X-chromosome and kidney function: evidence from a multi-trait genetic analysis of 908,697 individuals reveals sex-specific and sex-differential findings in genes regulated by androgen response elements
Nature Communications, vol. 15, pp. 586
Abstract
X-chromosomal genetic variants are understudied but can yield valuable insights into sexually dimorphic human traits and diseases. We performed a sex-stratified cross-ancestry X-chromosome-wide association meta-analysis of seven kidney-related traits (n = 908,697), identifying 23 loci genome-wide significantly associated with two of the traits: 7 for uric acid and 16 for estimated glomerular filtration rate (eGFR), including four novel eGFR loci containing the functionally plausible prioritized genes ACSL4, CLDN2, TSPAN6 and the female-specific DRP2. Further, we identified five novel sex-interactions, comprising male-specific effects at FAM9B and AR/EDA2R, and three sex-differential findings with larger genetic effect sizes in males at DCAF12L1 and MST4 and larger effect sizes in females at HPRT1. All prioritized genes in loci showing significant sex-interactions were located next to androgen response elements (ARE). Five ARE genes showed sex-differential expressions. This study contributes new insights into sex-dimorphisms of kidney traits along with new prioritized gene targets for further molecular research.
Authors 100
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Markus R. Scholz corresponding
Affiliation as printed
Institute for Medical Informatics, Statistics and Epidemiology, University of Leipzig, Leipzig, Germany. markus.scholz@imise.uni-leipzig.de
LIFE Research Center for Civilization Diseases, University of Leipzig, Leipzig, Germany. markus.scholz@imise.uni-leipzig.de
Institute for Medical Informatics, Statistics and Epidemiology, University of Leipzig, Leipzig, Germany
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Affiliation as printed
Institute for Medical Informatics, Statistics and Epidemiology, University of Leipzig, Leipzig, Germany
LIFE Research Center for Civilization Diseases, University of Leipzig, Leipzig, Germany
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Affiliation as printed
Institute for Medical Informatics, Statistics and Epidemiology, University of Leipzig, Leipzig, Germany
LIFE Research Center for Civilization Diseases, University of Leipzig, Leipzig, Germany
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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, Department of Data Driven Medicine, Faculty of Medicine and Medical Center-University of Freiburg, Freiburg, Germany
Institute of Genetic Epidemiology, Department of Data Driven Medicine, Faculty of Medicine and Medical Center–University of Freiburg, Freiburg, Germany
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Affiliation as printed
Institute for Medical Informatics, Statistics and Epidemiology, University of Leipzig, Leipzig, Germany
LIFE Research Center for Civilization Diseases, University of Leipzig, Leipzig, Germany
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Universitätsmedizin Greifswald · Universität Greifswald · German Centre for Cardiovascular Research
Affiliation as printed
DZHK (German Center for Cardiovascular Research), partner site Greifswald, Greifswald, Germany
Department of Psychiatry and Psychotherapy, University Medicine Greifswald, Greifswald, Germany
Institute for Community Medicine, University Medicine Greifswald, Greifswald, Germany
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Affiliation as printed
Institute for Medical Informatics, Statistics and Epidemiology, University of Leipzig, Leipzig, Germany
LIFE Research Center for Civilization Diseases, University of Leipzig, Leipzig, Germany
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University of Freiburg · University Medical Center Freiburg
Affiliation as printed
Institute of Genetic Epidemiology, Department of Data Driven Medicine, Faculty of Medicine and Medical Center-University of Freiburg, Freiburg, Germany
Institute of Genetic Epidemiology, Department of Data Driven Medicine, Faculty of Medicine and Medical Center–University of Freiburg, Freiburg, Germany
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University of Freiburg · University Medical Center Freiburg
Affiliation as printed
Institute of Genetic Epidemiology, Department of Data Driven Medicine, Faculty of Medicine and Medical Center-University of Freiburg, Freiburg, Germany
Institute of Genetic Epidemiology, Department of Data Driven Medicine, Faculty of Medicine and Medical Center–University of Freiburg, Freiburg, Germany
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University Hospital Regensburg · University of Regensburg
Affiliation as printed
Department of Genetic Epidemiology, University of Regensburg, Regensburg, Germany
Department of Nephrology, University Hospital Regensburg, Regensburg, Germany
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Universitätsmedizin Greifswald · 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, Greifswald, Germany
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Affiliation as printed
Division of Nephrology and Hypertension, Department of Medicine, University of Utah, Salt Lake City, UT, USA
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University of Mississippi Medical Center
Affiliation as printed
Division of Nephrology, Department of Medicine, University of Mississippi Medical Center, Jackson, MS, USA
Memory Impairment and Neurodegenerative Dementia (MIND) Center, University of Mississippi Medical Center, Jackson, MS, USA
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National University of Singapore · National University Health System
Affiliation as printed
Saw Swee Hock School of Public Health, National University of Singapore and National University Health System, Singapore, Singapore
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IRCCS Materno Infantile Burlo Garofolo
Affiliation as printed
Institute for Maternal and Child Health, IRCCS 'Burlo Garofolo', Trieste, Italy
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Washington University in St. Louis
Affiliation as printed
Division of Statistical Genomics, Department of Genetics, Washington University School of Medicine, St. Louis, MO, USA
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Institute of Genetics and Biophysics
Affiliation as printed
Institute of Genetics and Biophysics 'Adriano Buzzati-Traverso'-CNR, Naples, Italy
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Kyushu University · RIKEN Center for Integrative Medical Sciences
Affiliation as printed
Department of Ocular Pathology and Imaging Science, Graduate School of Medical Sciences, Kyushu University, Fukuoka, Japan
Laboratory for Statistical Analysis, RIKEN Center for Integrative Medical Sciences (IMS), Yokohama, Japan
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Norwegian University of Science and Technology · St Olav's University Hospital
Affiliation as printed
Department of Endocrinology, St. Olavs Hospital, Trondheim University Hospital, Trondheim, Norway
K. G. Jebsen Center for Genetic Epidemiology, Department of Public Health and Nursing, Faculty of Medicine and Health, Norwegian University of Science and Technology, Trondheim, Norway
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Affiliation as printed
Division of Nephrology, University of Washington, Seattle, WA, USA
Kidney Research Institute, University of Washington, Seattle, WA, USA
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Affiliation as printed
Cardiovascular Health Research Unit, Department of Medicine, University of Washington, Seattle, WA, USA
Department of Biostatistics, University of Washington, Seattle, WA, USA
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Institute of Genetics and Cancer · MRC Human Genetics Unit · University of Edinburgh
Affiliation as printed
Medical Research Council Human Genetics Unit, Institute of Genetics and Cancer, University of Edinburgh, Edinburgh, EH4 2XU, UK
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German Cancer Research Center · Heidelberg University
Affiliation as printed
Division of Clinical Epidemiology and Aging Research, German Cancer Research Center (DKFZ), Heidelberg, Germany
Network Aging Research, Heidelberg University, Heidelberg, Germany
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Norwegian University of Science and Technology · University of Bristol · St Olav's University Hospital · MRC Integrative Epidemiology Unit
Affiliation as printed
Clinic of Thoracic and Occupational Medicine, St. Olavs Hospital, Trondheim University Hospital, Trondheim, Norway
K. G. Jebsen Center for Genetic Epidemiology, Department of Public Health and Nursing, Faculty of Medicine and Health, Norwegian University of Science and Technology, Trondheim, Norway
MRC Integrative Epidemiology Unit, Population Health Sciences, Bristol Medical School, University of Bristol, Bristol, UK
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University Hospital Regensburg · Leipzig University
Affiliation as printed
Institute of Clinical Chemistry and Laboratory Medicine, University Hospital Regensburg, Regensburg, Germany
LIFE Research Center for Civilization Diseases, University of Leipzig, Leipzig, Germany
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University of North Carolina at Chapel Hill
Affiliation as printed
Department of Biostatistics, Gillings School of Global Public Health, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA
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Institute of Genetics and Cancer · University of Edinburgh
Affiliation as printed
Centre for Genomic and Experimental Medicine, Institute of Genetics & Cancer, University of Edinburgh, Edinburgh, EH4 2XU, UK
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Affiliation as printed
Centre for Global Health, Usher Institute, University of Edinburgh, Teviot Place, Edinburgh, EH8 9AG, Scotland
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UNSW Sydney · The George Institute for Global Health
Affiliation as printed
The George Institute for Global Health, University of New South Wales, Sydney, NSW, Australia
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Brigham and Women's Hospital · Harvard University
Affiliation as printed
Division of Preventive Medicine, Brigham and Women's Hospital, Boston, MA, USA
Harvard Medical School, Boston, MA, USA
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Singapore National Eye Center · Singapore Eye Research Institute
Affiliation as printed
Singapore Eye Research Institute, Singapore National Eye Center, Singapore, Singapore
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Singapore National Eye Center · Singapore Eye Research Institute
Affiliation as printed
Singapore Eye Research Institute, Singapore National Eye Center, Singapore, Singapore
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Affiliation as printed
Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Solna, Sweden
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National University of Singapore · Singapore National Eye Center · Singapore Eye Research Institute · Duke-NUS Medical School · National University Health System
Affiliation as printed
Department of Ophthalmology, Yong Loo Lin School of Medicine, National University of Singapore and National University Health System, Singapore, Singapore
Ophthalmology and Visual Sciences Academic Clinical Program (Eye ACP), Duke-NUS Medical School, Singapore, Singapore
Singapore Eye Research Institute, Singapore National Eye Center, Singapore, Singapore
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Charles University · Thomayer University Hospital
Affiliation as printed
Center for Cardiovascular Prevention, Charles University in Prague, First Faculty of Medicine and Thomayer University Hospital, Prague, Czech Republic
Department of Medicine II, Charles University in Prague, First Faculty of Medicine, Prague, Czech Republic
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University of Illinois Chicago
Affiliation as printed
Institute for Minority Health Research, University of Illinois at Chicago, Chicago, IL, USA
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Heidelberg University · Medizinische Fakultät Mannheim
Affiliation as printed
Vth Department of Medicine (Nephrology, Hypertensiology, Rheumatology, Endocrinology, Diabetology), Medical Faculty Mannheim, University of Heidelberg, Heidelberg, Germany
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University Hospital Leipzig · Leipzig University
Affiliation as printed
University Hospital for Children and Adolescents, Pediatric Research Unit, Medical Faculty, University Medical Center, University of Leipzig, Leipzig, Germany
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Vanderbilt University · VA Tennessee Valley Healthcare System · Vanderbilt University Medical Center
Affiliation as printed
Department of Veteran's Affairs, Tennessee Valley Healthcare System (626)/Vanderbilt University, Nashville, TN, USA
Division of Epidemiology, Department of Medicine, Vanderbilt Genetics Institute, Vanderbilt University Medical Center, Nashville, TN, USA
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Universitätsmedizin Greifswald · Universität Greifswald · German Centre for Cardiovascular Research
Affiliation as printed
DZHK (German Center for Cardiovascular Research), partner site Greifswald, Greifswald, Germany
Department of Anatomy and Cell Biology, University Medicine Greifswald, Greifswald, Germany
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Affiliation as printed
Massachusetts Veterans Epidemiology Research and Information Center, VA Cooperative Studies Program, VA Boston Healthcare System, Boston, MA, USA
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Vanderbilt University · Vanderbilt University Medical Center
Affiliation as printed
Biomedical Laboratory Research and Development, Tennessee Valley Healthcare System (626)/Vanderbilt University, Nashville, TN, USA
Division of Quantitative Sciences, Department of Obstetrics & Gynecology, Vanderbilt Genetics Institute, Vanderbilt Epidemiology Center, Institute for Medicine and Public Health, Vanderbilt University Medical Center, Nashville, TN, USA
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Affiliation as printed
Division of Preventive Medicine, Brigham and Women's Hospital, Boston, MA, USA
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QIMR Berghofer Medical Research Institute
Affiliation as printed
QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia
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deCODE Genetics (Iceland) · University of Iceland
Affiliation as printed
Iceland School of Engineering and Natural Sciences, University of Iceland, Reykjavik, Iceland
deCODE Genetics/Amgen, Inc., Reykjavik, Iceland
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Norwegian University of Science and Technology · St Olav's University Hospital
Affiliation as printed
Department of Clinical and Molecular Medicine, NTNU, Norwegian University of Science and Technology, Trondheim, Norway
Department of Nephrology, St. Olavs Hospital, Trondheim University Hospital, Trondheim, Norway
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Centre Hospitalier de l’Université de Montréal · Medpharmgene (Canada)
Affiliation as printed
Medpharmgene, Montreal, QC, Canada
Montreal University Hospital Research Center, CHUM, Montréal, QC, Canada
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University Medical Center Groningen · University of Groningen
Affiliation as printed
Interdisciplinary Centre Psychopathology and Emotion regulation (ICPE), Department of Psychiatry, University of Groningen, University Medical Center Groningen, Groningen, the Netherlands
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Institute of Genetics and Cancer · MRC Human Genetics Unit · University of Edinburgh
Affiliation as printed
Medical Research Council Human Genetics Unit, Institute of Genetics and Cancer, University of Edinburgh, Edinburgh, EH4 2XU, UK
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Affiliation as printed
Department of Genetic Epidemiology, University of Regensburg, Regensburg, Germany
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Vanderbilt University · VA Tennessee Valley Healthcare System · Vanderbilt University Medical Center
Affiliation as printed
Department of Veteran's Affairs, Tennessee Valley Healthcare System (626)/Vanderbilt University, Nashville, TN, USA
Division of Epidemiology, Department of Medicine, Vanderbilt Genetics Institute, Vanderbilt University Medical Center, Nashville, TN, USA
Vanderbilt Genetics Institute, Vanderbilt University Medical Center, Nashville, TN, USA
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Affiliation as printed
Division of Clinical Epidemiology and Aging Research, German Cancer Research Center (DKFZ), Heidelberg, Germany
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Affiliation as printed
deCODE Genetics/Amgen, Inc., Reykjavik, Iceland
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Tampere University · University of Tampere
Affiliation as printed
Tampere Centre for Skills Training and Simulation, Faculty of Medicine and Health Technology, Tampere University, Tampere, Finland
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Norwegian University of Science and Technology
Affiliation as printed
K. G. Jebsen Center for Genetic Epidemiology, Department of Public Health and Nursing, Faculty of Medicine and Health, Norwegian University of Science and Technology, Trondheim, Norway
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Affiliation as printed
Institute for Laboratory Medicine, University of Leipzig, Leipzig, Germany
LIFE Research Center for Civilization Diseases, University of Leipzig, Leipzig, Germany
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Beijing Tongren Hospital · Capital Medical University · Heidelberg University · Institute of Molecular and Clinical Ophthalmology Basel · Medizinische Fakultät Mannheim
Affiliation as printed
Beijing Institute of Ophthalmology, Key Laboratory of Ophthalmology and Visual Sciences, Beijing Tongren Hospital, Capital Medical University, Beijing, China
Department of Ophthalmology, Medical Faculty Mannheim, University Heidelberg, Mannheim, Germany
Institute of Molecular and Clinical Ophthalmology, Basel, Switzerland
Privatpraxis Prof Jonas und Dr Panda-Jonas, Heidelberg, Germany
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Affiliation as printed
Centre for Global Health, Usher Institute, University of Edinburgh, Teviot Place, Edinburgh, EH8 9AG, Scotland
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RIKEN Center for Integrative Medical Sciences · The University of Tokyo
Affiliation as printed
Laboratory for Statistical Analysis, RIKEN Center for Integrative Medical Sciences (IMS), Yokohama, Japan
Laboratory of Complex Trait Genomics, Department of Computational Biology and Medical Sciences, Graduate School of Frontier Sciences, The University of Tokyo, Tokyo, Japan
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Broad Institute · Harvard University · RIKEN Center for Integrative Medical Sciences
Affiliation as printed
Department of Biomedical Informatics, Harvard Medical School, Boston, MA, USA
Laboratory for Statistical Analysis, RIKEN Center for Integrative Medical Sciences (IMS), Yokohama, Japan
Program in Medical and Population Genetics, Broad Institute of Harvard and MIT, Cambridge, MA, USA
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Affiliation as printed
Finnish Medicines Agency, Kuopio, Finland
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Agency for Science, Technology and Research · National University of Singapore · National University Health System · Genome Institute of Singapore
Affiliation as printed
Department of Biochemistry, Yong Loo Lin School of Medicine, National University of Singapore and National University Health System, Singapore, Singapore
Genome Institute of Singapore, Agency for Science Technology and Research, Singapore, Singapore
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University Hospital Leipzig · Leipzig University
Affiliation as printed
LIFE Research Center for Civilization Diseases, University of Leipzig, Leipzig, Germany
University Hospital for Children and Adolescents, Pediatric Research Unit, Medical Faculty, University Medical Center, University of Leipzig, Leipzig, Germany
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Heidelberg University · Medizinische Fakultät Mannheim
Affiliation as printed
SYNLAB MVZ Humangenetik Mannheim, Mannheim, Germany
Vth Department of Medicine (Nephrology, Hypertensiology, Rheumatology, Endocrinology, Diabetology), Medical Faculty Mannheim, University of Heidelberg, Heidelberg, Germany
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Helmholtz Munich · University Hospital Leipzig · Leipzig University
Affiliation as printed
Helmholtz Institute for Metabolic, Obesity and Vascular Research (HI-MAG) of the Helmholtz Zentrum München at the University of Leipzig and University Hospital Leipzig, Leipzig, Germany
LIFE Research Center for Civilization Diseases, University of Leipzig, Leipzig, Germany
University Hospital for Children and Adolescents, Pediatric Research Unit, Medical Faculty, University Medical Center, University of Leipzig, Leipzig, Germany
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Affiliation as printed
Department of Medicine, University of Leipzig, Leipzig, Germany
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Charles University · Thomayer University Hospital
Affiliation as printed
Center for Cardiovascular Prevention, Charles University in Prague, First Faculty of Medicine and Thomayer University Hospital, Prague, Czech Republic
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Affiliation as printed
Department of Public Health Sciences, Loyola University Chicago, Maywood, IL, USA
Division of Nephrology and Hypertension, Loyola University Chicago, Chicago, IL, USA
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Heidelberg University · Medizinische Fakultät Mannheim
Affiliation as printed
Vth Department of Medicine (Nephrology, Hypertensiology, Rheumatology, Endocrinology, Diabetology), Medical Faculty Mannheim, University of Heidelberg, Heidelberg, Germany
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University of Helsinki · The University of Texas Rio Grande Valley · Finnish Institute for Health and Welfare
Affiliation as printed
Department of Human Genetics and South Texas Diabetes and Obesity Institute, University of Texas Rio Grande Valley School of Medicine, Brownsville, TX, USA
Finnish Institute for Health and Welfare, Helsinki, Finland
Research Program for Clinical and Molecular Metabolism, Faculty of Medicine, University of Helsinki, Helsinki, Finland
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Tampere University · Tampere University Hospital · University of Tampere
Affiliation as printed
Department of Clinical Physiology, Finnish Cardiovascular Research Center - Tampere, Faculty of Medicine and Health Technology, Tampere University, Tampere, Finland
Department of Clinical Physiology, Tampere University Hospital, Tampere, Finland
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University of Colorado Anschutz · University of Colorado Denver
Affiliation as printed
Division of Biomedical Informatics and Personalized Medicine, School of Medicine, University of Colorado Denver-Anschutz Medical Campus, Aurora, CO, USA
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University of Illinois Chicago
Affiliation as printed
Department of Medicine, University of Illinois at Chicago, Chicago, IL, USA
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Tampere University · Fimlab (Finland) · University of Tampere
Affiliation as printed
Department of Clinical Chemistry, Fimlab Laboratories, and The Wellbeing Services County of Pirkanmaa, Tampere, Finland
Department of Clinical Chemistry, Finnish Cardiovascular Research Center - Tampere, Faculty of Medicine and Health Technology, Tampere University, Tampere, Finland
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National University of Singapore · National University Health System
Affiliation as printed
Department of Ophthalmology, Yong Loo Lin School of Medicine, National University of Singapore and National University Health System, Singapore, Singapore
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University of North Carolina at Chapel Hill
Affiliation as printed
Department of Biostatistics, Gillings School of Global Public Health, University of North Carolina at Chapel Hill, Chapel Hill, NC, USA
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Agency for Science, Technology and Research · National University of Singapore · National University Health System · Genome Institute of Singapore
Affiliation as printed
Department of Medicine, Yong Loo Lin School of Medicine, National University of Singapore and National University Health System, Singapore, Singapore
Genome Institute of Singapore, Agency for Science Technology and Research, Singapore, Singapore
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Affiliation as printed
Institute for Medical Informatics, Statistics and Epidemiology, University of Leipzig, Leipzig, Germany
LIFE Research Center for Civilization Diseases, University of Leipzig, Leipzig, Germany
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Tampere University · Fimlab (Finland) · University of Tampere
Affiliation as printed
Department of Clinical Chemistry, Fimlab Laboratories, and The Wellbeing Services County of Pirkanmaa, Tampere, Finland
Department of Clinical Chemistry, Finnish Cardiovascular Research Center - Tampere, Faculty of Medicine and Health Technology, Tampere University, Tampere, Finland
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Affiliation as printed
Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Solna, Sweden
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QIMR Berghofer Medical Research Institute
Affiliation as printed
QIMR Berghofer Medical Research Institute, Brisbane, QLD, Australia
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Affiliation as printed
Laboratory of Clinical Genome Sequencing, Graduate School of Frontier Sciences, The University of Tokyo, Tokyo, Japan
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Amsterdam Neuroscience · Amsterdam Public Health · GGZ inGeest · Amsterdam University Medical Centers
Affiliation as printed
Department of Psychiatry, Amsterdam Public Health and Amsterdam Neuroscience, Amsterdam UMC/Vrije Universiteit and GGZ inGeest, Amsterdam, the Netherlands
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Tampere University · Fimlab (Finland) · University of Tampere
Affiliation as printed
Department of Clinical Chemistry, Fimlab Laboratories, and The Wellbeing Services County of Pirkanmaa, Tampere, Finland
Department of Clinical Chemistry, Finnish Cardiovascular Research Center - Tampere, Faculty of Medicine and Health Technology, Tampere University, Tampere, Finland
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Tampere University · Fimlab (Finland) · University of Tampere
Affiliation as printed
Department of Clinical Chemistry, Fimlab Laboratories, and The Wellbeing Services County of Pirkanmaa, Tampere, Finland
Department of Clinical Chemistry, Finnish Cardiovascular Research Center - Tampere, Faculty of Medicine and Health Technology, Tampere University, Tampere, Finland
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Affiliation as printed
University of Queensland, Brisbane, QLD, Australia
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Dennis O. Mook‐Kanamori Aachen Department of Clinical Epidemiology Department of Public Health and Primary Care
Leiden University Medical Center
Affiliation as printed
Department of Clinical Epidemiology, Leiden University Medical Center, Leiden, the Netherlands
Department of Public Health and Primary Care, Leiden University Medical Center, Leiden, the Netherlands
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Affiliation as printed
Center for Public Health Genomics, University of Virginia, Charlottesville, Charlottesville, VA, USA
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University of Graz · Medical University of Graz · Heidelberg University · Synlab (Germany) · Medizinische Fakultät Mannheim
Affiliation as printed
Clinical Institute of Medical and Chemical Laboratory Diagnostics, Medical University Graz, Graz, Austria
Synlab Academy, Synlab Holding Deutschland GmbH, Augsburg, Germany
Vth Department of Medicine (Nephrology, Hypertensiology, Rheumatology, Endocrinology, Diabetology), Medical Faculty Mannheim, University of Heidelberg, Heidelberg, Germany
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Universitätsmedizin Greifswald · 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
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Tampere University · Tampere University Hospital · University of Tampere
Affiliation as printed
Department of Cardiology, Finnish Cardiovascular Research Center - Tampere, Faculty of Medicine and Health Technology, Tampere University, Tampere, Finland
Department of Cardiology, Heart Center, Tampere University Hospital, Tampere, Finland
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University Medical Center Groningen · University of Groningen
Affiliation as printed
Department of Epidemiology, University of Groningen, University Medical Center Groningen, Groningen, the Netherlands
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Leiden University Medical Center
Affiliation as printed
Department of Internal Medicine, Section of Gerontology and Geriatrics, Leiden University Medical Center, Leiden, the Netherlands
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RIKEN Center for Integrative Medical Sciences · 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, Osaka, Japan
Laboratory for Systems Genetics, RIKEN Center for Integrative Medical Sciences (IMS), Yokohama, Japan
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National University Hospital of Iceland
Affiliation as printed
Department of Clinical Biochemistry, Landspitali University Hospital, Reykjavik, Iceland
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University Medical Center Groningen · University of Groningen
Affiliation as printed
Interdisciplinary Centre Psychopathology and Emotion regulation (ICPE), Department of Psychiatry, University of Groningen, University Medical Center Groningen, Groningen, the Netherlands
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Amsterdam Neuroscience · Amsterdam Public Health · GGZ inGeest · Amsterdam University Medical Centers
Affiliation as printed
Department of Psychiatry, Amsterdam Public Health and Amsterdam Neuroscience, Amsterdam UMC/Vrije Universiteit and GGZ inGeest, Amsterdam, the Netherlands
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University of Helsinki · Finnish Institute for Health and Welfare
Affiliation as printed
Finnish Institute for Health and Welfare, Helsinki, Finland
Research Program for Clinical and Molecular Metabolism, Faculty of Medicine, University of Helsinki, Helsinki, Finland
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Human Technopole · University of Edinburgh
Affiliation as printed
Biostatistics Unit - Population and Medical Genomics Programme, Genomics Research Centre, Human Technopole Palazzo Italia, Viale Rita Levi‑Montalcini, 1, 20157, Milan, Italy
Centre for Global Health, Usher Institute, University of Edinburgh, Teviot Place, Edinburgh, EH8 9AG, Scotland
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Affiliation as printed
University of Split Medical School, Split, Croatia
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