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Armitage determines Piwi−piRISC processing from precursor formation and quality control to inter‐organelle translocation

Piwi and piRNA form the piRNA‐induced silencing complex (piRISC) to repress transposons. In the current model, Armitage (Armi) brings the Piwi–piRISC precursor (pre‐piRISC) to mitochondria, where Zucchini‐dependent piRISC maturation occurs. Here, we show that Armi is necessary for Piwi–pre‐piRISC formation at Yb bodies and that Armi triggers the exit of Piwi–pre‐piRISC from Yb bodies and the translocation to mitochondria. Piwi–pre‐piRISC resist leaving Yb bodies until Armi binds Piwi–pre‐piRISC through the piRNA precursors. The lack of the Armi N‐terminus also blocks the Piwi–pre‐piRISC exit from Yb bodies. Thus, Armi determines Piwi–piRISC processing, in a multilayered manner, from precursor formation and quality control to inter‐organelle translocation for maturation.