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Mechanistically, the classic Smad-dependent TGF-?1 signaling functions through SMAD2/3 phosphorylation and the recruitment of SMAD4. 23 , 24 Alternatively, c-Jun N-terminal kinases (JNKs)/Jun proto-oncogene ( c-Jun ), mitogen-activated protein kinase, phosphatidylinositol 3-kinase-AKT-forkhead O (FOXO) and RHO function as the Smad-independent pathways, depending on cellular context.

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