About the Authors

Theru A. Sivakumaran

Contributed equally to this work with: Theru A. Sivakumaran, Robert P. Igo Jr

Affiliations Department of Epidemiology and Biostatistics, Case Western Reserve University, Cleveland, Ohio, United States of America, Division of Human Genetics, Cincinnati Children's Hospital Medical Center, Cincinnati, Ohio, United States of America

Robert P. Igo Jr

Contributed equally to this work with: Theru A. Sivakumaran, Robert P. Igo Jr

Affiliation Department of Epidemiology and Biostatistics, Case Western Reserve University, Cleveland, Ohio, United States of America

Jeffrey M. Kidd

Affiliation Department of Genome Sciences, University of Washington, Seattle, Washington, United States of America

Andy Itsara

Affiliation Department of Genome Sciences, University of Washington, Seattle, Washington, United States of America

Laura J. Kopplin

Affiliation Department of Genetics, Case Western Reserve University, Cleveland, Ohio, United States of America

Wei Chen

Affiliation Department of Biostatistics, Center for Statistical Genetics, University of Michigan School of Public Health, Ann Arbor, Michigan, United States of America

Stephanie A. Hagstrom

Affiliations Cole Eye Institute, Cleveland Clinic, Cleveland, Ohio, United States of America, Department of Ophthalmology, Cleveland Clinic Lerner College of Medicine of Case Western Reserve University, Cleveland, Ohio, United States of America

Neal S. Peachey

Affiliations Cole Eye Institute, Cleveland Clinic, Cleveland, Ohio, United States of America, Department of Ophthalmology, Cleveland Clinic Lerner College of Medicine of Case Western Reserve University, Cleveland, Ohio, United States of America, Research Service, Louis Stokes Cleveland Department of Veterans Affairs Medical Center, Cleveland, Ohio, United States of America

Peter J. Francis

Affiliation Casey Eye Institute, Oregon Health & Science University, Portland, Oregon, United States of America

Michael L. Klein

Affiliation Casey Eye Institute, Oregon Health & Science University, Portland, Oregon, United States of America

Emily Y. Chew

Affiliation Division of Epidemiology and Clinical Applications, National Eye Institute, Bethesda, Maryland, United States of America

Vedam L. Ramprasad

Affiliation SNONGC Department of Genetics and Molecular Biology, Vision Research Foundation, Sankara Nethralaya, Chennai, India

Wan-Ting Tay

Affiliation Singapore Eye Research Institute, Yong Loo Lin School of Medicine, National University of Singapore, Singapore, Singapore

Paul Mitchell

Affiliation Centre for Vision Research, University of Sydney, Sydney, Australia

Mark Seielstad

Affiliation Genome Institute of Singapore, Singapore, Singapore

Dwight E. Stambolian

Affiliation Departments of Ophthalmology and Genetics, University of Pennsylvania, Philadelphia, Pennsylvania, United States of America

Albert O. Edwards

Affiliation Institute of Molecular Biology, University of Oregon, Eugene, Oregon, United States of America

Kristine E. Lee

Affiliation Department of Ophthalmology and Visual Sciences, University of Wisconsin School of Medicine and Public Health, Madison, Wisconsin, United States of America

Dmitry V. Leontiev

Affiliation Department of Epidemiology and Biostatistics, Case Western Reserve University, Cleveland, Ohio, United States of America

Gyungah Jun

Affiliations Department of Epidemiology and Biostatistics, Case Western Reserve University, Cleveland, Ohio, United States of America, Department of Medicine, Boston University, Boston, Massachusetts, United States of America, Department of Ophthalmology, Boston University, Boston, Massachusetts, United States of America, Department of Biostatistics, Boston University, Boston, Massachusetts, United States of America

Yang Wang

Affiliation Department of Epidemiology and Biostatistics, Case Western Reserve University, Cleveland, Ohio, United States of America

Liping Tian

Affiliation Department of Epidemiology and Biostatistics, Case Western Reserve University, Cleveland, Ohio, United States of America

Feiyou Qiu

Affiliation Department of Epidemiology and Biostatistics, Case Western Reserve University, Cleveland, Ohio, United States of America

Alice K. Henning

Affiliation The EMMES Corporation, Rockville, Maryland, United States of America

Thomas LaFramboise

Affiliation Department of Genetics, Case Western Reserve University, Cleveland, Ohio, United States of America

Parveen Sen

Affiliation Department of Medical Retina, Vision Research Foundation, Sankara Nethralaya, Chennai, India

Manoharan Aarthi

Affiliation SNONGC Department of Genetics and Molecular Biology, Vision Research Foundation, Sankara Nethralaya, Chennai, India

Ronnie George

Affiliation Department of Glaucoma, Vision Research Foundation, Sankara Nethralaya, Chennai, India

Rajiv Raman

Affiliation Department of Medical Retina, Vision Research Foundation, Sankara Nethralaya, Chennai, India

Manmath Kumar Das

Affiliation Department of Medical Retina, Vision Research Foundation, Sankara Nethralaya, Chennai, India

Lingam Vijaya

Affiliation Department of Medical Retina, Vision Research Foundation, Sankara Nethralaya, Chennai, India

Govindasamy Kumaramanickavel

Affiliation SNONGC Department of Genetics and Molecular Biology, Vision Research Foundation, Sankara Nethralaya, Chennai, India

Tien Y. Wong

Affiliations Singapore Eye Research Institute, Yong Loo Lin School of Medicine, National University of Singapore, Singapore, Singapore, Centre for Eye Research Australia, University of Melbourne, East Melbourne, Australia

Anand Swaroop

Affiliations Neurobiology Neurodegeneration and Repair Laboratory, National Eye Institute, Bethesda, Maryland, United States of America, Kellogg Eye Center and Department of Human Genetics, University of Michigan, Ann Arbor, Michigan, United States of America

Goncalo R. Abecasis

Affiliation Department of Biostatistics, Center for Statistical Genetics, University of Michigan School of Public Health, Ann Arbor, Michigan, United States of America

Ronald Klein

Affiliation Department of Ophthalmology and Visual Sciences, University of Wisconsin School of Medicine and Public Health, Madison, Wisconsin, United States of America

Barbara E. K. Klein

Affiliation Department of Ophthalmology and Visual Sciences, University of Wisconsin School of Medicine and Public Health, Madison, Wisconsin, United States of America

Deborah A. Nickerson

Affiliation Department of Genome Sciences, University of Washington, Seattle, Washington, United States of America

Evan E. Eichler

Affiliations Department of Genome Sciences, University of Washington, Seattle, Washington, United States of America, Howard Hughes Medical Institute, University of Washington, Seattle, Washington, United States of America

Sudha K. Iyengar

ski@case.edu

Affiliations Department of Epidemiology and Biostatistics, Case Western Reserve University, Cleveland, Ohio, United States of America, Department of Genetics, Case Western Reserve University, Cleveland, Ohio, United States of America, Department of Ophthalmology, Case Western Reserve University, Cleveland, Ohio, United States of America

Competing Interests

Of two co-authors of this paper who have declared conflict, Alice Henning is employed by the EMMES corporation, and Dr. Evan E. Eichler is a member of the scientific advisory board for Pacific Biosciences, respectively. Their association with each respective organization does not alter the authors' adherence to all the PLoS ONE policies on sharing of data and materials.

Author Contributions

Conceived and designed the experiments: TAS SKI. Performed the experiments: TAS LT VLR. Analyzed the data: TAS RPI JMK AI LJK WC WTT DVL GJ YW FQ TL. Contributed reagents/materials/analysis tools: SAH NSP PJF MLK EYC PM MS DES AOE KEL AKH PS MA RG RR MKD LV GK TYW AS GRA RK BEKK DAN EEE SKI. Wrote the paper: TAS RPI EEE AS RK BEKK SKI.