In addition, although the three target genes of miR-33b, HMGA2, SALL4 and Twist1, are EMT inducers, our gain-of-function and loss-of-function assays in different breast cancer cell lines and a mammary epithelial cell line showed that miR-33b regulation of tumor metastasis is independent of the EMT-related program. Here, TWIST1 is linked to breast cancer.