Letter

Association analysis identifies 65 new breast cancer risk loci

Received:
Accepted:
Published online:

Abstract

Breast cancer risk is influenced by rare coding variants in susceptibility genes, such as BRCA1, and many common, mostly non-coding variants. However, much of the genetic contribution to breast cancer risk remains unknown. Here we report the results of a genome-wide association study of breast cancer in 122,977 cases and 105,974 controls of European ancestry and 14,068 cases and 13,104 controls of East Asian ancestry1. We identified 65 new loci that are associated with overall breast cancer risk at P < 5 × 10−8. The majority of credible risk single-nucleotide polymorphisms in these loci fall in distal regulatory elements, and by integrating in silico data to predict target genes in breast cells at each locus, we demonstrate a strong overlap between candidate target genes and somatic driver genes in breast tumours. We also find that heritability of breast cancer due to all single-nucleotide polymorphisms in regulatory features was 2–5-fold enriched relative to the genome-wide average, with strong enrichment for particular transcription factor binding sites. These results provide further insight into genetic susceptibility to breast cancer and will improve the use of genetic risk scores for individualized screening and prevention.

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References

  1. 1.

    et al. The OncoArray Consortium: a network for understanding the genetic architecture of common cancers. Cancer Epidemiol. Biomarkers Prev. 26, 126–135 (2017)

  2. 2.

    et al. Large-scale genotyping identifies 41 new loci associated with breast cancer risk. Nat. Genet. 45, 353–361 (2013)

  3. 3.

    et al. Genome-wide association study in East Asians identifies novel susceptibility loci for breast cancer. PLoS Genet. 8, e1002532 (2012)

  4. 4.

    et al. Genome-wide association analysis in East Asians identifies breast cancer susceptibility loci at 1q32.1, 5q14.3 and 15q26.1. Nat. Genet. 46, 886–890 (2014)

  5. 5.

    et al. A common deletion in the APOBEC3 genes and breast cancer risk. J. Natl Cancer Inst. 105, 573–579 (2013)

  6. 6.

    et al. Genome-wide association studies identify loci associated with age at menarche and age at natural menopause. Nat. Genet. 41, 724–728 (2009)

  7. 7.

    et al. PH domain-only protein PHLDA3 is a p53-regulated repressor of Akt. Cell 136, 535–550 (2009)

  8. 8.

    et al. Combinatorial effects of multiple enhancer variants in linkage disequilibrium dictate levels of gene expression to confer susceptibility to common traits. Genome Res. 24, 1–13 (2014)

  9. 9.

    , , & Global view of enhancer–promoter interactome in human cells. Proc. Natl Acad. Sci. USA 111, E2191–E2199 (2014)

  10. 10.

    et al. An atlas of active enhancers across human cell types and tissues. Nature 507, 455–461 (2014)

  11. 11.

    et al. Landscape of somatic mutations in 560 breast cancer whole-genome sequences. Nature 534, 47–54 (2016)

  12. 12.

    Cancer Genome Atlas Network. Comprehensive molecular portraits of human breast tumours. Nature 490, 61–70 (2012)

  13. 13.

    et al. Comprehensive molecular portraits of invasive lobular breast cancer. Cell 163, 506–519 (2015)

  14. 14.

    et al. The somatic mutation profiles of 2,433 breast cancers refine their genomic and transcriptomic landscapes. Nat. Commun. 7, 11479 (2016)

  15. 15.

    et al. Biological interpretation of genome-wide association studies using predicted gene functions. Nat. Commun. 6, 5890 (2015)

  16. 16.

    , , , & Enrichment map: a network-based method for gene-set enrichment visualization and interpretation. PLoS ONE 5, e13984 (2010)

  17. 17.

    & Fibroblast growth factor signalling: from development to cancer. Nat. Rev. Cancer 10, 116–129 (2010)

  18. 18.

    Targeting the PDGF signaling pathway in tumor treatment. Cell Commun. Signal. 11, 97 (2013)

  19. 19.

    & Wnt signaling and breast cancer. Cancer Biol. Ther. 3, 36–41 (2004)

  20. 20.

    et al. Partitioning heritability by functional annotation using genome-wide association summary statistics. Nat. Genet. 47, 1228–1235 (2015)

  21. 21.

    ., & Pidd, a new death-domain-containing protein, is induced by p53 and promotes apoptosis. Nat. Genet. 26, 122–127 (2000)

  22. 22.

    et al. CITED4 inhibits hypoxia-activated transcription in cancer cells, and its cytoplasmic location in breast cancer is associated with elevated expression of tumor cell hypoxia-inducible factor 1α. Cancer Res. 64, 6075–6081 (2004)

  23. 23.

    et al. Genome-wide association analysis of more than 120,000 individuals identifies 15 new susceptibility loci for breast cancer. Nat. Genet. 47, 373–380 (2015)

  24. 24.

    et al. A general approach for haplotype phasing across the full spectrum of relatedness. PLoS Genet. 10, e1004234 (2014)

  25. 25.

    , & A flexible and accurate genotype imputation method for the next generation of genome-wide association studies. PLoS Genet. 5, e1000529 (2009)

  26. 26.

    , , , & Fast and accurate genotype imputation in genome-wide association studies through pre-phasing. Nat. Genet. 44, 955–959 (2012)

  27. 27.

    ., ., ., . & MaCH: using sequence and genotype data to estimate haplotypes and unobserved genotypes. Genet. Epidemiol. 34, 816–834 (2010)

  28. 28.

    , & ProbABEL package for genome-wide association analysis of imputed data. BMC Bioinformatics 11, 134 (2010)

  29. 29.

    , & METAL: fast and efficient meta-analysis of genomewide association scans. Bioinformatics 26, 2190–2191 (2010)

  30. 30.

    , , & Joint analysis is more efficient than replication-based analysis for two-stage genome-wide association studies. Nat. Genet. 38, 209–213 (2006)

  31. 31.

    R Core Team. R: A Language and Environment for Statistical Computing (2016)

  32. 32.

    The ENCODE Project Consortium. A user’s guide to the encyclopedia of DNA elements (ENCODE). PLoS Biol. 9, e1001046 (2011)

  33. 33.

    et al. Integrative analysis of 111 reference human epigenomes. Nature 518, 317–330 (2015)

  34. 34.

    , & Evaluating the power to discriminate between highly correlated SNPs in genetic association studies. Genet. Epidemiol. 34, 463–468 (2010)

  35. 35.

    et al. Bayesian refinement of association signals for 14 loci in 3 common diseases. Nat. Genet. 44, 1294–1301 (2012)

  36. 36.

    et al. The genomic and transcriptomic architecture of 2,000 breast tumours reveals novel subgroups. Nature 486, 346–352 (2012)

  37. 37.

    et al. Fast and accurate inference of local ancestry in Latino populations. Bioinformatics 28, 1359–1367 (2012)

  38. 38.

    The 1000 Genomes Project Consortium. An integrated map of genetic variation from 1,092 human genomes. Nature 491, 56–65 (2012)

  39. 39.

    & RSEM: accurate transcript quantification from RNA-seq data with or without a reference genome. BMC Bioinformatics 12, 323 (2011)

  40. 40.

    et al. GISTIC2.0 facilitates sensitive and confident localization of the targets of focal somatic copy-number alteration in human cancers. Genome Biol. 12, R41 (2011)

  41. 41.

    et al. Integrative eQTL-based analyses reveal the biology of breast cancer risk loci. Cell 152, 633–641 (2013)

  42. 42.

    Matrix eQTL: ultra fast eQTL analysis via large matrix operations. Bioinformatics 28, 1353–1358 (2012)

  43. 43.

    et al. The UCSC Table Browser data retrieval tool. Nucleic Acids Res. 32, D493–D496 (2004)

  44. 44.

    et al. A 3D map of the human genome at kilobase resolution reveals principles of chromatin looping. Cell 159, 1665–1680 (2014)

  45. 45.

    & BEDTools: a flexible suite of utilities for comparing genomic features. Bioinformatics 26, 841–842 (2010)

  46. 46.

    et al. GENCODE: the reference human genome annotation for The ENCODE Project. Genome Res. 22, 1760–1774 (2012)

  47. 47.

    et al. Super-enhancers in the control of cell identity and disease. Cell 155, 934–947 (2013)

  48. 48.

    et al. metagene profiles analyses reveal regulatory element’s factor-specific recruitment patterns. PLOS Comput. Biol. 12, e1004751 (2016)

  49. 49.

    et al. A general framework for estimating the relative pathogenicity of human genetic variants. Nat. Genet. 46, 310–315 (2014)

  50. 50.

    et al. Predicting the functional, molecular, and phenotypic consequences of amino acid substitutions using hidden Markov models. Hum. Mutat. 34, 57–65 (2013)

  51. 51.

    & Identification of deleterious mutations within three human genomes. Genome Res. 19, 1553–1561 (2009)

  52. 52.

    , & Predicting the functional impact of protein mutations: application to cancer genomics. Nucleic Acids Res. 39, e118 (2011)

  53. 53.

    , , & MutationTaster2: mutation prediction for the deep-sequencing age. Nat. Methods 11, 361–362 (2014)

  54. 54.

    et al. A method and server for predicting damaging missense mutations. Nat. Methods 7, 248–249 (2010)

  55. 55.

    , , , & Predicting the functional effect of amino acid substitutions and indels. PLoS ONE 7, e46688 (2012)

  56. 56.

    , & Predicting the effects of coding non-synonymous variants on protein function using the SIFT algorithm. Nat. Protoc. 4, 1073–1081 (2009)

  57. 57.

    et al. Human Splicing Finder: an online bioinformatics tool to predict splicing signals. Nucleic Acids Res. 37, e67 (2009)

  58. 58.

    & Maximum entropy modeling of short sequence motifs with applications to RNA splicing signals. J. Comput. Biol. 11, 377–394 (2004)

  59. 59.

    et al. The NHGRI GWAS Catalog, a curated resource of SNP–trait associations. Nucleic Acids Res. 42, D1001–D1006 (2014)

  60. 60.

    et al. Evidence that breast cancer risk at the 2q35 locus is mediated through IGFBP5 regulation. Nat. Commun. 4, 4999 (2014)

  61. 61.

    et al. Reactome: a knowledgebase of biological pathways. Nucleic Acids Res. 33, D428–D432 (2005)

  62. 62.

    et al. PID: the pathway interaction database. Nucleic Acids Res. 37, D674–D679 (2009)

  63. 63.

    et al. Gene ontology: tool for the unification of biology. Nat. Genet. 25, 25–29 (2000)

  64. 64.

    et al. Computational prediction of human metabolic pathways from the complete human genome. Genome Biol. 6, R2 (2004)

  65. 65.

    et al. Gene set enrichment analysis: a knowledge-based approach for interpreting genome-wide expression profiles. Proc. Natl Acad. Sci. USA 102, 15545–15550 (2005)

  66. 66.

    et al. NetPath: a public resource of curated signal transduction pathways. Genome Biol. 11, R3 (2010)

  67. 67.

    et al. PANTHER: a library of protein families and subfamilies indexed by function. Genome Res. 13, 2129–2141 (2003)

  68. 68.

    , , , & Gene set analysis of genome-wide association studies: methodological issues and perspectives. Genomics 98, 1–8 (2011)

  69. 69.

    , & Analysing biological pathways in genome-wide association studies. Nat. Rev. Genet. 11, 843–854 (2010)

  70. 70.

    , & Pathway-based approaches for analysis of genomewide association studies. Am. J. Hum. Genet. 81, 1278–1283 (2007)

  71. 71.

    & PathAct: a novel method for pathway analysis using gene expression profiles. Bioinformation 9, 394–400 (2013)

  72. 72.

    et al. Gene set-based analysis of polymorphisms: finding pathways or biological processes associated to traits in genome-wide association studies. Nucleic Acids Res. 37, W340–W344 (2009)

  73. 73.

    , & Genome-wide pathway analysis of breast cancer. Tumour Biol. 35, 7699–7705 (2014)

  74. 74.

    , , , & dmGWAS: dense module searching for genome-wide association studies in protein–protein interaction networks. Bioinformatics 27, 95–102 (2011)

  75. 75.

    & Pathways of distinction analysis: a new technique for multi-SNP analysis of GWAS data. PLoS Genet. 7, e1002101 (2011)

  76. 76.

    et al. Cytoscape: a software environment for integrated models of biomolecular interaction networks. Genome Res. 13, 2498–2504 (2003)

Download references

Acknowledgements

We thank all the individuals who took part in these studies and all the researchers, clinicians, technicians and administrative staff who have enabled this work to be carried out. Genotyping of the OncoArray was principally funded from three sources: the PERSPECTIVE project, funded by the Government of Canada through Genome Canada and the Canadian Institutes of Health Research, the ‘Ministère de l’Économie, de la Science et de l’Innovation du Québec’ through Genome Québec, and the Quebec Breast Cancer Foundation; the NCI Genetic Associations and Mechanisms in Oncology (GAME-ON) initiative and Discovery, Biology and Risk of Inherited Variants in Breast Cancer (DRIVE) project (NIH Grants U19 CA148065 and X01HG007492); and Cancer Research UK (C1287/A10118 and C1287/A16563). BCAC is funded by Cancer Research UK (C1287/A16563), by the European Community’s Seventh Framework Programme under grant agreement 223175 (HEALTH-F2-2009-223175) (COGS) and by the European Union’s Horizon 2020 Research and Innovation Programme under grant agreements 633784 (B-CAST) and 634935 (BRIDGES). Genotyping of the iCOGS array was funded by the European Union (HEALTH-F2-2009-223175), Cancer Research UK (C1287/A10710), the Canadian Institutes of Health Research for the ‘CIHR Team in Familial Risks of Breast Cancer’ program, and the Ministry of Economic Development, Innovation and Export Trade of Quebec, grant PSR-SIIRI-701. Combining of the GWAS data was supported in part by The National Institute of Health (NIH) Cancer Post-Cancer GWAS initiative grant U19 CA 148065 (DRIVE, part of the GAME-ON initiative). For a full description of funding and acknowledgments, see Supplementary Note.

Author information

Author notes

    • Kyriaki Michailidou
    • , Sara Lindström
    • , Joe Dennis
    • , Jonathan Beesley
    • , Shirley Hui
    •  & Siddhartha Kar

    These authors contributed equally to this work.

    • Jacques Simard
    • , Peter Kraft
    •  & Douglas F. Easton

    These authors jointly supervised this work.

Affiliations

  1. Centre for Cancer Genetic Epidemiology, Department of Public Health and Primary Care, University of Cambridge, Cambridge, UK

    • Kyriaki Michailidou
    • , Joe Dennis
    • , Manjeet K. Bolla
    • , Qin Wang
    • , Andrew Lee
    • , Jamie Allen
    • , Antonis C. Antoniou
    • , Paul D. P. Pharoah
    •  & Douglas F. Easton
  2. Department of Electron Microscopy/Molecular Pathology, The Cyprus Institute of Neurology and Genetics, Nicosia, Cyprus

    • Kyriaki Michailidou
  3. Department of Epidemiology, University of Washington School of Public Health, Seattle, Washington, USA

    • Sara Lindström
  4. Program in Genetic Epidemiology and Statistical Genetics, Harvard T. H. Chan School of Public Health, Boston, Massachusetts, USA

    • Sara Lindström
    • , Xia Jiang
    • , Hilary Finucane
    • , Rulla Tamimi
    • , David J. Hunter
    •  & Peter Kraft
  5. Department of Genetics and Computational Biology, QIMR Berghofer Medical Research Institute, Brisbane, Australia

    • Jonathan Beesley
    • , Dylan Glubb
    • , Juliet D. French
    • , Xiao Qing Chen
    • , Karen McCue
    • , Stacey L. Edwards
    •  & Georgia Chenevix-Trench
  6. The Donnelly Centre, University of Toronto, Toronto, Ontario, Canada

    • Shirley Hui
    • , Asha Rostamianfar
    •  & Gary D. Bader
  7. Centre for Cancer Genetic Epidemiology, Department of Oncology, University of Cambridge, Cambridge, UK

    • Siddhartha Kar
    • , Jonathan Tyrer
    • , Ed Dicks
    • , Laura Fachal
    • , Maya Ghoussaini
    • , Caroline Baynes
    • , Don M. Conroy
    • , Patricia Harrington
    • , Craig Luccarini
    • , Valerie Rhenius
    • , Mitul Shah
    • , Paul D. P. Pharoah
    • , Alison M. Dunning
    •  & Douglas F. Easton
  8. Genomics Center, Centre Hospitalier Universitaire de Québec Research Center, Laval University, Québec City, Quebec, Canada

    • Audrey Lemaçon
    • , Penny Soucy
    • , Martine Dumont
    • , Arnaud Droit
    •  & Jacques Simard
  9. Department of Computational Biology, St Jude Children’s Research Hospital, Memphis, Tennessee, USA

    • Zhaoming Wang
  10. Cancer Genomics Research Laboratory (CGR), Division of Cancer Epidemiology and Genetics, National Cancer Institute, Rockville, Maryland, USA

    • Zhaoming Wang
    • , Belynda Hicks
    • , Kristine Jones
    •  & Bin Zhu
  11. Division of Molecular Pathology, The Netherlands Cancer Institute, Antoni van Leeuwenhoek Hospital, Amsterdam, The Netherlands

    • Renske Keeman
    • , Annegien Broeks
    • , Sten Cornelissen
    •  & Marjanka K. Schmidt
  12. Division of Cancer Epidemiology, German Cancer Research Center (DKFZ), Heidelberg, Germany

    • Ursula Eilber
    • , Myrto Barrdahl
    • , Sabine Behrens
    • , Rudolf Kaaks
    • , Anja Rudolph
    •  & Jenny Chang-Claude
  13. UQ Centre for Clinical Research, The University of Queensland, Brisbane, Australia

    • Amy E. McCart Reed
    •  & Sunil R. Lakhani
  14. Cancer Research UK Cambridge Research Institute, University of Cambridge, Li Ka Shing Centre, Cambridge, UK

    • Jason S. Carroll
  15. Department of Mathematics, Massachusetts Institute of Technology, Cambridge, Massachusetts, USA

    • Hilary Finucane
  16. Center for Inherited Disease Research (CIDR), Institute of Genetic Medicine, Johns Hopkins University School of Medicine, Baltimore, Maryland, USA

    • Marcia Adams
    • , Kimberly F. Doheny
    • , Elizabeth Pugh
    •  & Jane Romm
  17. Department of Clinical Genetics, VU University Medical Center, Amsterdam, The Netherlands

    • Muriel A. Adank
    • , Hanne Meijers-Heijboer
    •  & Quinten Waisfisz
  18. Center for Cancer Epidemiology and Prevention, The University of Chicago, Chicago, Illinois, USA

    • Habibul Ahsan
    •  & Ling Tong
  19. Department of Clinical Genetics, Helsinki University Hospital, University of Helsinki, Helsinki, Finland

    • Kristiina Aittomäki
  20. Department of Epidemiology, University of California Irvine, Irvine, California, USA

    • Hoda Anton-Culver
    •  & Argyrios Ziogas
  21. N. N. Alexandrov Research Institute of Oncology and Medical Radiology, Minsk, Belarus

    • Natalia N. Antonenkova
    •  & Natalia V. Bogdanova
  22. Division of Clinical Epidemiology and Aging Research, German Cancer Research Center (DKFZ), Heidelberg, Germany

    • Volker Arndt
    • , Hermann Brenner
    •  & Katja Butterbach
  23. Department of Public Health Sciences, and Cancer Research Institute, Queen’s University, Kingston, Ontario, Canada

    • Kristan J. Aronson
  24. Department of Breast Medical Oncology, University of Texas MD Anderson Cancer Center, Houston, Texas, USA

    • Banu Arun
  25. Cancer Prevention Program, Fred Hutchinson Cancer Research Center, Seattle, Washington, USA

    • Paul L. Auer
    •  & Ross Prentice
  26. Zilber School of Public Health, University of Wisconsin-Milwaukee, Milwaukee, Wisconsin, USA

    • Paul L. Auer
  27. McGill University and Génome Québec Innovation Centre, Montréal, Quebec, Canada

    • François Bacot
    • , Nathalie Hamel
    • , Daniel C. Tessier
    •  & Daniel Vincent
  28. Department of Gynaecology and Obstetrics, University Hospital Erlangen, Friedrich-Alexander University Erlangen-Nuremberg, Comprehensive Cancer Center Erlangen-EMN, Erlangen, Germany

    • Matthias W. Beckmann
    • , Peter A. Fasching
    • , Lothar Haeberle
    • , Alexander Hein
    • , Michael P. Lux
    •  & Matthias Ruebner
  29. Human Cancer Genetics Program, Spanish National Cancer Research Centre, Madrid, Spain

    • Javier Benitez
    •  & Anna González-Neira
  30. Centro de Investigación en Red de Enfermedades Raras (CIBERER), Valencia, Spain

    • Javier Benitez
  31. Institute of Biochemistry and Genetics, Ufa Scientific Center of Russian Academy of Sciences, Ufa, Russia

    • Marina Bermisheva
    •  & Elza Khusnutdinova
  32. Department of Population Sciences, Beckman Research Institute of City of Hope, Duarte, California, USA

    • Leslie Bernstein
    •  & Susan L. Neuhausen
  33. Department of Oncology, Helsinki University Hospital, University of Helsinki, Helsinki, Finland

    • Carl Blomqvist
  34. Department of Radiation Oncology, Hannover Medical School, Hannover, Germany

    • Natalia V. Bogdanova
    •  & Hans Christiansen
  35. Gynaecology Research Unit, Hannover Medical School, Hannover, Germany

    • Natalia V. Bogdanova
    • , Thilo Dörk
    • , Peter Hillemanns
    • , Tjoung-Won Park-Simon
    •  & Peter Schürmann
  36. Copenhagen General Population Study, Herlev and Gentofte Hospital, Copenhagen University Hospital, Herlev, Denmark

    • Stig E. Bojesen
    • , Sune F. Nielsen
    •  & Børge G. Nordestgaard
  37. Department of Clinical Biochemistry, Herlev and Gentofte Hospital, Copenhagen University Hospital, Herlev, Denmark

    • Stig E. Bojesen
    • , Sune F. Nielsen
    •  & Børge G. Nordestgaard
  38. Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark

    • Stig E. Bojesen
    •  & Børge G. Nordestgaard
  39. Division of Cancer Prevention and Genetics, Istituto Europeo di Oncologia, Milan, Italy

    • Bernardo Bonanni
  40. Department of Cancer Genetics, Institute for Cancer Research, Oslo University Hospital Radiumhospitalet, Oslo, Norway

    • Anne-Lise Børresen-Dale
    • , Grethe I. Grenaker Alnæs
    •  & Vessela N. Kristensen
  41. Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden

    • Judith S. Brand
    • , Kamila Czene
    • , Mikael Eriksson
    • , Marike Gabrielson
    • , Keith Humphreys
    • , Jingmei Li
    •  & Per Hall
  42. Dr Margarete Fischer-Bosch-Institute of Clinical Pharmacology, Stuttgart, Germany

    • Hiltrud Brauch
    •  & Wing-Yee Lo
  43. University of Tübingen, Tübingen, Germany

    • Hiltrud Brauch
    •  & Wing-Yee Lo
  44. German Cancer Consortium (DKTK), German Cancer Research Center (DKFZ), Heidelberg, Germany

    • Hiltrud Brauch
    •  & Hermann Brenner
  45. International Agency for Research on Cancer, Lyon, France

    • Paul Brennan
    • , Valerie Gaborieau
    •  & James McKay
  46. Division of Preventive Oncology, German Cancer Research Center (DKFZ) and National Center for Tumor Diseases (NCT), Heidelberg, Germany

    • Hermann Brenner
  47. Division of Cancer Epidemiology and Genetics, National Cancer Institute, Rockville, Maryland, USA

    • Louise Brinton
    • , Jonine Figueroa
    • , Robert N. Hoover
    • , Xiaohong R. Yang
    • , Montserrat García-Closas
    •  & Stephen J. Chanock
  48. Department of Cancer Epidemiology, Clinical Sciences, Lund University, Lund, Sweden

    • Per Broberg
    • , Carolina Ellberg
    • , Ute Krüger
    •  & Håkan Olsson
  49. Sheffield Institute for Nucleic Acids (SInFoNiA), Department of Oncology and Metabolism, University of Sheffield, Sheffield, UK

    • Ian W. Brock
    •  & Angela Cox
  50. Genome Sciences Centre, BC Cancer Agency, Vancouver, British Columbia, Canada

    • Angela Brooks-Wilson
  51. Department of Biomedical Physiology and Kinesiology, Simon Fraser University, Burnaby, British Columbia, Canada

    • Angela Brooks-Wilson
  52. Department of Gynecology and Obstetrics, University of Tübingen, Tübingen, Germany

    • Sara Y. Brucker
  53. Institute for Prevention and Occupational Medicine of the German Social Accident Insurance, Institute of the Ruhr University Bochum, Bochum, Germany

    • Thomas Brüning
  54. Department of Obstetrics and Gynecology, University of Heidelberg, Heidelberg, Germany

    • Barbara Burwinkel
    • , Andreas Schneeweiss
    •  & Harald Surowy
  55. Molecular Epidemiology Group, C080, German Cancer Research Center (DKFZ), Heidelberg, Germany

    • Barbara Burwinkel
    •  & Harald Surowy
  56. Division of Epidemiology, Department of Medicine, Vanderbilt Epidemiology Center, Vanderbilt-Ingram Cancer Center, Vanderbilt University School of Medicine, Nashville, Tennessee, USA

    • Qiuyin Cai
    • , Hui Cai
    • , Jirong Long
    • , Martha J. Shrubsole
    • , Xiao-Ou Shu
    •  & Wei Zheng
  57. Medical Oncology Department, Hospital Clínico San Carlos, IdISSC (Centro Investigacion Biomedica en Red), CIBERONC (Instituto de Investigación Sanitaria San Carlos), Madrid, Spain

    • Trinidad Caldés
    • , José A. García-Sáenz
    •  & Atocha Romero
  58. Genomic Epidemiology Group, German Cancer Research Center (DKFZ), Heidelberg, Germany

    • Federico Canzian
  59. Genomic Medicine Group, Galician Foundation of Genomic Medicine, Instituto de Investigación Sanitaria de Santiago de Compostela (IDIS), Complejo Hospitalario Universitario de Santiago, SERGAS, Santiago De Compostela, Spain

    • Angel Carracedo
    •  & Manuela Gago-Dominguez
  60. Centro de Investigación en Red de Enfermedades Raras (CIBERER) y Centro Nacional de Genotipado (CEGEN-PRB2), Universidad de Santiago de Compostela, Santiago De Compostela, Spain

    • Angel Carracedo
  61. Epidemiology Research Program, American Cancer Society, Atlanta, Georgia, USA

    • Brian D. Carter
    • , Susan M. Gapstur
    •  & Mia M. Gaudet
  62. Oncology and Genetics Unit, Instituto de Investigacion Biomedica (IBI) Orense-Pontevedra-Vigo, Xerencia de Xestion Integrada de Vigo-SERGAS, Vigo, Spain

    • Jose E. Castelao
  63. Hong Kong Hereditary Breast Cancer Family Registry, Happy Valley, Hong Kong

    • Tsun L. Chan
    • , Ava Kwong
    •  & Edmond S. K. Ma
  64. Department of Pathology, Hong Kong Sanatorium and Hospital, Happy Valley, Hong Kong

    • Tsun L. Chan
    • , Dona N. Ho
    •  & Edmond S. K. Ma
  65. Division of Cancer Prevention and Control, Roswell Park Cancer Institute, Buffalo, New York, USA

    • Ting-Yuan David Cheng
  66. Saw Swee Hock School of Public Health, National University of Singapore, Singapore, Singapore

    • Kee Seng Chia
    • , Mikael Hartman
    •  & Hui Miao
  67. Department of Biomedical Sciences, Seoul National University Graduate School, Seoul, South Korea

    • Ji-Yeob Choi
    • , Daehee Kang
    •  & Sue K. Park
  68. Cancer Research Institute, Seoul National University, Seoul, South Korea

    • Ji-Yeob Choi
    • , Daehee Kang
    •  & Sue K. Park
  69. Westmead Institute for Medical Research, University of Sydney, Sydney, Australia

    • Christine L. Clarke
  70. Department of Clinical Genetics, Erasmus University Medical Center, Rotterdam, The Netherlands

    • Margriet Collée
    •  & Ans M. W. van den Ouweland
  71. Cancer & Environment Group, Center for Research in Epidemiology and Population Health (CESP), INSERM, University Paris-Sud, University Paris-Saclay, Villejuif, France

    • Emilie Cordina-Duverger
    • , Pascal Guénel
    •  & Thérèse Truong
  72. Department of Epidemiology and Biostatistics, School of Public Health, Imperial College London, London, UK

    • David G. Cox
  73. INSERM U1052, Cancer Research Center of Lyon, Lyon, France

    • David G. Cox
  74. Academic Unit of Pathology, Department of Neuroscience, University of Sheffield, Sheffield, UK

    • Simon S. Cross
  75. Department of Laboratory Medicine and Pathology, Mayo Clinic, Rochester, Minnesota, USA

    • Julie M. Cunningham
    • , Jeffery Meyer
    • , Jason Vollenweider
    •  & Fergus J. Couch
  76. Department of Clinical Genetics, Fox Chase Cancer Center, Philadelphia, Pennsylvania, USA

    • Mary B. Daly
  77. Department of Pathology, Leiden University Medical Center, Leiden, The Netherlands

    • Peter Devilee
  78. Department of Human Genetics, Leiden University Medical Center, Leiden, The Netherlands

    • Peter Devilee
  79. Department of Non-Communicable Disease Epidemiology, London School of Hygiene and Tropical Medicine, London, UK

    • Isabel dos-Santos-Silva
    •  & Julian Peto
  80. Southampton Clinical Trials Unit, Faculty of Medicine, University of Southampton, Southampton, UK

    • Lorraine Durcan
    •  & Tom Maishman
  81. Cancer Sciences Academic Unit, Faculty of Medicine, University of Southampton, Southampton, UK

    • Lorraine Durcan
    • , Diana M. Eccles
    •  & Tom Maishman
  82. Department of Biomedical Sciences, Faculty of Science and Technology, University of Westminster, London, UK

    • Miriam Dwek
    •  & Nadege Presneau
  83. Institute of Human Genetics, University Hospital Erlangen, Friedrich-Alexander University Erlangen-Nuremberg, Comprehensive Cancer Center Erlangen-EMN, Erlangen, Germany

    • Arif B. Ekici
  84. Channing Division of Network Medicine, Department of Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, Massachusetts, USA

    • A. Heather Eliassen
    •  & Rulla Tamimi
  85. Department of Epidemiology, Harvard T. H. Chan School of Public Health, Boston, Massachusetts, USA

    • A. Heather Eliassen
    • , JoAnn E. Manson
    • , Rulla Tamimi
    • , Walter Willett
    • , David J. Hunter
    •  & Peter Kraft
  86. Oncology and Pathology, Department of Clinical Sciences, Lund University, Lund, Sweden

    • Carolina Ellberg
  87. Department of Genetics and Fundamental Medicine, Bashkir State University, Ufa, Russia

    • Mingajeva Elvira
    • , Elza Khusnutdinova
    •  & Darya Prokofyeva
  88. Institute for Medical Informatics, Statistics and Epidemiology, University of Leipzig, Leipzig, Germany

    • Christoph Engel
  89. LIFE—Leipzig Research Centre for Civilization Diseases, University of Leipzig, Leipzig, Germany

    • Christoph Engel
  90. David Geffen School of Medicine, Department of Medicine Division of Hematology and Oncology, University of California at Los Angeles, Los Angeles, California, USA

    • Peter A. Fasching
  91. Usher Institute of Population Health Sciences and Informatics, The University of Edinburgh Medical School, Edinburgh, UK

    • Jonine Figueroa
  92. Institute for Medical Biometrics and Epidemiology, University Medical Center Hamburg-Eppendorf, Hamburg, Germany

    • Dieter Flesch-Janys
  93. Department of Cancer Epidemiology, Clinical Cancer Registry, University Medical Center Hamburg-Eppendorf, Hamburg, Germany

    • Dieter Flesch-Janys
    •  & Kathrin Thöne
  94. Breast Cancer Now Toby Robins Research Centre, The Institute of Cancer Research, London, UK

    • Olivia Fletcher
    • , Nichola Johnson
    •  & Nick Orr
  95. Department of Breast Surgery, Herlev and Gentofte Hospital, Copenhagen University Hospital, Herlev, Denmark

    • Henrik Flyger
  96. School of Public Health, Curtin University, Perth, Australia

    • Lin Fritschi
  97. Moores Cancer Center, University of California San Diego, La Jolla, California, USA

    • Manuela Gago-Dominguez
    •  & Maria Elena Martinez
  98. Department of Epidemiology, Shanghai Cancer Institute, Shanghai, China

    • Yu-Tang Gao
  99. Department of Medical Oncology, University Hospital of Heraklion, Heraklion, Greece

    • Vassilios Georgoulias
    •  & Dimitrios Mavroudis
  100. Cancer Epidemiology and Intelligence Division, Cancer Council Victoria, Melbourne, Victoria, Australia

    • Graham G. Giles
    • , Robert J. MacInnis
    •  & Roger L. Milne
  101. Centre for Epidemiology and Biostatistics, Melbourne School of Population and Global Health, The University of Melbourne, Melbourne, Australia

    • Graham G. Giles
    • , John L. Hopper
    • , Robert J. MacInnis
    • , Enes Makalic
    • , Kelly-Anne Phillips
    • , Daniel F. Schmidt
    •  & Roger L. Milne
  102. Fred A. Litwin Center for Cancer Genetics, Lunenfeld-Tanenbaum Research Institute of Mount Sinai Hospital, Toronto, Ontario, Canada

    • Gord Glendon
    •  & Irene L. Andrulis
  103. Department of Medicine, McGill University, Montréal, Quebec, Canada

    • Mark S. Goldberg
  104. Division of Clinical Epidemiology, Royal Victoria Hospital, McGill University, Montréal, Quebec, Canada

    • Mark S. Goldberg
  105. Department of Dermatology, Huntsman Cancer Institute, University of Utah School of Medicine, Salt Lake City, Utah, USA

    • David E. Goldgar
  106. Department of Surgery, Oulu University Hospital, University of Oulu, Oulu, Finland

    • Mervi Grip
  107. Department of Genetics and Pathology, Pomeranian Medical University, Szczecin, Poland

    • Jacek Gronwald
    • , Anna Jakubowska
    • , Katarzyna Kaczmarek
    •  & Jan Lubinski
  108. Centre de Recherche du Centre Hospitalier de Université de Montréal (CHUM), Université de Montréal, Montréal, Quebec, Canada

    • Anne Grundy
  109. Center for Hereditary Breast and Ovarian Cancer, University Hospital of Cologne, Cologne, Germany

    • Eric Hahnen
    •  & Rita K. Schmutzler
  110. Center for Integrated Oncology (CIO), University Hospital of Cologne, Cologne, Germany

    • Eric Hahnen
    •  & Rita K. Schmutzler
  111. Center for Molecular Medicine Cologne (CMMC), University of Cologne, Cologne, Germany

    • Eric Hahnen
    •  & Rita K. Schmutzler
  112. Department of Preventive Medicine, Keck School of Medicine, University of Southern California, Los Angeles, California, USA

    • Christopher A. Haiman
    • , Eunjung Lee
    • , Grace Sheng
    • , Daniel O. Stram
    • , Chiu-Chen Tseng
    • , David Van Den Berg
    • , Anna H. Wu
    •  & Lucy Xia
  113. Institute of Environmental Medicine, Karolinska Institutet, Stockholm, Sweden

    • Niclas Håkansson
    •  & Alicja Wolk
  114. Molecular Genetics of Breast Cancer, German Cancer Research Center (DKFZ), Heidelberg, Germany

    • Ute Hamann
    • , Guanmengqian Huang
    • , Maria Kabisch
    •  & Diana Torres
  115. Department of Biostatistics & Epidemiology, University of Massachusetts, Amherst, Amherst, Massachusetts, USA

    • Susan Hankinson
  116. Department of Health Sciences Research, Mayo Clinic, Rochester, Minnesota, USA

    • Steven N. Hart
    • , Jenna Lilyquist
    • , Aaron Norman
    • , Janet E. Olson
    • , Curtis Olswold
    • , Christopher Scott
    •  & Celine Vachon
  117. Translational Cancer Research Area, University of Eastern Finland, Kuopio, Finland

    • Jaana M. Hartikainen
    • , Veli-Matti Kosma
    • , Arto Mannermaa
    •  & Maria Tengström
  118. Institute of Clinical Medicine, Pathology and Forensic Medicine, University of Eastern Finland, Kuopio, Finland

    • Jaana M. Hartikainen
    • , Veli-Matti Kosma
    •  & Arto Mannermaa
  119. Imaging Center, Department of Clinical Pathology, Kuopio University Hospital, Kuopio, Finland

    • Jaana M. Hartikainen
    • , Veli-Matti Kosma
    •  & Arto Mannermaa
  120. Department of Surgery, National University Health System, Singapore, Singapore

    • Mikael Hartman
    •  & Chuen Neng Lee
  121. School of Population Health, University of Western Australia, Perth, Australia

    • Jane Heyworth
  122. Department of Medical Oncology, Family Cancer Clinic, Erasmus MC Cancer Institute, Rotterdam, The Netherlands

    • Antoinette Hollestelle
    • , Maartje J. Hooning
    •  & Caroline Seynaeve
  123. Division of Breast Surgery, Department of Surgery, Kaohsiung Medical University, Kaohsiung, Taiwan

    • Ming-Feng Hou
  124. Institute of Biomedical Sciences, Academia Sinica, Taipei, Taiwan

    • Chia-Ni Hsiung
  125. Division of Epidemiology and Prevention, Aichi Cancer Center Research Institute, Nagoya, Japan

    • Junko Ishiguro
    •  & Hidemi Ito
  126. Department of Epidemiology, Nagoya University Graduate School of Medicine, Nagoya, Japan

    • Junko Ishiguro
    • , Hidemi Ito
    •  & Keitaro Matsuo
  127. Division of Epidemiology, Center for Public Health Sciences, National Cancer Center, Tokyo, Japan

    • Motoki Iwasaki
    •  & Taiki Yamaji
  128. Department of Breast Oncology, Aichi Cancer Center Hospital, Nagoya, Japan

    • Hiroji Iwata
  129. Department of Gynecology and Obstetrics, University Hospital Ulm, Ulm, Germany

    • Wolfgang Janni
  130. Department of Epidemiology, Cancer Prevention Institute of California, Fremont, California, USA

    • Esther M. John
  131. Department of Health Research and Policy–Epidemiology, Stanford University School of Medicine, Stanford, California, USA

    • Esther M. John
    •  & Alice S. Whittemore
  132. Stanford Cancer Institute, Stanford University School of Medicine, Stanford, California, USA

    • Esther M. John
    •  & Alice S. Whittemore
  133. Division of Genetics and Epidemiology, The Institute of Cancer Research, London, UK

    • Michael Jones
    • , Minouk J. Schoemaker
    •  & Anthony Swerdlow
  134. Department of Oncology, Oulu University Hospital, University of Oulu, Oulu, Finland

    • Arja Jukkola-Vuorinen
  135. Department of Preventive Medicine, Seoul National University College of Medicine, Seoul, South Korea

    • Daehee Kang
    •  & Sue K. Park
  136. Department of Surgery, Nagano Matsushiro General Hospital, Nagano, Japan

    • Yoshio Kasuga
  137. School of Medicine, National University of Ireland, Galway, Ireland

    • Michael J. Kerin
    •  & Nicola Miller
  138. Department of Obstetrics and Gynecology, Helsinki University Hospital, University of Helsinki, Helsinki, Finland

    • Sofia Khan
    • , Johanna I. Kiiski
    •  & Heli Nevanlinna
  139. Department of Surgery, Daerim Saint Mary’s Hospital, Seoul, South Korea

    • Sung-Won Kim
  140. Prosserman Centre for Health Research, Lunenfeld-Tanenbaum Research Institute of Mount Sinai Hospital, Toronto, Ontario, Canada

    • Julia A. Knight
  141. Division of Epidemiology, Dalla Lana School of Public Health, University of Toronto, Toronto, Ontario, Canada

    • Julia A. Knight
  142. Institute of Clinical Medicine, Faculty of Medicine, University of Oslo, Oslo, Norway

    • Vessela N. Kristensen
  143. Department of Clinical Molecular Biology, Oslo University Hospital, University of Oslo, Oslo, Norway

    • Vessela N. Kristensen
  144. Department of Surgery, The University of Hong Kong, Pok Fu Lam, Hong Kong

    • Ava Kwong
  145. Department of Surgery, Hong Kong Sanatorium and Hospital, Happy Valley, Hong Kong

    • Ava Kwong
  146. Vesalius Research Center, VIB, Leuven, Belgium

    • Diether Lambrechts
  147. Laboratory for Translational Genetics, Department of Oncology, University of Leuven, Leuven, Belgium

    • Diether Lambrechts
  148. University of Hawaii Cancer Center, Honolulu, Hawaii, USA

    • Loic Le Marchand
  149. Department of Surgery, Soonchunhyang University College of Medicine and Soonchunhyang University Hospital, Seoul, South Korea

    • Min Hyuk Lee
  150. Department of Surgery, University of Ulsan College of Medicine and Asan Medical Center, Seoul, South Korea

    • Jong Won Lee
  151. Department of Cardiac, Thoracic and Vascular Surgery, National University Health System, Singapore, Singapore

    • Chuen Neng Lee
  152. Clalit National Cancer Control Center, Carmel Medical Center and Technion Faculty of Medicine, Haifa, Israel

    • Flavio Lejbkowicz
    • , Mila Pinchev
    • , Gadi Rennert
    •  & Hedy S. Rennert
  153. Department of Molecular Medicine and Surgery, Karolinska Institutet, Stockholm, Sweden

    • Annika Lindblom
  154. Department of Cancer Epidemiology and Prevention, M. Sklodowska-Curie Memorial Cancer Center & Institute of Oncology, Warsaw, Poland

    • Jolanta Lissowska
  155. German Breast Group GmbH, Neu Isenburg, Germany

    • Sibylle Loibl
  156. Division of Health Sciences, Warwick Medical School, Warwick University, Coventry, UK

    • Artitaya Lophatananon
    • , Kenneth Muir
    •  & Sarah Stewart-Brown
  157. Institute of Population Health, University of Manchester, Manchester, UK

    • Artitaya Lophatananon
    •  & Kenneth Muir
  158. Division of Public Health Sciences, Epidemiology Program, Fred Hutchinson Cancer Research Center, Seattle, Washington, USA

    • Kathleen E. Malone
  159. Research Centre for Genetic Engineering and Biotechnology “Georgi D. Efremov”, Macedonian Academy of Sciences and Arts, Skopje, Republic of Macedonia

    • Ivana Maleva Kostovska
    •  & Dijana Plaseska-Karanfilska
  160. Unit of Medical Genetics, Department of Preventive and Predictive Medicine, Fondazione IRCCS (Istituto Di Ricovero e Cura a Carattere Scientifico) Istituto Nazionale dei Tumori (INT), Milan, Italy

    • Siranoush Manoukian
  161. Department of Medicine, Brigham and Women’s Hospital, Harvard Medical School, Boston, Massachusetts, USA

    • JoAnn E. Manson
  162. Department of Oncology–Pathology, Karolinska Institutet, Stockholm, Sweden

    • Sara Margolin
    •  & Camilla Wendt
  163. Cancer Research Malaysia, Subang Jaya, Selangor, Malaysia

    • Shivaani Mariapun
    •  & Soo H. Teo
  164. Department of Family Medicine and Public Health, University of California San Diego, La Jolla, California, USA

    • Maria Elena Martinez
  165. Division of Molecular Medicine, Aichi Cancer Center Research Institute, Nagoya, Japan

    • Keitaro Matsuo
  166. Anatomical Pathology, The Alfred Hospital, Melbourne, Australia

    • Catriona McLean
  167. Division of Gynaecology and Obstetrics, Technische Universität München, Munich, Germany

    • Alfons Meindl
  168. Servicio de Anatomía Patológica, Hospital Monte Naranco, Oviedo, Spain

    • Primitiva Menéndez
  169. Gynaecological Cancer Research Centre, Department of Women’s Cancer, Institute for Women’s Health, University College London, London, UK

    • Usha Menon
  170. Breast Cancer Research Unit, Cancer Research Institute, University Malaya Medical Centre, Kuala Lumpur, Malaysia

    • Nur Aishah Mohd Taib
    •  & Soo H. Teo
  171. Department of Laboratory Medicine and Pathobiology, University of Toronto, Toronto, Ontario, Canada

    • Anna Marie Mulligan
  172. Laboratory Medicine Program, University Health Network, Toronto, Ontario, Canada

    • Anna Marie Mulligan
  173. Université Paris Sorbonne Cité, INSERM UMR-S1147, Paris, France

    • Claire Mulot
  174. Leuven Multidisciplinary Breast Center, Department of Oncology, Leuven Cancer Institute, University Hospitals Leuven, Leuven, Belgium

    • Patrick Neven
    • , Ann Smeets
    •  & Hans Wildiers
  175. Department of Surgery, Seoul National University College of Medicine, Seoul, South Korea

    • Dong-Young Noh
  176. Center for Clinical Cancer Genetics and Global Health, The University of Chicago, Chicago, Illinois, USA

    • Olufunmilayo I. Olopade
  177. University of New Mexico Health Sciences Center, University of New Mexico, Albuquerque, New Mexico, USA

    • V. Shane Pankratz
  178. The Curtin UWA Centre for Genetic Origins of Health and Disease, Curtin University and University of Western Australia, Perth, Australia

    • Rachel Lloyd
    •  & Jennifer Stone
  179. Servicio de Cirugía General y Especialidades, Hospital Monte Naranco, Oviedo, Spain

    • Jose I. A. Perez
  180. IFOM, The FIRC (Italian Foundation for Cancer Research) Institute of Molecular Oncology, Milan, Italy

    • Paolo Peterlongo
  181. Peter MacCallum Cancer Center, Melbourne, Australia

    • Kelly-Anne Phillips
  182. Sir Peter MacCallum Department of Oncology, The University of Melbourne, Melbourne, Australia

    • Kelly-Anne Phillips
  183. Department of Medicine, St Vincent’s Hospital, The University of Melbourne, Fitzroy, Australia

    • Kelly-Anne Phillips
  184. Laboratory of Cancer Genetics and Tumor Biology, Cancer and Translational Medicine Research Unit, Biocenter Oulu, University of Oulu, Oulu, Finland

    • Katri Pylkäs
    •  & Robert Winqvist
  185. Laboratory of Cancer Genetics and Tumor Biology, Northern Finland Laboratory Centre Oulu, Oulu, Finland

    • Katri Pylkäs
    •  & Robert Winqvist
  186. Department of Gynecology and Obstetrics, Ludwig-Maximilians University of Munich, Munich, Germany

    • Brigitte Rack
  187. Unit of Molecular Bases of Genetic Risk and Genetic Testing, Department of Preventive and Predictive Medicine, Fondazione IRCCS (Istituto Di Ricovero e Cura a Carattere Scientifico) Istituto Nazionale dei Tumori (INT), Milan, Italy

    • Paolo Radice
  188. Section of Cancer Genetics, The Institute of Cancer Research, London, UK

    • Nazneen Rahman
    •  & Sheila Seal
  189. Medical Oncology Department, Hospital Universitario Puerta de Hierro, Madrid, Spain

    • Atocha Romero
  190. Department of Oncology, Mayo Clinic, Rochester, Minnesota, USA

    • Kathryn J. Ruddy
  191. Institute of Pathology, Staedtisches Klinikum Karlsruhe, Karlsruhe, Germany

    • Thomas Rüdiger
  192. Department of Surgery, The Netherlands Cancer Institute, Antoni van Leeuwenhoek Hospital, Amsterdam, The Netherlands

    • Emiel J. T. Rutgers
  193. Hereditary Cancer Clinic, University Hospital of Heraklion, Heraklion, Greece

    • Emmanouil Saloustros
  194. Epidemiology Branch, National Institute of Environmental Health Sciences, NIH, Research Triangle Park, North Carolina, USA

    • Dale P. Sandler
    •  & Jack A. Taylor
  195. National Cancer Institute, Bangkok, Thailand

    • Suleeporn Sangrajrang
  196. Research Oncology, Guy’s Hospital, King’s College London, London, UK

    • Elinor J. Sawyer
  197. National Center for Tumor Diseases, University of Heidelberg, Heidelberg, Germany

    • Andreas Schneeweiss
    •  & Christof Sohn
  198. Department of Epidemiology and Biostatistics, Case Western Reserve University, Cleveland, Ohio, USA

    • Fredrick Schumacher
  199. Division of Molecular Medicine, Pathology North, John Hunter Hospital, Newcastle, Australia

    • Rodney J. Scott
  200. Discipline of Medical Genetics, School of Biomedical Sciences and Pharmacy, Faculty of Health, University of Newcastle, Callaghan, Australia

    • Rodney J. Scott
  201. Department of Medicine, University of Kansas Medical Center, Kansas City, Kansas, USA

    • Priyanka Sharma
  202. School of Public Health, China Medical University, Taichung, Taiwan

    • Chen-Yang Shen
  203. Taiwan Biobank, Institute of Biomedical Sciences, Academia Sinica, Taipei, Taiwan

    • Chen-Yang Shen
  204. Division of Cancer Prevention, National Cancer Institute, Rockville, Maryland, USA

    • Mark E. Sherman
  205. Genetic Epidemiology Laboratory, Department of Pathology, The University of Melbourne, Melbourne, Australia

    • Melissa C. Southey
  206. Cancer Control Research, BC Cancer Agency, Vancouver, British Columbia, Canada

    • John J. Spinelli
  207. School of Population and Public Health, University of British Columbia, Vancouver, British Columbia, Canada

    • John J. Spinelli
  208. Saarland Cancer Registry, Saarbrücken, Germany

    • Christa Stegmaier
  209. Department of Obstetrics and Gynaecology, University of Melbourne and the Royal Women’s Hospital, Melbourne, Australia

    • Jennifer Stone
  210. Division of Breast Cancer Research, The Institute of Cancer Research, London, UK

    • Anthony Swerdlow
  211. Epigenetic and Stem Cell Biology Laboratory, National Institute of Environmental Health Sciences, NIH, Research Triangle Park, North Carolina, USA

    • Jack A. Taylor
  212. Cancer Center, Kuopio University Hospital, Kuopio, Finland

    • Maria Tengström
  213. Institute of Clinical Medicine, Oncology, University of Eastern Finland, Kuopio, Finland

    • Maria Tengström
  214. Department of Epidemiology, Mailman School of Public Health, Columbia University, New York, New York, USA

    • Mary Beth Terry
  215. National Cancer Institute, Ministry of Public Health, Nonthaburi, Thailand

    • Somchai Thanasitthichai
  216. Department of Surgery, Leiden University Medical Center, Leiden, The Netherlands

    • Rob A. E. M. Tollenaar
  217. Wellcome Trust Centre for Human Genetics and Oxford NIHR Biomedical Research Centre, University of Oxford, Oxford, UK

    • Ian Tomlinson
  218. Institute of Human Genetics, Pontificia Universidad Javeriana, Bogota, Colombia

    • Diana Torres
  219. Center for Public Health Sciences, National Cancer Center, Tokyo, Japan

    • Shoichiro Tsugane
  220. Frauenklinik der Stadtklinik Baden-Baden, Baden-Baden, Germany

    • Hans-Ulrich Ulmer
  221. Cancer Registry of Norway, Oslo, Norway

    • Giske Ursin
  222. Department of Nutrition, Institute of Basic Medical Sciences, University of Oslo, Oslo, Norway

    • Giske Ursin
  223. Department of Gynecology and Obstetrics, Helios Clinics Berlin-Buch, Berlin, Germany

    • Michael Untch
  224. Department of Clinical Genetics, Leiden University Medical Center, Leiden, The Netherlands

    • Christi J. van Asperen
  225. Family Cancer Clinic, The Netherlands Cancer Institute, Antoni van Leeuwenhoek Hospital, Amsterdam, The Netherlands

    • Lizet van der Kolk
  226. Division of Biomedical Genetics, University Medical Center Utrecht, Utrecht, The Netherlands

    • Rob B. van der Luijt
  227. Department of Gynaecology and Obstetrics, University of Ulm, Ulm, Germany

    • Shan Wang-Gohrke
  228. Biostatistics and Computational Biology Branch, National Institute of Environmental Health Sciences, NIH, Research Triangle Park, North Carolina, USA

    • Clarice R. Weinberg
  229. Department of Nutrition, Harvard T. H. Chan School of Public Health, Boston, Massachusetts, USA

    • Walter Willett
  230. Subang Jaya Medical Centre, Subang Jaya, Selangor, Malaysia

    • Cheng Har Yip
  231. Seoul National University College of Medicine, Seoul, South Korea

    • Keun-Young Yoo
  232. Armed Forces Capital Hospital, Seongnam, South Korea

    • Keun-Young Yoo
  233. Department of Surgery, Tri-Service General Hospital, National Defense Medical Center, Taipei, Taiwan

    • Jyh-Cherng Yu
  234. Shanghai Municipal Center for Disease Control and Prevention, Shanghai, China

    • Ying Zheng
  235. Department of Medicine, Institute for Human Genetics, UCSF Helen Diller Family Comprehensive Cancer Center, University of California San Francisco, San Francisco, California, USA

    • Elad Ziv
  236. Pathology Queensland, The Royal Brisbane and Women’s Hospital, Brisbane 4029, Australia

    • Sunil R. Lakhani
  237. Department of Molecular Genetics, University of Toronto, Toronto, Ontario, Canada

    • Irene L. Andrulis
  238. Center for Genomic Medicine, Department of Biomedical Data Science, Geisel School of Medicine, Dartmouth College, Hanover, New Hampshire, USA

    • Christopher I. Amos
  239. University Cancer Center Hamburg (UCCH), University Medical Center Hamburg-Eppendorf, Hamburg, Germany

    • Jenny Chang-Claude
  240. Department of Oncology, Södersjukhuset, Stockholm, Sweden

    • Per Hall
  241. Division of Psychosocial Research and Epidemiology, The Netherlands Cancer Institute, Antoni van Leeuwenhoek Hospital, Amsterdam, The Netherlands

    • Marjanka K. Schmidt

Consortia

  1. NBCS Collaborators

    Lists of participants and their affiliations appear in the Supplementary Information.

  2. ABCTB Investigators

    Lists of participants and their affiliations appear in the Supplementary Information.

  3. ConFab/AOCS Investigators

    Lists of participants and their affiliations appear in the Supplementary Information.

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Contributions

Writing group: K.Mi., S.Li., J.Bee., S.Hu., S.Ka., P.So., S.L.E., G.D.B., G.C.-T., J.Si., P.K. and D.F.E. Conceived the OncoArray and obtained financial support: C.I.A., J.Si., P.K. and D.F.E. Designed the OncoArray: J.D., E.D., A. Lee, Z.W., A.C.A., C.I.A., S.J.C., P.K. and D.F.E. Led the COGS project: P.Hal. Led the DRIVE project: D.J.H. Led the PERSPECTIVE project: J.Si. Led the working groups of BCAC: A.C.A., I.L.A., P.D.P.P., J.C.-C., R.L.M., M.G.-C., M.K.S. and A.M.D. Data management: J.D., M.K.B., Q.Wan., R.Ke., U.E., S.B., J.C.-C. and M.K.S. Bioinformatics analysis: J.D., J.Bee., A.Lem., P.So., J.A., M.Gh., J.C., A.D., A.E.M.R., S.R.L. Statistical analysis: K.Mi., S.Li., S.Hu., S.Ka., A.Ros., J.T., X.Q.C., L.Fa., X.J., H.Fi., G.D.B., P.K. and D.F.E. Functional analysis: D.G., X.Q.C., J.Bee., J.D.F., K.Mc., S.L.E. and G.C.-T. OncoArray genotyping: M.A., F.B., C.Ba., D.M.C., J.M.C., K.F.D., N.Ha., B.H., K.J., C.L., J.Me., E.P., J.R., G.S., D.C.T., D.V.D.B., D.V., J.V., L.X., B.Z. and A.M.D. Provided DNA samples and/or phenotypic data: M.A.A., H.A., K.A., H.A.-C., N.N.A., V.A., K.J.A., B.A., P.L.A., M.Ba., M.W.B., J.Ben., M.Be., L.Be., C.Bl., N.V.B., S.E.B., B.Bo., A.-L.B.-D., J.S.B., H.Bra., P.Bre., H.Bre., L.Br., P.Bro., I.W.B., A.B., A.B.-W., S.Y.B., T.B., B.Bu., K.B., H.Ca., Q.C., T.C., F.C., A.Ca., B.D.C., J.E.C., T.L.C., T.-Y.D.C., K.S.C., J.-Y.C., H.Ch., C.L.C., M.C., E.C.-D., S.C., A.Co., D.G.C., S.S.C., K.C., M.B.D., P.D., T.D., I.d.-S.-S., M.Du., L.D., M.Dw., D.M.E., A.B.E., A.H.E., C.El., M.El., C.En., M.Er., P.A.F., J.F., D.F.-J., O.F., H.Fl., L.Fr., V.Ga., M.Ga., M.G.-D., Y.-T.G., S.M.G., J.A.G.-S., M.M.G., V.Ge., G.G.G., G.G., M.S.G., D.E.G., A.G.-N., G.I.G.A., M.Gr., J.G., A.G., P.G., L.H., E.H., C.A.H., N.Hå., U.H., S.Ha., P.Har., S.N.H., J.M.H., M.H., A.He., J.H., P.Hi., D.N.H., A.Ho., M.J.H., R.N.H., J.L.H., M.-F.H., C.-N.H., G.H., K.H., J.I., H.It., M.I., H.Iw., A.J., W.J., E.M.J., N.J., M.J., A.J.-V., R.Ka., M.K., K.K., D.K., Y.K., M.J.K., S.Kh., E.K., J.I.K., S.-W.K., J.A.K., V.-M.K., I.M.K., V.N.K., U.K., A.K., D.L., L.L.M., C.N.L., E.L., J.W.L., M.H.L., F.L., J. Li, J.Lil., A.Li., J.Lis., R.L., W.-Y.L., S.Lo., J.Lo., A.Lo., J.Lu., M.P.L., E.S.K.M., R.J.M., T.M., E.M., K.E.M., A.Ma., S.Man., J.E.M., S.Marg., S.Mari., M.E.M., K.Ma., D.M., J.Mc., C.Mc., H.M.-H., A.Me., P.M., U.M., H.M., N.M., K.Mu., A.M.M., C.Mu., S.L.N., H.N., P.N., S.F.N., D.-Y.N., B.G.N., A.N., O.I.O., J.E.O., H.O., C.O., N.O., V.S.P., S.K.P., T.-W.P.-S., J.I.A.P., P.P., J.P., K.-A.P., M.P., D.P.-K., R.P., N.P., D.P., K.P., B.R., P.R., N.R., G.R., H.S.R., V.R., A.Rom., K.J.R., T.R., A.Ru., M.R., E.J.T.R., E.S., D.P.S., S.Sa., E.J.S., D.F.S., R.K.S., A.Sc., M.J.Sc., F.S., P.Sc., C.Sc., R.J.S., S.Se., C.Se., M.S., P.Sh., C.-Y.S., M.E.S., M.J.Sh., X.-O.S., A.Sm., C.So., M.C.S., J.J.S., C.St., S.S.-B., J.St., D.O.S., H.S., A.Sw., N.A.M.T., R.T., J.A.T., M.T., S.H.T., M.B.T., S.Th., K.T., R.A.E.M.T., I.T., L.T., D.T., T.T., C.-C.T., S.Ts., H.-U.U., M.U., G.U., C.V., C.J.v.A., A.M.W.v.d.O., L.v.d.K., R.B.v.d.L., Q.Wai., S.W.-G., C.R.W., C.W., A.S.W., H.W., W.W., R.W., A.W., A.H.W., T.Y., X.R.Y., C.H.Y., K.-Y.Y., J.-C.Y., W.Z., Y.Z., A.Z., E.Z., ABCTB Investigators, kConFab/AOCS Investigators, NBCS Collaborators, A.C.A., I.L.A., F.J.C., P.D.P.P., J.C.-C., P.Hal., D.J.H., R.L.M., M.G.-C., M.K.S., G.D.B., J.Si., P.K. and D.F.E. All authors read and approved the final version of the manuscript.

Competing interests

The authors declare no competing financial interests.

Corresponding author

Correspondence to Douglas F. Easton.

Publisher's note: Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Extended data

Supplementary information

PDF files

  1. 1.

    Supplementary Information

    This file contains Supplementary Tables 1, 2 and 24, a Supplementary Note, Funding Information, Study Acknowledgments, Members of Consortia listed as Authors, Supplementary References and a Supplementary Table Guide.

  2. 2.

    Reporting Summary

Excel files

  1. 1.

    Supplementary Data

    This file contains Supplementary Tables 3-5, 7, 9-12, 14-17, 21-23 and 25-33 – see the Supplementary Table Guide in the Supplementary Information document for full descriptions.

  2. 2.

    Supplementary Table 6

    This table contains sixty-five newly identified susceptibility loci for overall breast cancer.

  3. 3.

    Supplementary Table 8

    This table contains summary statistics for all variants for which the association with overall breast cancer in the combined dataset was significant at P<0.00001.

  4. 4.

    Supplementary Table 13

    This table contains a list of 2,221 credible variants at 65 novel loci, with annotations, UCSC Genome Browser links, and sources for genomic annotation data.

  5. 5.

    Supplementary Table 18

    This table displays eQTL associations significant at P<0.05 in the TCGA and METABRIC datasets, for credible risk variants from the analyses for overall breast cancer (see Methods), together with the corresponding results for the most significant eQTL association in the region for the same gene.

  6. 6.

    Supplementary Table 19

    This table contains summary INQUISIT gene prediction scores.

  7. 7.

    Supplementary Table 20

    This table contains detailed INQUISIT gene prediction scores.