About the Authors

Anatole Ghazalpour

Contributed equally to this work with: Anatole Ghazalpour, Sudheer Doss, Aldons J Lusis, Steve Horvath

Affiliation Department of Microbiology, Immunology, and Molecular Genetics, University of California Los Angeles, Los Angeles, California, United States of America

Sudheer Doss

Contributed equally to this work with: Anatole Ghazalpour, Sudheer Doss, Aldons J Lusis, Steve Horvath

Affiliation Department of Human Genetics, David Geffen School of Medicine, University of California Los Angeles, Los Angeles, California, United States of America

Bin Zhang

Affiliations Department of Human Genetics, David Geffen School of Medicine, University of California Los Angeles, Los Angeles, California, United States of America , Department of Biostatistics, School of Public Health, University of California Los Angeles, Los Angeles, California, United States of America

Susanna Wang

Affiliation Department of Human Genetics, David Geffen School of Medicine, University of California Los Angeles, Los Angeles, California, United States of America

Christopher Plaisier

Affiliation Department of Human Genetics, David Geffen School of Medicine, University of California Los Angeles, Los Angeles, California, United States of America

Ruth Castellanos

Affiliation Department of Microbiology, Immunology, and Molecular Genetics, University of California Los Angeles, Los Angeles, California, United States of America

Alec Brozell

Affiliation Department of Microbiology, Immunology, and Molecular Genetics, University of California Los Angeles, Los Angeles, California, United States of America

Eric E Schadt

Affiliation Rosetta Inpharmatics, Inc., Kirkland, Washington, United States of America

Thomas A Drake

Affiliation Department of Pathology and Laboratory Medicine, David Geffen School of Medicine, University of California Los Angeles, Los Angeles, California, United States of America

Aldons J Lusis

Contributed equally to this work with: Anatole Ghazalpour, Sudheer Doss, Aldons J Lusis, Steve Horvath

Affiliations Department of Microbiology, Immunology, and Molecular Genetics, University of California Los Angeles, Los Angeles, California, United States of America , Department of Human Genetics, David Geffen School of Medicine, University of California Los Angeles, Los Angeles, California, United States of America , Department of Medicine, David Geffen School of Medicine, University of California Los Angeles, Los Angeles, California, United States of America , Molecular Biology Institute, University of California Los Angeles, Los Angeles, California, United States of America

Steve Horvath

Contributed equally to this work with: Anatole Ghazalpour, Sudheer Doss, Aldons J Lusis, Steve Horvath

To whom correspondence should be addressed. E-mail: shorvath@mednet.ucla.edu

Affiliations Department of Human Genetics, David Geffen School of Medicine, University of California Los Angeles, Los Angeles, California, United States of America , Department of Biostatistics, School of Public Health, University of California Los Angeles, Los Angeles, California, United States of America

Competing Interests

The Molecular Biology Institute is a wholly-owned subsidiary of Merck & Co., Inc.

Author Contributions

AG, SD and SH conceived and designed the experiments. AG, SD, RC, and AB performed the experiments. AG, SD, CP, and SH analyzed the data. AG, SD, BZ, and SH contributed reagents/materials/analysis tools. BZ wrote major parts of the R software that will be made publicly available. SW was involved in F2 mouse cross data generation. CP helped to carry out the functional enrichment analysis. EES and TAD participated in data generation and conceptual discussion. AJL and SH co-directed the work. AG, SD, AJL, and SH wrote the paper.