institute for Human Genetics, University of California San Fransisco, San Francisco, California, USA</nlm:aff></affiliation></author><author><name sortKey="Steinhart, A Hillary" sort="Steinhart, A Hillary" uniqKey="Steinhart A" first="A. Hillary" last="Steinhart">A. Hillary Steinhart</name><affiliation><nlm:aff id="A56">University of Toronto, Mount Sinai Hospital Inflammatory Bowel Disease Centre, Toronto, Ontario, Canada</nlm:aff></affiliation></author><author><name sortKey="Targan, Stephan R" sort="Targan, Stephan R" uniqKey="Targan S" first="Stephan R." last="Targan">Stephan R. Targan</name><affiliation><nlm:aff id="A7">Cedars-Sinai Medical Center, Inflammatory Bowel and Immunobiology Research Institute, Los Angeles, California, USA</nlm:aff></affiliation></author><author><name sortKey="Van Den Berg, Leonard H" sort="Van Den Berg, Leonard H" uniqKey="Van Den Berg L" first="Leonard H." last="Van Den Berg">Leonard H. Van Den Berg</name><affiliation><nlm:aff id="A57">Rudolf Magnus Institute of Neuroscience, University Medical Center Utrecht, Department of Neurology, Utrecht, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Vatn, Morten" sort="Vatn, Morten" uniqKey="Vatn M" first="Morten" last="Vatn">Morten Vatn</name><affiliation><nlm:aff id="A58">Rikshospitalet University Hospital, Medical Department, Oslo, Norway</nlm:aff></affiliation></author><author><name sortKey="Verspaget, Hein" sort="Verspaget, Hein" uniqKey="Verspaget H" first="Hein" last="Verspaget">Hein Verspaget</name><affiliation><nlm:aff id="A59">Leiden University Medical Center, Experimental Gastroenterology, Leiden, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Walters, Thomas" sort="Walters, Thomas" uniqKey="Walters T" first="Thomas" last="Walters">Thomas Walters</name><affiliation><nlm:aff id="A60">The Hospital for Sick Children, University of Toronto, Toronto, Ontario, Canada</nlm:aff></affiliation></author><author><name sortKey="Wijmenga, Cisca" sort="Wijmenga, Cisca" uniqKey="Wijmenga C" first="Cisca" last="Wijmenga">Cisca Wijmenga</name><affiliation><nlm:aff id="A23">University Medical Center Groningen, Department of Genetics, Groningen, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Wilson, David C" sort="Wilson, David C" uniqKey="Wilson D" first="David C." last="Wilson">David C. Wilson</name><affiliation><nlm:aff id="A48">Royal Hospital for Sick Children, Paediatric Gastroenterology and Nutrition, Glasgow, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A61">Child Life and Health, University of Edinburgh, Scotland</nlm:aff></affiliation></author><author><name sortKey="Westra, Harm Jan" sort="Westra, Harm Jan" uniqKey="Westra H" first="Harm-Jan" last="Westra">Harm-Jan Westra</name><affiliation><nlm:aff id="A23">University Medical Center Groningen, Department of Genetics, Groningen, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Xavier, Ramnik J" sort="Xavier, Ramnik J" uniqKey="Xavier R" first="Ramnik J." last="Xavier">Ramnik J. Xavier</name><affiliation><nlm:aff id="A39">Massachusetts General Hospital, Harvard Medical School, Gastroenterology Unit Boston, Massachusetts, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A40">Center for Computational and Integrative Biology, Massachusetts General Hospital, Boston, Massachusetts, USA</nlm:aff></affiliation></author><author><name sortKey="Zhao, Zhen Z" sort="Zhao, Zhen Z" uniqKey="Zhao Z" first="Zhen Z." last="Zhao">Zhen Z. Zhao</name><affiliation><nlm:aff id="A30">Queensland Institute of Medical Research, Genetic Epidemiology, Brisbane, Australia</nlm:aff></affiliation></author><author><name sortKey="Ponsioen, Cyriel Y" sort="Ponsioen, Cyriel Y" uniqKey="Ponsioen C" first="Cyriel Y." last="Ponsioen">Cyriel Y. Ponsioen</name><affiliation><nlm:aff id="A62">Academic Medical Center, Department of Gastroenterology, Amsterdam, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Andersen, Vibeke" sort="Andersen, Vibeke" uniqKey="Andersen V" first="Vibeke" last="Andersen">Vibeke Andersen</name><affiliation><nlm:aff id="A63">Viborg Regional Hospital, Medical Department, Viborg, Denmark</nlm:aff></affiliation></author><author><name sortKey="Torkvist, Leif" sort="Torkvist, Leif" uniqKey="Torkvist L" first="Leif" last="Torkvist">Leif Torkvist</name><affiliation><nlm:aff id="A64">Karolinska Institutet, Department of Clinical Science Intervention and Technology, Stockholm, Sweden</nlm:aff></affiliation></author><author><name sortKey="Gazouli, Maria" sort="Gazouli, Maria" uniqKey="Gazouli M" first="Maria" last="Gazouli">Maria Gazouli</name><affiliation><nlm:aff id="A65">University of Athens, Department of Biology, School of Medicine, Athens, Greece</nlm:aff></affiliation></author><author><name sortKey="Anagnou, Nicholas P" sort="Anagnou, Nicholas P" uniqKey="Anagnou N" first="Nicholas P." last="Anagnou">Nicholas P. Anagnou</name><affiliation><nlm:aff id="A65">University of Athens, Department of Biology, School of Medicine, Athens, Greece</nlm:aff></affiliation></author><author><name sortKey="Karlsen, Tom H" sort="Karlsen, Tom H" uniqKey="Karlsen T" first="Tom H." last="Karlsen">Tom H. Karlsen</name><affiliation><nlm:aff id="A58">Rikshospitalet University Hospital, Medical Department, Oslo, Norway</nlm:aff></affiliation></author><author><name sortKey="Kupcinskas, Limas" sort="Kupcinskas, Limas" uniqKey="Kupcinskas L" first="Limas" last="Kupcinskas">Limas Kupcinskas</name><affiliation><nlm:aff id="A66">Kaunas University of medicine, Department of Gastroenterology, Kaunas, Lithuania</nlm:aff></affiliation></author><author><name sortKey="Sventoraityte, Jurgita" sort="Sventoraityte, Jurgita" uniqKey="Sventoraityte J" first="Jurgita" last="Sventoraityte">Jurgita Sventoraityte</name><affiliation><nlm:aff id="A66">Kaunas University of medicine, Department of Gastroenterology, Kaunas, Lithuania</nlm:aff></affiliation></author><author><name sortKey="Mansfield, John C" sort="Mansfield, John C" uniqKey="Mansfield J" first="John C." last="Mansfield">John C. Mansfield</name><affiliation><nlm:aff id="A67">Newcastle University, Institute of Human Genetics, Newcastle upon Tyne, UK</nlm:aff></affiliation></author><author><name sortKey="Kugathasan, Subra" sort="Kugathasan, Subra" uniqKey="Kugathasan S" first="Subra" last="Kugathasan">Subra Kugathasan</name><affiliation><nlm:aff id="A68">Emory School of Medicine, Department of Genetics and Department of Pediatrics, Atlanta, Georgia, USA</nlm:aff></affiliation></author><author><name sortKey="Silverberg, Mark S" sort="Silverberg, Mark S" uniqKey="Silverberg M" first="Mark S." last="Silverberg">Mark S. Silverberg</name><affiliation><nlm:aff id="A56">University of Toronto, Mount Sinai Hospital Inflammatory Bowel Disease Centre, Toronto, Ontario, Canada</nlm:aff></affiliation></author><author><name sortKey="Halfvarson, Jonas" sort="Halfvarson, Jonas" uniqKey="Halfvarson J" first="Jonas" last="Halfvarson">Jonas Halfvarson</name><affiliation><nlm:aff id="A69">Örebro University Hospital, Department of Medicine, Örebro, Sweden</nlm:aff></affiliation></author><author><name sortKey="Rotter, Jerome I" sort="Rotter, Jerome I" uniqKey="Rotter J" first="Jerome I." last="Rotter">Jerome I. Rotter</name><affiliation><nlm:aff id="A29">Cedars-Sinai Medical Center, Medical Genetics Institute, Los Angeles, California, USA</nlm:aff></affiliation></author><author><name sortKey="Mathew, Christopher G" sort="Mathew, Christopher G" uniqKey="Mathew C" first="Christopher G." last="Mathew">Christopher G. Mathew</name><affiliation><nlm:aff id="A46">King’s College London School of Medicine, Guy’s Hospital, Department of Medical and Molecular Genetics, London, UK</nlm:aff></affiliation></author><author><name sortKey="Griffiths, Anne M" sort="Griffiths, Anne M" uniqKey="Griffiths A" first="Anne M." last="Griffiths">Anne M. Griffiths</name><affiliation><nlm:aff id="A60">The Hospital for Sick Children, University of Toronto, Toronto, Ontario, Canada</nlm:aff></affiliation></author><author><name sortKey="Gearry, Richard" sort="Gearry, Richard" uniqKey="Gearry R" first="Richard" last="Gearry">Richard Gearry</name><affiliation><nlm:aff id="A14">University of Otago, Department of Medicine, Christchurch, New Zealand</nlm:aff></affiliation></author><author><name sortKey="Ahmad, Tariq" sort="Ahmad, Tariq" uniqKey="Ahmad T" first="Tariq" last="Ahmad">Tariq Ahmad</name><affiliation><nlm:aff id="A70">Peninsula College of Medicine and Dentistry, Barrack Road, Exeter, UK</nlm:aff></affiliation></author><author><name sortKey="Brant, Steven R" sort="Brant, Steven R" uniqKey="Brant S" first="Steven R." last="Brant">Steven R. Brant</name><affiliation><nlm:aff id="A15">Johns Hopkins University School of Medicine, Meyeroff Inflammatory Bowel Disease Center, Dept. of Medicine, Baltimore, Maryland, USA</nlm:aff></affiliation></author><author><name sortKey="Chamaillard, Mathias" sort="Chamaillard, Mathias" uniqKey="Chamaillard M" first="Mathias" last="Chamaillard">Mathias Chamaillard</name><affiliation><nlm:aff id="A71">Inserm, U1019, Lille, France</nlm:aff></affiliation></author><author><name sortKey="Satsangi, Jack" sort="Satsangi, Jack" uniqKey="Satsangi J" first="Jack" last="Satsangi">Jack Satsangi</name><affiliation><nlm:aff id="A4">University of Edinburgh, Western General Hospital, Gastrointestinal Unit, Molecular Medicine Centre, Edinburgh, UK</nlm:aff></affiliation></author><author><name sortKey="Cho, Judy H" sort="Cho, Judy H" uniqKey="Cho J" first="Judy H." last="Cho">Judy H. Cho</name><affiliation><nlm:aff id="A47">Yale University, Section of Digestive Diseases, Department of Medicine, New Haven, Connecticut, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A72">Yale University, Department of Genetics, Yale School of Medicine, New Haven CT</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="A5">Christian-Albrechts-University Kiel, Institute of Clinical Molecular Biology, Kiel, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="A73">Department for General Internal Medicine, Christian-Albrechts-University, Kiel, Germany</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="A74">Massachusetts General Hospital, Harvard Medical School, Center for Human Genetic Research, Boston, Massachusetts, USA</nlm:aff></affiliation></author><author><name sortKey="Barrett, Jeffrey C" sort="Barrett, Jeffrey C" uniqKey="Barrett J" first="Jeffrey C." last="Barrett">Jeffrey C. Barrett</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Parkes, Miles" sort="Parkes, Miles" uniqKey="Parkes M" first="Miles" last="Parkes">Miles Parkes</name><affiliation><nlm:aff id="A8">Addenbrooke’s Hospital, University of Cambridge, Gastroenterology Research Unit, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Annese, Vito" sort="Annese, Vito" uniqKey="Annese V" first="Vito" last="Annese">Vito Annese</name><affiliation><nlm:aff id="A10">IRCCS-CSS Hospital, Unit of Gastroenterology, San Giovanni Rotondo, Italy</nlm:aff></affiliation><affiliation><nlm:aff id="A37">AOU Careggi, Unit of Gastroenterology SOD2, Florence, Italy</nlm:aff></affiliation></author><author><name sortKey="Hakonarson, Hakon" sort="Hakonarson, Hakon" uniqKey="Hakonarson H" first="Hakon" last="Hakonarson">Hakon Hakonarson</name><affiliation><nlm:aff id="A13">The Children’s Hospital of Philadelphia, Department of Pediatrics, Center for Pediatric Inflammatory Bowel Disease, Philadelphia, Pennsylvania, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A75">Center for Applied Genomics, The Children’s Hospital of Philadelphia, Philadelphia, Pennsylvania, USA</nlm:aff></affiliation></author><author><name sortKey="Radford Smith, Graham" sort="Radford Smith, Graham" uniqKey="Radford Smith G" first="Graham" last="Radford-Smith">Graham Radford-Smith</name><affiliation><nlm:aff id="A76">Queensland Institute of Medical Research, IBD Research Group, Brisbane, Australia</nlm:aff></affiliation></author><author><name sortKey="Duerr, Richard H" sort="Duerr, Richard H" uniqKey="Duerr R" first="Richard H." last="Duerr">Richard H. Duerr</name><affiliation><nlm:aff id="A11">University of Pittsburgh School of Medicine, Division of Gastroenterology, Hepatology and Nutrition, Department of Medicine, Pittsburgh, Pennsylvania, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A77">University of Pittsburgh Graduate School of Public Health, 130 Desoto Street, Pittsburgh, PA, USA</nlm:aff></affiliation></author><author><name sortKey="Vermeire, Severine" sort="Vermeire, Severine" uniqKey="Vermeire S" first="Séverine" last="Vermeire">Séverine Vermeire</name><affiliation><nlm:aff id="A49">University Hospital Gasthuisberg, Division of Gastroenterology, Leuven, Belgium</nlm:aff></affiliation></author><author><name sortKey="Weersma, Rinse K" sort="Weersma, Rinse K" uniqKey="Weersma R" first="Rinse K." last="Weersma">Rinse K. Weersma</name><affiliation><nlm:aff id="A78">University Medical Center Groningen, Department of Gastroenterology, Groningen, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Rioux, John D" sort="Rioux, John D" uniqKey="Rioux J" first="John D." last="Rioux">John D. Rioux</name><affiliation><nlm:aff id="A2">Université de Montréal, Medicine, Montréal, Québec, Canada</nlm:aff></affiliation><affiliation><nlm:aff id="A3">Montreal Heart Institute, Research Center, Montréal, Québec, Canada</nlm:aff></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">PMC</idno><idno type="pmid">21297633</idno><idno type="pmc">3084597</idno><idno type="url">http://www.ncbi.nlm.nih.gov/pmc/articles/PMC3084597</idno><idno type="RBID">PMC:3084597</idno><idno type="doi">10.1038/ng.764</idno><date when="2011">2011</date><idno type="wicri:Area/Pmc/Corpus">002E05</idno><idno type="wicri:explorRef" wicri:stream="Pmc" wicri:step="Corpus" wicri:corpus="PMC">002E05</idno></publicationStmt><sourceDesc><biblStruct><analytic><title xml:lang="en" level="a" type="main">Meta-analysis identifies 29 additional ulcerative colitis risk loci, increasing the number of confirmed associations to 47</title><author><name sortKey="Anderson, Carl A" sort="Anderson, Carl A" uniqKey="Anderson C" first="Carl A." last="Anderson">Carl A. Anderson</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Boucher, Gabrielle" sort="Boucher, Gabrielle" uniqKey="Boucher G" first="Gabrielle" last="Boucher">Gabrielle Boucher</name><affiliation><nlm:aff id="A2">Université de Montréal, Medicine, Montréal, Québec, Canada</nlm:aff></affiliation><affiliation><nlm:aff id="A3">Montreal Heart Institute, Research Center, Montréal, Québec, Canada</nlm:aff></affiliation></author><author><name sortKey="Lees, Charlie W" sort="Lees, Charlie W" uniqKey="Lees C" first="Charlie W." last="Lees">Charlie W. Lees</name><affiliation><nlm:aff id="A4">University of Edinburgh, Western General Hospital, Gastrointestinal Unit, Molecular Medicine Centre, Edinburgh, UK</nlm:aff></affiliation></author><author><name sortKey="Franke, Andre" sort="Franke, Andre" uniqKey="Franke A" first="Andre" last="Franke">Andre Franke</name><affiliation><nlm:aff id="A5">Christian-Albrechts-University Kiel, Institute of Clinical Molecular Biology, Kiel, Germany</nlm:aff></affiliation></author><author><name sortKey="D Mato, Mauro" sort="D Mato, Mauro" uniqKey="D Mato M" first="Mauro" last="D Mato">Mauro D Mato</name><affiliation><nlm:aff id="A6">Karolinska Institute, Department of Biosciences and Nutrition, Stockholm, Sweden</nlm:aff></affiliation></author><author><name sortKey="Taylor, Kent D" sort="Taylor, Kent D" uniqKey="Taylor K" first="Kent D." last="Taylor">Kent D. Taylor</name><affiliation><nlm:aff id="A7">Cedars-Sinai Medical Center, Inflammatory Bowel and Immunobiology Research Institute, Los Angeles, California, USA</nlm:aff></affiliation></author><author><name sortKey="Lee, James C" sort="Lee, James C" uniqKey="Lee J" first="James C." last="Lee">James C. Lee</name><affiliation><nlm:aff id="A8">Addenbrooke’s Hospital, University of Cambridge, Gastroenterology Research Unit, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Goyette, Philippe" sort="Goyette, Philippe" uniqKey="Goyette P" first="Philippe" last="Goyette">Philippe Goyette</name><affiliation><nlm:aff id="A2">Université de Montréal, Medicine, Montréal, Québec, Canada</nlm:aff></affiliation><affiliation><nlm:aff id="A3">Montreal Heart Institute, Research Center, Montréal, Québec, Canada</nlm:aff></affiliation></author><author><name sortKey="Imielinski, Marcin" sort="Imielinski, Marcin" uniqKey="Imielinski M" first="Marcin" last="Imielinski">Marcin Imielinski</name><affiliation><nlm:aff id="A9">The Children’s Hospital of Philadelphia, Center for Applied Genomics, Philadelphia, Pennsylvania, USA</nlm:aff></affiliation></author><author><name sortKey="Latiano, Anna" sort="Latiano, Anna" uniqKey="Latiano A" first="Anna" last="Latiano">Anna Latiano</name><affiliation><nlm:aff id="A10">IRCCS-CSS Hospital, Unit of Gastroenterology, San Giovanni Rotondo, Italy</nlm:aff></affiliation></author><author><name sortKey="Lagace, Caroline" sort="Lagace, Caroline" uniqKey="Lagace C" first="Caroline" last="Lagacé">Caroline Lagacé</name><affiliation><nlm:aff id="A2">Université de Montréal, Medicine, Montréal, Québec, Canada</nlm:aff></affiliation><affiliation><nlm:aff id="A3">Montreal Heart Institute, Research Center, Montréal, Québec, Canada</nlm:aff></affiliation></author><author><name sortKey="Scott, Regan" sort="Scott, Regan" uniqKey="Scott R" first="Regan" last="Scott">Regan Scott</name><affiliation><nlm:aff id="A11">University of Pittsburgh School of Medicine, Division of Gastroenterology, Hepatology and Nutrition, Department of Medicine, Pittsburgh, Pennsylvania, USA</nlm:aff></affiliation></author><author><name sortKey="Amininejad, Leila" sort="Amininejad, Leila" uniqKey="Amininejad L" first="Leila" last="Amininejad">Leila Amininejad</name><affiliation><nlm:aff id="A12">Erasmus Hospital, Free University of Brussels, Department of Gastroenterology, Brussels, Belgium</nlm:aff></affiliation></author><author><name sortKey="Bumpstead, Suzannah" sort="Bumpstead, Suzannah" uniqKey="Bumpstead S" first="Suzannah" last="Bumpstead">Suzannah Bumpstead</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Baidoo, Leonard" sort="Baidoo, Leonard" uniqKey="Baidoo L" first="Leonard" last="Baidoo">Leonard Baidoo</name><affiliation><nlm:aff id="A11">University of Pittsburgh School of Medicine, Division of Gastroenterology, Hepatology and Nutrition, Department of Medicine, Pittsburgh, Pennsylvania, USA</nlm:aff></affiliation></author><author><name sortKey="Baldassano, Robert N" sort="Baldassano, Robert N" uniqKey="Baldassano R" first="Robert N." last="Baldassano">Robert N. Baldassano</name><affiliation><nlm:aff id="A13">The Children’s Hospital of Philadelphia, Department of Pediatrics, Center for Pediatric Inflammatory Bowel Disease, Philadelphia, Pennsylvania, USA</nlm:aff></affiliation></author><author><name sortKey="Barclay, Murray" sort="Barclay, Murray" uniqKey="Barclay M" first="Murray" last="Barclay">Murray Barclay</name><affiliation><nlm:aff id="A14">University of Otago, Department of Medicine, Christchurch, New Zealand</nlm:aff></affiliation></author><author><name sortKey="Bayless, Theodore M" sort="Bayless, Theodore M" uniqKey="Bayless T" first="Theodore M." last="Bayless">Theodore M. Bayless</name><affiliation><nlm:aff id="A15">Johns Hopkins University School of Medicine, Meyeroff Inflammatory Bowel Disease Center, Dept. of Medicine, Baltimore, Maryland, USA</nlm:aff></affiliation></author><author><name sortKey="Brand, Stephan" sort="Brand, Stephan" uniqKey="Brand S" first="Stephan" last="Brand">Stephan Brand</name><affiliation><nlm:aff id="A16">University Hospital Munich, Department of Medicine II, Munich, Germany</nlm:aff></affiliation></author><author><name sortKey="Buning, Carsten" sort="Buning, Carsten" uniqKey="Buning C" first="Carsten" last="Büning">Carsten Büning</name><affiliation><nlm:aff id="A17">Universitätsmedizin Berlin, Department of Gastroenterology, Charité, Campus Mitte, Berlin, Germany</nlm:aff></affiliation></author><author><name sortKey="Colombel, Jean Frederic" sort="Colombel, Jean Frederic" uniqKey="Colombel J" first="Jean-Frédéric" last="Colombel">Jean-Frédéric Colombel</name><affiliation><nlm:aff id="A18">Université de Lille Department of Hepato-Gastroenterology, Lille, France</nlm:aff></affiliation></author><author><name sortKey="Denson, Lee A" sort="Denson, Lee A" uniqKey="Denson L" first="Lee A." last="Denson">Lee A. Denson</name><affiliation><nlm:aff id="A19">Cincinnati Children’s Hospital Medical Center, Pediatric Gastroenterology, Cincinnati, Ohio, USA</nlm:aff></affiliation></author><author><name sortKey="De Vos, Martine" sort="De Vos, Martine" uniqKey="De Vos M" first="Martine" last="De Vos">Martine De Vos</name><affiliation><nlm:aff id="A20">Ghent University Hospital, Department of Hepatology and Gastroenterology, Ghent, Belgium</nlm:aff></affiliation></author><author><name sortKey="Dubinsky, Marla" sort="Dubinsky, Marla" uniqKey="Dubinsky M" first="Marla" last="Dubinsky">Marla Dubinsky</name><affiliation><nlm:aff id="A21">Cedars-Sinai Medical Center, Department of Pediatrics, Los Angeles, California, USA</nlm:aff></affiliation></author><author><name sortKey="Edwards, Cathryn" sort="Edwards, Cathryn" uniqKey="Edwards C" first="Cathryn" last="Edwards">Cathryn Edwards</name><affiliation><nlm:aff id="A22">Torbay Hospital, Department of Gastroenterology, Torbay, Devon, UK</nlm:aff></affiliation></author><author><name sortKey="Ellinghaus, David" sort="Ellinghaus, David" uniqKey="Ellinghaus D" first="David" last="Ellinghaus">David Ellinghaus</name><affiliation><nlm:aff id="A5">Christian-Albrechts-University Kiel, Institute of Clinical Molecular Biology, Kiel, Germany</nlm:aff></affiliation></author><author><name sortKey="Fehrmann, Rudolf S N" sort="Fehrmann, Rudolf S N" uniqKey="Fehrmann R" first="Rudolf S. N." last="Fehrmann">Rudolf S. N. Fehrmann</name><affiliation><nlm:aff id="A23">University Medical Center Groningen, Department of Genetics, Groningen, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Floyd, James A B" sort="Floyd, James A B" uniqKey="Floyd J" first="James A. B." last="Floyd">James A. B. Floyd</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Florin, Tim" sort="Florin, Tim" uniqKey="Florin T" first="Tim" last="Florin">Tim Florin</name><affiliation><nlm:aff id="A24">Mater Health Services, Department of Gastroenterology, Brisbane, Australia</nlm:aff></affiliation></author><author><name sortKey="Franchimont, Denis" sort="Franchimont, Denis" uniqKey="Franchimont D" first="Denis" last="Franchimont">Denis Franchimont</name><affiliation><nlm:aff id="A25">Erasmus Hospital, Free University of Brussels, Department of Gastroenterology, Brussels, Belgium</nlm:aff></affiliation></author><author><name sortKey="Franke, Lude" sort="Franke, Lude" uniqKey="Franke L" first="Lude" last="Franke">Lude Franke</name><affiliation><nlm:aff id="A23">University Medical Center Groningen, Department of Genetics, Groningen, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Georges, Michel" sort="Georges, Michel" uniqKey="Georges M" first="Michel" last="Georges">Michel Georges</name><affiliation><nlm:aff id="A26">University of Liège, Department of Genetics, Faculty of Veterinary Medicine, Liège, Belgium</nlm:aff></affiliation></author><author><name sortKey="Glas, Jurgen" sort="Glas, Jurgen" uniqKey="Glas J" first="Jürgen" last="Glas">Jürgen Glas</name><affiliation><nlm:aff id="A16">University Hospital Munich, Department of Medicine II, Munich, Germany</nlm:aff></affiliation></author><author><name sortKey="Glazer, Nicole L" sort="Glazer, Nicole L" uniqKey="Glazer N" first="Nicole L." last="Glazer">Nicole L. Glazer</name><affiliation><nlm:aff id="A27">University of Washington, Cardiovascular Health Research Unit, Department of Internal Medicine, Seattle, Washington, USA</nlm:aff></affiliation></author><author><name sortKey="Guthery, Stephen L" sort="Guthery, Stephen L" uniqKey="Guthery S" first="Stephen L." last="Guthery">Stephen L. Guthery</name><affiliation><nlm:aff id="A28">University of Utah School of Medicine, Department of Pediatrics, Salt Lake City, Utah, USA</nlm:aff></affiliation></author><author><name sortKey="Haritunians, Talin" sort="Haritunians, Talin" uniqKey="Haritunians T" first="Talin" last="Haritunians">Talin Haritunians</name><affiliation><nlm:aff id="A29">Cedars-Sinai Medical Center, Medical Genetics Institute, Los Angeles, California, USA</nlm:aff></affiliation></author><author><name sortKey="Hayward, Nicholas K" sort="Hayward, Nicholas K" uniqKey="Hayward N" first="Nicholas K." last="Hayward">Nicholas K. Hayward</name><affiliation><nlm:aff id="A30">Queensland Institute of Medical Research, Genetic Epidemiology, Brisbane, Australia</nlm:aff></affiliation></author><author><name sortKey="Hugot, Jean Pierre" sort="Hugot, Jean Pierre" uniqKey="Hugot J" first="Jean-Pierre" last="Hugot">Jean-Pierre Hugot</name><affiliation><nlm:aff id="A31">Université Paris Diderot & INSERM & Hopital Robert Debre APHP, Gastroenterology, Paris, France</nlm:aff></affiliation></author><author><name sortKey="Jobin, Gilles" sort="Jobin, Gilles" uniqKey="Jobin G" first="Gilles" last="Jobin">Gilles Jobin</name><affiliation><nlm:aff id="A2">Université de Montréal, Medicine, Montréal, Québec, Canada</nlm:aff></affiliation><affiliation><nlm:aff id="A32">Hôpital Maisonneuve-Rosemont, Dept of Gastroenterology, Montréal, Québec, Canada</nlm:aff></affiliation></author><author><name sortKey="Laukens, Debby" sort="Laukens, Debby" uniqKey="Laukens D" first="Debby" last="Laukens">Debby Laukens</name><affiliation><nlm:aff id="A20">Ghent University Hospital, Department of Hepatology and Gastroenterology, Ghent, Belgium</nlm:aff></affiliation></author><author><name sortKey="Lawrance, Ian" sort="Lawrance, Ian" uniqKey="Lawrance I" first="Ian" last="Lawrance">Ian Lawrance</name><affiliation><nlm:aff id="A33">The University of Western Australia, School of Medicine and Pharmacology, Fremantle, Australia</nlm:aff></affiliation></author><author><name sortKey="Lemann, Marc" sort="Lemann, Marc" uniqKey="Lemann M" first="Marc" last="Lémann">Marc Lémann</name><affiliation><nlm:aff id="A34">Université Paris Diderot, GETAID group, Paris, France</nlm:aff></affiliation></author><author><name sortKey="Levine, Arie" sort="Levine, Arie" uniqKey="Levine A" first="Arie" last="Levine">Arie Levine</name><affiliation><nlm:aff id="A35">Tel Aviv University, Pediatric Gastroenterology Unit, Wolfson Medical Center and Sackler School of Medicine, Tel Aviv, Israel</nlm:aff></affiliation></author><author><name sortKey="Libioulle, Cecile" sort="Libioulle, Cecile" uniqKey="Libioulle C" first="Cecile" last="Libioulle">Cecile Libioulle</name><affiliation><nlm:aff id="A36">Centre Hospitalier Universitaire Université de Liège, Division of Gastroenterology, Liège, Belgium</nlm:aff></affiliation></author><author><name sortKey="Louis, Edouard" sort="Louis, Edouard" uniqKey="Louis E" first="Edouard" last="Louis">Edouard Louis</name><affiliation><nlm:aff id="A36">Centre Hospitalier Universitaire Université de Liège, Division of Gastroenterology, Liège, Belgium</nlm:aff></affiliation></author><author><name sortKey="Mcgovern, Dermot P" sort="Mcgovern, Dermot P" uniqKey="Mcgovern D" first="Dermot P." last="Mcgovern">Dermot P. Mcgovern</name><affiliation><nlm:aff id="A7">Cedars-Sinai Medical Center, Inflammatory Bowel and Immunobiology Research Institute, Los Angeles, California, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A29">Cedars-Sinai Medical Center, Medical Genetics Institute, Los Angeles, California, USA</nlm:aff></affiliation></author><author><name sortKey="Milla, Monica" sort="Milla, Monica" uniqKey="Milla M" first="Monica" last="Milla">Monica Milla</name><affiliation><nlm:aff id="A37">AOU Careggi, Unit of Gastroenterology SOD2, Florence, Italy</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="A30">Queensland Institute of Medical Research, Genetic Epidemiology, Brisbane, Australia</nlm:aff></affiliation></author><author><name sortKey="Morley, Katherine I" sort="Morley, Katherine I" uniqKey="Morley K" first="Katherine I." last="Morley">Katherine I. Morley</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Mowat, Craig" sort="Mowat, Craig" uniqKey="Mowat C" first="Craig" last="Mowat">Craig Mowat</name><affiliation><nlm:aff id="A38">Ninewells Hospital and Medical School, Dept of Medicine, Dundee, UK</nlm:aff></affiliation></author><author><name sortKey="Ng, Aylwin" sort="Ng, Aylwin" uniqKey="Ng A" first="Aylwin" last="Ng">Aylwin Ng</name><affiliation><nlm:aff id="A39">Massachusetts General Hospital, Harvard Medical School, Gastroenterology Unit Boston, Massachusetts, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A40">Center for Computational and Integrative Biology, Massachusetts General Hospital, Boston, Massachusetts, USA</nlm:aff></affiliation></author><author><name sortKey="Newman, William" sort="Newman, William" uniqKey="Newman W" first="William" last="Newman">William Newman</name><affiliation><nlm:aff id="A41">University of Manchester Department of Medical Genetics, Manchester, UK</nlm:aff></affiliation></author><author><name sortKey="Ophoff, Roel A" sort="Ophoff, Roel A" uniqKey="Ophoff R" first="Roel A" last="Ophoff">Roel A. Ophoff</name><affiliation><nlm:aff id="A42">University Medical Center Utrecht, Department of Medical Genetics, Utrecht, Netherlands</nlm:aff></affiliation></author><author><name sortKey="Papi, Laura" sort="Papi, Laura" uniqKey="Papi L" first="Laura" last="Papi">Laura Papi</name><affiliation><nlm:aff id="A43">University of Florence, Institute of Human Genetics, Florence, Italy</nlm:aff></affiliation></author><author><name sortKey="Palmieri, Orazio" sort="Palmieri, Orazio" uniqKey="Palmieri O" first="Orazio" last="Palmieri">Orazio Palmieri</name><affiliation><nlm:aff id="A10">IRCCS-CSS Hospital, Unit of Gastroenterology, San Giovanni Rotondo, Italy</nlm:aff></affiliation></author><author><name sortKey="Peyrin Biroulet, Laurent" sort="Peyrin Biroulet, Laurent" uniqKey="Peyrin Biroulet L" first="Laurent" last="Peyrin-Biroulet">Laurent Peyrin-Biroulet</name><affiliation><nlm:aff id="A44">University Hospital of Nancy, Department of Hepato-Gastroenterology, Vandoeuvre-lès-Nancy, France</nlm:aff></affiliation></author><author><name sortKey="Panes, Julian" sort="Panes, Julian" uniqKey="Panes J" first="Julián" last="Panés">Julián Panés</name><affiliation><nlm:aff id="A45">Hospital Clínic de Barcelona, IDIBAPS, CIBERehd, Department of Gastroenterology, Barcelona, Spain</nlm:aff></affiliation></author><author><name sortKey="Phillips, Anne" sort="Phillips, Anne" uniqKey="Phillips A" first="Anne" last="Phillips">Anne Phillips</name><affiliation><nlm:aff id="A38">Ninewells Hospital and Medical School, Dept of Medicine, Dundee, UK</nlm:aff></affiliation></author><author><name sortKey="Prescott, Natalie J" sort="Prescott, Natalie J" uniqKey="Prescott N" first="Natalie J." last="Prescott">Natalie J. Prescott</name><affiliation><nlm:aff id="A46">King’s College London School of Medicine, Guy’s Hospital, Department of Medical and Molecular Genetics, London, UK</nlm:aff></affiliation></author><author><name sortKey="Proctor, Deborah D" sort="Proctor, Deborah D" uniqKey="Proctor D" first="Deborah D." last="Proctor">Deborah D. Proctor</name><affiliation><nlm:aff id="A47">Yale University, Section of Digestive Diseases, Department of Medicine, New Haven, Connecticut, USA</nlm:aff></affiliation></author><author><name sortKey="Roberts, Rebecca" sort="Roberts, Rebecca" uniqKey="Roberts R" first="Rebecca" last="Roberts">Rebecca Roberts</name><affiliation><nlm:aff id="A14">University of Otago, Department of Medicine, Christchurch, New Zealand</nlm:aff></affiliation></author><author><name sortKey="Russell, Richard" sort="Russell, Richard" uniqKey="Russell R" first="Richard" last="Russell">Richard Russell</name><affiliation><nlm:aff id="A48">Royal Hospital for Sick Children, Paediatric Gastroenterology and Nutrition, Glasgow, UK</nlm:aff></affiliation></author><author><name sortKey="Rutgeerts, Paul" sort="Rutgeerts, Paul" uniqKey="Rutgeerts P" first="Paul" last="Rutgeerts">Paul Rutgeerts</name><affiliation><nlm:aff id="A49">University Hospital Gasthuisberg, Division of Gastroenterology, Leuven, Belgium</nlm:aff></affiliation></author><author><name sortKey="Sanderson, Jeremy" sort="Sanderson, Jeremy" uniqKey="Sanderson J" first="Jeremy" last="Sanderson">Jeremy Sanderson</name><affiliation><nlm:aff id="A50">Guy’s & St Thomas’ NHS Foundation Trust, St Thomas’ Hospital, Dept Gastroenterology, London, UK</nlm:aff></affiliation></author><author><name sortKey="Sans, Miquel" sort="Sans, Miquel" uniqKey="Sans M" first="Miquel" last="Sans">Miquel Sans</name><affiliation><nlm:aff id="A51">Hospital Clínic de Barcelona, IDIBAPS, CIBERehd, Department of Gastroenterology, Barcelona, Spain</nlm:aff></affiliation></author><author><name sortKey="Schumm, Philip" sort="Schumm, Philip" uniqKey="Schumm P" first="Philip" last="Schumm">Philip Schumm</name><affiliation><nlm:aff id="A52">University of Chicago, Department of Health Studies, Chicago, Illinois, USA</nlm:aff></affiliation></author><author><name sortKey="Seibold, Frank" sort="Seibold, Frank" uniqKey="Seibold F" first="Frank" last="Seibold">Frank Seibold</name><affiliation><nlm:aff id="A53">University of Bern, Division of Gastroenterology, Inselspital, Bern, Switzerland</nlm:aff></affiliation></author><author><name sortKey="Sharma, Yashoda" sort="Sharma, Yashoda" uniqKey="Sharma Y" first="Yashoda" last="Sharma">Yashoda Sharma</name><affiliation><nlm:aff id="A47">Yale University, Section of Digestive Diseases, Department of Medicine, New Haven, Connecticut, USA</nlm:aff></affiliation></author><author><name sortKey="Simms, Lisa" sort="Simms, Lisa" uniqKey="Simms L" first="Lisa" last="Simms">Lisa Simms</name><affiliation><nlm:aff id="A30">Queensland Institute of Medical Research, Genetic Epidemiology, Brisbane, Australia</nlm:aff></affiliation></author><author><name sortKey="Seielstad, Mark" sort="Seielstad, Mark" uniqKey="Seielstad M" first="Mark" last="Seielstad">Mark Seielstad</name><affiliation><nlm:aff id="A54">Genome Institute of Singapore, Human Genetics, Singapore</nlm:aff></affiliation><affiliation><nlm:aff id="A55">Institute for Human Genetics, University of California San Fransisco, San Francisco, California, USA</nlm:aff></affiliation></author><author><name sortKey="Steinhart, A Hillary" sort="Steinhart, A Hillary" uniqKey="Steinhart A" first="A. Hillary" last="Steinhart">A. Hillary Steinhart</name><affiliation><nlm:aff id="A56">University of Toronto, Mount Sinai Hospital Inflammatory Bowel Disease Centre, Toronto, Ontario, Canada</nlm:aff></affiliation></author><author><name sortKey="Targan, Stephan R" sort="Targan, Stephan R" uniqKey="Targan S" first="Stephan R." last="Targan">Stephan R. Targan</name><affiliation><nlm:aff id="A7">Cedars-Sinai Medical Center, Inflammatory Bowel and Immunobiology Research Institute, Los Angeles, California, USA</nlm:aff></affiliation></author><author><name sortKey="Van Den Berg, Leonard H" sort="Van Den Berg, Leonard H" uniqKey="Van Den Berg L" first="Leonard H." last="Van Den Berg">Leonard H. Van Den Berg</name><affiliation><nlm:aff id="A57">Rudolf Magnus Institute of Neuroscience, University Medical Center Utrecht, Department of Neurology, Utrecht, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Vatn, Morten" sort="Vatn, Morten" uniqKey="Vatn M" first="Morten" last="Vatn">Morten Vatn</name><affiliation><nlm:aff id="A58">Rikshospitalet University Hospital, Medical Department, Oslo, Norway</nlm:aff></affiliation></author><author><name sortKey="Verspaget, Hein" sort="Verspaget, Hein" uniqKey="Verspaget H" first="Hein" last="Verspaget">Hein Verspaget</name><affiliation><nlm:aff id="A59">Leiden University Medical Center, Experimental Gastroenterology, Leiden, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Walters, Thomas" sort="Walters, Thomas" uniqKey="Walters T" first="Thomas" last="Walters">Thomas Walters</name><affiliation><nlm:aff id="A60">The Hospital for Sick Children, University of Toronto, Toronto, Ontario, Canada</nlm:aff></affiliation></author><author><name sortKey="Wijmenga, Cisca" sort="Wijmenga, Cisca" uniqKey="Wijmenga C" first="Cisca" last="Wijmenga">Cisca Wijmenga</name><affiliation><nlm:aff id="A23">University Medical Center Groningen, Department of Genetics, Groningen, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Wilson, David C" sort="Wilson, David C" uniqKey="Wilson D" first="David C." last="Wilson">David C. Wilson</name><affiliation><nlm:aff id="A48">Royal Hospital for Sick Children, Paediatric Gastroenterology and Nutrition, Glasgow, UK</nlm:aff></affiliation><affiliation><nlm:aff id="A61">Child Life and Health, University of Edinburgh, Scotland</nlm:aff></affiliation></author><author><name sortKey="Westra, Harm Jan" sort="Westra, Harm Jan" uniqKey="Westra H" first="Harm-Jan" last="Westra">Harm-Jan Westra</name><affiliation><nlm:aff id="A23">University Medical Center Groningen, Department of Genetics, Groningen, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Xavier, Ramnik J" sort="Xavier, Ramnik J" uniqKey="Xavier R" first="Ramnik J." last="Xavier">Ramnik J. Xavier</name><affiliation><nlm:aff id="A39">Massachusetts General Hospital, Harvard Medical School, Gastroenterology Unit Boston, Massachusetts, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A40">Center for Computational and Integrative Biology, Massachusetts General Hospital, Boston, Massachusetts, USA</nlm:aff></affiliation></author><author><name sortKey="Zhao, Zhen Z" sort="Zhao, Zhen Z" uniqKey="Zhao Z" first="Zhen Z." last="Zhao">Zhen Z. Zhao</name><affiliation><nlm:aff id="A30">Queensland Institute of Medical Research, Genetic Epidemiology, Brisbane, Australia</nlm:aff></affiliation></author><author><name sortKey="Ponsioen, Cyriel Y" sort="Ponsioen, Cyriel Y" uniqKey="Ponsioen C" first="Cyriel Y." last="Ponsioen">Cyriel Y. Ponsioen</name><affiliation><nlm:aff id="A62">Academic Medical Center, Department of Gastroenterology, Amsterdam, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Andersen, Vibeke" sort="Andersen, Vibeke" uniqKey="Andersen V" first="Vibeke" last="Andersen">Vibeke Andersen</name><affiliation><nlm:aff id="A63">Viborg Regional Hospital, Medical Department, Viborg, Denmark</nlm:aff></affiliation></author><author><name sortKey="Torkvist, Leif" sort="Torkvist, Leif" uniqKey="Torkvist L" first="Leif" last="Torkvist">Leif Torkvist</name><affiliation><nlm:aff id="A64">Karolinska Institutet, Department of Clinical Science Intervention and Technology, Stockholm, Sweden</nlm:aff></affiliation></author><author><name sortKey="Gazouli, Maria" sort="Gazouli, Maria" uniqKey="Gazouli M" first="Maria" last="Gazouli">Maria Gazouli</name><affiliation><nlm:aff id="A65">University of Athens, Department of Biology, School of Medicine, Athens, Greece</nlm:aff></affiliation></author><author><name sortKey="Anagnou, Nicholas P" sort="Anagnou, Nicholas P" uniqKey="Anagnou N" first="Nicholas P." last="Anagnou">Nicholas P. Anagnou</name><affiliation><nlm:aff id="A65">University of Athens, Department of Biology, School of Medicine, Athens, Greece</nlm:aff></affiliation></author><author><name sortKey="Karlsen, Tom H" sort="Karlsen, Tom H" uniqKey="Karlsen T" first="Tom H." last="Karlsen">Tom H. Karlsen</name><affiliation><nlm:aff id="A58">Rikshospitalet University Hospital, Medical Department, Oslo, Norway</nlm:aff></affiliation></author><author><name sortKey="Kupcinskas, Limas" sort="Kupcinskas, Limas" uniqKey="Kupcinskas L" first="Limas" last="Kupcinskas">Limas Kupcinskas</name><affiliation><nlm:aff id="A66">Kaunas University of medicine, Department of Gastroenterology, Kaunas, Lithuania</nlm:aff></affiliation></author><author><name sortKey="Sventoraityte, Jurgita" sort="Sventoraityte, Jurgita" uniqKey="Sventoraityte J" first="Jurgita" last="Sventoraityte">Jurgita Sventoraityte</name><affiliation><nlm:aff id="A66">Kaunas University of medicine, Department of Gastroenterology, Kaunas, Lithuania</nlm:aff></affiliation></author><author><name sortKey="Mansfield, John C" sort="Mansfield, John C" uniqKey="Mansfield J" first="John C." last="Mansfield">John C. Mansfield</name><affiliation><nlm:aff id="A67">Newcastle University, Institute of Human Genetics, Newcastle upon Tyne, UK</nlm:aff></affiliation></author><author><name sortKey="Kugathasan, Subra" sort="Kugathasan, Subra" uniqKey="Kugathasan S" first="Subra" last="Kugathasan">Subra Kugathasan</name><affiliation><nlm:aff id="A68">Emory School of Medicine, Department of Genetics and Department of Pediatrics, Atlanta, Georgia, USA</nlm:aff></affiliation></author><author><name sortKey="Silverberg, Mark S" sort="Silverberg, Mark S" uniqKey="Silverberg M" first="Mark S." last="Silverberg">Mark S. Silverberg</name><affiliation><nlm:aff id="A56">University of Toronto, Mount Sinai Hospital Inflammatory Bowel Disease Centre, Toronto, Ontario, Canada</nlm:aff></affiliation></author><author><name sortKey="Halfvarson, Jonas" sort="Halfvarson, Jonas" uniqKey="Halfvarson J" first="Jonas" last="Halfvarson">Jonas Halfvarson</name><affiliation><nlm:aff id="A69">Örebro University Hospital, Department of Medicine, Örebro, Sweden</nlm:aff></affiliation></author><author><name sortKey="Rotter, Jerome I" sort="Rotter, Jerome I" uniqKey="Rotter J" first="Jerome I." last="Rotter">Jerome I. Rotter</name><affiliation><nlm:aff id="A29">Cedars-Sinai Medical Center, Medical Genetics Institute, Los Angeles, California, USA</nlm:aff></affiliation></author><author><name sortKey="Mathew, Christopher G" sort="Mathew, Christopher G" uniqKey="Mathew C" first="Christopher G." last="Mathew">Christopher G. Mathew</name><affiliation><nlm:aff id="A46">King’s College London School of Medicine, Guy’s Hospital, Department of Medical and Molecular Genetics, London, UK</nlm:aff></affiliation></author><author><name sortKey="Griffiths, Anne M" sort="Griffiths, Anne M" uniqKey="Griffiths A" first="Anne M." last="Griffiths">Anne M. Griffiths</name><affiliation><nlm:aff id="A60">The Hospital for Sick Children, University of Toronto, Toronto, Ontario, Canada</nlm:aff></affiliation></author><author><name sortKey="Gearry, Richard" sort="Gearry, Richard" uniqKey="Gearry R" first="Richard" last="Gearry">Richard Gearry</name><affiliation><nlm:aff id="A14">University of Otago, Department of Medicine, Christchurch, New Zealand</nlm:aff></affiliation></author><author><name sortKey="Ahmad, Tariq" sort="Ahmad, Tariq" uniqKey="Ahmad T" first="Tariq" last="Ahmad">Tariq Ahmad</name><affiliation><nlm:aff id="A70">Peninsula College of Medicine and Dentistry, Barrack Road, Exeter, UK</nlm:aff></affiliation></author><author><name sortKey="Brant, Steven R" sort="Brant, Steven R" uniqKey="Brant S" first="Steven R." last="Brant">Steven R. Brant</name><affiliation><nlm:aff id="A15">Johns Hopkins University School of Medicine, Meyeroff Inflammatory Bowel Disease Center, Dept. of Medicine, Baltimore, Maryland, USA</nlm:aff></affiliation></author><author><name sortKey="Chamaillard, Mathias" sort="Chamaillard, Mathias" uniqKey="Chamaillard M" first="Mathias" last="Chamaillard">Mathias Chamaillard</name><affiliation><nlm:aff id="A71">Inserm, U1019, Lille, France</nlm:aff></affiliation></author><author><name sortKey="Satsangi, Jack" sort="Satsangi, Jack" uniqKey="Satsangi J" first="Jack" last="Satsangi">Jack Satsangi</name><affiliation><nlm:aff id="A4">University of Edinburgh, Western General Hospital, Gastrointestinal Unit, Molecular Medicine Centre, Edinburgh, UK</nlm:aff></affiliation></author><author><name sortKey="Cho, Judy H" sort="Cho, Judy H" uniqKey="Cho J" first="Judy H." last="Cho">Judy H. Cho</name><affiliation><nlm:aff id="A47">Yale University, Section of Digestive Diseases, Department of Medicine, New Haven, Connecticut, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A72">Yale University, Department of Genetics, Yale School of Medicine, New Haven CT</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="A5">Christian-Albrechts-University Kiel, Institute of Clinical Molecular Biology, Kiel, Germany</nlm:aff></affiliation><affiliation><nlm:aff id="A73">Department for General Internal Medicine, Christian-Albrechts-University, Kiel, Germany</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="A74">Massachusetts General Hospital, Harvard Medical School, Center for Human Genetic Research, Boston, Massachusetts, USA</nlm:aff></affiliation></author><author><name sortKey="Barrett, Jeffrey C" sort="Barrett, Jeffrey C" uniqKey="Barrett J" first="Jeffrey C." last="Barrett">Jeffrey C. Barrett</name><affiliation><nlm:aff id="A1">Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Parkes, Miles" sort="Parkes, Miles" uniqKey="Parkes M" first="Miles" last="Parkes">Miles Parkes</name><affiliation><nlm:aff id="A8">Addenbrooke’s Hospital, University of Cambridge, Gastroenterology Research Unit, Cambridge, UK</nlm:aff></affiliation></author><author><name sortKey="Annese, Vito" sort="Annese, Vito" uniqKey="Annese V" first="Vito" last="Annese">Vito Annese</name><affiliation><nlm:aff id="A10">IRCCS-CSS Hospital, Unit of Gastroenterology, San Giovanni Rotondo, Italy</nlm:aff></affiliation><affiliation><nlm:aff id="A37">AOU Careggi, Unit of Gastroenterology SOD2, Florence, Italy</nlm:aff></affiliation></author><author><name sortKey="Hakonarson, Hakon" sort="Hakonarson, Hakon" uniqKey="Hakonarson H" first="Hakon" last="Hakonarson">Hakon Hakonarson</name><affiliation><nlm:aff id="A13">The Children’s Hospital of Philadelphia, Department of Pediatrics, Center for Pediatric Inflammatory Bowel Disease, Philadelphia, Pennsylvania, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A75">Center for Applied Genomics, The Children’s Hospital of Philadelphia, Philadelphia, Pennsylvania, USA</nlm:aff></affiliation></author><author><name sortKey="Radford Smith, Graham" sort="Radford Smith, Graham" uniqKey="Radford Smith G" first="Graham" last="Radford-Smith">Graham Radford-Smith</name><affiliation><nlm:aff id="A76">Queensland Institute of Medical Research, IBD Research Group, Brisbane, Australia</nlm:aff></affiliation></author><author><name sortKey="Duerr, Richard H" sort="Duerr, Richard H" uniqKey="Duerr R" first="Richard H." last="Duerr">Richard H. Duerr</name><affiliation><nlm:aff id="A11">University of Pittsburgh School of Medicine, Division of Gastroenterology, Hepatology and Nutrition, Department of Medicine, Pittsburgh, Pennsylvania, USA</nlm:aff></affiliation><affiliation><nlm:aff id="A77">University of Pittsburgh Graduate School of Public Health, 130 Desoto Street, Pittsburgh, PA, USA</nlm:aff></affiliation></author><author><name sortKey="Vermeire, Severine" sort="Vermeire, Severine" uniqKey="Vermeire S" first="Séverine" last="Vermeire">Séverine Vermeire</name><affiliation><nlm:aff id="A49">University Hospital Gasthuisberg, Division of Gastroenterology, Leuven, Belgium</nlm:aff></affiliation></author><author><name sortKey="Weersma, Rinse K" sort="Weersma, Rinse K" uniqKey="Weersma R" first="Rinse K." last="Weersma">Rinse K. Weersma</name><affiliation><nlm:aff id="A78">University Medical Center Groningen, Department of Gastroenterology, Groningen, the Netherlands</nlm:aff></affiliation></author><author><name sortKey="Rioux, John D" sort="Rioux, John D" uniqKey="Rioux J" first="John D." last="Rioux">John D. Rioux</name><affiliation><nlm:aff id="A2">Université de Montréal, Medicine, Montréal, Québec, Canada</nlm:aff></affiliation><affiliation><nlm:aff id="A3">Montreal Heart Institute, Research Center, Montréal, Québec, Canada</nlm:aff></affiliation></author></analytic><series><title level="j">Nature genetics</title><idno type="ISSN">1061-4036</idno><idno type="eISSN">1546-1718</idno><imprint><date when="2011">2011</date></imprint></series></biblStruct></sourceDesc></fileDesc><profileDesc><textClass></textClass></profileDesc></teiHeader><front><div type="abstract" xml:lang="en"><p id="P1">Genome-wide association studies (GWAS) and candidate gene studies in ulcerative colitis (UC) have identified 18 susceptibility loci. We conducted a meta-analysis of 6 UC GWAS, comprising 6,687 cases and 19,718 controls, and followed-up the top association signals in 9,628 cases and 12,917 controls. We identified 29 additional risk loci (P<5×10<sup>-8</sup>), increasing the number of UC associated loci to 47. After annotating associated regions using GRAIL, eQTL data and correlations with non-synonymous SNPs, we identified many candidate genes providing potentially important insights into disease pathogenesis, including <italic>IL1R2, IL8RA/B, IL7R, IL12B, DAP, PRDM1, JAK2, IRF5, GNA12</italic> and <italic>LSP1</italic>. The total number of confirmed inflammatory bowel disease (IBD) risk loci is now 99, including a minimum of 28 shared association signals between Crohn’s disease (CD) and UC.</p></div></front><back><div1 type="bibliography"><listBibl><biblStruct><analytic><author><name sortKey="Xavier, Rj" uniqKey="Xavier R">RJ Xavier</name></author><author><name sortKey="Podolsky, Dk" uniqKey="Podolsky D">DK Podolsky</name></author></analytic></biblStruct><biblStruct></biblStruct><biblStruct><analytic><author><name sortKey="Festen, Ea" uniqKey="Festen E">EA Festen</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Franke, A" uniqKey="Franke A">A Franke</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Mcgovern, Dp" uniqKey="Mcgovern D">DP McGovern</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Franke, A" uniqKey="Franke A">A Franke</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Dideberg, V" uniqKey="Dideberg V">V Dideberg</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Asano, K" uniqKey="Asano K">K Asano</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Zhernakova, A" uniqKey="Zhernakova A">A Zhernakova</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Kugathasan, S" uniqKey="Kugathasan S">S Kugathasan</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Dubois, Pc" uniqKey="Dubois P">PC Dubois</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Kuester, D" uniqKey="Kuester D">D Kuester</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Koren, I" uniqKey="Koren I">I Koren</name></author><author><name sortKey="Reem, E" uniqKey="Reem E">E Reem</name></author><author><name sortKey="Kimchi, A" uniqKey="Kimchi A">A Kimchi</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Franke, A" uniqKey="Franke A">A Franke</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Turner, Ca" uniqKey="Turner C">CA Turner</name></author><author><name sortKey="Mack, Dh" uniqKey="Mack D">DH Mack</name></author><author><name sortKey="Davis, Mm" uniqKey="Davis M">MM Davis</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Martins, Ga" uniqKey="Martins G">GA Martins</name></author><author><name sortKey="Cimmino, L" uniqKey="Cimmino L">L Cimmino</name></author><author><name sortKey="Liao, J" uniqKey="Liao J">J Liao</name></author><author><name sortKey="Magnusdottir, E" uniqKey="Magnusdottir E">E Magnusdottir</name></author><author><name sortKey="Calame, K" uniqKey="Calame K">K Calame</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Kallies, A" uniqKey="Kallies A">A Kallies</name></author><author><name sortKey="Xin, A" uniqKey="Xin A">A Xin</name></author><author><name sortKey="Belz, Gt" uniqKey="Belz G">GT Belz</name></author><author><name sortKey="Nutt, Sl" uniqKey="Nutt S">SL Nutt</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Grenningloh, R" uniqKey="Grenningloh R">R Grenningloh</name></author><author><name sortKey="Kang, By" uniqKey="Kang B">BY Kang</name></author><author><name sortKey="Ho, Ic" uniqKey="Ho I">IC Ho</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Moisan, J" uniqKey="Moisan J">J Moisan</name></author><author><name sortKey="Grenningloh, R" uniqKey="Grenningloh R">R Grenningloh</name></author><author><name sortKey="Bettelli, E" uniqKey="Bettelli E">E Bettelli</name></author><author><name sortKey="Oukka, M" uniqKey="Oukka M">M Oukka</name></author><author><name sortKey="Ho, Ic" uniqKey="Ho I">IC Ho</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Sabath, E" uniqKey="Sabath E">E Sabath</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Bettelli, E" uniqKey="Bettelli E">E Bettelli</name></author><author><name sortKey="Korn, T" uniqKey="Korn T">T Korn</name></author><author><name sortKey="Oukka, M" uniqKey="Oukka M">M Oukka</name></author><author><name sortKey="Kuchroo, Vk" uniqKey="Kuchroo V">VK Kuchroo</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Steinberg, Mw" uniqKey="Steinberg M">MW Steinberg</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Maerten, P" uniqKey="Maerten P">P Maerten</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Mahida, Yr" uniqKey="Mahida Y">YR Mahida</name></author><author><name sortKey="Wu, K" uniqKey="Wu K">K Wu</name></author><author><name sortKey="Jewell, Dp" uniqKey="Jewell D">DP Jewell</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Williams, Ej" uniqKey="Williams E">EJ Williams</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Noble, Cl" uniqKey="Noble C">CL Noble</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Schluns, Ks" uniqKey="Schluns K">KS Schluns</name></author><author><name sortKey="Kieper, Wc" uniqKey="Kieper W">WC Kieper</name></author><author><name sortKey="Jameson, Sc" uniqKey="Jameson S">SC Jameson</name></author><author><name sortKey="Lefrancois, L" uniqKey="Lefrancois L">L Lefrancois</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Yamazaki, M" uniqKey="Yamazaki M">M Yamazaki</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Gregory, Sg" uniqKey="Gregory S">SG Gregory</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Fumagalli, M" uniqKey="Fumagalli M">M Fumagalli</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Pandey, Ak" uniqKey="Pandey A">AK Pandey</name></author></analytic></biblStruct><biblStruct><analytic><author><name sortKey="Liu, L" uniqKey="Liu L">L Liu</name></author></analytic></biblStruct></listBibl></div1></back></TEI><pmc article-type="research-article"><pmc-dir>properties open_access</pmc-dir>
<pmc-dir>properties manuscript</pmc-dir>
<front><journal-meta><journal-id journal-id-type="nlm-journal-id">9216904</journal-id><journal-id journal-id-type="pubmed-jr-id">2419</journal-id><journal-id journal-id-type="nlm-ta">Nat Genet</journal-id><journal-id journal-id-type="iso-abbrev">Nat. Genet.</journal-id><journal-title-group><journal-title>Nature genetics</journal-title></journal-title-group><issn pub-type="ppub">1061-4036</issn><issn pub-type="epub">1546-1718</issn></journal-meta><article-meta><article-id pub-id-type="pmid">21297633</article-id><article-id pub-id-type="pmc">3084597</article-id><article-id pub-id-type="doi">10.1038/ng.764</article-id><article-id pub-id-type="manuscript">NIHMS265993</article-id><article-categories><subj-group subj-group-type="heading"><subject>Article</subject></subj-group></article-categories><title-group><article-title>Meta-analysis identifies 29 additional ulcerative colitis risk loci, increasing the number of confirmed associations to 47</article-title></title-group><contrib-group><contrib contrib-type="author"><name><surname>Anderson</surname><given-names>Carl A.</given-names></name><xref ref-type="aff" rid="A1">1</xref></contrib><contrib contrib-type="author"><name><surname>Boucher</surname><given-names>Gabrielle</given-names></name><xref ref-type="aff" rid="A2">2</xref><xref ref-type="aff" rid="A3">3</xref><xref rid="FN2" ref-type="author-notes">79</xref></contrib><contrib contrib-type="author"><name><surname>Lees</surname><given-names>Charlie W.</given-names></name><xref ref-type="aff" rid="A4">4</xref><xref rid="FN2" ref-type="author-notes">79</xref></contrib><contrib contrib-type="author"><name><surname>Franke</surname><given-names>Andre</given-names></name><xref ref-type="aff" rid="A5">5</xref><xref rid="FN2" ref-type="author-notes">79</xref></contrib><contrib contrib-type="author"><name><surname>D’Amato</surname><given-names>Mauro</given-names></name><xref ref-type="aff" rid="A6">6</xref><xref rid="FN2" ref-type="author-notes">79</xref></contrib><contrib contrib-type="author"><name><surname>Taylor</surname><given-names>Kent D.</given-names></name><xref ref-type="aff" rid="A7">7</xref></contrib><contrib contrib-type="author"><name><surname>Lee</surname><given-names>James C.</given-names></name><xref ref-type="aff" rid="A8">8</xref></contrib><contrib contrib-type="author"><name><surname>Goyette</surname><given-names>Philippe</given-names></name><xref ref-type="aff" rid="A2">2</xref><xref ref-type="aff" rid="A3">3</xref></contrib><contrib contrib-type="author"><name><surname>Imielinski</surname><given-names>Marcin</given-names></name><xref ref-type="aff" rid="A9">9</xref></contrib><contrib contrib-type="author"><name><surname>Latiano</surname><given-names>Anna</given-names></name><xref ref-type="aff" rid="A10">10</xref></contrib><contrib contrib-type="author"><name><surname>Lagacé</surname><given-names>Caroline</given-names></name><xref ref-type="aff" rid="A2">2</xref><xref ref-type="aff" rid="A3">3</xref></contrib><contrib contrib-type="author"><name><surname>Scott</surname><given-names>Regan</given-names></name><xref ref-type="aff" rid="A11">11</xref></contrib><contrib contrib-type="author"><name><surname>Amininejad</surname><given-names>Leila</given-names></name><xref ref-type="aff" rid="A12">12</xref></contrib><contrib contrib-type="author"><name><surname>Bumpstead</surname><given-names>Suzannah</given-names></name><xref ref-type="aff" rid="A1">1</xref></contrib><contrib contrib-type="author"><name><surname>Baidoo</surname><given-names>Leonard</given-names></name><xref ref-type="aff" rid="A11">11</xref></contrib><contrib contrib-type="author"><name><surname>Baldassano</surname><given-names>Robert N.</given-names></name><xref ref-type="aff" rid="A13">13</xref></contrib><contrib contrib-type="author"><name><surname>Barclay</surname><given-names>Murray</given-names></name><xref ref-type="aff" rid="A14">14</xref></contrib><contrib contrib-type="author"><name><surname>Bayless</surname><given-names>Theodore M.</given-names></name><xref ref-type="aff" rid="A15">15</xref></contrib><contrib contrib-type="author"><name><surname>Brand</surname><given-names>Stephan</given-names></name><xref ref-type="aff" rid="A16">16</xref></contrib><contrib contrib-type="author"><name><surname>Büning</surname><given-names>Carsten</given-names></name><xref ref-type="aff" rid="A17">17</xref></contrib><contrib contrib-type="author"><name><surname>Colombel</surname><given-names>Jean-Frédéric</given-names></name><xref ref-type="aff" rid="A18">18</xref></contrib><contrib contrib-type="author"><name><surname>Denson</surname><given-names>Lee A.</given-names></name><xref ref-type="aff" rid="A19">19</xref></contrib><contrib contrib-type="author"><name><surname>De Vos</surname><given-names>Martine</given-names></name><xref ref-type="aff" rid="A20">20</xref></contrib><contrib contrib-type="author"><name><surname>Dubinsky</surname><given-names>Marla</given-names></name><xref ref-type="aff" rid="A21">21</xref></contrib><contrib contrib-type="author"><name><surname>Edwards</surname><given-names>Cathryn</given-names></name><xref ref-type="aff" rid="A22">22</xref></contrib><contrib contrib-type="author"><name><surname>Ellinghaus</surname><given-names>David</given-names></name><xref ref-type="aff" rid="A5">5</xref></contrib><contrib contrib-type="author"><name><surname>Fehrmann</surname><given-names>Rudolf S.N.</given-names></name><xref ref-type="aff" rid="A23">23</xref></contrib><contrib contrib-type="author"><name><surname>Floyd</surname><given-names>James A.B.</given-names></name><xref ref-type="aff" rid="A1">1</xref></contrib><contrib contrib-type="author"><name><surname>Florin</surname><given-names>Tim</given-names></name><xref ref-type="aff" rid="A24">24</xref></contrib><contrib contrib-type="author"><name><surname>Franchimont</surname><given-names>Denis</given-names></name><xref ref-type="aff" rid="A25">25</xref></contrib><contrib contrib-type="author"><name><surname>Franke</surname><given-names>Lude</given-names></name><xref ref-type="aff" rid="A23">23</xref></contrib><contrib contrib-type="author"><name><surname>Georges</surname><given-names>Michel</given-names></name><xref ref-type="aff" rid="A26">26</xref></contrib><contrib contrib-type="author"><name><surname>Glas</surname><given-names>Jürgen</given-names></name><xref ref-type="aff" rid="A16">16</xref></contrib><contrib contrib-type="author"><name><surname>Glazer</surname><given-names>Nicole L.</given-names></name><xref ref-type="aff" rid="A27">27</xref></contrib><contrib contrib-type="author"><name><surname>Guthery</surname><given-names>Stephen L.</given-names></name><xref ref-type="aff" rid="A28">28</xref></contrib><contrib contrib-type="author"><name><surname>Haritunians</surname><given-names>Talin</given-names></name><xref ref-type="aff" rid="A29">29</xref></contrib><contrib contrib-type="author"><name><surname>Hayward</surname><given-names>Nicholas K.</given-names></name><xref ref-type="aff" rid="A30">30</xref></contrib><contrib contrib-type="author"><name><surname>Hugot</surname><given-names>Jean-Pierre</given-names></name><xref ref-type="aff" rid="A31">31</xref></contrib><contrib contrib-type="author"><name><surname>Jobin</surname><given-names>Gilles</given-names></name><xref ref-type="aff" rid="A2">2</xref><xref ref-type="aff" rid="A32">32</xref></contrib><contrib contrib-type="author"><name><surname>Laukens</surname><given-names>Debby</given-names></name><xref ref-type="aff" rid="A20">20</xref></contrib><contrib contrib-type="author"><name><surname>Lawrance</surname><given-names>Ian</given-names></name><xref ref-type="aff" rid="A33">33</xref></contrib><contrib contrib-type="author"><name><surname>Lémann</surname><given-names>Marc</given-names></name><xref ref-type="aff" rid="A34">34</xref></contrib><contrib contrib-type="author"><name><surname>Levine</surname><given-names>Arie</given-names></name><xref ref-type="aff" rid="A35">35</xref></contrib><contrib contrib-type="author"><name><surname>Libioulle</surname><given-names>Cecile</given-names></name><xref ref-type="aff" rid="A36">36</xref></contrib><contrib contrib-type="author"><name><surname>Louis</surname><given-names>Edouard</given-names></name><xref ref-type="aff" rid="A36">36</xref></contrib><contrib contrib-type="author"><name><surname>McGovern</surname><given-names>Dermot P.</given-names></name><xref ref-type="aff" rid="A7">7</xref><xref ref-type="aff" rid="A29">29</xref></contrib><contrib contrib-type="author"><name><surname>Milla</surname><given-names>Monica</given-names></name><xref ref-type="aff" rid="A37">37</xref></contrib><contrib contrib-type="author"><name><surname>Montgomery</surname><given-names>Grant W.</given-names></name><xref ref-type="aff" rid="A30">30</xref></contrib><contrib contrib-type="author"><name><surname>Morley</surname><given-names>Katherine I.</given-names></name><xref ref-type="aff" rid="A1">1</xref></contrib><contrib contrib-type="author"><name><surname>Mowat</surname><given-names>Craig</given-names></name><xref ref-type="aff" rid="A38">38</xref></contrib><contrib contrib-type="author"><name><surname>Ng</surname><given-names>Aylwin</given-names></name><xref ref-type="aff" rid="A39">39</xref><xref ref-type="aff" rid="A40">40</xref></contrib><contrib contrib-type="author"><name><surname>Newman</surname><given-names>William</given-names></name><xref ref-type="aff" rid="A41">41</xref></contrib><contrib contrib-type="author"><name><surname>Ophoff</surname><given-names>Roel A</given-names></name><xref ref-type="aff" rid="A42">42</xref></contrib><contrib contrib-type="author"><name><surname>Papi</surname><given-names>Laura</given-names></name><xref ref-type="aff" rid="A43">43</xref></contrib><contrib contrib-type="author"><name><surname>Palmieri</surname><given-names>Orazio</given-names></name><xref ref-type="aff" rid="A10">10</xref></contrib><contrib contrib-type="author"><name><surname>Peyrin-Biroulet</surname><given-names>Laurent</given-names></name><xref ref-type="aff" rid="A44">44</xref></contrib><contrib contrib-type="author"><name><surname>Panés</surname><given-names>Julián</given-names></name><xref ref-type="aff" rid="A45">45</xref></contrib><contrib contrib-type="author"><name><surname>Phillips</surname><given-names>Anne</given-names></name><xref ref-type="aff" rid="A38">38</xref></contrib><contrib contrib-type="author"><name><surname>Prescott</surname><given-names>Natalie J.</given-names></name><xref ref-type="aff" rid="A46">46</xref></contrib><contrib contrib-type="author"><name><surname>Proctor</surname><given-names>Deborah D.</given-names></name><xref ref-type="aff" rid="A47">47</xref></contrib><contrib contrib-type="author"><name><surname>Roberts</surname><given-names>Rebecca</given-names></name><xref ref-type="aff" rid="A14">14</xref></contrib><contrib contrib-type="author"><name><surname>Russell</surname><given-names>Richard</given-names></name><xref ref-type="aff" rid="A48">48</xref></contrib><contrib contrib-type="author"><name><surname>Rutgeerts</surname><given-names>Paul</given-names></name><xref ref-type="aff" rid="A49">49</xref></contrib><contrib contrib-type="author"><name><surname>Sanderson</surname><given-names>Jeremy</given-names></name><xref ref-type="aff" rid="A50">50</xref></contrib><contrib contrib-type="author"><name><surname>Sans</surname><given-names>Miquel</given-names></name><xref ref-type="aff" rid="A51">51</xref></contrib><contrib contrib-type="author"><name><surname>Schumm</surname><given-names>Philip</given-names></name><xref ref-type="aff" rid="A52">52</xref></contrib><contrib contrib-type="author"><name><surname>Seibold</surname><given-names>Frank</given-names></name><xref ref-type="aff" rid="A53">53</xref></contrib><contrib contrib-type="author"><name><surname>Sharma</surname><given-names>Yashoda</given-names></name><xref ref-type="aff" rid="A47">47</xref></contrib><contrib contrib-type="author"><name><surname>Simms</surname><given-names>Lisa</given-names></name><xref ref-type="aff" rid="A30">30</xref></contrib><contrib contrib-type="author"><name><surname>Seielstad</surname><given-names>Mark</given-names></name><xref ref-type="aff" rid="A54">54</xref><xref ref-type="aff" rid="A55">55</xref></contrib><contrib contrib-type="author"><name><surname>Steinhart</surname><given-names>A. Hillary</given-names></name><xref ref-type="aff" rid="A56">56</xref></contrib><contrib contrib-type="author"><name><surname>Targan</surname><given-names>Stephan R.</given-names></name><xref ref-type="aff" rid="A7">7</xref></contrib><contrib contrib-type="author"><name><surname>van den Berg</surname><given-names>Leonard H.</given-names></name><xref ref-type="aff" rid="A57">57</xref></contrib><contrib contrib-type="author"><name><surname>Vatn</surname><given-names>Morten</given-names></name><xref ref-type="aff" rid="A58">58</xref></contrib><contrib contrib-type="author"><name><surname>Verspaget</surname><given-names>Hein</given-names></name><xref ref-type="aff" rid="A59">59</xref></contrib><contrib contrib-type="author"><name><surname>Walters</surname><given-names>Thomas</given-names></name><xref ref-type="aff" rid="A60">60</xref></contrib><contrib contrib-type="author"><name><surname>Wijmenga</surname><given-names>Cisca</given-names></name><xref ref-type="aff" rid="A23">23</xref></contrib><contrib contrib-type="author"><name><surname>Wilson</surname><given-names>David C.</given-names></name><xref ref-type="aff" rid="A48">48</xref><xref ref-type="aff" rid="A61">61</xref></contrib><contrib contrib-type="author"><name><surname>Westra</surname><given-names>Harm-Jan</given-names></name><xref ref-type="aff" rid="A23">23</xref></contrib><contrib contrib-type="author"><name><surname>Xavier</surname><given-names>Ramnik J.</given-names></name><xref ref-type="aff" rid="A39">39</xref><xref ref-type="aff" rid="A40">40</xref></contrib><contrib contrib-type="author"><name><surname>Zhao</surname><given-names>Zhen Z.</given-names></name><xref ref-type="aff" rid="A30">30</xref></contrib><contrib contrib-type="author"><name><surname>Ponsioen</surname><given-names>Cyriel Y.</given-names></name><xref ref-type="aff" rid="A62">62</xref></contrib><contrib contrib-type="author"><name><surname>Andersen</surname><given-names>Vibeke</given-names></name><xref ref-type="aff" rid="A63">63</xref></contrib><contrib contrib-type="author"><name><surname>Torkvist</surname><given-names>Leif</given-names></name><xref ref-type="aff" rid="A64">64</xref></contrib><contrib contrib-type="author"><name><surname>Gazouli</surname><given-names>Maria</given-names></name><xref ref-type="aff" rid="A65">65</xref></contrib><contrib contrib-type="author"><name><surname>Anagnou</surname><given-names>Nicholas P.</given-names></name><xref ref-type="aff" rid="A65">65</xref></contrib><contrib contrib-type="author"><name><surname>Karlsen</surname><given-names>Tom H.</given-names></name><xref ref-type="aff" rid="A58">58</xref></contrib><contrib contrib-type="author"><name><surname>Kupcinskas</surname><given-names>Limas</given-names></name><xref ref-type="aff" rid="A66">66</xref></contrib><contrib contrib-type="author"><name><surname>Sventoraityte</surname><given-names>Jurgita</given-names></name><xref ref-type="aff" rid="A66">66</xref></contrib><contrib contrib-type="author"><name><surname>Mansfield</surname><given-names>John C.</given-names></name><xref ref-type="aff" rid="A67">67</xref></contrib><contrib contrib-type="author"><name><surname>Kugathasan</surname><given-names>Subra</given-names></name><xref ref-type="aff" rid="A68">68</xref></contrib><contrib contrib-type="author"><name><surname>Silverberg</surname><given-names>Mark S.</given-names></name><xref ref-type="aff" rid="A56">56</xref></contrib><contrib contrib-type="author"><name><surname>Halfvarson</surname><given-names>Jonas</given-names></name><xref ref-type="aff" rid="A69">69</xref></contrib><contrib contrib-type="author"><name><surname>Rotter</surname><given-names>Jerome I.</given-names></name><xref ref-type="aff" rid="A29">29</xref></contrib><contrib contrib-type="author"><name><surname>Mathew</surname><given-names>Christopher G.</given-names></name><xref ref-type="aff" rid="A46">46</xref></contrib><contrib contrib-type="author"><name><surname>Griffiths</surname><given-names>Anne M.</given-names></name><xref ref-type="aff" rid="A60">60</xref></contrib><contrib contrib-type="author"><name><surname>Gearry</surname><given-names>Richard</given-names></name><xref ref-type="aff" rid="A14">14</xref></contrib><contrib contrib-type="author"><name><surname>Ahmad</surname><given-names>Tariq</given-names></name><xref ref-type="aff" rid="A70">70</xref></contrib><contrib contrib-type="author"><name><surname>Brant</surname><given-names>Steven R.</given-names></name><xref ref-type="aff" rid="A15">15</xref></contrib><contrib contrib-type="author"><name><surname>Chamaillard</surname><given-names>Mathias</given-names></name><xref ref-type="aff" rid="A71">71</xref></contrib><contrib contrib-type="author"><name><surname>Satsangi</surname><given-names>Jack</given-names></name><xref ref-type="aff" rid="A4">4</xref></contrib><contrib contrib-type="author"><name><surname>Cho</surname><given-names>Judy H.</given-names></name><xref ref-type="aff" rid="A47">47</xref><xref ref-type="aff" rid="A72">72</xref></contrib><contrib contrib-type="author"><name><surname>Schreiber</surname><given-names>Stefan</given-names></name><xref ref-type="aff" rid="A5">5</xref><xref ref-type="aff" rid="A73">73</xref></contrib><contrib contrib-type="author"><name><surname>Daly</surname><given-names>Mark J.</given-names></name><xref ref-type="aff" rid="A74">74</xref></contrib><contrib contrib-type="author"><name><surname>Barrett</surname><given-names>Jeffrey C.</given-names></name><xref ref-type="aff" rid="A1">1</xref></contrib><contrib contrib-type="author"><name><surname>Parkes</surname><given-names>Miles</given-names></name><xref ref-type="aff" rid="A8">8</xref></contrib><contrib contrib-type="author"><name><surname>Annese</surname><given-names>Vito</given-names></name><xref ref-type="aff" rid="A10">10</xref><xref ref-type="aff" rid="A37">37</xref></contrib><contrib contrib-type="author"><name><surname>Hakonarson</surname><given-names>Hakon</given-names></name><xref ref-type="aff" rid="A13">13</xref><xref ref-type="aff" rid="A75">75</xref><xref rid="FN3" ref-type="author-notes">80</xref></contrib><contrib contrib-type="author"><name><surname>Radford-Smith</surname><given-names>Graham</given-names></name><xref ref-type="aff" rid="A76">76</xref><xref rid="FN3" ref-type="author-notes">80</xref></contrib><contrib contrib-type="author"><name><surname>Duerr</surname><given-names>Richard H.</given-names></name><xref ref-type="aff" rid="A11">11</xref><xref ref-type="aff" rid="A77">77</xref><xref rid="FN3" ref-type="author-notes">80</xref></contrib><contrib contrib-type="author"><name><surname>Vermeire</surname><given-names>Séverine</given-names></name><xref ref-type="aff" rid="A49">49</xref><xref rid="FN3" ref-type="author-notes">80</xref></contrib><contrib contrib-type="author"><name><surname>Weersma</surname><given-names>Rinse K.</given-names></name><xref ref-type="aff" rid="A78">78</xref><xref rid="FN3" ref-type="author-notes">80</xref></contrib><contrib contrib-type="author"><name><surname>Rioux</surname><given-names>John D.</given-names></name><xref ref-type="aff" rid="A2">2</xref><xref ref-type="aff" rid="A3">3</xref></contrib></contrib-group><aff id="A1"><label>1</label>Wellcome Trust Sanger Institute, Wellcome Trust Genome Campus, Hinxton, Cambridge, UK</aff><aff id="A2"><label>2</label>Université de Montréal, Medicine, Montréal, Québec, Canada</aff><aff id="A3"><label>3</label>Montreal Heart Institute, Research Center, Montréal, Québec, Canada</aff><aff id="A4"><label>4</label>University of Edinburgh, Western General Hospital, Gastrointestinal Unit, Molecular Medicine Centre, Edinburgh, UK</aff><aff id="A5"><label>5</label>Christian-Albrechts-University Kiel, Institute of Clinical Molecular Biology, Kiel, Germany</aff><aff id="A6"><label>6</label>Karolinska Institute, Department of Biosciences and Nutrition, Stockholm, Sweden</aff><aff id="A7"><label>7</label>Cedars-Sinai Medical Center, Inflammatory Bowel and Immunobiology Research Institute, Los Angeles, California, USA</aff><aff id="A8"><label>8</label>Addenbrooke’s Hospital, University of Cambridge, Gastroenterology Research Unit, Cambridge, UK</aff><aff id="A9"><label>9</label>The Children’s Hospital of Philadelphia, Center for Applied Genomics, Philadelphia, Pennsylvania, USA</aff><aff id="A10"><label>10</label>IRCCS-CSS Hospital, Unit of Gastroenterology, San Giovanni Rotondo, Italy</aff><aff id="A11"><label>11</label>University of Pittsburgh School of Medicine, Division of Gastroenterology, Hepatology and Nutrition, Department of Medicine, Pittsburgh, Pennsylvania, USA</aff><aff id="A12"><label>12</label>Erasmus Hospital, Free University of Brussels, Department of Gastroenterology, Brussels, Belgium</aff><aff id="A13"><label>13</label>The Children’s Hospital of Philadelphia, Department of Pediatrics, Center for Pediatric Inflammatory Bowel Disease, Philadelphia, Pennsylvania, USA</aff><aff id="A14"><label>14</label>University of Otago, Department of Medicine, Christchurch, New Zealand</aff><aff id="A15"><label>15</label>Johns Hopkins University School of Medicine, Meyeroff Inflammatory Bowel Disease Center, Dept. of Medicine, Baltimore, Maryland, USA</aff><aff id="A16"><label>16</label>University Hospital Munich, Department of Medicine II, Munich, Germany</aff><aff id="A17"><label>17</label>Universitätsmedizin Berlin, Department of Gastroenterology, Charité, Campus Mitte, Berlin, Germany</aff><aff id="A18"><label>18</label>Université de Lille Department of Hepato-Gastroenterology, Lille, France</aff><aff id="A19"><label>19</label>Cincinnati Children’s Hospital Medical Center, Pediatric Gastroenterology, Cincinnati, Ohio, USA</aff><aff id="A20"><label>20</label>Ghent University Hospital, Department of Hepatology and Gastroenterology, Ghent, Belgium</aff><aff id="A21"><label>21</label>Cedars-Sinai Medical Center, Department of Pediatrics, Los Angeles, California, USA</aff><aff id="A22"><label>22</label>Torbay Hospital, Department of Gastroenterology, Torbay, Devon, UK</aff><aff id="A23"><label>23</label>University Medical Center Groningen, Department of Genetics, Groningen, the Netherlands</aff><aff id="A24"><label>24</label>Mater Health Services, Department of Gastroenterology, Brisbane, Australia</aff><aff id="A25"><label>25</label>Erasmus Hospital, Free University of Brussels, Department of Gastroenterology, Brussels, Belgium</aff><aff id="A26"><label>26</label>University of Liège, Department of Genetics, Faculty of Veterinary Medicine, Liège, Belgium</aff><aff id="A27"><label>27</label>University of Washington, Cardiovascular Health Research Unit, Department of Internal Medicine, Seattle, Washington, USA</aff><aff id="A28"><label>28</label>University of Utah School of Medicine, Department of Pediatrics, Salt Lake City, Utah, USA</aff><aff id="A29"><label>29</label>Cedars-Sinai Medical Center, Medical Genetics Institute, Los Angeles, California, USA</aff><aff id="A30"><label>30</label>Queensland Institute of Medical Research, Genetic Epidemiology, Brisbane, Australia</aff><aff id="A31"><label>31</label>Université Paris Diderot & INSERM & Hopital Robert Debre APHP, Gastroenterology, Paris, France</aff><aff id="A32"><label>32</label>Hôpital Maisonneuve-Rosemont, Dept of Gastroenterology, Montréal, Québec, Canada</aff><aff id="A33"><label>33</label>The University of Western Australia, School of Medicine and Pharmacology, Fremantle, Australia</aff><aff id="A34"><label>34</label>Université Paris Diderot, GETAID group, Paris, France</aff><aff id="A35"><label>35</label>Tel Aviv University, Pediatric Gastroenterology Unit, Wolfson Medical Center and Sackler School of Medicine, Tel Aviv, Israel</aff><aff id="A36"><label>36</label>Centre Hospitalier Universitaire Université de Liège, Division of Gastroenterology, Liège, Belgium</aff><aff id="A37"><label>37</label>AOU Careggi, Unit of Gastroenterology SOD2, Florence, Italy</aff><aff id="A38"><label>38</label>Ninewells Hospital and Medical School, Dept of Medicine, Dundee, UK</aff><aff id="A39"><label>39</label>Massachusetts General Hospital, Harvard Medical School, Gastroenterology Unit Boston, Massachusetts, USA</aff><aff id="A40"><label>40</label>Center for Computational and Integrative Biology, Massachusetts General Hospital, Boston, Massachusetts, USA</aff><aff id="A41"><label>41</label>University of Manchester Department of Medical Genetics, Manchester, UK</aff><aff id="A42"><label>42</label>University Medical Center Utrecht, Department of Medical Genetics, Utrecht, Netherlands</aff><aff id="A43"><label>43</label>University of Florence, Institute of Human Genetics, Florence, Italy</aff><aff id="A44"><label>44</label>University Hospital of Nancy, Department of Hepato-Gastroenterology, Vandoeuvre-lès-Nancy, France</aff><aff id="A45"><label>45</label>Hospital Clínic de Barcelona, IDIBAPS, CIBERehd, Department of Gastroenterology, Barcelona, Spain</aff><aff id="A46"><label>46</label>King’s College London School of Medicine, Guy’s Hospital, Department of Medical and Molecular Genetics, London, UK</aff><aff id="A47"><label>47</label>Yale University, Section of Digestive Diseases, Department of Medicine, New Haven, Connecticut, USA</aff><aff id="A48"><label>48</label>Royal Hospital for Sick Children, Paediatric Gastroenterology and Nutrition, Glasgow, UK</aff><aff id="A49"><label>49</label>University Hospital Gasthuisberg, Division of Gastroenterology, Leuven, Belgium</aff><aff id="A50"><label>50</label>Guy’s & St Thomas’ NHS Foundation Trust, St Thomas’ Hospital, Dept Gastroenterology, London, UK</aff><aff id="A51"><label>51</label>Hospital Clínic de Barcelona, IDIBAPS, CIBERehd, Department of Gastroenterology, Barcelona, Spain</aff><aff id="A52"><label>52</label>University of Chicago, Department of Health Studies, Chicago, Illinois, USA</aff><aff id="A53"><label>53</label>University of Bern, Division of Gastroenterology, Inselspital, Bern, Switzerland</aff><aff id="A54"><label>54</label>Genome Institute of Singapore, Human Genetics, Singapore</aff><aff id="A55"><label>55</label>Institute for Human Genetics, University of California San Fransisco, San Francisco, California, USA</aff><aff id="A56"><label>56</label>University of Toronto, Mount Sinai Hospital Inflammatory Bowel Disease Centre, Toronto, Ontario, Canada</aff><aff id="A57"><label>57</label>Rudolf Magnus Institute of Neuroscience, University Medical Center Utrecht, Department of Neurology, Utrecht, the Netherlands</aff><aff id="A58"><label>58</label>Rikshospitalet University Hospital, Medical Department, Oslo, Norway</aff><aff id="A59"><label>59</label>Leiden University Medical Center, Experimental Gastroenterology, Leiden, the Netherlands</aff><aff id="A60"><label>60</label>The Hospital for Sick Children, University of Toronto, Toronto, Ontario, Canada</aff><aff id="A61"><label>61</label>Child Life and Health, University of Edinburgh, Scotland</aff><aff id="A62"><label>62</label>Academic Medical Center, Department of Gastroenterology, Amsterdam, the Netherlands</aff><aff id="A63"><label>63</label>Viborg Regional Hospital, Medical Department, Viborg, Denmark</aff><aff id="A64"><label>64</label>Karolinska Institutet, Department of Clinical Science Intervention and Technology, Stockholm, Sweden</aff><aff id="A65"><label>65</label>University of Athens, Department of Biology, School of Medicine, Athens, Greece</aff><aff id="A66"><label>66</label>Kaunas University of medicine, Department of Gastroenterology, Kaunas, Lithuania</aff><aff id="A67"><label>67</label>Newcastle University, Institute of Human Genetics, Newcastle upon Tyne, UK</aff><aff id="A68"><label>68</label>Emory School of Medicine, Department of Genetics and Department of Pediatrics, Atlanta, Georgia, USA</aff><aff id="A69"><label>69</label>Örebro University Hospital, Department of Medicine, Örebro, Sweden</aff><aff id="A70"><label>70</label>Peninsula College of Medicine and Dentistry, Barrack Road, Exeter, UK</aff><aff id="A71"><label>71</label>Inserm, U1019, Lille, France</aff><aff id="A72"><label>72</label>Yale University, Department of Genetics, Yale School of Medicine, New Haven CT</aff><aff id="A73"><label>73</label>Department for General Internal Medicine, Christian-Albrechts-University, Kiel, Germany</aff><aff id="A74"><label>74</label>Massachusetts General Hospital, Harvard Medical School, Center for Human Genetic Research, Boston, Massachusetts, USA</aff><aff id="A75"><label>75</label>Center for Applied Genomics, The Children’s Hospital of Philadelphia, Philadelphia, Pennsylvania, USA</aff><aff id="A76"><label>76</label>Queensland Institute of Medical Research, IBD Research Group, Brisbane, Australia</aff><aff id="A77"><label>77</label>University of Pittsburgh Graduate School of Public Health, 130 Desoto Street, Pittsburgh, PA, USA</aff><aff id="A78"><label>78</label>University Medical Center Groningen, Department of Gastroenterology, Groningen, the Netherlands</aff><author-notes><corresp id="FN1">Correspondence should be addressed to C.A.A. (<email>carl.anderson@sanger.ac.uk</email>)or J.D.R. (<email>john.david.rioux@umontreal.ca</email>)</corresp><fn id="FN2" fn-type="equal"><label>79</label><p>These authors contributed equally to this work.</p></fn><fn id="FN3" fn-type="equal"><label>80</label><p>These authors contributed equally to this work.</p></fn></author-notes><pub-date pub-type="nihms-submitted"><day>5</day><month>4</month><year>2011</year></pub-date><pub-date pub-type="epub"><day>06</day><month>2</month><year>2011</year></pub-date><pub-date pub-type="ppub"><month>3</month><year>2011</year></pub-date><pub-date pub-type="pmc-release"><day>01</day><month>9</month><year>2011</year></pub-date><volume>43</volume><issue>3</issue><fpage>246</fpage><lpage>252</lpage><pmc-comment>elocation-id from pubmed: 10.1038/ng.764</pmc-comment>
<permissions><license xlink:href="http://www.nature.com/authors/editorial_policies/license.html#terms"><license-p>Users may view, print, copy, and download text and data-mine the content in such documents, for the purposes of academic research, subject always to the full Conditions of use:<ext-link ext-link-type="uri" xlink:href="http://www.nature.com/authors/editorial_policies/license.html#terms">http://www.nature.com/authors/editorial_policies/license.html#terms</ext-link></license-p></license></permissions><abstract><p id="P1">Genome-wide association studies (GWAS) and candidate gene studies in ulcerative colitis (UC) have identified 18 susceptibility loci. We conducted a meta-analysis of 6 UC GWAS, comprising 6,687 cases and 19,718 controls, and followed-up the top association signals in 9,628 cases and 12,917 controls. We identified 29 additional risk loci (P<5×10<sup>-8</sup>), increasing the number of UC associated loci to 47. After annotating associated regions using GRAIL, eQTL data and correlations with non-synonymous SNPs, we identified many candidate genes providing potentially important insights into disease pathogenesis, including <italic>IL1R2, IL8RA/B, IL7R, IL12B, DAP, PRDM1, JAK2, IRF5, GNA12</italic> and <italic>LSP1</italic>. The total number of confirmed inflammatory bowel disease (IBD) risk loci is now 99, including a minimum of 28 shared association signals between Crohn’s disease (CD) and UC.</p></abstract></article-meta></front><body><p id="P2">UC and CD represent the two major forms of inflammatory bowel disease (IBD: OMIM #266600), which together affect approximately 1:250 people in Europe, North America and Australasia. Clinical features, epidemiological data and genetic evidence suggest that UC and CD are related polygenic diseases. In contrast to CD, bowel inflammation in UC is limited to the colonic mucosa. While disease-related mortality is low, morbidity remains high and 10-20% of affected individuals will undergo colectomy. Though the precise etiology is unknown, the current hypothesis is a dysregulated mucosal immune response to commensal gut flora in genetically susceptible individuals<sup><xref rid="R1" ref-type="bibr">1</xref></sup>. Recent genome-wide and candidate-gene association studies have identified 18 UC susceptibility loci, including 7 that overlap with CD (e.g. <italic>IL23</italic> pathway genes, <italic>NKX2-3</italic> and <italic>IL10</italic>). Known UC specific loci (<italic>HNF4A, CDH1</italic> and <italic>LAMB1</italic>) have highlighted the role of defective barrier function in disease pathogenesis<sup><xref rid="R2" ref-type="bibr">2</xref></sup>.</p><p id="P3">The 18 confirmed UC loci explain approximately 11% of UC heritability (see <xref rid="SD2" ref-type="supplementary-material">Online Methods</xref>). To identify additional UC susceptibility loci and further elucidate disease pathogenesis, we combined data from six GWAS using genotype imputation and meta-analysis methodology (see <xref rid="SD2" ref-type="supplementary-material">Online Methods</xref>). The discovery panel consisted of 6,687 cases and 19,718 controls of European descent with data available for at least 1.1 million SNPs (<xref rid="SD2" ref-type="supplementary-material">Supplementary Table 1</xref>). A quantile-quantile plot of the meta-analysis test statistics showed a marked excess of significant associations in the tail of the distribution (<xref rid="SD2" ref-type="supplementary-material">Supplementary Figure 1</xref>). Although the majority (16/18) of previously confirmed UC loci are at a genome-wide significant level (P<5×10<sup>-8</sup>), two just failed to meet this threshold in the meta-analysis – 4q27<sup><xref rid="R3" ref-type="bibr">3</xref></sup>, and 22q13<sup><xref rid="R4" ref-type="bibr">4</xref></sup> (<xref rid="T1" ref-type="table">Table 1</xref>), though we still consider these to be true risk loci given the strength of association in the initial studies (<italic>P</italic>=1.35×10<sup>-10</sup> and <italic>P</italic>=4.21×10<sup>−8</sup> respectively). Fifty loci with P< 1×10<sup>-5</sup> and not previously associated with UC were followed up by genotyping the most associated SNP from each locus in an independent panel of 9,628 UC cases and 12,917 population controls (see <xref rid="SD2" ref-type="supplementary-material">Online Methods</xref> and <xref rid="SD2" ref-type="supplementary-material">Supplementary Table 2</xref>). Of these, 28 loci had evidence of association (P<0.05) in the follow-up panel and attained genome-wide significance (P<5×10<sup>-8</sup>) in the combined analysis of meta-analysis and follow-up cohorts (<xref rid="T2" ref-type="table">Table 2</xref> and <xref rid="SD2" ref-type="supplementary-material">Supplementary Table 3</xref>). In addition, although the locus on 1q32 failed follow-up genotyping (rs7554511) it had been previously tested for association to UC in an independent cohort (rs11584383: P = 1.2×10<sup>-5</sup>)<sup><xref rid="R5" ref-type="bibr">5</xref></sup>. This alternative tag SNP achieves genome-wide significancein our current meta-analysis (P=3.7×10<sup>-11</sup>) and therefore we consider this a confirmed UC locus, bringing the total number of new UC loci to 29. It should be noted that 12 of the 29 loci had documented nominal evidence of association (5×10<sup>-8</sup><sup><xref rid="R2" ref-type="bibr">2</xref></sup>, 1q32<sup><xref rid="R6" ref-type="bibr">6</xref></sup>, 5q33<sup><xref rid="R6" ref-type="bibr">6</xref></sup>, 6p21<sup><xref rid="R5" ref-type="bibr">5</xref></sup>, 7q32<sup><xref rid="R7" ref-type="bibr">7</xref></sup>, 9p24<sup><xref rid="R5" ref-type="bibr">5</xref>,<xref rid="R8" ref-type="bibr">8</xref></sup>, 9q34<sup><xref rid="R5" ref-type="bibr">5</xref>,<xref rid="R9" ref-type="bibr">9</xref></sup>, 10p11<sup><xref rid="R6" ref-type="bibr">6</xref></sup>, 11q23<sup><xref rid="R5" ref-type="bibr">5</xref></sup>, 13q12<sup><xref rid="R8" ref-type="bibr">8</xref></sup>, 13q13<sup><xref rid="R2" ref-type="bibr">2</xref></sup> and 20q13<sup><xref rid="R10" ref-type="bibr">10</xref></sup>). We also tested the 28 loci with follow-up genotype data for association with two clinically relevant disease sub-phenotypes (maximum disease extent and need for colectomy for medically refractory disease) but no significant associations were seen following correction for multiple testing (P<5.2×10<sup>-4</sup>) (<xref rid="SD2" ref-type="supplementary-material">Supplementary Table 4</xref>). In summary, there are 47 confirmed UC susceptibility loci, 18 from previous studies and 29 from the current study.</p><p id="P4">As a first step towards obtaining biological insight from the identification of these 47 loci, we examined the gene content of the associated regions (<xref rid="SD2" ref-type="supplementary-material">Supplementary Figure 2</xref>). Although three regions contained a single gene (5p15:<italic>DAP</italic>, 6q21:<italic>PRDM1</italic>, 10q24:<italic>NKX2-3</italic>), most (35/47) contain multiple genes and nine are not believed to contain any gene (<xref rid="T1" ref-type="table">Table 1</xref>). We attempted to identify plausible candidate genes by (a) using a literature-mining tool (GRAIL) to identify non-random, evidence-based links between genes, (b) searching an existing eQTL database<sup><xref rid="R11" ref-type="bibr">11</xref></sup> for correlations with our most associated SNPs (<xref rid="SD2" ref-type="supplementary-material">Supplementary Table 5</xref>), (c) using 1000 genomes data to identify non-synonymous SNPs in linkage disequilibrium (LD) (r<sup>2</sup>>0.5) with the most associated SNP in the locus (<xref rid="SD2" ref-type="supplementary-material">Supplementary Table 6</xref>), and (d) determining the gene in closest physical proximity to the most associated SNP (see <xref rid="SD2" ref-type="supplementary-material">Online Methods</xref>). These approaches (results summarized in <xref rid="T1" ref-type="table">Table 1</xref>, <xref rid="T2" ref-type="table">Table 2</xref> and <xref rid="SD2" ref-type="supplementary-material">Supplementary Table 7</xref>) consistently identified a single candidate gene in six of the associated regions (2q11:<italic>IL1R2</italic>, 5p15:<italic>IL7R</italic>, 7p22:<italic>GNA12</italic>, 10p11:<italic>CCNY</italic>, 1p31:<italic>IL23R</italic>, 16q22:<italic>ZFP90</italic>), potentially prioritizing which genes to follow up in future genetic and functional studies. Noteworthy candidate genes are described in <xref rid="BX1" ref-type="boxed-text">Box 1</xref>. Follow-up genotyping in even larger independent panels of cases and controls from a range of ethnicities may be needed to identify the genes containing causal variants.</p><boxed-text id="BX1" position="float" orientation="portrait"><caption><title>BOX 1 – Candidate genes within associated loci</title></caption><p id="P5"><bold><italic>TNFRSF14 / MMEL1</italic> (1p36)</bold>. <italic>TNFRSF14</italic> encodes a member of the TNF receptor superfamily. In a T cell transfer model of colitis, <italic>TNFRSF14</italic> expression by innate immune cells has an important role in preventing intestinal inflammation<sup><xref rid="R22" ref-type="bibr">22</xref></sup>. <italic>MMEL1</italic> encodes membrane metalloendopeptidase-like 1. This locus is associated with susceptibility to celiac disease and primary biliary cirrhosis; a nsSNP in MMEL1 was nominally associated with multiple sclerosis.</p><p id="P6"><bold><italic>TNFRSF9</italic> (1p36)</bold>: Tumour necrosis factor receptor superfamily member 9 is involved as a co-stimulator in the regulation of peripheral T cell activation, with enhanced proliferation and IL2 secretion. It is expressed by dendritic cells, granulocytes and endothelial cells at sites of inflammation. SCID mice transferred with naive CD4+ T cells from TNFRFSF9-deficient mice develop colitis of equal intensity as SCID mice transferred with wild type naïve T cells, but with amodified cytokine response<sup><xref rid="R23" ref-type="bibr">23</xref></sup>.</p><p id="P7"><bold><italic>IL1R2</italic> (2q11)</bold>: Interleukin 1 receptor, type II binds IL1a, IL1b and IL1R1, inhibiting the activity of these ligands. Two alternative splice transcripts of <italic>IL1R2</italic> have been reported. This protein serves to antagonise the action of IL1a and IL1b, pleiotropic cytokines with various roles in inflammatory processes. IL1b production by lamina propria macrophages is increased in patients with UC<sup><xref rid="R24" ref-type="bibr">24</xref></sup>.</p><p id="P8">This locus is immediately adjacent to a CD-associated locus containing <italic>IL18RAP, ILR1</italic> and other genes. It is unclear at present whether the CD-associated and UC-associated SNPs in these regions tag two separate loci or one locus. The lead CD SNP has a P=0.001 in our UC meta-analysis. There is a large recombination hotspot between <italic>IL1R2</italic> (UC) and <italic>IL1R1</italic> (CD).</p><p id="P9"><bold><italic>IL8RA / IL8RB</italic> (2q35)</bold>: <italic>IL8RA</italic> and <italic>IL8RB</italic> encode two receptors for interleukin-8, a powerful neutrophil chemotactic factor. <italic>IL8RA</italic> expression, limited to a subpopulation of lamina propria macrophages and germinal centre lymphocytes in the healthy colon, is increased in macrophages, lymphocytes and epithelium in UC<sup><xref rid="R25" ref-type="bibr">25</xref></sup>. <italic>IL8RB</italic> expression is more limited and not upregulated in UC. <italic>IL8</italic> expression is profoundly increased in colonic tissue from UC patients compared with controls; this increase is driven by inflammation<sup><xref rid="R26" ref-type="bibr">26</xref></sup>.</p><p id="P10"><bold><italic>DAP</italic> (5p15)</bold> encodes death-associated protein. The DAPs are a heterogenous group of polypeptides isolated in a screen for elements involved in the IFNγ – induced apoptosis of HeLa cells. <italic>DAP</italic> negatively regulates autophagy and is a substrate of mTOR<sup><xref rid="R13" ref-type="bibr">13</xref></sup>.</p><p id="P11"><bold><italic>IL7R</italic> (5p13)</bold> encodes the receptor for interleukin-7. IL7 is a key regulator of naïve and memory T cell survival, specifically the transition from effector to memory T cells<sup><xref rid="R27" ref-type="bibr">27</xref></sup>. T cells expressing high levels of IL7R are seen in human and murine colitis; selective depletion of these cells ameliorates established colitis <sup><xref rid="R28" ref-type="bibr">28</xref></sup>. <italic>IL7R</italic> is a confirmed multiple sclerosis susceptibility gene<sup><xref rid="R29" ref-type="bibr">29</xref></sup>. The gene may have undergone extensive evolutionary selective pressure by intestinal helminths<sup><xref rid="R30" ref-type="bibr">30</xref></sup>.</p><p id="P12"><bold><italic>PRDM1</italic> (6q21)</bold> encodes PR domain containing 1, with ZNF domain (synonym BLIMP1), the master transcriptional regulator of plasma cells and a transcriptional repressor of the IFN-β promoter. It plays important roles in the proliferation, survival and differentiation of B and T lymphocytes.</p><p id="P13"><bold><italic>GNA12</italic> (7p22)</bold> encodes guanine nucleotide binding protein (G protein) alpha 12, a membrane bound GTPase that plays an important role in tight junction assembly in epithelial cells, through interactions with ZO-1 and Src<sup><xref rid="R20" ref-type="bibr">20</xref></sup>.</p><p id="P14"><bold><italic>IRF5 (7q32)</italic></bold> encoding interferon regulatory factor 5, is a confirmed susceptibility gene for rheumatoid arthritis, SLE and primary biliary cirrhosis. This transcription factor regulates activity of type I interferons and induces cytokines including IL-6, IL-12 and TNFα, via TLR signaling. In response to mycobacterium tuberculosis infection of macrophages, Type I interferon expression is dependent on a pathway including <italic>IRF5, NOD2</italic> and <italic>RIP2</italic><sup><xref rid="R31" ref-type="bibr">31</xref></sup>.</p><p id="P15"><bold><italic>LSP1</italic> (11q15)</bold>: Lymphocyte-specific protein-1 is expressed by lymphocytes and macrophages, and also in endothelium wherein it is critical for normal neutrophil transmigration<sup><xref rid="R32" ref-type="bibr">32</xref></sup>.</p></boxed-text><p id="P16">Additional bioinformatic analyses were also performed on the entire set of genes in the associated regions to search for functional commonalities across this large number of loci (see <xref rid="SD2" ref-type="supplementary-material">Online Methods</xref>). Specifically, using a gene set enrichment approach the UC loci are seen to have more genes associated with cytokines and cytokine receptors (including IFNγ, several interleukins, five TNF and TNFR superfamily members), key regulators of cytokine-mediated signaling pathways, innate and adaptive immune response, macrophage activation and regulation of apoptosis than would be expected by chance (<xref rid="SD2" ref-type="supplementary-material">Supplementary Table 8 and Supplementary Figure 3</xref>). Enrichment analysis of the subset of candidate loci with no known association to other inflammatory diseases showed significant over-representation of gene sets associated with MAP kinase signaling, actin binding, calcium-dependent processes, fatty acid and lipid metabolism (<xref rid="SD2" ref-type="supplementary-material">Supplementary Table 8 and Supplementary Figure 3</xref>).</p><p id="P17">The 5p15 locus contains a single gene, <italic>DAP</italic> (death-associated protein), with the most associated SNP in this region having a strong eQTL effect on <italic>DAP</italic> expression (P=2.59×10<sup>-12</sup>)<sup><xref rid="R11" ref-type="bibr">11</xref></sup>. DAP kinase expression has been shown to increase with inflammation in UC<sup><xref rid="R12" ref-type="bibr">12</xref></sup>, and DAP itself has recently been identified as a novel substrate of mTOR (mammalian target of rapamycin)<sup><xref rid="R13" ref-type="bibr">13</xref></sup> and as a negative regulator of autophagy. While autophagic processes have previously been implicated in CD due to associations with <italic>ATG16L1</italic> and <italic>IRGM</italic><sup><xref rid="R14" ref-type="bibr">14</xref></sup>, this association with <italic>DAP</italic> suggests a possible link between autophagy and UC.</p><p id="P18">Association to loci containing <italic>PRDM1, IRF5</italic> and <italic>NKX2-3</italic> suggests an important role for transcriptional regulation in UC pathogenesis. A key example is BLIMP-1, encoded by the <italic>PRDM1</italic> gene, whose most important function is in B cells, as the master transcriptional regulator of plasma cells<sup><xref rid="R15" ref-type="bibr">15</xref></sup>. It also functions in T cells to attenuate IL-2 production upon antigen stimulation<sup><xref rid="R16" ref-type="bibr">16</xref></sup>, and topromote the development of short-lived effector cells and regulate clonal exhaustion in both CD4 and CD8 cells<sup><xref rid="R17" ref-type="bibr">17</xref></sup>. It is noteworthy that the 11q24 celiac disease susceptibility locus containing ETS1, a transcription factor essential for T-bet induced production of <italic>IFNγ</italic> and the development of colitis in animal models, just fails to reach genome-wide significance in our study (P=1.22×10<sup>-7</sup>, <xref rid="SD2" ref-type="supplementary-material">Supplementary Table 3b</xref>)<sup><xref rid="R18" ref-type="bibr">18</xref>,<xref rid="R19" ref-type="bibr">19</xref></sup>.</p><p id="P19">Identification of <italic>GNA12</italic> as the most likely candidate at the 7p22 locus suggests a role for intestinal barrier function as this gene is implicated in tight junction assembly in epithelial cells<sup><xref rid="R20" ref-type="bibr">20</xref></sup>. Barrier integrity appears to be a key pathway in UC pathogenesis given previous associations to loci containing <italic>HNF4A, CDH1</italic> and <italic>LAMB1</italic><sup><xref rid="R2" ref-type="bibr">2</xref>,<xref rid="R5" ref-type="bibr">5</xref></sup>.</p><p id="P20">Given the phenotypic overlap between UC and CD, we examined the evidence for association at all 47 UC loci in our recently completed CD GWAS meta-analysis comprising 6,333 cases and 15,056 controls<sup><xref rid="R14" ref-type="bibr">14</xref></sup> and, conversely, for evidence of association at all confirmed CD loci in our UC meta-analysis (<xref rid="T3" ref-type="table">Table 3</xref> and <xref rid="SD2" ref-type="supplementary-material">Supplementary Table 9</xref>). We find that, among the 99 confirmed IBD loci meeting genome-wide significance (P<5×10<sup>-8</sup>) either in UC and/or CD, 28 independent index SNPs have P<1×10<sup>-4</sup> in both scans. Interestingly, all index SNPs meeting these criteria showed the same direction of effect in both diseases, thus pointing to a minimum of 28 shared association signals between UC and CD. Multiple genes involved in the IL23 signaling pathway are included in this overlapping SNP list, specifically <italic>IL23R, JAK2, STAT3, IL12B (p40)</italic>, and <italic>PTPN2</italic>. The significance of these findings is underlined by the central role played by <italic>IL23</italic> in the induction of <italic>IL17</italic> by Th17 lymphocytes, its established role in other autoimmune disorders, and the intense interest in therapeutic manipulation of the <italic>IL23-IL23R</italic> interaction through blockade of the p40 or p19 IL23 subunits.</p><p id="P21">Loci not meeting these inclusion criteria cannot be formally discounted as shared loci, indeed many of the confirmed UC/CD loci with nominal association (1×10<sup>-4</sup>0.05) of association to CD are the three containing candidate genes that play a role in intestinal barrier function (<italic>GNA12, HNF4A</italic>, and <italic>LAMB1</italic>).</p> <p id="P22">In addition to loci shared with CD, 19 of the 47 UC risk loci are also associated with other immune-mediated diseases (<xref rid="T1" ref-type="table">Table 1</xref> and <xref rid="T2" ref-type="table">Table 2</xref>). In particular, these “shared loci” are enriched for genes involved in T-cell differentiation, specifically in the differentiation of T<sub>H</sub>1 and T<sub>H</sub>17 cells (e.g. loci encoding <italic>IL23R, IL21, IL10, IL7R, IFNG</italic>). Dysregulated auto-antigen specific T<sub>H</sub>1 responses are believed to be involved in organ-specific autoimmune diseases, and T<sub>H</sub>17 cells are increasingly recognized to contribute to host defense and induction of autoimmunity and tissue inflammation<sup><xref rid="R21" ref-type="bibr">21</xref></sup>. Another shared pathway between UC and other immune mediated diseases involves TNF-signaling (<italic>TNFRSF9, TNFRSF14, TNFSF15</italic>) with widespread immunological effects including NF-κB activation, a known key component of the inflammatory response in IBD.</p><p id="P23">The current study has more than doubled the number of confirmed UC susceptibility loci and we estimate that 16% of UC heritability is explained by these loci (see <xref rid="SD2" ref-type="supplementary-material">Online Methods</xref>). We have identified potentially causal genes at several loci but confirmation of causality awaits detailed fine-mapping, expression and functional studies. Dense fine-mapping and large-scale re-sequencing studies are underway with the goal of identifying the causal variation within many of these loci.</p><sec sec-type="supplementary-material" id="S1"><title>Supplementary Material</title><supplementary-material content-type="local-data" id="SD2"><label>1</label><media xlink:href="NIHMS265993-supplement-1.pdf" mimetype="application" mime-subtype="pdf" orientation="portrait" xlink:type="simple" id="d37e1973" position="anchor"></media></supplementary-material><supplementary-material content-type="local-data" id="SD3"><label>2</label><media xlink:href="NIHMS265993-supplement-2.xlsx" mimetype="application" mime-subtype="vnd.ms-excel" orientation="portrait" xlink:type="simple" id="d37e1977" position="anchor"></media></supplementary-material><supplementary-material content-type="local-data" id="SD4"><label>3</label><media xlink:href="NIHMS265993-supplement-3.xlsx" mimetype="application" mime-subtype="vnd.ms-excel" orientation="portrait" xlink:type="simple" id="d37e1981" position="anchor"></media></supplementary-material><supplementary-material content-type="local-data" id="SD5"><label>4</label><media xlink:href="NIHMS265993-supplement-4.xlsx" mimetype="application" mime-subtype="vnd.ms-excel" orientation="portrait" xlink:type="simple" id="d37e1985" position="anchor"></media></supplementary-material><supplementary-material content-type="local-data" id="SD6"><label>5</label><media xlink:href="NIHMS265993-supplement-5.xlsx" mimetype="application" mime-subtype="vnd.ms-excel" orientation="portrait" xlink:type="simple" id="d37e1989" position="anchor"></media></supplementary-material><supplementary-material content-type="local-data" id="SD7"><media xlink:href="NIHMS265993-supplement-supplement_1.pdf" mimetype="application" mime-subtype="pdf" orientation="portrait" xlink:type="simple" id="d37e1991" position="anchor"></media></supplementary-material></sec></body><back><ack id="S2"><p>In memoriam to Marc Lémann, who dedicated his life to his patients but died too soon.</p><p>We thank all subjects who contributed samples, and physicians and nursing staff who helped with recruitment globally. This study was supported by the German Ministry of Education and Research through the National Genome Research Network, the popgen biobank and infrastructure support through the DFG cluster of excellence ‘Inflammation at Interfaces. Italian case collections were supported by the Italian Group for IBD and the Italian Society for Paediatric Gastroenterology, Hepatology and Nutrition. We acknowledge funding provided by Royal Brisbane and Women’s Hospital Foundation; University of Queensland (Ferguson Fellowship); National Health and Medical Research Council, Australia and by the European Community (5th PCRDT). UK case collections were supported by the National Association for Colitis and Crohn’s disease, Wellcome Trust, Medical Research Council UK and Peninsular College of Medicine and Dentistry, Exeter. Activities in Sweden were supported by the Swedish Society of Medicine, the Bengt Ihre Foundation, the Karolinska Institutet, the Swedish National Program for IBD Genetics, the Swedish Organization for IBD, the Swedish Medical Research Council, the Soderbergh Foundation and the Swedish Cancer Foundation. Support for genotyping and genetic data analysis was provided by the Agency for Science Technology and Research (A*STAR), Singapore. We are grateful to the funders and investigators of the Epidemiological Investigation of Rheumatoid Arthritis for providing genotype data from healthy Swedish individuals.</p><p>The Wellcome Trust Case Control Consortium 2 project was supported by Wellcome Trust grant 083948/Z/07/Z. We also acknowledge the NIHR Biomedical Research Centre awards to Guy’s & St.Thomas’ NHS Trust/King’s College London and to Addenbrooke’s Hospital/University of Cambridge School of Clinical Medicine/University of Manchester and Central Manchester Foundation Trust. The NIDDK IBD Genetics Consortium is funded by the following grants: DK062431 (SRB), DK062422 (JHC), DK062420 (RHD), DK062432 (JDR), DK062423 (MSS), DK062413(DPBM), DK076984 (MJD), and DK084554 (MJD and DPBM), and DK062429 (JHC). JHC is also funded by the Crohn’s and Colitis Foundation of America; SLG by DK069513 and Primary Children’s Medical Center Foundation, and JDR by NIH/NIDDK grant DK064869. Cedars Sinai supported by NCRR grant M01-RR00425; NIH/NIDDK grant P01-DK046763; DK 063491; and Cedars-Sinai Medical Center Inflammatory Bowel Disease Research Funds. RW is supported by a clinical fellow grant (90700281) from the Netherlands Organization for Scientific Research; EL, DF and SV are senior clinical investigators for the Funds for Scientific Research (FWO/FNRS) Belgium. SB was supported by Deutsche Forschungsgemeinschaft (DFG BR 1912/5-1) and Else Kröner-Fresenius-Stiftung (P50/05/EKMS05/62). MC was supported by the Programme Hospitalier de Recherche Clinique. CAA is supported by Wellcome Trust grant WT091745/Z/10/Z. JCB is supported by Wellcome Trust grant WT089120/Z/09/Z. RKW is supported by a clinical fellowship grant (90.700.281) from the Netherlands Organization for Scientific Research (NWO). CW is supported by grants from the Celiac Disease Consortium (BSIK03009) and the Netherlands Organization for Scientific Research (NWO, VICI grant 918.66.620). LHvdB acknowledges funding from the Prinses Beatrix Fonds, the Adessium foundation and the Amyotrophic Lateral Sclerosis Association. LF received a Horizon Breakthrough grant from the Netherlands Genomics Initiative (93519031) and a VENI grant from NWO (ZonMW grant 916.10.135). RJX and AN are funded by DK83756, AI062773, DK043351 and the Helmsley Foundation.</p><p>Replication genotyping was supported by unrestricted grants from Abbott Laboratories Ltd, Giuliani SpA, Shire PLC and Ferring Pharmaceuticals. We thank the 1958 British Birth Cohort and Banco Nacional deADN, Salamanca, Spain who supplied control DNA samples. The IBSEN study group and the Norwegian Bone Marrow Donor Registry are acknowledged for contributing the Norwegian patient and control populations. The CHS research reported in this article was supported by contract numbers N01-HC-85079 through N01-HC- 85086, N01-HC-35129, N01 HC-15103, N01 HC-55222, N01-HC-75150, N01-HC- 45133, grant numbers U01 HL080295 and R01 HL087652 from the National Heart, Lung, and Blood Institute, with additional contribution from the National Institute of Neurological Disorders and Stroke. A full list of principal CHS investigators and institutions can be found at <ext-link ext-link-type="uri" xlink:href="http://www.chs-nhlbi.org/pi.htm">http://www.chs-nhlbi.org/pi.htm</ext-link>. We thank the members of the Quebec IBD Genetic Consortium, in particular A. Bitton, G. Aumais, E.J. Bernard, A. Cohen, C. Deslandres, R. Lahaie, D. Langelier and P. Paré. Other significant contributors: K. Hanigan, N. Huang, P. Webb, D. Whiteman, A. Rutherford, R. Gwilliam, J. Ghori, D Strachan, W. McCardle, W. Ouwehand, M. Newsky, S. Ehlers, I. Pauselius, K. Holm, C. Sina, M. Regueiro, A. Andriulli and M.C. Renda.</p></ack><fn-group><fn id="FN4" fn-type="con"><p><bold>Contribution of authors</bold> CWL, AF, KDT, JCL, MI, AL, LA, LB, RNB, MB, TMB, SB, CB, J-FC, LAD, MdV, MD, CE, RSNF, TF, DF, MG, JG, NLG, SLG, TH, NKH, J-PH, GJ, DL, IL, ML, AL, CLi, EL, DPM, MM, CM, AN, WN, RAO, LP, OP, LPB, JP, AP, NJP, DDP, RRo, RRu, PR, JS, MS, PS, FS, YS, MS, AHS, SRT, LHvdB, MV, HV, TW, CW, DCW, H-JW, CYP, VA, LT, MG, NPA, THK, LK, JS, JCM, SK, MSS, JH, JIR, CGM, AMG, RG, TA, SRB, MC, JS, JHC, SS, MP, VA, HH, GRS, RHD, SV, RKW and JDR established DNA collections, recruited patients or assembled phenotypic data; AF, MD’A, PG, CLa, RS, SB, CLi, DPM, GWM, LS, ZZZ, MC, RHD, and JDR conducted or supervised laboratory work; CAA, GB, DE, JABF, LF, KIM, AN, RAO, RJX, MJD, JCB, RKW, and JDR performed or supervised statistical analyses; CAA, GB, CWL, GRS, RHD, SV, RKW and JDR drafted the manuscript. All authors read and approved the final manuscript before submission.</p></fn><fn id="FN5" fn-type="conflict"><p>All authors declare no financial interest.</p></fn></fn-group><ref-list><ref id="R1"><label>1</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Xavier</surname><given-names>RJ</given-names></name><name><surname>Podolsky</surname><given-names>DK</given-names></name></person-group><article-title>Unravelling the pathogenesis of inflammatory bowel disease</article-title><source>Nature</source><volume>448</volume><fpage>427</fpage><lpage>34</lpage><year>2007</year><pub-id pub-id-type="pmid">17653185</pub-id></element-citation></ref><ref id="R2"><label>2</label><element-citation publication-type="journal"><collab>The UK IBD Genetics Consortium & The Wellcome Trust Case Control Consortium 2</collab><article-title>Genome-wide association study of ulcerative colitis identifies three new susceptibility loci, including the HNF4A region</article-title><source>Nat Genet</source><volume>41</volume><fpage>1330</fpage><lpage>4</lpage><year>2009</year><pub-id pub-id-type="pmid">19915572</pub-id></element-citation></ref><ref id="R3"><label>3</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Festen</surname><given-names>EA</given-names></name><etal></etal></person-group><article-title>Genetic variants in the region harbouring IL2/IL21 associated with ulcerative colitis</article-title><source>Gut</source><volume>58</volume><fpage>799</fpage><lpage>804</lpage><year>2009</year><pub-id pub-id-type="pmid">19201773</pub-id></element-citation></ref><ref id="R4"><label>4</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Franke</surname><given-names>A</given-names></name><etal></etal></person-group><article-title>Genome-wide association study for ulcerative colitis identifies risk loci at 7q22 and 22q13 (IL17REL)</article-title><source>Nat Genet</source><volume>42</volume><fpage>292</fpage><lpage>4</lpage><year>2010</year><pub-id pub-id-type="pmid">20228798</pub-id></element-citation></ref><ref id="R5"><label>5</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>McGovern</surname><given-names>DP</given-names></name><etal></etal></person-group><article-title>Genome-wide association identifies multiple ulcerative colitis susceptibility loci</article-title><source>Nat Genet</source><volume>42</volume><fpage>332</fpage><lpage>7</lpage><year>2010</year><pub-id pub-id-type="pmid">20228799</pub-id></element-citation></ref><ref id="R6"><label>6</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Franke</surname><given-names>A</given-names></name><etal></etal></person-group><article-title>Replication of signals from recent studies of Crohn’s disease identifies previously unknown disease loci for ulcerative colitis</article-title><source>Nat Genet</source><volume>40</volume><fpage>713</fpage><lpage>5</lpage><year>2008</year><pub-id pub-id-type="pmid">18438405</pub-id></element-citation></ref><ref id="R7"><label>7</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Dideberg</surname><given-names>V</given-names></name><etal></etal></person-group><article-title>An insertion-deletion polymorphism in the interferon regulatory Factor 5 (IRF5) gene confers risk of inflammatory bowel diseases</article-title><source>Hum Mol Genet</source><volume>16</volume><fpage>3008</fpage><lpage>16</lpage><year>2007</year><pub-id pub-id-type="pmid">17881657</pub-id></element-citation></ref><ref id="R8"><label>8</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Asano</surname><given-names>K</given-names></name><etal></etal></person-group><article-title>A genome-wide association study identifies three new susceptibility loci for ulcerative colitis in the Japanese population</article-title><source>Nat Genet</source><volume>41</volume><fpage>1325</fpage><lpage>9</lpage><year>2009</year><pub-id pub-id-type="pmid">19915573</pub-id></element-citation></ref><ref id="R9"><label>9</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Zhernakova</surname><given-names>A</given-names></name><etal></etal></person-group><article-title>Genetic analysis of innate immunity in Crohn’s disease and ulcerative colitis identifies two susceptibility loci harboring CARD9 and IL18RAP</article-title><source>Am J Hum Genet</source><volume>82</volume><fpage>1202</fpage><lpage>10</lpage><year>2008</year><pub-id pub-id-type="pmid">18439550</pub-id></element-citation></ref><ref id="R10"><label>10</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Kugathasan</surname><given-names>S</given-names></name><etal></etal></person-group><article-title>Loci on 20q13 and 21q22 are associated with pediatric-onset inflammatory bowel disease</article-title><source>Nat Genet</source><volume>40</volume><fpage>1211</fpage><lpage>5</lpage><year>2008</year><pub-id pub-id-type="pmid">18758464</pub-id></element-citation></ref><ref id="R11"><label>11</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Dubois</surname><given-names>PC</given-names></name><etal></etal></person-group><article-title>Multiple common variants for celiac disease influencing immune gene expression</article-title><source>Nat Genet</source><volume>42</volume><fpage>295</fpage><lpage>302</lpage><year>2010</year><pub-id pub-id-type="pmid">20190752</pub-id></element-citation></ref><ref id="R12"><label>12</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Kuester</surname><given-names>D</given-names></name><etal></etal></person-group><article-title>Aberrant methylation of DAPK in long-standing ulcerative colitis and ulcerative colitis-associated carcinoma</article-title><source>Pathol Res Pract</source><year>2010</year></element-citation></ref><ref id="R13"><label>13</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Koren</surname><given-names>I</given-names></name><name><surname>Reem</surname><given-names>E</given-names></name><name><surname>Kimchi</surname><given-names>A</given-names></name></person-group><article-title>DAP1, a novel substrate of mTOR, negatively regulates autophagy</article-title><source>Curr Biol</source><volume>20</volume><fpage>1093</fpage><lpage>8</lpage><year>2010</year><pub-id pub-id-type="pmid">20537536</pub-id></element-citation></ref><ref id="R14"><label>14</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Franke</surname><given-names>A</given-names></name><etal></etal></person-group><article-title>International Association Analysis Increases to 71 the Tally of Confirmed Crohn’s Disease Susceptibility Loci</article-title><source>Submitted</source></element-citation></ref><ref id="R15"><label>15</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Turner</surname><given-names>CA</given-names><suffix>Jr</suffix></name><name><surname>Mack</surname><given-names>DH</given-names></name><name><surname>Davis</surname><given-names>MM</given-names></name></person-group><article-title>Blimp-1, a novel zinc finger-containing protein that can drive the maturation of B lymphocytes into immunoglobulin-secreting cells</article-title><source>Cell</source><volume>77</volume><fpage>297</fpage><lpage>306</lpage><year>1994</year><pub-id pub-id-type="pmid">8168136</pub-id></element-citation></ref><ref id="R16"><label>16</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Martins</surname><given-names>GA</given-names></name><name><surname>Cimmino</surname><given-names>L</given-names></name><name><surname>Liao</surname><given-names>J</given-names></name><name><surname>Magnusdottir</surname><given-names>E</given-names></name><name><surname>Calame</surname><given-names>K</given-names></name></person-group><article-title>Blimp-1 directly represses Il2 and the Il2 activator Fos, attenuating T cell proliferation and survival</article-title><source>J Exp Med</source><volume>205</volume><fpage>1959</fpage><lpage>65</lpage><year>2008</year><pub-id pub-id-type="pmid">18725523</pub-id></element-citation></ref><ref id="R17"><label>17</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Kallies</surname><given-names>A</given-names></name><name><surname>Xin</surname><given-names>A</given-names></name><name><surname>Belz</surname><given-names>GT</given-names></name><name><surname>Nutt</surname><given-names>SL</given-names></name></person-group><article-title>Blimp-1 transcription factor is required for the differentiation of effector CD8(+) T cells and memory responses</article-title><source>Immunity</source><volume>31</volume><fpage>283</fpage><lpage>95</lpage><year>2009</year><pub-id pub-id-type="pmid">19664942</pub-id></element-citation></ref><ref id="R18"><label>18</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Grenningloh</surname><given-names>R</given-names></name><name><surname>Kang</surname><given-names>BY</given-names></name><name><surname>Ho</surname><given-names>IC</given-names></name></person-group><article-title>Ets-1, a functional cofactor of T-bet, is essential for Th1 inflammatory responses</article-title><source>J Exp Med</source><volume>201</volume><fpage>615</fpage><lpage>26</lpage><year>2005</year><pub-id pub-id-type="pmid">15728239</pub-id></element-citation></ref><ref id="R19"><label>19</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Moisan</surname><given-names>J</given-names></name><name><surname>Grenningloh</surname><given-names>R</given-names></name><name><surname>Bettelli</surname><given-names>E</given-names></name><name><surname>Oukka</surname><given-names>M</given-names></name><name><surname>Ho</surname><given-names>IC</given-names></name></person-group><article-title>Ets-1 is a negative regulator of Th17 differentiation</article-title><source>J Exp Med</source><volume>204</volume><fpage>2825</fpage><lpage>35</lpage><year>2007</year><pub-id pub-id-type="pmid">17967903</pub-id></element-citation></ref><ref id="R20"><label>20</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Sabath</surname><given-names>E</given-names></name><etal></etal></person-group><article-title>Galpha12 regulates protein interactions within the MDCK cell tight junction and inhibits tight-junction assembly</article-title><source>J Cell Sci</source><volume>121</volume><fpage>814</fpage><lpage>24</lpage><year>2008</year><pub-id pub-id-type="pmid">18285450</pub-id></element-citation></ref><ref id="R21"><label>21</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Bettelli</surname><given-names>E</given-names></name><name><surname>Korn</surname><given-names>T</given-names></name><name><surname>Oukka</surname><given-names>M</given-names></name><name><surname>Kuchroo</surname><given-names>VK</given-names></name></person-group><article-title>Induction and effector functions of T(H)17 cells</article-title><source>Nature</source><volume>453</volume><fpage>1051</fpage><lpage>7</lpage><year>2008</year><pub-id pub-id-type="pmid">18563156</pub-id></element-citation></ref><ref id="R22"><label>22</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Steinberg</surname><given-names>MW</given-names></name><etal></etal></person-group><article-title>A crucial role for HVEM and BTLA in preventing intestinal inflammation</article-title><source>J Exp Med</source><volume>205</volume><fpage>1463</fpage><lpage>76</lpage><year>2008</year><pub-id pub-id-type="pmid">18519647</pub-id></element-citation></ref><ref id="R23"><label>23</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Maerten</surname><given-names>P</given-names></name><etal></etal></person-group><article-title>Involvement of 4-1BB (CD137)-4-1BBligand interaction in the modulation of CD4 T cell-mediated inflammatory colitis</article-title><source>Clin Exp Immunol</source><volume>143</volume><fpage>228</fpage><lpage>36</lpage><year>2006</year><pub-id pub-id-type="pmid">16412046</pub-id></element-citation></ref><ref id="R24"><label>24</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Mahida</surname><given-names>YR</given-names></name><name><surname>Wu</surname><given-names>K</given-names></name><name><surname>Jewell</surname><given-names>DP</given-names></name></person-group><article-title>Enhanced production of interleukin 1-beta by mononuclear cells isolated from mucosa with active ulcerative colitis of Crohn’s disease</article-title><source>Gut</source><volume>30</volume><fpage>835</fpage><lpage>8</lpage><year>1989</year><pub-id pub-id-type="pmid">2787769</pub-id></element-citation></ref><ref id="R25"><label>25</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Williams</surname><given-names>EJ</given-names></name><etal></etal></person-group><article-title>Distribution of the interleukin-8 receptors, CXCR1 and CXCR2, in inflamed gut tissue</article-title><source>J Pathol</source><volume>192</volume><fpage>533</fpage><lpage>9</lpage><year>2000</year><pub-id pub-id-type="pmid">11113872</pub-id></element-citation></ref><ref id="R26"><label>26</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Noble</surname><given-names>CL</given-names></name><etal></etal></person-group><article-title>Regional variation in gene expression in the healthy colon is dysregulated in ulcerative colitis</article-title><source>Gut</source><volume>57</volume><fpage>1398</fpage><lpage>405</lpage><year>2008</year><pub-id pub-id-type="pmid">18523026</pub-id></element-citation></ref><ref id="R27"><label>27</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Schluns</surname><given-names>KS</given-names></name><name><surname>Kieper</surname><given-names>WC</given-names></name><name><surname>Jameson</surname><given-names>SC</given-names></name><name><surname>Lefrancois</surname><given-names>L</given-names></name></person-group><article-title>Interleukin-7 mediates the homeostasis of naive and memory CD8 T cells in vivo</article-title><source>Nat Immunol</source><volume>1</volume><fpage>426</fpage><lpage>32</lpage><year>2000</year><pub-id pub-id-type="pmid">11062503</pub-id></element-citation></ref><ref id="R28"><label>28</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Yamazaki</surname><given-names>M</given-names></name><etal></etal></person-group><article-title>Mucosal T cells expressing high levels of IL-7 receptor are potential targets for treatment of chronic colitis</article-title><source>J Immunol</source><volume>171</volume><fpage>1556</fpage><lpage>63</lpage><year>2003</year><pub-id pub-id-type="pmid">12874249</pub-id></element-citation></ref><ref id="R29"><label>29</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Gregory</surname><given-names>SG</given-names></name><etal></etal></person-group><article-title>Interleukin 7 receptor alpha chain (IL7R) shows allelic and functional association with multiple sclerosis</article-title><source>Nat Genet</source><volume>39</volume><fpage>1083</fpage><lpage>91</lpage><year>2007</year><pub-id pub-id-type="pmid">17660817</pub-id></element-citation></ref><ref id="R30"><label>30</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Fumagalli</surname><given-names>M</given-names></name><etal></etal></person-group><article-title>Parasites represent a major selective force for interleukin genes and shape the genetic predisposition to autoimmune conditions</article-title><source>J Exp Med</source><volume>206</volume><fpage>1395</fpage><lpage>408</lpage><year>2009</year><pub-id pub-id-type="pmid">19468064</pub-id></element-citation></ref><ref id="R31"><label>31</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Pandey</surname><given-names>AK</given-names></name><etal></etal></person-group><article-title>NOD2, RIP2 and IRF5 play a critical role in the type I interferon response to Mycobacterium tuberculosis</article-title><source>PLoS Pathog</source><volume>5</volume><fpage>e1000500</fpage><year>2009</year><pub-id pub-id-type="pmid">19578435</pub-id></element-citation></ref><ref id="R32"><label>32</label><element-citation publication-type="journal"><person-group person-group-type="author"><name><surname>Liu</surname><given-names>L</given-names></name><etal></etal></person-group><article-title>LSP1 is an endothelial gatekeeper of leukocyte transendothelial migration</article-title><source>J Exp Med</source><volume>201</volume><fpage>409</fpage><lpage>18</lpage><year>2005</year><pub-id pub-id-type="pmid">15684321</pub-id></element-citation></ref></ref-list></back><floats-group><table-wrap id="T1" position="float" orientation="landscape"><label>Table 1</label><caption><title>Association results and <italic>in silico</italic> analyses for the 18 previously confirmed (P<5×10<sup>-8</sup>) ulcerative colitis (UC) loci</title><p>Left-right association boundaries are given for each index SNP (see <xref rid="SD2" ref-type="supplementary-material">Online Methods</xref>). RAF = risk allele frequency. OR is estimated using the meta-analysis cohort only. Known associations represent phenotypes previously associated with the locus at P<5×10<sup>-8</sup>). AS = ankylosing spondylitis, Ast = Asthma, BD = Behçet’s disease, CD = Crohn’s disease, CeD = celiac disease, Graves’ disease = GD, HL = Hodgkin’s lymphoma, MS = multiple sclerosis, PBC = primary biliary sclerosis, Ps = psoriasis, RA = rheumatoid arthritis, SLE = systemic lupus erythematosus, T1D = type 1 diabetes and WBC = white blood cell count. Candidate genes of interest are listed. Those in bold were highlighted by <italic>in silico</italic> analyses (GRAIL connectivity and/or presence of an eQTL or nonsynonymous SNP. See <xref rid="SD2" ref-type="supplementary-material">Online Methods and Supplementary Table 7</xref> for more details).</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" rowspan="1" colspan="1">dbSNP ID</th><th align="left" rowspan="1" colspan="1">Chr.</th><th align="left" rowspan="1" colspan="1">Left-right(Mb)</th><th align="center" rowspan="1" colspan="1">Risk Allele</th><th align="center" rowspan="1" colspan="1">Allele frequency in controls</th><th align="center" rowspan="1" colspan="1">P-value (meta)</th><th align="center" rowspan="1" colspan="1">OR (95% CI)</th><th align="left" rowspan="1" colspan="1">Association reported with other phenotypes</th><th align="left" rowspan="1" colspan="1">Positional candidate genes of interest</th></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1">rs6426833</td><td align="left" rowspan="1" colspan="1">1p36</td><td align="left" rowspan="1" colspan="1">19.93-20.18</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.541</td><td align="center" rowspan="1" colspan="1">3.93 × 10<sup>-35</sup></td><td align="center" rowspan="1" colspan="1">1.30 (1.25-1.35)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">rs11209026</td><td align="left" rowspan="1" colspan="1">1p31</td><td align="left" rowspan="1" colspan="1">67.30-67.54</td><td align="center" rowspan="1" colspan="1">G</td><td align="center" rowspan="1" colspan="1">0.935</td><td align="center" rowspan="1" colspan="1">5.12 × 10<sup>-28</sup></td><td align="center" rowspan="1" colspan="1">1.74 (1.57-1.92)</td><td align="left" rowspan="1" colspan="1">CD, AS, BD, Ps</td><td align="left" rowspan="1" colspan="1"><bold><italic>IL23R</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs1801274</td><td align="left" rowspan="1" colspan="1">1q23</td><td align="left" rowspan="1" colspan="1">159.54-159.91</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.505</td><td align="center" rowspan="1" colspan="1">2.16 × 10<sup>-20</sup></td><td align="center" rowspan="1" colspan="1">1.21 (1.16-1.26)</td><td align="left" rowspan="1" colspan="1">SLE</td><td align="left" rowspan="1" colspan="1"><bold><italic>FCGR2A, FCGR2B, HSPA6</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs3024505</td><td align="left" rowspan="1" colspan="1">1q32</td><td align="left" rowspan="1" colspan="1">204.85-205.11</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.159</td><td align="center" rowspan="1" colspan="1">5.76 × 10<sup>-17</sup></td><td align="center" rowspan="1" colspan="1">1.25 (1.19-1.32)</td><td align="left" rowspan="1" colspan="1">CD, BD, SLE, T1D</td><td align="left" rowspan="1" colspan="1"><bold><italic>IL10, IL19</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs7608910</td><td align="left" rowspan="1" colspan="1">2p16</td><td align="left" rowspan="1" colspan="1">60.76-61.87</td><td align="center" rowspan="1" colspan="1">G</td><td align="center" rowspan="1" colspan="1">0.390</td><td align="center" rowspan="1" colspan="1">1.70 × 10<sup>-14</sup></td><td align="center" rowspan="1" colspan="1">1.19 (1.14-1.24)</td><td align="left" rowspan="1" colspan="1">CD, CeD, RA</td><td align="left" rowspan="1" colspan="1"><italic>PUS10</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">rs4676406</td><td align="left" rowspan="1" colspan="1">2q37</td><td align="left" rowspan="1" colspan="1">241.20-241.32</td><td align="center" rowspan="1" colspan="1">T</td><td align="center" rowspan="1" colspan="1">0.516</td><td align="center" rowspan="1" colspan="1">8.32 × 10<sup>-11</sup></td><td align="center" rowspan="1" colspan="1">1.14 (1.09-1.18)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"><bold><italic>GPR35</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs9822268</td><td align="left" rowspan="1" colspan="1">3p21</td><td align="left" rowspan="1" colspan="1">48.14-51.77</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.302</td><td align="center" rowspan="1" colspan="1">1.60 × 10<sup>-17</sup></td><td align="center" rowspan="1" colspan="1">1.21 (1.16-1.26)</td><td align="left" rowspan="1" colspan="1">CD</td><td align="left" rowspan="1" colspan="1"><bold>MST1, <italic>UBA7, APEH, AMIGO3, GMPPB, BSN</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs17388568</td><td align="left" rowspan="1" colspan="1">4q27</td><td align="left" rowspan="1" colspan="1">123.20-123.78</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.273</td><td align="center" rowspan="1" colspan="1">9.49 × 10<sup>-7</sup></td><td align="center" rowspan="1" colspan="1">1.12 (1.07-1.17)</td><td align="left" rowspan="1" colspan="1">CeD, T1D</td><td align="left" rowspan="1" colspan="1"><italic><bold>IL21, IL2</bold>, ADAD1</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">rs11739663</td><td align="left" rowspan="1" colspan="1">5p15</td><td align="left" rowspan="1" colspan="1">0.48-0.80</td><td align="center" rowspan="1" colspan="1">T</td><td align="center" rowspan="1" colspan="1">0.767</td><td align="center" rowspan="1" colspan="1">2.80 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.15 (1.09-1.21)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"><bold><italic>EXOC3</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs9268853</td><td align="left" rowspan="1" colspan="1">6p21</td><td align="left" rowspan="1" colspan="1">31.49-33.01</td><td align="center" rowspan="1" colspan="1">T</td><td align="center" rowspan="1" colspan="1">0.661</td><td align="center" rowspan="1" colspan="1">1.35 × 10<sup>-55</sup></td><td align="center" rowspan="1" colspan="1">1.40 (1.34-1.47)</td><td align="left" rowspan="1" colspan="1">CD, CeD, GrD, MS, PBC, RA, T1D</td><td align="left" rowspan="1" colspan="1"><bold><italic>HLA-DRB5, HLA-DQA1, HLA-DRB1, HLA-DRA, BTNL2</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs4510766</td><td align="left" rowspan="1" colspan="1">7q22</td><td align="left" rowspan="1" colspan="1">107.20-107.39</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.559</td><td align="center" rowspan="1" colspan="1">2.00 × 10<sup>-16</sup></td><td align="center" rowspan="1" colspan="1">1.20 (1.15-1.26)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">rs6584283</td><td align="left" rowspan="1" colspan="1">10q24</td><td align="left" rowspan="1" colspan="1">101.25-101.33</td><td align="center" rowspan="1" colspan="1">T</td><td align="center" rowspan="1" colspan="1">0.472</td><td align="center" rowspan="1" colspan="1">8.46 × 10<sup>-21</sup></td><td align="center" rowspan="1" colspan="1">1.21 (1.16-1.26)</td><td align="left" rowspan="1" colspan="1">CD</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">rs7134599</td><td align="left" rowspan="1" colspan="1">12q14</td><td align="left" rowspan="1" colspan="1">66.72-66.92</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.385</td><td align="center" rowspan="1" colspan="1">1.06 × 10<sup>-16</sup></td><td align="center" rowspan="1" colspan="1">1.19 (1.14-1.24)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"><bold><italic>IFNG, IL26</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs6499188</td><td align="left" rowspan="1" colspan="1">16q22</td><td align="left" rowspan="1" colspan="1">66.98-67.40</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.749</td><td align="center" rowspan="1" colspan="1">3.97 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.14 (1.09-1.20)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"><bold><italic>ZFP90</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs2872507</td><td align="left" rowspan="1" colspan="1">17q12</td><td align="left" rowspan="1" colspan="1">34.62-35.51</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.463</td><td align="center" rowspan="1" colspan="1">5.44 × 10<sup>-11</sup></td><td align="center" rowspan="1" colspan="1">1.15 (1.10-1.19)</td><td align="left" rowspan="1" colspan="1">CD, Ast, PBC, T1D, WBC</td><td align="left" rowspan="1" colspan="1"><bold><italic>IKZF3, ORMDL3, IKZF3, PNMT</italic>, ZPBP2, GSDML</bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs6017342</td><td align="left" rowspan="1" colspan="1">20q13</td><td align="left" rowspan="1" colspan="1">42.49-42.70</td><td align="center" rowspan="1" colspan="1">C</td><td align="center" rowspan="1" colspan="1">0.538</td><td align="center" rowspan="1" colspan="1">1.09 × 10<sup>-20</sup></td><td align="center" rowspan="1" colspan="1">1.20 (1.15-1.26)</td><td align="left" rowspan="1" colspan="1">HDL</td><td align="left" rowspan="1" colspan="1"><bold><italic>SERINC3</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs2836878</td><td align="left" rowspan="1" colspan="1">21q22</td><td align="left" rowspan="1" colspan="1">39.34-39.41</td><td align="center" rowspan="1" colspan="1">G</td><td align="center" rowspan="1" colspan="1">0.738</td><td align="center" rowspan="1" colspan="1">1.86 × 10<sup>-22</sup></td><td align="center" rowspan="1" colspan="1">1.25 (1.20-1.32)</td><td align="left" rowspan="1" colspan="1">AS</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">rs5771069</td><td align="left" rowspan="1" colspan="1">22q13</td><td align="left" rowspan="1" colspan="1">48.70-48.83</td><td align="center" rowspan="1" colspan="1">G</td><td align="center" rowspan="1" colspan="1">0.515</td><td align="center" rowspan="1" colspan="1">1.87 × 10<sup>-7</sup></td><td align="center" rowspan="1" colspan="1">1.11 (1.07-1.16)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"><bold><italic>PIM3, IL17REL</italic></bold></td></tr></tbody></table></table-wrap><table-wrap id="T2" position="float" orientation="landscape"><label>Table 2</label><caption><title>Association results and <italic>in silico</italic> analyses for the 29 newly confirmed ulcerative colitis (UC) loci</title><p>UC loci that meet genome-wide significance P<5×10<sup>-8</sup> in the combined analysis and P<0.05 in the replication study. Left-right association boundaries are given for each index SNP (see <xref rid="SD2" ref-type="supplementary-material">Online Methods</xref>). RAF = risk allele frequency. OR is estimated using the replication cohort only. Known associations represent phenotypes previously associated with the locus at P<5×10<sup>-8</sup>. AtD = atopic dermatitis, BMD = bone mineral density, CD = Crohn’s disease, CeD = celiac disease, Gli = glioma, Lep = leprosy, MS = multiple sclerosis, MyN = myeloproliferative neoplasms, PBC = primary biliary sclerosis, Ps = psoriasis, RA = rheumatoid arthritis and SLE = systemic lupus erythematosus. Candidate genes of interest are listed. Those in bold were highlighted by <italic>in silico</italic> analyses (GRAIL connectivity and/or presence of an eQTL or nonsynonymous SNP. See <xref rid="SD2" ref-type="supplementary-material">Online Methods and Supplementary Table 7</xref> for more details).</p></caption><table frame="hsides" rules="groups"><thead><tr><th align="left" rowspan="1" colspan="1">dbSNP ID</th><th align="left" rowspan="1" colspan="1">Chr.</th><th align="left" rowspan="1" colspan="1">Left-right(Mb)</th><th align="center" rowspan="1" colspan="1">Risk Allele</th><th align="center" rowspan="1" colspan="1">Allele frequency in controls</th><th align="center" rowspan="1" colspan="1">P-value (meta)</th><th align="center" rowspan="1" colspan="1">P-value (follow-up)</th><th align="center" rowspan="1" colspan="1">P-value (comb)</th><th align="center" rowspan="1" colspan="1">OR (95% CI)</th><th align="left" rowspan="1" colspan="1">Association reported with other phenotypes</th><th align="left" rowspan="1" colspan="1">Positional candidate genes of interest</th></tr></thead><tbody><tr><td align="left" rowspan="1" colspan="1">rs734999</td><td align="left" rowspan="1" colspan="1">1p36</td><td align="left" rowspan="1" colspan="1">2.39-2.80</td><td align="center" rowspan="1" colspan="1">C</td><td align="center" rowspan="1" colspan="1">0.524</td><td align="center" rowspan="1" colspan="1">1.21 × 10<sup>-9</sup></td><td align="center" rowspan="1" colspan="1">1.51 × 10<sup>-2</sup></td><td align="center" rowspan="1" colspan="1">3.34 × 10<sup>-9</sup></td><td align="center" rowspan="1" colspan="1">1.05 (1.01-1.09)</td><td align="left" rowspan="1" colspan="1">CeD, PBC</td><td align="left" rowspan="1" colspan="1"><bold><italic>TNFRSF14, MMEL1, PLCH2, C1orf93</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs35675666</td><td align="left" rowspan="1" colspan="1">1p36</td><td align="left" rowspan="1" colspan="1">7.83-8.13</td><td align="center" rowspan="1" colspan="1">G</td><td align="center" rowspan="1" colspan="1">0.829</td><td align="center" rowspan="1" colspan="1">1.09 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.13 × 10<sup>-2</sup></td><td align="center" rowspan="1" colspan="1">4.84 × 10<sup>-9</sup></td><td align="center" rowspan="1" colspan="1">1.08 (1.02-1.15)</td><td align="left" rowspan="1" colspan="1">CD</td><td align="left" rowspan="1" colspan="1"><italic><bold>TNFRSF9, ERFFI1,UTS2</bold>,PARK7</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">rs7524102</td><td align="left" rowspan="1" colspan="1">1p36</td><td align="left" rowspan="1" colspan="1">22.54-22.61</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.828</td><td align="center" rowspan="1" colspan="1">1.04 × 10<sup>-11</sup></td><td align="center" rowspan="1" colspan="1">2.06 × 10<sup>-4</sup></td><td align="center" rowspan="1" colspan="1">1.65 × 10<sup>-13</sup></td><td align="center" rowspan="1" colspan="1">1.10 (1.05-1.16)</td><td align="left" rowspan="1" colspan="1">BMD</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">rs7554511</td><td align="left" rowspan="1" colspan="1">1q32</td><td align="left" rowspan="1" colspan="1">199.06-199.33</td><td align="center" rowspan="1" colspan="1">C</td><td align="center" rowspan="1" colspan="1">0.721</td><td align="center" rowspan="1" colspan="1">2.04 × 10<sup>-13</sup></td><td align="center" rowspan="1" colspan="1">NA</td><td align="center" rowspan="1" colspan="1">NA</td><td align="center" rowspan="1" colspan="1">1.19 (1.14-1.25)</td><td align="left" rowspan="1" colspan="1">CD, CeD</td><td align="left" rowspan="1" colspan="1"><italic>C1orf106</italic></td></tr><tr><td align="left" rowspan="1" colspan="1">rs2310173</td><td align="left" rowspan="1" colspan="1">2q11</td><td align="left" rowspan="1" colspan="1">101.66-102.13</td><td align="center" rowspan="1" colspan="1">T</td><td align="center" rowspan="1" colspan="1">0.461</td><td align="center" rowspan="1" colspan="1">8.44 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">5.94 × 10<sup>-6</sup></td><td align="center" rowspan="1" colspan="1">3.17 × 10<sup>-12</sup></td><td align="center" rowspan="1" colspan="1">1.09 (1.05-1.14)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"><bold><italic>IL1R2</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs11676348</td><td align="left" rowspan="1" colspan="1">2q35</td><td align="left" rowspan="1" colspan="1">218.58-218.97</td><td align="center" rowspan="1" colspan="1">T</td><td align="center" rowspan="1" colspan="1">0.486</td><td align="center" rowspan="1" colspan="1">8.78 × 10<sup>-9</sup></td><td align="center" rowspan="1" colspan="1">6.18 × 10<sup>-4</sup></td><td align="center" rowspan="1" colspan="1">1.25 × 10<sup>-10</sup></td><td align="center" rowspan="1" colspan="1">1.07 (1.03-1.11)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"><bold><italic>IL8RA, SLC11A1, IL8RB, AAMP, ARPC</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs267939</td><td align="left" rowspan="1" colspan="1">5p15</td><td align="left" rowspan="1" colspan="1">10.72-10.90</td><td align="center" rowspan="1" colspan="1">C</td><td align="center" rowspan="1" colspan="1">0.368</td><td align="center" rowspan="1" colspan="1">9.67 × 10<sup>-7</sup></td><td align="center" rowspan="1" colspan="1">1.27 × 10<sup>-6</sup></td><td align="center" rowspan="1" colspan="1">6.01 × 10<sup>-12</sup></td><td align="center" rowspan="1" colspan="1">1.10 (1.06-1.15)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"><bold><italic>DAP</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs3194051</td><td align="left" rowspan="1" colspan="1">5p13</td><td align="left" rowspan="1" colspan="1">35.83-36.07</td><td align="center" rowspan="1" colspan="1">G</td><td align="center" rowspan="1" colspan="1">0.269</td><td align="center" rowspan="1" colspan="1">2.19 × 10<sup>-6</sup></td><td align="center" rowspan="1" colspan="1">2.06 × 10<sup>-3</sup></td><td align="center" rowspan="1" colspan="1">4.22 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.07 (1.02-1.12)</td><td align="left" rowspan="1" colspan="1">MS</td><td align="left" rowspan="1" colspan="1"><bold><italic>IL7R</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs6451493*</td><td align="left" rowspan="1" colspan="1">5p13</td><td align="left" rowspan="1" colspan="1">40.32-40.85</td><td align="center" rowspan="1" colspan="1">T</td><td align="center" rowspan="1" colspan="1">0.610</td><td align="center" rowspan="1" colspan="1">1.78 × 10<sup>-6</sup></td><td align="center" rowspan="1" colspan="1">2.09 × 10<sup>-4</sup></td><td align="center" rowspan="1" colspan="1">2.80 × 10<sup>-9</sup></td><td align="center" rowspan="1" colspan="1">1.08 (1.04-1.12)</td><td align="left" rowspan="1" colspan="1">CD, MS</td><td align="left" rowspan="1" colspan="1"><bold><italic>PTGER4</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs254560</td><td align="left" rowspan="1" colspan="1">5q31</td><td align="left" rowspan="1" colspan="1">134.41-134.53</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.397</td><td align="center" rowspan="1" colspan="1">3.06 × 10<sup>-7</sup></td><td align="center" rowspan="1" colspan="1">4.19 × 10<sup>-4</sup></td><td align="center" rowspan="1" colspan="1">1.25 × 10<sup>-9</sup></td><td align="center" rowspan="1" colspan="1">1.07 (1.03-1.12)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">rs6871626</td><td align="left" rowspan="1" colspan="1">5q33</td><td align="left" rowspan="1" colspan="1">158.46-158.86</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.334</td><td align="center" rowspan="1" colspan="1">1.02 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.40 × 10<sup>-14</sup></td><td align="center" rowspan="1" colspan="1">1.11 × 10<sup>-21</sup></td><td align="center" rowspan="1" colspan="1">1.17 (1.12-1.22)</td><td align="left" rowspan="1" colspan="1">CD, Ps, SLE</td><td align="left" rowspan="1" colspan="1"><bold><italic>IL12B</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs943072</td><td align="left" rowspan="1" colspan="1">6p21</td><td align="left" rowspan="1" colspan="1">43.88-43.92</td><td align="center" rowspan="1" colspan="1">G</td><td align="center" rowspan="1" colspan="1">0.092</td><td align="center" rowspan="1" colspan="1">1.05 × 10<sup>-6</sup></td><td align="center" rowspan="1" colspan="1">3.71 × 10<sup>-5</sup></td><td align="center" rowspan="1" colspan="1">2.37 × 10<sup>-10</sup></td><td align="center" rowspan="1" colspan="1">1.15 (1.08-1.23)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">rs6911490</td><td align="left" rowspan="1" colspan="1">6q21</td><td align="left" rowspan="1" colspan="1">106.51-106.67</td><td align="center" rowspan="1" colspan="1">T</td><td align="center" rowspan="1" colspan="1">0.210</td><td align="center" rowspan="1" colspan="1">3.51 × 10<sup>-7</sup></td><td align="center" rowspan="1" colspan="1">1.70 × 10<sup>-3</sup></td><td align="center" rowspan="1" colspan="1">1.01 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.08 (1.03-1.13)</td><td align="left" rowspan="1" colspan="1">CD, RA, SLE</td><td align="left" rowspan="1" colspan="1"><bold><italic>PDRM1</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs6920220</td><td align="left" rowspan="1" colspan="1">6q23</td><td align="left" rowspan="1" colspan="1">137.88-138.17</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.207</td><td align="center" rowspan="1" colspan="1">6.38 × 10<sup>-10</sup></td><td align="center" rowspan="1" colspan="1">1.94 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">8.05 × 10<sup>-17</sup></td><td align="center" rowspan="1" colspan="1">1.14 (1.09-1.20)</td><td align="left" rowspan="1" colspan="1">CeD, Ps, RA, SLE</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">rs798502</td><td align="left" rowspan="1" colspan="1">7p22</td><td align="left" rowspan="1" colspan="1">2.70-2.90</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.711</td><td align="center" rowspan="1" colspan="1">1.21 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">3.82 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">2.61 × 10<sup>-15</sup></td><td align="center" rowspan="1" colspan="1">1.13 (1.08-1.18)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"><bold><italic>GNA12</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs4728142</td><td align="left" rowspan="1" colspan="1">7q32</td><td align="left" rowspan="1" colspan="1">128.33-128.56</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.444</td><td align="center" rowspan="1" colspan="1">1.68 × 10<sup>-6</sup></td><td align="center" rowspan="1" colspan="1">1.25 × 10<sup>-3</sup></td><td align="center" rowspan="1" colspan="1">1.74 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.07 (1.03-1.11)</td><td align="left" rowspan="1" colspan="1">SLE, RA, PBC</td><td align="left" rowspan="1" colspan="1"><bold><italic>IRF5, TNPO3</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs10758669</td><td align="left" rowspan="1" colspan="1">9p24</td><td align="left" rowspan="1" colspan="1">4.93-5.28</td><td align="center" rowspan="1" colspan="1">C</td><td align="center" rowspan="1" colspan="1">0.350</td><td align="center" rowspan="1" colspan="1">8.52 × 10<sup>-13</sup></td><td align="center" rowspan="1" colspan="1">3.78 × 10<sup>-14</sup></td><td align="center" rowspan="1" colspan="1">2.22 × 10<sup>-25</sup></td><td align="center" rowspan="1" colspan="1">1.17 (1.12-1.21)</td><td align="left" rowspan="1" colspan="1">CD, MyN</td><td align="left" rowspan="1" colspan="1"><bold><italic>JAK2</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs4246905</td><td align="left" rowspan="1" colspan="1">9q32</td><td align="left" rowspan="1" colspan="1">116.48-116.74</td><td align="center" rowspan="1" colspan="1">C</td><td align="center" rowspan="1" colspan="1">0.713</td><td align="center" rowspan="1" colspan="1">4.77 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.44 × 10<sup>-5</sup></td><td align="center" rowspan="1" colspan="1">5.65 × 10<sup>-12</sup></td><td align="center" rowspan="1" colspan="1">1.10 (1.05-1.15)</td><td align="left" rowspan="1" colspan="1">CD, Lep</td><td align="left" rowspan="1" colspan="1"><bold><italic>TNFSF8, TNFSF15</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs10781499</td><td align="left" rowspan="1" colspan="1">9q34</td><td align="left" rowspan="1" colspan="1">138.27-138.55</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.411</td><td align="center" rowspan="1" colspan="1">6.95 × 10<sup>-13</sup></td><td align="center" rowspan="1" colspan="1">2.50 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">2.62 × 10<sup>-19</sup></td><td align="center" rowspan="1" colspan="1">1.12 (1.08-1.17)</td><td align="left" rowspan="1" colspan="1">CD</td><td align="left" rowspan="1" colspan="1"><bold><italic>CARD9, INPP5E, SDCCAG3, SEC16A, SNAPC4</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs12261843</td><td align="left" rowspan="1" colspan="1">10p11</td><td align="left" rowspan="1" colspan="1">35.22-35.94</td><td align="center" rowspan="1" colspan="1">G</td><td align="center" rowspan="1" colspan="1">0.286</td><td align="center" rowspan="1" colspan="1">2.35 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.22 × 10<sup>-3</sup></td><td align="center" rowspan="1" colspan="1">7.09 × 10<sup>-10</sup></td><td align="center" rowspan="1" colspan="1">1.07 (1.03-1.12)</td><td align="left" rowspan="1" colspan="1">CD</td><td align="left" rowspan="1" colspan="1"><bold><italic>CCNY</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs907611</td><td align="left" rowspan="1" colspan="1">11q15</td><td align="left" rowspan="1" colspan="1">1.82-1.93</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.317</td><td align="center" rowspan="1" colspan="1">2.49 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">3.58 × 10<sup>-4</sup></td><td align="center" rowspan="1" colspan="1">1.38 × 10<sup>-10</sup></td><td align="center" rowspan="1" colspan="1">1.08 (1.03-1.13)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"><bold><italic>LSP1</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs2155219</td><td align="left" rowspan="1" colspan="1">11q13</td><td align="left" rowspan="1" colspan="1">75.72-76.02</td><td align="center" rowspan="1" colspan="1">T</td><td align="center" rowspan="1" colspan="1">0.500</td><td align="center" rowspan="1" colspan="1">6.33 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.61 × 10<sup>-9</sup></td><td align="center" rowspan="1" colspan="1">5.39 × 10<sup>-16</sup></td><td align="center" rowspan="1" colspan="1">1.13 (1.08-1.17)</td><td align="left" rowspan="1" colspan="1">CD, AtD</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">rs678170</td><td align="left" rowspan="1" colspan="1">11q23</td><td align="left" rowspan="1" colspan="1">113.76-114.08</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.661</td><td align="center" rowspan="1" colspan="1">6.88 × 10<sup>-11</sup></td><td align="center" rowspan="1" colspan="1">2.50 × 10<sup>-5</sup></td><td align="center" rowspan="1" colspan="1">4.65 × 10<sup>-14</sup></td><td align="center" rowspan="1" colspan="1">1.09 (1.05-1.14)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">rs17085007</td><td align="left" rowspan="1" colspan="1">13q12</td><td align="left" rowspan="1" colspan="1">26.39-26.46</td><td align="center" rowspan="1" colspan="1">C</td><td align="center" rowspan="1" colspan="1">0.178</td><td align="center" rowspan="1" colspan="1">3.30 × 10<sup>-9</sup></td><td align="center" rowspan="1" colspan="1">4.66 × 10<sup>-9</sup></td><td align="center" rowspan="1" colspan="1">9.65 × 10<sup>-17</sup></td><td align="center" rowspan="1" colspan="1">1.16 (1.10-1.21)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">rs941823</td><td align="left" rowspan="1" colspan="1">13q13</td><td align="left" rowspan="1" colspan="1">39.90-39.95</td><td align="center" rowspan="1" colspan="1">C</td><td align="center" rowspan="1" colspan="1">0.756</td><td align="center" rowspan="1" colspan="1">3.93 × 10<sup>-7</sup></td><td align="center" rowspan="1" colspan="1">1.93 × 10<sup>-6</sup></td><td align="center" rowspan="1" colspan="1">3.82 × 10<sup>-12</sup></td><td align="center" rowspan="1" colspan="1">1.12 (1.07-1.17)</td><td align="left" rowspan="1" colspan="1"></td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">rs16940202</td><td align="left" rowspan="1" colspan="1">16q24</td><td align="left" rowspan="1" colspan="1">84.53-84.58</td><td align="center" rowspan="1" colspan="1">C</td><td align="center" rowspan="1" colspan="1">0.180</td><td align="center" rowspan="1" colspan="1">1.27 × 10<sup>-12</sup></td><td align="center" rowspan="1" colspan="1">1.42 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">5.96 × 10<sup>-19</sup></td><td align="center" rowspan="1" colspan="1">1.15 (1.10-1.21)</td><td align="left" rowspan="1" colspan="1">MS</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">rs2297441</td><td align="left" rowspan="1" colspan="1">20q13</td><td align="left" rowspan="1" colspan="1">61.66-61.98</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.766</td><td align="center" rowspan="1" colspan="1">5.78 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">2.68 × 10<sup>-4</sup></td><td align="center" rowspan="1" colspan="1">1.70 × 10<sup>-10</sup></td><td align="center" rowspan="1" colspan="1">1.09 (1.04-1.15)</td><td align="left" rowspan="1" colspan="1">CD, Gli</td><td align="left" rowspan="1" colspan="1"><bold><italic>SLC2A4RG, STMN3, ZBTB46, ZGPAT, RTEL1, TNFRSF6B</italic></bold></td></tr><tr><td align="left" rowspan="1" colspan="1">rs1297265</td><td align="left" rowspan="1" colspan="1">21q21</td><td align="left" rowspan="1" colspan="1">15.62-15.77</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.564</td><td align="center" rowspan="1" colspan="1">1.73 × 10<sup>-7</sup></td><td align="center" rowspan="1" colspan="1">5.02 × 10<sup>-7</sup></td><td align="center" rowspan="1" colspan="1">6.99 × 10<sup>-13</sup></td><td align="center" rowspan="1" colspan="1">1.11 (1.06-1.15)</td><td align="left" rowspan="1" colspan="1">CD</td><td align="left" rowspan="1" colspan="1"></td></tr><tr><td align="left" rowspan="1" colspan="1">rs2838519</td><td align="left" rowspan="1" colspan="1">21q22</td><td align="left" rowspan="1" colspan="1">44.41-44.52</td><td align="center" rowspan="1" colspan="1">G</td><td align="center" rowspan="1" colspan="1">0.390</td><td align="center" rowspan="1" colspan="1">2.26 × 10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">7.10 × 10<sup>-4</sup></td><td align="center" rowspan="1" colspan="1">6.41 × 10<sup>-11</sup></td><td align="center" rowspan="1" colspan="1">1.14 (1.05-1.22)</td><td align="left" rowspan="1" colspan="1">CD</td><td align="left" rowspan="1" colspan="1"><bold><italic>ICOSLG</italic></bold></td></tr></tbody></table></table-wrap><table-wrap id="T3" position="float" orientation="landscape"><label>Table 3</label><caption><title>Shared association signals between UC and CD</title><p>A shared association is defined as a confirmed association (P<sub>combined</sub> < 5×10<sup>-8</sup>) in either UC or CD and P<sub>meta</sub> < 1×10-4 in the other form of IBD. For more details and comparative results across all 99 reported IBD risk loci see <xref rid="SD2" ref-type="supplementary-material">Supplementary table 9</xref>.</p></caption><table frame="box" rules="groups"><thead><tr><th align="center" colspan="2" valign="bottom" rowspan="1">LOCUS</th><th align="center" valign="bottom" rowspan="1" colspan="1">GENE</th><th align="center" colspan="3" valign="bottom" rowspan="1">INDEX SNP</th><th align="center" colspan="2" valign="bottom" rowspan="1">CD-META (6333/15056)</th><th align="center" colspan="2" valign="bottom" rowspan="1">UC-META (6687/19718)</th></tr><tr><th align="center" rowspan="1" colspan="1">Chr.</th><th align="center" rowspan="1" colspan="1">Left-Right (Mb)</th><th align="center" rowspan="1" colspan="1">Candidate</th><th align="center" rowspan="1" colspan="1">SNP</th><th align="center" rowspan="1" colspan="1">Risk Allele</th><th align="center" rowspan="1" colspan="1">Allele frequency in controls</th><th align="center" rowspan="1" colspan="1">p-value</th><th align="center" rowspan="1" colspan="1">OR (95% CI)</th><th align="center" rowspan="1" colspan="1">p-value</th><th align="center" rowspan="1" colspan="1">OR (95% CI)</th></tr></thead><tbody><tr><td align="center" rowspan="1" colspan="1">1p31</td><td align="center" rowspan="1" colspan="1">67.30-67.54</td><td align="center" rowspan="1" colspan="1"><italic>IL23R</italic></td><td align="center" rowspan="1" colspan="1">rs11209026</td><td align="center" rowspan="1" colspan="1">G</td><td align="center" rowspan="1" colspan="1">0.94</td><td align="center" rowspan="1" colspan="1">1.00×10<sup>-64</sup></td><td align="center" rowspan="1" colspan="1">2.67 (2.37-3.01)</td><td align="center" rowspan="1" colspan="1">5.12×10<sup>-28</sup></td><td align="center" rowspan="1" colspan="1">1.74 (1.57-1.92)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">1q32</td><td align="center" rowspan="1" colspan="1">199.0 -199.33</td><td align="center" rowspan="1" colspan="1"><italic>KIF21B</italic></td><td align="center" rowspan="1" colspan="1">rs7554511</td><td align="center" rowspan="1" colspan="1">C</td><td align="center" rowspan="1" colspan="1">0.72</td><td align="center" rowspan="1" colspan="1">1.58×10<sup>-7</sup></td><td align="center" rowspan="1" colspan="1">1.14 (1.08-1.19)</td><td align="center" rowspan="1" colspan="1">2.04×10<sup>-13</sup></td><td align="center" rowspan="1" colspan="1">1.19 (1.14-1.25)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">1q32</td><td align="center" rowspan="1" colspan="1">204.85-205.11</td><td align="center" rowspan="1" colspan="1"><italic>IL10</italic></td><td align="center" rowspan="1" colspan="1">rs3024505</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.16</td><td align="center" rowspan="1" colspan="1">8.32×10<sup>-9</sup></td><td align="center" rowspan="1" colspan="1">1.18 (1.12-1.25)</td><td align="center" rowspan="1" colspan="1">5.76×10<sup>-17</sup></td><td align="center" rowspan="1" colspan="1">1.25 (1.19-1.32)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">2p16</td><td align="center" rowspan="1" colspan="1">60.76-61.87</td><td align="center" rowspan="1" colspan="1"><italic>REL</italic></td><td align="center" rowspan="1" colspan="1">rs7608910</td><td align="center" rowspan="1" colspan="1">G</td><td align="center" rowspan="1" colspan="1">0.39</td><td align="center" rowspan="1" colspan="1">3.11×10<sup>-7</sup></td><td align="center" rowspan="1" colspan="1">1.14 (1.09-1.21)</td><td align="center" rowspan="1" colspan="1">1.70×10<sup>-14</sup></td><td align="center" rowspan="1" colspan="1">1.19 (1.14-1.24)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">2q11</td><td align="center" rowspan="1" colspan="1">101.66-102.13</td><td align="center" rowspan="1" colspan="1"><italic>IL1R2</italic></td><td align="center" rowspan="1" colspan="1">rs2310173</td><td align="center" rowspan="1" colspan="1">T</td><td align="center" rowspan="1" colspan="1">0.46</td><td align="center" rowspan="1" colspan="1">8.31×10<sup>-5</sup></td><td align="center" rowspan="1" colspan="1">1.09 (1.04-1.14)</td><td align="center" rowspan="1" colspan="1">8.44×10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.12 (1.07-1.16)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">3p21</td><td align="center" rowspan="1" colspan="1">48.14-51.77</td><td align="center" rowspan="1" colspan="1"><italic>MST1</italic></td><td align="center" rowspan="1" colspan="1">rs3197999</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.30</td><td align="center" rowspan="1" colspan="1">6.17×10<sup>-17</sup></td><td align="center" rowspan="1" colspan="1">1.22 (1.16-1.27)</td><td align="center" rowspan="1" colspan="1">1.86×10<sup>-17</sup></td><td align="center" rowspan="1" colspan="1">1.21 (1.16-1.26)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">5p13</td><td align="center" rowspan="1" colspan="1">40.32-40.85</td><td align="center" rowspan="1" colspan="1"><italic>PTGER4</italic></td><td align="center" rowspan="1" colspan="1">rs6451493</td><td align="center" rowspan="1" colspan="1">T</td><td align="center" rowspan="1" colspan="1">0.61</td><td align="center" rowspan="1" colspan="1">1.61×10<sup>-27</sup></td><td align="center" rowspan="1" colspan="1">1.35 (1.28-1.43)</td><td align="center" rowspan="1" colspan="1">1.78×10<sup>-6</sup></td><td align="center" rowspan="1" colspan="1">1.12 (1.07-1.17)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1"></td><td align="center" rowspan="1" colspan="1"></td><td align="center" rowspan="1" colspan="1"><italic>IL12B</italic></td><td align="center" rowspan="1" colspan="1">rs6871626</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.33</td><td align="center" rowspan="1" colspan="1">6.08×10<sup>-12</sup></td><td align="center" rowspan="1" colspan="1">1.15 (1.10-1.20)</td><td align="center" rowspan="1" colspan="1">1.02×10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.12 (1.08-1.17)</td></tr><tr><td align="center" rowspan="1" colspan="1">5q33</td><td align="center" rowspan="1" colspan="1">158.46-158.86</td><td align="center" rowspan="1" colspan="1"><italic>IL12B</italic></td><td align="center" rowspan="1" colspan="1">rs6556412 (r<sup>2</sup>=0.03)</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.34</td><td align="center" rowspan="1" colspan="1">5.37×10<sup>-14</sup></td><td align="center" rowspan="1" colspan="1">1.18 (1.13-1.23)</td><td align="center" rowspan="1" colspan="1">1.69×10<sup>-5</sup></td><td align="center" rowspan="1" colspan="1">1.09 (1.05-1.14)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">6p22</td><td align="center" rowspan="1" colspan="1">20.60-21.25</td><td align="center" rowspan="1" colspan="1"><italic>CDKAL1</italic></td><td align="center" rowspan="1" colspan="1">rs6908425</td><td align="center" rowspan="1" colspan="1">C</td><td align="center" rowspan="1" colspan="1">0.78</td><td align="center" rowspan="1" colspan="1">1.41×10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.17 (1.11-1.23)</td><td align="center" rowspan="1" colspan="1">7.75×10<sup>-5</sup></td><td align="center" rowspan="1" colspan="1">1.11 (1.05-1.16)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">6q21</td><td align="center" rowspan="1" colspan="1">106.51-106.67</td><td align="center" rowspan="1" colspan="1"><italic>PRDM1</italic></td><td align="center" rowspan="1" colspan="1">rs6911490</td><td align="center" rowspan="1" colspan="1">T</td><td align="center" rowspan="1" colspan="1">0.21</td><td align="center" rowspan="1" colspan="1">4.28×10<sup>-7</sup></td><td align="center" rowspan="1" colspan="1">1.12 (1.07-1.18)</td><td align="center" rowspan="1" colspan="1">3.51×10<sup>-7</sup></td><td align="center" rowspan="1" colspan="1">1.13 (1.07-1.18)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">9p24</td><td align="center" rowspan="1" colspan="1">4.93-5.29</td><td align="center" rowspan="1" colspan="1"><italic>JAK2</italic></td><td align="center" rowspan="1" colspan="1">rs10758669</td><td align="center" rowspan="1" colspan="1">C</td><td align="center" rowspan="1" colspan="1">0.35</td><td align="center" rowspan="1" colspan="1">1.00×10<sup>-13</sup></td><td align="center" rowspan="1" colspan="1">1.18 (1.13-1.23)</td><td align="center" rowspan="1" colspan="1">8.52×10<sup>-13</sup></td><td align="center" rowspan="1" colspan="1">1.16 (1.11-1.21)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">9q32</td><td align="center" rowspan="1" colspan="1">116.48-116.74</td><td align="center" rowspan="1" colspan="1"><italic>TNFSF15</italic></td><td align="center" rowspan="1" colspan="1">rs4246905</td><td align="center" rowspan="1" colspan="1">C</td><td align="center" rowspan="1" colspan="1">0.71</td><td align="center" rowspan="1" colspan="1">1.33×10<sup>-15</sup></td><td align="center" rowspan="1" colspan="1">1.21 (1.15-1.28)</td><td align="center" rowspan="1" colspan="1">4.77×10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.13 (1.08-1.18)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">9q34</td><td align="center" rowspan="1" colspan="1">138.27-138.55</td><td align="center" rowspan="1" colspan="1"><italic>CARD9</italic></td><td align="center" rowspan="1" colspan="1">rs10781499</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.40</td><td align="center" rowspan="1" colspan="1">3.49×10<sup>-18</sup></td><td align="center" rowspan="1" colspan="1">1.20 (1.15-1.26)</td><td align="center" rowspan="1" colspan="1">6.95×10<sup>-13</sup></td><td align="center" rowspan="1" colspan="1">1.16 (1.11-1.21)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">10p11</td><td align="center" rowspan="1" colspan="1">35.22-35.94</td><td align="center" rowspan="1" colspan="1"><italic>CREM/CCNY</italic></td><td align="center" rowspan="1" colspan="1">rs12261843</td><td align="center" rowspan="1" colspan="1">G</td><td align="center" rowspan="1" colspan="1">0.29</td><td align="center" rowspan="1" colspan="1">1.87×10<sup>-9</sup></td><td align="center" rowspan="1" colspan="1">1.15 (1.10-1.20)</td><td align="center" rowspan="1" colspan="1">2.35×10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.13 (1.08-1.18)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">10q21</td><td align="center" rowspan="1" colspan="1">63.97-64.43</td><td align="center" rowspan="1" colspan="1"><italic>ZNF365</italic></td><td align="center" rowspan="1" colspan="1">rs10761659</td><td align="center" rowspan="1" colspan="1">G</td><td align="center" rowspan="1" colspan="1">0.54</td><td align="center" rowspan="1" colspan="1">4.37×10<sup>-22</sup></td><td align="center" rowspan="1" colspan="1">1.23 (1.18-1.28)</td><td align="center" rowspan="1" colspan="1">7.39×10<sup>-6</sup></td><td align="center" rowspan="1" colspan="1">1.10 (1.05-1.14)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">10q24</td><td align="center" rowspan="1" colspan="1">101.25-101.33</td><td align="center" rowspan="1" colspan="1"><italic>NKX2.3</italic></td><td align="center" rowspan="1" colspan="1">rs6584283</td><td align="center" rowspan="1" colspan="1">T</td><td align="center" rowspan="1" colspan="1">0.47</td><td align="center" rowspan="1" colspan="1">7.18×10<sup>-20</sup></td><td align="center" rowspan="1" colspan="1">1.21 (1.16-1.27)</td><td align="center" rowspan="1" colspan="1">8.46×10<sup>-21</sup></td><td align="center" rowspan="1" colspan="1">1.21 (1.16-1.26)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">11q13</td><td align="center" rowspan="1" colspan="1">75.72-76.02</td><td align="center" rowspan="1" colspan="1"><italic>C11Orf30</italic></td><td align="center" rowspan="1" colspan="1">rs2155219</td><td align="center" rowspan="1" colspan="1">T</td><td align="center" rowspan="1" colspan="1">0.50</td><td align="center" rowspan="1" colspan="1">1.58×10<sup>-12</sup></td><td align="center" rowspan="1" colspan="1">1.16 (1.11-1.21)</td><td align="center" rowspan="1" colspan="1">6.33×10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.12 (1.07-1.16)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">15q22</td><td align="center" rowspan="1" colspan="1">65.2-65.27</td><td align="center" rowspan="1" colspan="1"><italic>SMAD3</italic></td><td align="center" rowspan="1" colspan="1">rs17293632</td><td align="center" rowspan="1" colspan="1">T</td><td align="center" rowspan="1" colspan="1">0.24</td><td align="center" rowspan="1" colspan="1">1.41×10<sup>-13</sup></td><td align="center" rowspan="1" colspan="1">1.19 (1.14-1.25)</td><td align="center" rowspan="1" colspan="1">9.52×10<sup>-6</sup></td><td align="center" rowspan="1" colspan="1">1.11 (1.06-1.16)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">17q12</td><td align="center" rowspan="1" colspan="1">34.62-35.51</td><td align="center" rowspan="1" colspan="1"><italic>ORMDL3</italic></td><td align="center" rowspan="1" colspan="1">rs2872507</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.46</td><td align="center" rowspan="1" colspan="1">1.51×10<sup>-9</sup></td><td align="center" rowspan="1" colspan="1">1.14 (1.09-1.19)</td><td align="center" rowspan="1" colspan="1">5.44×10<sup>-11</sup></td><td align="center" rowspan="1" colspan="1">1.15 (1.10-1.19)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">18p11</td><td align="center" rowspan="1" colspan="1">12.73-12.92</td><td align="center" rowspan="1" colspan="1"><italic>PTPN2</italic></td><td align="center" rowspan="1" colspan="1">rs1893217</td><td align="center" rowspan="1" colspan="1">G</td><td align="center" rowspan="1" colspan="1">0.16</td><td align="center" rowspan="1" colspan="1">1.29×10<sup>-14</sup></td><td align="center" rowspan="1" colspan="1">1.25 (1.18-1.32)</td><td align="center" rowspan="1" colspan="1">4.78×10<sup>-5</sup></td><td align="center" rowspan="1" colspan="1">1.12 (1.06-1.18)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">19p13</td><td align="center" rowspan="1" colspan="1">10.26-10.5</td><td align="center" rowspan="1" colspan="1"><italic>TYK2</italic></td><td align="center" rowspan="1" colspan="1">rs12720356</td><td align="center" rowspan="1" colspan="1">C</td><td align="center" rowspan="1" colspan="1">0.08</td><td align="center" rowspan="1" colspan="1">9.20×10<sup>-10</sup></td><td align="center" rowspan="1" colspan="1">1.22 (1.14-1.31)</td><td align="center" rowspan="1" colspan="1">3.90×10<sup>-6</sup></td><td align="center" rowspan="1" colspan="1">1.17 (1.09-1.26)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">19q13</td><td align="center" rowspan="1" colspan="1">38.42-38.47</td><td align="center" rowspan="1" colspan="1">-</td><td align="center" rowspan="1" colspan="1">rs736289</td><td align="center" rowspan="1" colspan="1">T</td><td align="center" rowspan="1" colspan="1">0.61</td><td align="center" rowspan="1" colspan="1">2.69×10<sup>-7</sup></td><td align="center" rowspan="1" colspan="1">1.11 (1.06-1.16)</td><td align="center" rowspan="1" colspan="1">1.89×10<sup>-5</sup></td><td align="center" rowspan="1" colspan="1">1.08 (1.03-1.12)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">20q13</td><td align="center" rowspan="1" colspan="1">61.66-61.98</td><td align="center" rowspan="1" colspan="1"><italic>RTEL1/SLC2A4RG</italic></td><td align="center" rowspan="1" colspan="1">rs2297441</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.76</td><td align="center" rowspan="1" colspan="1">1.83×10<sup>-11</sup></td><td align="center" rowspan="1" colspan="1">1.19 (1.13-1.25)</td><td align="center" rowspan="1" colspan="1">5.78×10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.14 (1.09-1.20)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">21q21</td><td align="center" rowspan="1" colspan="1">15.62-15.77</td><td align="center" rowspan="1" colspan="1">-</td><td align="center" rowspan="1" colspan="1">rs1297265</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.57</td><td align="center" rowspan="1" colspan="1">1.41×10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.16 (1.10-1.22)</td><td align="center" rowspan="1" colspan="1">1.73×10<sup>-7</sup></td><td align="center" rowspan="1" colspan="1">1.11 (1.06-1.16)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">21q22</td><td align="center" rowspan="1" colspan="1">39.34-39.41</td><td align="center" rowspan="1" colspan="1">-</td><td align="center" rowspan="1" colspan="1">rs2836878</td><td align="center" rowspan="1" colspan="1">G</td><td align="center" rowspan="1" colspan="1">0.74</td><td align="center" rowspan="1" colspan="1">3.22×10<sup>-6</sup></td><td align="center" rowspan="1" colspan="1">1.12 (1.06-1.17)</td><td align="center" rowspan="1" colspan="1">1.86×10<sup>-22</sup></td><td align="center" rowspan="1" colspan="1">1.25 (1.20-1.32)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">21q22</td><td align="center" rowspan="1" colspan="1">44.41-44.52</td><td align="center" rowspan="1" colspan="1"><italic>ICOSLG</italic></td><td align="center" rowspan="1" colspan="1">rs2838519</td><td align="center" rowspan="1" colspan="1">G</td><td align="center" rowspan="1" colspan="1">0.39</td><td align="center" rowspan="1" colspan="1">2.09×10<sup>-14</sup></td><td align="center" rowspan="1" colspan="1">1.18 (1.13-1.23)</td><td align="center" rowspan="1" colspan="1">2.26×10<sup>-8</sup></td><td align="center" rowspan="1" colspan="1">1.12 (1.08-1.17)</td></tr><tr><td align="left" colspan="10" valign="bottom" rowspan="1"><hr></hr></td></tr><tr><td align="center" rowspan="1" colspan="1">22q11</td><td align="center" rowspan="1" colspan="1">20.14-20.39</td><td align="center" rowspan="1" colspan="1"><italic>YDJC</italic></td><td align="center" rowspan="1" colspan="1">rs181359</td><td align="center" rowspan="1" colspan="1">A</td><td align="center" rowspan="1" colspan="1">0.19</td><td align="center" rowspan="1" colspan="1">6.31×10<sup>-13</sup></td><td align="center" rowspan="1" colspan="1">1.21 (1.15-1.28)</td><td align="center" rowspan="1" colspan="1">2.73×10<sup>-5</sup></td><td align="center" rowspan="1" colspan="1">1.11 (1.06-1.17)</td></tr></tbody></table></table-wrap></floats-group></pmc>
