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Gene Review

hep  -  hemipterous

Drosophila melanogaster

Synonyms: CG2190, CG4353, D-MKK7, DHEP/MKK7, DJNKK, ...
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High impact information on hep

  • Further, hep is required for expression in the dorsal epithelium edges of another dorsal closure gene, puckered [1].
  • hemipterous encodes a novel Drosophila MAP kinase kinase, required for epithelial cell sheet movement [1].
  • Here, we show that the small GTPases Dcdc42, Drac1, and the Hep JNKK control dpp expression in this migratory process [2].
  • Dorsal closure can also be blocked by dominant negative Drosophila cdc42, which has been shown to act upstream of JNKK in vertebrates [3].
  • DJNK is phosphorylated and activated by the Drosophila MAP kinase kinase HEP [4].

Biological context of hep


Anatomical context of hep

  • We show that hep controls the expression of puckered (puc) in the follicular epithelium in a cell-autonomous manner [8].
  • By contrast, constitutively active hopscotch and hemipterous give massive activation of lamellocyte formation with little or no increase in total hemocyte numbers [9].

Regulatory relationships of hep

  • In this report, we demonstrate that Drosophila TAK1 functions as both the Drosophila IkappaB kinase-activating kinase and the JNK kinase-activating kinase [10].

Other interactions of hep

  • The study of mammalian signaling pathways to JNK is complicated by the significant degree of redundancy among upstream JNK regulators, especially at the level of JNK kinase kinases (JNKKK) [11].
  • Furthermore, we show that the small GTPase Dcdc42, but not Drac1, is an activator of puc expression in a hep-dependent manner in imaginal discs [12].
  • Consistent with these observations, expression of LF in Drosophila inhibited the Hep/c-Jun N-terminal kinase pathway during embryonic dorsal closure and the related process of adult thoracic closure [13].
  • Further, dMLK directly phosphorylates Hep, dMKK4 and also their mammalian counterparts, MKK7 and SEK1, in an in vitro kinase assay [14].


  1. hemipterous encodes a novel Drosophila MAP kinase kinase, required for epithelial cell sheet movement. Glise, B., Bourbon, H., Noselli, S. Cell (1995) [Pubmed]
  2. Coupling of Jun amino-terminal kinase and Decapentaplegic signaling pathways in Drosophila morphogenesis. Glise, B., Noselli, S. Genes Dev. (1997) [Pubmed]
  3. The Drosophila Jun-N-terminal kinase is required for cell morphogenesis but not for DJun-dependent cell fate specification in the eye. Riesgo-Escovar, J.R., Jenni, M., Fritz, A., Hafen, E. Genes Dev. (1996) [Pubmed]
  4. A JNK signal transduction pathway that mediates morphogenesis and an immune response in Drosophila. Sluss, H.K., Han, Z., Barrett, T., Davis, R.J., Ip, Y.T. Genes Dev. (1996) [Pubmed]
  5. A novel SAPK/JNK kinase, MKK7, stimulated by TNFalpha and cellular stresses. Moriguchi, T., Toyoshima, F., Masuyama, N., Hanafusa, H., Gotoh, Y., Nishida, E. EMBO J. (1997) [Pubmed]
  6. Defective dorsal closure and loss of epidermal decapentaplegic expression in Drosophila fos mutants. Zeitlinger, J., Kockel, L., Peverali, F.A., Jackson, D.B., Mlodzik, M., Bohmann, D. EMBO J. (1997) [Pubmed]
  7. Dissecting JNK signaling, one KKKinase at a time. Stronach, B. Dev. Dyn. (2005) [Pubmed]
  8. The Drosophila JNK pathway controls the morphogenesis of the egg dorsal appendages and micropyle. Suzanne, M., Perrimon, N., Noselli, S. Dev. Biol. (2001) [Pubmed]
  9. A directed screen for genes involved in Drosophila blood cell activation. Zettervall, C.J., Anderl, I., Williams, M.J., Palmer, R., Kurucz, E., Ando, I., Hultmark, D. Proc. Natl. Acad. Sci. U.S.A. (2004) [Pubmed]
  10. Immune activation of NF-kappaB and JNK requires Drosophila TAK1. Silverman, N., Zhou, R., Erlich, R.L., Hunter, M., Bernstein, E., Schneider, D., Maniatis, T. J. Biol. Chem. (2003) [Pubmed]
  11. D-MEKK1, the Drosophila orthologue of mammalian MEKK4/MTK1, and Hemipterous/D-MKK7 mediate the activation of D-JNK by cadmium and arsenite in Schneider cells. Ryabinina, O.P., Subbian, E., Iordanov, M.S. BMC Cell Biol. (2006) [Pubmed]
  12. The Drosophila JNK pathway controls the morphogenesis of imaginal discs during metamorphosis. Agnès, F., Suzanne, M., Noselli, S. Development (1999) [Pubmed]
  13. Anthrax lethal factor and edema factor act on conserved targets in Drosophila. Guichard, A., Park, J.M., Cruz-Moreno, B., Karin, M., Bier, E. Proc. Natl. Acad. Sci. U.S.A. (2006) [Pubmed]
  14. Drosophila mixed lineage kinase/slipper, a missing biochemical link in Drosophila JNK signaling. Sathyanarayana, P., Barthwal, M.K., Lane, M.E., Acevedo, S.F., Skoulakis, E.M., Bergmann, A., Rana, A. Biochim. Biophys. Acta (2003) [Pubmed]
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