Author information: (1)QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia. Puya.Gharahkhani@qimrberghofer.edu.au. (2)Division of Research, Kaiser Permanente Northern California (KPNC), Oakland, CA, USA. (3)Twin Research and Genetic Epidemiology, King's College London, London, UK. (4)NIHR Biomedical Research Centre, Moorfields Eye Hospital NHS Foundation Trust and UCL Institute of Ophthalmology, London, UK. (5)Department of Public Health and Primary Care, Institute of Public Health, University of Cambridge School of Clinical Medicine, Cambridge, UK. (6)Geisinger Research, Biomedical and Translational Informatics Institute, Danville, PA, USA. (7)QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia. (8)Menzies Institute for Medical Research, University of Tasmania, Hobart, TAS, Australia. (9)Centre for Eye Research Australia, University of Melbourne, Melbourne, VIC, Australia. (10)Department of Ophthalmology, Harvard Medical School, Boston, MA, USA. (11)Department of Population and Quantitative Health Sciences, Case Western Reserve University School of Medicine, Cleveland, OH, USA. (12)Department of Ophthalmology, Flinders University, Bedford Park, SA, Australia. (13)Geisinger Research, Biomedical and Translational Informatics Institute, Rockville, MD, USA. (14)Cleveland Institute for Computational Biology, Case Western Reserve University School of Medicine, Cleveland, OH, USA. (15)Depatment of Ophthalmology, Erasmus MC, Rotterdam, The Netherlands. (16)Department of Epidemiology, Erasmus MC, Rotterdam, The Netherlands. (17)The Rotterdam Eye Hospital, Rotterdam, The Netherlands. (18)Department of Clinical Genetics, Erasmus MC, Rotterdam, The Netherlands. (19)Department of Ophthalmology and Visual Sciences, University of Wisconsin-Madison, Madison, WI, USA. (20)Medical Research Council Human Genetics Unit, Institute of Genetics and Molecular Medicine, University of Edinburgh, Edinburgh, UK. (21)Institute for Molecular Medicine Finland, HiLIFE, University of Helsinki, Helsinki, Finland. (22)Broad Institute of the Massachusetts Institute of Technology and Harvard University, Cambridge, MA, USA. (23)Analytic and Translational Genetics Unit, Massachusetts General Hospital and Harvard Medical School, Boston, MA, USA. (24)Institute of Human Genetics, Universitätsklinikum Erlangen, Friedrich-Alexander-Universität, Erlangen-Nürnberg, Erlangen, Germany. (25)Department of Ophthalmology, KPNC, Redwood City, CA, USA. (26)Department of Ophthalmology, School of Medicine, University of California San Francisco (UCSF), San Francisco, CA, USA. (27)Department of Ophthalmology, Kings College London, London, United Kingdom. (28)Institute of Ophthalmology, University College London, London, UK. (29)University Hospital Southampton NHS Foundation Trust, Southampton, UK. (30)Faculty of Medicine, University of Southampton, Southampton, UK. (31)Department of Ophthalmology, Inselspital, University Hospital Bern, University of Bern, Bern, Germany. (32)Department of Ophthalmology, University Medical Center Mainz, Mainz, Germany. (33)Institute of Medical Biostatistics, Epidemiology and Informatics, University Medical Center Mainz, Mainz, Germany. (34)Department of Ophthalmology and Visual Sciences, The Chinese University of Hong Kong, Hong Kong, China. (35)Singapore Eye Research Institute, Singapore National Eye Certre, Singapore, Singapore. (36)Ophthalmology & Visual Sciences Academic Clinical Program, Duke-NUS Medical School, Singapore, Singapore. (37)Department of Ophthalmology, Yong Loo Lin School of Medicine, National University of Singapore, Singapore, Singapore. (38)Duke-National University of Singapore Medical School, Singapore, Republic of Singapore. (39)Tohoku Medical Megabank Organization, Tohoku University, 2-1 Seiryo-machi, Aoba-ku, Sendai, Miyagi, Japan. (40)RIKEN Center for Advanced Intelligence Project, 1-4-1 Nihonbashi, Chuo-ku, Tokyo, Japan. (41)Department of Ophthalmology, Tohoku University Graduate School of Medicine, 1-1, Seiryo-machi, Aoba-ku, Sendai, Miyagi, Japan. (42)Department of Retinal Disease Control, Tohoku University Graduate School of Medicine, 1-1, Seiryo-machi, Aoba-ku, Sendai, Miyagi, Japan. (43)Department of Advanced Ophthalmic Medicine, Tohoku University Graduate School of Medicine, 1-1, Seiryo-machi, Aoba-ku, Sendai, Miyagi, Japan. (44)Department of Ophthalmic Imaging and Information Analytics, Tohoku University Graduate School of Medicine, 1-1, Seiryo-machi, Aoba-ku, Sendai, Miyagi, Japan. (45)European Molecular Biology Laboratory, European Bioinformatics Institute (EMBL-EBI), Wellcome Genome Campus, Hinxton, Cambridge, UK. (46)23 and Me Inc., San Francisco, CA, USA. (47)Department of Ophthalmology, University of Ibadan, Ibadan, Nigeria. (48)Division of Ophthalmology, Department of Neurosciences, University of the Witwatersrand, Johannesburg, South Africa. (49)Unit of Ophthalmology, Department of Surgery, University of Ghana Medical School, Accra, Ghana. (50)Sydney Brenner Institute for Molecular Bioscience, Faculty of Health Sciences, University of the Witwatersrand, Johannesburg, South Africa. (51)Laboratory for Statistical Analysis, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan. (52)Laboratory of Complex Trait Genomics, Department of Computational Biology and Medical Sciences, Graduate School of Frontier Sciences, The University of Tokyo, Tokyo, Japan. (53)Department of Ophthalmology, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan. (54)Laboratory for Genotyping Development, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan. (55)National Institute for Health Research (NIHR) Biomedical Research Centre at Moorfields Eye Hospital National Health Service Foundation Trust & UCL Institute of Ophthalmology, London, UK. (56)UCL Institute of Ophthalmology, University College London, London, UK. (57)Cardiff Centre for Vision Sciences, College of Biomedical and Life Sciences, Maindy Road, Cardiff University, Cardiff, UK. (58)Program in Genetic Epidemiology and Statistical Genetics, Harvard T.H. Chan School of Public Health, Boston, MA, USA. (59)Channing Division of Network Medicine, Brigham and Women's Hospital, Harvard Medical School, Boston, MA, USA. (60)Centre for Vision Research, Department of Ophthalmology and Westmead Institute for Medical Research, University of Sydney, Sydney, NSW, Australia. (61)Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland. (62)Psychiatric & Neurodevelopmental Genetics Unit, Departments of Psychiatry and Neurology, Massachusetts General Hospital, Boston, MA, USA. (63)Broad Institute of MIT and Harvard, Cambridge, MA, USA. (64)Nuffield Department of Population Health, University of Oxford, Oxford, UK. (65)Department of Ophthalmology, Icahn School of Medicine at Mount Sinai, New York, NY, 10029, USA. (66)Department of Ophthalmology, Radboud University Medical Center, Nijmegen, The Netherlands. (67)Institute for Molecular and Clinical Ophthalmology, Basel, Switzerland. (68)Department of Medicine, Duke University, Durham, NC, USA. (69)Department of Ophthalmology, Duke University, Durham, NC, USA. (70)Singapore Eye Research Institute, Singapore, Singapore. (71)Duke-NUS Medical School, Singapore, Singapore. (72)Division of Human Genetics, Genome Institute of Singapore, Singapore, Singapore. (73)Centre for Ophthalmology and Visual Science, University of Western Australia, Lions Eye Institute, Nedlands, WA, Australia. (74)RIKEN Center for Integrative Medical Sciences, Yokohama, Japan. (75)Department of Ophthalmology, Flinders University, Flinders Medical Centre, Bedford Park, SA, Australia. (#)Contributed equally