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All data generated or analysed during this study are included in this
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Acknowledgements We thank D. Hegan and A. Dhawan for their assistance. This work was
supported by the NIH (grants R01ES005775 and R35CA197574 to P.M.G., and R01CA215453 to R.S.B.)
and by the American Cancer Society (Research Scholar Grant to R.S.B.). P.L.S. was supported by the
NIH National Institute of General Medical Sciences training grant T32GM007223.
Author contributions P.L.S. designed and performed experiments and contributed to all
aspects of the study. S.O. designed experiments and performed and analysed imaging studies
and laser micro-stripe irradiation. J.D. and N.E. performed DNA repair and cell biology assays
and contributed to data analysis and compiling of the manuscript. Y.L. designed and
performed the tumour growth delay assays. B.S., X.B. and K.N. provided reagents and
contributed to the experiments. L.M. and M.C.K. designed and performed end-resection
assays. J.L. performed LC/MS assays. P.M.G. and R.S.B. conceptualized and supervised the
study and interpreted the data. P.M.G., R.S.B., M.C.K. and P.L.S. wrote the manuscript.Competing interests R.S.B., P.M.G. and P.L.S. are inventors on US patent application no.
62/344,678 submitted by and assigned to Yale University, which covers compositions and
methods for targeting and treating HDR-deficient tumours. P.M.G and R.S.B are founders of and
consultants for Cybrexa Therapeutics. P.M.G. is a consultant for pHLIP. Neither company has
any conceptual or financial connection to this manuscript.Additional information
Supplementary information is available for this paper at https://doi.org/10.1038/s41586-020-
2363-0.
Correspondence and requests for materials should be addressed to R.S.B. or P.M.G.
Peer review information Nature thanks Brendan Price, Yue Xiong and the other, anonymous,
reviewer(s) for their contribution to the peer review of this work.
Reprints and permissions information is available at http://www.nature.com/reprints.