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Abstract
A Correction to this paper has been published: https://doi.org/10.1038/s41467-021-22613-2
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 ; Evans, Daniel S 2 ; Arking, Dan E 3 ; Tesi Niccolò 4
 
; Evans, Daniel S 2 ; Arking, Dan E 3 ; Tesi Niccolò 4  
 ; Nygaard, Marianne 5
 
; Nygaard, Marianne 5  
 ; Liu, Xiaomin 6
 
; Liu, Xiaomin 6  
 ; Wojczynski, Mary K 7 ; Biggs, Mary L 8 ; van der Spek Ashley 9
 
; Wojczynski, Mary K 7 ; Biggs, Mary L 8 ; van der Spek Ashley 9  
 ; Atzmon Gil 10 ; Ware, Erin B 11
 
; Atzmon Gil 10 ; Ware, Erin B 11  
 ; Sarnowski Chloé 12 ; Smith, Albert V 13 ; Seppälä Ilkka 14
 
; Sarnowski Chloé 12 ; Smith, Albert V 13 ; Seppälä Ilkka 14  
 ; Cordell, Heather J 15
 
; Cordell, Heather J 15  
 ; Dose Janina 16 ; Amin Najaf 9 ; Arnold, Alice M 17 ; Ayers, Kristin L 18 ; Barzilai Nir 19 ; Becker, Elizabeth J 20 ; Beekman, Marian 21
 
; Dose Janina 16 ; Amin Najaf 9 ; Arnold, Alice M 17 ; Ayers, Kristin L 18 ; Barzilai Nir 19 ; Becker, Elizabeth J 20 ; Beekman, Marian 21  
 ; Blanché Hélène 22
 
; Blanché Hélène 22  
 ; Christensen, Kaare 23
 
; Christensen, Kaare 23  
 ; Christiansen, Lene 24 ; Collerton, Joanna C 25 ; Cubaynes, Sarah 26 ; Cummings, Steven R 2 ; Davies, Karen 27 ; Debrabant Birgit 28 ; Deleuze Jean-François 29 ; Duncan, Rachel 30 ; Faul, Jessica D 11 ; Franceschi, Claudio 31 ; Galan, Pilar 32
 
; Christiansen, Lene 24 ; Collerton, Joanna C 25 ; Cubaynes, Sarah 26 ; Cummings, Steven R 2 ; Davies, Karen 27 ; Debrabant Birgit 28 ; Deleuze Jean-François 29 ; Duncan, Rachel 30 ; Faul, Jessica D 11 ; Franceschi, Claudio 31 ; Galan, Pilar 32  
 ; Vilmundur, Gudnason 33
 
; Vilmundur, Gudnason 33  
 ; Harris, Tamara B 34 ; Huisman Martijn 35 ; Hurme, Mikko A 36 ; Jagger, Carol 30
 
; Harris, Tamara B 34 ; Huisman Martijn 35 ; Hurme, Mikko A 36 ; Jagger, Carol 30  
 ; Jansen, Iris 37 ; Jylhä Marja 38 ; Kähönen Mika 39 ; Karasik, David 40
 
; Jansen, Iris 37 ; Jylhä Marja 38 ; Kähönen Mika 39 ; Karasik, David 40  
 ; Kardia, Sharon L, R 41 ; Kingston, Andrew 30 ; Kirkwood Thomas B L 42 ; Launer, Lenore J 34 ; Lehtimäki Terho 14 ; Lieb, Wolfgang 43 ; Leo-Pekka, Lyytikäinen 14 ; Martin-Ruiz, Carmen 27
 
; Kardia, Sharon L, R 41 ; Kingston, Andrew 30 ; Kirkwood Thomas B L 42 ; Launer, Lenore J 34 ; Lehtimäki Terho 14 ; Lieb, Wolfgang 43 ; Leo-Pekka, Lyytikäinen 14 ; Martin-Ruiz, Carmen 27  
 ; Min Junxia 44 ; Nebel Almut 16 ; Newman, Anne B 45
 
; Min Junxia 44 ; Nebel Almut 16 ; Newman, Anne B 45  
 ; Nie Chao 46 ; Nohr, Ellen A 47 ; Orwoll, Eric S 48
 
; Nie Chao 46 ; Nohr, Ellen A 47 ; Orwoll, Eric S 48  
 ; Perls, Thomas T 49 ; Province, Michael A 7 ; Psaty, Bruce M 50 ; Raitakari, Olli T 51 ; Reinders Marcel J T 52
 
; Perls, Thomas T 49 ; Province, Michael A 7 ; Psaty, Bruce M 50 ; Raitakari, Olli T 51 ; Reinders Marcel J T 52  
 ; Robine Jean-Marie 53
 
; Robine Jean-Marie 53  
 ; Rotter, Jerome I 54
 
; Rotter, Jerome I 54  
 ; Sebastiani, Paola 12 ; Smith, Jennifer 55
 
; Sebastiani, Paola 12 ; Smith, Jennifer 55  
 ; Sørensen Thorkild I A 56
 
; Sørensen Thorkild I A 56  
 ; Taylor, Kent D 57
 
; Taylor, Kent D 57  
 ; Uitterlinden, André G 58
 
; Uitterlinden, André G 58  
 ; van der Flier Wiesje 59 ; van der Lee Sven J 60
 
; van der Flier Wiesje 59 ; van der Lee Sven J 60  
 ; van Duijn Cornelia M 61 ; Diana, van Heemst 62 ; Vaupel, James W 63 ; Weir, David 11
 
; van Duijn Cornelia M 61 ; Diana, van Heemst 62 ; Vaupel, James W 63 ; Weir, David 11  
 ; Ye, Kenny 64
 
; Ye, Kenny 64  
 ; Zeng, Yi 65 ; Zheng Wanlin 2 ; Henne, Holstege 4 ; Kiel, Douglas P 66
 
; Zeng, Yi 65 ; Zheng Wanlin 2 ; Henne, Holstege 4 ; Kiel, Douglas P 66  
 ; Lunetta, Kathryn L 12
 
; Lunetta, Kathryn L 12  
 ; Eline, Slagboom P 21 ; Murabito, Joanne M 67
 
; Eline, Slagboom P 21 ; Murabito, Joanne M 67 1 Max Planck Institute for Biology of Ageing, Cologne, Germany (GRID:grid.419502.b) (ISNI:0000 0004 0373 6590); Leiden University Medical Center, Molecular Epidemiology, Department of Biomedical Data Sciences, Leiden, The Netherlands (GRID:grid.10419.3d) (ISNI:0000000089452978)
2 California Pacific Medical Center Research Institute, San Francisco, USA (GRID:grid.17866.3e) (ISNI:0000000098234542)
3 Johns Hopkins University School of Medicine, McKusick-Nathans Institute of Genetic Medicine, Department of Genetic Medicine, Baltimore, USA (GRID:grid.21107.35) (ISNI:0000 0001 2171 9311)
4 Vrije Universiteit Amsterdam, Amsterdam UMC, Alzheimer Center Amsterdam, Department of Neurology, Amsterdam Neuroscience, Amsterdam, The Netherlands (GRID:grid.12380.38) (ISNI:0000 0004 1754 9227); Amsterdam UMC, Department of Clinical Genetics, Amsterdam, The Netherlands (GRID:grid.509540.d) (ISNI:0000 0004 6880 3010); Delft University of Technology, Delft Bioinformatics Lab, Delft, The Netherlands (GRID:grid.5292.c) (ISNI:0000 0001 2097 4740)
5 University of Southern Denmark, The Danish Aging Research Center, Department of Public Health, Odense C, Denmark (GRID:grid.10825.3e) (ISNI:0000 0001 0728 0170)
6 BGI-Shenzhen, Shenzhen, China (GRID:grid.21155.32) (ISNI:0000 0001 2034 1839); China National Genebank, BGI-Shenzhen, Shenzhen, China (GRID:grid.21155.32) (ISNI:0000 0001 2034 1839)
7 Washington University School of Medicine, Division of Statistical Genomics, Department of Genetics, Saint Louis, USA (GRID:grid.4367.6) (ISNI:0000 0001 2355 7002)
8 University of Washington, Department of Biostatistics, Seattle, USA (GRID:grid.34477.33) (ISNI:0000000122986657); University of Washington, Cardiovascular Health Research Unit, Department of Medicine, Seattle, USA (GRID:grid.34477.33) (ISNI:0000000122986657)
9 Erasmus MC, Department of Epidemiology, Rotterdam, The Netherlands (GRID:grid.5645.2) (ISNI:000000040459992X)
10 University of Haifa, Department of Biology, Faculty of Natural Science, Haifa, Israel (GRID:grid.18098.38) (ISNI:0000 0004 1937 0562); Albert Einstein College of Medicine, Departments of Medicine and Genetics, Bronx, USA (GRID:grid.251993.5) (ISNI:0000000121791997)
11 University of Michigan, Institute for Social Research, Survey Research Center, Ann Arbor, USA (GRID:grid.214458.e) (ISNI:0000000086837370)
12 Boston University School of Public Health, Department of Biostatistics, Boston, USA (GRID:grid.189504.1) (ISNI:0000 0004 1936 7558)
13 University of Michigan, School of Public Health, Department of Biostatistics, Ann Arbor, USA (GRID:grid.214458.e) (ISNI:0000000086837370); Icelandic Heart Association, Kópavogur, Iceland (GRID:grid.420802.c) (ISNI:0000 0000 9458 5898)
14 Tampere University, Department of Clinical Chemistry, Fimlab Laboratories and Finnish Cardiovascular Research Center—Tampere, Faculty of Medicine and Health Technology, Tampere, Finland (GRID:grid.502801.e) (ISNI:0000 0001 2314 6254)
15 Newcastle University, Institute of Genetic Medicine, Newcastle upon Tyne, UK (GRID:grid.1006.7) (ISNI:0000 0001 0462 7212)
16 Kiel University, Institute of Clinical Molecular Biology, Kiel, Germany (GRID:grid.9764.c) (ISNI:0000 0001 2153 9986)
17 University of Washington, Department of Biostatistics, Seattle, USA (GRID:grid.34477.33) (ISNI:0000000122986657)
18 Sema4, a Mount Sinai venture, Stamford, USA (GRID:grid.416167.3)
19 Albert Einstein College of Medicine, Departments of Medicine and Genetics, Bronx, USA (GRID:grid.251993.5) (ISNI:0000000121791997)
20 Boston University, Bioinformatics Program, Boston, USA (GRID:grid.189504.1) (ISNI:0000 0004 1936 7558)
21 Leiden University Medical Center, Molecular Epidemiology, Department of Biomedical Data Sciences, Leiden, The Netherlands (GRID:grid.10419.3d) (ISNI:0000000089452978)
22 Fondation Jean Dausset—CEPH, Paris, France (GRID:grid.417836.f) (ISNI:0000 0004 0639 125X)
23 University of Southern Denmark, The Danish Aging Research Center, Department of Public Health, Odense C, Denmark (GRID:grid.10825.3e) (ISNI:0000 0001 0728 0170); Odense University Hospital, Clinical Biochemistry and Pharmacology, Odense C, Denmark (GRID:grid.7143.1) (ISNI:0000 0004 0512 5013); Odense University Hospital, Department of Clinical Genetics, Odense C, Denmark (GRID:grid.7143.1) (ISNI:0000 0004 0512 5013)
24 University of Southern Denmark, The Danish Aging Research Center, Department of Public Health, Odense C, Denmark (GRID:grid.10825.3e) (ISNI:0000 0001 0728 0170); Copenhagen University Hospital, Rigshospitalet, Department of Clinical Immunology, Copenhagen, Denmark (GRID:grid.475435.4)
25 Newcastle University, Institute of Health & Society, Newcastle upon Tyne, UK (GRID:grid.1006.7) (ISNI:0000 0001 0462 7212)
26 MMDN, Univ. Montpellier, EPHE, Unité Inserm 1198, PSL Research University, Montpellier, France (GRID:grid.121334.6) (ISNI:0000 0001 2097 0141)
27 Newcastle University, Institute of Neuroscience, Newcastle upon Tyne, UK (GRID:grid.1006.7) (ISNI:0000 0001 0462 7212)
28 University of Southern Denmark, Department of Public Health, Odense C, Denmark (GRID:grid.10825.3e) (ISNI:0000 0001 0728 0170)
29 Fondation Jean Dausset—CEPH, Paris, France (GRID:grid.417836.f) (ISNI:0000 0004 0639 125X); Centre National de Recherche en Génomique Humaine, CEA-Institut de Biologie François Jacob, Evry, France (GRID:grid.418135.a) (ISNI:0000 0004 0641 3404)
30 Newcastle University, Institute of Health & Society, Newcastle upon Tyne, UK (GRID:grid.1006.7) (ISNI:0000 0001 0462 7212); Newcastle University, Newcastle University Institute for Ageing, Newcastle upon Tyne, UK (GRID:grid.1006.7) (ISNI:0000 0001 0462 7212)
31 Lobachevsky State University of Nizhny Novgorod, Department of Applied Mathematics and Centre of Bioinformatics, Nizhny Novgorod, Russia (GRID:grid.28171.3d) (ISNI:0000 0001 0344 908X); IRCCS Institute of Neurological Sciences of Bologna (ISNB), Bologna, Italy (GRID:grid.492077.f)
32 EREN, UMR U1153 Inserm/U1125 Inra/Cnam/Paris 13, Université Paris 13, CRESS, Bobigny, France (GRID:grid.11318.3a) (ISNI:0000000121496883)
33 Icelandic Heart Association, Kópavogur, Iceland (GRID:grid.420802.c) (ISNI:0000 0000 9458 5898); University of Iceland, Faculty of Medicine, Reykjavik, Iceland (GRID:grid.14013.37) (ISNI:0000 0004 0640 0021)
34 National Institute on Aging, NIH, Laboratory of Epidemiology and Population Sciences, Bethesda, USA (GRID:grid.419475.a) (ISNI:0000 0000 9372 4913)
35 Vrije Universiteit Amsterdam, Amsterdam UMC, Department of Epidemiology and Biostatistics, Amsterdam, The Netherlands (GRID:grid.12380.38) (ISNI:0000 0004 1754 9227); Amsterdam Public Health Research Institute, Amsterdam, The Netherlands (GRID:grid.16872.3a) (ISNI:0000 0004 0435 165X)
36 Tampere University, Department of Microbiology and Immunology, Faculty of Medicine and Health Technology, Tampere, Finland (GRID:grid.502801.e) (ISNI:0000 0001 2314 6254)
37 Vrije Universiteit Amsterdam, Amsterdam UMC, Alzheimer Center Amsterdam, Department of Neurology, Amsterdam Neuroscience, Amsterdam, The Netherlands (GRID:grid.12380.38) (ISNI:0000 0004 1754 9227); Vrije Universiteit Amsterdam, Department of Complex Trait Genetics, Center for Neurogenomics and Cognitive Research, Amsterdam, The Netherlands (GRID:grid.12380.38) (ISNI:0000 0004 1754 9227)
38 Tampere University, Faculty of Social Sciences (Health Sciences) and Gerontology Research Center (GEREC), Tampere, Finland (GRID:grid.502801.e) (ISNI:0000 0001 2314 6254)
39 Tampere University, Department of Clinical Physiology, Tampere University Hospital and Finnish Cardiovascular Research Center—Tampere, Faculty of Medicine and Health Technology, Tampere, Finland (GRID:grid.502801.e) (ISNI:0000 0001 2314 6254)
40 Azrieli Faculty of Medicine, Bar Ilan University, Safed, Israel (GRID:grid.22098.31) (ISNI:0000 0004 1937 0503); Hinda and Arthur Marcus Institute for Aging Research, Hebrew SeniorLife, Boston, USA (GRID:grid.497274.b) (ISNI:0000 0004 0627 5136)
41 University of Michigan, School of Public Health, Epidemiology, Ann Arbor, USA (GRID:grid.214458.e) (ISNI:0000000086837370)
42 Newcastle University, Newcastle University Institute for Ageing, Newcastle upon Tyne, UK (GRID:grid.1006.7) (ISNI:0000 0001 0462 7212)
43 Kiel University, Institute of Epidemiology and Biobank PopGen, Kiel, Germany (GRID:grid.9764.c) (ISNI:0000 0001 2153 9986)
44 Zhejiang University, Institute of Translational Medicine, School of Medicine, Hangzhou, China (GRID:grid.13402.34) (ISNI:0000 0004 1759 700X)
45 University of Pittsburgh, Department of Epidemiology, Graduate School of Public Health, Pittsburgh, USA (GRID:grid.21925.3d) (ISNI:0000 0004 1936 9000)
46 BGI-Shenzhen, Shenzhen, China (GRID:grid.21155.32) (ISNI:0000 0001 2034 1839)
47 University of Southern Denmark, Research Unit of Gynecology and Obstetrics, Department of Clinical Research, Odense C, Denmark (GRID:grid.10825.3e) (ISNI:0000 0001 0728 0170)
48 Oregon Health Sciences University, Bone and Mineral Unit, Portland, USA (GRID:grid.5288.7) (ISNI:0000 0000 9758 5690)
49 Boston University School of Medicine, Department of Medicine, Geriatrics Section, Boston Medical Center, Boston, USA (GRID:grid.189504.1) (ISNI:0000 0004 1936 7558)
50 University of Washington, Cardiovascular Health Research Unit, Department of Medicine, Seattle, USA (GRID:grid.34477.33) (ISNI:0000000122986657); University of Washington, Department of Epidemiology, Seattle, USA (GRID:grid.34477.33) (ISNI:0000000122986657); University of Washington, Department of Health Services, Seattle, USA (GRID:grid.34477.33) (ISNI:0000000122986657); Kaiser Permanente Washington Health Research Institute, Seattle, USA (GRID:grid.488833.c) (ISNI:0000 0004 0615 7519)
51 Turku University Hospital, Department of Clinical Physiology and Nuclear Medicine, Turku, Finland (GRID:grid.410552.7) (ISNI:0000 0004 0628 215X); University of Turku, Research Centre of Applied and Preventive Cardiovascular Medicine, Turku, Finland (GRID:grid.1374.1) (ISNI:0000 0001 2097 1371)
52 Delft University of Technology, Delft Bioinformatics Lab, Delft, The Netherlands (GRID:grid.5292.c) (ISNI:0000 0001 2097 4740)
53 MMDN, Univ. Montpellier, EPHE, Unité Inserm 1198, PSL Research University, Montpellier, France (GRID:grid.121334.6) (ISNI:0000 0001 2097 0141); CERMES3, UMR CNRS 8211—Unité Inserm 988—EHESS—Université Paris Descartes, Paris, France (GRID:grid.469410.e) (ISNI:0000 0001 0429 0824)
54 Los Angeles Biomedical Research Institute at Harbor-UCLA Medical Center, Institute for Translational Genomics and Population Sciences, Torrance, USA (GRID:grid.279946.7) (ISNI:0000 0004 0521 0744); Harbor-UCLA Medical Center, Division of Genetic Outcomes, Department of Pediatrics, Torrance, USA (GRID:grid.239844.0) (ISNI:0000 0001 0157 6501)
55 University of Michigan, Institute for Social Research, Survey Research Center, Ann Arbor, USA (GRID:grid.214458.e) (ISNI:0000000086837370); University of Michigan, School of Public Health, Epidemiology, Ann Arbor, USA (GRID:grid.214458.e) (ISNI:0000000086837370)
56 University of Copenhagen, Novo Nordisk Foundation Center for Basic Metabolic Research, Section of Metabolic Genetics, and Department of Public Health, Section of Epidemiology, Faculty of Health and Medical Sciences, Copenhagen N, Denmark (GRID:grid.5254.6) (ISNI:0000 0001 0674 042X); Bristol University, MRC Integrative Epidemiology Unit, Bristol, UK (GRID:grid.5337.2) (ISNI:0000 0004 1936 7603)
57 Los Angeles Biomedical Research Institute at Harbor-UCLA Medical Center, Institute for Translational Genomics and Population Sciences, Torrance, USA (GRID:grid.279946.7) (ISNI:0000 0004 0521 0744); Harbor-UCLA Medical Center, Department of Pediatrics, Torrance, USA (GRID:grid.239844.0) (ISNI:0000 0001 0157 6501)
58 Erasmus MC, Department of Epidemiology, Rotterdam, The Netherlands (GRID:grid.5645.2) (ISNI:000000040459992X); Erasmus MC, Department of Internal Medicine, Rotterdam, The Netherlands (GRID:grid.5645.2) (ISNI:000000040459992X)
59 Vrije Universiteit Amsterdam, Amsterdam UMC, Alzheimer Center Amsterdam, Department of Neurology, Amsterdam Neuroscience, Amsterdam, The Netherlands (GRID:grid.12380.38) (ISNI:0000 0004 1754 9227); Vrije Universiteit Amsterdam, Amsterdam UMC, Department of Epidemiology and Biostatistics, Amsterdam, The Netherlands (GRID:grid.12380.38) (ISNI:0000 0004 1754 9227)
60 Vrije Universiteit Amsterdam, Amsterdam UMC, Alzheimer Center Amsterdam, Department of Neurology, Amsterdam Neuroscience, Amsterdam, The Netherlands (GRID:grid.12380.38) (ISNI:0000 0004 1754 9227); Amsterdam UMC, Department of Clinical Genetics, Amsterdam, The Netherlands (GRID:grid.509540.d) (ISNI:0000 0004 6880 3010)
61 Erasmus MC, Department of Epidemiology, Rotterdam, The Netherlands (GRID:grid.5645.2) (ISNI:000000040459992X); Nuffield Department of Population Health, University of Oxford, Oxford, UK (GRID:grid.4991.5) (ISNI:0000 0004 1936 8948)
62 Leiden University Medical Center, Department of Gerontology and Geriatrics, Leiden, The Netherlands (GRID:grid.10419.3d) (ISNI:0000000089452978)
63 Max Planck Institute for Demographic Research, Rostock, Germany (GRID:grid.419511.9) (ISNI:0000 0001 2033 8007)
64 Albert Einstein College of Medicine, Department of Epidemiology and Population Health, Bronx, USA (GRID:grid.251993.5) (ISNI:0000000121791997)
65 Peking University, Center for Healthy Aging and Development Studies, National School of Development and Raissun Institute for Advanced Studies, Beijing, China (GRID:grid.11135.37) (ISNI:0000 0001 2256 9319); Medical School of Duke University, Center for the Study of Aging and Human Development and Geriatrics Division, Durham, USA (GRID:grid.26009.3d) (ISNI:0000 0004 1936 7961)
66 Hinda and Arthur Marcus Institute for Aging Research, Hebrew SeniorLife, Boston, USA (GRID:grid.497274.b) (ISNI:0000 0004 0627 5136); Beth Israel Deaconess Medical Center and Harvard Medical School, Department of Medicine, Boston, USA (GRID:grid.239395.7) (ISNI:0000 0000 9011 8547); Broad Institute of MIT & Harvard, Cambridge, USA (GRID:grid.66859.34)
67 NHLBI’s and Boston University’s Framingham Heart Study, Framingham, USA (GRID:grid.189504.1) (ISNI:0000 0004 1936 7558); Boston University School of Medicine, Section of General Internal Medicine, Department of Medicine, Boston, USA (GRID:grid.189504.1) (ISNI:0000 0004 1936 7558)




