SMAD4 and colorectal carcinoma: Furthermore, overexpression of PLS3 in CRC cells significantly increased the expression levels of EMT-related transcription factors (TWIST, SNAI1, SLUG, SMAD4, and ZEB1), EMT markers (vimentin, FN1, and N-cadherin), and TGF-β, enhancing the invasiveness of CRC cells [28].