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Effects of USP4, USP15, and USP9X on TGF-β type I receptor and SMADs. As depicted schematically, USP4 associates with and deubiquitinates TβRI and recruits other DUBs. USP15 binds TβRI via SMAD7 and competes with SMURF2 to balance TβRI ubiquitination. The transcriptional activity of R-SMAD can be restricted by the SMURFs via monoubiquitination and reversed by USP15. USP9X reverses SMAD4 monoubiquitination that can be conjugated by Ectodermin/TIF1γ E3 ligase
