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USP4 deubiquitinates SMAD4. 3 / 4
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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)."

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"However, USP4 was also identified to regulate deubiquitination of Smad4 and was excluded in this study as the content of USP4 in HUVECs was much lower than USP10."

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