Gene Symbol: snk
Description: snake
Alias: CG7996, Dmel\CG7996, SNK, SP26, Snk, c-SP26, l(3)87Dg, me(3)4, snake, CG7996-PA, CG7996-PB, CG7996-PC, snk-PA, snk-PB, snk-PC
Species: fruit fly

Top Publications

  1. Trivedi B. Biography of Kathryn V. Anderson. Proc Natl Acad Sci U S A. 2005;102:5910-2 pubmed
  2. Stein D, Roth S, Vogelsang E, Nusslein Volhard C. The polarity of the dorsoventral axis in the Drosophila embryo is defined by an extracellular signal. Cell. 1991;65:725-35 pubmed
    ..Presumably, local activation of the Toll protein by the ligand initiates the formation of the nuclear concentration gradient of the dorsal protein, thereby determining dorsoventral pattern. ..
  3. Coates D, Siviter R, Isaac R. Exploring the Caenorhabditis elegans and Drosophila melanogaster genomes to understand neuropeptide and peptidase function. Biochem Soc Trans. 2000;28:464-9 pubmed
  4. Ray R, Arora K, Nusslein Volhard C, Gelbart W. The control of cell fate along the dorsal-ventral axis of the Drosophila embryo. Development. 1991;113:35-54 pubmed
    ..Two of the genes, dpp and twi, are unaffected by mutations in any of the tested zygotic dorsal-ventral genes, suggesting that dpp and twi are the primary patterning genes for dorsal ectoderm and mesoderm, respectively...
  5. Dutton F, Chovnick A. The l(3)S12 locus of Drosophila melanogaster: heterochromatic position effects and stage-specific misexpression of the gene in P element transposons. Genetics. 1991;128:103-18 pubmed
    ..Heterochromatic position effect on this locus has been demonstrated. Available evidence suggests that the l(3)S12 gene may be involved in protein synthesis, perhaps encoding a ribosomal protein. ..
  6. Stein D. Pattern formation: The link between ovary and embryo. Curr Biol. 1995;5:1360-3 pubmed
  7. Holley S, Neul J, Attisano L, Wrana J, Sasai Y, O Connor M, et al. The Xenopus dorsalizing factor noggin ventralizes Drosophila embryos by preventing DPP from activating its receptor. Cell. 1996;86:607-17 pubmed
    ..These results suggest a mechanistic basis for the action of Spemann's organizer during Xenopus development and provide further support for the conservation of dorsal-ventral patterning mechanisms between arthropods and chordates. ..
  8. Curtis D, Bender W. Gene conversion in Drosophila and the effects of the meiotic mutants mei-9 and mei-218. Genetics. 1991;127:739-46 pubmed
    ..Some unusual conversion events were found in the mei-218 and mei-9 samples, although most events were indistinguishable from wild-type tracts in their length and continuity. ..
  9. Rose T, LeMosy E, Cantwell A, Banerjee Roy D, Skeath J, Di Cera E. Three-dimensional models of proteases involved in patterning of the Drosophila Embryo. Crucial role of predicted cation binding sites. J Biol Chem. 2003;278:11320-30 pubmed
    Three-dimensional models of the catalytic domains of Nudel (Ndl), Gastrulation Defective (Gd), Snake (Snk), and Easter (Ea), and their complexes with substrate suggest a possible organization of the enzyme cascade controlling the ..

More Information


  1. Bergmann A, Stein D, Geisler R, Hagenmaier S, Schmid B, Fernandez N, et al. A gradient of cytoplasmic Cactus degradation establishes the nuclear localization gradient of the dorsal morphogen in Drosophila. Mech Dev. 1996;60:109-23 pubmed
    ..Genetic evidence indicates that degradation of Cactus is required, but not sufficient to translocates Dorsal completely into the nucleus. ..
  2. Pawson T, Bernstein A. Receptor tyrosine kinases: genetic evidence for their role in Drosophila and mouse development. Trends Genet. 1990;6:350-6 pubmed
  3. Anderson K, Schneider D, Morisato D, Jin Y, Ferguson E. Extracellular morphogens in Drosophila embryonic dorsal-ventral patterning. Cold Spring Harb Symp Quant Biol. 1992;57:409-17 pubmed
  4. Qiu P, Pan P, Govind S. A role for the Drosophila Toll/Cactus pathway in larval hematopoiesis. Development. 1998;125:1909-20 pubmed
    ..These findings are discussed in light of similar hematopoietic functions of Rel/I(kappa)B-family proteins in mice. ..
  5. Stein D, Nusslein Volhard C. Multiple extracellular activities in Drosophila egg perivitelline fluid are required for establishment of embryonic dorsal-ventral polarity. Cell. 1992;68:429-40 pubmed
    ..separate activities present in the perivitelline fluid that can restore dorsal-ventral polarity to mutant easter, snake, and spätzle embryos, respectively...
  6. Hudson J, Podos S, Keith K, Simpson S, Ferguson E. The Drosophila Medea gene is required downstream of dpp and encodes a functional homolog of human Smad4. Development. 1998;125:1407-20 pubmed
    ..Moreover, injection of human SMAD4 mRNA into embryos lacking all Medea activity conferred phenotypic rescue of the dorsal-ventral pattern, demonstrating conservation of function between the two gene products. ..
  7. Jackson P, Hoffmann F. Embryonic expression patterns of the Drosophila decapentaplegic gene: separate regulatory elements control blastoderm expression and lateral ectodermal expression. Dev Dyn. 1994;199:28-44 pubmed
  8. Smith C, DeLotto R. Ventralizing signal determined by protease activation in Drosophila embryogenesis. Nature. 1994;368:548-51 pubmed
    ..The snake and easter genes are required for the production of the ligand and they encode the proenzyme form of secreted ..
  9. Chasan R, Jin Y, Anderson K. Activation of the easter zymogen is regulated by five other genes to define dorsal-ventral polarity in the Drosophila embryo. Development. 1992;115:607-16 pubmed
    ..We propose that dorsal-ventral asymmetry is initiated by a ventrally-localized molecule in the vitelline membrane that nucleates an easter zymogen activation complex, leading to the production of ventrally active easter enzyme. ..
  10. Beutler B, Poltorak A. Toll we meet again. Nat Immunol. 2001;2:9-10 pubmed
  11. Morisato D, Anderson K. The spätzle gene encodes a component of the extracellular signaling pathway establishing the dorsal-ventral pattern of the Drosophila embryo. Cell. 1994;76:677-88 pubmed
    ..We propose that proteolytic processing of the spätzle protein is confined to the ventral side of the embryo and that the localization of processed spätzle determines where the receptor, Toll, is active. ..
  12. Tian S, LeMosy E. Mutagenesis of the cysteine-rich clip domain in the Drosophila patterning protease, Snake. Arch Biochem Biophys. 2008;475:169-74 pubmed publisher
    ..b>Snake functions in a cascade that patterns the Drosophila embryo, and provides an accessible model for exploring the ..
  13. Roth S, Hiromi Y, Godt D, Nusslein Volhard C. cactus, a maternal gene required for proper formation of the dorsoventral morphogen gradient in Drosophila embryos. Development. 1991;112:371-88 pubmed
    ..We discuss models of how the action of the dorsal group genes might counteract the cactus function ventrally. ..
  14. Carroll S, Winslow G, Twombly V, Scott M. Genes that control dorsoventral polarity affect gene expression along the anteroposterior axis of the Drosophila embryo. Development. 1987;99:327-32 pubmed
  15. LeMosy E, Tan Y, Hashimoto C. Activation of a protease cascade involved in patterning the Drosophila embryo. Proc Natl Acad Sci U S A. 2001;98:5055-60 pubmed
    ..Production of this signal requires the serine proteases Gastrulation Defective (GD), Snake, and Easter, which genetic studies suggest act sequentially in a cascade that is activated locally in response to ..
  16. Dingwall C. Transport across the nuclear envelope: enigmas and explanations. Bioessays. 1991;13:213-8 pubmed
  17. Ponomareff G, Giordano H, DeLotto Y, DeLotto R. Interallelic complementation at the Drosophila melanogaster gastrulation defective locus defines discrete functional domains of the protein. Genetics. 2001;159:635-45 pubmed
    ..An allele-specific interaction between an autoactivating form of Snake (Snk) and a gd allele altered in the protease domain suggests that Gd directly activates Snk in a protease ..
  18. Sorrentino R, Melk J, Govind S. Genetic analysis of contributions of dorsal group and JAK-Stat92E pathway genes to larval hemocyte concentration and the egg encapsulation response in Drosophila. Genetics. 2004;166:1343-56 pubmed
  19. Roth S. Drosophila development: the secrets of delayed induction. Curr Biol. 1998;8:R906-10 pubmed
    ..This asymmetry, in turn, depends on the expression specifically in ventral follicle cells of pipe, the product of which seems likely to be a glycosaminoglycan-modifying enzyme. ..
  20. Stein D, Charatsi I, Cho Y, Zhang Z, Nguyen J, DeLotto R, et al. Localization and activation of the Drosophila protease easter require the ER-resident saposin-like protein seele. Curr Biol. 2010;20:1953-8 pubmed publisher
    ..Gastrulation Defective processes Snake, which then cleaves Easter, which then processes Spätzle into the activating ligand for the Toll receptor...
  21. Hecht P, Anderson K. Extracellular proteases and embryonic pattern formation. Trends Cell Biol. 1992;2:197-202 pubmed
  22. Dissing M, Giordano H, DeLotto R. Autoproteolysis and feedback in a protease cascade directing Drosophila dorsal-ventral cell fate. EMBO J. 2001;20:2387-93 pubmed
    Three serine protease zymogens, Gastrulation defective (GD), Snake (Snk) and Easter (Ea), and a nerve growth factor-like growth factor ligand precursor, Spaetzle, are required for specification of dorsal- ventral cell fate during ..
  23. Grosshans J, Schnorrer F, Nusslein Volhard C. Oligomerisation of Tube and Pelle leads to nuclear localisation of dorsal. Mech Dev. 1999;81:127-38 pubmed
  24. LeMosy E. Spatially dependent activation of the patterning protease, Easter. FEBS Lett. 2006;580:2269-72 pubmed
    ..Here, biochemical studies demonstrate that this spatial control occurs at or above the level of Easter zymogen activation, rather than through direct promotion of Easter's catalytic activity against the signaling ligand. ..
  25. Belvin M, Anderson K. A conserved signaling pathway: the Drosophila toll-dorsal pathway. Annu Rev Cell Dev Biol. 1996;12:393-416 pubmed
  26. Letsou A, Alexander S, Orth K, Wasserman S. Genetic and molecular characterization of tube, a Drosophila gene maternally required for embryonic dorsoventral polarity. Proc Natl Acad Sci U S A. 1991;88:810-4 pubmed
    ..These results suggest that tube represents a class of protein active in signal transduction at two stages of development. ..
  27. DeLotto R. Gastrulation defective, a complement factor C2/B-like protease, interprets a ventral prepattern in Drosophila. EMBO Rep. 2001;2:721-6 pubmed
    ..A homology to the mammalian complement factors C2 and B is also described. ..
  28. Gay N, Keith F. Formation of a gradient of the Drosophila dorsal morphogen by differential nuclear localisation. Bioessays. 1990;12:181-2 pubmed
  29. Roth S. Axis determination. Proteolytic generation of a morphogen. Curr Biol. 1994;4:755-7 pubmed
    ..Ventral activation of a transmembrane receptor, Toll, is a crucial step in dorsoventral axis establishment in Drosophila embryos. The ventral ligand for Toll seems to be a proteolytic fragment of the Spätzle protein. ..
  30. Rushlow C, Bender W, Chovnick A. Studies on the mechanism of heterochromatic position effect at the rosy locus of Drosophila melanogaster. Genetics. 1984;108:603-15 pubmed
    ..Histone region deletion heterozygosity and butyrate-feeding experiments failed to exhibit specific suppression of position effect at the rosy locus. ..
  31. Rushlow C, Levine M. Role of the zerknüllt gene in dorsal-ventral pattern formation in Drosophila. Adv Genet. 1990;27:277-307 pubmed
  32. Stein D, Stevens L. Establishment of dorsal-ventral and terminal pattern in the Drosophila embryo. Curr Opin Genet Dev. 1991;1:247-54 pubmed
  33. Rushlow C. Dorsoventral patterning: a serpin pinned down at last. Curr Biol. 2004;14:R16-8 pubmed
    ..At long last, the prediction that a protease inhibitor is involved in this pathway has been shown to be true, with the identification of a serpin that plays a key part in Drosophila embryonic patterning. ..
  34. Cho Y, Stevens L, Stein D. Pipe-dependent ventral processing of Easter by Snake is the defining step in Drosophila embryo DV axis formation. Curr Biol. 2010;20:1133-7 pubmed publisher
    ..Gastrulation Defective cleaves and activates Snake, which processes and activates Easter, which cleaves Spätzle to form the activating ligand for the Toll receptor...
  35. Clark S, Chovnick A. Studies of normal and position-affected expression of rosy region genes in Drosophila melanogaster. Genetics. 1986;114:819-40 pubmed
    ..rearrangements are examined with respect to position effects on expression of the rosy region genes l(3)12, rosy, snake and piccolo, as well as suppressor effects...
  36. Neul J, Ferguson E. Spatially restricted activation of the SAX receptor by SCW modulates DPP/TKV signaling in Drosophila dorsal-ventral patterning. Cell. 1998;95:483-94 pubmed
    ..We propose that spatial regulation of the SAX pathway modulates TKV signaling to create positional information over the embryonic ectoderm. ..
  37. Lemaitre B, Nicolas E, Michaut L, Reichhart J, Hoffmann J. The dorsoventral regulatory gene cassette spätzle/Toll/cactus controls the potent antifungal response in Drosophila adults. Cell. 1996;86:973-83 pubmed
    ..Antibacterial genes are induced either by a distinct pathway involving the immune deficiency gene (imd) or by combined activation of both imd and dorsoventral pathways. ..
  38. Leclerc V, Pelte N, El Chamy L, Martinelli C, Ligoxygakis P, Hoffmann J, et al. Prophenoloxidase activation is not required for survival to microbial infections in Drosophila. EMBO Rep. 2006;7:231-5 pubmed
    ..This raises the question with regard to the precise function of phenoloxidase activation in defence, if any. ..
  39. Ligoxygakis P, Roth S, Reichhart J. A serpin regulates dorsal-ventral axis formation in the Drosophila embryo. Curr Biol. 2003;13:2097-102 pubmed
    ..Since this serpin has been recently shown to restrain an immune reaction in the blood of Drosophila, it demonstrates that proteolysis can be regulated by the same serpin in different biological contexts. ..
  40. Tang H, Kambris Z, Lemaitre B, Hashimoto C. Two proteases defining a melanization cascade in the immune system of Drosophila. J Biol Chem. 2006;281:28097-104 pubmed
    ..We have also shown that the melanization reaction activated by MP1 and MP2 plays an important role in augmenting the effectiveness of other immune reactions, thereby promoting resistance of Drosophila to microbial infection. ..
  41. Cho Y, Stevens L, Sieverman K, Nguyen J, Stein D. A ventrally localized protease in the Drosophila egg controls embryo dorsoventral polarity. Curr Biol. 2012;22:1013-8 pubmed publisher
    ..Gastrulation Defective (GD) cleaves and activates Snake (Snk). Activated Snk cleaves and activates Easter (Ea), exclusively on the ventral side of the embryo...
  42. Lemaitre B, Nicolas E, Michaut L, Reichhart J, Hoffmann J. Pillars article: the dorsoventral regulatory gene cassette spätzle/Toll/cactus controls the potent antifungal response in Drosophila adults. Cell. 1996. 86: 973-983. J Immunol. 2012;188:5210-20 pubmed
    ..Antibacterial genes are induced either by a distinct pathway involving the immune deficiency gene (imd) or by combined activation of both imd and dorsoventral pathways. ..
  43. Stein D, Cho Y, Zhang Z, Stevens L. No requirement for localized Nudel protein expression in Drosophila embryonic axis determination. Fly (Austin). 2008;2:220-8 pubmed
    ..Gastrulation Defective, Snake, and Easter comprise a serine protease cascade that operates in the perivitelline space to generate active ligand ..
  44. Steen P, Tian S, Tully S, Cravatt B, LeMosy E. Activation of Snake in a serine protease cascade that defines the dorsoventral axis is atypical and pipe-independent in Drosophila embryos. FEBS Lett. 2010;584:3557-60 pubmed publisher
    ..spatial regulation of an extracellular serine protease cascade composed of Nudel, Gastrulation Defective (GD), Snake, and Easter...
  45. Han J, Lee S, Tan Y, LeMosy E, Hashimoto C. Gastrulation defective is a serine protease involved in activating the receptor toll to polarize the Drosophila embryo. Proc Natl Acad Sci U S A. 2000;97:9093-7 pubmed
    ..Our data support a model in which the GD protease catalyzes the ventral activation of the proteolytic cascade that produces the Toll ligand. ..
  46. Grosshans J, Bergmann A, Haffter P, Nusslein Volhard C. Activation of the kinase Pelle by Tube in the dorsoventral signal transduction pathway of Drosophila embryo. Nature. 1994;372:563-6 pubmed
    ..In addition, Pelle and Tube interact directly with one another. We propose that Tube is a direct activator of the protein kinase Pelle. ..
  47. Wu L, Anderson K. Related signaling networks in Drosophila that control dorsoventral patterning in the embryo and the immune response. Cold Spring Harb Symp Quant Biol. 1997;62:97-103 pubmed
  48. Decotto E, Ferguson E. A positive role for Short gastrulation in modulating BMP signaling during dorsoventral patterning in the Drosophila embryo. Development. 2001;128:3831-41 pubmed
  49. DeLotto R, Spierer P. A gene required for the specification of dorsal-ventral pattern in Drosophila appears to encode a serine protease. Nature. 1986;323:688-92 pubmed
    The maternal effect gene snake is required for the establishment of the dorsal-ventral axis during the embryonic development of Drosophila...
  50. Aquadro C, Lado K, Noon W. The rosy region of Drosophila melanogaster and Drosophila simulans. I. Contrasting levels of naturally occurring DNA restriction map variation and divergence. Genetics. 1988;119:875-88 pubmed
    A 40-kb region around the rosy and snake loci was analyzed for restriction map variation among 60 lines of Drosophila melanogaster and 30 lines of Drosophila simulans collected together at a single locality in Raleigh, North Carolina...
  51. Anderson K, Nusslein Volhard C. Information for the dorsal--ventral pattern of the Drosophila embryo is stored as maternal mRNA. Nature. 1984;311:223-7 pubmed
    ..Injection of RNA isolated from wild-type embryos into mutants at six loci partially restores dorsal-ventral polarity. For the mutant snake, injection of poly(A)+ RNA restores a complete dorsal-ventral pattern.
  52. LeMosy E, Kemler D, Hashimoto C. Role of Nudel protease activation in triggering dorsoventral polarization of the Drosophila embryo. Development. 1998;125:4045-53 pubmed
  53. Smith C, DeLotto R. A common domain within the proenzyme regions of the Drosophila snake and easter proteins and Tachypleus proclotting enzyme defines a new subfamily of serine proteases. Protein Sci. 1992;1:1225-6 pubmed
  54. Sukumari Ramesh S, LeMosy E. Gastrulation defective protease interacts with anionic components of the Drosophila ovary extracellular matrix. Protein Pept Lett. 2009;16:437-43 pubmed
    The Drosophila proteases Gastrulation Defective and Snake function in embryonic polarity establishment and bind heparin, a surrogate for anionic species present in the extracellular matrix...
  55. Lemaitre B, Meister M, Govind S, Georgel P, Steward R, Reichhart J, et al. Functional analysis and regulation of nuclear import of dorsal during the immune response in Drosophila. EMBO J. 1995;14:536-45 pubmed
    ..These data underline the complexity of the Drosophila immune response. Finally, we observed that like other rel proteins, dl can control the level of its own transcription. ..
  56. Levashina E, Ohresser S, Lemaitre B, Imler J. Two distinct pathways can control expression of the gene encoding the Drosophila antimicrobial peptide metchnikowin. J Mol Biol. 1998;278:515-27 pubmed
    ..5 kb of metchnikowin gene upstream sequences indicates that this fragment is able to confer full immune inducibility and tissue specificity of expression on the transgene. ..
  57. LeMosy E, Hong C, Hashimoto C. Signal transduction by a protease cascade. Trends Cell Biol. 1999;9:102-7 pubmed
    ..Studies of the proteases and somatically expressed proteins involved in this signalling process suggest a working model for how the protease cascade is locally activated hours after the ovarian somatic cells have degenerated. ..
  58. Levashina E, Langley E, Green C, Gubb D, Ashburner M, Hoffmann J, et al. Constitutive activation of toll-mediated antifungal defense in serpin-deficient Drosophila. Science. 1999;285:1917-9 pubmed
    ..Hence, Spn43Ac negatively regulates the Toll signaling pathway, and Toll does not function as a pattern recognition receptor in the Drosophila host defense...
  59. Baker D, Meadows L, Wang J, Dow J, Russell S. Variable sexually dimorphic gene expression in laboratory strains of Drosophila melanogaster. BMC Genomics. 2007;8:454 pubmed
    ..Much of this variation reflects sex-specific challenges associated with divergent physiological trade-offs, morphology and regulatory pathways operating within males and females. ..
  60. Anderson K, Jurgens G, Nusslein Volhard C. Establishment of dorsal-ventral polarity in the Drosophila embryo: genetic studies on the role of the Toll gene product. Cell. 1985;42:779-89 pubmed
    ..Complementation studies show specific trans interactions between copies of the Toll product. Double mutant phenotypes suggest that the products of several other dorsal-group genes regulate the activity of Toll. ..
  61. Charatsi I, Luschnig S, Bartoszewski S, NUSSLEIN VOLHARD C, Moussian B. Krapfen/dMyd88 is required for the establishment of dorsoventral pattern in the Drosophila embryo. Mech Dev. 2003;120:219-26 pubmed
    ..Tube in turn interacts with its downstream effector Pelle through death domain association. We therefore suggest that upon Toll activation, Kra associates with Pelle and Tube, in an heterotrimeric complex. ..