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• W2038598921 abstract "The proneural genes of the achaete-scute complex (AS-C; achaete [ac], scute [sc], and lethal of scute [l'sc]) and atonal (ato) encode basic helix-loop-helix (bHLH) transcriptional activators that initiate neurogenesis in Drosophila (3Jarman A.P. Grau Y. Jan L.Y. Jan Y.N. Cell. 1993; 73: 1307-1321Abstract Full Text PDF PubMed Scopus (440) Google Scholar, 9Modolell J. Perspect. Dev. Neurobiol. 1997; 4: 285-296PubMed Google Scholar). The spatial pattern of their activity is refined by multiple levels of negative control: their transcription (at least in the case of ac) is directly repressed by the bHLH protein Hairy, while their regulatory activity is antagonized by direct protein–protein interaction with the inhibitory HLH protein Extramacrochaetae (Emc) (9Modolell J. Perspect. Dev. Neurobiol. 1997; 4: 285-296PubMed Google Scholar). Subsequently, during lateral inhibition, a Notch pathway–mediated cell–cell signaling process, proneural gene expression appears to be directly repressed by bHLH proteins encoded by several Notch-regulated genes of the Enhancer of split complex [E(spl)-C] (11Ohsako S. Hyer J. Panganiban G. Oliver I. Caudy M. Genes Dev. 1994; 8: 2743-2755Crossref PubMed Scopus (228) Google Scholar, 14Van Doren M. Bailey A. Esnayra J. Ede K. Posakony J. Genes Dev. 1994; 8: 2729-2742Crossref PubMed Scopus (219) Google Scholar, 4Jimenez G. Ish-Horowicz D. Mol. Cell. Biol. 1997; 17: 4355-4362PubMed Google Scholar). The novel non-bHLH protein encoded by the m4 gene of the E(spl)-C (5Klämbt C. Knust E. Tietze K. Campos-Ortega J. EMBO J. 1989; 8: 203-210Crossref PubMed Scopus (199) Google Scholar), which is similarly activated by Notch signaling, may also be involved in regulation of the proneural genes; it is related to another small protein with an apparent role in lateral inhibition, Bearded (Brd) (7Leviten M.W. Posakony J.W. Dev. Biol. 1996; 176: 264-283Crossref PubMed Scopus (30) Google Scholar, 8Leviten M.W. Lai E.C. Posakony J.W. Development. 1997; 124: 4039-4051PubMed Google Scholar). The 3′ untranslated regions (3′ UTRs) of Brd, hairy, and many genes of the E(spl)-C contain a novel class of sequence motif, the GY box (GYB, GUCUUCC); emc contains the variant sequence GUUUUCC (6Lai E.C. Posakony J.W. Development. 1997; 124: 4847-4856PubMed Google Scholar, 8Leviten M.W. Lai E.C. Posakony J.W. Development. 1997; 124: 4039-4051PubMed Google Scholar; Figure 1A). Recently, we have also recognized that the 3′ UTRs of three proneural genes include a second type of sequence element, the proneural box (PB, AAUGGAAGACAAU; Figure 1A). The full 13 nt PB is found once each in ac, l'sc, and ato, along with a second, variant version in both l'sc and ato (Figure 1A). The presence of these motifs in such distantly related paralogs as hairy and certain bHLH genes of the E(spl)-C (for the GYB), and ato and two genes of the AS-C (for the PB), indicates that both classes of sequence element are subject to strong selection. Furthermore, both the PB and the GYB are conserved in the orthologs of ac and E(spl)m4 from the distantly related Drosophilids D. virilis and D. hydei, respectively (Figure 1A), though these 3′ UTRs are otherwise quite divergent from their D. melanogaster counterparts. These findings strongly suggest functional roles for both of these sequence elements. Intriguingly, the central 7 nt of the PB and the GYB are exactly complementary, and are often located within extensive regions of RNA:RNA duplex predicted to form between PB- and GYB-containing 3′ UTRs (Figure 1B). Indeed, using in vitro assays we have observed RNA duplex formation between the ato/Brd and ato/m4 3′ UTR pairs that is PB- and GYB-dependent (our unpublished results). It is noteworthy that the predicted duplex interactions involving the GYB of Brd are significantly stronger than those involving the GYBs of the other transcripts. For example, Brd and ato are perfectly complementary over 18 contiguous nucleotides (Figure 1B). This difference in the degree of PB:GYB-associated complementarity is likely to have functional consequences. In C. elegans, small antisense RNAs encoded by lin-4 mediate translational repression of lin-14 and lin-28 transcripts by binding to complementary sequences in their 3′ UTRs (13Slack F. Ruvkun G. Annu. Rev. Genet. 1997; 31: 611-634Crossref PubMed Scopus (88) Google Scholar). We suggest that, in Drosophila, PB- and GYB-bearing transcripts may likewise participate in a regulatory mechanism mediated by RNA:RNA duplexes, but with the feature that both partners are mRNAs that also direct the synthesis of functionally interacting proteins. The opportunity to form such duplexes clearly exists, as transcripts from proneural genes and their regulators very frequently accumulate in coincident or overlapping patterns (12Singson A. Leviten M.W. Bang A.G. Hua X.H. Posakony J.W. Genes Dev. 1994; 8: 2058-2071Crossref PubMed Scopus (116) Google Scholar). Moreover, while 7 nt is the minimum length of complementarity between any PB and any GYB, the longest possible uninterrupted duplex between a given GYB-bearing transcript and a given proneural partner is almost always considerably longer (8–12 nt). It is worth noting that in a lin-4/lin-14 duplex that has been shown to be sufficient for proper regulation in vivo, the longest region of uninterrupted complementarity is only 7 nt (1Ha I. Wightman B. Ruvkun G. Genes Dev. 1996; 10: 3041-3050Crossref PubMed Scopus (176) Google Scholar). The formation of the postulated RNA duplexes may serve to regulate proneural gene function, consistent with the known roles of hairy, emc, and the bHLH genes of the E(spl)-C. This might explain occasional C-to-U transitions in the GYB sequence (in emc and D. hydei m4; Figure 1A); these variants retain complementarity with the PB due to G:U base-pairing. It is equally plausible that GYB-containing transcripts are regulated by duplex formation. A third very interesting possibility is that RNA:RNA duplexes formed between PB- and GYB-containing transcripts function to initiate a downstream regulatory activity affecting as-yet-unknown targets. Ample precedent exists establishing the trans-regulatory potency of double-stranded RNA (2Jacobs B.L. Langland J.O. Virology. 1996; 219: 339-349Crossref PubMed Scopus (504) Google Scholar, 10Nicholson A.W. Progr. Nucleic Acid Res. Mol. Biol. 1996; 52: 1-65Crossref PubMed Google Scholar, 15Williams B.R. Biochem. Soc. Trans. 1997; 25: 509-513Crossref PubMed Scopus (135) Google Scholar). In any case, the apparent capacity of transcripts from the proneural genes and their regulators to form duplexes in their 3′ UTRs suggests further complexity in the already complex regulatory interactions that control Drosophila neurogenesis." @default.
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• W2038598921 title "Regulation of Drosophila Neurogenesis byRNA:RNA Duplexes?" @default.
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