The piwi gene of Drosophila belongs to a novel class of evolutionarily conserved genes (the piwi or Argonaute family). The Piwi familyμ is required for germ- and stem-cell development in invertebrates, and two Piwi members, MIWI and MILI (Miwi like; Piwil2; Piwi-like protein 2), are essential for spermatogenesis in mouse. MIWI and MILI partner with Piwi-interacting RNAs (piRNAs) which are 26?C31 nucleotides (nt) in lengthμclearly distinct from the 21?C23 nt of microRNAs (miRNAs) or short interfering RNAs (siRNAs). A search for murine small RNAs that might program Piwi proteins for transposon suppression revealed developmentally regulated piRNA loci, some of which resemble transposon master control loci of Drosophila. MILI might be programmed by distinct piRNA populations at different stages of germ cell development, suggesting MILI plays an important role in meiotic prophase progression.
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