A

Multi-trait analysis characterizes the genetics of thyroid function and identifies causal associations with clinical implications

Nature Communications, vol. 15, pp. 888

Abstract

To date only a fraction of the genetic footprint of thyroid function has been clarified. We report a genome-wide association study meta-analysis of thyroid function in up to 271,040 individuals of European ancestry, including reference range thyrotropin (TSH), free thyroxine (FT4), free and total triiodothyronine (T3), proxies for metabolism (T3/FT4 ratio) as well as dichotomized high and low TSH levels. We revealed 259 independent significant associations for TSH (61% novel), 85 for FT4 (67% novel), and 62 novel signals for the T3 related traits. The loci explained 14.1%, 6.0%, 9.5% and 1.1% of the total variation in TSH, FT4, total T3 and free T3 concentrations, respectively. Genetic correlations indicate that TSH associated loci reflect the thyroid function determined by free T3, whereas the FT4 associations represent the thyroid hormone metabolism. Polygenic risk score and Mendelian randomization analyses showed the effects of genetically determined variation in thyroid function on various clinical outcomes, including cardiovascular risk factors and diseases, autoimmune diseases, and cancer. In conclusion, our results improve the understanding of thyroid hormone physiology and highlight the pleiotropic effects of thyroid function on various diseases.

Authors 98

  1. Erasmus MC · Radboud University Medical Center

    Affiliation as printed

    Academic Center for Thyroid Diseases, Department of Internal Medicine, Erasmus Medical Center, Rotterdam, The Netherlands

    Department of Internal Medicine, Division of Endocrinology, Radboud University Medical Center, Nijmegen, The Netherlands

  2. University of Freiburg

    Affiliation as printed

    Institute of Genetic Epidemiology, Faculty of Medicine and Medical Center - University of Freiburg, Freiburg, Germany

  3. University of Freiburg

    Affiliation as printed

    Institute of Genetic Epidemiology, Faculty of Medicine and Medical Center - University of Freiburg, Freiburg, Germany

  4. University of Utah

    Affiliation as printed

    The University of Utah, Salt Lake City, UT, USA

  5. RIKEN Center for Integrative Medical Sciences · The University of Osaka

    Affiliation as printed

    Department of Statistical Genetics, Osaka University Graduate School of Medicine, Suita, Japan

    Laboratory for Systems Genetics, RIKEN Center for Integrative Medical Sciences, Kanagawa, Japan

  6. Queen Mary University of London · William Harvey Research Institute

    Affiliation as printed

    Digital Environment Research Institute, Queen Mary University of London, London, UK

    William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London, United Kingdom

  7. Radboud University Medical Center

    Affiliation as printed

    Department for Health Evidence, Radboud University Medical Center, Nijmegen, The Netherlands

  8. Queen Mary University of London · William Harvey Research Institute

    Affiliation as printed

    William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London, United Kingdom

  9. University of Copenhagen · Copenhagen University Hospital · Rigshospitalet

    Affiliation as printed

    Department of Biomedical Sciences, University of Copenhagen, Copenhagen, Denmark

    Laboratory for Molecular Cardiology, Department of Cardiology, Copenhagen University Hospital - Rigshospitalet, Copenhagen, Denmark

  10. Novo Nordisk Foundation

    Affiliation as printed

    Department of Obesity and Nutritional Sciences, The Novo Nordisk Foundation, Hellerup, Denmark

  11. Norwegian University of Science and Technology · St Olav's University Hospital

    Affiliation as printed

    Department of Endocrinology, Clinic of Medicine, St. Olavs Hospital, Trondheim University Hospital, Trondheim, Norway

    K.G. Jebsen Center for Genetic Epidemiology, Department of Public Health and Nursing, NTNU, Norwegian University of Science and Technology, Trondheim, Norway

  12. Affiliation as printed

    Geriatric Unit, Azienda Sanitaria Toscana Centro, Florence, Italy

  13. Leiden University Medical Center

    Affiliation as printed

    Department of Biomedical Data Sciences, Section Molecular Epidemiology, Leiden University Medical Center, Leiden, The Netherlands

  14. The University of Western Australia

    Affiliation as printed

    School of Biomedical Sciences, The University of Western Australia, Perth, WA, 6009, Australia

  15. University of Copenhagen · Novo Nordisk Foundation Center for Basic Metabolic Research

    Affiliation as printed

    Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

  16. Western General Hospital · Institute of Genetics and Cancer · MRC Human Genetics Unit · University of Edinburgh

    Affiliation as printed

    MRC Human Genetics Unit, Institute of Genetics and Cancer, University of Edinburgh, Western General Hospital, Edinburgh, United Kingdom

  17. University of Washington

    Affiliation as printed

    Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, WA, USA

  18. Sir Charles Gairdner Hospital

    Affiliation as printed

    Department of Endocrinology and Diabetes, Sir Charles Gairdner Hospital, Nedlands, WA, 6009, Australia

  19. Norwegian University of Science and Technology

    Affiliation as printed

    HUNT Research Centre, Department of Public Health and Nursing, NTNU, Norwegian University of Science and Technology, Levanger, 7600, Norway

    K.G. Jebsen Center for Genetic Epidemiology, Department of Public Health and Nursing, NTNU, Norwegian University of Science and Technology, Trondheim, Norway

  20. Sir Charles Gairdner Hospital

    Affiliation as printed

    Department of Endocrinology and Diabetes, Sir Charles Gairdner Hospital, Nedlands, WA, 6009, Australia

  21. University of Pennsylvania

    Affiliation as printed

    Division of Endocrinology, Diabetes, and Metabolism, University of Pennsylvania, Philadelphia, PA, USA

  22. University of Parma · Ospedale di Parma

    Affiliation as printed

    Azienda Ospedaliero-Universitaria di Parma, Parma, Italy

    Oncological Endocrinology, University of Parma, Parma, Italy

  23. Harvard University · Erasmus MC

    Affiliation as printed

    Department of Epidemiology, Erasmus MC, University Medical Centre, Rotterdam, The Netherlands

    Department of Epidemiology, Harvard T.H. Chan School of Public Health, Boston, MA, USA

    Department of Internal Medicine, Erasmus Medical Center, Rotterdam, The Netherlands

  24. Brigham and Women's Hospital · Harvard University

    Affiliation as printed

    Division of Preventive Medicine, Brigham and Women's Hospital, Boston, USA

    Harvard Medical School, Boston, USA

  25. IRCCS Materno Infantile Burlo Garofolo

    Affiliation as printed

    Institute for Maternal and Child Health - IRCCS "Burlo Garofolo", Trieste, Italy

    Institute for Maternal and Child Health – IRCCS "Burlo Garofolo", Trieste, Italy

  26. Leiden University Medical Center

    Affiliation as printed

    Department of Biomedical Data Sciences, Section Molecular Epidemiology, Leiden University Medical Center, Leiden, The Netherlands

  27. QIMR Berghofer Medical Research Institute

    Affiliation as printed

    QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia

  28. University of Sassari · Institute of Genetic and Biomedical Research · National Research Council

    Affiliation as printed

    Istituto di Ricerca Genetica e Biomedica, Consiglio Nazionale delle Ricerche, 09042, Monserrato (CA), Italy

    Università di Sassari, Dipartimento di Scienze Biomediche, V.le San Pietro, 07100, Sassari (SS), Italy

  29. University of Edinburgh

    Affiliation as printed

    Lothian Birth Cohorts, Department of Psychology, University of Edinburgh, EH8 9JZ, Edinburgh, United Kingdom

  30. Aarhus University Hospital · Steno Diabetes Center Aarhus

    Affiliation as printed

    Steno Diabetes Center Aarhus, Aarhus University Hospital, Palle Juul-Jensens Blvd. 11, Entrance A, 8200, Aarhus, Denmark

  31. Johns Hopkins University · Johns Hopkins Medicine

    Affiliation as printed

    Division of Endocrinology, Diabetes, and Metabolism, Johns Hopkins School of Medicine, Baltimore, MD, 21205, USA

  32. University of Copenhagen · Boston Children's Hospital · Harvard University · Zealand University Hospital Køge

    Affiliation as printed

    Department of Clinical Biochemistry, Zealand University Hospital, Køge, Denmark

    Department of Laboratory Medicine, Boston Children's Hospital, Boston, MA, USA

    Faculty of Medical Science, Department of Clinical Medicine, University of Copenhagen, Copenhagen, Denmark

    Harvard Medical School, Boston, USA

  33. University of Helsinki · National University of Singapore

    Affiliation as printed

    Department of General Practice and Primary health Care, University of Helsinki, Helsinki, Finland

    National University Singapore, Yong Loo Lin School of Medicine, Department of Obstetrics and Gynecology, Singapore, Singapore

  34. National Institute on Aging

    Affiliation as printed

    Longitudinal Study Section, National Institute on Aging, Baltimore, MD, USA

  35. Regeneron (United States)

    Affiliation as printed

    Regeneron Pharmaceuticals, Inc., Tarrytown, New York, USA

  36. Eurac Research · Helmholtz Munich · Institute for Biomedicine · University of Lübeck

    Affiliation as printed

    Institute for Biomedicine (affiliated with the University of Lübeck), Eurac Research, Bolzano, Italy

    Research Unit Molecular Epidemiology, Institute of Epidemiology, Helmholtz Zentrum München, Neuherberg, Germany

  37. Brigham and Women's Hospital · Helmholtz Munich

    Affiliation as printed

    Research Unit Molecular Epidemiology, Institute of Epidemiology, Helmholtz Zentrum München, Neuherberg, Germany

    Division of Preventive Medicine, Brigham and Women's Hospital, Boston, USA

  38. Brigham and Women's Hospital · Eurac Research · Institute for Biomedicine · University of Lübeck

    Affiliation as printed

    Division of Preventive Medicine, Brigham and Women's Hospital, Boston, USA

    Institute for Biomedicine (affiliated with the University of Lübeck), Eurac Research, Bolzano, Italy

  39. Eurac Research · University of Michigan · Michigan Medicine · Institute for Biomedicine · University of Lübeck

    Affiliation as printed

    Institute for Biomedicine (affiliated with the University of Lübeck), Eurac Research, Bolzano, Italy

    Department of Internal Medicine, Cardiology, University of Michigan, Ann Arbor, MI, 48109, USA

  40. University of Copenhagen · University of Michigan · Michigan Medicine · Novo Nordisk Foundation Center for Basic Metabolic Research

    Affiliation as printed

    Department of Internal Medicine, Cardiology, University of Michigan, Ann Arbor, MI, 48109, USA

    Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

  41. University of Copenhagen · Novo Nordisk Foundation Center for Basic Metabolic Research · University of Split

    Affiliation as printed

    Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

    Department of Medical Biology, University of Split, School of Medicine, Split, Croatia

  42. University of Copenhagen · Novo Nordisk Foundation Center for Basic Metabolic Research · University of Split

    Affiliation as printed

    Department of Medical Biology, University of Split, School of Medicine, Split, Croatia

    Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

  43. University of Copenhagen · University of Washington · Novo Nordisk Foundation Center for Basic Metabolic Research

    Affiliation as printed

    Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

    Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, WA, USA

  44. University of Washington · Barcelona Institute for Global Health · University of Edinburgh

    Affiliation as printed

    Barcelona Institute for Global Health, Barcelona, Spain

    Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, WA, USA

    Lothian Birth Cohorts, Department of Psychology, University of Edinburgh, EH8 9JZ, Edinburgh, United Kingdom

  45. Copenhagen University Hospital · Rigshospitalet · University of Edinburgh

    Affiliation as printed

    Lothian Birth Cohorts, Department of Psychology, University of Edinburgh, EH8 9JZ, Edinburgh, United Kingdom

    Laboratory for Molecular Cardiology, Department of Cardiology, Copenhagen University Hospital - Rigshospitalet, Copenhagen, Denmark

  46. University of Copenhagen · Western General Hospital · Copenhagen University Hospital · Rigshospitalet · Institute of Genetics and Cancer · MRC Human Genetics Unit · University of Edinburgh

    Affiliation as printed

    Department of Clinical Medicine, University of Copenhagen, Copenhagen, Denmark

    Laboratory for Molecular Cardiology, Department of Cardiology, Copenhagen University Hospital - Rigshospitalet, Copenhagen, Denmark

    MRC Human Genetics Unit, Institute of Genetics and Cancer, University of Edinburgh, Western General Hospital, Edinburgh, United Kingdom

  47. The University of Western Australia · Western General Hospital · Institute of Genetics and Cancer · Pathwest Laboratory Medicine · MRC Human Genetics Unit · University of Edinburgh

    Affiliation as printed

    MRC Human Genetics Unit, Institute of Genetics and Cancer, University of Edinburgh, Western General Hospital, Edinburgh, United Kingdom

    Pathwest Laboratory Medicine WA, Nedlands, WA, 6009, Australia

    School of Population and Global Health, The University of Western Australia, Crawley, WA, 6009, Australia

  48. The University of Western Australia · Universitätsmedizin Greifswald · Universität Greifswald · German Centre for Cardiovascular Research · Pathwest Laboratory Medicine

    Affiliation as printed

    Pathwest Laboratory Medicine WA, Nedlands, WA, 6009, Australia

    School of Population and Global Health, The University of Western Australia, Crawley, WA, 6009, Australia

    DZHK (German Center for Cardiovascular Research), partner site Greifswald, Greifswald, Germany

    Institute for Community Medicine, University Medicine Greifswald, 17475, Greifswald, Germany

  49. Universitätsmedizin Greifswald · Leiden University Medical Center · Universität Greifswald · German Centre for Cardiovascular Research

    Affiliation as printed

    DZHK (German Center for Cardiovascular Research), partner site Greifswald, Greifswald, Germany

    Institute for Community Medicine, University Medicine Greifswald, 17475, Greifswald, Germany

    Department of Cardiology, Leiden University Medical Center, Leiden, the Netherlands

  50. Norwegian University of Science and Technology · Oulu University Hospital · Leiden University Medical Center · Finnish Institute for Health and Welfare · Netherlands Heart Institute · University of Oulu

    Affiliation as printed

    Department of Cardiology, Leiden University Medical Center, Leiden, the Netherlands

    Netherlands Heart Institute, Utrecht, the Netherlands

    Clinical Medicine Research Unit, MRC Oulu, Oulu University Hospital and University of Oulu, Oulu, Finland

    Department of Clinical and Molecular Medicine, Norwegian University of Science and Technology, Trondheim, Norway

    Finnish Institute for Health and Welfare, Population Health Unit, Helsinki and Oulu, Oulu, Finland

  51. University of Copenhagen · Norwegian University of Science and Technology · Oulu University Hospital · Finnish Institute for Health and Welfare · University of Oulu

    Affiliation as printed

    Clinical Medicine Research Unit, MRC Oulu, Oulu University Hospital and University of Oulu, Oulu, Finland

    Department of Clinical and Molecular Medicine, Norwegian University of Science and Technology, Trondheim, Norway

    Finnish Institute for Health and Welfare, Population Health Unit, Helsinki and Oulu, Oulu, Finland

    Department of Biomedical Sciences, University of Copenhagen, Copenhagen, Denmark

  52. University of Copenhagen · University of California, San Francisco · Frederiksberg Hospital · Bispebjerg Hospital · Center for Clinical Research and Prevention

    Affiliation as printed

    Center of Physiological Research, University of California San Francisco, San Francisco, USA

    Department of Biomedical Sciences, University of Copenhagen, Copenhagen, Denmark

    Center for Clinical Research and Prevention, Bispebjerg and Frederiksberg Hospital, Copenhagen, Denmark

  53. Frederiksberg Hospital · Bispebjerg Hospital · Radboud University Medical Center · Center for Clinical Research and Prevention

    Affiliation as printed

    Center for Clinical Research and Prevention, Bispebjerg and Frederiksberg Hospital, Copenhagen, Denmark

    Department for Health Evidence, Radboud University Medical Center, Nijmegen, The Netherlands

  54. Leiden University Medical Center · Radboud University Medical Center

    Affiliation as printed

    Department for Health Evidence, Radboud University Medical Center, Nijmegen, The Netherlands

    Department of Urology, Radboud University Medical Center, Nijmegen, The Netherlands

    Departments of Rheumatology and Clinical Epidemiology, Leiden University Medical Center, Leiden, The Netherlands

  55. Leiden University Medical Center · Helmholtz Munich

    Affiliation as printed

    Departments of Rheumatology and Clinical Epidemiology, Leiden University Medical Center, Leiden, The Netherlands

    Research Unit Molecular Epidemiology, Institute of Epidemiology, Helmholtz Zentrum München, Neuherberg, Germany

  56. University of Helsinki · Helmholtz Munich

    Affiliation as printed

    Research Unit Molecular Epidemiology, Institute of Epidemiology, Helmholtz Zentrum München, Neuherberg, Germany

    Department of Psychology and Logopedics, Faculty of Medicine, University of Helsinki, Helsinki, Finland

  57. University of Helsinki · Queen Mary University of London · University of Cambridge · MRC Epidemiology Unit

    Affiliation as printed

    Department of Psychology and Logopedics, Faculty of Medicine, University of Helsinki, Helsinki, Finland

    MRC Epidemiology Unit, Institute of Metabolic Science, University of Cambridge School of Clinical Medicine, Cambridge, CB2 0QQ, UK

    Precision Healthcare University Research Institute, Queen Mary University of London, London, UK

  58. Queen Mary University of London · University of Cambridge · Ghent University Hospital · Berlin Institute of Health at Charité - Universitätsmedizin Berlin · MRC Epidemiology Unit

    Affiliation as printed

    Computational Medicine, Berlin Institute of Health at Charité - Universitätsmedizin Berlin, Berlin, Germany

    MRC Epidemiology Unit, Institute of Metabolic Science, University of Cambridge School of Clinical Medicine, Cambridge, CB2 0QQ, UK

    Precision Healthcare University Research Institute, Queen Mary University of London, London, UK

    Department of Endocrinology, Ghent University Hospital, C. Heymanslaan 10, 9000, Ghent, Belgium

  59. NHS Tayside · Ghent University Hospital

    Affiliation as printed

    Department of Endocrinology, Ghent University Hospital, C. Heymanslaan 10, 9000, Ghent, Belgium

    NHS Tayside, Scotland, Dundee, UK

  60. Boston University · NHS Tayside

    Affiliation as printed

    NHS Tayside, Scotland, Dundee, UK

    Department of Biostatistics, Boston University, Boston, MA, USA

  61. Boston University · University of Edinburgh

    Affiliation as printed

    Department of Biostatistics, Boston University, Boston, MA, USA

    Lothian Birth Cohorts, Department of Psychology, University of Edinburgh, EH8 9JZ, Edinburgh, United Kingdom

  62. University of California, San Francisco · University of Edinburgh

    Affiliation as printed

    Lothian Birth Cohorts, Department of Psychology, University of Edinburgh, EH8 9JZ, Edinburgh, United Kingdom

    Center of Physiological Research, University of California San Francisco, San Francisco, USA

  63. University of Copenhagen · University of California, San Francisco · Novo Nordisk Foundation Center for Basic Metabolic Research

    Affiliation as printed

    Center of Physiological Research, University of California San Francisco, San Francisco, USA

    Department of Clinical Medicine, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

    Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

  64. University of Copenhagen · University of Cambridge · MRC Epidemiology Unit · Novo Nordisk Foundation Center for Basic Metabolic Research

    Affiliation as printed

    Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

    MRC Epidemiology Unit, Institute of Metabolic Science, University of Cambridge School of Clinical Medicine, Cambridge, CB2 0QQ, UK

  65. QIMR Berghofer Medical Research Institute · University of Cambridge · MRC Epidemiology Unit

    Affiliation as printed

    MRC Epidemiology Unit, Institute of Metabolic Science, University of Cambridge School of Clinical Medicine, Cambridge, CB2 0QQ, UK

    QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia

  66. QIMR Berghofer Medical Research Institute · University of Split

    Affiliation as printed

    QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia

    Department of Medical Biology, University of Split, School of Medicine, Split, Croatia

  67. Erasmus MC · University of Split

    Affiliation as printed

    Department of Medical Biology, University of Split, School of Medicine, Split, Croatia

    Academic Center for Thyroid Diseases, Department of Internal Medicine, Erasmus Medical Center, Rotterdam, The Netherlands

  68. Erasmus MC · Technical University of Munich

    Affiliation as printed

    Academic Center for Thyroid Diseases, Department of Internal Medicine, Erasmus Medical Center, Rotterdam, The Netherlands

    Institute for Human Genetics, Technical University of Munich, Munich, Germany

  69. Leiden University Medical Center · Technical University of Munich

    Affiliation as printed

    Institute for Human Genetics, Technical University of Munich, Munich, Germany

    Department of Biomedical Data Sciences, Section Molecular Epidemiology, Leiden University Medical Center, Leiden, The Netherlands

  70. University of Maryland, Baltimore · Leiden University Medical Center

    Affiliation as printed

    Department of Biomedical Data Sciences, Section Molecular Epidemiology, Leiden University Medical Center, Leiden, The Netherlands

    University of Maryland School of Medicine, Division of Endocrinology, Diabetes and Nutrition, Baltimore, USA

  71. University of Maryland, Baltimore · Frederiksberg Hospital · Bispebjerg Hospital · Center for Clinical Research and Prevention · Baltimore VA Medical Center

    Affiliation as printed

    Geriatrics Research and Education Clinical Center, Baltimore Veterans Administration Medical Center, Baltimore, MD, 21201, USA

    University of Maryland School of Medicine, Division of Endocrinology, Diabetes and Nutrition, Baltimore, USA

    Center for Clinical Research and Prevention, Bispebjerg and Frederiksberg Hospital, Copenhagen, Denmark

  72. Brigham and Women's Hospital · Harvard University · Frederiksberg Hospital · Bispebjerg Hospital · Center for Clinical Research and Prevention

    Affiliation as printed

    Center for Clinical Research and Prevention, Bispebjerg and Frederiksberg Hospital, Copenhagen, Denmark

    Division of Preventive Medicine, Brigham and Women's Hospital, Boston, USA

    Harvard Medical School, Boston, USA

  73. Brigham and Women's Hospital · Harvard University · Institute of Genetic and Biomedical Research · National Research Council

    Affiliation as printed

    Division of Preventive Medicine, Brigham and Women's Hospital, Boston, USA

    Harvard Medical School, Boston, USA

    Istituto di Ricerca Genetica e Biomedica, Consiglio Nazionale delle Ricerche, 09042, Monserrato (CA), Italy

  74. Institute of Genetic and Biomedical Research · German Centre for Cardiovascular Research · National Research Council

    Affiliation as printed

    Istituto di Ricerca Genetica e Biomedica, Consiglio Nazionale delle Ricerche, 09042, Monserrato (CA), Italy

    DZHK (German Center for Cardiovascular Research), partner site Greifswald, Greifswald, Germany

  75. Universitätsmedizin Greifswald · Radboud University Medical Center · Universität Greifswald · German Centre for Cardiovascular Research

    Affiliation as printed

    DZHK (German Center for Cardiovascular Research), partner site Greifswald, Greifswald, Germany

    Institute of Clinical Chemistry and Laboratory Medicine, University Medicine Greifswald, Greifswald, Germany

    Department of Internal Medicine, Division of Endocrinology, Radboud University Medical Center, Nijmegen, The Netherlands

  76. Leiden University Medical Center · Radboud University Medical Center

    Affiliation as printed

    Department of Internal Medicine, Division of Endocrinology, Radboud University Medical Center, Nijmegen, The Netherlands

    Department of Internal Medicine, Section of Gerontology and Geriatrics, Leiden University Medical Center, Leiden, the Netherlands

  77. University of Copenhagen · Leiden University Medical Center · Novo Nordisk Foundation Center for Basic Metabolic Research

    Affiliation as printed

    Department of Internal Medicine, Section of Gerontology and Geriatrics, Leiden University Medical Center, Leiden, the Netherlands

    Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

  78. University of Copenhagen · Novo Nordisk Foundation Center for Basic Metabolic Research · The University of Tokyo · The University of Osaka

    Affiliation as printed

    Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

    Department of Genome Informatics, Graduate School of Medicine, the University of Tokyo, Tokyo, Japan

    Laboratory of Statistical Immunology, Immunology Frontier Research Center (WPI-IFReC), Osaka University, Suita, Japan

    Premium Research Institute for Human Metaverse Medicine (WPI-PRIMe), Osaka University, Suita, Japan

  79. Institute of Genetic and Biomedical Research · RIKEN Center for Integrative Medical Sciences · National Research Council · The University of Tokyo · The University of Osaka

    Affiliation as printed

    Department of Genome Informatics, Graduate School of Medicine, the University of Tokyo, Tokyo, Japan

    Department of Statistical Genetics, Osaka University Graduate School of Medicine, Suita, Japan

    Laboratory for Systems Genetics, RIKEN Center for Integrative Medical Sciences, Kanagawa, Japan

    Laboratory of Statistical Immunology, Immunology Frontier Research Center (WPI-IFReC), Osaka University, Suita, Japan

    Premium Research Institute for Human Metaverse Medicine (WPI-PRIMe), Osaka University, Suita, Japan

    Istituto di Ricerca Genetica e Biomedica, Consiglio Nazionale delle Ricerche, 09042, Monserrato (CA), Italy

  80. Queen Mary University of London · Institute of Genetic and Biomedical Research · William Harvey Research Institute · National Research Council

    Affiliation as printed

    Istituto di Ricerca Genetica e Biomedica, Consiglio Nazionale delle Ricerche, 09042, Monserrato (CA), Italy

    William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London, United Kingdom

  81. Queen Mary University of London · University of Dundee · William Harvey Research Institute

    Affiliation as printed

    William Harvey Research Institute, Barts and The London School of Medicine and Dentistry, Queen Mary University of London, London, United Kingdom

    Division of Population Health Genomics, School of Medicine, University of Dundee, DD19SY, Dundee, UK

  82. University of Copenhagen · Eurac Research · Institute for Biomedicine · Novo Nordisk Foundation Center for Basic Metabolic Research · University of Lübeck

    Affiliation as printed

    Institute for Biomedicine (affiliated with the University of Lübeck), Eurac Research, Bolzano, Italy

    Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

  83. University of Copenhagen · Gentofte Hospital · Helmholtz Munich · Novo Nordisk Foundation Center for Basic Metabolic Research · Institut für Medizinische Informationsverarbeitung, Biometrie und Epidemiologie · Ludwig-Maximilians-Universität München

    Affiliation as printed

    Center for Clinical Metabolic Research, Herlev-Gentofte University Hospital, Copenhagen, Denmark

    Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

    Chair of Epidemiology, Institute for Medical Information Processing, Biometry and Epidemiology, Medical Faculty, Ludwig-Maximilians-Universität München, Munich, Germany

    Institute of Epidemiology, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, Germany

  84. Queen Mary University of London · University of Cambridge · Helmholtz Munich · MRC Epidemiology Unit · Institut für Medizinische Informationsverarbeitung, Biometrie und Epidemiologie · Ludwig-Maximilians-Universität München

    Affiliation as printed

    Chair of Epidemiology, Institute for Medical Information Processing, Biometry and Epidemiology, Medical Faculty, Ludwig-Maximilians-Universität München, Munich, Germany

    Institute of Epidemiology, Helmholtz Zentrum München, German Research Center for Environmental Health, Neuherberg, Germany

    MRC Epidemiology Unit, Institute of Metabolic Science, University of Cambridge School of Clinical Medicine, Cambridge, CB2 0QQ, UK

    Precision Healthcare University Research Institute, Queen Mary University of London, London, UK

  85. Queen Mary University of London · University of Cambridge · Algebra Bernays University · Berlin Institute of Health at Charité - Universitätsmedizin Berlin · MRC Epidemiology Unit · University of Split

    Affiliation as printed

    Computational Medicine, Berlin Institute of Health at Charité - Universitätsmedizin Berlin, Berlin, Germany

    MRC Epidemiology Unit, Institute of Metabolic Science, University of Cambridge School of Clinical Medicine, Cambridge, CB2 0QQ, UK

    Precision Healthcare University Research Institute, Queen Mary University of London, London, UK

    Algebra University College, Zagreb, Croatia

    Department of Public Health, University of Split, School of Medicine, Split, Croatia

  86. Eurac Research · Algebra Bernays University · Institute for Biomedicine · University of Split · University of Lübeck

    Affiliation as printed

    Algebra University College, Zagreb, Croatia

    Department of Public Health, University of Split, School of Medicine, Split, Croatia

    Institute for Biomedicine (affiliated with the University of Lübeck), Eurac Research, Bolzano, Italy

  87. Eurac Research · University of Washington · Institute for Biomedicine · University of Lübeck

    Affiliation as printed

    Institute for Biomedicine (affiliated with the University of Lübeck), Eurac Research, Bolzano, Italy

    Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, WA, USA

  88. University of Washington · Klinički Bolnički Centar Split · University of Split

    Affiliation as printed

    Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, WA, USA

    Departments of Epidemiology and Health Systems and Population Health, University of Washington, Seattle, WA, USA

    Department of Nuclear Medicine, University Hospital Split, Split, Croatia

  89. Johns Hopkins University · Klinički Bolnički Centar Split · University of Split

    Affiliation as printed

    Department of Nuclear Medicine, University Hospital Split, Split, Croatia

    Department of Epidemiology, Johns Hopkins Bloomberg School of Public Health, Baltimore, MD, 21205, USA

  90. Johns Hopkins University · University of Edinburgh

    Affiliation as printed

    Department of Epidemiology, Johns Hopkins Bloomberg School of Public Health, Baltimore, MD, 21205, USA

    Lothian Birth Cohorts, Department of Psychology, University of Edinburgh, EH8 9JZ, Edinburgh, United Kingdom

  91. Jewish General Hospital · Lady Davis Institute for Medical Research · University of Edinburgh

    Affiliation as printed

    Lothian Birth Cohorts, Department of Psychology, University of Edinburgh, EH8 9JZ, Edinburgh, United Kingdom

    Lady Davis Institute, Jewish General Hospital, Montreal, Quebec, H3T 1E2, Canada

  92. Brigham and Women's Hospital · Harvard University · Jewish General Hospital · Lady Davis Institute for Medical Research

    Affiliation as printed

    Lady Davis Institute, Jewish General Hospital, Montreal, Quebec, H3T 1E2, Canada

    Division of Preventive Medicine, Brigham and Women's Hospital, Boston, USA

    Harvard Medical School, Boston, USA

  93. Brigham and Women's Hospital · Harvard University · University of Edinburgh

    Affiliation as printed

    Division of Preventive Medicine, Brigham and Women's Hospital, Boston, USA

    Harvard Medical School, Boston, USA

    Lothian Birth Cohorts, Department of Psychology, University of Edinburgh, EH8 9JZ, Edinburgh, United Kingdom

  94. University of Maryland, Baltimore · Alzheimer Scotland · University of Edinburgh

    Affiliation as printed

    Alzheimer Scotland Dementia Research Centre, University of Edinburgh, Edinburgh, United Kingdom

    Lothian Birth Cohorts, Department of Psychology, University of Edinburgh, EH8 9JZ, Edinburgh, United Kingdom

    University of Maryland School of Medicine, Division of Endocrinology, Diabetes and Nutrition, Baltimore, USA

  95. University of Copenhagen · University of Maryland, Baltimore · Copenhagen University Hospital · Rigshospitalet

    Affiliation as printed

    University of Maryland School of Medicine, Division of Endocrinology, Diabetes and Nutrition, Baltimore, USA

    Department of Biomedical Sciences, University of Copenhagen, Copenhagen, Denmark

    Laboratory for Molecular Cardiology, Department of Cardiology, Copenhagen University Hospital - Rigshospitalet, Copenhagen, Denmark

  96. University of Copenhagen · University of Freiburg · University Medical Center Freiburg · Copenhagen University Hospital · Rigshospitalet

    Affiliation as printed

    Department of Biomedical Sciences, University of Copenhagen, Copenhagen, Denmark

    Laboratory for Molecular Cardiology, Department of Cardiology, Copenhagen University Hospital - Rigshospitalet, Copenhagen, Denmark

    Department of Medicine IV – Nephrology and Primary Care, Faculty of Medicine and Medical Center – University of Freiburg, Freiburg, Germany

  97. Johns Hopkins University · University of Freiburg · University Medical Center Freiburg

    Affiliation as printed

    Department of Medicine IV - Nephrology and Primary Care, Faculty of Medicine and Medical Center - University of Freiburg, Freiburg, Germany

    Institute of Genetic Epidemiology, Faculty of Medicine and Medical Center - University of Freiburg, Freiburg, Germany

    Department of Epidemiology, Johns Hopkins Bloomberg School of Public Health, Baltimore, MD, 21205, USA

  98. Johns Hopkins University · Queen Mary University of London

    Affiliation as printed

    Department of Epidemiology, Johns Hopkins Bloomberg School of Public Health, Baltimore, MD, 21205, USA

    Wolfson Institute of Population Health, Queen Mary University of London, London, UK

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