A comprehensive 1000 Genomes–based genome-wide association meta-analysis of coronary artery disease (original) (raw)
Change history
14 September 2015
In the version of this article initially published online, there was a typographical error in the third sentence of the abstract. The corrected sentence should read: "In addition to confirming most known CAD-associated loci, we identified ten new loci (eight additive and two recessive) that contain candidate causal genes newly implicating biological processes in vessel walls." The error has been corrected for the print, PDF and HTML versions of this article.
References
- Kessler, T., Erdmann, J. & Schunkert, H. Genetics of coronary artery disease and myocardial infarction—2013. Curr. Cardiol. Rep. 15, 368 (2013).
Article PubMed Google Scholar - O'Donnell, C.J. & Nabel, E.G. Genomics of cardiovascular disease. N. Engl. J. Med. 365, 2098–2109 (2011).
Article CAS PubMed Google Scholar - CARDIoGRAMplusC4D Consortium. Large-scale association analysis identifies new risk loci for coronary artery disease. Nat. Genet. 45, 25–33 (2013).
- Coronary Artery Disease Genetics (C4D) Consortium. A genome-wide association study in Europeans and South Asians identifies five new loci for coronary artery disease. Nat. Genet. 43, 339–344 (2011).
- 1000 Genomes Project Consortium. An integrated map of genetic variation from 1,092 human genomes. Nature 491, 56–65 (2012).
- Wang, F. et al. Genome-wide association identifies a susceptibility locus for coronary artery disease in the Chinese Han population. Nat. Genet. 43, 345–349 (2011).
Article CAS PubMed Google Scholar - IBC 50K CAD Consortium. Large-scale gene-centric analysis identifies novel variants for coronary artery disease. PLoS Genet. 7, e1002260 (2011).
- Clarke, R. et al. Genetic variants associated with Lp(a) lipoprotein level and coronary disease. N. Engl. J. Med. 361, 2518–2528 (2009).
Article CAS PubMed Google Scholar - Bennet, A.M. et al. Association of apolipoprotein E genotypes with lipid levels and coronary risk. J. Am. Med. Assoc. 298, 1300–1311 (2007).
Article CAS Google Scholar - Benn, M., Nordestgaard, B.G., Grande, P., Schnohr, P. & Tybjaerg-Hansen, A. PCSK9 R46L, low-density lipoprotein cholesterol levels, and risk of ischemic heart disease: 3 independent studies and meta-analyses. J. Am. Coll. Cardiol. 55, 2833–2842 (2010).
Article CAS PubMed Google Scholar - Cohen, J.C., Boerwinkle, E., Mosley, T.H. Jr. & Hobbs, H.H. Sequence variations in PCSK9, low LDL, and protection against coronary heart disease. N. Engl. J. Med. 354, 1264–1272 (2006).
Article CAS PubMed Google Scholar - Myocardial Infarction Genetics Consortium. A PCSK9 missense variant associated with a reduced risk of early-onset myocardial infarction. N. Engl. J. Med. 358, 2299–2300 (2008).
- Peloso, G.M. et al. Association of low-frequency and rare coding-sequence variants with blood lipids and coronary heart disease in 56,000 whites and blacks. Am. J. Hum. Genet. 94, 223–232 (2014).
Article CAS PubMed PubMed Central Google Scholar - Davies, R.W. et al. A genome-wide association study for coronary artery disease identifies a novel susceptibility locus in the major histocompatibility complex. Circ Cardiovasc Genet 5, 217–225 (2012).
Article CAS PubMed PubMed Central Google Scholar - Wellcome Trust Case Control Consortium. Genome-wide association study of 14,000 cases of seven common diseases and 3,000 shared controls. Nature 447, 661–678 (2007).
- Dickson, S.P., Wang, K., Krantz, I., Hakonarson, H. & Goldstein, D.B. Rare variants create synthetic genome-wide associations. PLoS Biol. 8, e1000294 (2010).
Article CAS PubMed PubMed Central Google Scholar - Yang, J., Lee, S.H., Goddard, M.E. & Visscher, P.M. GCTA: a tool for genome-wide complex trait analysis. Am. J. Hum. Genet. 88, 76–82 (2011).
Article CAS PubMed PubMed Central Google Scholar - Wang, K., Li, M. & Hakonarson, H. ANNOVAR: functional annotation of genetic variants from high-throughput sequencing data. Nucleic Acids Res. 38, e164 (2010).
Article PubMed PubMed Central Google Scholar - Tang, T. et al. hNOA1 interacts with complex I and DAP3 and regulates mitochondrial respiration and apoptosis. J. Biol. Chem. 284, 5414–5424 (2009).
Article CAS PubMed PubMed Central Google Scholar - Chong, J.A. et al. REST: a mammalian silencer protein that restricts sodium channel gene expression to neurons. Cell 80, 949–957 (1995).
Article CAS PubMed Google Scholar - Cheong, A. et al. Downregulated REST transcription factor is a switch enabling critical potassium channel expression and cell proliferation. Mol. Cell 20, 45–52 (2005).
Article CAS PubMed Google Scholar - Hao, K. et al. Lung eQTLs to help reveal the molecular underpinnings of asthma. PLoS Genet. 8, e1003029 (2012).
Article CAS PubMed PubMed Central Google Scholar - Salvi, E. et al. Genomewide association study using a high-density single nucleotide polymorphism array and case-control design identifies a novel essential hypertension susceptibility locus in the promoter region of endothelial NO synthase. Hypertension 59, 248–255 (2012).
Article CAS PubMed Google Scholar - Erdmann, J. et al. Dysfunctional nitric oxide signalling increases risk of myocardial infarction. Nature 504, 432–436 (2013).
Article CAS PubMed Google Scholar - Casas, J.P. et al. Endothelial nitric oxide synthase gene polymorphisms and cardiovascular disease: a HuGE review. Am. J. Epidemiol. 164, 921–935 (2006).
Article PubMed Google Scholar - Chacón-Martínez, C.A. et al. The switch-associated protein 70 (SWAP-70) bundles actin filaments and contributes to the regulation of F-actin dynamics. J. Biol. Chem. 288, 28687–28703 (2013).
Article CAS PubMed PubMed Central Google Scholar - Zeller, T. et al. Genetics and beyond—the transcriptome of human monocytes and disease susceptibility. PLoS ONE 5, e10693 (2010).
Article CAS PubMed PubMed Central Google Scholar - Fairfax, B.P. et al. Innate immune activity conditions the effect of regulatory variants upon monocyte gene expression. Science 343, 1246949 (2014).
Article CAS PubMed PubMed Central Google Scholar - Grundberg, E. et al. Mapping _cis_- and _trans_-regulatory effects across multiple tissues in twins. Nat. Genet. 44, 1084–1089 (2012).
Article CAS PubMed PubMed Central Google Scholar - Ashcroft, G.S. et al. Mice lacking Smad3 show accelerated wound healing and an impaired local inflammatory response. Nat. Cell Biol. 1, 260–266 (1999).
Article CAS PubMed Google Scholar - Samani, N.J. et al. Genomewide association analysis of coronary artery disease. N. Engl. J. Med. 357, 443–453 (2007).
Article CAS PubMed PubMed Central Google Scholar - Silvestre, J.S. et al. Lactadherin promotes VEGF-dependent neovascularization. Nat. Med. 11, 499–506 (2005).
Article CAS PubMed Google Scholar - Hanayama, R. et al. Identification of a factor that links apoptotic cells to phagocytes. Nature 417, 182–187 (2002).
Article CAS PubMed Google Scholar - Miyata, K. et al. Elevated mature macrophage expression of human ABHD2 gene in vulnerable plaque. Biochem. Biophys. Res. Commun. 365, 207–213 (2008).
Article CAS PubMed Google Scholar - Jain, M., Bhat, G.P., Vijayraghavan, K. & Inamdar, M.S. Rudhira/BCAS3 is a cytoskeletal protein that controls Cdc42 activation and directional cell migration during angiogenesis. Exp. Cell Res. 318, 753–767 (2012).
Article CAS PubMed Google Scholar - Kim, J.Y., Ahn, H.J., Ryu, J.H., Suk, K. & Park, J.H. BH3-only protein Noxa is a mediator of hypoxic cell death induced by hypoxia-inducible factor 1α. J. Exp. Med. 199, 113–124 (2004).
Article CAS PubMed PubMed Central Google Scholar - Global Lipids Genetics Consortium. Discovery and refinement of loci associated with lipid levels. Nat. Genet. 45, 1274–1283 (2013).
- Lango Allen, H. et al. Hundreds of variants clustered in genomic loci and biological pathways affect human height. Nature 467, 832–838 (2010).
Article CAS PubMed PubMed Central Google Scholar - Morris, A.P. et al. Large-scale association analysis provides insights into the genetic architecture and pathophysiology of type 2 diabetes. Nat. Genet. 44, 981–990 (2012).
Article CAS PubMed PubMed Central Google Scholar - Scott, R.A. et al. Large-scale association analyses identify new loci influencing glycemic traits and provide insight into the underlying biological pathways. Nat. Genet. 44, 991–1005 (2012).
Article CAS PubMed PubMed Central Google Scholar - Speliotes, E.K. et al. Association analyses of 249,796 individuals reveal 18 new loci associated with body mass index. Nat. Genet. 42, 937–948 (2010).
Article CAS PubMed PubMed Central Google Scholar - Pearce, L.R. et al. KSR2 mutations are associated with obesity, insulin resistance, and impaired cellular fuel oxidation. Cell 155, 765–777 (2013).
Article CAS PubMed PubMed Central Google Scholar - Schork, N.J., Murray, S.S., Frazer, K.A. & Topol, E.J. Common vs. rare allele hypotheses for complex diseases. Curr. Opin. Genet. Dev. 19, 212–219 (2009).
Article CAS PubMed PubMed Central Google Scholar - Lettre, G., Lange, C. & Hirschhorn, J.N. Genetic model testing and statistical power in population-based association studies of quantitative traits. Genet. Epidemiol. 31, 358–362 (2007).
Article PubMed Google Scholar - Do, R. et al. Exome sequencing identifies rare LDLR and APOA5 alleles conferring risk for myocardial infarction. Nature 518, 102–106 (2015).
Article CAS PubMed Google Scholar - TG and HDL Working Group of the Exome Sequencing Project. Loss-of-function mutations in APOC3, triglycerides, and coronary disease. N. Engl. J. Med. 371, 22–31 (2014).
- Myocardial Infarction Genetics Consortium Investigators. Inactivating mutations in NPC1L1 and protection from coronary heart disease. N. Engl. J. Med. 371, 2072–2082 (2014).
- Maurano, M.T. et al. Systematic localization of common disease-associated variation in regulatory DNA. Science 337, 1190–1195 (2012).
CAS PubMed PubMed Central Google Scholar - Libby, P., Ridker, P.M. & Hansson, G.K. Progress and challenges in translating the biology of atherosclerosis. Nature 473, 317–325 (2011).
Article CAS PubMed Google Scholar - Reilly, M.P. et al. Identification of ADAMTS7 as a novel locus for coronary atherosclerosis and association of ABO with myocardial infarction in the presence of coronary atherosclerosis: two genome-wide association studies. Lancet 377, 383–392 (2011).
Article CAS PubMed PubMed Central Google Scholar - Dichgans, M. et al. Shared genetic susceptibility to ischemic stroke and coronary artery disease: a genome-wide analysis of common variants. Stroke 45, 24–36 (2014).
Article CAS PubMed Google Scholar - Keating, B.J. et al. Concept, design and implementation of a cardiovascular gene-centric 50 k SNP array for large-scale genomic association studies. PLoS ONE 3, e3583 (2008).
Article CAS PubMed PubMed Central Google Scholar - Voight, B.F. et al. The Metabochip, a custom genotyping array for genetic studies of metabolic, cardiovascular, and anthropometric traits. PLoS Genet. 8, e1002793 (2012).
Article CAS PubMed PubMed Central Google Scholar - Schunkert, H. et al. Large-scale association analysis identifies 13 new susceptibility loci for coronary artery disease. Nat. Genet. 43, 333–338 (2011).
Article CAS PubMed PubMed Central Google Scholar - Miyata, K. et al. Increase of smooth muscle cell migration and of intimal hyperplasia in mice lacking the α/β hydrolase domain containing 2 gene. Biochem. Biophys. Res. Commun. 329, 296–304 (2005).
Article CAS PubMed Google Scholar - Bobik, A. Transforming growth factor-βs and vascular disorders. Arterioscler. Thromb. Vasc. Biol. 26, 1712–1720 (2006).
Article CAS PubMed Google Scholar - Mallat, Z. et al. Inhibition of transforming growth factor-β signaling accelerates atherosclerosis and induces an unstable plaque phenotype in mice. Circ. Res. 89, 930–934 (2001).
Article CAS PubMed Google Scholar - Yang, Z. et al. Infarct-sparing effect of A2A-adenosine receptor activation is due primarily to its action on lymphocytes. Circulation 111, 2190–2197 (2005).
Article CAS PubMed Google Scholar - Aziz, M., Jacob, A., Matsuda, A. & Wang, P. Review: milk fat globule–EGF factor 8 expression, function and plausible signal transduction in resolving inflammation. Apoptosis 16, 1077–1086 (2011).
Article CAS PubMed Google Scholar - Yang, J. et al. Genomic inflation factors under polygenic inheritance. Eur. J. Hum. Genet. 19, 807–812 (2011).
Article PubMed PubMed Central Google Scholar - Howie, B., Fuchsberger, C., Stephens, M., Marchini, J. & Abecasis, G.R. Fast and accurate genotype imputation in genome-wide association studies through pre-phasing. Nat. Genet. 44, 955–959 (2012).
Article CAS PubMed PubMed Central Google Scholar - Devlin, B. & Roeder, K. Genomic control for association studies. Biometrics 55, 997–1004 (1999).
Article CAS PubMed Google Scholar - Mägi, R. & Morris, A.P. GWAMA: software for genome-wide association meta-analysis. BMC Bioinformatics 11, 288 (2010).
Article PubMed PubMed Central Google Scholar - Cochran, W.G. The combination of estimates from different experiments. Biometrics 10, 101–129 (1954).
Article Google Scholar - Higgins, J.P. & Thompson, S.G. Quantifying heterogeneity in a meta-analysis. Stat. Med. 21, 1539–1558 (2002).
Article PubMed Google Scholar - DerSimonian, R. & Laird, N. Meta-analysis in clinical trials. Control. Clin. Trials 7, 177–188 (1986).
Article CAS PubMed Google Scholar - Newson, R.B. Frequentist _q_-values for multiple-test procedures. Stata J. 10, 568–584 (2010).
Article Google Scholar - Benjamini, Y. & Yekutieli, D. The control of the false-discovery rate in multiple testing under dependency. Ann. Stat. 29, 1165–1188 (2001).
Article Google Scholar - Yang, J. et al. Conditional and joint multiple-SNP analysis of GWAS summary statistics identifies additional variants influencing complex traits. Nat. Genet. 44, 369–375 (2012).
Article CAS PubMed PubMed Central Google Scholar - So, H.C., Gui, A.H., Cherny, S.S. & Sham, P.C. Evaluating the heritability explained by known susceptibility variants: a survey of ten complex diseases. Genet. Epidemiol. 35, 310–317 (2011).
Article PubMed Google Scholar - Heid, I.M. et al. Meta-analysis identifies 13 new loci associated with waist-hip ratio and reveals sexual dimorphism in the genetic basis of fat distribution. Nat. Genet. 42, 949–960 (2010).
Article CAS PubMed PubMed Central Google Scholar - International Consortium for Blood Pressure Genome-Wide Association Studies. Genetic variants in novel pathways influence blood pressure and cardiovascular disease risk. Nature 478, 103–109 (2011).
- Wain, L.V. et al. Genome-wide association study identifies six new loci influencing pulse pressure and mean arterial pressure. Nat. Genet. 43, 1005–1011 (2011).
Article CAS PubMed PubMed Central Google Scholar - Welter, D. et al. The NHGRI GWAS Catalog, a curated resource of SNP-trait associations. Nucleic Acids Res. 42, D1001–D1006 (2014).
Article CAS PubMed Google Scholar - Fehrmann, R.S. et al. _Trans_-eQTLs reveal that independent genetic variants associated with a complex phenotype converge on intermediate genes, with a major role for the HLA. PLoS Genet. 7, e1002197 (2011).
Article CAS PubMed PubMed Central Google Scholar - Garnier, S. et al. Genome-wide haplotype analysis of cis expression quantitative trait loci in monocytes. PLoS Genet. 9, e1003240 (2013).
Article CAS PubMed PubMed Central Google Scholar - Gibbs, J.R. et al. Abundant quantitative trait loci exist for DNA methylation and gene expression in human brain. PLoS Genet. 6, e1000952 (2010).
Article CAS PubMed PubMed Central Google Scholar - Liang, L. et al. A cross-platform analysis of 14,177 expression quantitative trait loci derived from lymphoblastoid cell lines. Genome Res. 23, 716–726 (2013).
Article CAS PubMed PubMed Central Google Scholar - Westra, H.J. et al. Systematic identification of trans eQTLs as putative drivers of known disease associations. Nat. Genet. 45, 1238–1243 (2013).
Article CAS PubMed PubMed Central Google Scholar - Busch, S.J., Barnhart, R.L., Martin, G.A., Flanagan, M.A. & Jackson, R.L. Differential regulation of hepatic triglyceride lipase and 3-hydroxy-3-methylglutaryl-CoA reductase gene expression in a human hepatoma cell line, HepG2. J. Biol. Chem. 265, 22474–22479 (1990).
CAS PubMed Google Scholar - Park, H.J. et al. Human umbilical vein endothelial cells and human dermal microvascular endothelial cells offer new insights into the relationship between lipid metabolism and angiogenesis. Stem Cell Rev. 2, 93–102 (2006).
Article CAS PubMed Google Scholar - Durrani, S., Konoplyannikov, M., Ashraf, M. & Haider, K.H. Skeletal myoblasts for cardiac repair. Regen. Med. 5, 919–932 (2010).
Article PubMed Google Scholar - ENCODE Project Consortium. An integrated encyclopedia of DNA elements in the human genome. Nature 489, 57–74 (2012).
Acknowledgements
We sincerely thank the participants and the medical, nursing, technical and administrative staff in each of the studies who have contributed to this project. We are grateful for support from our funders; more detailed acknowledgments are included in the Supplementary Note.
Author information
Author notes
- Majid Nikpay, Anuj Goel, Hong-Hee Won, Leanne M Hall, Christina Willenborg, Stavroula Kanoni and Danish Saleheen: These authors contributed equally to this work.
- Hugh Watkins, Sekar Kathiresan, Ruth McPherson, Panos Deloukas, Heribert Schunkert, Nilesh J Samani and Martin Farrall: These authors jointly supervised this work.
Authors and Affiliations
- Ruddy Canadian Cardiovascular Genetics Centre, University of Ottawa Heart Institute, Ottawa, Ontario, Canada
Majid Nikpay, Alexandre F Stewart & Ruth McPherson - Division of Cardiovascular Medicine, Radcliffe Department of Medicine, University of Oxford, Oxford, UK
Anuj Goel, Theodosios Kyriakou, Christopher Grace, Shapour Jalilzadeh, Hugh Watkins & Martin Farrall - Wellcome Trust Centre for Human Genetics, University of Oxford, Oxford, UK
Anuj Goel, Theodosios Kyriakou, Christopher Grace, Natalie R van Zuydam, Shapour Jalilzadeh, Cecilia M Lindgren, Andrew P Morris, Erik Ingelsson, Hugh Watkins & Martin Farrall - Broad Institute of MIT and Harvard University, Cambridge, Massachusetts, USA
Hong-Hee Won, Andrew Bjonnes, Richa Saxena, Cecilia M Lindgren, Tõnu Esko & Sekar Kathiresan - Cardiovascular Research Center, Massachusetts General Hospital, Boston, Massachusetts, USA
Hong-Hee Won & Sekar Kathiresan - Center for Human Genetic Research, Massachusetts General Hospital, Boston, Massachusetts, USA
Hong-Hee Won, Andrew Bjonnes, Richa Saxena & Sekar Kathiresan - Department of Medicine, Harvard Medical School, Boston, Massachusetts, USA
Hong-Hee Won & Sekar Kathiresan - Department of Cardiovascular Sciences, University of Leicester, Leicester, UK
Leanne M Hall, Christopher P Nelson, Thomas R Webb & Nilesh J Samani - Institut für Integrative und Experimentelle Genomik, Universität zu Lübeck, Lübeck, Germany
Christina Willenborg & Jeanette Erdmann - DZHK (German Research Center for Cardiovascular Research), partner site Hamburg-Lübeck-Kiel, Lübeck, Germany
Christina Willenborg, Inke R König & Jeanette Erdmann - William Harvey Research Institute, Barts and the London School of Medicine and Dentistry, Queen Mary University of London, London, UK
Stavroula Kanoni, Kathleen E Stirrups & Panos Deloukas - Perelman School of Medicine, University of Pennsylvania, Philadelphia, Pennsylvania, USA
Danish Saleheen & Wei Zhao - Center for Non-Communicable Diseases, Karachi, Pakistan
Danish Saleheen, Philippe Frossard, Asif Rasheed & Maria Samuel - National Institute for Health Research (NIHR) Leicester Cardiovascular Biomedical Research Unit, Glenfield Hospital, Leicester, UK
Christopher P Nelson, Thomas R Webb, Alison H Goodall & Nilesh J Samani - Nuffield Department of Population Health, Clinical Trial Service Unit and Epidemiological Studies Unit (CTSU), University of Oxford, Oxford, UK
Jemma C Hopewell, King Wai Lau, Rory Collins & Robert Clarke - Deutsches Herzzentrum München, Technische Universität München, Munich, Germany
Lingyao Zeng, Thorsten Kessler, Christian Hengstenberg & Heribert Schunkert - DZHK (German Centre for Cardiovascular Research), partner site Munich Heart Alliance, Munich, Germany
Lingyao Zeng, Christian Gieger, Thomas Meitinger, Annette Peters, Christian Hengstenberg & Heribert Schunkert - Department of Epidemiology, Erasmus University Medical Center, Rotterdam, the Netherlands
Abbas Dehghan, Andre Uitterlinden, Paul S de Vries, Oscar H Franco & Albert Hofman - Estonian Genome Center, University of Tartu, Tartu, Estonia
Maris Alver, Evelin Mihailov, Natalia Pervjakova, Tõnu Esko, Andres Metspalu & Markus Perola - Institute of Molecular and Cell Biology, University of Tartu, Tartu, Estonia
Maris Alver & Andres Metspalu - Division of Biomedical Statistics and Informatics, Department of Health Sciences Research, Mayo Clinic, Rochester, Minnesota, USA
Sebastian M Armasu & Mariza de Andrade - Department of Health, National Institute for Health and Welfare, Helsinki, Finland
Kirsi Auro, Natalia Pervjakova & Markus Perola - Institute for Molecular Medicine Finland (FIMM), University of Helsinki, Helsinki, Finland
Kirsi Auro, Natalia Pervjakova, Emmi Tikkanen, Markus Perola & Samuli Ripatti - Diabetes and Obesity Research Program, University of Helsinki, Helsinki, Finland
Kirsi Auro, Natalia Pervjakova & Markus Perola - Division of Preventive Medicine, Brigham and Women's Hospital, Boston, Massachusetts, USA
Daniel I Chasman, Lynda M Rose, Julie E Buring & Paul M Ridker - Harvard Medical School, Boston, Massachusetts, USA
Daniel I Chasman & Paul M Ridker - State Key Laboratory of Cardiovascular Disease, Fuwai Hospital, National Center of Cardiovascular Diseases, Chinese Academy of Medical Sciences and Peking Union Medical College, Beijing, China
Shufeng Chen, Xiangfeng Lu, Xueli Yang, Laiyuan Wang & Dongfeng Gu - Robertson Center for Biostatistics, University of Glasgow, Glasgow, UK
Ian Ford - Department of Epidemiology, Gillings School of Global Public Health, University of North Carolina, Chapel Hill, North Carolina, USA
Nora Franceschini - Institute of Epidemiology II, Helmholtz Zentrum München–German Research Center for Environmental Health, Neuherberg, Germany
Christian Gieger & Annette Peters - Research Unit of Molecular Epidemiology, Helmholtz Zentrum München–German Research Center for Environmental Health, Neuherberg, Germany
Christian Gieger - Department of Medical Sciences, Molecular Epidemiology, Uppsala University, Uppsala, Sweden
Stefan Gustafsson & Erik Ingelsson - Science for Life Laboratory, Uppsala University, Uppsala, Sweden
Stefan Gustafsson & Erik Ingelsson - Wellcome Trust Sanger Institute, Hinxton, UK
Jie Huang, John Danesh & Samuli Ripatti - National Heart, Lung, and Blood Institute's Framingham Heart Study, Framingham, Massachusetts, USA
Shih-Jen Hwang, L Adrienne Cupples & Christopher J O'Donnell - Department of Biostatistics, Boston University School of Public Health, Boston, Massachusetts, USA
Shih-Jen Hwang & L Adrienne Cupples - Center for Genome Science, Korea National Institute of Health, Chungcheongbuk-do, Korea
Yun Kyoung Kim, Bok-Ghee Han & Bong-Jo Kim - Vth Department of Medicine (Nephrology, Hypertensiology, Endocrinology, Diabetology, Rheumatology), Medical Faculty of Mannheim, University of Heidelberg, Mannheim, Germany
Marcus E Kleber & Winfried März - Charles Bronfman Institute for Personalized Medicine, Icahn School of Medicine at Mount Sinai, New York, New York, USA
Yingchang Lu, Omri Gottesman, Erwin P Bottinger & Ruth J F Loos - Genetics of Obesity and Related Metabolic Traits Program, Icahn School of Medicine at Mount Sinai, New York, New York, USA
Yingchang Lu & Ruth J F Loos - Department of Clinical Chemistry, Fimlab Laboratories, Tampere, Finland
Leo-Pekka Lyytikäinen, Pekka J Karhunen & Terho Lehtimäki - Department of Clinical Chemistry, University of Tampere School of Medicine, Tampere, Finland
Leo-Pekka Lyytikäinen & Terho Lehtimäki - Human Genetics Center, School of Public Health, University of Texas Health Science Center at Houston, Houston, Texas, USA
Alanna C Morrison & Eric Boerwinkle - Department of Biostatistics and Epidemiology, University of Pennsylvania, Philadelphia, Pennsylvania, USA
Liming Qu - Division of Cardiovascular Medicine, Department of Medicine, Stanford University, Stanford, California, USA
Elias Salfati, Erik Ingelsson, Thomas Quertermous & Themistocles L Assimes - Department of Anesthesia, Critical Care and Pain Medicine, Massachusetts General Hospital, Harvard Medical School, Boston, Massachusetts, USA
Richa Saxena - Institute for Medical Informatics, Statistics and Epidemiology, Medical Faculty, University of Leipzig, Leipzig, Germany
Markus Scholz - LIFE Research Center of Civilization Diseases, Leipzig, Germany
Markus Scholz, Frank Beutner & Joachim Thiery - Icelandic Heart Association, Kopavogur, Iceland
Albert V Smith & Vilmundur Gudnason - Faculty of Medicine, University of Iceland, Reykjavik, Iceland
Albert V Smith & Vilmundur Gudnason - Department of Public Health, University of Helsinki, Helsinki, Finland
Emmi Tikkanen & Samuli Ripatti - Department of Epidemiology and Biostatistics, Imperial College London, London, UK
Weihua Zhang & John C Chambers - Department of Cardiology, Ealing Hospital National Health Service (NHS) Trust, Middlesex, UK
Weihua Zhang, John C Chambers & Jaspal S Kooner - Medical Research Institute, University of Dundee, Dundee, UK
Natalie R van Zuydam & Colin N Palmer - Department of Medicine, Population Health Research Institute, Hamilton Health Sciences, McMaster University, Hamilton, Ontario, Canada
Sonia S Anand - Platform for Genome Analytics, Institutes of Neurogenetics and Integrative and Experimental Genomics, University of Lübeck, Lübeck, Germany
Lars Bertram - Neuroepidemiology and Ageing Research Unit, School of Public Health, Faculty of Medicine, Imperial College of Science, Technology and Medicine, London, UK
Lars Bertram - Heart Center Leipzig, Cardiology, University of Leipzig, Leipzig, Germany
Frank Beutner - Department of Dietetics-Nutrition, Harokopio University, Athens, Greece
George Dedoussis - INSERM, UMRS 1138, Centre de Recherche des Cordeliers, Paris, France
Dominique Gauguier - Department of Cardiovascular Sciences, University of Leicester, Glenfield Hospital, Leicester, UK
Alison H Goodall - School of Medicine, Lebanese American University, Beirut, Lebanon
Marc Haber & Pierre A Zalloua - Hypertension Division, Fuwai Hospital, National Center for Cardiovascular Diseases, Chinese Academy of Medical Sciences and Peking Union Medical College, Beijing, China
Jianfeng Huang - Klinikum Rechts der Isar, Munich, Germany
Thorsten Kessler - Institut für Medizinische Biometrie und Statistik, Universität zu Lübeck, Lübeck, Germany
Inke R König - Department of Public Health and Caring Sciences, Geriatrics, Uppsala University, Uppsala, Sweden
Lars Lannfelt - Institut für Epidemiologie, Christian Albrechts Universität zu Kiel, Kiel, Germany
Wolfgang Lieb - Department of Medical Sciences, Cardiovascular Epidemiology, Uppsala University, Uppsala, Sweden
Lars Lind - Transplantation Laboratory, Haartman Institute, University of Helsinki, Helsinki, Finland
Marja-Liisa Lokki - Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden
Patrik K Magnusson & Nancy L Pedersen - Punjab Institute of Cardiology, Lahore, Pakistan
Nadeem H Mallick - All India Institute of Medical Sciences, New Delhi, India
Narinder Mehra - Institut für Humangenetik, Helmholtz Zentrum München–German Research Center for Environmental Health, Neuherberg, Germany
Thomas Meitinger - Institute of Human Genetics, Technische Universität München, Munich, Germany
Thomas Meitinger - Red Crescent Institute of Cardiology, Hyderabad, Pakistan
Fazal-ur-Rehman Memon & Asif Rasheed - Department of Biostatistics, University of Liverpool, Liverpool, UK
Andrew P Morris - Department of Cardiology, Department of Medicine, Helsinki University Central Hospital, Helsinki, Finland
Markku S Nieminen & Juha Sinisalo - Second Department of Cardiology, Attikon Hospital, School of Medicine, University of Athens, Athens, Greece
Loukianos S Rallidis - Department of Medicine, Duke University Medical Center, Durham, North Carolina, USA
Svati H Shah & Christopher B Granger - Department of Haematology, University of Cambridge, Cambridge, UK
Kathleen E Stirrups - Department of Cardiology, Leiden University Medical Center, Leiden, the Netherlands
Stella Trompet & J Wouter Jukema - Department of Gerontology and Geriatrics, Leiden University Medical Center, Leiden, the Netherlands
Stella Trompet - National Human Genome Center at Beijing, Beijing, China
Laiyuan Wang - National Institute of Cardiovascular Diseases, Karachi, Pakistan
Khan S Zaman - Division of Cardiology, Azienda Ospedaliero Universitaria di Parma, Parma, Italy
Diego Ardissino - Associazione per lo Studio della Trombosi in Cardiologia, Pavia, Italy
Diego Ardissino - Human Genome Sequencing Center, Baylor College of Medicine, Houston, Texas, USA
Eric Boerwinkle - Department of Genetics, Washington University School of Medicine, St. Louis, Missouri, USA
Ingrid B Borecki & Mary F Feitosa - Imperial College Healthcare NHS Trust, London, UK
John C Chambers & Jaspal S Kooner - Department of Public Health and Primary Care, University of Cambridge, Cambridge, UK
John Danesh - Berlin Aging Study II,
Ilja Demuth - Research Group on Geriatrics, Charité Universitätsmedizin Berlin, Berlin, Germany
Ilja Demuth - Institute of Medical and Human Genetics, Charité Universitätsmedizin Berlin, Berlin, Germany
Ilja Demuth - Grupo de Epidemiología y Genética Cardiovascular, Institut Hospital del Mar d'Investigacions Mèdiques (IMIM), Barcelona, Spain
Roberto Elosua - MedStar Heart and Vascular Institute, MedStar Washington Hospital Center, Washington, DC, USA
Stephen E Epstein - Division of Endocrinology and Basic and Translational Obesity Research, Boston Children's Hospital, Boston, Massachusetts, USA
Tõnu Esko - Department of Genetics, Harvard Medical School, Boston, Massachusetts, USA
Tõnu Esko - Department of Cardiovascular Research, Istituto di Ricerca e Cura a Carattere Scientifico (IRCCS) Istituto di Ricerche Farmacologiche Mario Negri, Milan, Italy
Maria Grazia Franzosi - Leeds Institute of Genetics, Health and Therapeutics, University of Leeds, Leeds, UK
Alistair S Hall - Department of Medicine Solna, Cardiovascular Genetics and Genomics Group, Atherosclerosis Research Unit, Karolinska Institutet, Stockholm, Sweden
Anders Hamsten - Laboratory of Epidemiology, Demography and Biometry, National Institute on Aging, US National Institutes of Health, Bethesda, Maryland, USA
Tamara B Harris - Department of Cellular and Molecular Medicine, Cleveland Clinic, Cleveland, Ohio, USA
Stanley L Hazen - Kaiser Permanente Division of Research, Oakland, California, USA
Carlos Iribarren - Durrer Center for Cardiogenetic Research, Amsterdam, the Netherlands
J Wouter Jukema - Interuniversity Cardiology Institute of the Netherlands, Utrecht, the Netherlands
J Wouter Jukema - Department of Forensic Medicine, University of Tampere School of Medicine, Tampere, Finland
Pekka J Karhunen - Cardiovascular Science, National Heart and Lung Institute, Imperial College London, London, UK
Jaspal S Kooner - Division of Cardiovascular Diseases, Department of Medicine, Mayo Clinic, Rochester, Minnesota, USA
Iftikhar J Kullo - Mindich Child Health and Development Institute, Icahn School of Medicine at Mount Sinai, New York, New York, USA
Ruth J F Loos - Department of Clinical Sciences, Hypertension and Cardiovascular Disease, Lund University, University Hospital Malmö, Malmö, Sweden
Olle Melander - Synlab Academy, Synlab Services, Mannheim, Germany
Winfried März - Clinical Institute of Medical and Chemical Laboratory Diagnostics, Medical University of Graz, Graz, Austria
Winfried März - Stanford Cardiovascular Institute, Stanford University, Stanford, California, USA
Thomas Quertermous & Themistocles L Assimes - Department of Genetics, Perelman School of Medicine at the University of Pennsylvania, Philadelphia, Pennsylvania, USA
Daniel J Rader - Cardiovascular Institute, Perelman School of Medicine at the University of Pennsylvania, Philadelphia, Pennsylvania, USA
Daniel J Rader & Muredach P Reilly - University of Ottawa Heart Institute, Ottawa, Ontario, Canada
Robert Roberts - Department of Chronic Disease Prevention, National Institute for Health and Welfare, Helsinki, Finland
Veikko Salomaa - Department of Pediatrics, College of Medicine, University of Oklahoma Health Sciences Center, Oklahoma City, Oklahoma, USA
Dharambir K Sanghera - Department of Pharmaceutical Sciences, College of Pharmacy, University of Oklahoma Health Sciences Center, Oklahoma City, Oklahoma, USA
Dharambir K Sanghera - Oklahoma Center for Neuroscience, Oklahoma City, Oklahoma, USA
Dharambir K Sanghera - Public Health Sciences Division, Fred Hutchinson Cancer Research Center, Seattle, Washington, USA
Stephen M Schwartz - Department of Epidemiology, University of Washington, Seattle, Washington, USA
Stephen M Schwartz - Department of Prosthetic Dentistry, Center for Dental and Oral Medicine, University Medical Center Hamburg-Eppendorf, Hamburg, Germany
Udo Seedorf - Institute of Cardiovascular and Medical Sciences, Faculty of Medicine, University of Glasgow, Glasgow, UK
David J Stott - Institute for Laboratory Medicine, Clinical Chemistry and Molecular Diagnostics, University Hospital Leipzig, Medical Faculty, Leipzig, Germany
Joachim Thiery - Harvard School of Public Health, Boston, Massachusetts, USA
Pierre A Zalloua - and Blood Institute Division of Intramural Research, National Heart, Lung, Bethesda, Maryland, USA
Christopher J O'Donnell - Cardiology Division, Massachusetts General Hospital, Boston, Massachusetts, USA
Christopher J O'Donnell - Department of Health Sciences, University of Leicester, Leicester, UK
John R Thompson - Princess Al-Jawhara Al-Brahim Centre of Excellence in Research of Hereditary Disorders (PACER-HD), King Abdulaziz University, Jeddah, Saudi Arabia
Panos Deloukas
Consortia
the CARDIoGRAMplusC4D Consortium
- Majid Nikpay
- , Anuj Goel
- , Hong-Hee Won
- , Leanne M Hall
- , Christina Willenborg
- , Stavroula Kanoni
- , Danish Saleheen
- , Theodosios Kyriakou
- , Christopher P Nelson
- , Jemma C Hopewell
- , Thomas R Webb
- , Lingyao Zeng
- , Abbas Dehghan
- , Maris Alver
- , Sebastian M Armasu
- , Kirsi Auro
- , Andrew Bjonnes
- , Daniel I Chasman
- , Shufeng Chen
- , Ian Ford
- , Nora Franceschini
- , Christian Gieger
- , Christopher Grace
- , Stefan Gustafsson
- , Jie Huang
- , Shih-Jen Hwang
- , Yun Kyoung Kim
- , Marcus E Kleber
- , King Wai Lau
- , Xiangfeng Lu
- , Yingchang Lu
- , Leo-Pekka Lyytikäinen
- , Evelin Mihailov
- , Alanna C Morrison
- , Natalia Pervjakova
- , Liming Qu
- , Lynda M Rose
- , Elias Salfati
- , Richa Saxena
- , Markus Scholz
- , Albert V Smith
- , Emmi Tikkanen
- , Andre Uitterlinden
- , Xueli Yang
- , Weihua Zhang
- , Wei Zhao
- , Mariza de Andrade
- , Paul S de Vries
- , Natalie R van Zuydam
- , Sonia S Anand
- , Lars Bertram
- , Frank Beutner
- , George Dedoussis
- , Philippe Frossard
- , Dominique Gauguier
- , Alison H Goodall
- , Omri Gottesman
- , Marc Haber
- , Bok-Ghee Han
- , Jianfeng Huang
- , Shapour Jalilzadeh
- , Thorsten Kessler
- , Inke R König
- , Lars Lannfelt
- , Wolfgang Lieb
- , Lars Lind
- , Cecilia M Lindgren
- , Marja-Liisa Lokki
- , Patrik K Magnusson
- , Nadeem H Mallick
- , Narinder Mehra
- , Thomas Meitinger
- , Fazal-ur-Rehman Memon
- , Andrew P Morris
- , Markku S Nieminen
- , Nancy L Pedersen
- , Annette Peters
- , Loukianos S Rallidis
- , Asif Rasheed
- , Maria Samuel
- , Svati H Shah
- , Juha Sinisalo
- , Kathleen E Stirrups
- , Stella Trompet
- , Laiyuan Wang
- , Khan S Zaman
- , Diego Ardissino
- , Eric Boerwinkle
- , Ingrid B Borecki
- , Erwin P Bottinger
- , Julie E Buring
- , John C Chambers
- , Rory Collins
- , L Adrienne Cupples
- , John Danesh
- , Ilja Demuth
- , Roberto Elosua
- , Stephen E Epstein
- , Tõnu Esko
- , Mary F Feitosa
- , Oscar H Franco
- , Maria Grazia Franzosi
- , Christopher B Granger
- , Dongfeng Gu
- , Vilmundur Gudnason
- , Alistair S Hall
- , Anders Hamsten
- , Tamara B Harris
- , Stanley L Hazen
- , Christian Hengstenberg
- , Albert Hofman
- , Erik Ingelsson
- , Carlos Iribarren
- , J Wouter Jukema
- , Pekka J Karhunen
- , Bong-Jo Kim
- , Jaspal S Kooner
- , Iftikhar J Kullo
- , Terho Lehtimäki
- , Ruth J F Loos
- , Olle Melander
- , Andres Metspalu
- , Winfried März
- , Colin N Palmer
- , Markus Perola
- , Thomas Quertermous
- , Daniel J Rader
- , Paul M Ridker
- , Samuli Ripatti
- , Robert Roberts
- , Veikko Salomaa
- , Dharambir K Sanghera
- , Stephen M Schwartz
- , Udo Seedorf
- , Alexandre F Stewart
- , David J Stott
- , Joachim Thiery
- , Pierre A Zalloua
- , Christopher J O'Donnell
- , Muredach P Reilly
- , Themistocles L Assimes
- , John R Thompson
- , Jeanette Erdmann
- , Robert Clarke
- , Hugh Watkins
- , Sekar Kathiresan
- , Ruth McPherson
- , Panos Deloukas
- , Heribert Schunkert
- , Nilesh J Samani
- & Martin Farrall
Contributions
Cohort oversight: D.A., E.B., I.B.B., E.P.B., J.E.B., J.C.C., R. Collins, L.A.C., J.D., I.D., R.E., S.E.E., T.E., M.F.F., O.H.F., M.G.F., C.B.G., D. Gu, V.G., A.S.H., A. Hamsten, T.B.H., S.L.H., C.H., A. Hofman, E.I., C.I., J.W.J., P.J.K., B.-J.K., J.S.K., I.J.K., T.L., R.J.F.L., O.M., A.M., W.M., C.N.P., M.P., T.Q., D.J.R., P.M.R., S.R., R.R., V.S., D.K.S., S.M.S., U.S., A.F.S., D.J.S., J.T., P.A.Z., C.J.O'D., M.P.R., T.L.A., J.R.T., J.E., H.W., S. Kathiresan, R.M., P.D., H.S., N.J.S. and M.F. Cohort genotyping: H.-H.W., S. Kanoni, D.S., J.C.H., Jie Huang, M.E.K., Y.L., L.-P.L., A.U., S.S.A., L.B., G.D., D. Gauguier, A.H.G., M.H., B.-G.H., S.J., L. Lind, C.M.L., M.-L.L., P.K.M., A.P.M., M.S.N., N.L.P., J.S., K.E.S., S.T., L.W., I.B.B., J.C.C., R. Collins, M.F.F., A. Hofman, E.I., J.S.K., T.L., R.R., D.K.S., A.F.S., R. Clarke, P.D. and N.J.S. Cohort phenotyping: D.S., J.C.H., A.D., M.A., K.A., Y.K.K., E.M., L.M.R., S.S.A., F.B., G.D., P.F., A.H.G., O.G., Jianfeng Huang, T. Kessler, I.R.K., L. Lannfelt, W.L., L. Lind, C.M.L., P.K.M., N.H.M., N.M., T.M., F.-ur-R.M., A.P.M., N.L.P., A.P., L.S.R., A.R., M. Samuel, S.H.S., K.S.Z., D.A., J.E.B., J.C.C., R. Collins, R.E., C.B.G., V.G., A.S.H., A. Hamsten, S.L.H., E.I., J.W.J., P.J.K., J.S.K., I.J.K., O.M., A.M., M.P., R.R., D.K.S., A.F.S., D.J.S., P.A.Z., M.P.R., R. Clarke, S. Kathiresan, H.S. and N.J.S. Cohort data analyst: M.N., A.G., H.-H.W., L.M.H., C.W., S. Kanoni, D.S., T. Kyriakou, C.P.N., J.C.H., T.R.W., L.Z., A.D., M.A., S.M.A., K.A., A.B., D.I.C., S.C., I.F., N.F., C. Gieger, C. Grace, S.G., Jie Huang, S.-J.H., Y.K.K., M.E.K., K.W.L., X.L., Y.L., L.-P.L., E.M., A.C.M., N.P., L.Q., L.M.R., E.S., R.S., M. Scholz, A.V.S., E.T., A.U., X.Y., W. Zhang, W. Zhao, M.d.A., P.S.d.V., N.R.v.Z., M.F.F., J.R.T. and M.F. Meta-analysis: M.N., A.G., H.-H.W., L.M.H., C.P.N., J.R.T. and M.F. Variant annotation: M.N., A.G., H.-H.W., T. Kyriakou, J.C.H. and T.R.W. Manuscript drafting: M.N., A.G., H.-H.W., L.M.H., T. Kyriakou, J.C.H., H.W., S. Kathiresan, R.M., H.S., N.J.S. and M.F. Project steering committee: M.N., A.G., H.-H.W., L.M.H., S. Kanoni., J.C.H., D.I.C., M.E.K., N.R.v.Z., C.N.P., R.R., C.J.O'D., M.P.R., T.L.A., J.R.T., J.E., R. Clarke, H.W., S. Kathiresan, R.M., P.D., H.S., N.J.S. and M.F. (secretariat: J.C.H. and R. Clarke). CARDIoGRAMplusC4D executive committee: J.D., D. Gu, A. Hamsten, J.S.K., R.R., H.W., S. Kathiresan, P.D., H.S. and N.J.S.
Corresponding authors
Correspondence toHugh Watkins, Sekar Kathiresan, Ruth McPherson or Martin Farrall.
Ethics declarations
Competing interests
The author declare no competing financial interests.
Integrated supplementary information
Supplementary Figure 1 Venn diagram showing case-control overlap between 1000G GWAS and Metabochip studies.
Venn diagram showing the number of cases (top) and controls (bottom) that overlap between the present 1000G GWAS meta-analysis study and the Metabochip study (Nat. Genet. 45, 25–33, 2013). There is a 57.5% overlap of our cases and a 40.1% overlap of our controls with the previously published study.
Supplementary Figure 2 A Manhattan plot summarizing the 1000 Genomes CAD additive association results.
The meta-analysis statistics have been adjusted for overdispersion (genomic control parameter = 1.18) and have been capped to P = 1 × 10−20. The genome-wide significance threshold is shown as a horizontal blue line at P < 5 × 10−8. Novel CAD loci are presented with red stacks and gene names (Table 1). Previously reported loci showing genome-wide significance are shown in brown, and those showing nominal significance (P < 0.05) in our meta-analysis are shown in blue (Supplementary Table 2).
Supplementary Figure 3 Comparing effect sizes for the MI subphenotype and the inclusive CAD phenotype.
Point estimates of effect sizes (odds ratios) are shown by open circles, with 95% confidence intervals represented by solid lines. The line of identity is shown as a dashed line. Loci showing marked differences in effect sizes are shown in blue.
Supplementary Figure 4 Heat map of the number of variants with power >90% to detect genome-wide significant association.
Heat map summarizing the number of variants that were calculated to be powered at ≥90% in the meta-analysis to detect a genome-wide significant association with an additive susceptibility variant with odds ratio (OR) = 1.3. Each cell is shaded from white to black to represent larger and smaller numbers of variants, respectively. The modal cell covers variants in the sector with 0.05 < MAF < 0.075 and imputation quality >0.95.
Supplementary Figure 5 Allele frequency analysis to identify strand flipping and data formatting issues.
Allele frequency analysis to identify systematic allele mismatching in individual studies due to strand flipping and other data formatting issues. (a) Proportion of variants that align with the 1000 Genomes phase 1 v3 training set minor allele after assignment to bins on the basis of MAF. The blue plot shows a typical analysis for studies with well-matched alleles, such that there is 100% concordance for lower-frequency alleles (MAF < 0.2) that declines to 50% for more frequent alleles. The red trace for study TH (Supplementary Table 1 of studies with study code) shows a marked discordance in allele frequencies that was resolved before inclusion in the meta-analysis. (b) Surface plot of 28 studies with 2 studies showing systematic strand flipping and further studies showing more subtly different allele frequency patterns. (c) Allele frequency analysis for data submitted to the meta-analysis (i.e., after any systematic mismatching issues had been resolved). Six studies of East and South Asian, Hispanic and African-American ancestries show MAF distortions that contrast with those of the remaining 42 European-ancestry studies.
Supplementary Figure 6 Quantile-quantile plots of the double–genomic controlled CAD meta-analysis results.
Shaded areas represent 95% confidence intervals. (a,b) Plots showing all additive and recessive results. (c,d) Plots showing additive and recessive results after removing variants from known loci. Supplementary Table 15 refers to the genomic control correction of each study before the final meta-analysis.
Supplementary Figure 7 Comparison of GCTA joint association analysis with standard multiple–logistic regression analysis in four studies.
We have investigated the accuracy of the GCTA joint association analysis by comparing the approximate GCTA results with a standard multiple–logistic regression analysis in 4 studies (MIGEN, PROCARDIS, OHGS and Interheart) for the 202 FDR variants. The figure shows scatterplots of the regression coefficients (left column), standard errors (center column) and –log10 (P values) (right column) for each variant; for each scatterplot, the x axis shows the standard multiple–logistic regression result, and the y axis shows the corresponding GCTA COJO result. The regression coefficients and standard errors for the majority (95%) of the variants are very accurately approximated as their results lie close to the line of identity (y = x) shown in red. The –log10 (P values) for the two analyses were positively correlated (0.86 < ρ < 0.93).
Supplementary information
Source data
Rights and permissions
About this article
Cite this article
the CARDIoGRAMplusC4D Consortium. A comprehensive 1000 Genomes–based genome-wide association meta-analysis of coronary artery disease.Nat Genet 47, 1121–1130 (2015). https://doi.org/10.1038/ng.3396
- Received: 13 January 2015
- Accepted: 14 August 2015
- Published: 07 September 2015
- Issue Date: October 2015
- DOI: https://doi.org/10.1038/ng.3396