MicroRNAs (miRNAs) are one of the most important regulators of gene expression.Their biogenesis starts with the transcription of primary miRNA transcripts (pri-miRNAs).In plants, the RNase Ⅲ enzyme DICER-LIKER 1 (DCL1), together with its partner, the double-stranded RNA-binding protein HYPONASTIC LEAVES 1 (HYL1), and the zinc-finger protein SERRATE (SE), process the imperfect miRNA-residing stem loop of pri-miRNAs to release the miRNA/miRNA* duplexes in the nucleus, which are then methylated by HUA1 ENHANCER 1 (Rogers and Chen, 2013;Song et al., 2019).Then, most miRNAs are sorted into AGONAUTE 1 to mediate mRNA-cleavage or translation inhibition based on sequence complementarity (Rogers and Chen, 2013;Song et al., 2019).Studies have shown that miRNA biogenesis is controlled through modulating pri-miRNA transcription, stability and processing, which is often coupled with various developmental and environmental signals.