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      Hundreds of variants clustered in genomic loci and biological pathways affect human height

      research-article
      Author(s): 1 , 2 , 3 , 4 , 5 , 6 , 7 , 1 , 2 , 3 , 4 , 8 , 8 , 9 , 10 , 11 , 12 , 13 , 1 , 8 , 11 , 14 , 15 , 10 , 16 , 17 , 17 , 18 , 7 , 19 , 20 , 21 ,   13 , 22 , 23 , 24 , 25 , 13 , 26 , 27 , 28 , 29 , 30 , 31 , 32 , 20 , 30 , 33 , 34 , 5 , 9 , 10 , 22 , 2 , 4 , 35 , 36 , 3 , 37 , 38 , 39 , 40 , 41 , 42 , 43 , 44 , 37 , 45 , 46 , 18 , 13 , 47 , 48 , 31 , 49 , 50 , 51 , 52 , 53 , 54 , 41 , 42 , 55 , 56 , 57 , 58 , 13 , 47 , 13 , 47 , 20 , 59 , 11 , 14 , 15 , 60 , 61 , 32 , 62 , 63 , 64 , 63 , 64 , 1 , 63 , 64 , 38 , 9 , 10 , 2 , 24 , 25 , 65 , 5 , 66 , 67 , 68 ,   69 , 70 , 71 , 72 , 17 , 26 , 27 , 73 , 74 , 16 , 67 , 75 , 63 , 64 , 58 , 26 , 27 , 70 , 76 , 77 , 78 , 13 , 31 , 79 , 80 , 81 , 14 , 82 , 83 , 84 , 9 , 10 , 85 , 86 , 87 , 7 , 69 , 88 , 89 , 9 , 10 , 77 , 26 , 27 , 51 , 81 , 90 , 91 , 92 , 93 , 94 , 95 , 96 , 97 , 69 , 98 , 99 , 100 , 101 , 102 , 103 , 104 , 52 , 105 , 72 , 106 , 107 , 95 , 70 , 108 , 109 , 35 , 31 , 110 , 100 , 32 , 111 , 56 , 107 , 112 , 113 , 114 , 115 , 116 , 87 , 117 , 118 , 87 , 119 , 120 , 13 , 121 , 122 , 95 , 123 , 124 , 106 , 125 , 126 , 127 , 128 , 129 , 97 , 130 , 131 , 132 , 8 , 20 , 63 , 93 , 52 , 94 , 56 , 133 , 38 , 13 , 121 , 134 , 135 , 38 , 39 , 40 , 3 , 4 , 63 , 64 , 2 , 3 , 4 , 136 , 11 , 41 , 137 , 108 , 109 , 138 , 139 , 41 , 42 , 140 , 85 , 141 , 24 , 97 , 92 , 142 , 74 , 74 , 143 , 144 , 145 , 63 , 123 , 146 , 11 , 14 , 147 , 148 , 149 , 136 , 150 , 151 , 152 , 153 , 154 , 126 , 155 , 109 , 156 , 131 , 131 , 148 , 157 , 77 , 158 , 70 , 109 , 159 , 66 , 67 , 5 , 6 , 124 , 41 , 160 , 72 , 161 , 162 , 163 , 164 , 165 , 164 , 166 , 167 , 168 , 169 , 170 , 171 , 172 , 173 , 174 , 37 , 175 , 176 , 78 , 177 , 21 , 178 , 7 , 17 , 31 , 60 , 60 , 1 , 179 , 180 , 130 , 181 , 107 , 182 , 87 , 183 , 51 , 54 , 17 , 184 , 2 , 4 , 185 , 52 , 44 , 81 ,   160 , 186 , 24 , 25 , 35 , 153 , 187 , 69 , 111 , 125 , 188 , 183 , 44 , 17 , 128 , 129 , 132 , 189 , 190 , 95 , 191 , 192 , 54 , 51 , 116 , 84 , 88 , 18 , 130 , 13 , 121 , 93 , 91 , 127 , 22 , 23 , 45 , 193 , 38 , 39 , 40 , 55 , 194 , 195 , 32 , 22 , 23 , 45 , 72 , 196 , 31 , 80 , 197 , 37 , 175 , 2 , 4 , 198 , 199 , 20 , 200 , 17 , 63 , 64 , 82 , 201 , 2 , 3 , 4 , 19 , 7 , 32 , 8 , 13 , 47 , 202 , 60 , 13 , 47 , 8 , 20 , 200 , 1 , 9 , 10 , 203
      Publication date (Electronic):
      Journal: Nature

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          Abstract

          Most common human traits and diseases have a polygenic pattern of inheritance: DNA sequence variants at many genetic loci influence phenotype. Genome-wide association (GWA) studies have identified >600 variants associated with human traits 1, but these typically explain small fractions of phenotypic variation, raising questions about the utility of further studies. Here, using 183,727 individuals, we show that hundreds of genetic variants, in at least 180 loci, influence adult height, a highly heritable and classic polygenic trait 2, 3. The large number of loci reveals patterns with important implications for genetic studies of common human diseases and traits. First, the 180 loci are not random, but instead are enriched for genes that are connected in biological pathways ( P=0.016), and that underlie skeletal growth defects ( P<0.001). Second, the likely causal gene is often located near the most strongly associated variant: in 13 of 21 loci containing a known skeletal growth gene, that gene was closest to the associated variant. Third, at least 19 loci have multiple independently associated variants, suggesting that allelic heterogeneity is a frequent feature of polygenic traits, that comprehensive explorations of already-discovered loci should discover additional variants, and that an appreciable fraction of associated loci may have been identified. Fourth, associated variants are enriched for likely functional effects on genes, being over-represented amongst variants that alter amino acid structure of proteins and expression levels of nearby genes. Our data explain ∼10% of the phenotypic variation in height, and we estimate that unidentified common variants of similar effect sizes would increase this figure to ∼16% of phenotypic variation (∼20% of heritable variation). Although additional approaches are needed to fully dissect the genetic architecture of polygenic human traits, our findings indicate that GWA studies can identify large numbers of loci that implicate biologically relevant genes and pathways.

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          Rare Variants Create Synthetic Genome-Wide Associations

          Samuel P. Dickson, Kai Wang, Ian D. Krantz … (2010)
          Introduction Efforts to fine map the causal variants responsible for genome-wide association studies (GWAS) signals have been largely predicated on the common disease common variant theory, postulating a common variant as the culprit for observed associations. This has led to extensive resequencing efforts that have been largely unsuccessful [1]–[5]. Here, we explore the possibility that part of the reason for this may be that the disease class causing an observed association may consist of multiple low-frequency variants across large regions of the genome—a phenomenon we call synthetic association. For convenience, these less common variants will be referred to here as “rare,” but we emphasize that we use this term loosely, only to refer to variants less common than those routinely studied in GWAS. The basic idea of how synthetic associations emerge in this model is illustrated in Figure 1, which shows how rare variants, by chance, can occur disproportionately in some parts of a gene genealogy. Any variant “higher up in the genealogy” that partitions those parts of the genealogy containing more disease variants than average will be identified as disease-associated. It is well appreciated that a noncausal variant will show association with a causal variant if the two are in strong linkage disequilibrium (LD). We use the previously introduced term synthetic association [6], however, to describe how such indirect association can occur between a common variant and at least one and possibly many rarer causal variants. Using the term synthetic as opposed to indirect emphasizes that the properties of the association signal are very different when the responsible variant or variants are much less frequent than the marker that carries the signal, as we detail below. 10.1371/journal.pbio.1000294.g001 Figure 1 Example genealogies showing causal variants and the strongest association for a common variant. (A) A genealogy with 10,000 original haplotypes was generated with 3,000 cases and 3,000 controls, genotype relative risk (γ) = 4, and nine causal variants. The branches containing the strongest synthetic association are indicated in blue. The branches containing the rare causal variants are in red. (B) A second genealogy was generated using the same parameters. These genealogies demonstrate two scenarios with genome-wide significant synthetic associations: the first (upper genealogy) had a high risk allele frequency (RAF = 0.49), and the second (lower genealogy) had a low RAF (0.08). To assess the tendency of rare disease-causing variants to create synthetic signals of association that are credited to single polymorphisms that are much more common in the population than the causal variants, we have simulated 10,000 haplotypes based on a coalescent model in a region either with or without recombination (Materials and Methods). We assumed that gene variants that influence disease have an allele frequency between 0.005 and 0.02, which is generally below the range of reliable detection (either by inclusion or indirect representation) using the genome-wide association platforms currently in use. We assumed a baseline probability of disease of φ for individuals with none of the rare genetic risk factors. The presence of at least one rare risk allele at the locus increased the probability of disease from φ to γ. We considered two values of φ (0.01, 0.1) and chose values of the penetrance γ such that the genotypic relative risk (GRR) of the rare causal variants varied incrementally between 2 and 6, where GRR is the ratio γ/φ. These values were chosen to explore the space around a GRR of 4, a threshold above which consistent linkage signals would be expected [7]. We simulated scenarios with one, three, five, seven, and nine rare causal variants. Results Across the conditions we have studied, not only is it possible to achieve genome-wide significance for common variants when one or more rare variants are the only contributors to disease, it is often the likely outcome (Figure 2). Overall, 30% of the simulations were able to detect an association with a common SNP at genome-wide significance (p 5%, Hardy-Weinberg equilibrium p-value >1×10−6, SNP call rate >95%), using the PLINK software [40]. For the sickle cell anemia GWAS, we compared 194 cases and 7,407 controls of inferred African ancestry via multidimensional scaling, with a genomic control inflation factor of 1.01. For hearing loss, we performed a GWAS on 418 cases and 6,892 control subjects, all of whom were of genetically inferred European ancestry via multidimensional scaling, with a genomic control inflation factor of 1.02.
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            Common genetic variation and human traits.

            David B Goldstein (2009)
            Article 0 comments Cited 240 times – based on 0 reviews     Review now
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              Many sequence variants affecting diversity of adult human height.

              Daniel Gudbjartsson, G Walters, Gudmar Thorleifsson … (2008)
              Adult human height is one of the classical complex human traits. We searched for sequence variants that affect height by scanning the genomes of 25,174 Icelanders, 2,876 Dutch, 1,770 European Americans and 1,148 African Americans. We then combined these results with previously published results from the Diabetes Genetics Initiative on 3,024 Scandinavians and tested a selected subset of SNPs in 5,517 Danes. We identified 27 regions of the genome with one or more sequence variants showing significant association with height. The estimated effects per allele of these variants ranged between 0.3 and 0.6 cm and, taken together, they explain around 3.7% of the population variation in height. The genes neighboring the identified loci cluster in biological processes related to skeletal development and mitosis. Association to three previously reported loci are replicated in our analyses, and the strongest association was with SNPs in the ZBTB38 gene.
              Article 0 comments Cited 229 times – based on 0 reviews     Review now
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                Author and article information

                Contributors
                : On behalf of : Procardis Consortium
                Journal
                Journal ID (nlm-journal-id): 0410462
                Journal ID (pubmed-jr-id): 6011
                Journal ID (nlm-ta): Nature
                Title: Nature
                ISSN (Print): 0028-0836
                ISSN (Electronic): 1476-4687
                Publication date Nihms-submitted: 10 August 2010
                Publication date (Electronic): 29 September 2010
                Publication date (Print): 14 October 2010
                Publication date PMC-release: 14 April 2011
                Volume: 467
                Issue: 7317
                Pages: 832-838
                Affiliations
                [1 ] Genetics of Complex Traits, Peninsula College of Medicine and Dentistry, University of Exeter, Exeter, EX1 2LU, UK
                [2 ] Department of Epidemiology, Erasmus MC, Rotterdam, 3015GE, The Netherlands
                [3 ] Department of Internal Medicine, Erasmus MC, Rotterdam, 3015GE, The Netherlands
                [4 ] Netherlands Genomics Initiative (NGI)-sponsored Netherlands Consortium for Healthy Aging (NCHA)
                [5 ] Montreal Heart Institute, Montreal, Quebec, H1T 1C8, Canada
                [6 ] Department of Medicine, Université de Montréal, Montreal, Quebec, H3T 1J4, Canada
                [7 ] Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Department of Health and Human Services, Bethesda, Maryland 20892, USA
                [8 ] Department of Biostatistics, Center for Statistical Genetics, University of Michigan, Ann Arbor, Michigan 48109, USA
                [9 ] Divisions of Genetics and Endocrinology and Program in Genomics, Children's Hospital, Boston, Massachusetts 02115, USA
                [10 ] Metabolism Initiative and Program in Medical and Population Genetics, Broad Institute, Cambridge, Massachusetts 02142, USA
                [11 ] Program in Medical and Population Genetics, Broad Institute of Harvard and Massachusetts Institute of Technology, Cambridge, Massachusetts 02142, USA
                [12 ] Division of Rheumatology, Immunology and Allergy, Brigham and Women's Hospital, Harvard Medical School, Boston, Massachusetts 02115 USA
                [13 ] Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, OX3 7BN, UK
                [14 ] Center for Human Genetic Research, Massachusetts General Hospital, Boston, Massachusetts 02114, USA
                [15 ] Department of Molecular Biology, Massachusetts General Hospital, Boston, Massachusetts 02114, USA
                [16 ] Division of Gastroenterology, Massachusetts General Hospital, Boston, Massachusetts 02114, USA
                [17 ] Wellcome Trust Sanger Institute, Hinxton, Cambridge, CB10 1SA, UK
                [18 ] Department of Twin Research and Genetic Epidemiology, King's College London, Lambeth Palace Rd, London, SE1 7EH, UK
                [19 ] Queensland Statistical Genetics Laboratory, Queensland Institute of Medical Research, Queensland 4006, Australia
                [20 ] deCODE Genetics, 101 Reykjavik, Iceland
                [21 ] Department of Neurology, Boston University School of Medicine, Boston, Massachusetts 02118, USA
                [22 ] Department of Nutrition, Harvard School of Public Health, Boston, Massachusetts 02115, USA
                [23 ] Channing Laboratory, Department of Medicine, Brigham and Women's Hospital and Harvard Medical School, Boston, Massachusetts 02115, USA
                [24 ] Icelandic Heart Association, Kopavogur, Iceland
                [25 ] University of Iceland, Reykjavik, Iceland
                [26 ] McGill University and Genome Québec Innovation Centre, Montréal, Québec H3A 1A4, Canada
                [27 ] Department of Human Genetics, McGill University Health Centre, McGill University, Montréal, Québec H3G 1A4, Canada
                [28 ] Department of Medical Genetics, McGill University Health Centre, McGill University, Montréal, Québec H3G 1A4, Canada
                [29 ] Departments of Epidemiology and Biostatistics, Harvard School of Public Health, Cambridge, Massachusetts 02138, USA
                [30 ] Regensburg University Medical Center, Department of Epidemiology and Preventive Medicine, 93053 Regensburg, Germany
                [31 ] Institute of Epidemiology, Helmholtz Zentrum München - German Research Center for Environmental Health, 85764 Neuherberg, Germany
                [32 ] MRC Epidemiology Unit, Institute of Metabolic Science, Addenbrooke's Hospital, Cambridge, CB2 0QQ, UK
                [33 ] University of Melbourne, Parkville 3010, Australia
                [34 ] Department of Primary Industries, Melbourne, Victoria 3001, Australia
                [35 ] Department of Genetics and Pathology, Rudbeck Laboratory, University of Uppsala, SE-75185 Uppsala, Sweden
                [36 ] Department of Cancer Research and Molecular Medicine, Faculty of Medicine, Norwegian University of Science and Technology (NTNU), Trondheim, N-7489, Norway
                [37 ] Department of Genetics, Washington University School of Medicine, St Louis, Missouri 63110, USA
                [38 ] Estonian Genome Center, University of Tartu, Tartu 50410, Estonia
                [39 ] Estonian Biocenter, Tartu 51010, Estonia
                [40 ] Institute of Molecular and Cell Biology, University of Tartu, Tartu 51010, Estonia
                [41 ] Department of Medical Genetics, University of Lausanne, 1005 Lausanne, Switzerland
                [42 ] Swiss Institute of Bioinformatics, 1015 Lausanne, Switzerland
                [43 ] Clinical Pharmacology, William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary, University of London, London, UK
                [44 ] Clinical Pharmacology and Barts and The London Genome Centre, William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, Charterhouse Square, London EC1M 6BQ, UK
                [45 ] Department of Epidemiology, Harvard School of Public Health, Boston, Massachusetts 02115, USA
                [46 ] Department of Biostatistics, Harvard School of Public Health, Boston, Massachusetts 02115, USA
                [47 ] Oxford Centre for Diabetes, Endocrinology and Metabolism, University of Oxford, Oxford, OX3 7LJ, UK
                [48 ] Hudson Alpha Institute for Biotechnology, Huntsville, Alabama 35806, USA
                [49 ] Interfaculty Institute for Genetics and Functional Genomics, Ernst-Moritz-Arndt-University Greifswald, 17487 Greifswald, Germany
                [50 ] Cardiovascular Health Resarch Unit and Department of Medicine, University of Washington, Seattle, Washington 98101, USA
                [51 ] MRC Human Genetics Unit, Institute for Genetics and Molecular Medicine, Western General Hospital, Edinburgh, EH4 2XU, Scotland, UK
                [52 ] Department of Biological Psychology, VU University Amsterdam, 1081 BT Amsterdam, The Netherlands
                [53 ] Core Genotyping Facility, SAIC-Frederick, Inc., NCI-Frederick, Frederick, Maryland 21702, USA
                [54 ] Department of Medicine, Stanford University School of Medicine, Stanford, California 94305, USA
                [55 ] Department of Epidemiology, School of Public Health, University of North Carolina at Chapel Hill, Chapel Hill, North Carolina 27514 USA
                [56 ] Andrija Stampar School of Public Health, Medical School, University of Zagreb, 10000 Zagreb, Croatia
                [57 ] Gen-Info Ltd, 10000 Zagreb, Croatia
                [58 ] Universität zu Lübeck, Institut für Medizinische Biometrie und Statistik, 23562 Lübeck, Germany
                [59 ] Department of Oncology, University of Cambridge, Cambridge, CB1 8RN, UK
                [60 ] Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, 171 77 Stockholm, Sweden
                [61 ] Center for Human Genomics, Wake Forest University, Winston-Salem, North Carolina 27157, USA
                [62 ] Neurogenetics Laboratory, Queensland Institute of Medical Research, Queensland 4006, Australia
                [63 ] Institute for Molecular Medicine Finland (FIMM), University of Helsinki, 00014, Helsinki, Finland
                [64 ] National Institute for Health and Welfare, Department of Chronic Disease Prevention, Unit of Public Health Genomics, 00014, Helsinki, Finland
                [65 ] Department of Genome Sciences, University of Washington, Seattle, 98195 Washington, USA
                [66 ] Division of Preventive Medicine, Brigham and Women's Hospital, Boston, Massachusetts 02215, USA
                [67 ] Harvard Medical School, Boston, Massachusetts 02115, USA
                [68 ] Program in Molecular and Genetic Epidemiology, Harvard School of Public Health, Boston, Massachusetts 02115, USA
                [69 ] Department of Epidemiology and Biostatistics, School of Public Health, Faculty of Medicine, Imperial College London, London, W2 1PG, UK
                [70 ] Centre for Genetic Epidemiology and Biostatistics, University of Western Australia, Crawley, Western Australia 6009, Australia
                [71 ] Royal National Hospital for Rheumatic Diseases and University of Bath, Bath, BA1 1RL, UK
                [72 ] Department of Medicine, University of Maryland School of Medicine, Baltimore, Maryland 21201, USA
                [73 ] Genetics Department, Rosetta Inpharmatics, a Wholly Owned Subsidiary of Merck & Co. Inc., Seattle, Washington 98109, USA
                [74 ] Department of Internal Medicine, University of Oulu, 90014 Oulu, Finland
                [75 ] MGH Weight Center, Massachusetts General Hospital, Boston, Massachusetts 02114, USA
                [76 ] Stanford University School of Medicine, Stanford, California 93405, USA
                [77 ] Department of Internal Medicine, Institute of Medicine, Sahlgrenska Academy, University of Gothenburg, 413 45 Gothenburg, Sweden
                [78 ] Departments of Biostatistics, University of Washington, Seattle, Washington 98195, USA
                [79 ] Ludwig-Maximilians-University, Department of Medicine I, University Hospital Grosshadern, 81377 Munich, Germany
                [80 ] Ludwig-Maximilians-Universität, Institute of Medical Informatics, Biometry and Epidemiology, Chair of Epidemiology, 81377 Munich, Germany
                [81 ] Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts 02118, USA
                [82 ] The Broad Institute of Harvard and MIT, Cambridge, Massachusetts 02142, USA
                [83 ] Department of Psychiatry, Harvard Medical School, Boston, Massachusetts 02115, USA
                [84 ] Department of Cardiovascular Sciences, University of Leicester, Glenfield Hospital, Leicester, LE3 9QP, UK
                [85 ] University of Milan, Department of Medicine, Surgery and Dentistry, 20139 Milano, Italy
                [86 ] KOS Genetic Srl, 20123 Milan, Italy
                [87 ] Istituto di Neurogenetica e Neurofarmacologia del CNR, Monserrato, 09042, Cagliari, Italy
                [88 ] Leicester NIHR Biomedical Research Unit in Cardiovascular Disease, Glenfield Hospital, Leicester, LE3 9QP, UK
                [89 ] Department of Health Sciences, University of Leicester, University Road, Leicester, LE1 7RH, UK
                [90 ] Universität zu Lübeck, Institut für Medizinische Biometrie und Statistik, 23562 Lübeck, Germany
                [91 ] Lund University Diabetes Centre, Department of Clinical Sciences, Lund University, 20502 Malmö, Sweden
                [92 ] Multidisciplinary Cardiovascular Research Centre (MCRC), Leeds Institute of Genetics, Health and Therapeutics (LIGHT), University of Leeds, Leeds LS2 9JT, UK
                [93 ] Centre for Population Health Sciences, University of Edinburgh, Teviot Place, Edinburgh, EH8 9AG, Scotland
                [94 ] University Vita-Salute San Raffaele, Division of Nephrology and Dialysis, 20132 Milan, Italy
                [95 ] Institute of Genetic Medicine, European Academy Bozen/Bolzano (EURAC), Bolzano/Bozen, 39100, Italy. Affiliated Institute of the University of Lübeck, Lübeck, Germany
                [96 ] British Heart Foundation Glasgow Cardiovascular Research Centre, University of Glasgow, Glasgow, G12 8TA, UK
                [97 ] Northshore University Healthsystem, Evanston, Ilinois 60201, USA
                [98 ] Cardiovascular Epidemiology and Genetics, Institut Municipal D'investigacio Medica and CIBER Epidemiología y Salud Pública, Barcelona, Spain
                [99 ] Department of General Practice and Primary health Care, University of Helsinki, Helsinki, Finland
                [100 ] National Institute for Health and Welfare, 00271 Helsinki, Finland
                [101 ] Helsinki University Central Hospital, Unit of General Practice, 00280 Helsinki, Finland
                [102 ] Folkhalsan Research Centre, 00250 Helsinki, Finland
                [103 ] Vasa Central Hospital, 65130 Vasa, Finland
                [104 ] Center for Neurobehavioral Genetics, University of California, Los Angeles, California 90095, USA
                [105 ] Hypertension and Cardiovascular Prevention Center, University of Sassari, 07100 Sassari, Italy
                [106 ] Department of Clinical Sciences/Obstetrics and Gynecology, University of Oulu, 90014 Oulu, Finland
                [107 ] National Institute for Health and Welfare, Department of Chronic Disease Prevention, Chronic Disease Epidemiology and Prevention Unit, 00014, Helsinki, Finland
                [108 ] PathWest Laboratory of Western Australia, Department of Molecular Genetics, J Block, QEII Medical Centre, Nedlands, Western Australia 6009, Australia
                [109 ] Busselton Population Medical Research Foundation Inc., Sir Charles Gairdner Hospital, Nedlands, Western Australia 6009, Australia
                [110 ] National Institute for Health and Welfare, Department of Chronic Disease Prevention, Population Studies Unit, 20720 Turku, Finland
                [111 ] Institute of Health Sciences, University of Oulu, 90014 Oulu, Finland
                [112 ] Interdisciplinary Centre for Clinical Research, University of Leipzig, 04103 Leipzig, Germany
                [113 ] Finnish Institute of Occupational Health, 90220 Oulu, Finland
                [114 ] Human Genetics, Genome Institute of Singapore, Singapore 138672, Singapore
                [115 ] Transplantation Laboratory, Haartman Institute, University of Helsinki, 00014, Helsinki, Finland
                [116 ] Croatian Centre for Global Health, School of Medicine, University of Split, Split 21000, Croatia
                [117 ] Institute of Human Genetics, Klinikum rechts der Isar der Technischen Universität München, 81675 Munich, Germany
                [118 ] Institute of Human Genetics, Helmholtz Zentrum München - German Research Center for Environmental Health, 85764 Neuherberg, Germany
                [119 ] Department of Pathology and Molecular Medicine, McMaster University, Hamilton, Ontario L8N3Z5, Canada
                [120 ] Amgen, Cambridge, Massachusetts 02139, USA
                [121 ] Department of Cardiovascular Medicine, University of Oxford, Level 6 West Wing, John Radcliffe Hospital, Headley Way, Headington, Oxford, OX3 9DU
                [122 ] Institut für Klinische Chemie und Laboratoriumsmedizin, Universität Greifswald, 17475 Greifswald, Germany
                [123 ] Finnish Twin Cohort Study, Department of Public Health, University of Helsinki, 00014, Helsinki, Finland
                [124 ] Obesity Research unit, Department of Psychiatry, Helsinki University Central Hospital, Helsinki, Finland
                [125 ] National Institute for Health and Welfare, 90101 Oulu, Finland
                [126 ] Department of Clinical Sciences, Lund University, 20502 Malmö, Sweden
                [127 ] Medical Genetics Institute, Cedars-Sinai Medical Center, Los Angeles, California 90048, USA
                [128 ] Department of Haematology, University of Cambridge, Cambridge CB2 0PT, UK
                [129 ] NHS Blood and Transplant, Cambridge Centre, Cambridge, CB2 0PT, UK
                [130 ] Institut für Community Medicine, 17489 Greifswald, Germany
                [131 ] Division of Cardiology, Cardiovascular Laboratory, Helsinki University Central Hospital, 00029 Helsinki, Finland
                [132 ] Department of Psychiatry/EMGO Institute, VU University Medical Center, 1081 BT Amsterdam, The Netherlands
                [133 ] Department of Psychiatrics, Lapland Central Hospital, 96101 Rovaniemi, Finland
                [134 ] Cardiovascular Medicine, University of Oxford, Wellcome Trust Centre for Human Genetics, Oxford, OX3 7BN, UK
                [135 ] Avon Longitudinal Study of Parents and Children (ALSPAC) Laboratory, Department of Social Medicine, University of Bristol, Bristol, BS8 2BN, UK
                [136 ] Comprehensive Cancer Center East, 6501 BG Nijmegen, The Netherlands
                [137 ] Service of Medical Genetics, Centre Hospitalier Universitaire Vaudois (CHUV) University Hospital, 1011 Lausanne, Switzerland
                [138 ] School of Pathology and Laboratory Medicine, University of Western Australia, Nedlands, Western Australia 6009, Australia
                [139 ] Department of Physiology and Biophysics, Keck School of Medicine, University of Southern California, Los Angeles, California 90033, USA
                [140 ] National Human Genome Research Institute, National Institutes of Health, Bethesda, Maryland 20892, USA
                [141 ] Department of Endocrinology, Radboud University Nijmegen Medical Centre, 6500 HB Nijmegen, The Netherlands
                [142 ] Atherosclerosis Research Unit, Department of Medicine, Solna, Karolinska Institutet, Karolinska University Hospital, 171 76 Stockholm, Sweden
                [143 ] Division of Research, Kaiser Permanente Northern California, Oakland, California 94612, USA
                [144 ] Department of Epidemiology and Biostatistics, University of California, San Francisco, San Francisco, California 94107, USA
                [145 ] Department of Clinical Physiology, University of Tampere and Tampere University Hospital, 33520 Tampere, Finland
                [146 ] National Institute for Health and Welfare, Department of Mental Health and Substance Abuse Services, Unit for Child and Adolescent Mental Health, 00271 Helsinki, Finland
                [147 ] Cardiovascular Research Center and Cardiology Division, Massachusetts General Hospital, Boston, Massachusetts 02114, USA
                [148 ] Framingham Heart Study of the National, Heart, Lung, and Blood Institute and Boston University, Framingham, Massachusetts 01702, USA
                [149 ] Department of Medicine, Harvard Medical School, Boston, Massachusetts 02115, USA
                [150 ] Department of Epidemiology, Biostatistics and HTA, Radboud University Nijmegen Medical Centre, 6500 HB Nijmegen, The Netherlands
                [151 ] Department of Urology, Radboud University Nijmegen Medical Centre, 6500 HB Nijmegen, The Netherlands
                [152 ] Zentrum für Zahn-, Mund- und Kieferheilkunde, 17489 Greifswald, Germany
                [153 ] Laboratory of Epidemiology, Demography, Biometry, National Institute on Aging, National Institutes of Health, Bethesda, Maryland 20892, USA
                [154 ] Department of Clinical Chemistry, University of Tampere and Tampere University Hospital, 33520 Tampere, Finland
                [155 ] Department of Medicine, Division of Geriatrics, University of Mississippi Medical Center, Jackson, Mississippi 39216, USA
                [156 ] School of Medicine and Pharmacology, University of Western Australia, Perth, Western Australia 6009, Australia
                [157 ] National, Lung, and Blood Institute, National Institutes of Health, Framingham, Massachusetts 01702, USA
                [158 ] Department of Clinical Genetics, Erasmus MC, Rotterdam, 3015GE, The Netherlands
                [159 ] Research Centre of Applied and Preventive Cardiovascular Medicine, University of Turku and the Department of Clinical Physiology, Turku University Hospital, 20520 Turku, Finland
                [160 ] Universität zu Lübeck, Medizinische Klinik II, 23562 Lübeck, Germany
                [161 ] Geriatrics Research and Education Clinical Center, Baltimore Veterans Administration Medical Center, Baltimore, Maryland 21201, USA
                [162 ] Cardiovascular Health Research Unit, University of Washington, Seattle, Washington 98101, USA
                [163 ] Departments of Medicine and Epidemiology, University of Washington, Seattle, Washington 98195, USA
                [164 ] Department of Medicine, University of Leipzig, 04103 Leipzig, Germany
                [165 ] LIFE Study Centre, University of Leipzig, Leipzig, Germany
                [166 ] Coordination Centre for Clinical Trials, University of Leipzig, Härtelstr. 16-18, 04103 Leipzig, Germany
                [167 ] National Institute for Health and Welfare, Diabetes Prevention Unit, 00271 Helsinki, Finland
                [168 ] Hjelt Institute, Department of Public Health, University of Helsinki, 00014 Helsinki, Finland
                [169 ] South Ostrobothnia Central Hospital, 60220 Seinajoki, Finland
                [170 ] Department of Human Genetics and Center of Medical Systems Biology, Leiden University Medical Center, 2333 ZC Leiden, the Netherlands
                [171 ] Department of Medicine, University of Turku and Turku University Hospital, 20520 Turku, Finland
                [172 ] Department of Psychiatry and Midwest Alcoholism Research Center, Washington University School of Medicine, St Louis, Missouri 63108, USA
                [173 ] Genetic Epidemiology Laboratory, Queensland Institute of Medical Research, Queensland 4006, Australia
                [174 ] Molecular Epidemiology Laboratory, Queensland Institute of Medical Research, Queensland 4006, Australia
                [175 ] Division of Biostatistics, Washington University School of Medicine, St. Louis, Missouri 63110, USA
                [176 ] Department of Forensic Molecular Biology, Erasmus MC, Rotterdam, 3015GE, The Netherlands
                [177 ] Collaborative Health Studies Coordinating Center, Seattle, Washington 98115, USA
                [178 ] Human Genetics Center and Institute of Molecular Medicine and Division of Epidemiology, University of Texas Health Science Center, Houston, Texas 77030, USA
                [179 ] Klinik und Poliklinik für Innere Medizin II, Universität Regensburg, 93053 Regensburg, Germany
                [180 ] Regensburg University Medical Center, Innere Medizin II, 93053 Regensburg, Germany
                [181 ] Centre National de Genotypage, Evry, Paris 91057, France
                [182 ] Christian-Albrechts-University, University Hospital Schleswig-Holstein, Institute for Clinical Molecular Biology and Department of Internal Medicine I, Schittenhelmstrasse 12, 24105 Kiel
                [183 ] Genetics Division, GlaxoSmithKline, King of Prussia, Pennsylvania 19406, USA
                [184 ] University of Cambridge Metabolic Research Labs, Institute of Metabolic Science Addenbrooke's Hospital, CB2 OQQ, Cambridge, UK
                [185 ] Department of Genetics, University of North Carolina, Chapel Hill, North Carolina 27599, USA
                [186 ] Division of Intramural Research, National Heart, Lung and Blood Institute, Framingham Heart Study, Framingham, Massachusetts 01702, USA
                [187 ] New York University Medical Center, New York, New York 10016, USA
                [188 ] Biocenter Oulu, University of Oulu, 90014 Oulu, Finland
                [189 ] Department of Psychiatry, Leiden University Medical Centre, 2300 RC Leiden, The Netherlands
                [190 ] Department of Psychiatry, University Medical Centre Groningen, 9713 GZ Groningen, The Netherlands
                [191 ] Department of Neurology, General Central Hospital, Bolzano, Italy
                [192 ] Department of Neurology, University of Lübeck, Lübeck, Germany
                [193 ] Department of Epidemiology and Population Health, Albert Einstein College of Medicine, Bronx, New York 10461, USA
                [194 ] Carolina Center for Genome Sciences, School of Public Health, University of North Carolina Chapel Hill, Chapel Hill, North Carolina 27514, USA
                [195 ] Laboratory of Genetics, National Institute on Aging, Baltimore, Maryland 21224, USA
                [196 ] Division of Community Health Sciences, St George's, University of London, London, SW17 0RE, UK
                [197 ] Klinikum Grosshadern, 81377 Munich, Germany
                [198 ] Pacific Biosciences, Menlo Park, California 94025, USA
                [199 ] Sage Bionetworks, Seattle, Washington 98109, USA
                [200 ] Faculty of Medicine, University of Iceland, 101 Reykjavík, Iceland
                [201 ] Department of Medical Genetics, University of Helsinki, 00014 Helsinki, Finland
                [202 ] NIHR Oxford Biomedical Research Centre, Churchill Hospital, Oxford, OX3 7LJ, UK
                [203 ] Department of Genetics, Harvard Medical School, Boston, Massachusetts 02115, USA
                Author notes
                Corresponding Authors: Michael Weedon, Genetics of Complex Traits, Peninsula College of Medicine and Dentistry, University of Exeter, Exeter, EX1 2LU, UK, michael.weedon@ 123456pms.ac.uk , Gonçalo Abecasis, Department of Biostatistics, Center for Statistical Genetics, University of Michigan, Ann Arbor, Michigan 48109, USA, goncalo@ 123456umich.edu , Kari Stefansson, deCODE Genetics, 101 Reykjavik, Iceland, kstefans@ 123456decode.is , Timothy Frayling, Genetics of Complex Traits, Peninsula College of Medicine and Dentistry, University of Exeter, Exeter, EX1 2LU, UK, tim.frayling@ 123456pms.ac.uk , Joel Hirschhorn, Children's Hospital, Harvard Medical School, Broad Institute, Boston, Massachusetts 02115, USA, joelh@ 123456broadinstitute.org
                [*]

                These authors contributed equally

                Article
                Manuscript ID: nihpa225625
                DOI: 10.1038/nature09410
                PMC ID: 2955183
                PubMed ID: 20881960
                SO-VID: a9921794-87e2-4751-b66a-a693e25ca014
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                Award ID: R01 HL087647-01 ||HL
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                Award ID: R01 HL087641-01 ||HL
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                Funded by: National Cancer Institute : NCI
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                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 HL059367-10 ||HL
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 HL043851-10 ||HL
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 HG002651-05 ||HG
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 DK075787-03 ||DK
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 DK075681-02 ||DK
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 DK073490-01 ||DK
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 DK072193-05 ||DK
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 DK068336-01 ||DK
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 DK058845-11 ||DK
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 DA012854-09 ||DA
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 CA104021-02 ||CA
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 CA067262-14 ||CA
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 CA065725-14 ||CA
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 CA050385-20 ||CA
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 CA047988-20 ||CA
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 AG031890-02 ||AG
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 AA014041-05 ||AA
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 AA013326-05 ||AA
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 AA013321-05 ||AA
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 AA013320-04 ||AA
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: R01 AA007535-08 ||AA
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: P01 CA087969-12 ||CA
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: K23 DK080145-01 ||DK
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: K08 AR055688-04 ||AR
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: K08 AR055688-03 ||AR
                Funded by: National Institute of Mental Health : NIMH
                Funded by: National Heart, Lung, and Blood Institute : NHLBI
                Funded by: National Human Genome Research Institute : NHGRI
                Funded by: National Institute of General Medical Sciences : NIGMS
                Funded by: National Institute of Diabetes and Digestive and Kidney Diseases : NIDDK
                Funded by: National Cancer Institute : NCI
                Funded by: National Institute on Drug Abuse : NIDA
                Funded by: National Institute on Aging : NIA
                Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
                Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
                Award ID: F32 DK079466-01 ||DK
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