A

PRANA: A Deep Learning Method for Adapting Polygenic Risk Scores to Diverse Ethnic Groups

medRxiv

Abstract

Abstract Polygenic risk scores (PRSs), which quantify inherited susceptibility to complex traits and diseases, have emerged as valuable tools for risk stratification and precision medicine. Despite their promise, PRS developed on European cohorts often demonstrate substantially reduced predictive accuracy in non-European populations, due to differences in genetic architecture. The disproportionate representation of European ancestry cohorts in genome-wide association studies (GWAS) leads to inequitable deployment of PRS technologies across diverse populations. Here, we introduce PRANA (Polygenic Risk Adaptation via Neural-network Architecture), a deep learning framework that adapts an existing PRS developed on one population to other ancestries. Unlike methods that require large-scale GWAS in the target population, PRANA leverages pre-trained PRS models derived from European cohorts and adapts them using modestly sized cohorts from the target population. We evaluated PRANA on seven complex traits in South Asian, East Asian and Ashkenazi Jewish populations, as well as in selected smaller East Asian subpopulations where the scarcity of training data poses a particular challenge. PRANA mostly improved predictive performance of the baseline PRS models by 5%-20% in terms of effect size (β) and Nagelkerke’s R², and, in most cases, outperformed existing cross-ancestry multi-PRS approaches. These results highlight PRANA as a scalable and practical strategy to reduce disparities in genomic risk prediction and advance the equitable application of PRS in diverse populations.

Authors 100

  1. Tel Aviv University

    Affiliation as printed

    Blavatnik School of Computer Science and AI, Tel Aviv University, Tel Aviv, Israel

    Gray Faculty of Medical & Health Sciences, Tel Aviv University, Tel Aviv, Israel

  2. University of Cambridge

    Affiliation as printed

    Centre for Cancer Genetic Epidemiology, Department of Public Health and Primary Care, University of Cambridge, Cambridge, UK, CB1 8RN, 2 Worts’ Causeway

  3. University of Cambridge

    Affiliation as printed

    Centre for Cancer Genetic Epidemiology, Department of Public Health and Primary Care, University of Cambridge, Cambridge, UK, CB1 8RN, 2 Worts’ Causeway

  4. University of Cambridge

    Affiliation as printed

    Centre for Cancer Genetic Epidemiology, Department of Public Health and Primary Care, University of Cambridge, Cambridge, UK, CB1 8RN, 2 Worts’ Causeway

  5. Mount Sinai Hospital · University of Toronto · Lunenfeld-Tanenbaum Research Institute

    Affiliation as printed

    Department of Molecular Genetics, University of Toronto, Toronto, Ontario, Canada, M5S 1A8, 1 King’s College Circle

    Fred A. Litwin Center for Cancer Genetics, Lunenfeld-Tanenbaum Research Institute of Mount Sinai Hospital, Toronto, Ontario, Canada, M5G 1X5, 600 University Avenue

  6. N.N. Alexandrov National Cancer Centre

    Affiliation as printed

    N.N. Alexandrov Research Institute of Oncology and Medical Radiology, Minsk, Belarus, 223040, Settlement of Lesnoy-2

  7. Hong Kong Sanatorium and Hospital

    Affiliation as printed

    Molecular Pathology Division, Department of Pathology, Hong Kong Sanatorium and Hospital, Hong Kong, 2 Village Rd

  8. Lund University · Skåne University Hospital

    Affiliation as printed

    Oncology, Department of Clinical Sciences in Lund, Lund University, Lund, Sweden, 221 85, Barngatan 4, Skånes universitetssjukhus

  9. National Institutes of Health · National Cancer Institute · Division of Cancer Epidemiology and Genetics

    Affiliation as printed

    Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, MD, USA, 20850, 9609 Medical Center Dr

  10. German Cancer Research Center

    Affiliation as printed

    Division of Cancer Epidemiology, German Cancer Research Center (DKFZ), Heidelberg, Germany, 69120, Im Neuenheimer Feld 280

  11. Institute of Biochemistry and Genetics of Ufa Scientific Centre

    Affiliation as printed

    Institute of Biochemistry and Genetics—Subdivision of the Ufa Federal Research Centre of the Russian Academy of Sciences, Ufa, Russia, 450054, 71 prosp. Oktyabrya

  12. American Cancer Society

    Affiliation as printed

    Department of Population Science, American Cancer Society, Atlanta, GA, USA, 30303, 250 Williams Street NW

  13. Medizinische Hochschule Hannover · N.N. Alexandrov National Cancer Centre

    Affiliation as printed

    Department of Radiation Oncology, Hannover Medical School, Hannover, Germany, 30625, Carl-Neuberg-Straße 1

    Gynaecology Research Unit, Hannover Medical School, Hannover, Germany, 30625, Carl-Neuberg-Straße 1

    N.N. Alexandrov Research Institute of Oncology and Medical Radiology, Minsk, Belarus, 223040, Settlement of Lesnoy-2

  14. University of Copenhagen · Herlev Hospital · Copenhagen University Hospital · Gentofte Hospital

    Affiliation as printed

    Copenhagen General Population Study, Herlev and Gentofte Hospital, Copenhagen University Hospital, Herlev, Denmark, 2730, Herlev Ringvej 75

    Department of Clinical Biochemistry, Herlev and Gentofte Hospital, Copenhagen University Hospital, Herlev, Denmark, 2730, Herlev Ringvej 75

    Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark, 2200, Blegdamsvej 3B

  15. German Cancer Research Center

    Affiliation as printed

    Cancer Prevention Graduate School, German Cancer Research Center (DKFZ), Heidelberg, Germany, 69120, Im Neuenheimer Feld 280

  16. University of Sheffield

    Affiliation as printed

    Division of Clinical Medicine, School of Medicine and Population Health, University of Shefield, Shefield, UK, S10 2TN, Western Bank

  17. Institute for Prevention and Occupational Medicine

    Affiliation as printed

    Institute for Prevention and Occupational Medicine of the German Social Accident Insurance, Institute of the Ruhr University Bochum, Bochum, Germany, 44789, Bürkle-de-la-Camp-Platz 1

  18. Manchester Academic Health Science Centre · St Mary's Hospital · Manchester University NHS Foundation Trust

    Affiliation as printed

    North West Genomics Laboratory Hub, Manchester Centre for Genomic Medicine, St Mary’s Hospital, Manchester University NHS Foundation Trust, Manchester Academic Health Science Centre, Manchester, UK, M13 9WL, Oxford Road

  19. University of Utah · Huntsman Cancer Institute

    Affiliation as printed

    Department of Internal Medicine and Huntsman Cancer Institute, University of Utah, Salt Lake City, UT, USA, 84112, 2000 Circle of Hope

  20. Servicio Gallego de Salud

    Affiliation as printed

    Oncology and Genetics Unit, Instituto de Investigación Sanitaria Galicia Sur (IIS-GS), SERGAS, Vigo, Spain, 36312, Estrada Clara Campoamor n° 341

  21. Seoul National University

    Affiliation as printed

    Cancer Research Institute, Seoul National University, Seoul, Korea, 03080, 103 Daehak-ro, Jongno-gu Seoul

    Department of Biomedical Sciences, Seoul National University Graduate School, Seoul, Korea, 03080, 103 Daehak-ro, Jongno-gu Seoul

    Institute of Health Policy and Management, Seoul National University Medical Research Center, Seoul, Korea, 03080, 103 Daehak-ro, Jongno-gu Seoul

  22. Boston Children's Hospital · Harvard University

    Affiliation as printed

    Boston Children’s Hospital, Harvard Medical School, Boston, MA, USA

  23. University of Utah · Huntsman Cancer Institute

    Affiliation as printed

    Department of Internal Medicine and Huntsman Cancer Institute, University of Utah, Salt Lake City, UT, USA, 84112, 2000 Circle of Hope

  24. Mayo Clinic

    Affiliation as printed

    Department of Laboratory Medicine and Pathology, Mayo Clinic, Rochester, MN, USA, 55905, 200 First St. SW

  25. Karolinska Institutet

    Affiliation as printed

    Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden, 171 65, Karolinska Univ Hospital

  26. Fox Chase Cancer Center

    Affiliation as printed

    Department of Clinical Genetics, Fox Chase Cancer Center, Philadelphia, PA, USA, 19111, 333 Cottman Ave

  27. Leiden University Medical Center

    Affiliation as printed

    Department of Human Genetics, Leiden University Medical Center, Leiden, the Netherlands, 2333 ZA, Albinusdreef 2, P.O. Box 9600, 2300 RC

    Department of Pathology, Leiden University Medical Center, Leiden, the Netherlands, 2333 ZA, Albinusdreef 2

  28. Medizinische Hochschule Hannover

    Affiliation as printed

    Gynaecology Research Unit, Hannover Medical School, Hannover, Germany, 30625, Carl-Neuberg-Straße 1

  29. Brigham and Women's Hospital · Harvard University

    Affiliation as printed

    Channing Division of Network Medicine, Department of Medicine, Brigham and Women’s Hospital and Harvard Medical School, Boston, MA, USA, 02115, 181 Longwood Avenue, 3rd Floor

    Department of Epidemiology, Harvard T.H. Chan School of Public Health, Boston, MA, USA, 02115, 677 Huntington Avenue

    Department of Nutrition, Harvard T.H. Chan School of Public Health, Boston, MA, USA, 02115, 677 Huntington Ave

  30. Karolinska Institutet

    Affiliation as printed

    Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden, 171 65, Karolinska Univ Hospital

  31. Manchester Academic Health Science Centre · University of Manchester · St Mary's Hospital · Manchester University NHS Foundation Trust

    Affiliation as printed

    Division of Evolution and Genomic Sciences, School of Biological Sciences, Faculty of Biology, Medicine and Health, University of Manchester, Manchester Academic Health Science Centre, Manchester, UK, M13 9WL, Oxford Road

    North West Genomics Laboratory Hub, Manchester Centre for Genomic Medicine, St Mary’s Hospital, Manchester University NHS Foundation Trust, Manchester Academic Health Science Centre, Manchester, UK, M13 9WL, Oxford Road

  32. Friedrich-Alexander-Universität Erlangen-Nürnberg · Universitätsklinikum Erlangen · Comprehensive Cancer Center Erlangen

    Affiliation as printed

    Department of Gynecology and Obstetrics, Comprehensive Cancer Center Erlangen-EMN, Friedrich-Alexander University Erlangen-Nuremberg, University Hospital Erlangen, Erlangen, Germany, 91054, Maximiliansplatz 2

  33. American Cancer Society

    Affiliation as printed

    Department of Population Science, American Cancer Society, Atlanta, GA, USA, 30303, 250 Williams Street NW

  34. Complejo Hospitalario Universitario de Santiago · Instituto de Investigación Sanitaria de Santiago · Servicio Gallego de Salud

    Affiliation as printed

    Cancer Genetics and Epidemiology Group IDIS, Fundación Instituto de Investigación Sanitaria de Santiago de Compostela (FIDIS), Complejo Hospitalario Universitario de Santiago, SERGAS, Santiago de Compostela, Spain, 15706, Travesía da Choupana S/N

  35. Institute of Cancer Research

    Affiliation as printed

    Division of Genetics and Epidemiology, The Institute of Cancer Research, London, UK, SW7 3RP

  36. University of Southern California

    Affiliation as printed

    Department of Preventive Medicine, Keck School of Medicine, University of Southern California, Los Angeles, CA, USA, 90033, 1975 Zonal Ave

  37. German Cancer Research Center

    Affiliation as printed

    Molecular Genetics of Breast Cancer, German Cancer Research Center (DKFZ), Heidelberg, Germany, 69120, Im Neuenheimer Feld 580

  38. National University of Singapore · National University Hospital · National University Health System

    Affiliation as printed

    Department of Surgery, National University Hospital and National University Health System, Singapore City, Singapore, 119228, E Kent Ridge Rd

    Department of Surgery, Yong Loo Lin School of Medicine, National University of Singapore and National University Health System, Singapore City, Singapore, 119228, 21 Lower Kent Ridge Rd

    Saw Swee Hock School of Public Health, National University of Singapore and National University Health System, Singapore City, Singapore, 117549, 21 Lower Kent Ridge Rd

  39. Université de Montréal

    Affiliation as printed

    Département de médicine sociale et préventive, Université de Montréal, Montréal, Québec, Canada, H2X 0A9, Tour Saint-Antoine, 850 rue Saint-Denis, 2e étage, bureau S02.434

    Health Innovation and Evaluation Hub, Université de Montréal Hospital Research Centre (CRCHUM), Montréal, Québec, Canada

  40. 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 City, Singapore, 117549, 21 Lower Kent Ridge Rd

  41. Erasmus MC · Erasmus MC Cancer Institute · Erasmus University Rotterdam

    Affiliation as printed

    Department of Clinical Genetics, Erasmus University Medical Center, Rotterdam, the Netherlands, 3015 CN, Wytemaweg 80, P.O. Box 2040, 3000 CA

    Department of Medical Oncology, Erasmus MC Cancer Institute, Rotterdam, the Netherlands, 3015 GD, Dr. Molewaterplein 40, P.O. Box 2040, 3000 CA

  42. Dr. Margarete Fischer-Bosch-Institute of Clinical Pharmacology · University of Tübingen

    Affiliation as printed

    Dr. Margarete Fischer-Bosch-Institute of Clinical Pharmacology, Stuttgart, Germany, 70376, Auerbachstr. 112

    University of Tübingen, Tübingen, Germany, 72074, Geschwister-Scholl-Platz

  43. University of Manchester

    Affiliation as printed

    Division of Cancer Sciences, University of Manchester, Manchester, UK, M13 9PL

  44. Aichi Cancer Center · Nagoya University

    Affiliation as printed

    Division of Cancer Epidemiology, Nagoya University Graduate School of Medicine, Nagoya, Japan, 466-8550, 65 Tsurumai-cho, Showa-ku

    Division of Cancer Information and Control, Aichi Cancer Center Research Institute, Nagoya, Japan, 464-8681, 1-1 Kanokoden, Chikusa-ku

  45. National Cancer Centre Japan

    Affiliation as printed

    Division of Epidemiology, National Cancer Center Institute for Cancer Control, Tokyo, Japan, 104-0045, 5-1-1 Tsukiji, Chuo-ku

  46. Pomeranian Medical University

    Affiliation as printed

    Department of Genetics and Pathology, Pomeranian Medical University, Szczecin, Poland, 71-252, Unii Lubelskiej 1

    Independent Laboratory of Molecular Biology and Genetic Diagnostics, Pomeranian Medical University, Szczecin, Poland, 71-252

  47. Lund University · Skåne University Hospital

    Affiliation as printed

    Oncology, Department of Clinical Sciences in Lund, Lund University, Lund, Sweden, 221 85, Barngatan 4, Skånes universitetssjukhus

  48. Memorial Sloan Kettering Cancer Center

    Affiliation as printed

    Clinical Genetics Research Lab, Department of Cancer Biology and Genetics, Memorial Sloan Kettering Cancer Center, New York, NY, USA, 10065, 1275 York Avenue

    Clinical Genetics Service, Department of Medicine, Memorial Sloan Kettering Cancer Center, New York, NY, USA, 10065, 1275 York Avenue

  49. German Cancer Research Center

    Affiliation as printed

    Division of Cancer Epidemiology, German Cancer Research Center (DKFZ), Heidelberg, Germany, 69120, Im Neuenheimer Feld 280

  50. Seoul National University

    Affiliation as printed

    Cancer Research Institute, Seoul National University, Seoul, Korea, 03080, 103 Daehak-ro, Jongno-gu Seoul

    Department of Preventive Medicine, Seoul National University College of Medicine, Seoul, Korea, 03080, 103 Daehak-ro, Jongno-gu Seoul

  51. Institute of Biochemistry and Genetics of Ufa Scientific Centre · Ufa University of Science and Technology

    Affiliation as printed

    Department of Genetics and Fundamental Medicine, Ufa University of Science and Technology, Ufa, Russia, 450076, ul. Zaki Validi 32

    Institute of Biochemistry and Genetics—Subdivision of the Ufa Federal Research Centre of the Russian Academy of Sciences, Ufa, Russia, 450054, 71 prosp. Oktyabrya

  52. Macedonian Academy of Sciences and Arts

    Affiliation as printed

    Research Centre for Genetic Engineering and Biotechnology “Georgi D. Efremov”, Macedonian Academy of Sciences and Arts, Skopje, Republic of North Macedonia, 1000, Boulevard Krste Petkov Misirkov

  53. National Institutes of Health · National Cancer Institute · Division of Cancer Epidemiology and Genetics

    Affiliation as printed

    Division of Cancer Epidemiology and Genetics, National Cancer Institute, National Institutes of Health, Bethesda, MD, USA, 20850, 9609 Medical Center Dr

  54. Oslo University Hospital · University of Oslo

    Affiliation as printed

    Department of Medical Genetics, Oslo University Hospital and University of Oslo, Oslo, Norway, 0379, Ullernchausseen 70

  55. Stanford Medicine · Stanford Cancer Institute · Stanford University

    Affiliation as printed

    Department of Epidemiology and Population Health, Stanford University School of Medicine, Stanford, CA, USA, 94305, 259 Campus Drive

    Department of Medicine, Division of Oncology, Stanford Cancer Institute, Stanford University School of Medicine, Stanford, CA, USA, 94304, 780 Welch Road, Suite CJ250C

  56. Hong Kong Sanatorium and Hospital · University of Hong Kong

    Affiliation as printed

    Department of Surgery and Cancer Genetics Center, Hong Kong Sanatorium and Hospital, Hong Kong, 2 Village Rd

    Department of Surgery, The University of Hong Kong, Hong Kong, 102 Pokfulam Road

    Hong Kong Hereditary Breast Cancer Family Registry, Hong Kong, 18 A Kung Ngam Village Road

  57. VIB-KU Leuven Center for Cancer Biology · KU Leuven

    Affiliation as printed

    Laboratory for Translational Genetics, Department of Human Genetics, KU Leuven, Leuven, Belgium, 3000, Oude Markt 13

    VIB Center for Cancer Biology, VIB, Leuven, Belgium, 3001, Herestraat 46, Box 912

  58. Mayo Clinic

    Affiliation as printed

    Department of Quantitative Health Sciences, Division of Epidemiology, Mayo Clinic, Rochester, MN, USA, 55905, 200 First Street SW, Harwick 6

  59. University of Hawaii Cancer Center

    Affiliation as printed

    Epidemiology Program, University of Hawaii Cancer Center, Honolulu, HI, USA, 96813, 701 Ilalo St

  60. The University of Melbourne · University of Cambridge · Monash Health · Monash University

    Affiliation as printed

    Centre for Cancer Genetic Epidemiology, Department of Public Health and Primary Care, University of Cambridge, Cambridge, UK, CB1 8RN, 2 Worts’ Causeway

    Centre for Epidemiology and Biostatistics, Melbourne School of Population and Global Health, The University of Melbourne, Melbourne, Victoria, Australia, 3010, 207 Bouverie Street

    Precision Medicine, School of Clinical Sciences at Monash Health, Monash University, Clayton, Victoria, Australia, 3168, 246 Clayton Road

  61. Agency for Science, Technology and Research · Genome Institute of Singapore

    Affiliation as printed

    Human Genetics Division, Genome Institute of Singapore, Agency for Science, Technology and Research (A*STAR), Singapore City, Singapore, 138672, 60 Biopolis St

  62. University of Manchester

    Affiliation as printed

    Division of Population Health, Health Services Research and Primary Care, School of Health Sciences, Faculty of Biology, Medicine and Health, The University of Manchester, Manchester, UK, M13 9PL, Oxford Road

  63. University of Eastern Finland · Kuopio University Hospital

    Affiliation as printed

    Biobank of Eastern Finland, Kuopio University Hospital, Kuopio, Finland

    Institute of Clinical Medicine, Pathology and Forensic Medicine, University of Eastern Finland, Kuopio, Finland, 70210, Yliopistonranta 1

    Translational Cancer Research Area, University of Eastern Finland, Kuopio, Finland, 70210, Yliopistonranta 1

  64. Aichi Cancer Center · Nagoya University

    Affiliation as printed

    Division of Cancer Epidemiology and Prevention, Aichi Cancer Center Research Institute, Nagoya, Japan, 464-8681, 1-1 Kanokoden, Chikusa-ku

    Division of Cancer Epidemiology, Nagoya University Graduate School of Medicine, Nagoya, Japan, 466-8550, 65 Tsurumai-cho, Showa-ku

  65. Huntsman Cancer Institute

    Affiliation as printed

    Huntsman Cancer Institute, Salt Lake City, UT, USA, 84112, 2000 Circle of Hope

  66. University of Manchester

    Affiliation as printed

    Division of Population Health, Health Services Research and Primary Care, School of Health Sciences, Faculty of Biology, Medicine and Health, The University of Manchester, Manchester, UK, M13 9PL, Oxford Road

  67. Universitair Ziekenhuis Leuven

    Affiliation as printed

    Leuven Multidisciplinary Breast Center, Department of Oncology, Leuven Cancer Institute, University Hospitals Leuven, Leuven, Belgium, 3000, Oude Markt 13

  68. University Medical Center Hamburg-Eppendorf

    Affiliation as printed

    Institute for Medical Biometry and Epidemiology, University Medical Center Hamburg-Eppendorf, Hamburg, Germany, 20246, Martinistraße 52

    Institute for Occupational and Maritime Medicine, University Medical Center Hamburg-Eppendorf, Hamburg, Germany, 20246, Martinistraße 52

  69. National Institutes of Health · National Institute of Environmental Health Sciences

    Affiliation as printed

    Epidemiology Branch, National Institute of Environmental Health Sciences, NIH, Research Triangle Park, NC, USA, 27709, 111 T.W. Alexander Drive

  70. Memorial Sloan Kettering Cancer Center

    Affiliation as printed

    Clinical Genetics Research Lab, Department of Cancer Biology and Genetics, Memorial Sloan Kettering Cancer Center, New York, NY, USA, 10065, 1275 York Avenue

    Clinical Genetics Service, Department of Medicine, Memorial Sloan Kettering Cancer Center, New York, NY, USA, 10065, 1275 York Avenue

  71. University of Sheffield

    Affiliation as printed

    Division of Clinical Medicine, School of Medicine and Population Health, University of Shefield, Shefield, UK, S10 2TN, Western Bank

  72. American Cancer Society

    Affiliation as printed

    Department of Population Science, American Cancer Society, Atlanta, GA, USA, 30303, 250 Williams Street NW

  73. European Institute of Oncology

    Affiliation as printed

    Laboratory of Medical Genetics, Cytogenetics and Molecular Genetics, IEO, European Institute of Oncology IRCCS, Milan, Italy

  74. Macedonian Academy of Sciences and Arts

    Affiliation as printed

    Research Centre for Genetic Engineering and Biotechnology “Georgi D. Efremov”, Macedonian Academy of Sciences and Arts, Skopje, Republic of North Macedonia, 1000, Boulevard Krste Petkov Misirkov

  75. Pomeranian Medical University

    Affiliation as printed

    Department of Genetics and Pathology, Pomeranian Medical University, Szczecin, Poland, 71-252, Unii Lubelskiej 1

  76. Fondazione IRCCS Istituto Nazionale dei Tumori

    Affiliation as printed

    Unit of “Predictive Medicine: Molecular Bases of Genetic Risk”, Department of Experimental Research, Fondazione IRCCS Istituto Nazionale dei Tumori (INT), Milan, Italy, 20133, Via Giacomo Venezian 1

  77. Medizinische Hochschule Hannover

    Affiliation as printed

    Gynaecology Research Unit, Hannover Medical School, Hannover, Germany, 30625, Carl-Neuberg-Straße 1

  78. German Cancer Research Center · Shaukat Khanum Memorial Cancer Hospital and Research Center

    Affiliation as printed

    Department of Basic Sciences, Shaukat Khanum Memorial Cancer Hospital and Research Centre (SKMCH & RC), Lahore, Pakistan, 54000, Shaukat Khanum Hospital Road

    Molecular Genetics of Breast Cancer, German Cancer Research Center (DKFZ), Heidelberg, Germany, 69120, Im Neuenheimer Feld 580

  79. Hospital Universitario Puerta de Hierro Majadahonda

    Affiliation as printed

    Medical Oncology Department, Hospital Universitario Puerta de Hierro, Madrid, Spain, 28222, Calle Manuel de Falla, 1

  80. University of Thessaly

    Affiliation as printed

    Division of Oncology, Faculty of Medicine, School of Health Sciences, University of Thessaly, Larissa, Greece, 411 10, Mezourlo

  81. National Institutes of Health · National Institute of Environmental Health Sciences

    Affiliation as printed

    Epidemiology Branch, National Institute of Environmental Health Sciences, NIH, Research Triangle Park, NC, USA, 27709, 111 T.W. Alexander Drive

  82. German Cancer Research Center

    Affiliation as printed

    Division of Clinical Epidemiology of Early Cancer Detection, German Cancer Research Center (DKFZ), Heidelberg, Germany, 69120, Im Neuenheimer Feld 280

  83. Centre hospitalier universitaire de Québec · Centre hospitalier de l'Université Laval · Université Laval

    Affiliation as printed

    Genomics Center, Centre Hospitalier Universitaire de Québec – Université Laval Research Center, Québec City, Québec, Canada, G1V 4G2, 2705 Laurier Boulevard

  84. The University of Melbourne · Cancer Council Victoria · Monash Health · Monash University

    Affiliation as printed

    Cancer Epidemiology Division, Cancer Council Victoria, Melbourne, Victoria, Australia, 3004, 615 St Kilda Road

    Department of Clinical Pathology, The University of Melbourne, Melbourne, Victoria, Australia, 3010, Cnr Grattan Street and Royal Parade

    Precision Medicine, School of Clinical Sciences at Monash Health, Monash University, Clayton, Victoria, Australia, 3168, 246 Clayton Road

  85. The University of Melbourne · The University of Western Australia

    Affiliation as printed

    Centre for Epidemiology and Biostatistics, Melbourne School of Population and Global Health, The University of Melbourne, Melbourne, Victoria, Australia, 3010, 207 Bouverie Street

    Genetic Epidemiology Group, School of Population and Global Health, University of Western Australia, Perth, Western Australia, Australia, 6000, 35 Stirling Hwy

  86. Centre National de la Recherche Scientifique · Inserm · Université de Versailles Saint-Quentin-en-Yvelines · Université Paris Cité · Université Paris-Saclay · Institut Gustave Roussy · Centre de recherche en Epidémiologie et Santé des Populations · Mathématiques Appliquées à Paris 5

    Affiliation as printed

    Team “Exposome and Heredity”, CESP, Gustave Roussy, INSERM, University Paris-Saclay, UVSQ, Villejuif, France, 94805, 39 rue Camille Desmoulins

    Université Paris Cité, CNRS, MAP5, Paris, France, 75006

  87. National Institutes of Health · National Institute of Environmental Health Sciences

    Affiliation as printed

    Epidemiology Branch, National Institute of Environmental Health Sciences, NIH, Research Triangle Park, NC, USA, 27709, 111 T.W. Alexander Drive

    Epigenetic and Stem Cell Biology Laboratory, National Institute of Environmental Health Sciences, NIH, Research Triangle Park, NC, USA, 27709, 111 T.W. Alexander Drive

  88. American Cancer Society

    Affiliation as printed

    Department of Population Science, American Cancer Society, Atlanta, GA, USA, 30303, 250 Williams Street NW

  89. Inserm · Université de Versailles Saint-Quentin-en-Yvelines · Université Paris-Saclay · Institut Gustave Roussy · Centre de recherche en Epidémiologie et Santé des Populations

    Affiliation as printed

    Team “Exposome and Heredity”, CESP, Gustave Roussy, INSERM, University Paris-Saclay, UVSQ, Villejuif, France, 94805, 39 rue Camille Desmoulins

  90. National Institutes of Health · National Institute of Environmental Health Sciences

    Affiliation as printed

    Biostatistics and Computational Biology Branch, National Institute of Environmental Health Sciences, NIH, Research Triangle Park, NC, USA, 27709, 111 T.W. Alexander Drive

  91. Mayo Clinic

    Affiliation as printed

    Department of Oncology, Mayo Clinic, Rochester, MN, USA, 55905, 200 First Street SW

  92. National Cancer Centre Japan

    Affiliation as printed

    Division of Epidemiology, National Cancer Center Institute for Cancer Control, Tokyo, Japan, 104-0045, 5-1-1 Tsukiji, Chuo-ku

  93. Vanderbilt University · Vanderbilt-Ingram Cancer Center

    Affiliation as printed

    Division of Epidemiology, Department of Medicine, Vanderbilt Epidemiology Center, Vanderbilt-Ingram Cancer Center, Vanderbilt University School of Medicine, Nashville, TN, USA, 37232, 1161 21st Ave S # D3300

  94. University of Cambridge

    Affiliation as printed

    Centre for Cancer Genetic Epidemiology, Department of Oncology, University of Cambridge, Cambridge, UK, CB1 8RN, 2 Worts’ Causeway

  95. University of Cambridge

    Affiliation as printed

    Centre for Cancer Genetic Epidemiology, Department of Public Health and Primary Care, University of Cambridge, Cambridge, UK, CB1 8RN, 2 Worts’ Causeway

  96. University of Cambridge

    Affiliation as printed

    Centre for Cancer Genetic Epidemiology, Department of Oncology, University of Cambridge, Cambridge, UK, CB1 8RN, 2 Worts’ Causeway

    Centre for Cancer Genetic Epidemiology, Department of Public Health and Primary Care, University of Cambridge, Cambridge, UK, CB1 8RN, 2 Worts’ Causeway

  97. University of Cambridge · Cyprus Institute of Neurology and Genetics

    Affiliation as printed

    Centre for Cancer Genetic Epidemiology, Department of Public Health and Primary Care, University of Cambridge, Cambridge, UK, CB1 8RN, 2 Worts’ Causeway

    Department of Biostatistics, The Cyprus Institute of Neurology and Genetics, Nicosia, Cyprus, 2371, 6 Iroon Avenue,2371 Ayios Dometios, Nicosia, P.O.Box 23462, 1683, Nicosia, Cyprus

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