ATR and cancer: Notably, APE1 forms biomolecular condensates in vitro and in nucleoli independent of its nuclease activity to promote the ATR-Chk1 DDR pathway activation in cancer cells, and APE1 is proposed as a new direct activator of the ATR kinase, in addition to TopBP1 and ETAA1 (Li and Yan, 2023; Li et al., 2022).