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Abstract
Atopic dermatitis (AD) is a common inflammatory skin condition and prior genome-wide association studies (GWAS) have identified 71 associated loci. In the current study we conducted the largest AD GWAS to date (discovery N = 1,086,394, replication N = 3,604,027), combining previously reported cohorts with additional available data. We identified 81 loci (29 novel) in the European-only analysis (which all replicated in a separate European analysis) and 10 additional loci in the multi-ancestry analysis (3 novel). Eight variants from the multi-ancestry analysis replicated in at least one of the populations tested (European, Latino or African), while two may be specific to individuals of Japanese ancestry. AD loci showed enrichment for DNAse I hypersensitivity and eQTL associations in blood. At each locus we prioritised candidate genes by integrating multi-omic data. The implicated genes are predominantly in immune pathways of relevance to atopic inflammation and some offer drug repurposing opportunities.
The genetic basis of atopic dermatitis is not fully understood. Here, the authors find 91 genetic loci associated with atopic dermatitis in a GWAS of >1million individuals, which highlight the importance of systemic immune regulation.
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1 University of Bristol, Medical Research Council Integrative Epidemiology Unit, Bristol Medical School, Bristol, England (GRID:grid.5337.2) (ISNI:0000 0004 1936 7603); University of Bristol, Population Health Sciences, Bristol Medical School, Bristol, England (GRID:grid.5337.2) (ISNI:0000 0004 1936 7603)
2 Helmholtz Zentrum München - German Research Center for Environmental Health, Institute of Epidemiology, Neuherberg, Germany (GRID:grid.4567.0) (ISNI:0000 0004 0483 2525); University of Munich Medical Center, Division of Metabolic and Nutritional Medicine, Dr. von Hauner Children’s Hospital, Munich, Germany (GRID:grid.5252.0) (ISNI:0000 0004 1936 973X); Ludwig-Maximilians University Munich, Pettenkofer School of Public Health, Munich, Germany (GRID:grid.5252.0) (ISNI:0000 0004 1936 973X)
3 23andMe, Inc., Sunnyvale, USA (GRID:grid.420283.f) (ISNI:0000 0004 0626 0858)
4 University of Tartu, Estonian Genome Centre, Institute of Genomics, Tartu, Estonia (GRID:grid.10939.32) (ISNI:0000 0001 0943 7661)
5 University of Tartu, Estonian Genome Centre, Institute of Genomics, Tartu, Estonia (GRID:grid.10939.32) (ISNI:0000 0001 0943 7661); University of Tartu, Core Facility of Genomics, Tartu, Estonia (GRID:grid.10939.32) (ISNI:0000 0001 0943 7661)
6 RIKEN Center for Integrative Medical Sciences, Laboratory for Statistical and Translational Genetics, Yokohama, Japan (GRID:grid.509459.4) (ISNI:0000 0004 0472 0267); Tokyo Medical and Dental University (TMDU), Department of Rheumatology, Graduate School of Medical and Dental Sciences, Tokyo, Japan (GRID:grid.265073.5) (ISNI:0000 0001 1014 9130)
7 Norwegian University of Science and Technology, K.G. Jebsen Center for Genetic Epidemiology, Department of Public Health and Nursing, NTNU, Trondheim, Norway (GRID:grid.5947.f) (ISNI:0000 0001 1516 2393); Norwegian University of Science and Technology, HUNT Research Centre, Department of Public Health and Nursing, NTNU, Levanger, Norway (GRID:grid.5947.f) (ISNI:0000 0001 1516 2393); Trondheim University Hospital, Clinic of Medicine, St. Olavs Hospital, Trondheim, Norway (GRID:grid.52522.32) (ISNI:0000 0004 0627 3560)
8 Norwegian University of Science and Technology, K.G. Jebsen Center for Genetic Epidemiology, Department of Public Health and Nursing, NTNU, Trondheim, Norway (GRID:grid.5947.f) (ISNI:0000 0001 1516 2393); NTNU Norwegian University of Science and Technology, Department of Clinical and Molecular Medicine, Trondheim, Norway (GRID:grid.5947.f) (ISNI:0000 0001 1516 2393); Norwegian University of Science and Technology, BioCore - Bioinformatics Core Facility, NTNU, Trondheim, Norway (GRID:grid.5947.f) (ISNI:0000 0001 1516 2393); Trondheim University Hospital, Clinic of Laboratory Medicine, St. Olavs Hospital, Trondheim, Norway (GRID:grid.52522.32) (ISNI:0000 0004 0627 3560)
9 University of Gothenburg, Department of Obstetrics and Gynecology, Institute of Clinical Sciences, Sahlgrenska Academy, Gothenburg, Sweden (GRID:grid.8761.8) (ISNI:0000 0000 9919 9582)
10 Universidad de La Laguna, Genomics and Health Group, Department of Biochemistry, Microbiology, Cell Biology and Genetics, La Laguna, Spain (GRID:grid.10041.34) (ISNI:0000 0001 2106 0879)
11 University of Copenhagen, Novo Nordisk Foundation Center for Basic Metabolic Research, Faculty of Health and Medical Sciences, København, Denmark (GRID:grid.5254.6) (ISNI:0000 0001 0674 042X)
12 Max-Delbrück-Center for Molecular Medicine, Berlin, Germany (GRID:grid.419491.0) (ISNI:0000 0001 1014 0849); Charité-Universitätsmedizin Berlin, Clinic for Pediatric Allergy, Experimental and Clinical Research Center, Berlin, Germany (GRID:grid.6363.0) (ISNI:0000 0001 2218 4662)
13 Swiss Tropical and Public Health Institute, Basel, Switzerland (GRID:grid.416786.a) (ISNI:0000 0004 0587 0574); University of Basel, Basel, Switzerland (GRID:grid.6612.3) (ISNI:0000 0004 1937 0642)
14 University of Leicester, Department of Health Sciences, Leicester, UK (GRID:grid.9918.9) (ISNI:0000 0004 1936 8411)
15 University of Regensburg, Department of Epidemiology and Preventive Medicine, Regensburg, Germany (GRID:grid.7727.5) (ISNI:0000 0001 2190 5763)
16 University Hospital Schleswig-Holstein, Department of Dermatology and Allergy, Kiel, Germany (GRID:grid.412468.d) (ISNI:0000 0004 0646 2097)
17 University of Surrey, School of Biosciences and Medicine, Guildford, UK (GRID:grid.5475.3) (ISNI:0000 0004 0407 4824)
18 Karolinska Institutet, Department of Clinical Science and Education Södersjukhuset, Solna, Sweden (GRID:grid.4714.6) (ISNI:0000 0004 1937 0626)
19 Children’s Hospital of Philadelphia, Center for Applied Genomics, Philadelphia, USA (GRID:grid.239552.a) (ISNI:0000 0001 0680 8770); University of Pennsylvania, Department of Genetics, Perelman School of Medicine, Philadelphia, USA (GRID:grid.25879.31) (ISNI:0000 0004 1936 8972); Rhythm Pharmaceuticals, Boston, USA (GRID:grid.476681.a)
20 Children’s Hospital of Philadelphia, Center for Applied Genomics, Philadelphia, USA (GRID:grid.239552.a) (ISNI:0000 0001 0680 8770)
21 University Medical Center Rotterdam, The Generation R Study Group, Erasmus MC, Rotterdam, The Netherlands (GRID:grid.5645.2) (ISNI:0000 0004 0459 992X); University Medical Center Rotterdam, Department of Internal Medicine, Erasmus MC, Rotterdam, The Netherlands (GRID:grid.5645.2) (ISNI:0000 0004 0459 992X)
22 University Medical Center Rotterdam, The Generation R Study Group, Erasmus MC, Rotterdam, The Netherlands (GRID:grid.5645.2) (ISNI:0000 0004 0459 992X); University Medical Center Rotterdam, Department of Dermatology, Erasmus MC, Rotterdam, The Netherlands (GRID:grid.5645.2) (ISNI:0000 0004 0459 992X)
23 University of Groningen, University Medical Center Groningen, Department of Pediatric Pulmonology and Pediatric Allergy, Beatrix Children’s Hospital, Groningen, The Netherlands (GRID:grid.4494.d) (ISNI:0000 0000 9558 4598); University of Groningen, University Medical Center Groningen, GRIAC Research Institute, Groningen, The Netherlands (GRID:grid.4494.d) (ISNI:0000 0000 9558 4598); CiiM, a joint venture between Hannover Medical School and the Helmholtz Centre for Infection Research, Centre for Individualized Infection Medicine, Hannover, Germany (GRID:grid.512472.7); Centre for Experimental and Clinical Infection Research, a joint venture between the Hannover Medical School and the Helmholtz Centre for Infection Research, TWINCORE, Hannover, Germany (GRID:grid.452370.7) (ISNI:0000 0004 0408 1805)
24 University of Groningen, University Medical Center Groningen, Department of Pediatric Pulmonology and Pediatric Allergy, Beatrix Children’s Hospital, Groningen, The Netherlands (GRID:grid.4494.d) (ISNI:0000 0000 9558 4598); University of Groningen, University Medical Center Groningen, GRIAC Research Institute, Groningen, The Netherlands (GRID:grid.4494.d) (ISNI:0000 0000 9558 4598)
25 University of Southampton, MRC Lifecourse Epidemiology Centre, Southampton, UK (GRID:grid.5491.9) (ISNI:0000 0004 1936 9297)
26 University of Southampton, Faculty of Medicine, Southampton, UK (GRID:grid.5491.9) (ISNI:0000 0004 1936 9297); University of Southampton, Institute of Developmental Sciences, Southampton, UK (GRID:grid.5491.9) (ISNI:0000 0004 1936 9297)
27 Department of Bioinformatics & Data Mining, Måløv, Denmark (GRID:grid.5491.9); Statens Serum Institut, Department of Epidemiology Research, Copenhagen, Denmark (GRID:grid.6203.7) (ISNI:0000 0004 0417 4147); Lund University Diabetes Centre, Department of Clinical Sciences, Malmö, Sweden (GRID:grid.4514.4) (ISNI:0000 0001 0930 2361); University of Helsinki, Institute for Molecular Medicine Finland (FIMM), Helsinki, Finland (GRID:grid.7737.4) (ISNI:0000 0004 0410 2071)
28 University of Newcastle, School of Medicine and Public Health, Newcastle, Australia (GRID:grid.266842.c) (ISNI:0000 0000 8831 109X); Hunter Medical Research Institute, Newcastle, Australia (GRID:grid.413648.c)
29 Helmholtz Zentrum München - German Research Center for Environmental Health, Institute of Epidemiology, Neuherberg, Germany (GRID:grid.4567.0) (ISNI:0000 0004 0483 2525); University of Munich Medical Center, Division of Metabolic and Nutritional Medicine, Dr. von Hauner Children’s Hospital, Munich, Germany (GRID:grid.5252.0) (ISNI:0000 0004 1936 973X)
30 Henry Ford Health, Center for Individualized and Genomic Medicine Research (CIGMA), Department of Medicine, Detroit, USA (GRID:grid.239864.2) (ISNI:0000 0000 8523 7701)
31 Brain and Skin Aging, IUF – Leibniz Research Institute for Environmental Medicine, Environmental Epidemiology of Lung, Düsseldorf, Germany (GRID:grid.435557.5) (ISNI:0000 0004 0518 6318)
32 University of Southampton, Human Development and Health, Faculty of Medicine, Southampton, UK (GRID:grid.5491.9) (ISNI:0000 0004 1936 9297); University Hospital Southampton, NIHR Southampton Biomedical Research Centre, Southampton, UK (GRID:grid.123047.3) (ISNI:0000000103590315)
33 University of Southampton, Human Development and Health, Faculty of Medicine, Southampton, UK (GRID:grid.5491.9) (ISNI:0000 0004 1936 9297)
34 University of Memphis, Division of Epidemiology, Biostatistics, and Environmental Health, School of Public Health, Memphis, USA (GRID:grid.56061.34) (ISNI:0000 0000 9560 654X)
35 University of Cape Town, Division of Immunology, Department of Pathology, Faculty of Health Sciences, Cape Town, South Africa (GRID:grid.7836.a) (ISNI:0000 0004 1937 1151)
36 Institute for Human Genetics, UCSF, San Francisco, USA (GRID:grid.266102.1) (ISNI:0000 0001 2297 6811); UCSF, Department of Epidemiology and Biostatistics, San Francisco, USA (GRID:grid.266102.1) (ISNI:0000 0001 2297 6811)
37 Regeneron Genetics Center, Tarrytown, USA (GRID:grid.418961.3) (ISNI:0000 0004 0472 2713)
38 Kaiser Permanente Northern California, Division of Research, Oakland, USA (GRID:grid.280062.e) (ISNI:0000 0000 9957 7758)
39 University of Bergen, Center for Diabetes Research, Department of Clinical Science, Bergen, Norway (GRID:grid.7914.b) (ISNI:0000 0004 1936 7443); Haukeland University Hospital, Children and Youth Clinic, Bergen, Norway (GRID:grid.412008.f) (ISNI:0000 0000 9753 1393)
40 University of Oslo, NORMENT Centre, Institute of Clinical Medicine, Oslo, Norway (GRID:grid.5510.1) (ISNI:0000 0004 1936 8921); Oslo University Hospital, Division of Mental Health and Addiction, Oslo, Norway (GRID:grid.55325.34) (ISNI:0000 0004 0389 8485)
41 University of Bergen, Center for Diabetes Research, Department of Clinical Science, Bergen, Norway (GRID:grid.7914.b) (ISNI:0000 0004 1936 7443); Haukeland University Hospital, Department of Medical Genetics, Bergen, Norway (GRID:grid.412008.f) (ISNI:0000 0000 9753 1393)
42 Karolinska Institutet, Department of Medical Epidemiology and Biostatistics, Stockholm, Sweden (GRID:grid.4714.6) (ISNI:0000 0004 1937 0626); Karolinska University Hospital, Pediatric Lung and Allergy Unit, Astrid Lindgren Children’s Hospital, Stockholm, Sweden (GRID:grid.24381.3c) (ISNI:0000 0000 9241 5705)
43 Karolinska Institutet, Department of Medical Epidemiology and Biostatistics, Stockholm, Sweden (GRID:grid.4714.6) (ISNI:0000 0004 1937 0626)
44 University of California San Francisco, Department of Medicine, San Francisco, USA (GRID:grid.266102.1) (ISNI:0000 0001 2297 6811); University of California San Francisco, Department of Bioengineering and Therapeutic Sciences, San Francisco, USA (GRID:grid.266102.1) (ISNI:0000 0001 2297 6811)
45 University of Copenhagen, Department of Dermatology, Bispebjerg Hospital, Copenhagen, Denmark (GRID:grid.5254.6) (ISNI:0000 0001 0674 042X)
46 Bispebjerg and Frederiksberg Hospital, Center for Clinical Research and Prevention, Frederiksberg, Denmark (GRID:grid.512917.9)
47 University Medicine Greifswald, Clinic and Polyclinic of Dermatology, Greifswald, Germany (GRID:grid.5603.0)
48 University Medicine Greifswald, Department of Functional Genomics, Interfaculty Institute for Genetics and Functional Genomics, Greifswald, Germany (GRID:grid.5603.0)
49 Charité-Universitätsmedizin Berlin, Department of Pediatric Respiratory Medicine, Immunology, and Critical Care Medicine, Berlin, Germany (GRID:grid.6363.0) (ISNI:0000 0001 2218 4662)
50 University of Bonn, School of Medicine & University Hospital Bonn, Institute of Human Genetics, Bonn, Germany (GRID:grid.10388.32) (ISNI:0000 0001 2240 3300)
51 Max-Delbrück-Center for Molecular Medicine, Berlin, Germany (GRID:grid.419491.0) (ISNI:0000 0001 1014 0849); Charite-Universitätsmedizin Berlin, Berlin, Germany (GRID:grid.6363.0) (ISNI:0000 0001 2218 4662)
52 Kings College London, Department of Twin Research & Genetics Epidemiology, London, UK (GRID:grid.13097.3c) (ISNI:0000 0001 2322 6764)
53 Kings College London, Department of Twin Research & Genetics Epidemiology, London, UK (GRID:grid.13097.3c) (ISNI:0000 0001 2322 6764); West Herts NHS Trust, Dermatology Department, Watford, UK (GRID:grid.13097.3c)
54 VU University, Dept Biological Psychology, Netherlands Twin Register, Amsterdam, the Netherlands (GRID:grid.12380.38) (ISNI:0000 0004 1754 9227); VU University, Institute for Health and Care Research (EMGO), Amsterdam, the Netherlands (GRID:grid.466632.3) (ISNI:0000 0001 0686 3219)
55 VU University, Dept Biological Psychology, Netherlands Twin Register, Amsterdam, the Netherlands (GRID:grid.12380.38) (ISNI:0000 0004 1754 9227)
56 Helmholtz Zentrum München - German Research Center for Environmental Health, Institute of Genetic Epidemiology, Neuherberg, Germany (GRID:grid.4567.0) (ISNI:0000 0004 0483 2525); LMU Munich, IBE, Faculty of Medicine, Munich, Germany (GRID:grid.5252.0) (ISNI:0000 0004 1936 973X); University Medical Center, Johannes Gutenberg University, Institute of Medical Biostatistics, Epidemiology and Informatics (IMBEI), Mainz, Germany (GRID:grid.5802.f) (ISNI:0000 0001 1941 7111)
57 University of Copenhagen, COPSAC, Copenhagen Prospective Studies on Asthma in Childhood, Herlev and Gentofte Hospital, Copenhagen, Denmark (GRID:grid.5254.6) (ISNI:0000 0001 0674 042X); Steno Diabetes Center Copenhagen, Herlev, Denmark (GRID:grid.419658.7) (ISNI:0000 0004 0646 7285); University of Copenhagen, Department of Biology, Copenhagen, Denmark (GRID:grid.5254.6) (ISNI:0000 0001 0674 042X)
58 University of Copenhagen, COPSAC, Copenhagen Prospective Studies on Asthma in Childhood, Herlev and Gentofte Hospital, Copenhagen, Denmark (GRID:grid.5254.6) (ISNI:0000 0001 0674 042X); Slagelse Hospital, Department of Pediatrics, Slagelse, Denmark (GRID:grid.452905.f)
59 University of Copenhagen, COPSAC, Copenhagen Prospective Studies on Asthma in Childhood, Herlev and Gentofte Hospital, Copenhagen, Denmark (GRID:grid.5254.6) (ISNI:0000 0001 0674 042X)
60 University of Valencia, FISABIO-University Jaume I-University of Valencia, Nursing School, Valencia, Spain (GRID:grid.5338.d) (ISNI:0000 0001 2173 938X); Joint Research Unit of Epidemiology and Environmental Health, CIBERESP, Valencia, Spain (GRID:grid.5338.d)
61 ISGlobal, Institute for Global Health, Barcelona, Spain (GRID:grid.434607.2) (ISNI:0000 0004 1763 3517); Universitat Pompeu Fabra (UPF), Barcelona, Spain (GRID:grid.5612.0) (ISNI:0000 0001 2172 2676); CIBER Epidemiología y Salud Pública, Madrid, Spain (GRID:grid.466571.7) (ISNI:0000 0004 1756 6246)
62 Brigham & Women’s Hospital and Harvard Medical School, Channing Division of Network Medicine, Boston, USA (GRID:grid.62560.37) (ISNI:0000 0004 0378 8294)
63 University of Southampton, Clinical and Experimental Sciences, Faculty of Medicine, Southampton, UK (GRID:grid.5491.9) (ISNI:0000 0004 1936 9297); David Hide Asthma and Allergy Research Centre, Isle of Wight, UK (GRID:grid.512470.5)
64 RIKEN Center for Integrative Medical Sciences, Laboratory for Statistical and Translational Genetics, Yokohama, Japan (GRID:grid.509459.4) (ISNI:0000 0004 0472 0267); Clinical Research Center, Shizuoka General Hospital, Shizuoka, Japan (GRID:grid.415804.c) (ISNI:0000 0004 1763 9927); University of Shizuoka, Department of Applied Genetics, School of Pharmaceutical Sciences, Shizuoka, Japan (GRID:grid.469280.1) (ISNI:0000 0000 9209 9298)
65 University of California San Francisco, Department of Dermatology, San Francisco, USA (GRID:grid.266102.1) (ISNI:0000 0001 2297 6811)
66 Norwegian University of Science and Technology, K.G. Jebsen Center for Genetic Epidemiology, Department of Public Health and Nursing, NTNU, Trondheim, Norway (GRID:grid.5947.f) (ISNI:0000 0001 1516 2393); Trondheim University Hospital, Department of Dermatology, Clinic of Orthopaedy, Rheumatology and Dermatology, St. Olavs Hospital, Trondheim, Norway (GRID:grid.52522.32) (ISNI:0000 0004 0627 3560)
67 Norwegian University of Science and Technology, K.G. Jebsen Center for Genetic Epidemiology, Department of Public Health and Nursing, NTNU, Trondheim, Norway (GRID:grid.5947.f) (ISNI:0000 0001 1516 2393); Norwegian University of Science and Technology, HUNT Research Centre, Department of Public Health and General Practice, Levanger, Norway (GRID:grid.5947.f) (ISNI:0000 0001 1516 2393)
68 University of Gothenburg, Department of Obstetrics and Gynecology, Institute of Clinical Sciences, Sahlgrenska Academy, Gothenburg, Sweden (GRID:grid.8761.8) (ISNI:0000 0000 9919 9582); Norwegian Institute of Public Health, Department of Genetics and Bioinformatics, Oslo, Norway (GRID:grid.418193.6) (ISNI:0000 0001 1541 4204)
69 Universidad de La Laguna, Genomics and Health Group, Department of Biochemistry, Microbiology, Cell Biology and Genetics, La Laguna, Spain (GRID:grid.10041.34) (ISNI:0000 0001 2106 0879); Instituto de Salud Carlos III, CIBER de Enfermedades Respiratorias, Madrid, Spain (GRID:grid.413448.e) (ISNI:0000 0000 9314 1427); Universidad de La Laguna, San Cristóbal de La Laguna, Instituto de Tecnologías Biomédicas (ITB), Santa Cruz de Tenerife, Spain (GRID:grid.10041.34) (ISNI:0000 0001 2106 0879)
70 University of London, Cranmer Terrace, Population Health Research Institute, St George’s, London, UK (GRID:grid.4464.2) (ISNI:0000 0001 2161 2573)
71 Bispebjerg and Frederiksberg Hospital, Center for Clinical Research and Prevention, Frederiksberg, Denmark (GRID:grid.512917.9); University of Copenhagen, Department of Clinical Medicine, Faculty of Health and Medical Sciences, Copenhagen, Denmark (GRID:grid.5254.6) (ISNI:0000 0001 0674 042X)
72 University Hospital Schleswig-Holstein, Department of Dermatology, Allergology and Venereology, Kiel, Germany (GRID:grid.412468.d) (ISNI:0000 0004 0646 2097)
73 School of Public Health,Imperial College London, Department of Epidemiology and Biostatistics, MRC-PHE Centre for Environment & Health, London, UK (GRID:grid.7445.2) (ISNI:0000 0001 2113 8111); University of Oulu, Center for Life Course Health Research, Faculty of Medicine, Oulu, Finland (GRID:grid.10858.34) (ISNI:0000 0001 0941 4873); University of Oulu, Biocenter Oulu, Oulu, Finland (GRID:grid.10858.34) (ISNI:0000 0001 0941 4873)
74 Children’s Hospital of Philadelphia, Center for Applied Genomics, Philadelphia, USA (GRID:grid.239552.a) (ISNI:0000 0001 0680 8770); University of Pennsylvania, Department of Pediatrics, Divisions of Human Genetics and Pulmonary Medicine, Perelman School of Medicine, Philadelphia, USA (GRID:grid.25879.31) (ISNI:0000 0004 1936 8972); University of Iceland, Faculty of Medicine, Reykjavík, Iceland (GRID:grid.14013.37) (ISNI:0000 0004 0640 0021)
75 Trinity College, Department of Clinical Medicine, Dublin, Ireland (GRID:grid.8217.c) (ISNI:0000 0004 1936 9705)
76 Imperial College London, Respiratory Epidemiology, Occupational Medicine and Public Health, National Heart and Lung Institute, London, United Kingdom (GRID:grid.7445.2) (ISNI:0000 0001 2113 8111); Medical Research Council and Public Health England Centre for Environment and Health, London, United Kingdom (GRID:grid.14105.31) (ISNI:0000000122478951)
77 University Medical Center Rotterdam, Department of Dermatology, Erasmus MC, Rotterdam, The Netherlands (GRID:grid.5645.2) (ISNI:0000 0004 0459 992X)
78 University Medical Center Rotterdam, Department of Pediatrics, division of Respiratory Medicine and Allergology, Erasmus MC, Rotterdam, The Netherlands (GRID:grid.5645.2) (ISNI:0000 0004 0459 992X); University Medical Center Rotterdam, Department of Pediatrics, division of Neonatology, Erasmus MC, Rotterdam, The Netherlands (GRID:grid.5645.2) (ISNI:0000 0004 0459 992X)
79 University of Groningen, University Medical Center Groningen, GRIAC Research Institute, Groningen, The Netherlands (GRID:grid.4494.d) (ISNI:0000 0000 9558 4598); University of Groningen, University Medical Center Groningen, Department of Epidemiology, Groningen, The Netherlands (GRID:grid.4494.d) (ISNI:0000 0000 9558 4598)
80 University of Southampton and University Hospital Southampton NHS Foundation Trust, MRC Lifecourse Epidemiology Centre and NIHR Southampton Biomedical Research Centre, Southampton, UK (GRID:grid.430506.4) (ISNI:0000 0004 0465 4079)
81 Statens Serum Institut, Department of Epidemiology Research, Copenhagen, Denmark (GRID:grid.6203.7) (ISNI:0000 0004 0417 4147)
82 The University of Queensland, Children’s Health and Environment Program, Child Health Research Centre, South Brisbane, Australia (GRID:grid.1003.2) (ISNI:0000 0000 9320 7537); The University of Queensland, Australian Infectious Diseases Research Centre, St Lucia, Australia (GRID:grid.1003.2) (ISNI:0000 0000 9320 7537)
83 University of Western Australia, Telethon Kids Institute, Perth, Australia (GRID:grid.1012.2) (ISNI:0000 0004 1936 7910)
84 The University of Manchester, Manchester Academic Health Science Centre, and Manchester University NHS Foundation Trust, Division of Immunology, Immunity to Infection and Respiratory Medicine, School of Biological Sciences, Manchester, England (GRID:grid.498924.a) (ISNI:0000 0004 0430 9101)
85 Leibniz Research Institute for Environmental Medicine, Environmental Epidemiology of Lung, Brain and Skin Aging, Düsseldorf, Germany (GRID:grid.435557.5) (ISNI:0000 0004 0518 6318)
86 Icahn School of Medicine at Mount Sinai, Division of Allergy and Immunology, Department of Pediatrics, and Department of Genetics and Genomic Sciences, New York, USA (GRID:grid.59734.3c) (ISNI:0000 0001 0670 2351)
87 University of Southampton, Human Development and Health, Faculty of Medicine, Southampton, UK (GRID:grid.5491.9) (ISNI:0000 0004 1936 9297); University of Southampton, Clinical and Experimental Sciences, Faculty of Medicine, Southampton, UK (GRID:grid.5491.9) (ISNI:0000 0004 1936 9297)
88 University of Edinburgh, Centre for Genomics and Experimental Medicine, Institute for Genetics and Cancer, Edinburgh, Scotland (GRID:grid.4305.2) (ISNI:0000 0004 1936 7988)
89 Helmholtz Zentrum München - German Research Center for Environmental Health, Institute of Epidemiology, Neuherberg, Germany (GRID:grid.4567.0) (ISNI:0000 0004 0483 2525); German Center for Lung Research (DZL), Munich, Germany (GRID:grid.452624.3)