microRNA cell-to-cell and vascular movement requires the Arabidopsis EXPORTIN5 ortholog

In a recent EMBO Journastudy the Voinnet group (IMBP) identifies the Arabidopsis EXPORTIN5 ortholog as the first component required for both cell-​to-cell and long-​distance movement of plant microRNAs. This work clarifies a process that, while agronomically important, had remained cryptic so far.

In the reporter line used, an artificial (a)microRNA produced only in the phloem companion-​cells is reprogrammed to target a photosynthesis-​related mRNA. The ensuing photo-​bleaching expands well beyond the veins, indicating amicroRNA movement.
In the reporter line used, an artificial (a)microRNA produced only in the phloem companion-​cells is reprogrammed to target a photosynthesis-​related mRNA. The ensuing photo-​bleaching expands well beyond the veins, indicating amicroRNA movement.

Gene regulation by RNA silencing mediated by small RNAs (sRNAs) is widespread in eukaryotic life including in plants. Remarkably, a class of plant sRNAs called microRNAs act not only in the cells where they are produced, but also at a distance, by travelling cell-​to-cell and through the vasculature. Mobile miRNAs act as key positional signals during development but also as long-​distance stress regulators: being used as systemic signals, they can adapt, in near-​real time, the gene expression landscape of stress-​distal growing points, thereby contributing to the remarkable phenotypic plasticity of plants.
Since the underlying mechanisms have remained unknown, the Voinnet group devised a visual reporter for microRNA movement and conducted a forward genetic screen for movement-​deficient mutants, which isolated many alleles of HASTY (HST), the Arabidopsis XPO5 ortholog. They found that, independently of its exportin activity, HST cell-​autonomously regulates cell-​to-cell and vasculature-​dependent microRNA movement in microRNA-​emitting, but not microRNA-​receiving, tissues. This control is likely coupled with microRNA biogenesis in the nucleus of miRNA-​emitting cells. HST is not required for movement of a diffusible protein or small interfering (si)RNAs, indicating a miRNA-​specific function. This discovery prompts the intriguing and somewhat counterintuitive idea that sRNA movement can be cell-​autonomously regulated including via nuclear cues integrated in silencing-​incipient cells.

Link to the publication in external pageEMBO Journal

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