About the Authors

Jared O'Connell

Contributed equally to this work with: Jared O'Connell, Deepti Gurdasani

Affiliations Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom, Department of Statistics, University of Oxford, Oxford, United Kingdom

Deepti Gurdasani

Contributed equally to this work with: Jared O'Connell, Deepti Gurdasani

Affiliations Wellcome Trust Sanger Institute, Hinxton, United Kingdom, Department of Public Health and Primary Care, University of Cambridge, Cambridge, United Kingdom

Olivier Delaneau

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

Nicola Pirastu

Affiliation Institute for Maternal and Child Health - IRCCS Burlo Garofolo, University of Trieste, Trieste, Italy

Sheila Ulivi

Affiliation Institute for Maternal and Child Health - IRCCS Burlo Garofolo, Trieste, Italy

Massimiliano Cocca

Affiliation Division of Genetics and Cell Biology, San Raffaele Scientific Institute, Milano, Italy

Michela Traglia

Affiliation Division of Genetics and Cell Biology, San Raffaele Scientific Institute, Milano, Italy

Jie Huang

Affiliation Wellcome Trust Sanger Institute, Hinxton, United Kingdom

Jennifer E. Huffman

Affiliation MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, United Kingdom

Igor Rudan

Affiliation Centre for Population Health Sciences, University of Edinburgh, Edinburgh, United Kingdom

Ruth McQuillan

Affiliation Centre for Population Health Sciences, University of Edinburgh, Edinburgh, United Kingdom

Ross M. Fraser

Affiliation Centre for Population Health Sciences, University of Edinburgh, Edinburgh, United Kingdom

Harry Campbell

Affiliation Centre for Population Health Sciences, University of Edinburgh, Edinburgh, United Kingdom

Ozren Polasek

Affiliation Faculty of Medicine, University of Split, Split, Croatia

Gershim Asiki

Affiliation Medical Research Council/Uganda Virus Research Institute (MRC/UVRI), Uganda Research Unit on AIDS, Entebbe, Uganda

Kenneth Ekoru

Affiliation Laboratoire Génomique, Bioinformatique, et Applications (EA4627), Conservatoire National des Arts et Métiers, Paris, France

Caroline Hayward

Affiliation MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, United Kingdom

Alan F. Wright

Affiliation MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, United Kingdom

Veronique Vitart

Affiliation MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, United Kingdom

Pau Navarro

Affiliation MRC Human Genetics Unit, MRC Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, United Kingdom

Jean-Francois Zagury

Affiliation Laboratoire Génomique, Bioinformatique, et Applications (EA4627), Conservatoire National des Arts et Métiers, Paris, France

James F. Wilson

Affiliation Centre for Population Health Sciences, University of Edinburgh, Edinburgh, United Kingdom

Daniela Toniolo

Affiliation Division of Genetics and Cell Biology, San Raffaele Scientific Institute, Milano, Italy

Paolo Gasparini

Affiliation Institute for Maternal and Child Health - IRCCS Burlo Garofolo, University of Trieste, Trieste, Italy

Nicole Soranzo

Affiliation Wellcome Trust Sanger Institute, Hinxton, United Kingdom

Manjinder S. Sandhu

Affiliations Wellcome Trust Sanger Institute, Hinxton, United Kingdom, Department of Public Health and Primary Care, University of Cambridge, Cambridge, United Kingdom

Jonathan Marchini

marchini@stats.ox.ac.uk

Affiliations Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, United Kingdom, Department of Statistics, University of Oxford, Oxford, United Kingdom

Competing Interests

The authors have declared that no competing interests exist.

Author Contributions

Conceived and designed the experiments: JO JM OD JFZ NS MSS PG DT JFW. Performed the experiments: JO DG. Analyzed the data: JO DG. Wrote the paper: JM JO. Provided access and support for the ORCADES, VIS, Split and Korcula datasets: JEH RMF HC CH AFW RM VV PN JH IR OP JFW. Provided access and support for the GPC dataset: DG GA KE MSS. Provided access and support for the Val Borbera, FVG and CARL cohorts: NP SU MC MT JH PG DT NS.