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Enucleation, a rate-limiting step of erythrocyte maturation, is tightly regulated via coordination of multiple transcription factors (FOXO3, E2F2, etc.), miRNAs (miR-30a, miR-191, miR-181a), cytoskeleton proteins (F-actin, dynein, tropomodulin, etc.), and kinases (p38 MAPK, p38 mitogen-activated protein kinase) [33]

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