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      Is Open Access

      A saturated map of common genetic variants associated with human height

      research-article
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340 , 58 , 166 , 478 , 222 , 253 , 480 , 23andMe Research Team, VA Million Veteran Program, DiscovEHR (DiscovEHR and MyCode Community Health Initiative), eMERGE (Electronic Medical Records and Genomics Network), Lifelines Cohort Study, The PRACTICAL Consortium, Understanding Society Scientific Group, 5 , 73 , 223 , 82 , 26 , 214 , 497 , 85 , 482 , 5 , 57 , 483 , 29 , 231 , 338 , 484 , 485 , 33 , 463 , 21 , 4 , 486 , 9 , 481 , 3 , 23 , 26 , 40 , 207 , 208 , 487 , 19 , 9 , 231 , 338 , 27 , 190 , 82 , 9 , 3 , 17 , 18 , 1 , 488 , 489 , 28 , 234 , 235 , 12 , 21 , 16 , 268 , 490 , 491 , 15 , 169 , 456 , 24 , 6 , 7 , 492 , 493 , 498 , 499 , , 24 , , 1 , , 2 , 494 , 495 ,
      Nature
      Nature Publishing Group UK
      Genome-wide association studies, Quantitative trait, Genetic markers

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          Abstract

          Common single-nucleotide polymorphisms (SNPs) are predicted to collectively explain 40–50% of phenotypic variation in human height, but identifying the specific variants and associated regions requires huge sample sizes 1 . Here, using data from a genome-wide association study of 5.4 million individuals of diverse ancestries, we show that 12,111 independent SNPs that are significantly associated with height account for nearly all of the common SNP-based heritability. These SNPs are clustered within 7,209 non-overlapping genomic segments with a mean size of around 90 kb, covering about 21% of the genome. The density of independent associations varies across the genome and the regions of increased density are enriched for biologically relevant genes. In out-of-sample estimation and prediction, the 12,111 SNPs (or all SNPs in the HapMap 3 panel 2 ) account for 40% (45%) of phenotypic variance in populations of European ancestry but only around 10–20% (14–24%) in populations of other ancestries. Effect sizes, associated regions and gene prioritization are similar across ancestries, indicating that reduced prediction accuracy is likely to be explained by linkage disequilibrium and differences in allele frequency within associated regions. Finally, we show that the relevant biological pathways are detectable with smaller sample sizes than are needed to implicate causal genes and variants. Overall, this study provides a comprehensive map of specific genomic regions that contain the vast majority of common height-associated variants. Although this map is saturated for populations of European ancestry, further research is needed to achieve equivalent saturation in other ancestries.

          Abstract

          A large genome-wide association study of more than 5 million individuals reveals that 12,111 single-nucleotide polymorphisms account for nearly all the heritability of height attributable to common genetic variants.

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          Most cited references67

          • Record: found
          • Abstract: found
          • Article: found
          Is Open Access

          A global reference for human genetic variation

          The 1000 Genomes Project set out to provide a comprehensive description of common human genetic variation by applying whole-genome sequencing to a diverse set of individuals from multiple populations. Here we report completion of the project, having reconstructed the genomes of 2,504 individuals from 26 populations using a combination of low-coverage whole-genome sequencing, deep exome sequencing, and dense microarray genotyping. We characterized a broad spectrum of genetic variation, in total over 88 million variants (84.7 million single nucleotide polymorphisms (SNPs), 3.6 million short insertions/deletions (indels), and 60,000 structural variants), all phased onto high-quality haplotypes. This resource includes >99% of SNP variants with a frequency of >1% for a variety of ancestries. We describe the distribution of genetic variation across the global sample, and discuss the implications for common disease studies.
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            • Article: found
            Is Open Access

            The UK Biobank resource with deep phenotyping and genomic data

            The UK Biobank project is a prospective cohort study with deep genetic and phenotypic data collected on approximately 500,000 individuals from across the United Kingdom, aged between 40 and 69 at recruitment. The open resource is unique in its size and scope. A rich variety of phenotypic and health-related information is available on each participant, including biological measurements, lifestyle indicators, biomarkers in blood and urine, and imaging of the body and brain. Follow-up information is provided by linking health and medical records. Genome-wide genotype data have been collected on all participants, providing many opportunities for the discovery of new genetic associations and the genetic bases of complex traits. Here we describe the centralized analysis of the genetic data, including genotype quality, properties of population structure and relatedness of the genetic data, and efficient phasing and genotype imputation that increases the number of testable variants to around 96 million. Classical allelic variation at 11 human leukocyte antigen genes was imputed, resulting in the recovery of signals with known associations between human leukocyte antigen alleles and many diseases.
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              GCTA: a tool for genome-wide complex trait analysis.

              For most human complex diseases and traits, SNPs identified by genome-wide association studies (GWAS) explain only a small fraction of the heritability. Here we report a user-friendly software tool called genome-wide complex trait analysis (GCTA), which was developed based on a method we recently developed to address the "missing heritability" problem. GCTA estimates the variance explained by all the SNPs on a chromosome or on the whole genome for a complex trait rather than testing the association of any particular SNP to the trait. We introduce GCTA's five main functions: data management, estimation of the genetic relationships from SNPs, mixed linear model analysis of variance explained by the SNPs, estimation of the linkage disequilibrium structure, and GWAS simulation. We focus on the function of estimating the variance explained by all the SNPs on the X chromosome and testing the hypotheses of dosage compensation. The GCTA software is a versatile tool to estimate and partition complex trait variation with large GWAS data sets.
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                Author and article information

                Contributors
                l.yengo@imb.uq.edu.au
                yokada@sg.med.osaka-u.ac.jp
                A.R.Wood@exeter.ac.uk
                peter.visscher@uq.edu.au
                Joel.Hirschhorn@childrens.harvard.edu
                Journal
                Nature
                Nature
                Nature
                Nature Publishing Group UK (London )
                0028-0836
                1476-4687
                12 October 2022
                12 October 2022
                2022
                : 610
                : 7933
                : 704-712
                Affiliations
                [1 ]GRID grid.1003.2, ISNI 0000 0000 9320 7537, Institute for Molecular Bioscience, , The University of Queensland, ; Brisbane, Queensland Australia
                [2 ]GRID grid.2515.3, ISNI 0000 0004 0378 8438, Division of Endocrinology, , Boston Children’s Hospital, ; Boston, MA USA
                [3 ]GRID grid.66859.34, ISNI 0000 0004 0546 1623, Program in Medical and Population Genetics, , Broad Institute of MIT and Harvard, ; Cambridge, MA USA
                [4 ]GRID grid.4868.2, ISNI 0000 0001 2171 1133, William Harvey Research Institute, Barts and the London School of Medicine and Dentistry, , Queen Mary University of London, ; London, UK
                [5 ]GRID grid.38142.3c, ISNI 000000041936754X, Harvard Medical School, ; Boston, MA USA
                [6 ]GRID grid.509459.4, ISNI 0000 0004 0472 0267, Laboratory for Statistical Analysis, RIKEN Center for Integrative Medical Sciences, ; Yokohama, Japan
                [7 ]GRID grid.136593.b, ISNI 0000 0004 0373 3971, Department of Statistical Genetics, , Osaka University Graduate School of Medicine, ; Osaka, Japan
                [8 ]GRID grid.38142.3c, ISNI 000000041936754X, Divisions of Genetics and Rheumatology, Brigham and Women’s Hospital and Department of Medicine, , Harvard Medical School, ; Boston, MA USA
                [9 ]GRID grid.10698.36, ISNI 0000000122483208, Department of Epidemiology, Gillings School of Global Public Health, , University of North Carolina at Chapel Hill, ; Chapel Hill, NC USA
                [10 ]GRID grid.5254.6, ISNI 0000 0001 0674 042X, COPSAC, Copenhagen Prospective Studies on Asthma in Childhood, Herlev and Gentofte Hospital, , University of Copenhagen, ; Copenhagen, Denmark
                [11 ]GRID grid.5170.3, ISNI 0000 0001 2181 8870, Section for Bioinformatics, Department of Health Technology, , Technical University of Denmark, ; Copenhagen, Denmark
                [12 ]GRID grid.420283.f, ISNI 0000 0004 0626 0858, 23andMe, ; Sunnyvale, CA USA
                [13 ]GRID grid.280893.8, ISNI 0000 0004 0419 5175, Department of Veterans Affairs, , Eastern Colorado Healthcare System, ; Aurora, CO USA
                [14 ]GRID grid.430503.1, ISNI 0000 0001 0703 675X, Division of Biomedical Informatics and Personalized Medicine, , University of Colorado Anschutz Medical Campus, ; Aurora, CO USA
                [15 ]GRID grid.48336.3a, ISNI 0000 0004 1936 8075, Division of Cancer Epidemiology and Genetics, , National Cancer Institute, ; Rockville, MD USA
                [16 ]GRID grid.214458.e, ISNI 0000000086837370, Department of Internal Medicine, Division of Cardiology, , University of Michigan, ; Ann Arbor, MI USA
                [17 ]GRID grid.62560.37, ISNI 0000 0004 0378 8294, Division of Genetics, Department of Medicine, , Brigham and Women’s Hospital, ; Boston, MA USA
                [18 ]GRID grid.38142.3c, ISNI 000000041936754X, Department of Medicine, , Harvard Medical School, ; Boston, MA USA
                [19 ]GRID grid.10698.36, ISNI 0000000122483208, Department of Genetics, , University of North Carolina at Chapel Hill, ; Chapel Hill, NC USA
                [20 ]GRID grid.266683.f, ISNI 0000 0001 2166 5835, Department of Biostatistics and Epidemiology, School of Public Health and Health Sciences, , University of Massachusetts, ; Amherst, MA USA
                [21 ]GRID grid.214458.e, ISNI 0000000086837370, Department of Biostatistics and Center for Statistical Genetics, , University of Michigan School of Public Health, ; Ann Arbor, MI USA
                [22 ]GRID grid.241167.7, ISNI 0000 0001 2185 3318, Department of Biostatistics and Data Science, , Wake Forest School of Medicine, ; Winston-Salem, NC USA
                [23 ]GRID grid.4991.5, ISNI 0000 0004 1936 8948, Big Data Institute, Li Ka Shing Centre for Health Information and Discovery, , University of Oxford, ; Oxford, UK
                [24 ]GRID grid.8391.3, ISNI 0000 0004 1936 8024, Genetics of Complex Traits, College of Medicine and Health, , University of Exeter, ; Exeter, UK
                [25 ]GRID grid.5254.6, ISNI 0000 0001 0674 042X, Center for Health Data Science, Faculty of Health and Medical Sciences, , University of Copenhagen, ; Copenhagen, Denmark
                [26 ]GRID grid.4991.5, ISNI 0000 0004 1936 8948, Wellcome Centre for Human Genetics, Nuffield Department of Medicine, , University of Oxford, ; Oxford, UK
                [27 ]GRID grid.4991.5, ISNI 0000 0004 1936 8948, Nuffield Department of Population Health, , University of Oxford, ; Oxford, UK
                [28 ]GRID grid.10939.32, ISNI 0000 0001 0943 7661, Institute of Genomics, Estonian Genome Centre, , University of Tartu, ; Tartu, Estonia
                [29 ]GRID grid.5645.2, ISNI 000000040459992X, Department of Internal Medicine, Erasmus MC, , University Medical Center Rotterdam, ; Rotterdam, The Netherlands
                [30 ]GRID grid.62562.35, ISNI 0000000100301493, Division of Biostatistics and Epidemiology, , RTI International, ; Durham, NC USA
                [31 ]GRID grid.9851.5, ISNI 0000 0001 2165 4204, Center for Primary Care and Public Health, , University of Lausanne, ; Lausanne, Switzerland
                [32 ]GRID grid.419765.8, ISNI 0000 0001 2223 3006, Swiss Institute of Bioinformatics, ; Lausanne, Switzerland
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                [35 ]GRID grid.419658.7, ISNI 0000 0004 0646 7285, Steno Diabetes Center Copenhagen, ; Herlev, Denmark
                [36 ]GRID grid.5254.6, ISNI 0000 0001 0674 042X, Department of Biology, The Bioinformatics Center, , University of Copenhagen, ; Copenhagen, Denmark
                [37 ]GRID grid.177174.3, ISNI 0000 0001 2242 4849, Department of Ophthalmology, Graduate School of Medical Sciences, , Kyushu University, ; Fukuoka, Japan
                [38 ]GRID grid.266100.3, ISNI 0000 0001 2107 4242, Department of Family Medicine, , University of California, San Diego, ; La Jolla, CA USA
                [39 ]GRID grid.19006.3e, ISNI 0000 0000 9632 6718, Department of Human Genetics, , David Geffen School of Medicine at UCLA, ; Los Angeles, CA USA
                [40 ]GRID grid.5254.6, ISNI 0000 0001 0674 042X, Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, , University of Copenhagen, ; Copenhagen, Denmark
                [41 ]GRID grid.4494.d, ISNI 0000 0000 9558 4598, Department of Epidemiology, , University of Groningen, University Medical Center Groningen, ; Groningen, The Netherlands
                [42 ]GRID grid.411036.1, ISNI 0000 0001 1498 685X, Department of Bioinformatics, , Isfahan University of Medical Sciences, ; Isfahan, Iran
                [43 ]GRID grid.4973.9, ISNI 0000 0004 0646 7373, Institute of Biological Psychiatry, Mental Health Services, , Copenhagen University Hospital, ; Copenhagen, Denmark
                [44 ]GRID grid.10698.36, ISNI 0000000122483208, Thurston Arthritis Research Center, , University of North Carolina at Chapel Hill, ; Chapel Hill, NC USA
                [45 ]GRID grid.417634.3, ISNI 0000 0004 0496 8123, Genomic Research on Complex diseases (GRC-Group), , CSIR-Centre for Cellular and Molecular Biology, ; Hyderabad, India
                [46 ]GRID grid.214458.e, ISNI 0000000086837370, Department of Epidemiology, , University of Michigan School of Public Health, ; Ann Arbor, MI USA
                [47 ]GRID grid.7737.4, ISNI 0000 0004 0410 2071, Institute for Molecular Medicine Finland (FIMM), HiLIFE, , University of Helsinki, ; Helsinki, Finland
                [48 ]GRID grid.94365.3d, ISNI 0000 0001 2297 5165, Molecular Genetics Section, Center for Precision Health Research, National Human Genome Research Institute, , National Institutes of Health, ; Bethesda, MD USA
                [49 ]GRID grid.239552.a, ISNI 0000 0001 0680 8770, Center for Applied Genomics, , Children’s Hospital of Philadelphia, ; Philadelphia, PA USA
                [50 ]Quantinuum Research, Wayne, PA USA
                [51 ]GRID grid.25879.31, ISNI 0000 0004 1936 8972, Department of Genetics, , University of Pennsylvania, ; Philadelphia, PA USA
                [52 ]GRID grid.9918.9, ISNI 0000 0004 1936 8411, Department of Cardiovascular Sciences, , University of Leicester, ; Leicester, UK
                [53 ]GRID grid.412925.9, ISNI 0000 0004 0400 6581, NIHR Leicester Biomedical Research Centre, , Glenfield Hospital, ; Leicester, UK
                [54 ]GRID grid.34477.33, ISNI 0000000122986657, Cardiovascular Health Research Unit, Department of Medicine, , University of Washington, ; Seattle, WA USA
                [55 ]GRID grid.475435.4, Department of Clinical Immunology, , Copenhagen University Hospital, Rigshospitalet, ; Copenhagen, Denmark
                [56 ]GRID grid.5254.6, ISNI 0000 0001 0674 042X, NovoNordic Center for Protein Research, , Copenhagen University, ; Copenhagen, Denmark
                [57 ]GRID grid.62560.37, ISNI 0000 0004 0378 8294, Department of Medicine, , Brigham and Women’s Hospital, ; Boston, MA USA
                [58 ]GRID grid.412807.8, ISNI 0000 0004 1936 9916, Division of Epidemiology, Department of Medicine, , Vanderbilt University Medical Center, ; Nashville, TN USA
                [59 ]GRID grid.4305.2, ISNI 0000 0004 1936 7988, Centre for Genomic and Experimental Medicine, Institute of Genetics and Cancer, , University of Edinburgh, ; Edinburgh, UK
                [60 ]GRID grid.4777.3, ISNI 0000 0004 0374 7521, Centre for Public Health, , Queen’s University of Belfast, ; Belfast, UK
                [61 ]GRID grid.418712.9, ISNI 0000 0004 1760 7415, Institute for Maternal and Child Health – IRCCS, Burlo Garofolo, ; Trieste, Italy
                [62 ]GRID grid.419272.b, ISNI 0000 0000 9960 1711, Ocular Epidemiology, Singapore Eye Research Institute, , Singapore National Eye Centre, ; Singapore, Singapore
                [63 ]GRID grid.4280.e, ISNI 0000 0001 2180 6431, Department of Ophthalmology, , National University of Singapore and National University Health System, ; Singapore, Singapore
                [64 ]GRID grid.482251.8, ISNI 0000 0004 0633 7958, Institute of Biomedical Sciences, Academia Sinica, ; Taipei, Taiwan
                [65 ]GRID grid.19188.39, ISNI 0000 0004 0546 0241, Graduate Institute of Medical Genomics and Proteomics, Medical College, , National Taiwan University, ; Taipei, Taiwan
                [66 ]GRID grid.412094.a, ISNI 0000 0004 0572 7815, Department of Internal Medicine, , National Taiwan University Hospital, ; Taipei, Taiwan
                [67 ]GRID grid.25879.31, ISNI 0000 0004 1936 8972, Department of Pathology and Laboratory Medicine, , University of Pennsylvania, ; Philadelphia, PA USA
                [68 ]GRID grid.239552.a, ISNI 0000 0001 0680 8770, Center for Spatial and Functional Genomics, Division of Human Genetics, , Children’s Hospital of Philadelphia, ; Philadelphia, PA USA
                [69 ]GRID grid.66859.34, ISNI 0000 0004 0546 1623, Cardiovascular Disease Initiative, , Broad Institute of MIT and Harvard, ; Cambridge, MA USA
                [70 ]GRID grid.59784.37, ISNI 0000000406229172, Institute of Population Health Sciences, , National Health Research Institutes, ; Zhunan, Taiwan
                [71 ]GRID grid.23856.3a, ISNI 0000 0004 1936 8390, Department of Kinesiology, Faculty of Medicine, , Université Laval, ; Québec City, Quebec Canada
                [72 ]GRID grid.1003.2, ISNI 0000 0000 9320 7537, Diamantina Institute, , The University of Queensland, ; Brisbane, Queensland Australia
                [73 ]GRID grid.62560.37, ISNI 0000 0004 0378 8294, Division of Preventive Medicine, , Brigham and Women’s Hospital, ; Boston, MA USA
                [74 ]GRID grid.4367.6, ISNI 0000 0001 2355 7002, Division of Statistical Genomics, Department of Genetics, , Washington University School of Medicine, ; St Louis, MO USA
                [75 ]GRID grid.9764.c, ISNI 0000 0001 2153 9986, Institute of Clinical Molecular Biology, , Christian-Albrechts University of Kiel, ; Kiel, Germany
                [76 ]GRID grid.7700.0, ISNI 0000 0001 2190 4373, Vth Department of Medicine, Medical Faculty Mannheim, , Heidelberg University, ; Mannheim, Germany
                [77 ]GRID grid.11450.31, ISNI 0000 0001 2097 9138, Dipartimento di Scienze Mediche Chirurgiche e Sperimentali, , Università degli Studi di Sassari, ; Sassari, Italy
                [78 ]GRID grid.5645.2, ISNI 000000040459992X, Department of Epidemiology, Erasmus MC, , University Medical Center Rotterdam, ; Rotterdam, The Netherlands
                [79 ]GRID grid.5475.3, ISNI 0000 0004 0407 4824, Section of Statistical Multi-omics, Department of Clinical and Experimental Medicine, , University of Surrey, ; Guildford, UK
                [80 ]GRID grid.189504.1, ISNI 0000 0004 1936 7558, Department of Biostatistics, , Boston University School of Public Health, ; Boston, MA USA
                [81 ]GRID grid.410370.1, ISNI 0000 0004 4657 1992, Center for Data and Computational Sciences, , VA Boston Healthcare System, ; Boston, MA USA
                [82 ]GRID grid.7727.5, ISNI 0000 0001 2190 5763, Department of Genetic Epidemiology, , University of Regensburg, ; Regensburg, Germany
                [83 ]GRID grid.411941.8, ISNI 0000 0000 9194 7179, Department of Internal Medicine II, , University Hospital Regensburg, ; Regensburg, Germany
                [84 ]GRID grid.15823.3d, ISNI 0000 0004 0622 2843, Department of Nutrition and Dietetics, School of Health and Education, , Harokopio University of Athens, ; Athens, Greece
                [85 ]GRID grid.412860.9, ISNI 0000 0004 0459 1231, Center for Precision Medicine, Wake Forest School of Medicine, , Medical Center Boulevard, ; Winston-Salem, NC USA
                [86 ]GRID grid.185448.4, ISNI 0000 0004 0637 0221, Genome Institute of Singapore, , Agency for Science, Technology and Research, ; Singapore, Singapore
                [87 ]GRID grid.428397.3, ISNI 0000 0004 0385 0924, Health Services and Systems Research, , Duke-NUS Medical School, ; Singapore, Singapore
                [88 ]GRID grid.5330.5, ISNI 0000 0001 2107 3311, Institute of Human Genetics, Universitätsklinikum Erlangen, , Friedrich-Alexander-Universität Erlangen-Nürnberg, ; Erlangen, Germany
                [89 ]GRID grid.83440.3b, ISNI 0000000121901201, Institute of Cardiovascular Science, Faculty of Population Health, , University College London, ; London, UK
                [90 ]GRID grid.214458.e, ISNI 0000000086837370, Survey Research Center, Institute for Social Research, , University of Michigan, ; Ann Arbor, MI USA
                [91 ]GRID grid.452651.1, ISNI 0000 0004 0627 7633, Computational Genomics Department, , National Institute of Genomic Medicine, ; Mexico City, Mexico
                [92 ]GRID grid.5361.1, ISNI 0000 0000 8853 2677, Institute of Genetic Epidemiology, , Medical University of Innsbruck, ; Innsbruck, Austria
                [93 ]GRID grid.5133.4, ISNI 0000 0001 1941 4308, Department of Medicine, Surgery and Health Sciences, , University of Trieste, ; Trieste, Italy
                [94 ]GRID grid.9764.c, ISNI 0000 0001 2153 9986, Institute of Medical Informatics and Statistics, , Kiel University, ; Kiel, Germany
                [95 ]GRID grid.511439.b, Eurac Research, , Institute for Biomedicine, Affiliated Institute of the University of Lübeck, ; Bolzano, Italy
                [96 ]GRID grid.10417.33, ISNI 0000 0004 0444 9382, Radboud University Medical Center, Radboud Institute for Health Sciences, Department for Health Evidence, ; Nijmegen, The Netherlands
                [97 ]GRID grid.251313.7, ISNI 0000 0001 2169 2489, Jackson Heart Study, Department of Medicine, , University of Mississippi, ; Jackson, MS USA
                [98 ]GRID grid.410745.3, ISNI 0000 0004 1765 1045, Nanjing University of Chinese Medicine, ; Nanjing, China
                [99 ]GRID grid.4567.0, ISNI 0000 0004 0483 2525, Research Unit of Molecular Epidemiology, Institute of Epidemiology, , Helmholtz Zentrum München Research Center for Environmental Health, ; Neuherberg, Germany
                [100 ]GRID grid.417834.d, Institute of Epidemiology, Helmholtz Zentrum München Research Center for Environmental Health, ; Neuherberg, Germany
                [101 ]GRID grid.6203.7, ISNI 0000 0004 0417 4147, Department of Epidemiology Research, , Statens Serum Institut, ; Copenhagen, Denmark
                [102 ]GRID grid.4991.5, ISNI 0000 0004 1936 8948, Cardiovascular Medicine, Radcliffe Department of Medicine, , University of Oxford, John Radcliffe Hospital, ; Oxford, UK
                [103 ]GRID grid.1049.c, ISNI 0000 0001 2294 1395, Genetic Epidemiology, , QIMR Berghofer Medical Research Institute, ; Brisbane, Queensland Australia
                [104 ]GRID grid.7048.b, ISNI 0000 0001 1956 2722, Department of Biomedicine (Human Genetics) and iSEQ Center, , Aarhus University, ; Aarhus, Denmark
                [105 ]GRID grid.452548.a, ISNI 0000 0000 9817 5300, The Lundbeck Foundation Initiative for Integrative Psychiatric Research, iPSYCH, ; Aarhus, Denmark
                [106 ]GRID grid.7048.b, ISNI 0000 0001 1956 2722, BiRC—Bioinformatics Research Centre, , Aarhus University, ; Aarhus, Denmark
                [107 ]GRID grid.513199.6, The Institute for Translational Genomics and Population Sciences, Department of Pediatrics, , The Lundquist Institute for Biomedical Innovation at Harbor-UCLA Medical Center, ; Torrance, CA USA
                [108 ]GRID grid.8993.b, ISNI 0000 0004 1936 9457, Department of Medical Sciences, , Uppsala University, ; Uppsala, Sweden
                [109 ]GRID grid.270240.3, ISNI 0000 0001 2180 1622, Division of Public Health Sciences, , Fred Hutchinson Cancer Research Center, ; Seattle, WA USA
                [110 ]GRID grid.475435.4, Danish Headache Center, Department of Neurology, , Copenhagen University Hospital, Rigshospitalet, ; Rigshospitalet, Copenhagen Denmark
                [111 ]GRID grid.14758.3f, ISNI 0000 0001 1013 0499, Department of Public Health and Welfare, , Finnish Institute for Health and Welfare, ; Helsinki, Finland
                [112 ]GRID grid.5337.2, ISNI 0000 0004 1936 7603, MRC Integrative Epidemiology Unit, , University of Bristol, ; Bristol, UK
                [113 ]GRID grid.5337.2, ISNI 0000 0004 1936 7603, Bristol Dental School, , University of Bristol, ; Bristol, UK
                [114 ]GRID grid.189504.1, ISNI 0000 0004 1936 7558, Department of Neurology, , Boston University School of Medicine, ; Boston, MA USA
                [115 ]GRID grid.510954.c, ISNI 0000 0004 0444 3861, Framingham Heart Study, ; Framingham, MA USA
                [116 ]GRID grid.452356.3, ISNI 0000 0004 0518 1285, Department of Genetics and Bioinformatics, , Dasman Diabetes Institute, ; Kuwait City, Kuwait
                [117 ]GRID grid.4514.4, ISNI 0000 0001 0930 2361, Department of Clinical Sciences in Malmö, , Lund University, ; Malmö, Sweden
                [118 ]GRID grid.410370.1, ISNI 0000 0004 4657 1992, Veterans Affairs Boston Healthcare System, ; Boston, MA USA
                [119 ]GRID grid.11598.34, ISNI 0000 0000 8988 2476, Clinical Division of Neurogeriatrics, Department of Neurology, , Medical University of Graz, ; Graz, Austria
                [120 ]GRID grid.11598.34, ISNI 0000 0000 8988 2476, Institute for Medical Informatics, Statistics and Documentation, , Medical University of Graz, ; Graz, Austria
                [121 ]GRID grid.266842.c, ISNI 0000 0000 8831 109X, School of Medicine and Public Health, , University of Newcastle, ; Callaghan, New South Wales Australia
                [122 ]GRID grid.9647.c, ISNI 0000 0004 7669 9786, Institute for Medical Informatics, Statistics and Epidemiology, , University of Leipzig, Medical Faculty, ; Leipzig, Germany
                [123 ]GRID grid.9647.c, ISNI 0000 0004 7669 9786, LIFE Research Center for Civilization Diseases, , University of Leipzig, Medical Faculty, ; Leipzig, Germany
                [124 ]GRID grid.12380.38, ISNI 0000 0004 1754 9227, Department of Biological Psychology, Behaviour and Movement Sciences, , Vrije Universiteit Amsterdam, ; Amsterdam, The Netherlands
                [125 ]GRID grid.38142.3c, ISNI 000000041936754X, Department of Epidemiology, , Harvard T.H. Chan School of Public Health, ; Boston, MA USA
                [126 ]GRID grid.263817.9, ISNI 0000 0004 1773 1790, School of Public Health and Emergency Management, , Southern University of Science and Technology, ; Shenzhen, China
                [127 ]GRID grid.11135.37, ISNI 0000 0001 2256 9319, Institute for Global Health and Development, , Peking University, ; Beijing, China
                [128 ]GRID grid.66859.34, ISNI 0000 0004 0546 1623, Programs in Metabolism and Medical and Population Genetics, , Broad Institute of Harvard and MIT, ; Cambridge, MA USA
                [129 ]GRID grid.9486.3, ISNI 0000 0001 2159 0001, Departamento de Medicina Genómica y Toxicología Ambiental, Instituto de Investigaciones Biomédicas, , Universidad Nacional Autónoma de México Ciudad Universitaria, ; Mexico City, Mexico
                [130 ]GRID grid.416850.e, ISNI 0000 0001 0698 4037, Unidad de Biología Molecular y Medicina Genómica, , Instituto Nacional de Ciencias Médicas y Nutrición, ; Mexico City, Mexico
                [131 ]GRID grid.416121.1, ISNI 0000 0004 0573 0539, Division of Endocrine and Metabolism, , Tri-Service General Hospital Songshan Branch, ; Taipei, Taiwan
                [132 ]GRID grid.410726.6, ISNI 0000 0004 1797 8419, Shanghai Institute of Nutrition and Health, , University of Chinese Academy of Sciences, Chinese Academy of Sciences, ; Shanghai, China
                [133 ]GRID grid.415482.e, ISNI 0000 0004 0647 4899, Division of Genome Science, Department of Precision Medicine, , National Institute of Health, ; Cheongju, Republic of Korea
                [134 ]GRID grid.419953.3, ISNI 0000 0004 1756 0784, Biomedical Technology Research Center, , Tokushima Research Institute, Otsuka Pharmaceutical Co., ; Tokushima, Japan
                [135 ]GRID grid.45203.30, ISNI 0000 0004 0489 0290, Research Institute, National Center for Global Health and Medicine, ; Tokyo, Japan
                [136 ]GRID grid.418213.d, ISNI 0000 0004 0390 0098, Department of Molecular Epidemiology, , German Institute of Human Nutrition Potsdam-Rehbruecke, ; Nuthetal, Germany
                [137 ]GRID grid.452622.5, German Center for Diabetes Research (DZD), ; Neuherberg, Germany
                [138 ]GRID grid.12380.38, ISNI 0000 0004 1754 9227, Department of Complex Trait Genetics, Center for Neurogenomics and Cognitive Research, Amsterdam Neuroscience, , Vrije Universiteit Amsterdam, ; Amsterdam, The Netherlands
                [139 ]GRID grid.16872.3a, ISNI 0000 0004 0435 165X, Department of Child and Adolescent Psychiatry and Pediatric Psychology, Section Complex Trait Genetics, Amsterdam Neuroscience, , Vrije Universiteit Medical Center, ; Amsterdam, The Netherlands
                [140 ]GRID grid.12650.30, ISNI 0000 0001 1034 3451, Department of Biobank Research, , Umeå University, ; Umeå, Sweden
                [141 ]GRID grid.12650.30, ISNI 0000 0001 1034 3451, Department of Odontology, , Umeå University, ; Umeå, Sweden
                [142 ]GRID grid.508836.0, Institute of Molecular and Clinical Ophthalmology Basel, ; Basel, Switzerland
                [143 ]Privatpraxis Prof Jonas und Dr Panda-Jonas, Heidelberg, Germany
                [144 ]GRID grid.7700.0, ISNI 0000 0001 2190 4373, Department of Ophthalmology, Medical Faculty Mannheim, , Heidelberg University, ; Mannheim, Germany
                [145 ]GRID grid.414373.6, ISNI 0000 0004 1758 1243, Beijing Institute of Ophthalmology, Beijing Tongren Eye Center, Beijing Tongren Hospital, , Capital Medical University, Beijing Ophthalmology and Visual Sciences Key Laboratory, ; Beijing, China
                [146 ]GRID grid.4973.9, ISNI 0000 0004 0646 7373, Center for Clinical Research and Prevention, , Copenhagen University Hospital - Bispebjerg and Frederiksberg, ; Copenhagen, Denmark
                [147 ]GRID grid.5254.6, ISNI 0000 0001 0674 042X, Department of Public Health, Faculty of Health and Medical Sciences, , University of Copenhagen, ; Copenhagen, Denmark
                [148 ]GRID grid.45202.31, ISNI 0000 0000 8631 5388, Faculty of Medicine, , University of Kelaniya, ; Ragama, Sri Lanka
                [149 ]GRID grid.136593.b, ISNI 0000 0004 0373 3971, Department of Geriatric and General Medicine, , Osaka University Graduate School of Medicine, ; Suita, Japan
                [150 ]GRID grid.258799.8, ISNI 0000 0004 0372 2033, Center for Genomic Medicine, , Kyoto University Graduate School of Medicine, ; Kyoto, Japan
                [151 ]GRID grid.25879.31, ISNI 0000 0004 1936 8972, Department of Psychiatry, , University of Pennsylvania, ; Philadelphia, PA USA
                [152 ]GRID grid.4305.2, ISNI 0000 0004 1936 7988, Centre for Global Health, Usher Institute, , University of Edinburgh, ; Edinburgh, UK
                [153 ]GRID grid.4305.2, ISNI 0000 0004 1936 7988, Centre for Cardiovascular Sciences, Queens Medical Research Institute, , University of Edinburgh, ; Edinburgh, UK
                [154 ]GRID grid.264381.a, ISNI 0000 0001 2181 989X, Medical Research Institute, Kangbuk Samsung Hospital, , Sungkyunkwan University School of Medicine, ; Seoul, Republic of Korea
                [155 ]GRID grid.264381.a, ISNI 0000 0001 2181 989X, Department of Clinical Research Design and Evaluation (SAIHST), , Sungkyunkwan University, ; Seoul, Republic of Korea
                [156 ]SYNLAB MVZ Humangenetik Mannheim, Mannheim, Germany
                [157 ]GRID grid.4514.4, ISNI 0000 0001 0930 2361, Department of Clinical Sciences, Genetic and Molecular Epidemiology Unit, , Lund University, ; Malmö, Sweden
                [158 ]GRID grid.17063.33, ISNI 0000 0001 2157 2938, Department of Computer Science, , University of Toronto, ; Toronto, Ontario Canada
                [159 ]GRID grid.17063.33, ISNI 0000 0001 2157 2938, Department of Anthropology, , University of Toronto at Mississauga, ; Mississauga, Ontario Canada
                [160 ]GRID grid.1024.7, ISNI 0000000089150953, Genomics Research Centre, Centre for Genomics and Personalised Health, School of Biomedical Sciences, , Queensland University of Technology, ; Kelvin Grove, Queensland Australia
                [161 ]Oneomics, Soonchunhyang Mirai Medical Center, Bucheon-si, Republic of Korea
                [162 ]GRID grid.94365.3d, ISNI 0000 0001 2297 5165, Laboratory of Neurogenetics, National Institute on Aging, National Institutes of Health, ; Bethesda, MD USA
                [163 ]GRID grid.94365.3d, ISNI 0000 0001 2297 5165, Center for Alzheimer’s and Related Dementias, , National Institutes of Health, ; Bethesda, MD USA
                [164 ]GRID grid.511118.d, Data Tecnica International, ; Glen Echo, MD USA
                [165 ]GRID grid.419407.f, ISNI 0000 0004 4665 8158, Cancer Genomics Research Laboratory, , Leidos Biomedical Research, ; Rockville, MD USA
                [166 ]GRID grid.67105.35, ISNI 0000 0001 2164 3847, Department of Population and Quantitative Health Sciences, , Case Western Reserve University, ; Cleveland, OH USA
                [167 ]GRID grid.168645.8, ISNI 0000 0001 0742 0364, Department of Medicine, , University of Massachusetts Chan Medical School, ; Worcester, MA USA
                [168 ]GRID grid.410764.0, ISNI 0000 0004 0573 0731, Center for Geriatrics and Gerontology, , Taichung Veterans General Hospital, ; Taichung, Taiwan
                [169 ]GRID grid.482476.b, ISNI 0000 0000 8995 9090, Montreal Heart Institute, ; Montreal, Quebec Canada
                [170 ]GRID grid.10417.33, ISNI 0000 0004 0444 9382, Departments of Ophthalmology and Human Genetics, , Radboud University Nijmegen Medical Center, ; Nijmegen, The Netherlands
                [171 ]GRID grid.5335.0, ISNI 0000000121885934, MRC Epidemiology Unit, , University of Cambridge School of Clinical Medicine, ; Cambridge, UK
                [172 ]GRID grid.7914.b, ISNI 0000 0004 1936 7443, Department of Clinical Science, Center for Diabetes Research, , University of Bergen, ; Bergen, Norway
                [173 ]GRID grid.4514.4, ISNI 0000 0001 0930 2361, Department of Clinical Sciences, , Lund University Diabetes Centre, ; Malmö, Sweden
                [174 ]GRID grid.511163.1, ISNI 0000 0004 0518 4910, Department of Clinical Chemistry, , Fimlab Laboratories, ; Tampere, Finland
                [175 ]GRID grid.502801.e, ISNI 0000 0001 2314 6254, Department of Clinical Chemistry, Finnish Cardiovascular Research Center - Tampere, Faculty of Medicine and Health Technology, , Tampere University, ; Tampere, Finland
                [176 ]GRID grid.412330.7, ISNI 0000 0004 0628 2985, Department of Cardiology, Heart Center, , Tampere University Hospital, ; Tampere, Finland
                [177 ]GRID grid.5216.0, ISNI 0000 0001 2155 0800, National and Kapodistrian University of Athens, Dromokaiteio Psychiatric Hospital, ; Athens, Greece
                [178 ]GRID grid.13097.3c, ISNI 0000 0001 2322 6764, Department of Twin Research and Genetic Epidemiology, , King’s College London, ; London, UK
                [179 ]GRID grid.420545.2, ISNI 0000 0004 0489 3985, NIHR Biomedical Research Centre at Guy’s and St Thomas’ Foundation Trust, ; London, UK
                [180 ]GRID grid.27755.32, ISNI 0000 0000 9136 933X, Center for Public Health Genomics, , University of Virginia School of Medicine, ; Charlottesville, VA USA
                [181 ]GRID grid.417068.c, ISNI 0000 0004 0624 9907, MRC Human Genetics Unit, Institute of Genetics and Cancer, , University of Edinburgh, Western General Hospital, ; Edinburgh, UK
                [182 ]GRID grid.55602.34, ISNI 0000 0004 1936 8200, Department of Psychiatry and Department of Community Health and Epidemiology, , Dalhousie University, ; Halifax, Nova Scotia Canada
                [183 ]GRID grid.5252.0, ISNI 0000 0004 1936 973X, Institute of Psychiatric Phenomics and Genomics (IPPG), , University Hospital, LMU Munich, ; Munich, Germany
                [184 ]GRID grid.410718.b, ISNI 0000 0001 0262 7331, Institute for Medical Informatics, Biometry and Epidemiology, , University Hospital Essen, ; Essen, Germany
                [185 ]GRID grid.32224.35, ISNI 0000 0004 0386 9924, Diabetes Unit, , Massachusetts General Hospital, ; Boston, MA USA
                [186 ]GRID grid.32224.35, ISNI 0000 0004 0386 9924, Center for Genomic Medicine, , Massachusetts General Hospital, ; Boston, MA USA
                [187 ]GRID grid.509540.d, ISNI 0000 0004 6880 3010, Department of Psychiatry, Amsterdam Public Health and Amsterdam Neuroscience, , Amsterdam UMC and Vrije Universiteit, ; Amsterdam, The Netherlands
                [188 ]Biomedical and Translational Informatics Institute, Geisinger, Danville, PA USA
                [189 ]GRID grid.25879.31, ISNI 0000 0004 1936 8972, Department of Genetics, Institute for Biomedical Informatics, Perelman School of Medicine, , University of Pennsylvania, ; Philadelphia, PA USA
                [190 ]GRID grid.4991.5, ISNI 0000 0004 1936 8948, MRC Population Health Research Unit, Nuffield Department of Population Health, , University of Oxford, ; Oxford, UK
                [191 ]GRID grid.5337.2, ISNI 0000 0004 1936 7603, Population Health Sciences, Bristol Medical School, , University of Bristol, ; Bristol, UK
                [192 ]GRID grid.4562.5, ISNI 0000 0001 0057 2672, Institute for Cardiogenetics, , University of Lübeck, DZHK (German Research Centre for Cardiovascular Research) partner site Hamburg/Lübeck/Kiel and University Heart Center Lübeck, ; Lübeck, Germany
                [193 ]GRID grid.27476.30, ISNI 0000 0001 0943 978X, Public Health Informatics Unit, Department of Integrated Health Sciences, , Nagoya University Graduate School of Medicine, ; Nagoya, Japan
                [194 ]GRID grid.8761.8, ISNI 0000 0000 9919 9582, Centre for Bone and Arthritis Research, Department of Internal Medicine and Clinical Nutrition, Institute of Medicine, Sahlgrenska Academy, , University of Gothenburg, ; Gothenburg, Sweden
                [195 ]GRID grid.8761.8, ISNI 0000 0000 9919 9582, Bioinformatics Core Facility, Sahlgrenska Academy, , University of Gothenburg, ; Gothenburg, Sweden
                [196 ]GRID grid.35541.36, ISNI 0000000121053345, Korea Institute of Science and Technology, , Gangneung Institute of Natural Products, ; Gangneung, Republic of Korea
                [197 ]GRID grid.459623.f, ISNI 0000 0004 0587 0347, Department of Clinical Biochemistry, , Lillebaelt Hospital, ; Kolding, Denmark
                [198 ]GRID grid.10306.34, ISNI 0000 0004 0606 5382, Department of Human Genetics, , Wellcome Sanger Institute, ; Hinxton, UK
                [199 ]GRID grid.10419.3d, ISNI 0000000089452978, Department of Internal Medicine, Section of Gerontology and Geriatrics, , Leiden University Medical Center, ; Leiden, The Netherlands
                [200 ]GRID grid.5326.2, ISNI 0000 0001 1940 4177, Institute of Genetics and Biophysics A. Buzzati-Traverso, CNR, ; Naples, Italy
                [201 ]GRID grid.416811.b, ISNI 0000 0004 0631 6436, Department of Clinical Biochemistry and Immunology, , Hospital of Southern Jutland, ; Aabenraa, Denmark
                [202 ]GRID grid.7048.b, ISNI 0000 0001 1956 2722, The National Centre for Register-based Research, , University of Aarhus, ; Aarhus, Denmark
                [203 ]GRID grid.1374.1, ISNI 0000 0001 2097 1371, Centre for Population Health Research, , University of Turku and Turku University Hospital, ; Turku, Finland
                [204 ]GRID grid.1374.1, ISNI 0000 0001 2097 1371, Research Centre of Applied and Preventive Cardiovascular Medicine, , University of Turku, ; Turku, Finland
                [205 ]GRID grid.38603.3e, ISNI 0000 0004 0644 1675, Medical School, University of Split, ; Split, Croatia
                [206 ]GRID grid.509547.a, Algebra University College, ; Zagreb, Croatia
                [207 ]GRID grid.59734.3c, ISNI 0000 0001 0670 2351, The Charles Bronfman Institute for Personalized Medicine, , Icahn School of Medicine at Mount Sinai, ; New York, NY USA
                [208 ]GRID grid.59734.3c, ISNI 0000 0001 0670 2351, Department of Environmental Medicine and Public Health, , Icahn School of Medicine at Mount Sinai, ; New York, NY USA
                [209 ]GRID grid.11205.37, ISNI 0000 0001 2152 8769, Aragon Institute of Engineering Research, , University of Zaragoza, ; Zaragoza, Spain
                [210 ]GRID grid.429738.3, ISNI 0000 0004 1763 291X, Centro de Investigación Biomédica en Red en Bioingeniería, Biomateriales y Nanomedicina (CIBER-BBN), ; Madrid, Spain
                [211 ]GRID grid.497620.e, Center for Non-Communicable Diseases, ; Karachi, Pakistan
                [212 ]GRID grid.4494.d, ISNI 0000 0000 9558 4598, Department of Psychiatry, Interdisciplinary Center Psychopathology and Emotion Regulation, , University of Groningen, University Medical Center Groningen, ; Groningen, The Netherlands
                [213 ]GRID grid.4567.0, ISNI 0000 0004 0483 2525, Institute of Translational Genomics, , Helmholtz Zentrum München, German Research Center for Environmental Health, ; Neuherberg, Germany
                [214 ]GRID grid.415719.f, ISNI 0000 0004 0488 9484, Present Address: Oxford Centre for Diabetes, Endocrinology and Metabolism, Radcliffe Department of Medicine, , University of Oxford, Churchill Hospital, ; Oxford, UK
                [215 ]GRID grid.413648.c, Hunter Medical Research Institute, ; New Lambton Heights, New South Wales Australia
                [216 ]GRID grid.266842.c, ISNI 0000 0000 8831 109X, School of Medicine and Public Health, College of Health, Medicine and Wellbeing, The University of Newcastle, ; New Lambton Heights, New South Wales Australia
                [217 ]GRID grid.419543.e, ISNI 0000 0004 1760 3561, IRCCS Neuromed, ; Pozzilli, Italy
                [218 ]GRID grid.411024.2, ISNI 0000 0001 2175 4264, Department of Medicine, Division of Endocrinology, Diabetes and Nutrition, , University of Maryland School of Medicine, ; Baltimore, MD USA
                [219 ]GRID grid.411024.2, ISNI 0000 0001 2175 4264, Program for Personalized and Genomic Medicine, , University of Maryland School of Medicine, ; Baltimore, MD USA
                [220 ]GRID grid.413396.a, ISNI 0000 0004 1768 8905, Unit of Genomics of Complex Diseases, , Sant Pau Biomedical Research Institute (IIB Sant Pau), ; Barcelona, Spain
                [221 ]GRID grid.4714.6, ISNI 0000 0004 1937 0626, Cardiovascular Medicine Unit, Department of Medicine, , Karolinska Institutet, Center for Molecular Medicine, ; Stockholm, Sweden
                [222 ]GRID grid.94365.3d, ISNI 0000 0001 2297 5165, Laboratory of Epidemiology and Population Sciences, National Institute on Aging, , National Institutes of Health, ; Baltimore, MD USA
                [223 ]GRID grid.256753.0, ISNI 0000 0004 0470 5964, Department of Biomedical Science, , Hallym University, ; Chuncheon, Republic of Korea
                [224 ]GRID grid.5326.2, ISNI 0000 0001 1940 4177, Istituto di Ricerca Genetica e Biomedica, , Consiglio Nazionale delle Ricerche (CNR), ; Cagliari, Italy
                [225 ]GRID grid.10419.3d, ISNI 0000000089452978, Department of Cell and Chemical Biology, , Leiden University Medical Center, ; Leiden, The Netherlands
                [226 ]GRID grid.509540.d, ISNI 0000 0004 6880 3010, Epidemiology and Data Science, , Amsterdam UMC, location Vrije Universiteit Amsterdam, ; Amsterdam, The Netherlands
                [227 ]Amsterdam Cardiovascular Sciences, Amsterdam, The Netherlands
                [228 ]GRID grid.10419.3d, ISNI 0000000089452978, Department of Clinical Epidemiology, , Leiden University Medical Center, ; Leiden, The Netherlands
                [229 ]GRID grid.420802.c, ISNI 0000 0000 9458 5898, Icelandic Heart Association, ; Kópavogur, Iceland
                [230 ]GRID grid.10306.34, ISNI 0000 0004 0606 5382, Wellcome Sanger Institute, ; Hinxton, UK
                [231 ]GRID grid.421812.c, ISNI 0000 0004 0618 6889, deCODE Genetics/Amgen, ; Reykjavik, Iceland
                [232 ]GRID grid.7914.b, ISNI 0000 0004 1936 7443, Mohn Nutrition Research Laboratory, Department of Clinical Science, , University of Bergen, ; Bergen, Norway
                [233 ]GRID grid.164971.c, ISNI 0000 0001 1089 6558, Department of Public Health Sciences, Parkinson School of Health Sciences and Public Health, , Loyola University Chicago, ; Maywood, IL USA
                [234 ]GRID grid.280747.e, ISNI 0000 0004 0419 2556, VA Palo Alto Health Care System, ; Palo Alto, CA USA
                [235 ]GRID grid.168010.e, ISNI 0000000419368956, Department of Medicine, , Stanford University School of Medicine, ; Stanford, CA USA
                [236 ]GRID grid.5603.0, Institute for Community Medicine, , University Medicine Greifswald, ; Greifswald, Germany
                [237 ]GRID grid.452396.f, ISNI 0000 0004 5937 5237, DZHK (German Centre for Cardiovascular Research), partner site Greifswald, ; Greifswald, Germany
                [238 ]GRID grid.266102.1, ISNI 0000 0001 2297 6811, Cardiology Division, Department of Pediatrics, , University of California, San Francisco, ; Oakland, CA USA
                [239 ]GRID grid.5335.0, ISNI 0000000121885934, Centre for Cancer Genetic Epidemiology, , University of Cambridge, ; Cambridge, UK
                [240 ]GRID grid.5335.0, ISNI 0000000121885934, Department of Public Health and Primary Care, , University of Cambridge, ; Cambridge, UK
                [241 ]GRID grid.10419.3d, ISNI 0000000089452978, Department of Cardiology, , Leiden University Medical Center, ; Leiden, The Netherlands
                [242 ]GRID grid.5477.1, ISNI 0000000120346234, Central Diagnostics Laboratory, Division Laboratories, Pharmacy and Biomedical Genetics, , University Medical Center Utrecht, Utrecht University, ; Utrecht, The Netherlands
                [243 ]GRID grid.10419.3d, ISNI 0000000089452978, Department of Human Genetics, , Leiden University Medical Center, ; Leiden, The Netherlands
                [244 ]GRID grid.7177.6, ISNI 0000000084992262, Laboratory Genetic Metabolic Diseases, , Department of Clinical Chemistry, Amsterdam UMC, University of Amsterdam, ; Amsterdam, The Netherlands
                [245 ]GRID grid.7177.6, ISNI 0000000084992262, Core Facility Metabolomics, Amsterdam UMC, , University of Amsterdam, ; Amsterdam, The Netherlands
                [246 ]GRID grid.5477.1, ISNI 0000000120346234, Department of Cardiology, Division Heart and Lungs, , University Medical Center Utrecht, Utrecht University, ; Utrecht, The Netherlands
                [247 ]GRID grid.4494.d, ISNI 0000 0000 9558 4598, Department of Cardiology, University of Groningen, , University Medical Center Groningen, ; Groningen, The Netherlands
                [248 ]GRID grid.33199.31, ISNI 0000 0004 0368 7223, Department of Epidemiology and Biostatistics, School of Public Health, , Tongji Medical College, Huazhong University of Science and Technology, ; Wuhan, China
                [249 ]GRID grid.4868.2, ISNI 0000 0001 2171 1133, NIHR Barts Cardiovascular Biomedical Research Centre, Barts and The London School of Medicine and Dentistry, , Queen Mary University of London, ; London, UK
                [250 ]GRID grid.24696.3f, ISNI 0000 0004 0369 153X, Department of Ophthalmology, Beijing Tongren Hospital, , Capital Medical University, ; Beijing, China
                [251 ]GRID grid.7445.2, ISNI 0000 0001 2113 8111, Department of Epidemiology and Biostatistics, MRC-PHE Centre for Environment and Health, School of Public Health, , Imperial College London, ; London, UK
                [252 ]GRID grid.22937.3d, ISNI 0000 0000 9259 8492, Department of Dermatology, , Medical University of Vienna, ; Vienna, Austria
                [253 ]GRID grid.55325.34, ISNI 0000 0004 0389 8485, Department of Research and Innovation, Division of Clinical Neuroscience, , Oslo University Hospital, ; Oslo, Norway
                [254 ]GRID grid.55325.34, ISNI 0000 0004 0389 8485, Department of Neurology, , Oslo University Hospital, ; Oslo, Norway
                [255 ]GRID grid.83440.3b, ISNI 0000000121901201, MRC Unit for Lifelong Health and Ageing at UCL, Institute of Cardiovascular Science, , University College London, ; London, UK
                [256 ]GRID grid.5963.9, Institute of Genetic Epidemiology, Faculty of Medicine and Medical Center, , University of Freiburg, ; Freiburg, Germany
                [257 ]GRID grid.5963.9, Department of Medicine IV – Nephrology and Primary Care, Faculty of Medicine and Medical Center, , University of Freiburg, ; Freiburg, Germany
                [258 ]GRID grid.267308.8, ISNI 0000 0000 9206 2401, Institute of Molecular Medicine, McGovern Medical School, , University of Texas Health Science Center at Houston, ; Houston, TX USA
                [259 ]GRID grid.410781.b, ISNI 0000 0001 0706 0776, Department of Medical Biochemistry, , Kurume University School of Medicine, ; Kurume, Japan
                [260 ]GRID grid.240684.c, ISNI 0000 0001 0705 3621, Rush Alzheimer’s Disease Center, , Rush University Medical Center, ; Chicago, IL USA
                [261 ]GRID grid.240684.c, ISNI 0000 0001 0705 3621, Department of Neurological Sciences, , Rush University Medical Center, ; Chicago, IL USA
                [262 ]GRID grid.416973.e, ISNI 0000 0004 0582 4340, Department of Genetic Medicine, , Weill Cornell Medicine-Qatar, ; Doha, Qatar
                [263 ]GRID grid.7155.6, ISNI 0000 0001 2260 6941, Department of Computer and Systems Engineering, , Alexandria University, ; Alexandria, Egypt
                [264 ]GRID grid.6936.a, ISNI 0000000123222966, Department of Cardiology, German Heart Centre Munich, , Technical University Munich, ; Munich, Germany
                [265 ]GRID grid.439803.5, Department of Cardiology, Ealing Hospital, , London North West University Healthcare NHS Trust, ; London, UK
                [266 ]GRID grid.7445.2, ISNI 0000 0001 2113 8111, Department of Epidemiology and Biostatistics, , Imperial College London, ; London, UK
                [267 ]GRID grid.5335.0, ISNI 0000000121885934, Cardiovascular Epidemiology Unit, Department of Public Health and Primary Care, , University of Cambridge, Strangeways Research Laboratory, ; Cambridge, UK
                [268 ]GRID grid.214458.e, ISNI 0000000086837370, Department of Computational Medicine and Bioinformatics, , University of Michigan, ; Ann Arbor, MI USA
                [269 ]GRID grid.32224.35, ISNI 0000 0004 0386 9924, Analytic and Translational Genetics Unit, , Massachusetts General Hospital, ; Boston, MA USA
                [270 ]GRID grid.66859.34, ISNI 0000 0004 0546 1623, Stanley Center for Psychiatric Research, , Broad Institute of MIT and Harvard, ; Cambridge, MA USA
                [271 ]GRID grid.10698.36, ISNI 0000000122483208, Department of Nutrition, Gillings School of Global Public Health, , University of North Carolina at Chapel Hill, ; Chapel Hill, NC USA
                [272 ]GRID grid.10698.36, ISNI 0000000122483208, Carolina Population Center, , University of North Carolina at Chapel Hill, ; Chapel Hill, NC USA
                [273 ]GRID grid.5645.2, ISNI 000000040459992X, Department of Clinical Genetics, , Erasmus MC, ; Rotterdam, The Netherlands
                [274 ]GRID grid.5645.2, ISNI 000000040459992X, Department of Radiology and Nuclear Medicine, , Erasmus MC, ; Rotterdam, The Netherlands
                [275 ]GRID grid.440617.0, ISNI 0000 0001 2162 5606, Latin American Brain Health (BrainLat), , Universidad Adolfo Ibáñez, ; Santiago, Chile
                [276 ]GRID grid.416850.e, ISNI 0000 0001 0698 4037, Unidad de Investigacion de Enfermedades Metabolicas and Direction of Nutrition, , Instituto Nacional de Ciencias Medicas y Nutricion, ; Mexico City, Mexico
                [277 ]GRID grid.419886.a, ISNI 0000 0001 2203 4701, Escuela de Medicina y Ciencias de la Salud, , Tecnologico de Monterrey, ; Monterrey, Mexico
                [278 ]GRID grid.266539.d, ISNI 0000 0004 1936 8438, Department of Epidemiology and Dean’s Office, College of Public Health, , University of Kentucky, ; Lexington, KY USA
                [279 ]GRID grid.83440.3b, ISNI 0000000121901201, Institute of Cardiovascular Science, Faculty of Population Health Sciences, , University College London, ; London, UK
                [280 ]GRID grid.83440.3b, ISNI 0000000121901201, Health Data Research UK and Institute of Health Informatics, , University College London, ; London, UK
                [281 ]GRID grid.5947.f, ISNI 0000 0001 1516 2393, KG Jebsen Center for Genetic Epidemiology, Department of Public Health and Nursing, Faculty of Medicine and Health Sciences, , Norwegian University of Science and Technology, NTNU, ; Trondheim, Norway
                [282 ]GRID grid.5947.f, ISNI 0000 0001 1516 2393, HUNT Research Centre, Department of Public Health and Nursing, , Norwegian University of Science and Technology, ; Levanger, Norway
                [283 ]GRID grid.52522.32, ISNI 0000 0004 0627 3560, Department of Endocrinology, Clinic of Medicine, St. Olavs Hospital, , Trondheim University Hospital, ; Trondheim, Norway
                [284 ]GRID grid.411941.8, ISNI 0000 0000 9194 7179, Department of Nephrology, , University Hospital Regensburg, ; Regensburg, Germany
                [285 ]Geriatric Unit, Azienda Toscana Centro, Florence, Italy
                [286 ]GRID grid.10706.30, ISNI 0000 0004 0498 924X, Systems Genomics Laboratory, School of Biotechnology, , Jawaharlal Nehru University (JNU), ; New Delhi, India
                [287 ]GRID grid.5326.2, ISNI 0000 0001 1940 4177, Institute of Molecular Genetics, , National Research Council of Italy, ; Pavia, Italy
                [288 ]GRID grid.267308.8, ISNI 0000 0000 9206 2401, Human Genetics Center and Department of Epidemiology, , University of Texas Health Science Center at Houston, ; Houston, TX USA
                [289 ]Department of Nephrology and Rheumatology, Kliniken Südostbayern, Regensburg, Germany
                [290 ]KfH Kidney Center Traunstein, Traunstein, Germany
                [291 ]GRID grid.7048.b, ISNI 0000 0001 1956 2722, Center for Genomics and Personalized Medicine (CGPM), , Aarhus University, ; Aarhus, Denmark
                [292 ]GRID grid.7048.b, ISNI 0000 0001 1956 2722, Bioinformatics Research Centre, , Aarhus University, ; Aarhus, Denmark
                [293 ]GRID grid.267101.3, ISNI 0000 0001 0672 9351, USC-Office of Population Studies Foundation, , University of San Carlos, ; Cebu City, Philippines
                [294 ]GRID grid.267101.3, ISNI 0000 0001 0672 9351, Department of Nutrition and Dietetics, , University of San Carlos, ; Cebu City, Philippines
                [295 ]GRID grid.250514.7, ISNI 0000 0001 2159 6024, Human Genomics Laboratory, , Pennington Biomedical Research Center, ; Baton Rouge, LA USA
                [296 ]GRID grid.241167.7, ISNI 0000 0001 2185 3318, Department of Biochemistry, , Wake Forest School of Medicine, Medical Center Boulevard, ; Winston-Salem, NC USA
                [297 ]GRID grid.459623.f, ISNI 0000 0004 0587 0347, Department of Clinical Biochemistry, , Lillebaelt Hospital, ; Vejle, Denmark
                [298 ]GRID grid.10825.3e, ISNI 0000 0001 0728 0170, Institute of Regional Health Research, , University of Southern Denmark, ; Odense, Denmark
                [299 ]GRID grid.52522.32, ISNI 0000 0004 0627 3560, Clinic of Medicine, St. Olavs Hospital, , Trondheim University Hospital, ; Trondheim, Norway
                [300 ]GRID grid.59025.3b, ISNI 0000 0001 2224 0361, Lee Kong Chian School of Medicine, , Nanyang Technological University, ; Singapore, Singapore
                [301 ]GRID grid.7445.2, ISNI 0000 0001 2113 8111, Imperial College Healthcare NHS Trust, , Imperial College London, ; London, UK
                [302 ]GRID grid.411962.9, ISNI 0000 0004 1761 157X, Adjunct Faculty, JSS University Academy of Higher Education and Research (JSSAHER), , JSS (Deemed to be) University, ; Mysuru, India
                [303 ]GRID grid.428397.3, ISNI 0000 0004 0385 0924, Ophthalmology and Visual Sciences Academic Clinical Program (Eye ACP), , Duke-NUS Medical School, ; Singapore, Singapore
                [304 ]GRID grid.55325.34, ISNI 0000 0004 0389 8485, Department of Medical Genetics, , Oslo University Hospital, ; Oslo, Norway
                [305 ]GRID grid.414168.e, ISNI 0000 0004 0627 3595, Department of Medical Research, , Bærum Hospital, Vestre Viken Hospital Trust, ; Gjettum, Norway
                [306 ]GRID grid.411024.2, ISNI 0000 0001 2175 4264, Department of Neurology, Division of Vascular Neurology, , University of Maryland School of Medicine, ; Baltimore, MD USA
                [307 ]GRID grid.280711.d, ISNI 0000 0004 0419 6661, Baltimore Veterans Affairs Medical Center, Department of Neurology, ; Baltimore, MD USA
                [308 ]GRID grid.419157.f, ISNI 0000 0001 1091 9430, Unidad de Investigación Médica en Bioquímica, Hospital de Especialidades, Centro Médico Nacional Siglo XXI, , Instituto Mexicano del Seguro Social, ; Mexico City, Mexico
                [309 ]GRID grid.11450.31, ISNI 0000 0001 2097 9138, Dipartimento di Scienze Biomediche, , Università degli Studi di Sassari, ; Sassari, Italy
                [310 ]GRID grid.414785.b, ISNI 0000 0004 0609 0182, Intermountain Heart Institute, , Intermountain Medical Center, ; Murray, UT USA
                [311 ]GRID grid.25879.31, ISNI 0000 0004 1936 8972, Department of Surgery, , University of Pennsylvania, ; Philadelphia, PA USA
                [312 ]GRID grid.410355.6, ISNI 0000 0004 0420 350X, Corporal Michael J. Crescenz VA Medical Center, ; Philadelphia, PA USA
                [313 ]GRID grid.7692.a, ISNI 0000000090126352, Department of Vascular Surgery, , University Medical Center Utrecht, University of Utrecht, ; Utrecht, The Netherlands
                [314 ]GRID grid.4818.5, ISNI 0000 0001 0791 5666, Department of Human Nutrition, , Wageningen University, ; Wageningen, The Netherlands
                [315 ]GRID grid.239585.0, ISNI 0000 0001 2285 2675, Center for Translational and Computational Neuroimmunology, Department of Neurology, , Columbia University Medical Center, ; New York, NY USA
                [316 ]GRID grid.5335.0, ISNI 0000000121885934, Department of Oncology, , University of Cambridge, ; Cambridge, UK
                [317 ]GRID grid.16872.3a, ISNI 0000 0004 0435 165X, Department of General Practice, Amsterdam Public Health Institute, , Amsterdam UMC, location VUmc, ; Amsterdam, The Netherlands
                [318 ]GRID grid.38142.3c, ISNI 000000041936754X, Department of Nutrition, , Harvard T.H. Chan School of Public Health, ; Boston, MA USA
                [319 ]GRID grid.32224.35, ISNI 0000 0004 0386 9924, Cardiac Arrhythmia Service, , Massachusetts General Hospital, ; Boston, MA USA
                [320 ]GRID grid.32224.35, ISNI 0000 0004 0386 9924, Cardiovascular Research Center, , Massachusetts General Hospital, ; Boston, MA USA
                [321 ]GRID grid.4514.4, ISNI 0000 0001 0930 2361, Department of Clinical Sciences in Malmö, Division of Geriatric Medicine, , Lund University, ; Malmö, Sweden
                [322 ]GRID grid.1057.3, ISNI 0000 0000 9472 3971, Molecular Cardiology Division, , Victor Chang Cardiac Research Institute, ; Darlinghurst, New South Wales Australia
                [323 ]GRID grid.437825.f, ISNI 0000 0000 9119 2677, Cardiology Department, , St Vincent’s Hospital, ; Darlinghurst, New South Wales Australia
                [324 ]GRID grid.1005.4, ISNI 0000 0004 4902 0432, Faculty of Medicine, , UNSW Sydney, ; Kensington, New South Wales Australia
                [325 ]GRID grid.94365.3d, ISNI 0000 0001 2297 5165, Translational Gerontology Branch, National Institute on Aging, , National Institutes of Health, ; Baltimore, MD USA
                [326 ]GRID grid.8756.c, ISNI 0000 0001 2193 314X, Robertson Center for Biostatistics, , University of Glasgow, ; Glasgow, UK
                [327 ]GRID grid.267308.8, ISNI 0000 0000 9206 2401, Human Genetics Center, School of Public Health, , University of Texas Health Science Center at Houston, ; Houston, TX USA
                [328 ]GRID grid.12650.30, ISNI 0000 0001 1034 3451, Department of Public Health and Clinical Medicine, , Umeå University, ; Umeå, Sweden
                [329 ]GRID grid.241167.7, ISNI 0000 0001 2185 3318, Department of Internal Medicine, , Wake Forest School of Medicine, Medical Center Boulevard, ; Winston-Salem, NC USA
                [330 ]GRID grid.1008.9, ISNI 0000 0001 2179 088X, Faculty of Veterinary and Agricultural Science, , University of Melbourne, ; Parkville, Victoria Australia
                [331 ]GRID grid.511012.6, ISNI 0000 0001 0744 2459, Agriculture Victoria Research, , Department of Jobs, Precincts and Regions, ; Bundoora, Victoria Australia
                [332 ]GRID grid.10698.36, ISNI 0000000122483208, Injury Prevention Research Center, , University of North Carolina at Chapel Hill, ; Chapel Hill, NC USA
                [333 ]GRID grid.10698.36, ISNI 0000000122483208, Division of Physical Therapy, , University of North Carolina at Chapel Hill, ; Chapel Hill, NC USA
                [334 ]GRID grid.415771.1, ISNI 0000 0004 1773 4764, Centro de Investigacion en Salud Poblacional Instituto Nacional de Salud Publica and Centro de Estudios en Diabetes, ; Cuernavaca, Mexico
                [335 ]GRID grid.239552.a, ISNI 0000 0001 0680 8770, Division of Human Genetics, , Children’s Hospital of Philadelphia, ; Philadelphia, PA USA
                [336 ]GRID grid.25879.31, ISNI 0000 0004 1936 8972, Departments of Pediatrics and Genetics, Perelman School of Medicine, , University of Pennsylvania, ; Philadelphia, PA USA
                [337 ]GRID grid.239552.a, ISNI 0000 0001 0680 8770, Division of Endocrinology and Diabetes, , Children’s Hospital of Philadelphia, ; Philadelphia, PA USA
                [338 ]GRID grid.14013.37, ISNI 0000 0004 0640 0021, Faculty of Medicine, , University of Iceland, ; Reykjavik, Iceland
                [339 ]GRID grid.42505.36, ISNI 0000 0001 2156 6853, Department of Preventive Medicine, , Keck School of Medicine of USC, ; Los Angeles, CA USA
                [340 ]GRID grid.25879.31, ISNI 0000 0004 1936 8972, Department of Pediatrics, Perelman School of Medicine, , University of Pennsylvania, ; Philadelphia, PA USA
                [341 ]GRID grid.239552.a, ISNI 0000 0001 0680 8770, Division of Pulmonary Medicine, , Children’s Hospital of Philadelphia, ; Philadelphia, PA USA
                [342 ]GRID grid.8391.3, ISNI 0000 0004 1936 8024, Institute of Biomedical and Clinical Science, , University of Exeter Medical School, ; Exeter, UK
                [343 ]GRID grid.34477.33, ISNI 0000000122986657, Cardiovascular Health Research Unit, Department of Epidemiology, , University of Washington, ; Seattle, WA USA
                [344 ]GRID grid.4280.e, ISNI 0000 0001 2180 6431, Department of Paediatrics, Yong Loo Lin School of Medicine, , National University of Singapore, ; Singapore, Singapore
                [345 ]GRID grid.410759.e, ISNI 0000 0004 0451 6143, Khoo Teck Puat - National University Children’s Medical Institute, , National University Health System, ; Singapore, Singapore
                [346 ]GRID grid.22937.3d, ISNI 0000 0000 9259 8492, Department of Internal Medicine II, Division of Cardiology, , Medical University of Vienna, ; Vienna, Austria
                [347 ]GRID grid.1009.8, ISNI 0000 0004 1936 826X, Menzies Research Institute Tasmania, , University of Tasmania, ; Hobart, Tasmania Australia
                [348 ]GRID grid.1008.9, ISNI 0000 0001 2179 088X, Centre for Eye Research Australia, Royal Victorian Eye and Ear Hospital, , University of Melbourne, ; Melbourne, Victoria Australia
                [349 ]GRID grid.1012.2, ISNI 0000 0004 1936 7910, Lions Eye Institute, Centre for Ophthalmology and Vision Science, , University of Western Australia, ; Perth, Western Australia Australia
                [350 ]GRID grid.32224.35, ISNI 0000 0004 0386 9924, Cardiology Division, , Massachusetts General Hospital, ; Boston, MA USA
                [351 ]GRID grid.464306.3, ISNI 0000 0004 0410 5707, Department of Genetics, Shanghai-MOST Key Laboratory of Heath and Disease Genomics, , Chinese National Human Genome Center and Shanghai Industrial Technology Institute, ; Shanghai, China
                [352 ]GRID grid.223827.e, ISNI 0000 0001 2193 0096, Department of Internal Medicine, , University of Utah, ; Salt Lake City, UT USA
                [353 ]GRID grid.1026.5, ISNI 0000 0000 8994 5086, Australian Centre for Precision Health, Clinical and Health Sciences, , University of South Australia, ; Adelaide, South Australia Australia
                [354 ]GRID grid.430453.5, ISNI 0000 0004 0565 2606, South Australian Health and Medical Research Institute, ; Adelaide, South Australia Australia
                [355 ]GRID grid.410804.9, ISNI 0000000123090000, Department of Environmental and Preventive Medicine, , Jichi Medical University School of Medicine, ; Shimotsuke, Japan
                [356 ]GRID grid.251993.5, ISNI 0000000121791997, Department of Epidemiology and Population Health, , Albert Einstein College of Medicine, ; Bronx, NY USA
                [357 ]GRID grid.261331.4, ISNI 0000 0001 2285 7943, Division of Endocrinology, Diabetes and Metabolism, School of Medicine, , Ohio State University, ; Columbus, OH USA
                [358 ]GRID grid.10858.34, ISNI 0000 0001 0941 4873, Center for Life Course Health Research, Faculty of Medicine, , University of Oulu, ; Oulu, Finland
                [359 ]GRID grid.412326.0, ISNI 0000 0004 4685 4917, Unit of Primary Health Care, , Oulu University Hospital, OYS, ; Oulu, Finland
                [360 ]GRID grid.7728.a, ISNI 0000 0001 0724 6933, Department of Life Sciences, College of Health and Life Sciences, , Brunel University London, ; Uxbridge, UK
                [361 ]GRID grid.268099.c, ISNI 0000 0001 0348 3990, The Eye Hospital, School of Ophthalmology and Optometry, , Wenzhou Medical University, ; Wenzhou, China
                [362 ]GRID grid.10419.3d, ISNI 0000000089452978, Einthoven Laboratory for Experimental Vascular Medicine, LUMC, ; Leiden, The Netherlands
                [363 ]GRID grid.411737.7, Netherlands Heart Institute, ; Utrecht, The Netherlands
                [364 ]GRID grid.412330.7, ISNI 0000 0004 0628 2985, Department of Clinical Physiology, , Tampere University Hospital, ; Tampere, Finland
                [365 ]GRID grid.502801.e, ISNI 0000 0001 2314 6254, Department of Clinical Physiology, Finnish Cardiovascular Research Center - Tampere, Faculty of Medicine and Health Technology, , Tampere University, ; Tampere, Finland
                [366 ]GRID grid.26999.3d, ISNI 0000 0001 2151 536X, Laboratory of Complex Trait Genomics, Department of Computational Biology and Medical Sciences, Graduate School of Frontier Sciences, , The University of Tokyo, ; Tokyo, Japan
                [367 ]GRID grid.411947.e, ISNI 0000 0004 0470 4224, Department of Ophthalmology, , The Catholic University of Korea Incheon St. Mary’s Hospital, ; Incheon, Republic of Korea
                [368 ]GRID grid.415719.f, ISNI 0000 0004 0488 9484, NIHR Oxford Biomedical Research Centre, Churchill Hospital, ; Oxford, UK
                [369 ]GRID grid.452396.f, ISNI 0000 0004 5937 5237, German Centre for Cardiovascular Research (DZHK), partner site Munich Heart Alliance, ; Munich, Germany
                [370 ]GRID grid.10417.33, ISNI 0000 0004 0444 9382, Radboud University Medical Center, Radboud Institute for Health Sciences, Department of Urology, ; Nijmegen, The Netherlands
                [371 ]GRID grid.15444.30, ISNI 0000 0004 0470 5454, Corneal Dystrophy Research Institute, , Yonsei University College of Medicine, ; Seoul, Republic of Korea
                [372 ]Saevit Eye Hospital, Goyang, Republic of Korea
                [373 ]GRID grid.255649.9, ISNI 0000 0001 2171 7754, Department of Biochemistry, College of Medicine, , Ewha Womans University, ; Seoul, Republic of Korea
                [374 ]GRID grid.13648.38, ISNI 0000 0001 2180 3484, Department of Cardiology, , University Heart and Vascular Center UKE Hamburg, ; Hamburg, Germany
                [375 ]GRID grid.6572.6, ISNI 0000 0004 1936 7486, Institute of Cardiovascular Sciences, College of Medical and Dental Sciences, , University of Birmingham, ; Birmingham, UK
                [376 ]GRID grid.452396.f, ISNI 0000 0004 5937 5237, German Center for Cardiovascular Research, partner site Hamburg/Kiel/Lübeck, ; Hamburg, Germany
                [377 ]GRID grid.476464.3, ISNI 0000 0004 0431 535X, Atrial Fibrillation NETwork, ; Münster, Germany
                [378 ]GRID grid.83440.3b, ISNI 0000000121901201, Department of Epidemiology and Public Health, UCL Institute of Epidemiology and Health Care, , University College London, ; London, UK
                [379 ]GRID grid.4280.e, ISNI 0000 0001 2180 6431, Healthy Longevity Translational Research Programme, Yong Loo Lin School of Medicine, , National University of Singapore, ; Singapore, Singapore
                [380 ]GRID grid.15485.3d, ISNI 0000 0000 9950 5666, University of Helsinki and Department of Medicine, Helsinki University Hospital, ; Helsinki, Finland
                [381 ]GRID grid.452540.2, Minerva Foundation Institute for Medical Research, ; Helsinki, Finland
                [382 ]GRID grid.415451.0, ISNI 0000 0004 0622 6078, Department of Preventive Cardiology, Lipoprotein Apheresis Unit and Lipid Disorders Clinic, , Metropolitan Hospital, ; Athens, Greece
                [383 ]GRID grid.7445.2, ISNI 0000 0001 2113 8111, MRC-PHE Centre for Environment and Health, , Imperial College London, ; London, UK
                [384 ]GRID grid.7445.2, ISNI 0000 0001 2113 8111, National Heart and Lung Institute, , Imperial College London, ; London, UK
                [385 ]GRID grid.9647.c, ISNI 0000 0004 7669 9786, Medical Department III – Endocrinology, Nephrology, Rheumatology, , University of Leipzig Medical Center, ; Leipzig, Germany
                [386 ]GRID grid.8356.8, ISNI 0000 0001 0942 6946, Institute for Social and Economic Research, , University of Essex, ; Colchester, UK
                [387 ]GRID grid.9668.1, ISNI 0000 0001 0726 2490, Institute of Clinical Medicine, Internal Medicine, , University of Eastern Finland and Kuopio University Hospital, ; Kuopio, Finland
                [388 ]GRID grid.430503.1, ISNI 0000 0001 0703 675X, Department of Medicine, , University of Colorado at Denver, ; Aurora, CO USA
                [389 ]GRID grid.484013.a, ISNI 0000 0004 6879 971X, Berlin Institute of Health at Charité – Universitätsmedizin Berlin, ; Berlin, Germany
                [390 ]GRID grid.410445.0, ISNI 0000 0001 2188 0957, Epidemiology Program, University of Hawaii Cancer Center, ; Honolulu, HI USA
                [391 ]GRID grid.255649.9, ISNI 0000 0001 2171 7754, Department of Internal Medicine, , Ewha Womans University School of Medicine, ; Seoul, Republic of Korea
                [392 ]GRID grid.11135.37, ISNI 0000 0001 2256 9319, Department of Epidemiology and Biostatistics, , Peking University Health Science Center, ; Beijing, China
                [393 ]GRID grid.11135.37, ISNI 0000 0001 2256 9319, Peking University Center for Public Health and Epidemic Preparedness and Response, ; Beijing, China
                [394 ]GRID grid.9764.c, ISNI 0000 0001 2153 9986, Institute of Epidemiology and Biobank Popgen, , Kiel University, ; Kiel, Germany
                [395 ]GRID grid.410726.6, ISNI 0000 0004 1797 8419, Key Laboratory of Systems Health Science of Zhejiang Province, Hangzhou Institute for Advanced Study, , University of Chinese Academy of Sciences, Chinese Academy of Sciences, ; Hangzhou, China
                [396 ]GRID grid.5254.6, ISNI 0000 0001 0674 042X, Department of Clinical Medicine, Faculty of Health and Medical Sciences, , University of Copenhagen, ; Copenhagen, Denmark
                [397 ]GRID grid.412584.e, ISNI 0000 0004 0434 9816, Division of Cardiovascular Medicine and Abboud Cardiovascular Research Center, , University of Iowa Hospitals and Clinics, ; Iowa City, IA USA
                [398 ]GRID grid.250674.2, ISNI 0000 0004 0626 6184, Alliance for Human Development, , Lunenfeld-Tanenbaum Research Institute, Sinai Health System, ; Toronto, Ontario Canada
                [399 ]GRID grid.4714.6, ISNI 0000 0004 1937 0626, Department of Medical Epidemiology and Biostatistics, , Karolinska Institutet, ; Stockholm, Sweden
                [400 ]GRID grid.266102.1, ISNI 0000 0001 2297 6811, Division of Cardiology, , University of California, San Francisco, ; San Francisco, CA USA
                [401 ]GRID grid.8515.9, ISNI 0000 0001 0423 4662, Department of Medicine, , Internal Medicine, Lausanne University Hospital, ; Lausanne, Switzerland
                [402 ]GRID grid.9851.5, ISNI 0000 0001 2165 4204, University of Lausanne, ; Lausanne, Switzerland
                [403 ]GRID grid.461810.a, ISNI 0000 0004 0572 0285, SYNLAB Academy, SYNLAB Holding Deutschland, ; Mannheim, Germany
                [404 ]GRID grid.11598.34, ISNI 0000 0000 8988 2476, Clinical Institute of Medical and Chemical Laboratory Diagnostics, , Medical University of Graz, ; Graz, Austria
                [405 ]GRID grid.26009.3d, ISNI 0000 0004 1936 7961, Department of Medicine, Division of Cardiology, , Duke University School of Medicine, ; Durham, NC USA
                [406 ]GRID grid.26009.3d, ISNI 0000 0004 1936 7961, Duke Molecular Physiology Institute, , Duke University School of Medicine, ; Durham, NC USA
                [407 ]GRID grid.1049.c, ISNI 0000 0001 2294 1395, Psychiatric Genetics, QIMR Berghofer Medical Research Institute, ; Brisbane, Queensland Australia
                [408 ]GRID grid.8761.8, ISNI 0000 0000 9919 9582, Geriatric Medicine, Institute of Medicine, , Sahlgrenska Academy, University of Gothenburg, ; Gothenburg, Sweden
                [409 ]GRID grid.280711.d, ISNI 0000 0004 0419 6661, Geriatrics Research and Education Clinical Center, Baltimore Veterans Administration Medical Center, ; Baltimore, MD USA
                [410 ]GRID grid.1013.3, ISNI 0000 0004 1936 834X, Centre for Vision Research and Department of Ophthalmology, Westmead Millennium Institute of Medical Research, , University of Sydney, ; Sydney, New South Wales Australia
                [411 ]GRID grid.10419.3d, ISNI 0000000089452978, Department of Public Health and Primary Care, , Leiden University Medical Center, ; Leiden, The Netherlands
                [412 ]GRID grid.4305.2, ISNI 0000 0004 1936 7988, Usher Institute of Population Health Sciences and Informatics, University of Edinburgh, ; Edinburgh, UK
                [413 ]GRID grid.25879.31, ISNI 0000 0004 1936 8972, Electrophysiology Section, Division of Cardiovascular Medicine, Perelman School of Medicine, University of Pennsylvania, ; Philadelphia, PA USA
                [414 ]GRID grid.10698.36, ISNI 0000000122483208, Department of Medicine, , University of North Carolina at Chapel Hill, ; Chapel Hill, NC USA
                [415 ]GRID grid.418193.6, ISNI 0000 0001 1541 4204, Department of Chronic Diseases, , Norwegian Institute of Public Health, ; Oslo, Norway
                [416 ]GRID grid.55325.34, ISNI 0000 0004 0389 8485, Department of Pain Management and Research, , Oslo University Hospital, ; Oslo, Norway
                [417 ]GRID grid.10388.32, ISNI 0000 0001 2240 3300, Institute of Human Genetics, School of Medicine and University Hospital Bonn, ; Bonn, Germany
                [418 ]GRID grid.1649.a, ISNI 000000009445082X, Sahlgrenska University Hospital, Department of Drug Treatment, ; Gothenburg, Sweden
                [419 ]GRID grid.452651.1, ISNI 0000 0004 0627 7633, Laboratorio de Inmunogenómica y Enfermedades Metabólicas, , Instituto Nacional de Medicina Genómica, CDMX, ; Mexico City, Mexico
                [420 ]GRID grid.1374.1, ISNI 0000 0001 2097 1371, Paavo Nurmi Centre, Sports and Exercise Medicine Unit, Department of Physical Activity and Health, , University of Turku, ; Turku, Finland
                [421 ]GRID grid.19006.3e, ISNI 0000 0000 9632 6718, Institute for Precision Health, , David Geffen School of Medicine at UCLA, ; Los Angeles, CA USA
                [422 ]GRID grid.8241.f, ISNI 0000 0004 0397 2876, Pat MacPherson Centre for Pharmacogenetics and Pharmacogenomics, Division of Population Health and Genomics, School of Medicine, University of Dundee, Ninewells Hospital and Medical School, ; Dundee, UK
                [423 ]GRID grid.23856.3a, ISNI 0000 0004 1936 8390, Centre Nutrition, Santé et Société (NUTRISS), Institute of Nutrition and Functional Foods, , Université Laval, ; Québec City, Quebec Canada
                [424 ]GRID grid.5252.0, ISNI 0000 0004 1936 973X, IBE-Chair of Epidemiology, LMU Munich, ; Neuherberg, Germany
                [425 ]GRID grid.420468.c, Population, Policy and Practice, UCL Great Ormond Street Hospital Institute of Child Health, ; London, UK
                [426 ]GRID grid.25879.31, ISNI 0000 0004 1936 8972, Department of Medicine, , University of Pennsylvania, ; Philadelphia, PA USA
                [427 ]GRID grid.410552.7, ISNI 0000 0004 0628 215X, Department of Clinical Physiology and Nuclear Medicine, , Turku University Hospital, ; Turku, Finland
                [428 ]Hero DMC Heart Institute, Dyanand Medical College, Ludhiana, India
                [429 ]GRID grid.5216.0, ISNI 0000 0001 2155 0800, Second Department of Cardiology, Medical School, , National and Kapodistrian University of Athens, University General Hospital Attikon, ; Athens, Greece
                [430 ]GRID grid.4367.6, ISNI 0000 0001 2355 7002, Division of Biostatistics, , Washington University School of Medicine, ; St Louis, MO USA
                [431 ]GRID grid.417993.1, ISNI 0000 0001 2260 0793, Genetics, Merck Sharp & Dohme, ; Kenilworth, NJ USA
                [432 ]GRID grid.34477.33, ISNI 0000000122986657, Department of Epidemiology, , University of Washington, ; Seattle, WA USA
                [433 ]GRID grid.289247.2, ISNI 0000 0001 2171 7818, Department of Endocrinology and Metabolism, , Kyung Hee University School of Medicine, ; Seoul, Korea
                [434 ]GRID grid.7737.4, ISNI 0000 0004 0410 2071, Department of Public Health, Clinicum, , Faculty of Medicine, University of Helsinki, ; Helsinki, Finland
                [435 ]GRID grid.412807.8, ISNI 0000 0004 1936 9916, Departments of Medicine, Pharmacology, and Biomedical Informatics, , Vanderbilt University Medical Center, ; Nashville, TN USA
                [436 ]GRID grid.16872.3a, ISNI 0000 0004 0435 165X, Department of Epidemiology and Data Science, Amsterdam Public Health Institute, Amsterdam Cardiovascular Sciences Institute, , Amsterdam UMC, location VUmc, ; Amsterdam, The Netherlands
                [437 ]GRID grid.21729.3f, ISNI 0000000419368729, Department of Cardiology and Department of Medicine, , Columbia University, ; New York, NY USA
                [438 ]GRID grid.266902.9, ISNI 0000 0001 2179 3618, Department of Pediatrics, Section of Genetics, College of Medicine, , University of Oklahoma Health Sciences Center, ; Oklahoma City, OK USA
                [439 ]GRID grid.266902.9, ISNI 0000 0001 2179 3618, Department of Pharmaceutical Sciences, , University of Oklahoma Health Sciences Center, ; Oklahoma City, OK USA
                [440 ]GRID grid.266902.9, ISNI 0000 0001 2179 3618, Department of Physiology, , University of Oklahoma Health Sciences Center, ; Oklahoma City, OK USA
                [441 ]GRID grid.266902.9, ISNI 0000 0001 2179 3618, Oklahoma Center for Neuroscience, , University of Oklahoma Health Sciences Center, ; Oklahoma City, OK USA
                [442 ]GRID grid.8756.c, ISNI 0000 0001 2193 314X, Institute of Cardiovascular and Medical Sciences, , University of Glasgow, ; Glasgow, UK
                [443 ]GRID grid.11598.34, ISNI 0000 0000 8988 2476, Gottfried Schatz Research Center (for Cell Signaling, Metabolism and Aging), , Medical University of Graz, ; Graz, Austria
                [444 ]GRID grid.11348.3f, ISNI 0000 0001 0942 1117, Institute of Nutritional Science, , University of Potsdam, ; Nuthetal, Germany
                [445 ]GRID grid.6936.a, ISNI 0000000123222966, Deutsches Herzzentrum München, Cardiology, Deutsches Zentrum für Herz- und Kreislaufforschung (DZHK) – Munich Heart Alliance, and Technische Universität München, ; München, Germany
                [446 ]GRID grid.266842.c, ISNI 0000 0000 8831 109X, School of Biomedical Science and Pharmacy, , University of Newcastle, ; New Lambton Heights, New South Wales Australia
                [447 ]GRID grid.412807.8, ISNI 0000 0004 1936 9916, Department of Medicine, , Vanderbilt University Medical Center, ; Nashville, TN USA
                [448 ]GRID grid.428366.d, ISNI 0000 0004 1773 9952, Central University of Punjab, ; Bathinda, India
                [449 ]GRID grid.5252.0, ISNI 0000 0004 1936 973X, Department of Medicine I, , University Hospital, LMU Munich, ; Munich, Germany
                [450 ]GRID grid.411843.b, ISNI 0000 0004 0623 9987, Department of Cardiology, Clinical Sciences, , Lund University and Skåne University Hospital, ; Lund, Sweden
                [451 ]GRID grid.1649.a, ISNI 000000009445082X, The Wallenberg Laboratory, Department of Molecular and Clinical Medicine, Institute of Medicine, Gothenburg University and the Department of Cardiology, Sahlgrenska University Hospital, ; Gothenburg, Sweden
                [452 ]GRID grid.4514.4, ISNI 0000 0001 0930 2361, Wallenberg Center for Molecular Medicine and Lund University Diabetes Center, , Lund University, ; Lund, Sweden
                [453 ]GRID grid.264200.2, ISNI 0000 0000 8546 682X, Population Health Research Institute, , St George’s, University of London, ; London, UK
                [454 ]GRID grid.10419.3d, ISNI 0000000089452978, Molecular Epidemiology Section, Department of Biomedical Data Sciences, , Leiden University Medical Center, ; Leiden, The Netherlands
                [455 ]GRID grid.168010.e, ISNI 0000000419368956, Department of Genetics, , Stanford University School of Medicine, ; Stanford, CA USA
                [456 ]GRID grid.14848.31, ISNI 0000 0001 2292 3357, Department of Medicine, Faculty of Medicine, , Université de Montréal, ; Montreal, Quebec Canada
                [457 ]GRID grid.7737.4, ISNI 0000 0004 0410 2071, Helsinki University Central Hospital, Research Program for Clinical and Molecular Metabolism, , University of Helsinki, ; Helsinki, Finland
                [458 ]GRID grid.428673.c, ISNI 0000 0004 0409 6302, Folkhälsan Research Center, ; Helsinki, Finland
                [459 ]GRID grid.7737.4, ISNI 0000 0004 0410 2071, Department of Public Health, , University of Helsinki, ; Helsinki, Finland
                [460 ]GRID grid.412125.1, ISNI 0000 0001 0619 1117, Diabetes Research Group, , King Abdulaziz University, ; Jeddah, Saudi Arabia
                [461 ]GRID grid.9486.3, ISNI 0000 0001 2159 0001, Unidad de Biología Molecular y Medicina Genómica, Instituto de Investigaciones Biomédicas, , UNAM, ; Mexico City, Mexico
                [462 ]GRID grid.416850.e, ISNI 0000 0001 0698 4037, Instituto Nacional de Ciencias Médicas y Nutrición Salvador Zubirán, ; Mexico City, Mexico
                [463 ]GRID grid.4280.e, ISNI 0000 0001 2180 6431, Yong Loo Lin School of Medicine, , National University of Singapore and National University Health System, ; Singapore, Singapore
                [464 ]GRID grid.253615.6, ISNI 0000 0004 1936 9510, Milken Institute School of Public Health, , The George Washington University, ; Washington, DC USA
                [465 ]GRID grid.7177.6, ISNI 0000000084992262, Department Geriatric Medicine, Amsterdam Public Health, , Amsterdam UMC location University of Amsterdam, ; Amsterdam, The Netherlands
                [466 ]GRID grid.509540.d, ISNI 0000 0004 6880 3010, Department of Epidemiology and Data Science, , Amsterdam UMC, ; Amsterdam, The Netherlands
                [467 ]GRID grid.5603.0, Interfaculty Institute for Genetics and Functional Genomics, , University Medicine Greifswald, ; Greifswald, Germany
                [468 ]GRID grid.419157.f, ISNI 0000 0001 1091 9430, Unidad de Investigación Médica en Epidemiología Clínica, Hospital de Especialidades, Centro Médico Nacional Siglo XXI, , Instituto Mexicano del Seguro Social, ; Mexico City, Mexico
                [469 ]GRID grid.1006.7, ISNI 0000 0001 0462 7212, Institute of Cellular Medicine, , Newcastle University, ; Newcastle upon Tyne, UK
                [470 ]GRID grid.42505.36, ISNI 0000 0001 2156 6853, Department of Population and Public Health Sciences, , Keck School of Medicine of USC, ; Los Angeles, CA USA
                [471 ]GRID grid.42505.36, ISNI 0000 0001 2156 6853, Department of Physiology and Neuroscience, , Keck School of Medicine of USC, ; Los Angeles, CA USA
                [472 ]GRID grid.42505.36, ISNI 0000 0001 2156 6853, USC Diabetes and Obesity Research Institute, , Keck School of Medicine of USC, ; Los Angeles, CA USA
                [473 ]GRID grid.5254.6, ISNI 0000 0001 0674 042X, Lundbeck Foundation Center for GeoGenetics, GLOBE Institute, , University of Copenhagen, ; Copenhagen, Denmark
                [474 ]GRID grid.254145.3, ISNI 0000 0001 0083 6092, School of Chinese Medicine, China Medical University, ; Taichung, Taiwan
                [475 ]GRID grid.46534.30, ISNI 0000 0004 1793 8046, Diabetes Unit, KEM Hospital and Research Centre, ; Pune, India
                [476 ]GRID grid.410781.b, ISNI 0000 0001 0706 0776, Kurume University School of Medicine, ; Kurume, Japan
                [477 ]GRID grid.21925.3d, ISNI 0000 0004 1936 9000, Division of Cancer Control and Population Sciences, UPMC Hillman Cancer Center, , University of Pittsburgh, ; Pittsburgh, PA USA
                [478 ]GRID grid.21925.3d, ISNI 0000 0004 1936 9000, Department of Epidemiology, Graduate School of Public Health, , University of Pittsburgh, ; Pittsburgh, PA USA
                [479 ]GRID grid.6936.a, ISNI 0000000123222966, TUM School of Medicine, , Technical University of Munich and Klinikum Rechts der Isar, ; Munich, Germany
                [480 ]GRID grid.5510.1, ISNI 0000 0004 1936 8921, Institute of Clinical Medicine, Faculty of Medicine, , University of Oslo, ; Oslo, Norway
                [481 ]GRID grid.280776.c, ISNI 0000 0004 0394 1447, Department of Population Health Sciences, , Geisinger, ; Danville, PA USA
                [482 ]GRID grid.412807.8, ISNI 0000 0004 1936 9916, Vanderbilt Genetics Institute, Division of Genetic Medicine, , Vanderbilt University Medical Center, ; Nashville, TN USA
                [483 ]GRID grid.410370.1, ISNI 0000 0004 4657 1992, Department of Medicine, , Veterans Affairs Boston Healthcare System, ; Boston, MA USA
                [484 ]GRID grid.189967.8, ISNI 0000 0001 0941 6502, Department of Epidemiology, , Emory University Rollins School of Public Health, ; Atlanta, GA USA
                [485 ]GRID grid.484294.7, Atlanta VA Health Care System, ; Decatur, GA USA
                [486 ]GRID grid.412125.1, ISNI 0000 0001 0619 1117, Princess Al-Jawhara Al-Brahim Centre of Excellence in Research of Hereditary Disorders (PACER-HD), , King Abdulaziz University, ; Jeddah, Saudi Arabia
                [487 ]GRID grid.59734.3c, ISNI 0000 0001 0670 2351, The Mindich Child Health and Development Institute, , Icahn School of Medicine at Mount Sinai, ; New York, NY USA
                [488 ]GRID grid.494629.4, ISNI 0000 0004 8008 9315, School of Life Sciences, Westlake University, ; Hangzhou, China
                [489 ]GRID grid.494629.4, ISNI 0000 0004 8008 9315, Westlake Laboratory of Life Sciences and Biomedicine, ; Hangzhou, China
                [490 ]GRID grid.214458.e, ISNI 0000000086837370, Department of Human Genetics, , University of Michigan, ; Ann Arbor, MI USA
                [491 ]GRID grid.4367.6, ISNI 0000 0001 2355 7002, McDonnell Genome Institute and Department of Medicine, , Washington University School of Medicine, ; St Louis, MO USA
                [492 ]GRID grid.136593.b, ISNI 0000 0004 0373 3971, Laboratory of Statistical Immunology, Immunology Frontier Research Center (WPI-IFReC), ; Osaka, Japan
                [493 ]GRID grid.136593.b, ISNI 0000 0004 0373 3971, Integrated Frontier Research for Medical Science Division, Institute for Open and Transdisciplinary Research Initiatives, , Osaka University, ; Osaka, Japan
                [494 ]GRID grid.66859.34, ISNI 0000 0004 0546 1623, Programs in Metabolism and Medical and Population Genetics, , Broad Institute of MIT and Harvard, ; Cambridge, MA USA
                [495 ]GRID grid.38142.3c, ISNI 000000041936754X, Departments of Pediatrics and Genetics, , Harvard Medical School, ; Boston, MA USA
                [496 ]GRID grid.264047.3, ISNI 0000 0001 0738 3196, Present Address: Department of Mathematics and Statistics, , St Cloud State University, ; St Cloud, MN USA
                [497 ]GRID grid.418158.1, ISNI 0000 0004 0534 4718, Genentech, ; South San Francisco, CA USA
                [498 ]GRID grid.509459.4, ISNI 0000 0004 0472 0267, Present Address: Laboratory for Systems Genetics, , RIKEN Center for Integrative Medical Sciences, ; Kanagawa, Japan
                [499 ]GRID grid.26999.3d, ISNI 0000 0001 2151 536X, Department of Genome Informatics, Graduate School of Medicine, , The University of Tokyo, ; Tokyo, Japan
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                Article
                5275
                10.1038/s41586-022-05275-y
                9605867
                36224396
                8a5f20da-3d5e-42be-a9b4-9b94b25aba94
                © The Author(s) 2022

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                : 19 December 2021
                : 24 August 2022
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                © The Author(s), under exclusive licence to Springer Nature Limited 2022

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                genome-wide association studies,quantitative trait,genetic markers
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                genome-wide association studies, quantitative trait, genetic markers

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