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Gene Review

os  -  outstretched

Drosophila melanogaster

Synonyms: CG5993, Dmel\CG5993, OS, Os, UPD, ...
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Disease relevance of os

  • Ventral unpaired median somata provide immunoreactive processes within the subesophageal ganglion and tritocerebrum [1].

High impact information on os

  • We have used this technique to test the proposal of Jack and Judd that the zeste locus of Drosophila melanogaster cannot repress the activity of unpaired alleles at the white locus [2].
  • We show that a localized source of the pathway ligand, Unpaired, is present at the midline of the developing eye, which is capable of activating the JAK/STAT pathway over long distances [3].
  • Polarity determination in the Drosophila eye: a novel role for unpaired and JAK/STAT signaling [3].
  • When the z1 protein is over-produced in vivo, it reduces the expression of an unpaired copy of white, indicating that the normal requirement for chromosome pairing is simply a device to increase the size of the aggregate bound to the white regulatory region [4].
  • These studies suggest that the topo III enzyme behaves as a structure-specific endonuclease in vivo, providing a reversible DNA cleavage activity that is specific for unpaired regions in the DNA [5].

Biological context of os

  • These phenotypes are enhanced by reduction of unpaired activity, suggesting that Unpaired is a necessary ligand for the JAK pathway in oogenesis [6].
  • Notch activation at the dorsal-ventral boundary of the Drosophila eye-head primordium leads to the expression of the secreted protein Unpaired, a ligand of the JAK-STAT pathway that induces cell proliferation in the undifferentiated tissue [7].
  • Additionally, the unpaired kinetochores accumulated attachments to both poles (merotelic attachments), congressed (again) to a pseudometaphase plate, and reacquired associations with checkpoint proteins more characteristic of prometaphase kinetochores [8].
  • The sequence between the two HJs is identical on both strands, allowing the HJs to migrate without the generation of unpaired regions of DNA, whereas the distance between the HJs is on the order of gene conversion tracts thus far measured in Drosophila and mouse model systems [9].
  • Eukaryotic 3'-tRNase can endonucleolytically remove a 3' end trailer by cleaving on the 3' side of the discriminator base (the unpaired nucleotide 3' of the last base pair of the acceptor stem) [10].

Anatomical context of os

  • Here we demonstrate that the graded activity of the Janus kinase (JAK) pathway, stimulated by the Unpaired ligand, patterns the anterior-posterior axis of the follicular epithelium [11].
  • Molecular control of spermatogonial stem cell self-renewal by glycoprotein unpaired, a cytokine homolog, in Drosophila melanogaster [12].
  • In a second part we describe in detail modulation of ionic currents in three particularly well investigated preparations, i.e. Drosophila photoreceptor, cockroach DUM (dorsal unpaired median) neuron and locust jumping muscle [13].
  • The Drosophila eye field that gives rise to the visual system and dorsal head epidermis forms an unpaired anlage located in the dorsal head ectoderm [14].

Associations of os with chemical compounds

  • It does not contain an unpaired 5' terminal G residue, as reported for Drosophila and sheep histidine tRNAs [15].
  • Comparison between the current d(GGGA)(2) structure and the published crystal d(GAAA)(2) structure implies that the alignment of the unpaired purine bases plays an important role in determining the minor groove width of the purine-rich d(GPuPuA)(2) motif [16].
  • Nonassociative control procedures (CS Alone, US Alone and Explicitly Unpaired) produced slight decreases in avoidance responses, but these affected both odors equally and did not alter our associative learning index (A) [17].
  • Interestingly, the alignment revealed that an unpaired adenine, previously thought to be invariant, is replaced by a guanine in four SECIS elements [18].

Other interactions of os

  • The difference in effect between localized expression of ligand (Unpaired) and dominant active JAK (Hopscotch) demonstrates that the ligand plays a cell non-autonomous role in hindgut cell rearrangement [19].
  • GOF activity requires the X-linked unpaired (upd) gene, which encodes a ligand for the Drosophila JAK/STAT signaling pathway. upd also functions as a numerator element in regulating wild-type Sxl-Pe reporters [20].
  • Reduction of DBHD gene activity suppresses the GSC overproliferation phenotype associated with overexpression of either unpaired (upd) or decapentaplegic (dpp) [21].
  • Depletion of Pc gene products by Pc3 mutation strongly enhances the induction phenomenon, as shown by (1) the increase of the number of wing disc cells in which induction of the homologous allele is detectable, and (2) the induction of not only a paired normal allele but also an unpaired one [22].
  • We also observed that en-/+ heterozygotes could induce a ci phenotype in situations where the ci+ locus is either unpaired or hemizygous [23].

Analytical, diagnostic and therapeutic context of os

  • Since an isolated alpha chain is unstable unless it is associated with a beta chain, this design permits rapid separation of alpha,beta-heterodimer from unpaired beta chain in a single step of Ni-NTA Agarose chromatography yielding 90% pure alpha,beta-TCR [24].
  • In the control group, olfactory and gustatory stimulations were unpaired to prevent a learning process from developing [25].


  1. Comparison of octopamine-like immunoreactivity in the brains of the fruit fly and blow fly. Sinakevitch, I., Strausfeld, N.J. J. Comp. Neurol. (2006) [Pubmed]
  2. Chimaeras of Drosophila melanogaster obtained by injection of haploid nuclei. Santamaría, P., Gans, M. Nature (1980) [Pubmed]
  3. Polarity determination in the Drosophila eye: a novel role for unpaired and JAK/STAT signaling. Zeidler, M.P., Perrimon, N., Strutt, D.I. Genes Dev. (1999) [Pubmed]
  4. Self-association of the Drosophila zeste protein is responsible for transvection effects. Bickel, S., Pirrotta, V. EMBO J. (1990) [Pubmed]
  5. Preferential cleavage of plasmid-based R-loops and D-loops by Drosophila topoisomerase IIIbeta. Wilson-Sali, T., Hsieh, T.S. Proc. Natl. Acad. Sci. U.S.A. (2002) [Pubmed]
  6. JAK signaling is somatically required for follicle cell differentiation in Drosophila. McGregor, J.R., Xi, R., Harrison, D.A. Development (2002) [Pubmed]
  7. Growth control in the proliferative region of the Drosophila eye-head primordium: The elbow-noc gene complex. Luque, C.M., Milán, M. Dev. Biol. (2007) [Pubmed]
  8. Cyclin B destruction triggers changes in kinetochore behavior essential for successful anaphase. Parry, D.H., Hickson, G.R., O'Farrell, P.H. Curr. Biol. (2003) [Pubmed]
  9. A novel, topologically constrained DNA molecule containing a double Holliday junction: design, synthesis, and initial biochemical characterization. Plank, J.L., Hsieh, T.S. J. Biol. Chem. (2006) [Pubmed]
  10. The 3' end CCA of mature tRNA is an antideterminant for eukaryotic 3'-tRNase. Mohan, A., Whyte, S., Wang, X., Nashimoto, M., Levinger, L. RNA (1999) [Pubmed]
  11. A gradient of JAK pathway activity patterns the anterior-posterior axis of the follicular epithelium. Xi, R., McGregor, J.R., Harrison, D.A. Dev. Cell (2003) [Pubmed]
  12. Molecular control of spermatogonial stem cell self-renewal by glycoprotein unpaired, a cytokine homolog, in Drosophila melanogaster. Tulina, N.M. Dokl. Biol. Sci. (2003) [Pubmed]
  13. Non-synaptic ion channels in insects--basic properties of currents and their modulation in neurons and skeletal muscles. Wicher, D., Walther, C., Wicher, C. Prog. Neurobiol. (2001) [Pubmed]
  14. Antagonistic relationship between Dpp and EGFR signaling in Drosophila head patterning. Chang, T., Shy, D., Hartenstein, V. Dev. Biol. (2003) [Pubmed]
  15. Isolation and nucleotide sequence of a mouse histidine tRNA gene. Han, J.H., Harding, J.D. Nucleic Acids Res. (1982) [Pubmed]
  16. Quadruple intercalated G-6 stack: a possible motif in the fold-back structure of the Drosophila centromeric dodeca-satellite? Chou, S.H., Chin, K.H. J. Mol. Biol. (2001) [Pubmed]
  17. Classical conditioning and retention in normal and mutant Drosophila melanogaster. Tully, T., Quinn, W.G. J. Comp. Physiol. A (1985) [Pubmed]
  18. Structural analysis of new local features in SECIS RNA hairpins. Fagegaltier, D., Lescure, A., Walczak, R., Carbon, P., Krol, A. Nucleic Acids Res. (2000) [Pubmed]
  19. Localized JAK/STAT signaling is required for oriented cell rearrangement in a tubular epithelium. Johansen, K.A., Iwaki, D.D., Lengyel, J.A. Development (2003) [Pubmed]
  20. The JAK/STAT signaling pathway is required for the initial choice of sexual identity in Drosophila melanogaster. Jinks, T.M., Polydorides, A.D., Calhoun, G., Schedl, P. Mol. Cell (2000) [Pubmed]
  21. The Drosophila homolog of the human tumor suppressor gene BHD interacts with the JAK-STAT and Dpp signaling pathways in regulating male germline stem cell maintenance. Singh, S.R., Zhen, W., Zheng, Z., Wang, H., Oh, S.W., Liu, W., Zbar, B., Schmidt, L.S., Hou, S.X. Oncogene (2006) [Pubmed]
  22. Transvection in the Drosophila Ultrabithorax gene: a Cbx1 mutant allele induces ectopic expression of a normal allele in trans. Castelli-Gair, J.E., Micol, J.L., García-Bellido, A. Genetics (1990) [Pubmed]
  23. Engrailed gene dosage determines whether certain recessive cubitus interruptus alleles exhibit dominance of the adult wing phenotype in Drosophila. Locke, J., Hanna, S. Dev. Genet. (1996) [Pubmed]
  24. Soluble HIV-specific T cell receptor: expression, purification and analysis of the specificity. Anikeeva, N., Lebedeva, T., Sumaroka, M., Kalams, S.A., Sykulev, Y. J. Immunol. Methods (2003) [Pubmed]
  25. A new attempt to assess the effect of learning processes on the cholinergic system: studies on fruitflies and honeybees. Fresquet, N., Fournier, D., Gauthier, M. Comp. Biochem. Physiol. B, Biochem. Mol. Biol. (1998) [Pubmed]
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