Abstract

Blood pressure (BP) is a major risk factor for cardiovascular disease and more than 200 genetic loci associated with BP are known. Here, we perform a multi-stage genome-wide association study for BP (max N = 289,038) principally in East Asians and meta-analysis in East Asians and Europeans. We report 19 new genetic loci and ancestry-specific BP variants, conforming to a common ancestry-specific variant association model. At 10 unique loci, distinct non-rare ancestry-specific variants colocalize within the same linkage disequilibrium block despite the significantly discordant effects for the proxy shared variants between the ethnic groups. The genome-wide transethnic correlation of causal-variant effect-sizes is 0.898 and 0.851 for systolic and diastolic BP, respectively. Some of the ancestry-specific association signals are also influenced by a selective sweep. Our results provide new evidence for the role of common ancestry-specific variants and natural selection in ethnic differences in complex traits such as BP.

Details

Title
Interethnic analyses of blood pressure loci in populations of East Asian and European descent
Author
Takeuchi, Fumihiko 1   VIAFID ORCID Logo  ; Akiyama, Masato 2 ; Matoba, Nana 2   VIAFID ORCID Logo  ; Katsuya, Tomohiro 3 ; Nakatochi, Masahiro 4   VIAFID ORCID Logo  ; Tabara, Yasuharu 5 ; Narita, Akira 6 ; Woei-Yuh Saw 7 ; Moon, Sanghoon 8 ; Spracklen, Cassandra N 9 ; Jin-Fang, Chai 10 ; Young-Jin, Kim 8 ; Zhang, Liang 11 ; Wang, Chaolong 12 ; Li, Huaixing 13 ; Li, Honglan 14 ; Wu, Jer-Yuarn 15 ; Dorajoo, Rajkumar 16   VIAFID ORCID Logo  ; Nierenberg, Jovia L 17 ; Ya Xing Wang 18   VIAFID ORCID Logo  ; He, Jing 19 ; Bennett, Derrick A 20 ; Takahashi, Atsushi 21   VIAFID ORCID Logo  ; Momozawa, Yukihide 22 ; Hirata, Makoto 23   VIAFID ORCID Logo  ; Matsuda, Koichi 24 ; Rakugi, Hiromi 25   VIAFID ORCID Logo  ; Nakashima, Eitaro 26 ; Isono, Masato 27 ; Shirota, Matsuyuki 6 ; Hozawa, Atsushi 6 ; Ichihara, Sahoko 28 ; Matsubara, Tatsuaki 29 ; Yamamoto, Ken 30   VIAFID ORCID Logo  ; Kohara, Katsuhiko 31 ; Igase, Michiya 32   VIAFID ORCID Logo  ; Han, Sohee 8 ; Gordon-Larsen, Penny 33   VIAFID ORCID Logo  ; Huang, Wei 34 ; Lee, Nanette R 35 ; Adair, Linda S 33 ; Hwang, Mi Yeong 8 ; Lee, Juyoung 8 ; Miao, Li Chee 11 ; Charumathi Sabanayagam 36   VIAFID ORCID Logo  ; Zhao, Wanting 37 ; Liu, Jianjun 38   VIAFID ORCID Logo  ; Reilly, Dermot F 39   VIAFID ORCID Logo  ; Sun, Liang 13 ; Huo, Shaofeng 13 ; Edwards, Todd L 19   VIAFID ORCID Logo  ; Long, Jirong 19 ; Li-Ching, Chang 40 ; Chen, Chien-Hsiun 40 ; Jian-Min, Yuan 41 ; Koh, Woon-Puay 42 ; Friedlander, Yechiel 43 ; Kelly, Tanika N 17 ; Wen Bin Wei 44 ; Xu, Liang 18 ; Cai, Hui 19 ; Yong-Bing, Xiang 14 ; Lin, Kuang 20 ; Clarke, Robert 20 ; Walters, Robin G 20   VIAFID ORCID Logo  ; Millwood, Iona Y 45 ; Li, Liming 46 ; Chambers, John C 47 ; Kooner, Jaspal S 48 ; Elliott, Paul 49 ; Pim van der Harst 50   VIAFID ORCID Logo  ; Loh, Marie 51 ; Verweij, Niek 52 ; Zhang, Weihua 53 ; Lehne, Benjamin 47 ; Irene Mateo Leach 52 ; Drong, Alexander 54 ; Abbott, James 55 ; Tan, Sian-Tsung 56 ; Scott, William R 57 ; Campanella, Gianluca 47 ; Chadeau-Hyam, Marc 47 ; Afzal, Uzma 53 ; Tõnu Esko 58 ; Harris, Sarah E 59 ; Hartiala, Jaana 60 ; Kleber, Marcus E 61 ; Saxena, Richa 62 ; Stewart, Alexandre F R 63 ; Ahluwalia, Tarunveer S 64 ; Aits, Imke 65 ; Alexessander Da Silva Couto Alves 66 ; Das, Shikta 66 ; Hopewell, Jemma C 67 ; Koivula, Robert W 68 ; Leo-Pekka Lyytikäinen 69 ; Postmus, Iris 70 ; Raitakari, Olli T 71 ; Scott, Robert A 72 ; Sorice, Rossella 73 ; Tragante, Vinicius 74 ; Traglia, Michela 75 ; White, Jon 76 ; Barroso, Inês 77 ; Bjonnes, Andrew 62 ; Collins, Rory 74 ; Davies, Gail 78 ; Delgado, Graciela 61 ; Doevendans, Pieter A 74 ; Franke, Lude 79 ; Gansevoort, Ron T 80 ; Grammer, Tanja B 61 ; Grarup, Niels 61 ; Grewal, Jagvir 53 ; Hartikainen, Anna-Liisa 81 ; Hazen, Stanley L 82 ; Hsu, Chris 83 ; Husemoen, Lise L N 84 ; Justesen, Johanne M 64 ; Kumari, Meena 85 ; Lieb, Wolfgang 65 ; Liewald, David C M 78 ; Mihailov, Evelin 86 ; Milani, Lili 86 ; Mills, Rebecca 87 ; Mononen, Nina 69 ; Nikus, Kjell 88 ; Nutile, Teresa 73 ; Parish, Sarah 67 ; Rolandsson, Olov 89 ; Ruggiero, Daniela 73 ; Sala, Cinzia F 90 ; Snieder, Harold 91 ; Spasø, Thomas H W 64 ; Spiering, Wilko 92 ; Starr, John M 93 ; Stott, David J 94 ; Stram, Daniel O 83 ; Szymczak, Silke 95 ; Wilson Tang, W H 82 ; Trompet, Stella 96 ; Turjanmaa, Väinö 97 ; Vaarasmaki, Marja 98 ; van Gilst, Wiek H 52 ; van Veldhuisen, Dirk J 52 ; Viikari, Jorma S 99 ; Asselbergs, Folkert W 100 ; Ciullo, Marina 73 ; Franke, Andre 95 ; Franks, Paul W 101 ; Franks, Steve 102 ; Gross, Myron D 103 ; Hansen, Torben 64 ; Jarvelin, Marjo-Riitta 104 ; Jørgensen, Torben 84 ; Jukema, Wouter J 105 ; Kähönen, Mika 97 ; Kivimaki, Mika 85 ; Lehtimäki, Terho 69 ; Linneberg, Allan 84 ; Pedersen, Oluf 64 ; Samani, Nilesh J 106 ; Toniolo, Daniela 107 ; Allayee, Hooman 60 ; Deary, Ian J 108 ; März, Winfried 109 ; Metspalu, Andres 86 ; Wijmenga, Cisca 79 ; Wolffenbuttel, Bruce H W 110 ; Vineis, Paolo 47 ; Kyrtopoulos, Soterios A 111 ; Kleinjans, Jos C S 112 ; McCarthy, Mark I 113 ; Scott, James 48 ; Chen, Zhengming 20 ; Sasaki, Makoto 114   VIAFID ORCID Logo  ; Xiao-Ou, Shu 19 ; Jonas, Jost B 115   VIAFID ORCID Logo  ; He, Jiang 116 ; Chew-Kiat Heng 117   VIAFID ORCID Logo  ; Yuan-Tsong, Chen 40 ; Zheng, Wei 19   VIAFID ORCID Logo  ; Xu, Lin 13 ; Yik-Ying Teo 118 ; E-Shyong, Tai 119 ; Ching-Yu, Cheng 36 ; Tien Yin Wong 36 ; Sim, Xueling 10   VIAFID ORCID Logo  ; Mohlke, Karen L 9   VIAFID ORCID Logo  ; Yamamoto, Masayuki 6 ; Bong-Jo, Kim 8 ; Miki, Tetsuro 32 ; Nabika, Toru 120 ; Yokota, Mitsuhiro 121 ; Kamatani, Yoichiro 122   VIAFID ORCID Logo  ; Kubo, Michiaki 123 ; Kato, Norihiro 1 

 Medical Genomics Center, National Center for Global Health and Medicine, Tokyo, Japan; Department of Gene Diagnostics and Therapeutics, Research Institute, National Center for Global Health and Medicine, Tokyo, Japan 
 Laboratory for Statistical Analysis, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan 
 Department of Clinical Gene Therapy, Osaka University Graduate School of Medicine, Suita, Japan; Department of Geriatric and General Medicine, Osaka University Graduate School of Medicine, Suita, Japan 
 Data Coordinating Center, Department of Advanced Medicine, Nagoya University Hospital, Nagoya, Japan 
 Center for Genomic Medicine, Kyoto University Graduate School of Medicine, Kyoto, Japan 
 Tohoku Medical Megabank Organization, Tohoku University, Sendai, Japan 
 Saw Swee Hock School of Public Health, National University of Singapore and National University Health System, Singapore, Singapore; Life Sciences Institute, National University of Singapore, Singapore, Singapore 
 Division of Genome Research, Center for Genome Science, National Institute of Health, Chungcheongbuk-do, Republic of Korea 
 Department of Genetics, University of North Carolina, Chapel Hill, USA 
10  Saw Swee Hock School of Public Health, National University of Singapore and National University Health System, Singapore, Singapore 
11  Singapore Eye Research Institute, Singapore National Eye Centre, Singapore, Singapore 
12  Genome Institute of Singapore, Agency for Science, Technology and Research, Singapore, Singapore; Duke-NUS Medical School, Singapore, Singapore; Department of Epidemiology and Biostatistics, School of Public Health, Tongji Medical College, Huazhong University of Science and Technology, Wuhan, China 
13  CAS Key Laboratory of Nutrition, Metabolism and Food safety, Shanghai Institute of Nutrition and Health, Shanghai Institutes for Biological Sciences, University of the Chinese Academy of Sciences, Chinese Academy of Sciences, Shanghai, China 
14  State Key Laboratory of Oncogene and Related Genes and Department of Epidemiology, Shanghai Cancer Institute, Renji Hospital, Shanghai Jiaotong University School of Medicine, Shanghai, China 
15  Institute of Biomedical Sciences, Academia Sinica, Taipei, Taiwan; School of Chinese Medicine, China Medical University, Taichung, Taiwan 
16  Genome Institute of Singapore, Agency for Science, Technology and Research, Singapore, Singapore 
17  Department of Epidemiology, Tulane University School of Public Health and Tropical Medicine, New Orleans, USA 
18  Beijing Institute of Ophthalmology, Beijing Key Laboratory of Ophthalmology and Visual Sciences, Beijing Tongren Eye Center, Beijing Tongren Hospital, Capital Medical University, Beijing, China 
19  Division of Epidemiology, Vanderbilt University Medical Center, Nashville, USA 
20  Clinical Trial Service Unit and Epidemiological Studies Unit, Nuffield Department of Population Health, University of Oxford, Oxford, UK 
21  Laboratory for Statistical Analysis, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan; Department of Genomic Medicine, Research Institute, National Cerebral and Cardiovascular Center, Osaka, Japan 
22  Laboratory for Genotyping Development, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan 
23  Institute of Medical Science, The University of Tokyo, Tokyo, Japan 
24  Graduate School of Frontier Sciences, The University of Tokyo, Kashiwa, Japan 
25  Department of Geriatric and General Medicine, Osaka University Graduate School of Medicine, Suita, Japan 
26  Division of Endocrinology and Diabetes, Department of Internal Medicine, Nagoya University Graduate School of Medicine, Nagoya, Japan; Department of Diabetes and Endocrinology, Chubu Rosai Hospital, Nagoya, Japan 
27  Department of Gene Diagnostics and Therapeutics, Research Institute, National Center for Global Health and Medicine, Tokyo, Japan 
28  Department of Environmental and Preventive Medicine, Jichi Medical University School of Medicine, Shimotsuke, Japan 
29  Department of Internal Medicine, School of Dentistry, Aichi Gakuin University, Nagoya, Japan 
30  Department of Medical Biochemistry, Kurume University School of Medicine, Kurume, Japan 
31  Faculty of Collaborative Regional Innovation, Ehime University, Matsuyama, Ehime, Japan 
32  Department of Geriatric Medicine, Ehime University Graduate School of Medicine, Toon, Ehime, Japan 
33  Department of Nutrition, Gillings School of Global Public Health, University of North Carolina at Chapel Hill, Chapel Hill, USA; Carolina Population Center, University of North Carolina at Chapel Hill, Chapel Hill, USA 
34  Department of Genetics, Shanghai-MOST Key Laboratory of Health and Disease Genomics, Chinese National Human Genome Center and Shanghai Industrial Technology Institute (SITI), Shanghai, China 
35  USC-Office of Population Studies Foundation, University of San Carlos, Cebu City, Philippines; Department of Anthropology, Sociology and History, University of San Carlos, Cebu City, Philippines 
36  Singapore Eye Research Institute, Singapore National Eye Centre, Singapore, Singapore; Ophthalmology & Visual Sciences Academic Clinical Program (Eye ACP), Duke-NUS Medical School, Singapore, Singapore; Department of Ophthalmology, Yong Loo Lin School of Medicine, National University of Singapore, Singapore, Singapore 
37  Singapore Eye Research Institute, Singapore National Eye Centre, Singapore, Singapore; Duke-NUS Medical School, Singapore, Singapore 
38  Genome Institute of Singapore, Agency for Science, Technology and Research, Singapore, Singapore; Yong Loo Lin School of Medicine, National University of Singapore, Singapore, Singapore 
39  Merck Sharp Dohme Corp, Kenilworth, USA 
40  Institute of Biomedical Sciences, Academia Sinica, Taipei, Taiwan 
41  Department of Epidemiology, Graduate School of Public Health, University of Pittsburgh, Pittsburgh, USA; Division of Cancer Control and Population Sciences, UPMC Hillman Cancer, University of Pittsburgh, Pittsburgh, USA 
42  Saw Swee Hock School of Public Health, National University of Singapore and National University Health System, Singapore, Singapore; Health Services and Systems Research, Duke-NUS Medical School, Singapore, Singapore 
43  Unit of Epidemiology, Hebrew University-Hadassah Braun School of Public Health, Jerusalem, Israel 
44  Beijing Tongren Eye Center, Beijing Tongren Hospital, Capital Medical University, Beijing, China 
45  Clinical Trial Service Unit and Epidemiological Studies Unit, Nuffield Department of Population Health, University of Oxford, Oxford, UK; MRC Population Health Research Unit, Nuffield Department of Population Health, University of Oxford, Oxford, UK 
46  Chinese Academy of Medical Sciences, Beijing, China; Department of Epidemiology and Biostatistics, School of Public Health, Peking University, Beijing, China 
47  Department of Epidemiology and Biostatistics, Imperial College London, London, UK 
48  National Heart and Lung Institute, Imperial College London, London, UK 
49  Medical Research Council-Public Health England (MRC-PHE) Centre for Environment and Health, Department of Epidemiology and Biostatistics, School of Public Health, Faculty of Medicine, Imperial College London, London, UK; Imperial College Biomedical Research Centre, Imperial College London, London, UK; UK-Dementia Research Institute at Imperial College London, London, UK; National Institute for Health Research (NIHR) Health Protection Research Unit on Health Impacts of Environmental Hazards, London, UK; Health Data Research UK-London, London, UK 
50  Department of Cardiology, University of Groningen, University Medical Center Groningen, Groningen, Netherlands 
51  Institute of Health Sciences, University of Oulu, Oulu, Finland; Department of Epidemiology and Biostatistics, Imperial College London, London, UK 
52  Department of Cardiology, University Medical Center Groningen, University of Groningen, Groningen, Netherlands 
53  Department of Epidemiology and Biostatistics, Imperial College London, London, UK; Ealing Hospital NHS Trust, Middlesex, UK 
54  Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, UK 
55  Bioinformatics Support Service, Imperial College London, South Kensington, London, UK 
56  Ealing Hospital NHS Trust, Middlesex, UK; National Heart and Lung Institute, Imperial College London, London, UK 
57  Department of Epidemiology and Biostatistics, Imperial College London, London, UK; National Heart and Lung Institute, Imperial College London, London, UK 
58  Estonian Genome Center, University of Tartu, Tartu, Estonia; Division of Endocrinology, Children’s Hospital Boston, Boston, MA, USA; Department of Genetics, Harvard Medical School, Boston, MA, USA; Program in Medical and Population Genetics, Broad Institute, Cambridge, MA, USA 
59  Medical Genetics Section, University of Edinburgh Molecular Medicine Centre and MRC Institute of Genetics and Molecular Medicine, Western General Hospital, Edinburgh, UK; Centre for Cognitive Aging and Cognitive Epidemiology, University of Edinburgh, Edinburgh, UK 
60  Department of Preventive Medicine, USC Keck School of Medicine, Los Angeles, CA, USA; Institute for Genetic Medicine, USC Keck School of Medicine, Los Angeles, CA, USA 
61  Medical Clinic V, Mannheim Medical Faculty, University of Heidelberg, Mannheim, Germany 
62  Massachusetts General Hospital, Harvard Medical School, Boston, MA, USA 
63  University of Ottawa Heart Institute, Cardiovascular Research Methods Centre, Ontario, Canada; Ruddy Canadian Cardiovascular Genetics Centre, Ontario, Canada 
64  Novo Nordisk Foundation Centre for Basic Metabolic Research, Section of Metabolic Genetics, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark 
65  Institute of Epidemiology and Biobank Popgen, Christian-Albrechts-University of Kiel, Kiel, Germany 
66  Department of Epidemiology and Biostatistics, MRC Health Protection Agency (HPE) Centre for Environment and Health, School of Public Health, Imperial College London, London, UK 
67  Clinical Trial Service Unit & Epidemiological Studies Unit, University of Oxford, Oxford, UK 
68  Department of Clinical Sciences, Genetic and Molecular Epidemiology Unit, Skåne University Hospital Malmö, Malmö, Sweden 
69  Department of Clinical Chemistry, Fimlab Laboratories, Tampere, Finland; Department of Clinical Chemistry, University of Tampere School of Medicine, Tampere, Finland 
70  Department of Gerontology and Geriatrics, Leiden University Medical Center, Leiden, Netherlands; Netherlands Consortium for Healthy Ageing, Leiden, Netherlands 
71  Department of Clinical Physiology and Nuclear Medicine, Turku University Hospital, Turku, Finland; Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku, Turku, Finland 
72  MRC Epidemiology Unit, Institute of Metabolic Science, University of Cambridge, Cambridge, UK 
73  Institute of Genetics and Biophysics A. Buzzati-Traverso, CNR, Naples, Italy 
74  Department of Cardiology, Division Heart and Lungs, University Medical Center Utrecht, Utrecht, Netherlands 
75  Division of Genetics and Cell Biology, San Raffaele Scientific Institute, Milano, Italy; Institute for Maternal and Child Health—IRCCS ‘‘Burlo Garofolo’’—Trieste, Trieste, Italy 
76  UCL Genetics Institute, Department of Genetics, Environment and Evolution, UCL, London, UK 
77  Metabolic Disease Group, The Wellcome Trust Sanger Institute, Cambridge, UK; NIHR Cambridge Biomedical Research Centre, Institute of Metabolic Science, Addenbrooke’s Hospital, Cambridge, UK; University of Cambridge Metabolic Research Laboratories, Institute of Metabolic Science, Addenbrooke’s Hospital, Cambridge, UK 
78  Department of Psychology, University of Edinburgh, Edinburgh, UK 
79  Department of Genetics, University Medical Center, University of Groningen, Groningen, Netherlands 
80  Department of Internal Medicine, University Medical Center Groningen, University of Groningen, Groningen, Netherlands 
81  Department of Obstetrics and Gynecology, University Hospital of Oulu, University of Oulu, Oulu, Finland; Department of Clinical Sciences/Obsterics and Gynecology, University of Oulu, Oulu, Finland 
82  Center for Cardiovascular Diagnostics and Prevention, Cleveland Clinic, Cleveland, OH, USA; Department of Cellular and Molecular Medicine, Lerner Research Institute, Cleveland Clinic, Cleveland, OH, USA 
83  Keck School of Medicine, University of Southern California, Los Angeles, CA, USA 
84  Research Centre for Prevention and Health, Glostrup University Hospital, Glostrup, Denmark 
85  Department of Epidemiology and Public Health, UCL, London, UK 
86  Estonian Genome Center, University of Tartu, Tartu, Estonia 
87  Ealing Hospital NHS Trust, Middlesex, UK 
88  Heart Centre, Department of Cardiology, Tampere University Hospital, and University of Tampere School of Medicine, Tampere, Finland 
89  Department of Public Health & Clinical Medicine, Section for Family Medicine, Umeå universitet, Umeå, Sweden 
90  Division of Genetics and Cell Biology, San Raffaele Scientific Institute, Milano, Italy 
91  Department of Epidemiology, University Medical Center Groningen, University of Groningen, Groningen, Netherlands 
92  Department of Vascular Medicine, University Medical Center Utrecht, Utrecht, Netherlands 
93  Centre for Cognitive Aging and Cognitive Epidemiology, University of Edinburgh, Edinburgh, UK; Alzheimer Scotland Dementia Research Centre, University of Edinburgh, Edinburgh, UK 
94  Academic Section of Geriatric Medicine, Institute of Cardiovascular and Medical Sciences, Faculty of Medicine, University of Glasgow, Glasgow, UK 
95  Institute of Clinical Molecular Biology, Christian-Albrechts-University of Kiel, Kiel, Germany 
96  Department of Cardiology, Leiden University Medical Center, Leiden, Netherlands 
97  Department of Clinical Physiology, Tampere University Hospital, Tampere, Finland; Department of Clinical Physiology, University of Tampere School of Medicine, Tampere, Finland 
98  Department of Obstetrics and Gynecology, Oulu University Hospital, Oulu, Finland 
99  Department of Medicine, Turku University Hospital, Turku, Finland; Department of Medicine, University of Turku, Turku, Finland 
100  Department of Cardiology, Division Heart and Lungs, University Medical Center Utrecht, Utrecht, Netherlands; Durrer Center for Cardiogenetic Research, ICIN-Netherlands Heart Institute, Utrecht, Netherlands; Institute of Cardiovascular Science, Faculty of Population Health Sciences, University College London, London, UK 
101  Department of Clinical Sciences, Genetic and Molecular Epidemiology Unit, Skåne University Hospital Malmö, Malmö, Sweden; Department of Public Health & Clinical Medicine, Section for Family Medicine, Umeå universitet, Umeå, Sweden; Department of Nutrition, Harvard School of Public Health, Boston, MA, USA 
102  Institute of Reproductive and Developmental Biology, Imperial College London, Hammersmith Hospital, London, UK 
103  School of Medicine, University of Minnesota, Minneapolis, MN, USA 
104  Department of Epidemiology and Biostatistics, Imperial College London, London, UK; Department of Epidemiology and Biostatistics, MRC Health Protection Agency (HPE) Centre for Environment and Health, School of Public Health, Imperial College London, London, UK; Biocenter Oulu, University of Oulu, Oulu, Finland; Unit of Primary Care, Oulu University Hospital, Oulu, Finland; Department of Children and Young People and Families, National Institute for Health and Welfare, Oulu, Finland 
105  Department of Cardiology, Leiden University Medical Center, Leiden, Netherlands; Durrer Center for Cardiogenetic Research, ICIN-Netherlands Heart Institute, Utrecht, Netherlands; Interuniversity Cardiology Institute of the Netherlands, Utrecht, Netherlands 
106  Department of Cardiovascular Sciences, University of Leicester, Glenfield Hospital, Leicester, UK; National Institute for Health Research Leicester Cardiovascular Biomedical Research Unit, Glenfield Hospital, Leicester, UK 
107  Division of Genetics and Cell Biology, San Raffaele Scientific Institute, Milano, Italy; Institute of Molecular GeneticsCNR, Pavia, Italy 
108  Centre for Cognitive Aging and Cognitive Epidemiology, University of Edinburgh, Edinburgh, UK; Department of Psychology, University of Edinburgh, Edinburgh, UK 
109  Medical Clinic V, Mannheim Medical Faculty, University of Heidelberg, Mannheim, Germany; Clinical Institute of Medical and Chemical Laboratory Diagnostics, Medical University of Graz, Graz, Austria; Synlab Academy, Synlab Services GmbH, Mannheim, Germany 
110  Department of Endocrinology, University Medical Center Groningen, University of Groningen, Groningen, Netherlands 
111  National Hellenic Research Foundation, Institute of Biological Research and Biotechnology, Athens, Greece 
112  Department of Toxicogenomics, Maastricht University, Maastricht, Netherlands 
113  Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, UK; Oxford Centre for Diabetes Endocrinology and Metabolism, University of Oxford, Oxford, UK 
114  Iwate Tohoku Medical Megabank Organization, Iwate, Japan 
115  Beijing Institute of Ophthalmology, Beijing Key Laboratory of Ophthalmology and Visual Sciences, Beijing Tongren Eye Center, Beijing Tongren Hospital, Capital Medical University, Beijing, China; Department of Ophthalmology, Medical Faculty Mannheim, University Heidelberg, Germany, Mannheim, Germany 
116  Department of Epidemiology, Tulane University School of Public Health and Tropical Medicine, New Orleans, USA; Department of Medicine, Tulane University School of Medicine, New Orleans, LA, USA 
117  Department of Paediatrics, Yong Loo Lin School of Medicine, National University of Singapore, Singapore, Singapore; Khoo Teck Puat-National University Children’s Medical Institute, National University Health System, Singapore, Singapore 
118  Saw Swee Hock School of Public Health, National University of Singapore and National University Health System, Singapore, Singapore; Life Sciences Institute, National University of Singapore, Singapore, Singapore; NUS Graduate School for Integrative Science and Engineering, National University of Singapore, Singapore, Singapore 
119  Saw Swee Hock School of Public Health, National University of Singapore and National University Health System, Singapore, Singapore; Duke-NUS Medical School, Singapore, Singapore; Yong Loo Lin School of Medicine, National University of Singapore, Singapore, Singapore 
120  Department of Functional Pathology, Shimane University Faculty of Medicine, Izumo, Japan 
121  Department of Genome Science, School of Dentistry, Aichi Gakuin University, Nagoya, Japan 
122  Laboratory for Statistical Analysis, RIKEN Center for Integrative Medical Sciences, Yokohama, Japan; Kyoto-McGill International Collaborative School in Genomic Medicine, Kyoto University Graduate School of Medicine, Kyoto, Japan 
123  RIKEN Center for Integrative Medical Sciences, Yokohama, Japan 
Pages
1-16
Publication year
2018
Publication date
Nov 2018
Publisher
Nature Publishing Group
e-ISSN
20411723
Source type
Scholarly Journal
Language of publication
English
ProQuest document ID
2139097911
Copyright
© 2018. This work is published under http://creativecommons.org/licenses/by/4.0/ (the “License”). Notwithstanding the ProQuest Terms and Conditions, you may use this content in accordance with the terms of the License.