Wellcome Trust 223743/Z/21/Z Medical Research Council MC_UU_00034/4 MC_UU_00034/7 MR/R021562/1 MC_UU_00034/2 MC_UU_12014/8 MC_UU_00034/4 European Research Council (ERC) 101001634 EPSRC V011359/1 (P)