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The  Drosophila oskar  ( osk ) mRNA is unusual in that it has both coding and noncoding functions. As an mRNA,  osk  encodes a protein required for embryonic patterning and germ cell formation. Independent of that function, the absence of  osk  mRNA disrupts formation of the karyosome and blocks progression through oogenesis. Here we show that loss of  osk  mRNA also affects the distribution of regulatory proteins, relaxing their association with large RNPs within the germline, and allowing them to accumulate in the somatic follicle cells. This and other noncoding functions of the  osk  mRNA are mediated by multiple sequence elements with distinct roles. One role, provided by numerous binding sites in two distinct regions of the  osk  3′ UTR, is to sequester the translational regulator Bruno (Bru), which itself controls translation of  osk  mRNA. This defines a novel regulatory circuit, with Bru restricting the activity of  osk , and  osk  in turn restricting the activity of Bru. Other functional elements, which do not bind Bru and are positioned close to the 3′ end of the RNA, act in the oocyte and are essential. Despite the different roles played by the different types of elements contributing to RNA function, mutation of any leads to accumulation of the germline regulatory factors in the follicle cells.

oskar RNA plays multiple noncoding roles to support oogenesis and maintain integrity of the germline/soma distinction