Control of transcription and miRNA biogenesis by TGF-β/BMP signaling. At the core of this pathway, ligand binding to Type I/II heterodimeric receptors induces phosphorylation and nuclear translocation of SMAD transcription factors, which directly regulate the expression of protein-coding and miRNA genes. Mammalian SMAD proteins can also bind to the double-stranded stems of certain pri-miRNAs that bear CAGAC motifs, and promote their cleavage by the RNase III enzyme Drosha and its dsRBD partner DGCR8. This enhances the production of CAGAC-bearing pre-miRNAs, which are exported to the cytoplasm, cleaved by Dicer, and loaded into Argonaute (Ago) complexes to repress target genes.