@Article{Ilik_Maticzka_Georgiev-mutua_exclu_stem-2017,
  author =	 {Ilik, Ibrahim Avsar and Maticzka, Daniel and Georgiev, 
                  Plamen and Gutierrez, Noel Marie and Backofen, Rolf and 
                  Akhtar, Asifa},
  title =	 {A mutually exclusive stem-loop arrangement in {roX2} {RNA} 
                  is essential for {X}-chromosome regulation in {Drosophila}},
  journal =	 {Genes Dev},
  year =	 {2017},
  volume =	 {31},
  number =	 {19},
  pages =	 {1973-1987},
  user =	 {maticzkd},
  pmid =	 {29066499},
  doi = 	 {10.1101/gad.304600.117},
  issn = 	 {0890-9369},
  issn = 	 {1549-5477},
  abstract =	 {The X chromosome provides an ideal model system to study 
                  the contribution of RNA-protein interactions in epigenetic 
                  regulation. In male flies, roX long noncoding RNAs (lncRNAs) 
                  harbor several redundant domains to interact with the 
                  ubiquitin ligase male-specific lethal 2 (MSL2) and the RNA 
                  helicase Maleless (MLE) for X-chromosomal regulation. 
                  However, how these interactions provide the mechanics of 
                  spreading remains unknown. By using the uvCLAP (UV 
                  cross-linking and affinity purification) methodology, which 
                  provides unprecedented information about RNA secondary 
                  structures in vivo, we identified the minimal functional 
                  unit of roX2 RNA. By using wild-type and various MLE mutant 
                  derivatives, including a catalytically inactive MLE 
                  derivative, MLE(GET), we show that the minimal roX RNA 
                  contains two mutually exclusive stem-loops that exist in a 
                  peculiar structural arrangement: When one stem-loop is 
                  unwound by MLE, an alternate structure can form, likely 
                  trapping MLE in this perpetually structured region. We show 
                  that this functional unit is necessary for dosage 
                  compensation, as mutations that disrupt this formation lead 
                  to male lethality. Thus, we propose that roX2 lncRNA 
                  contains an MLE-dependent affinity switch to enable 
                  reversible interactions of the MSL complex to allow dosage 
                  compensation of the X chromosome.}
}