The Piwi/Argonaute protein family plays important roles in stem-cell self-renewal, RNA silencing and translational regulation in various organisms. The family members are defined by conserved both PAZ (Piwi Argonaut and Zwille) and Piwi domains participate in RNA binding. Piwi-interacting RNA (piRNA) is a class of small RNA molecules that is expressed uniquely in mammalian testes and forms RNA-protein complexes with Piwi proteins. These small regulatory RNAs have been implicated in spermatogenesis, repression of retrotransposon transposition in germline cells, epigenetic regulation and positive regulation of translation and mRNA stability. MIWI (PIWIL1) and MILI (PIWIL2) are mouse homologues of Piwi. MILI is expressed from primordial germ cells (PGC) in embryo, whereas MIWI is detected later from the mid-pachytene stage to early round spermatids. Their expression overlaps in the mid-pachytene and early round spermatid stage. The expression patterns and knockout phenotypes of mammalian Piwi members indicated MILI is an essential factor for meiotic differentiation during spermatogenesis. It was also reported that MILI (PIWIL2) is widely expressed in various tumors and acts as an oncogene by inhibiting apoptosis and promoting proliferation through the activation of Stat3/Bcl-xL pathway.
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