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

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

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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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      PLINK: a tool set for whole-genome association and population-based linkage analyses.

      Whole-genome association studies (WGAS) bring new computational, as well as analytic, challenges to researchers. Many existing genetic-analysis tools are not designed to handle such large data sets in a convenient manner and do not necessarily exploit the new opportunities that whole-genome data bring. To address these issues, we developed PLINK, an open-source C/C++ WGAS tool set. With PLINK, large data sets comprising hundreds of thousands of markers genotyped for thousands of individuals can be rapidly manipulated and analyzed in their entirety. As well as providing tools to make the basic analytic steps computationally efficient, PLINK also supports some novel approaches to whole-genome data that take advantage of whole-genome coverage. We introduce PLINK and describe the five main domains of function: data management, summary statistics, population stratification, association analysis, and identity-by-descent estimation. In particular, we focus on the estimation and use of identity-by-state and identity-by-descent information in the context of population-based whole-genome studies. This information can be used to detect and correct for population stratification and to identify extended chromosomal segments that are shared identical by descent between very distantly related individuals. Analysis of the patterns of segmental sharing has the potential to map disease loci that contain multiple rare variants in a population-based linkage analysis.
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        Finding the missing heritability of complex diseases.

        Genome-wide association studies have identified hundreds of genetic variants associated with complex human diseases and traits, and have provided valuable insights into their genetic architecture. Most variants identified so far confer relatively small increments in risk, and explain only a small proportion of familial clustering, leading many to question how the remaining, 'missing' heritability can be explained. Here we examine potential sources of missing heritability and propose research strategies, including and extending beyond current genome-wide association approaches, to illuminate the genetics of complex diseases and enhance its potential to enable effective disease prevention or treatment.
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          A second generation human haplotype map of over 3.1 million SNPs.

          We describe the Phase II HapMap, which characterizes over 3.1 million human single nucleotide polymorphisms (SNPs) genotyped in 270 individuals from four geographically diverse populations and includes 25-35% of common SNP variation in the populations surveyed. The map is estimated to capture untyped common variation with an average maximum r2 of between 0.9 and 0.96 depending on population. We demonstrate that the current generation of commercial genome-wide genotyping products captures common Phase II SNPs with an average maximum r2 of up to 0.8 in African and up to 0.95 in non-African populations, and that potential gains in power in association studies can be obtained through imputation. These data also reveal novel aspects of the structure of linkage disequilibrium. We show that 10-30% of pairs of individuals within a population share at least one region of extended genetic identity arising from recent ancestry and that up to 1% of all common variants are untaggable, primarily because they lie within recombination hotspots. We show that recombination rates vary systematically around genes and between genes of different function. Finally, we demonstrate increased differentiation at non-synonymous, compared to synonymous, SNPs, resulting from systematic differences in the strength or efficacy of natural selection between populations.
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            Author and article information

            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

            Contributors
            On behalf of : Procardis Consortium
            Journal
            0410462
            6011
            Nature
            Nature
            0028-0836
            1476-4687
            10 August 2010
            29 September 2010
            14 October 2010
            14 April 2011
            : 467
            : 7317
            : 832-838
            2955183
            20881960
            10.1038/nature09410
            nihpa225625

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            Funding
            Funded by: National Institute of Mental Health : NIMH
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            Funded by: National Institute of General Medical Sciences : NIGMS
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            Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
            Award ID: U01 MH079470-03 ||MH
            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
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            Funded by: National Cancer Institute : NCI
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            Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
            Award ID: U01 MH079469-03 ||MH
            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
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            Funded by: National Cancer Institute : NCI
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            Award ID: U01 HL084756-03 ||HL
            Funded by: National Institute of Mental Health : NIMH
            Funded by: National Heart, Lung, and Blood Institute : NHLBI
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            Award ID: U01 HL084729-03 ||HL
            Funded by: National Institute of Mental Health : NIMH
            Funded by: National Heart, Lung, and Blood Institute : NHLBI
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            Funded by: National Institute of General Medical Sciences : NIGMS
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            Award ID: U01 HL080295-04 ||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
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            Award ID: U01 HL072515-06 ||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
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            Award ID: U01 HL069757-10 ||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
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            Funded by: National Institute on Aging : NIA
            Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
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            Award ID: U01 HG005214-02 ||HG
            Funded by: National Institute of Mental Health : NIMH
            Funded by: National Heart, Lung, and Blood Institute : NHLBI
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            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
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            Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
            Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
            Award ID: U01 HG004402-02 ||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
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            Award ID: U01 HG004399-02 ||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
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            Funded by: National Institute on Alcohol Abuse and Alcoholism : NIAAA
            Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
            Award ID: U01 GM074518-05 ||GM
            Funded by: National Institute of Mental Health : NIMH
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            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
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            Award ID: U01 DK062370-08 ||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
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            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: U01 CA098233-08 ||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: U01 CA049449-21 ||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: RL1 MH083268-05 ||MH
            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: RC2 HG005581-02 ||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 MH081800-01 ||MH
            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 MH067257-04 ||MH
            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 MH061675-09 ||MH
            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 MH060879-08 ||MH
            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 MH060870-09 ||MH
            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 MH059588-08 ||MH
            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 MH059587-09 ||MH
            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 MH059586-08 ||MH
            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 MH059571-05 ||MH
            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 MH059566-08 ||MH
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            Funded by: National Heart, Lung, and Blood Institute : NHLBI
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            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 MH059565-06 ||MH
            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 MH059160-04 ||MH
            Funded by: National Institute of Mental Health : NIMH
            Funded by: National Heart, Lung, and Blood Institute : NHLBI
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            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 HL088119-01 ||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 HL087700-03 ||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
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            Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
            Award ID: R01 HL087679-01 ||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
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            Award ID: R01 HL087676-01 ||HL
            Funded by: National Institute of Mental Health : NIMH
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            Funded by: National Cancer Institute : NCI
            Funded by: National Institute on Drug Abuse : NIDA
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            Award ID: R01 HL087652-01 ||HL
            Funded by: National Institute of Mental Health : NIMH
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            Funded by: National Institute of General Medical Sciences : NIGMS
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            Funded by: National Cancer Institute : NCI
            Funded by: National Institute on Drug Abuse : NIDA
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            Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
            Award ID: R01 HL087647-01 ||HL
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            Funded by: National Institute on Drug Abuse : NIDA
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            Award ID: R01 HL087641-01 ||HL
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            Funded by: National Cancer Institute : NCI
            Funded by: National Institute on Drug Abuse : NIDA
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            Award ID: R01 HL086694-02 ||HL
            Funded by: National Institute of Mental Health : NIMH
            Funded by: National Heart, Lung, and Blood Institute : NHLBI
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            Funded by: National Cancer Institute : NCI
            Funded by: National Institute on Drug Abuse : NIDA
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            Funded by: National Institute of Arthritis and Musculoskeletal and Skin Diseases : NIAMS
            Award ID: R01 HL071981-07 ||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
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            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
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            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
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            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
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            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
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            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
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            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
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            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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