About the Authors

Charlie W Lees

Contributed equally to this work with: Charlie W Lees, William J Zacharias

To whom correspondence should be addressed. E-mail: Charlie.lees@ed.ac.uk (CWL); dgumucio@umich.edu (DLG)

Affiliations Gastrointestinal Unit, Molecular Medicine Centre, University of Edinburgh, Edinburgh, United Kingdom , Immunobiology Group, Medical Research Council (MRC) Centre for Inflammation Research, Queens Medical Research Institute, University of Edinburgh, Edinburgh, United Kingdom

William J Zacharias

Contributed equally to this work with: Charlie W Lees, William J Zacharias

Affiliation Department of Cell and Developmental Biology, University of Michigan Medical School, Ann Arbor, Michigan, United States of America

Mark Tremelling

Affiliation Inflammatory Bowel Disease (IBD) Research Group, Addenbrooke's Hospital, University of Cambridge, Cambridge, United Kingdom

Colin L Noble

Affiliation Gastrointestinal Unit, Molecular Medicine Centre, University of Edinburgh, Edinburgh, United Kingdom

Elaine R Nimmo

Affiliation Gastrointestinal Unit, Molecular Medicine Centre, University of Edinburgh, Edinburgh, United Kingdom

Albert Tenesa

Affiliation Colon Cancer Genetics Group, MRC Human Genetics Unit, Western General Hospital, Edinburgh, United Kingdom

Jennine Cornelius

Affiliation Department of Pathology, Genentech, Inc, South San Francisco, California, United States of America

Leif Torkvist

Affiliation Department of Medical & Surgical Gastroenterology, Karolinska University Hospital, Stockholm, Sweden

John Kao

Affiliation Department of Internal Medicine, University of Michigan Medical School, Ann Arbor, Michigan, United States of America

Susan Farrington

Affiliation Colon Cancer Genetics Group, MRC Human Genetics Unit, Western General Hospital, Edinburgh, United Kingdom

Hazel E Drummond

Affiliation Gastrointestinal Unit, Molecular Medicine Centre, University of Edinburgh, Edinburgh, United Kingdom

Gwo-Tzer Ho

Affiliation Gastrointestinal Unit, Molecular Medicine Centre, University of Edinburgh, Edinburgh, United Kingdom

Ian D. R Arnott

Affiliation Gastrointestinal Unit, Molecular Medicine Centre, University of Edinburgh, Edinburgh, United Kingdom

Henry D Appelman

Affiliation Department of Pathology, University of Michigan Medical School, Ann Arbor, Michigan, United States of America

Lauri Diehl

Affiliation Department of Pathology, Genentech, Inc, South San Francisco, California, United States of America

Harry Campbell

Affiliation Colon Cancer Genetics Group, MRC Human Genetics Unit, Western General Hospital, Edinburgh, United Kingdom

Malcolm G Dunlop

Affiliation Colon Cancer Genetics Group, MRC Human Genetics Unit, Western General Hospital, Edinburgh, United Kingdom

Miles Parkes

Affiliation Inflammatory Bowel Disease (IBD) Research Group, Addenbrooke's Hospital, University of Cambridge, Cambridge, United Kingdom

Sarah E. M Howie

Affiliation Immunobiology Group, Medical Research Council (MRC) Centre for Inflammation Research, Queens Medical Research Institute, University of Edinburgh, Edinburgh, United Kingdom

Deborah L Gumucio

To whom correspondence should be addressed. E-mail: Charlie.lees@ed.ac.uk (CWL); dgumucio@umich.edu (DLG)

These authors are joint senior authors on this work.

Affiliation Department of Cell and Developmental Biology, University of Michigan Medical School, Ann Arbor, Michigan, United States of America

Jack Satsangi

These authors are joint senior authors on this work.

Affiliation Gastrointestinal Unit, Molecular Medicine Centre, University of Edinburgh, Edinburgh, United Kingdom

Competing Interests

Lauri Diehl is a paid employee of Genentech, Inc and a stockholder in that company. While this paper does not describe any Genentech product, it is possible that these findings may be relevant to a pathway or pathways of interest to the company for current or future therapeutic agents. None of the other authors have any competing interests to declare.

Author Contributions

CWL, WJZ, CLN, ERN, LD, DLG, and JS designed the experiments/the study. CWL, WJZ, MT, CLN, JC, LT, JK, HED, IDRA, HDA, LD, MP, and SEMH collected data or did experiments for the study. CWL, WJZ, MT, AT, JK, LD, and DLG analyzed the data. CWL, MT, CLN, LT, G-TH, IRDA, SF, MGD and MP enrolled patients. CWL and WJZ wrote the first draft of the paper. CWL, WJZ, MT, AT, HC, MGD, SEMH, DLG, and JS contributed to writing the paper.