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We are grateful to H. Finucane and Y. Reshef for helpful discussions. This research was conducted using the UK Biobank Resource under application 10438 and was supported by US National Institutes of Health grants R01 HG006399, R01 GM105857 and R01 MH107649 (A.L.P.), a Burroughs Wellcome Fund Career Award at the Scientific Interfaces and the Next Generation Fund at the Broad Institute of MIT and Harvard (P.-R.L.), and a Boehringer Ingelheim Fonds fellowship (A.P.S.). Computational analyses were performed on the Orchestra High-Performance Compute Cluster at Harvard Medical School, which is partially supported by grant NCRR 1S10RR028832-01.

Author information


  1. Division of Genetics, Department of Medicine, Brigham and Women’s Hospital and Harvard Medical School, Boston, MA, USA

    • Po-Ru Loh
  2. Program in Medical and Population Genetics, Broad Institute of MIT and Harvard, Cambridge, MA, USA

    • Po-Ru Loh
    • , Steven Gazal
    • , Armin P. Schoech
    •  & Alkes L. Price
  3. Bioinformatics Interdepartmental Program, University of California, Los Angeles, Los Angeles, CA, USA

    • Gleb Kichaev
  4. Department of Epidemiology, Harvard T.H. Chan School of Public Health, Boston, MA, USA

    • Steven Gazal
    • , Armin P. Schoech
    •  & Alkes L. Price
  5. Department of Biostatistics, Harvard T.H. Chan School of Public Health, Boston, MA, USA

    • Armin P. Schoech
    •  & Alkes L. Price


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P.-R.L. and A.L.P. designed the study. P.-R.L., G.K., S.G. and A.P.S. performed analyses. All authors wrote the manuscript.

Competing interests

The authors declare no competing interests.

Corresponding authors

Correspondence to Po-Ru Loh or Alkes L. Price.

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    Supplementary Figures 1 and 2, Supplementary Tables 1–9 and Supplementary Note

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