TERT and hepatocellular carcinoma: Importantly, PJA1 knockdown significantly increases binding of SMAD3 and SPTBN1 to the TERT promoter, thereby transcriptionally increasing TERT gene expression in HCC cells, suggesting that PJA1 may also regulate TERT activity through TGF-β and/or CTCF-mediated epigenetic regulation (Figure 1) [22].