After treatment with TUG1 ASO for 24 h, FCM showed significant decrease in DNA replication rate (i.e. EdU incorporation, Supplementary Fig. 4f) and increased numbers of γ-H2AX-positive cells in the S phase of HeLa/Fucci2 and other cancer cell lines (Fig. 6b, Supplementary Fig. 8a, b). Here, H2AX is linked to cancer.