[1] Univ Oxford, John Radcliffe Hosp, Weatherall Inst Mol Med, Oxford OX3 9DS, England
[2] Univ Oxford, Canc Res UK, Oxford OX3 7DQ, England
[3] Univ Oxford, Med Res Council Oxford Inst Radiat Oncol, Dept Oncol, Oxford OX3 7DQ, England
[4] Univ Cambridge, Wellcome Trust, Tennis Court Rd, Cambridge CB2 1QN, England
[5] Univ Cambridge, Canc Res UK Gurdon Inst, Tennis Court Rd, Cambridge CB2 1QN, England
[6] Univ Cambridge, Dept Biochem, Tennis Court Rd, Cambridge CB2 1GA, England