Overview of the miRNA biogenesis pathway. Pri-miRNAs are transcribed by RNA polymerase II in the nucleus. The first step is mediated by a microprocessor, comprised of Drosha and DGCR8, to produce pre-miRNA. Next, pre-miRNAs are exported to the cytoplasm by EXP5. The second step is mediated by Dicer, TRBP and AGO1–4, during which pre-miRNA are processed. Finally, the remaining strand of miRNA is loaded on AGO to form RISC.