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Abstract
GWAS have identified >200 risk loci for Inflammatory Bowel Disease (IBD). The majority of disease associations are known to be driven by regulatory variants. To identify the putative causative genes that are perturbed by these variants, we generate a large transcriptome data set (nine disease-relevant cell types) and identify 23,650 cis-eQTL. We show that these are determined by ∼9720 regulatory modules, of which ∼3000 operate in multiple tissues and ∼970 on multiple genes. We identify regulatory modules that drive the disease association for 63 of the 200 risk loci, and show that these are enriched in multigenic modules. Based on these analyses, we resequence 45 of the corresponding 100 candidate genes in 6600 Crohn disease (CD) cases and 5500 controls, and show with burden tests that they include likely causative genes. Our analyses indicate that ≥10-fold larger sample sizes will be required to demonstrate the causality of individual genes using this approach.
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Title
IBD risk loci are enriched in multigenic regulatory modules encompassing putative causative genes
Author
Momozawa, Yukihide 1 ; Dmitrieva, Julia 2 ; Théâtre, Emilie 2 ; Deffontaine, Valérie 2 ; Rahmouni, Souad 2 ; Charloteaux, Benoît 2 ; Crins, François 2 ; Docampo, Elisa 2 ; Elansary, Mahmoud 2 ; Ann-Stephan Gori 2 ; Lecut, Christelle 3 ; Mariman, Rob 2 ; Mni, Myriam 2 ; Oury, Cécile 3 ; Altukhov, Ilya 4 ; Alexeev, Dmitry 5 ; Aulchenko, Yuri 6 ; Amininejad, Leila 7 ; Bouma, Gerd 8 ; Hoentjen, Frank 9 ; Löwenberg, Mark 10 ; Oldenburg, Bas 11 ; Pierik, Marieke J 12 ; Andrea E vander Meulen-de Jong 13 ; C Janneke van der Woude 14 ; Visschedijk, Marijn C 15 ; Abraham, Clara 16 ; Jean-Paul Achkar 17 ; Ahmad, Tariq 18 ; Ananthakrishnan, Ashwin N 19 ; Andersen, Vibeke 20 ; Anderson, Carl A 21 ; Andrews, Jane M 22 ; Annese, Vito 23 ; Aumais, Guy 24 ; Baidoo, Leonard 25 ; Baldassano, Robert N 26 ; Bampton, Peter A 27 ; Barclay, Murray 28 ; Barrett, Jeffrey C 21 ; Bayless, Theodore M 29 ; Bethge, Johannes 30 ; Bitton, Alain 31 ; Boucher, Gabrielle 32 ; Brand, Stephan 33 ; Brandt, Berenice 30 ; Brant, Steven R 29 ; Carsten Büning 34 ; Chew, Angela 35 ; Cho, Judy H 36 ; Cleynen, Isabelle 37 ; Cohain, Ariella 38 ; Croft, Anthony 39 ; Daly, Mark J 40 ; Mauro D’Amato 41 ; Danese, Silvio 42 ; De Jong, Dirk 9 ; Goda Denapiene 43 ; Denson, Lee A 44 ; Devaney, Kathy L 45 ; Dewit, Olivier 46 ; Renata D’Inca 47 ; Dubinsky, Marla 48 ; Duerr, Richard H 49 ; Edwards, Cathryn 50 ; Ellinghaus, David 51 ; Essers, Jonah 52 ; Ferguson, Lynnette R 53 ; Festen, Eleonora A 15 ; Fleshner, Philip 54 ; Florin, Tim 55 ; Franke, Andre 51 ; Fransen, Karin 56 ; Gearry, Richard 57 ; Gieger, Christian 58 ; Jürgen Glas 59 ; Goyette, Philippe 32 ; Green, Todd 60 ; Griffiths, Anne M 61 ; Guthery, Stephen L 62 ; Hakonarson, Hakon 63 ; Halfvarson, Jonas 63 ; Hanigan, Katherine 39 ; Haritunians, Talin 54 ; Hart, Ailsa 64 ; Hawkey, Chris 65 ; Hayward, Nicholas K 66 ; Hedl, Matija 16 ; Henderson, Paul 67 ; Hu, Xinli 68 ; Huang, Hailiang 40 ; Hui, Ken Y 36 ; Imielinski, Marcin 26 ; Ippoliti, Andrew 54 ; Jonaitis, Laimas 69 ; Jostins, Luke 70 ; Karlsen, Tom H 71 ; Kennedy, Nicholas A 72 ; Mohammed Azam Khan 73 ; Kiudelis, Gediminas 69 ; Krupa Krishnaprasad 74 ; Kugathasan, Subra 75 ; Limas Kupcinskas 76 ; Latiano, Anna 77 ; Laukens, Debby 78 ; Lawrance, Ian C 79 ; Lee, James C 80 ; Lees, Charlie W 72 ; Leja, Marcis 81 ; Johan Van Limbergen 61 ; Lionetti, Paolo 82 ; Liu, Jimmy Z 21 ; Mahy, Gillian 83 ; Mansfield, John 84 ; Massey, Dunecan 80 ; Mathew, Christopher G 85 ; McGovern, Dermot P B 54 ; Milgrom, Raquel 86 ; Mitrovic, Mitja 87 ; Montgomery, Grant W 66 ; Mowat, Craig 88 ; Newman, William 73 ; Ng, Aylwin 89 ; Ng, Siew C 90 ; Sok Meng Evelyn Ng 16 ; Nikolaus, Susanna 30 ; Kaida Ning 16 ; Nöthen, Markus 91 ; Oikonomou, Ioannis 16 ; Palmieri, Orazio 77 ; Parkes, Miles 80 ; Phillips, Anne 88 ; Ponsioen, Cyriel Y 10 ; Potocnik, Urõs 92 ; Prescott, Natalie J 93 ; Proctor, Deborah D 16 ; Radford-Smith, Graham 94 ; Rahier, Jean-Francois 95 ; Raychaudhuri, Soumya 68 ; Regueiro, Miguel 25 ; Rieder, Florian 96 ; Rioux, John D 97 ; Ripke, Stephan 40 ; Roberts, Rebecca 28 ; Russell, Richard K 98 ; Sanderson, Jeremy D 99 ; Sans, Miquel 100 ; Satsangi, Jack 72 ; Schadt, Eric E 38 ; Schreiber, Stefan 101 ; Schulte, Dominik 30 ; Schumm, L Philip 102 ; Regan, Scott 25 ; Seielstad, Mark 103 ; Sharma, Yashoda 16 ; Silverberg, Mark S 86 ; Simms, Lisa A 39 ; Skieceviciene, Jurgita 69 ; Spain, Sarah L 104 ; A Hillary Steinhart 86 ; Stempak, Joanne M 86 ; Stronati, Laura 105 ; Sventoraityte, Jurgita 74 ; Targan, Stephan R 54 ; Taylor, Kirstin M 99 ; Anje ter Velde 10 ; Torkvist, Leif 106 ; Tremelling, Mark 107 ; Suzanne van Sommeren 15 ; Vasiliauskas, Eric 54 ; Verspaget, Hein W 13 ; Walters, Thomas 108 ; Wang, Kai 26 ; Wang, Ming-Hsi 109 ; Wei, Zhi 110 ; Whiteman, David 66 ; Wijmenga, Cisca 56 ; Wilson, David C 67 ; Winkelmann, Juliane 111 ; Xavier, Ramnik J 112 ; Zhang, Bin 38 ; Zhang, Clarence K 113 ; Zhang, Hu 114 ; Zhang, Wei 16 ; Zhao, Hongyu 113 ; Zhao, Zhen Z 66 ; Lathrop, Mark 115 ; Hugot, Jean-Pierre 116 ; Weersma, Rinse K 15 ; De Vos, Martine 78 ; Franchimont, Denis 7 ; Vermeire, Severine 37 ; Kubo, Michiaki 117 ; Louis, Edouard 118 ; Georges, Michel 2
1 Unit of Animal Genomics, WELBIO, GIGA-R & Faculty of Veterinary Medicine, University of Liège (B34), Liège, Belgium; Laboratory for Genotyping Development, RIKEN Center for Integrative Medical Science, Yokohama, Kanagawa, Japan
2 Unit of Animal Genomics, WELBIO, GIGA-R & Faculty of Veterinary Medicine, University of Liège (B34), Liège, Belgium
3 Laboratory of Thrombosis and Hemostasis, GIGA-R, University of Liège (B34), Liège, Belgium
4 Moscow Institute of Physics and Technology, Dolgoprudny, Russian Federation
5 Novosibirsk State University, Novosibirsk, Russian Federation
6 PolyOmica, ‘s-Hertogenbosch, The Netherlands; Institute of Cytology and Genetics SD RAS, Novosibirsk, Russia; Centre for Global Health Research, Usher Institute of Population Health Sciences and Informatics, University of Edinburgh, Teviot Place, Edinburgh, UK
7 Gastroentérologie Médicale, Faculté de Médicine, Université Libre de Bruxelles, Anderlecht, Belgium
8 Department of Gastroenterology and Hepatology, VU University Medical Centre, Amsterdam, The Netherlands
9 Department of Gastroenterology and Hepatology, University Medical Centre St. Radboud, Nijmegen, The Netherlands
10 Department of Gastroenterology and Hepatology, Amsterdam Medical Centre, Amsterdam, The Netherlands
11 Department of Gastroenterology and Hepatology, University Medical Centre Utrecht, Utrecht, The Netherlands
12 Department of Gastroenterology and Hepatology, University Medical Centre Maastricht, Maastricht, The Netherlands
13 Department of Gastroenterology and Hepatology, Leiden University Medical Centre, Leiden, The Netherlands
14 Department of Gastroenterology and Hepatology, Erasmus Medical Centre, Rotterdam, The Netherlands
15 Department of Gastroenterology and Hepatology, University of Groningen and University Medical Center Groningen, Groningen, The Netherlands
16 Section of Digestive Diseases, Department of Internal Medicine, Yale School of Medicine, New Haven, CT, USA
17 Department of Gastroenterology and Hepatology, Digestive Disease Institute, Cleveland Clinic, Cleveland, OH, USA; Department of Pathobiology, Lerner Research Institute, Cleveland Clinic, Cleveland, OH, USA
18 Peninsula College of Medicine and Dentistry, Exeter, UK
19 Gastroenterology Unit, Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA; Division of Medical Sciences, Harvard Medical School, Boston, MA, USA
20 Focused Research Unit for Molecular Diagnostic and Clinical Research (MOK), IRS-Center Sonderjylland, Hospital of Southern Jutland, Åbenrå, Denmark; Institute of Molecular Medicine, University of Southern Denmark, Odense, Denmark; Institute of Regional Health Research, University of Southern Denmark, Odense, Denmark
21 Wellcome Trust Sanger Institute, Wellcome Genome Campus, Hinxton, Cambridgeshire, UK
22 Inflammatory Bowel Disease Service, Department of Gastroenterology and Hepatology, Royal Adelaide Hospital, Adelaide, Australia
23 Unit of Gastroenterology, Istituto di Ricovero e Cura a Carattere Scientifico-Casa Sollievo della Sofferenza (IRCCS-CSS) Hospital, San Giovanni Rotondo, Italy; Strutture Organizzative Dipartimentali (SOD) Gastroenterologia 2, Azienda Ospedaliero Universitaria (AOU) Careggi, Florence, Italy
24 Facult de Médecine, Universit de Montréal, Montréal, QC, Canada; Department of Gastroenterology, Hôpital Maisonneuve-Rosemont, Montréal, QC, Canada
25 Division of Gastroenterology, Hepatology and Nutrition, Department of Medicine, University of Pittsburgh School of Medicine, Pittsburgh, PA, USA
26 Center for Applied Genomics, The Children’s Hospital of Philadelphia, Philadelphia, PA, USA
27 Department of Gastroenterology and Hepatology, Flinders Medical Centre and School of Medicine, Flinders University, Adelaide, Australia
28 Department of Medicine, University of Otago, Christchurch, New Zealand
29 Meyerhoff Inflammatory Bowel Disease Center, Department of Medicine, Johns Hopkins University School of Medicine, Baltimore, MD, USA
30 Department for General Internal Medicine, Christian-Albrechts-University, Kiel, Germany
31 Division of Gastroenterology, Royal Victoria Hospital, Montréal, QC, Canada
32 Research Center, Montreal Heart Institute, Montréal, QC, Canada
33 Department of Medicine II, Ludwig-Maximilians-University Hospital Munich-Grosshadern, Munich, Germany
34 Department of Gastroenterology, Campus Charité Mitte, Universitatsmedizin Berlin, Berlin, Germany
35 Harry Perkins Institute for Medical Research, School of Medicine and Pharmacology, University of Western Australia, Murdoch, WA, Australia; IBD Unit, Fremantle Hospital, Fremantle, Australia
36 Department of Genetics, Yale School of Medicine, New Haven, CT, USA
37 Translational Research in Gastrointestinal Disorders, Department of Clinical and Experimental Medicine, KU Leuven, Leuven, Belgium
38 Department of Genetics and Genomic Sciences, Mount Sinai School of Medicine, New York, NY, USA
39 Inflammatory Bowel Diseases, Genetics and Computational Biology, Queensland Institute of Medical Research, Brisbane, Australia
40 Analytic and Translational Genetics Unit, Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA; Broad Institute of MIT and Harvard, Cambridge, MA, USA
41 Clinical Epidemiology Unit, Department of Medicine Solna, Karolinska Institutet, Stockholm, Sweden; Department of Gastrointestinal and Liver Diseases, BioDonostia Health Research Institute, San Sebastián, Spain; IKERBASQUE, Basque Foundation for Science, Bilbao, Spain
42 IBD Center, Department of Gastroenterology, Istituto Clinico Humanitas, Milan, Italy
43 Center of Hepatology, Gastroenterology and Dietetics, Vilnius University, Vilnius, Lithuania
44 Pediatric Gastroenterology, Cincinnati Children’s Hospital Medical Center, Cincinnati, OH, USA
45 Gastroenterology Unit, Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA
46 Department of Gastroenterology, Université Catholique de Louvain (UCL) Cliniques Universitaires Saint-Luc, Brussels, Belgium
47 Division of Gastroenterology, University Hospital Padua, Padua, Italy
48 Department of Pediatrics, Cedars Sinai Medical Center, Los Angeles, CA, USA
49 Division of Gastroenterology, Hepatology and Nutrition, Department of Medicine, University of Pittsburgh School of Medicine, Pittsburgh, PA, USA; Department of Human Genetics, University of Pittsburgh Graduate School of Public Health, Pittsburgh, PA, USA
50 Department of Gastroenterology, Torbay Hospital, Torbay, Devon, UK
51 Institute of Clinical Molecular Biology, Christian-Albrechts-University of Kiel, Kiel, Germany
52 Center for Human Genetic Research, Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA; Pediatrics, Harvard Medical School, Boston, MA, USA
53 Faculty of Medical & Health Sciences, School of Medical Sciences, The University of Auckland, Auckland, New Zealand
54 F. Widjaja Foundation Inflammatory Bowel and Immunobiology Research Institute, Cedars-Sinai Medical Center, Los Angeles, CA, USA
55 Department of Gastroenterology, Mater Health Services, Brisbane, Australia
56 Department of Genetics, University Medical Center Groningen, Groningen, The Netherlands
57 Department of Medicine, University of Otago, Christchurch, New Zealand; Department of Gastroenterology, Christchurch Hospital, Christchurch, New Zealand
58 Institute of Genetic Epidemiology, Helmholtz Zentrum München—German Research Center for Environmental Health, Neuherberg, Germany
59 Department of Medicine II, Ludwig-Maximilians-University Hospital Munich-Grosshadern, Munich, Germany; Department of Preventive Dentistry and Periodontology, Ludwig-Maximilians-University Hospital Munich-Grosshadern, Munich, Germany
60 Broad Institute of MIT and Harvard, Cambridge, MA, USA; Center for Human Genetic Research, Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA
61 Division of Pediatric Gastroenterology, Hepatology and Nutrition, Hospital for Sick Children, Toronto, ON, Canada
62 Department of Pediatrics, University of Utah School of Medicine, Salt Lake City, UT, USA
63 Department of Gastroenterology, Faculty of Medicine and Health, Örebro University, Örebro, Sweden
64 Department of Medicine, St. Mark’s Hospital, Harrow, Middlesex, UK
65 Nottingham Digestive Diseases Centre, Queens Medical Centre, Nottingham, UK
66 Molecular Epidemiology, Genetics and Computational Biology, Queensland Institute of Medical Research, Brisbane, Australia
67 Paediatric Gastroenterology and Nutrition, Royal Hospital for Sick Children, Edinburgh, UK; Child Life and Health, University of Edinburgh, Edinburgh, Scotland, UK
68 Division of Rheumatology Immunology and Allergy, Brigham and Women’s Hospital, Boston, MA, USA
69 Academy of Medicine, Lithuanian University of Health Sciences, Kaunas, Lithuania
70 Wellcome Trust Centre for Human Genetics, University of Oxford, Headington, UK; Christ Church, University of Oxford, St Aldates, UK
71 Research Institute of Internal Medicine, Department of Transplantation Medicine, Division of Cancer, Surgery and Transplantation, Oslo University Hospital Rikshospitalet, Oslo, Norway; Norwegian PSC Research Center, Department of Transplantation Medicine, Division of Cancer, Surgery and Transplantation, Oslo University Hospital Rikshospitalet, Oslo, Norway; K.G. Jebsen Inflammation Research Centre, Institute of Clinical Medicine, University of Oslo, Oslo, Norway
72 Gastrointestinal Unit, Western General Hospital University of Edinburgh, Edinburgh, UK
73 Genetic Medicine, Manchester Academic Health Science Centre, Manchester, UK; The Manchester Centre for Genomic Medicine, University of Manchester, Manchester, UK
74 QIMR Berghofer Medical Research Institute, Royal Brisbane Hospital, Brisbane, Australia
75 Department of Pediatrics, Emory University School of Medicine, Atlanta, GA, USA
76 Department of Gastroenterology, Kaunas University of Medicine, Kaunas, Lithuania
77 Unit of Gastroenterology, Istituto di Ricovero e Cura a Carattere Scientifico-Casa Sollievo della Sofferenza (IRCCS-CSS) Hospital, San Giovanni Rotondo, Italy
78 Department of Gastroenterology, University Hospital, Gent, Belgium
79 Harry Perkins Institute for Medical Research, School of Medicine and Pharmacology, University of Western Australia, Murdoch, WA, Australia; Centre for Inflammatory Bowel Diseases, Saint John of God Hospital, Subiaco, WA, Australia
80 Inflammatory Bowel Disease Research Group, Addenbrooke’s Hospital, Cambridge, UK
81 Faculty of medicine, University of Latvia, Riga, Latvia
82 Dipartimento di Neuroscienze, Psicologia, Area del Farmaco e Salute del Bambino, Universitê di Firenze Strutture Organizzative Dipartimentali (SOD) Gastroenterologia e Nutrizione Ospedale Pediatrico Meyer, Firenze, Italy
83 Department of Gastroenterology, The Townsville Hospital, Townsville, Australia
84 Institute of Human Genetics, Newcastle University, Newcastle upon Tyne, UK
85 Department of Medical and Molecular Genetics, King’s College London, London, UK; Sydney Brenner Institute for Molecular Bioscience, University of the Witwatersrand, Johannesburg, South Africa
86 Inflammatory Bowel Disease Centre, Mount Sinai Hospital, Toronto, ON, Canada
87 Department of Genetics, University Medical Center Groningen, Groningen, The Netherlands; Center for Human Molecular Genetics and Pharmacogenomics, Faculty of Medicine, University of Maribor, Maribor, Slovenia
88 Department of Medicine, Ninewells Hospital and Medical School, Dundee, UK
89 Gastroenterology Unit, Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA; Center for Computational and Integrative Biology, Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA
90 Department of Medicine and Therapeutics, Institute of Digestive Disease, Chinese University of Hong Kong, Hong Kong, Hong Kong
91 Department of Genomics Life & Brain Center, University Hospital Bonn, Bonn, Germany
92 Center for Human Molecular Genetics and Pharmacogenomics, Faculty of Medicine, University of Maribor, Maribor, Slovenia; Faculty for Chemistry and Chemical Engineering, University of Maribor, Maribor, Slovenia
93 Department of Medical and Molecular Genetics, King’s College London, London, UK
94 Inflammatory Bowel Diseases, Genetics and Computational Biology, Queensland Institute of Medical Research, Brisbane, Australia; Department of Gastroenterology, Royal Brisbane and Womens Hospital, Brisbane, Australia
95 Department of Gastroenterology, Université Catholique de Louvain (UCL) Centre Hospitalier Universitaire (CHU) Mont-Godinne, Mont-Godinne, Belgium
96 Department of Gastroenterology and Hepatology, Digestive Disease Institute, Cleveland Clinic, Cleveland, OH, USA
97 Facult de Médecine, Universit de Montréal, Montréal, QC, Canada; Research Center, Montreal Heart Institute, Montréal, QC, Canada
98 Paediatric Gastroenterology and Nutrition, Royal Hospital for Sick Children, Edinburgh, UK
99 Department of Gastroenterology, Guy’s & St. Thomas’ NHS Foundation Trust, St.-Thomas Hospital, London, UK
100 Department of Digestive Diseases, Hospital Quiron Teknon, Barcelona, Spain
101 Department for General Internal Medicine, Christian-Albrechts-University, Kiel, Germany; Institute of Clinical Molecular Biology, Christian-Albrechts-University of Kiel, Kiel, Germany
102 Department of Public Health Sciences, University of Chicago, Chicago, IL, USA
103 Human Genetics, Genome Institute of Singapore, Singapore, Singapore; Institute for Human Genetics, University of California, San Francisco, CA, USA
104 Wellcome Trust Sanger Institute, Wellcome Genome Campus, Hinxton, Cambridgeshire, UK; Open Targets, Wellcome Trust Genome Campus, Hinxton, Cambridgeshire, UK
105 Department of Biology of Radiations and Human Health, Agenzia Nazionale per le Nuove Tecnologie l’energia e lo Sviluppo Economico Sostenibile (ENEA), Rome, Italy
106 Department of Clinical Science Intervention and Technology, Karolinska Institutet, Stockholm, Sweden
107 Gastroenterology & General Medicine, Norfolk and Norwich University Hospital, Norwich, UK
108 Division of Pediatric Gastroenterology, Hepatology and Nutrition, Hospital for Sick Children, Toronto, ON, Canada; Faculty of Medicine, University of Toronto, Toronto, ON, Canada
109 Department of Gastroenterology and Hepatology, Digestive Disease Institute, Cleveland Clinic, Cleveland, OH, USA; Meyerhoff Inflammatory Bowel Disease Center, Department of Medicine, Johns Hopkins University School of Medicine, Baltimore, MD, USA
110 Department of Computer Science, New Jersey Institute of Technology, Newark, NJ, USA
111 Institute of Human Genetics, Technische Universität München, Munich, Germany; Department of Neurology, Technische Universität München, Munich, Germany
112 Gastroenterology Unit, Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA; Broad Institute of MIT and Harvard, Cambridge, MA, USA
113 Department of Biostatistics, School of Public Health, Yale University, New Haven, CT, USA
114 Department of Gastroenterology, West China Hospital, Chengdu, Sichuan, China; State Key Laboratory of Biotherapy, Sichuan University West China University of Medical Sciences (WCUMS), Chengdu, Sichuan, China
115 McGill University Centre for Molecular and Computational Genomics, Montreal, QC, Canada
116 UMR 1149 INSERM/Université Paris-Diderot Sorbonne Paris-Cité, Assistance Publique Hôpitaux de Paris, Paris, France
117 Laboratory for Genotyping Development, RIKEN Center for Integrative Medical Science, Yokohama, Kanagawa, Japan
118 CHU-Liège and Unit of Gastroenterology, GIGA-R & Faculty of Medicine, University of Liège, Liège, Belgium
1 Unit of Animal Genomics, WELBIO, GIGA-R & Faculty of Veterinary Medicine, University of Liège (B34), Liège, Belgium; Laboratory for Genotyping Development, RIKEN Center for Integrative Medical Science, Yokohama, Kanagawa, Japan
2 Unit of Animal Genomics, WELBIO, GIGA-R & Faculty of Veterinary Medicine, University of Liège (B34), Liège, Belgium
3 Laboratory of Thrombosis and Hemostasis, GIGA-R, University of Liège (B34), Liège, Belgium
4 Moscow Institute of Physics and Technology, Dolgoprudny, Russian Federation
5 Novosibirsk State University, Novosibirsk, Russian Federation
6 PolyOmica, ‘s-Hertogenbosch, The Netherlands; Institute of Cytology and Genetics SD RAS, Novosibirsk, Russia; Centre for Global Health Research, Usher Institute of Population Health Sciences and Informatics, University of Edinburgh, Teviot Place, Edinburgh, UK
7 Gastroentérologie Médicale, Faculté de Médicine, Université Libre de Bruxelles, Anderlecht, Belgium
8 Department of Gastroenterology and Hepatology, VU University Medical Centre, Amsterdam, The Netherlands
9 Department of Gastroenterology and Hepatology, University Medical Centre St. Radboud, Nijmegen, The Netherlands
10 Department of Gastroenterology and Hepatology, Amsterdam Medical Centre, Amsterdam, The Netherlands
11 Department of Gastroenterology and Hepatology, University Medical Centre Utrecht, Utrecht, The Netherlands
12 Department of Gastroenterology and Hepatology, University Medical Centre Maastricht, Maastricht, The Netherlands
13 Department of Gastroenterology and Hepatology, Leiden University Medical Centre, Leiden, The Netherlands
14 Department of Gastroenterology and Hepatology, Erasmus Medical Centre, Rotterdam, The Netherlands
15 Department of Gastroenterology and Hepatology, University of Groningen and University Medical Center Groningen, Groningen, The Netherlands
16 Section of Digestive Diseases, Department of Internal Medicine, Yale School of Medicine, New Haven, CT, USA
17 Department of Gastroenterology and Hepatology, Digestive Disease Institute, Cleveland Clinic, Cleveland, OH, USA; Department of Pathobiology, Lerner Research Institute, Cleveland Clinic, Cleveland, OH, USA
18 Peninsula College of Medicine and Dentistry, Exeter, UK
19 Gastroenterology Unit, Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA; Division of Medical Sciences, Harvard Medical School, Boston, MA, USA
20 Focused Research Unit for Molecular Diagnostic and Clinical Research (MOK), IRS-Center Sonderjylland, Hospital of Southern Jutland, Åbenrå, Denmark; Institute of Molecular Medicine, University of Southern Denmark, Odense, Denmark; Institute of Regional Health Research, University of Southern Denmark, Odense, Denmark
21 Wellcome Trust Sanger Institute, Wellcome Genome Campus, Hinxton, Cambridgeshire, UK
22 Inflammatory Bowel Disease Service, Department of Gastroenterology and Hepatology, Royal Adelaide Hospital, Adelaide, Australia
23 Unit of Gastroenterology, Istituto di Ricovero e Cura a Carattere Scientifico-Casa Sollievo della Sofferenza (IRCCS-CSS) Hospital, San Giovanni Rotondo, Italy; Strutture Organizzative Dipartimentali (SOD) Gastroenterologia 2, Azienda Ospedaliero Universitaria (AOU) Careggi, Florence, Italy
24 Facult de Médecine, Universit de Montréal, Montréal, QC, Canada; Department of Gastroenterology, Hôpital Maisonneuve-Rosemont, Montréal, QC, Canada
25 Division of Gastroenterology, Hepatology and Nutrition, Department of Medicine, University of Pittsburgh School of Medicine, Pittsburgh, PA, USA
26 Center for Applied Genomics, The Children’s Hospital of Philadelphia, Philadelphia, PA, USA
27 Department of Gastroenterology and Hepatology, Flinders Medical Centre and School of Medicine, Flinders University, Adelaide, Australia
28 Department of Medicine, University of Otago, Christchurch, New Zealand
29 Meyerhoff Inflammatory Bowel Disease Center, Department of Medicine, Johns Hopkins University School of Medicine, Baltimore, MD, USA
30 Department for General Internal Medicine, Christian-Albrechts-University, Kiel, Germany
31 Division of Gastroenterology, Royal Victoria Hospital, Montréal, QC, Canada
32 Research Center, Montreal Heart Institute, Montréal, QC, Canada
33 Department of Medicine II, Ludwig-Maximilians-University Hospital Munich-Grosshadern, Munich, Germany
34 Department of Gastroenterology, Campus Charité Mitte, Universitatsmedizin Berlin, Berlin, Germany
35 Harry Perkins Institute for Medical Research, School of Medicine and Pharmacology, University of Western Australia, Murdoch, WA, Australia; IBD Unit, Fremantle Hospital, Fremantle, Australia
36 Department of Genetics, Yale School of Medicine, New Haven, CT, USA
37 Translational Research in Gastrointestinal Disorders, Department of Clinical and Experimental Medicine, KU Leuven, Leuven, Belgium
38 Department of Genetics and Genomic Sciences, Mount Sinai School of Medicine, New York, NY, USA
39 Inflammatory Bowel Diseases, Genetics and Computational Biology, Queensland Institute of Medical Research, Brisbane, Australia
40 Analytic and Translational Genetics Unit, Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA; Broad Institute of MIT and Harvard, Cambridge, MA, USA
41 Clinical Epidemiology Unit, Department of Medicine Solna, Karolinska Institutet, Stockholm, Sweden; Department of Gastrointestinal and Liver Diseases, BioDonostia Health Research Institute, San Sebastián, Spain; IKERBASQUE, Basque Foundation for Science, Bilbao, Spain
42 IBD Center, Department of Gastroenterology, Istituto Clinico Humanitas, Milan, Italy
43 Center of Hepatology, Gastroenterology and Dietetics, Vilnius University, Vilnius, Lithuania
44 Pediatric Gastroenterology, Cincinnati Children’s Hospital Medical Center, Cincinnati, OH, USA
45 Gastroenterology Unit, Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA
46 Department of Gastroenterology, Université Catholique de Louvain (UCL) Cliniques Universitaires Saint-Luc, Brussels, Belgium
47 Division of Gastroenterology, University Hospital Padua, Padua, Italy
48 Department of Pediatrics, Cedars Sinai Medical Center, Los Angeles, CA, USA
49 Division of Gastroenterology, Hepatology and Nutrition, Department of Medicine, University of Pittsburgh School of Medicine, Pittsburgh, PA, USA; Department of Human Genetics, University of Pittsburgh Graduate School of Public Health, Pittsburgh, PA, USA
50 Department of Gastroenterology, Torbay Hospital, Torbay, Devon, UK
51 Institute of Clinical Molecular Biology, Christian-Albrechts-University of Kiel, Kiel, Germany
52 Center for Human Genetic Research, Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA; Pediatrics, Harvard Medical School, Boston, MA, USA
53 Faculty of Medical & Health Sciences, School of Medical Sciences, The University of Auckland, Auckland, New Zealand
54 F. Widjaja Foundation Inflammatory Bowel and Immunobiology Research Institute, Cedars-Sinai Medical Center, Los Angeles, CA, USA
55 Department of Gastroenterology, Mater Health Services, Brisbane, Australia
56 Department of Genetics, University Medical Center Groningen, Groningen, The Netherlands
57 Department of Medicine, University of Otago, Christchurch, New Zealand; Department of Gastroenterology, Christchurch Hospital, Christchurch, New Zealand
58 Institute of Genetic Epidemiology, Helmholtz Zentrum München—German Research Center for Environmental Health, Neuherberg, Germany
59 Department of Medicine II, Ludwig-Maximilians-University Hospital Munich-Grosshadern, Munich, Germany; Department of Preventive Dentistry and Periodontology, Ludwig-Maximilians-University Hospital Munich-Grosshadern, Munich, Germany
60 Broad Institute of MIT and Harvard, Cambridge, MA, USA; Center for Human Genetic Research, Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA
61 Division of Pediatric Gastroenterology, Hepatology and Nutrition, Hospital for Sick Children, Toronto, ON, Canada
62 Department of Pediatrics, University of Utah School of Medicine, Salt Lake City, UT, USA
63 Department of Gastroenterology, Faculty of Medicine and Health, Örebro University, Örebro, Sweden
64 Department of Medicine, St. Mark’s Hospital, Harrow, Middlesex, UK
65 Nottingham Digestive Diseases Centre, Queens Medical Centre, Nottingham, UK
66 Molecular Epidemiology, Genetics and Computational Biology, Queensland Institute of Medical Research, Brisbane, Australia
67 Paediatric Gastroenterology and Nutrition, Royal Hospital for Sick Children, Edinburgh, UK; Child Life and Health, University of Edinburgh, Edinburgh, Scotland, UK
68 Division of Rheumatology Immunology and Allergy, Brigham and Women’s Hospital, Boston, MA, USA
69 Academy of Medicine, Lithuanian University of Health Sciences, Kaunas, Lithuania
70 Wellcome Trust Centre for Human Genetics, University of Oxford, Headington, UK; Christ Church, University of Oxford, St Aldates, UK
71 Research Institute of Internal Medicine, Department of Transplantation Medicine, Division of Cancer, Surgery and Transplantation, Oslo University Hospital Rikshospitalet, Oslo, Norway; Norwegian PSC Research Center, Department of Transplantation Medicine, Division of Cancer, Surgery and Transplantation, Oslo University Hospital Rikshospitalet, Oslo, Norway; K.G. Jebsen Inflammation Research Centre, Institute of Clinical Medicine, University of Oslo, Oslo, Norway
72 Gastrointestinal Unit, Western General Hospital University of Edinburgh, Edinburgh, UK
73 Genetic Medicine, Manchester Academic Health Science Centre, Manchester, UK; The Manchester Centre for Genomic Medicine, University of Manchester, Manchester, UK
74 QIMR Berghofer Medical Research Institute, Royal Brisbane Hospital, Brisbane, Australia
75 Department of Pediatrics, Emory University School of Medicine, Atlanta, GA, USA
76 Department of Gastroenterology, Kaunas University of Medicine, Kaunas, Lithuania
77 Unit of Gastroenterology, Istituto di Ricovero e Cura a Carattere Scientifico-Casa Sollievo della Sofferenza (IRCCS-CSS) Hospital, San Giovanni Rotondo, Italy
78 Department of Gastroenterology, University Hospital, Gent, Belgium
79 Harry Perkins Institute for Medical Research, School of Medicine and Pharmacology, University of Western Australia, Murdoch, WA, Australia; Centre for Inflammatory Bowel Diseases, Saint John of God Hospital, Subiaco, WA, Australia
80 Inflammatory Bowel Disease Research Group, Addenbrooke’s Hospital, Cambridge, UK
81 Faculty of medicine, University of Latvia, Riga, Latvia
82 Dipartimento di Neuroscienze, Psicologia, Area del Farmaco e Salute del Bambino, Universitê di Firenze Strutture Organizzative Dipartimentali (SOD) Gastroenterologia e Nutrizione Ospedale Pediatrico Meyer, Firenze, Italy
83 Department of Gastroenterology, The Townsville Hospital, Townsville, Australia
84 Institute of Human Genetics, Newcastle University, Newcastle upon Tyne, UK
85 Department of Medical and Molecular Genetics, King’s College London, London, UK; Sydney Brenner Institute for Molecular Bioscience, University of the Witwatersrand, Johannesburg, South Africa
86 Inflammatory Bowel Disease Centre, Mount Sinai Hospital, Toronto, ON, Canada
87 Department of Genetics, University Medical Center Groningen, Groningen, The Netherlands; Center for Human Molecular Genetics and Pharmacogenomics, Faculty of Medicine, University of Maribor, Maribor, Slovenia
88 Department of Medicine, Ninewells Hospital and Medical School, Dundee, UK
89 Gastroenterology Unit, Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA; Center for Computational and Integrative Biology, Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA
90 Department of Medicine and Therapeutics, Institute of Digestive Disease, Chinese University of Hong Kong, Hong Kong, Hong Kong
91 Department of Genomics Life & Brain Center, University Hospital Bonn, Bonn, Germany
92 Center for Human Molecular Genetics and Pharmacogenomics, Faculty of Medicine, University of Maribor, Maribor, Slovenia; Faculty for Chemistry and Chemical Engineering, University of Maribor, Maribor, Slovenia
93 Department of Medical and Molecular Genetics, King’s College London, London, UK
94 Inflammatory Bowel Diseases, Genetics and Computational Biology, Queensland Institute of Medical Research, Brisbane, Australia; Department of Gastroenterology, Royal Brisbane and Womens Hospital, Brisbane, Australia
95 Department of Gastroenterology, Université Catholique de Louvain (UCL) Centre Hospitalier Universitaire (CHU) Mont-Godinne, Mont-Godinne, Belgium
96 Department of Gastroenterology and Hepatology, Digestive Disease Institute, Cleveland Clinic, Cleveland, OH, USA
97 Facult de Médecine, Universit de Montréal, Montréal, QC, Canada; Research Center, Montreal Heart Institute, Montréal, QC, Canada
98 Paediatric Gastroenterology and Nutrition, Royal Hospital for Sick Children, Edinburgh, UK
99 Department of Gastroenterology, Guy’s & St. Thomas’ NHS Foundation Trust, St.-Thomas Hospital, London, UK
100 Department of Digestive Diseases, Hospital Quiron Teknon, Barcelona, Spain
101 Department for General Internal Medicine, Christian-Albrechts-University, Kiel, Germany; Institute of Clinical Molecular Biology, Christian-Albrechts-University of Kiel, Kiel, Germany
102 Department of Public Health Sciences, University of Chicago, Chicago, IL, USA
103 Human Genetics, Genome Institute of Singapore, Singapore, Singapore; Institute for Human Genetics, University of California, San Francisco, CA, USA
104 Wellcome Trust Sanger Institute, Wellcome Genome Campus, Hinxton, Cambridgeshire, UK; Open Targets, Wellcome Trust Genome Campus, Hinxton, Cambridgeshire, UK
105 Department of Biology of Radiations and Human Health, Agenzia Nazionale per le Nuove Tecnologie l’energia e lo Sviluppo Economico Sostenibile (ENEA), Rome, Italy
106 Department of Clinical Science Intervention and Technology, Karolinska Institutet, Stockholm, Sweden
107 Gastroenterology & General Medicine, Norfolk and Norwich University Hospital, Norwich, UK
108 Division of Pediatric Gastroenterology, Hepatology and Nutrition, Hospital for Sick Children, Toronto, ON, Canada; Faculty of Medicine, University of Toronto, Toronto, ON, Canada
109 Department of Gastroenterology and Hepatology, Digestive Disease Institute, Cleveland Clinic, Cleveland, OH, USA; Meyerhoff Inflammatory Bowel Disease Center, Department of Medicine, Johns Hopkins University School of Medicine, Baltimore, MD, USA
110 Department of Computer Science, New Jersey Institute of Technology, Newark, NJ, USA
111 Institute of Human Genetics, Technische Universität München, Munich, Germany; Department of Neurology, Technische Universität München, Munich, Germany
112 Gastroenterology Unit, Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA; Broad Institute of MIT and Harvard, Cambridge, MA, USA
113 Department of Biostatistics, School of Public Health, Yale University, New Haven, CT, USA
114 Department of Gastroenterology, West China Hospital, Chengdu, Sichuan, China; State Key Laboratory of Biotherapy, Sichuan University West China University of Medical Sciences (WCUMS), Chengdu, Sichuan, China
115 McGill University Centre for Molecular and Computational Genomics, Montreal, QC, Canada
116 UMR 1149 INSERM/Université Paris-Diderot Sorbonne Paris-Cité, Assistance Publique Hôpitaux de Paris, Paris, France
117 Laboratory for Genotyping Development, RIKEN Center for Integrative Medical Science, Yokohama, Kanagawa, Japan
118 CHU-Liège and Unit of Gastroenterology, GIGA-R & Faculty of Medicine, University of Liège, Liège, Belgium
Pages
1-18
Publication year
2018
Publication date
Jun 2018
e-ISSN
20411723
Source type
Scholarly Journal
Language of publication
English
ProQuest document ID
2058239460
Copyright
© 2018. This work is published under http://creativecommons.org/licenses/by/4.0/ (the “License”). Notwithstanding the ProQuest Terms and Conditions, you may use this content in accordance with the terms of the License.