Recent findings demonstrate that the autophagy regulator UVRAG (UV radiation resistance-associated) is subjected to non-proteolytic ubiquitylation by the HECT E3 ligase SMURF1 (SMAD specific E3 ubiquitin-protein ligase 1), which promotes autophagosome maturation and inhibits HCC tumor growth in vivo115 (Fig. 4). Here, SMURF1 is linked to neoplasm.