About the Authors

Ian P. M. Tomlinson

Contributed equally to this work with: Ian P. M. Tomlinson, Luis G. Carvajal-Carmona

iant@well.ox.ac.uk (IPMT); richard.houlston@icr.ac.uk (RSH); malcolm.dunlop@hgu.mrc.ac.uk (MGD)

Affiliation Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom

Luis G. Carvajal-Carmona

Contributed equally to this work with: Ian P. M. Tomlinson, Luis G. Carvajal-Carmona

Affiliation Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom

Sara E. Dobbins

Affiliation Section of Cancer Genetics, Institute of Cancer Research, Sutton, United Kingdom

Albert Tenesa

Affiliation Colon Cancer Genetics Group, Institute of Genetics and Molecular Medicine, University of Edinburgh and Medical Research Council Human Genetics Unit, Edinburgh, United Kingdom

Angela M. Jones

Affiliation Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom

Kimberley Howarth

Affiliation Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom

Claire Palles

Affiliation Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom

Peter Broderick

Affiliation Section of Cancer Genetics, Institute of Cancer Research, Sutton, United Kingdom

Emma E. M. Jaeger

Affiliation Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom

Susan Farrington

Affiliation Colon Cancer Genetics Group, Institute of Genetics and Molecular Medicine, University of Edinburgh and Medical Research Council Human Genetics Unit, Edinburgh, United Kingdom

Annabelle Lewis

Affiliation Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom

James G. D. Prendergast

Affiliation Colon Cancer Genetics Group, Institute of Genetics and Molecular Medicine, University of Edinburgh and Medical Research Council Human Genetics Unit, Edinburgh, United Kingdom

Alan M. Pittman

Affiliation Section of Cancer Genetics, Institute of Cancer Research, Sutton, United Kingdom

Evropi Theodoratou

Affiliation The University of Edinburgh Medical School, Edinburgh, United Kingdom

Bianca Olver

Affiliation Section of Cancer Genetics, Institute of Cancer Research, Sutton, United Kingdom

Marion Walker

Affiliation Colon Cancer Genetics Group, Institute of Genetics and Molecular Medicine, University of Edinburgh and Medical Research Council Human Genetics Unit, Edinburgh, United Kingdom

Steven Penegar

Affiliation Section of Cancer Genetics, Institute of Cancer Research, Sutton, United Kingdom

Ella Barclay

Affiliation Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom

Nicola Whiffin

Affiliation Section of Cancer Genetics, Institute of Cancer Research, Sutton, United Kingdom

Lynn Martin

Affiliation Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom

Stephane Ballereau

Affiliation Colon Cancer Genetics Group, Institute of Genetics and Molecular Medicine, University of Edinburgh and Medical Research Council Human Genetics Unit, Edinburgh, United Kingdom

Amy Lloyd

Affiliation Section of Cancer Genetics, Institute of Cancer Research, Sutton, United Kingdom

Maggie Gorman

Affiliation Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom

Steven Lubbe

Affiliation Section of Cancer Genetics, Institute of Cancer Research, Sutton, United Kingdom

The COGENT Consortium

Memberships of the consortia are provided in Text S1.

The CORGI Collaborators

Memberships of the consortia are provided in Text S1.

The EPICOLON Consortium

Memberships of the consortia are provided in Text S1.

Bryan Howie

Affiliation Department of Statistics, University of Oxford, Oxford, United Kingdom

Jonathan Marchini

Affiliation Department of Statistics, University of Oxford, Oxford, United Kingdom

Clara Ruiz-Ponte

Affiliation Galician Public Foundation of Genomic Medicine (FPGMX), Centro de Investigacion Biomedica en Red de Enfermedades Raras (CIBERER), Genomics Medicine Group, Hospital Clinico, Santiago de Compostela, University of Santiago de Compostela, Galicia, Spain

Ceres Fernandez-Rozadilla

Affiliation Galician Public Foundation of Genomic Medicine (FPGMX), Centro de Investigacion Biomedica en Red de Enfermedades Raras (CIBERER), Genomics Medicine Group, Hospital Clinico, Santiago de Compostela, University of Santiago de Compostela, Galicia, Spain

Antoni Castells

Affiliation Department of Gastroenterology, Hospital Clinic, CIBERehd, IDIBAPS, University of Barcelona, Barcelona, Catalonia, Spain

Angel Carracedo

Affiliation Galician Public Foundation of Genomic Medicine (FPGMX), Centro de Investigacion Biomedica en Red de Enfermedades Raras (CIBERER), Genomics Medicine Group, Hospital Clinico, Santiago de Compostela, University of Santiago de Compostela, Galicia, Spain

Sergi Castellvi-Bel

Affiliation Department of Gastroenterology, Hospital Clinic, CIBERehd, IDIBAPS, University of Barcelona, Barcelona, Catalonia, Spain

David Duggan

Affiliation Translational Genomics Research Institute, Phoenix, Arizona, United States of America

David Conti

Affiliation Department of Preventive Medicine, University of Southern California, Los Angeles, California, United States of America

Jean-Baptiste Cazier

Affiliation Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom

Harry Campbell

Affiliation Public Health Sciences, University of Edinburgh, Edinburgh, United Kingdom

Oliver Sieber

Affiliation Ludwig Colon Cancer Initiative Laboratory, Ludwig Institute for Cancer Research, Royal Melbourne Hospital, Parkville, Australia

Lara Lipton

Affiliation Ludwig Colon Cancer Initiative Laboratory, Ludwig Institute for Cancer Research, Royal Melbourne Hospital, Parkville, Australia

Peter Gibbs

Affiliation Ludwig Colon Cancer Initiative Laboratory, Ludwig Institute for Cancer Research, Royal Melbourne Hospital, Parkville, Australia

Nicholas G. Martin

Affiliation Genetic and Molecular Epidemiology Laboratories, Queensland Institute of Medical Research, Brisbane, Australia

Grant W. Montgomery

Affiliation Genetic and Molecular Epidemiology Laboratories, Queensland Institute of Medical Research, Brisbane, Australia

Joanne Young

Affiliation Familial Cancer Laboratory, Queensland Institute of Medical Research, Brisbane, Australia

Paul N. Baird

Affiliation Centre for Eye Research Australia, University of Melbourne, Melbourne, Australia

Steven Gallinger

Affiliation Samuel Lunenfeld Research Institute, Mount Sinai Hospital, Toronto, Canada

Polly Newcomb

Affiliation Fred Hutchinson Cancer Research Center, Seattle, Washington, United States of America

John Hopper

Affiliation Centre for Molecular, Environmental, Genetic, and Analytic Epidemiology, The University of Melbourne, Australia

Mark A. Jenkins

Affiliation Centre for Molecular, Environmental, Genetic, and Analytic Epidemiology, The University of Melbourne, Australia

Lauri A. Aaltonen

Affiliation Department of Medical Genetics, Genome-Scale Biology Research Program, Biomedicum Helsinki, University of Helsinki, Helsinki, Finland

David J. Kerr

Affiliation Department of Clinical Pharmacology, University of Oxford, Oxford, United Kingdom

Jeremy Cheadle

Affiliation Institute of Medical Genetics, School of Medicine, Cardiff University, Cardiff, United Kingdom

Paul Pharoah

Affiliation Cancer Research UK Laboratories, Strangeways Research Laboratory, Department of Oncology, University of Cambridge, Cambridge, United Kingdom

Graham Casey

Affiliation Department of Preventive Medicine, University of Southern California, Los Angeles, California, United States of America

Richard S. Houlston

iant@well.ox.ac.uk (IPMT); richard.houlston@icr.ac.uk (RSH); malcolm.dunlop@hgu.mrc.ac.uk (MGD)

Affiliation Section of Cancer Genetics, Institute of Cancer Research, Sutton, United Kingdom

Malcolm G. Dunlop

iant@well.ox.ac.uk (IPMT); richard.houlston@icr.ac.uk (RSH); malcolm.dunlop@hgu.mrc.ac.uk (MGD)

Affiliation Colon Cancer Genetics Group, Institute of Genetics and Molecular Medicine, University of Edinburgh and Medical Research Council Human Genetics Unit, Edinburgh, United Kingdom

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: IPM Tomlinson, RS Houlston, MG Dunlop. Performed the experiments: LG Carvajal-Carmona, AM Jones, K Howarth, P Broderick, EEM Jaeger, S Farrington, A Lewis, JGD Pendergast, AM Pittman, B Olver, M Walker, N Whiffin, C Ruiz-Ponte, C Fernandez-Rozadilla. Analyzed the data: IPM Tomlinson, LG Carvajal-Carmona, SE Dobbins, A Tenesa, AM Jones, C Palles, E Theodoratou, S Ballereau, A Lloyd, J-B Cazier. Contributed reagents/materials/analysis tools: S Penegar, E Barclay, L Martin, M Gorman, S Lubbe, B Howie, J Marchini, A Castells, A Carracedo, S Castellvi-Bel, D Duggan, D Conti, H Campbell, O Sieber, L Lipton, P Gibbs, NG Martin, GW Montgomery, J Young, PN Baird, P Newcomb, J Hopper, MA Jenkins, LA Aaltonen, DJ Kerr, J Cheadle, P Pharoah, G Casey, S Gallinger. Wrote the paper: IPM Tomlinson, RS Houlston, MG Dunlop.