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USP4 leads to the deubiquitination of SMAD4. 3 / 3
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"Interestingly, these speculations have been confirmed that HECW1 can promote the proliferation, migration and invasion of cancer cells by inducing the ubiquitination and degradation of SMAD4 (34), while USP4 can mediate the deubiquitination of SMAD4, leading to the expression of apoptotic proteins (35) ."

"We demonstrated that ubiquitin-specific protease (USP) 4 strongly induces activin/BMP signaling by removing the inhibitory monoubiquitination from SMAD4."

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"USP4 inhibits SMAD4 monoubiquitination and promotes activin and BMP signaling."