JOSD2 and cholangiocarcinoma: This observation was consistent with our previous study, where we identified YAP/TAZ as the substrates for JOSD2 in cholangiocarcinoma.20 While phosphorylation of LKB1 at Ser428 and AMPKα at Thr172, two well-established readouts of LKB1/AMPK pathway, were predominantly upregulated upon JOSD2 knockdown (Fig. 2c and Supplementary Fig. S3c).