However, Fig. 4 shows that the restoration of SPAG9 expression in miR-141-transfected cells significantly reversed the inhibitory effects of miR-141 on the migration and invasion of HCC cells, indicating that the SPAG9-JNK-MMP9 axis is also engaged in the anti-metastasis effect of miR-141 on HCC. Here, SPAG9 is linked to hepatocellular carcinoma.