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Data availability

The data that support the findings of this study are shown in Supplementary Table 1 and are available from the corresponding authors upon reasonable request.


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We thank E. Tiensuu Janson, S. Enroth and C. Polychronakos for review of the manuscript. K. Ström is acknowledged for collection of samples from the ULSAM cohort. The study was sponsored by funding from the European Research Council (ERC) Starting Grant, the Swedish Research Council, the Olle Enqvist Byggmästare Foundation, the Kjell and Märta Beijers Foundation to L.A.F. and by the Swedish Cancer Society, the Swedish Research Council, Konung Gustav V:s och Drottning Viktorias Frimurarestiftelse and the Science for Life Laboratory, Uppsala, to J.P.D. Genotyping and sequencing were performed by the SNP&SEQ Technology Platform in Uppsala, which is part of the Science for Life Laboratory at Uppsala University and is supported as a national infrastructure by the Swedish Research Council.

Author information


  1. Departments of Immunology, Genetics and Pathology, and Science for Life Laboratory, Uppsala University, Uppsala, Sweden

    • Lars A. Forsberg
    • , Jonatan Halvardson
    • , Edyta Rychlicka-Buniowska
    • , Marcus Danielsson
    • , Behrooz Torabi Moghadam
    • , Jonas Mattisson
    • , Chiara Rasi
    • , Hanna Davies
    •  & Jan P. Dumanski
  2. Beijer Laboratory of Genome Research, Uppsala University, Uppsala, Sweden

    • Lars A. Forsberg
  3. Department of Medical Sciences, Uppsala University, Uppsala, Sweden

    • Lars Lind
  4. Department of Public Health and Caring Sciences/Geriatrics, Uppsala University, Uppsala, Sweden

    • Vilmantas Giedraitis
    • , Lars Lannfelt
    • , Lena Kilander
    •  & Martin Ingelsson
  5. Faculty of Pharmacy, Medical University of Gdansk, Gdansk, Poland

    • Jan P. Dumanski


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Competing interests

J.P.D. and L.A.F. are cofounders and shareholders in Cray Innovation AB.

Corresponding authors

Correspondence to Lars A. Forsberg or Jan P. Dumanski.

Supplementary information

  1. Supplementary Text and Figures

    Supplementary Figures 1–5, Supplementary Table 1 and Supplementary Note

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