Gene Symbol: Sos
Description: Son of sevenless
Alias: 34Ea, BG:DS00941.4, CG7793, Dmel\CG7793, E(sev)2A, E(var)189, EC2-5, EK2-8, EY2-3, SOS, Sos1, Su(tor)2-2, br24, br25, dSos, dme-SOS, l(2)34Ea, l(2)Sos, l(2)br24, l(2)br25, l(2)k05224, l(2)k06321, sos, Son of sevenless, CG7793-PA, Son-of-sevenless, Sos-PA, lethal 24 in the black-reduced region, lethal 25 in the black-reduced region
Species: fruit fly

Top Publications

  1. Yang L, Bashaw G. Son of sevenless directly links the Robo receptor to rac activation to control axon repulsion at the midline. Neuron. 2006;52:595-607 pubmed
    b>Son of sevenless (Sos) is a dual specificity guanine nucleotide exchange factor (GEF) that regulates both Ras and Rho family GTPases and thus is uniquely poised to integrate signals that affect both gene expression and cytoskeletal ..
  2. Rebay I, Chen F, Hsiao F, Kolodziej P, Kuang B, Laverty T, et al. A genetic screen for novel components of the Ras/Mitogen-activated protein kinase signaling pathway that interact with the yan gene of Drosophila identifies split ends, a new RNA recognition motif-containing protein. Genetics. 2000;154:695-712 pubmed
    ..We also isolated mutations in five previously uncharacterized genes, one of which, split ends, we have characterized molecularly and have shown to encode a member of the RRM family of RNA-binding proteins. ..
  3. Kolodkin A, Pickup A, Lin D, Goodman C, Banerjee U. Characterization of Star and its interactions with sevenless and EGF receptor during photoreceptor cell development in Drosophila. Development. 1994;120:1731-45 pubmed
    ..mutations in Star act as suppressors of R7 development in a sensitized genetic background involving the Son of sevenless (Sos) locus, and overexpression of Star enhances R7 development in this genetic background...
  4. Therrien M, Morrison D, Wong A, Rubin G. A genetic screen for modifiers of a kinase suppressor of Ras-dependent rough eye phenotype in Drosophila. Genetics. 2000;156:1231-42 pubmed
    ..One of them corresponds to the kismet locus, which encodes a putative chromatin remodeling factor. The relevance of these loci with respect to the function of KSR and the Ras1 pathway in general is discussed. ..
  5. Perrimon N. Signalling pathways initiated by receptor protein tyrosine kinases in Drosophila. Curr Opin Cell Biol. 1994;6:260-6 pubmed
    ..In addition to molecules previously identified from vertebrate studies, i.e. Grb2, Sos, Ras-Gap, p21ras, Raf, MEK and MAPK, genetic studies have suggested that two novel proteins, the protein tyrosine ..
  6. Diaz Benjumea F, Hafen E. The sevenless signalling cassette mediates Drosophila EGF receptor function during epidermal development. Development. 1994;120:569-78 pubmed
    In Drosophila, Drk, an SH2 adaptor protein, Sos, a putative activator of Ras1, Ras1, raf and rolled/MAP kinase have been shown to be required for signalling from the sevenless and the torso receptor tyrosine kinase...
  7. Pflumm M, Botchan M. Orc mutants arrest in metaphase with abnormally condensed chromosomes. Development. 2001;128:1697-707 pubmed
    ..Thus, cells have a mitotic checkpoint that senses chromosome integrity. These studies also suggest that the density of functional replication origins and completion of S phase are requirements for proper chromosomal condensation. ..
  8. Herbst R, Carroll P, Allard J, Schilling J, Raabe T, Simon M. Daughter of sevenless is a substrate of the phosphotyrosine phosphatase Corkscrew and functions during sevenless signaling. Cell. 1996;85:899-909 pubmed
    ..Analysis of dos mutations indicates that DOS is a positive component of the SEV signaling pathway and suggests that DOS dephosphorylation by CSW may be a key event during signaling by SEV. ..
  9. Murillo Maldonado J, Zeineddine F, Stock R, THACKERAY J, Riesgo Escovar J. Insulin receptor-mediated signaling via phospholipase C-? regulates growth and differentiation in Drosophila. PLoS ONE. 2011;6:e28067 pubmed publisher
    ..Overall, our results provide evidence that PLC-? acts during development at a time when growth ends and differentiation begins, and is important for proper coordination of these two processes. ..

More Information


  1. Egan S, Giddings B, Brooks M, Buday L, Sizeland A, Weinberg R. Association of Sos Ras exchange protein with Grb2 is implicated in tyrosine kinase signal transduction and transformation. Nature. 1993;363:45-51 pubmed
    ..Grb2-Sem-5 binds directly to murine Sos1, a Ras exchange factor, through two SH3 domains...
  2. Perrimon N. The torso receptor protein-tyrosine kinase signaling pathway: an endless story. Cell. 1993;74:219-22 pubmed
  3. Altshuler D, Lillien L. Control of photoreceptor development. Curr Opin Neurobiol. 1992;2:16-22 pubmed
    ..In this review the strategies used to generate rods in the vertebrate retina are compared with those described for photoreceptor development in the Drosophila retina. ..
  4. Rogge R, Cagan R, Majumdar A, Dulaney T, Banerjee U. Neuronal development in the Drosophila retina: the sextra gene defines an inhibitory component in the developmental pathway of R7 photoreceptor cells. Proc Natl Acad Sci U S A. 1992;89:5271-5 pubmed
    ..Additionally, sxt functions in R7 precursors, but the sevenless signal is essential for specification of this fate, since loss of sextra alone is unable to impart a neural fate to this cell. ..
  5. Thomas B, Gunning D, Cho J, Zipursky L. Cell cycle progression in the developing Drosophila eye: roughex encodes a novel protein required for the establishment of G1. Cell. 1994;77:1003-14 pubmed
    ..e., Ras1 and Star). rux encodes a novel protein of 335 amino acids. We propose that rux functions as a negative regulator of G1 progression in the developing eye. ..
  6. Rawlings J, Rennebeck G, Harrison S, Xi R, Harrison D. Two Drosophila suppressors of cytokine signaling (SOCS) differentially regulate JAK and EGFR pathway activities. BMC Cell Biol. 2004;5:38 pubmed
    ..The non-canonical properties of Socs44A may be representative of the class of less characterized vertebrate SOCS with which it shares greatest similarity. ..
  7. Knust E. Drosophila morphogenesis: follow-my-leader in epithelia. Curr Biol. 1996;6:379-81 pubmed
    ..The discovery of a second MAPK kinase in Drosophila provides new insights into mechanisms that control the coordinated movement of epithelial cells. ..
  8. Read R, Bach E, Cagan R. Drosophila C-terminal Src kinase negatively regulates organ growth and cell proliferation through inhibition of the Src, Jun N-terminal kinase, and STAT pathways. Mol Cell Biol. 2004;24:6676-89 pubmed
    ..In particular, blockade of STAT function in dCsk mutants severely reduced Src-dependent overgrowth and activated apoptosis of mutant tissue. Our data provide in vivo evidence that Src activity requires JNK and STAT function. ..
  9. Hayashi T, Kojima T, Saigo K. Specification of primary pigment cell and outer photoreceptor fates by BarH1 homeobox gene in the developing Drosophila eye. Dev Biol. 1998;200:131-45 pubmed
  10. Pawson T. Signal transduction--a conserved pathway from the membrane to the nucleus. Dev Genet. 1993;14:333-8 pubmed
  11. Slováková J, Speicher S, Sanchez Soriano N, Prokop A, Carmena A. The actin-binding protein Canoe/AF-6 forms a complex with Robo and is required for Slit-Robo signaling during axon pathfinding at the CNS midline. J Neurosci. 2012;32:10035-44 pubmed publisher
  12. Freeman M. The spitz gene is required for photoreceptor determination in the Drosophila eye where it interacts with the EGF receptor. Mech Dev. 1994;48:25-33 pubmed
    ..These data suggest that photoreceptors other than R7 use a Ras1 signalling pathway activated by the spitz/Egfr interaction, in a manner analogous to the Ras1 pathway activated by boss/sevenless in photoreceptor R7. ..
  13. Bai J, Chiu W, Wang J, Tzeng T, Perrimon N, Hsu J. The cell adhesion molecule Echinoid defines a new pathway that antagonizes the Drosophila EGF receptor signaling pathway. Development. 2001;128:591-601 pubmed
    ..Together, our results support a model whereby ED defines an independent pathway that antagonizes EGFR signaling by regulating the activity, but not the level, of the TTK88 transcriptional repressor. ..
  14. Fath I, Apiou F, Schweighoffer F, Chevallier Multon M, Ciora T, Dutrillaux B, et al. Identification of two human homologues to Drosophila SOS (son of sevenless) localized on two different chromosomes. Nucleic Acids Res. 1993;21:4398 pubmed
  15. Doyle H, Bishop J. Torso, a receptor tyrosine kinase required for embryonic pattern formation, shares substrates with the sevenless and EGF-R pathways in Drosophila. Genes Dev. 1993;7:633-46 pubmed
    ..Two of these correspond to mutations in ras-1 and Son of sevenless (Sos), which also function in the sevenless and EGF receptor (Der) tyrosine kinase pathways...
  16. Gubb D, Ashburner M, Roote J, Davis T. A novel transvection phenomenon affecting the white gene of Drosophila melanogaster. Genetics. 1990;126:167-76 pubmed
    ..This effect of rearrangements has been used as the basis for a screen, gamma-ray induced aberrations with at least one breakpoint opposite the TE site were recovered by their suppression of the zeste phenotype. ..
  17. Pazman C, Mayes C, Fanto M, Haynes S, Mlodzik M. Rasputin, the Drosophila homologue of the RasGAP SH3 binding protein, functions in ras- and Rho-mediated signaling. Development. 2000;127:1715-25 pubmed
    ..These data suggest that Rin is not a general component of polarity generation, but serves a function specific to Ras and RhoA signaling pathways. ..
  18. Lu T, Doherty J, Freeman M. DRK/DOS/SOS converge with Crk/Mbc/dCed-12 to activate Rac1 during glial engulfment of axonal debris. Proc Natl Acad Sci U S A. 2014;111:12544-9 pubmed publisher
    ..kinase (DRK) and daughter of sevenless (DOS) (mammalian homologs, Grb2 and Gab2, respectively) and the GEF son of sevenless (SOS) (mammalian homolog, mSOS) are required for efficient activation of glia after axotomy and ..
  19. Amoyel M, Anderson J, Suisse A, Glasner J, Bach E. Socs36E Controls Niche Competition by Repressing MAPK Signaling in the Drosophila Testis. PLoS Genet. 2016;12:e1005815 pubmed publisher
    ..Socs36E is node through which these processes are linked, demonstrating that negative feedback inhibition integrates multiple aspects of stem cell behavior. ..
  20. Sawamoto K, Okabe M, Tanimura T, Mikoshiba K, Nishida Y, Okano H. The Drosophila secreted protein Argos regulates signal transduction in the Ras/MAPK pathway. Dev Biol. 1996;178:13-22 pubmed
    ..argos mutation enhanced the overproduction of R7 neurons caused by gain-of-function alleles of Son of sevenless and Dsor1...
  21. Baxevanis A, Arents G, Moudrianakis E, Landsman D. A variety of DNA-binding and multimeric proteins contain the histone fold motif. Nucleic Acids Res. 1995;23:2685-91 pubmed
    ..It is proposed that these proteins may share a similar three-dimensional conformation despite the lack of significant sequence similarity. ..
  22. Ni J, Liu L, Binari R, Hardy R, Shim H, Cavallaro A, et al. A Drosophila resource of transgenic RNAi lines for neurogenetics. Genetics. 2009;182:1089-100 pubmed publisher
  23. Lu X, Li Y. Drosophila Src42A is a negative regulator of RTK signaling. Dev Biol. 1999;208:233-43 pubmed
    ..Our study provides the first evidence that Src42A defines a negative regulatory pathway parallel to Ras1 in the RTK signaling cascade. A possible model for Src42A function is discussed. ..
  24. Kunda P, Craig G, Dominguez V, Baum B. Abi, Sra1, and Kette control the stability and localization of SCAR/WAVE to regulate the formation of actin-based protrusions. Curr Biol. 2003;13:1867-75 pubmed
    ..By controlling the stability, localization, and function of SCAR, these proteins may help to ensure that Arp2/3 activation and the generation of actin-based protrusions remain strictly dependant on local GTPase signaling. ..
  25. Oishi K, Zhang H, Gault W, Wang C, Tan C, Kim I, et al. Phosphatase-defective LEOPARD syndrome mutations in PTPN11 gene have gain-of-function effects during Drosophila development. Hum Mol Genet. 2009;18:193-201 pubmed publisher
  26. Greenwald I, Rubin G. Making a difference: the role of cell-cell interactions in establishing separate identities for equivalent cells. Cell. 1992;68:271-81 pubmed
  27. Boutla A, Delidakis C, Tabler M. Developmental defects by antisense-mediated inactivation of micro-RNAs 2 and 13 in Drosophila and the identification of putative target genes. Nucleic Acids Res. 2003;31:4973-80 pubmed
    ..These genes, which include Sos and Myd88, represent putative targets for miRNA regulation...
  28. Bergwitz C, Wee M, Sinha S, Huang J, Derobertis C, Mensah L, et al. Genetic determinants of phosphate response in Drosophila. PLoS ONE. 2013;8:e56753 pubmed publisher
    ..We identified a number of candidate genes potentially important for the cellular response to phosphate. Evaluation of 51 genes in live flies revealed some that affect larval development, adult life span and hemolymph phosphate levels. ..
  29. Kramer H. Patrilocal* cell-cell interactions: sevenless captures its bride. Trends Cell Biol. 1993;3:103-5 pubmed
  30. Livingston B, Wilt F. Cell interactions in invertebrate development. Curr Opin Cell Biol. 1993;5:839-43 pubmed
  31. Meisner H, Daga A, Buxton J, Fernandez B, Chawla A, Banerjee U, et al. Interactions of Drosophila Cbl with epidermal growth factor receptors and role of Cbl in R7 photoreceptor cell development. Mol Cell Biol. 1997;17:2217-25 pubmed
  32. Dickson B. Nuclear factors in sevenless signalling. Trends Genet. 1995;11:106-11 pubmed
  33. Innocenti P, Chenoweth S. Interspecific divergence of transcription networks along lines of genetic variance in Drosophila: dimensionality, evolvability, and constraint. Mol Biol Evol. 2013;30:1358-67 pubmed publisher
  34. Hafen E, Dickson B, Raabe T, Brunner D, Oellers N, van der Straten A. Genetic analysis of the sevenless signal transduction pathway of Drosophila. Dev Suppl. 1993;:41-6 pubmed
    ..In contrast to the receptor and its ligand, these components are shared between different signal transduction pathways not only in Drosophila but homologous components are also involved in signal transduction in other organisms. ..
  35. Perrimon N, Desplan C. Signal transduction in the early Drosophila embryo: when genetics meets biochemistry. Trends Biochem Sci. 1994;19:509-13 pubmed
    ..This pathway involves a membrane-bound receptor tyrosine kinase (RTK) that is similar to other Drosophila RTKs, such as sevenless, and the mammalian RTKs, such as the epidermal growth factor or platelet-derived growth factor receptors. ..
  36. Bhadra U, Pal Bhadra M, Birchler J. Histone acetylation and gene expression analysis of sex lethal mutants in Drosophila. Genetics. 2000;155:753-63 pubmed
    ..In both cases we find relatively little effect upon X chromosomal gene expression. ..
  37. Lim Y, Tsuda L, Inoue Y, Irie K, Adachi Yamada T, Hata M, et al. Dominant mutations of Drosophila MAP kinase kinase and their activities in Drosophila and yeast MAP kinase cascades. Genetics. 1997;146:263-73 pubmed
    ..The results suggest that they function as suppressors due to increased reactivity to upstream factors. ..
  38. Karlovich C, Bonfini L, McCollam L, Rogge R, Daga A, Czech M, et al. In vivo functional analysis of the Ras exchange factor son of sevenless. Science. 1995;268:576-9 pubmed
    The Son of sevenless (Sos) protein functions as a guanine nucleotide transfer factor for Ras and interacts with the receptor tyrosine kinase Sevenless through the protein Drk, a homolog of mammalian Grb2...
  39. Hales K, Korey C, Larracuente A, Roberts D. Genetics on the Fly: A Primer on the Drosophila Model System. Genetics. 2015;201:815-42 pubmed publisher
  40. den Hertog J. Protein-tyrosine phosphatases in development. Mech Dev. 1999;85:3-14 pubmed
    ..Here I will introduce PTP-mediated signalling and discuss recent findings regarding the function of PTPs in development. ..
  41. Fritz J, VanBerkum M. Calmodulin and son of sevenless dependent signaling pathways regulate midline crossing of axons in the Drosophila CNS. Development. 2000;127:1991-2000 pubmed
    ..Here, a genetic approach in Drosophila has demonstrated that both Calmodulin and Son of sevenless signaling pathways are used to regulate which axons cross the midline...
  42. Lin M, Greenberg M. Orchestral maneuvers in the axon: trio and the control of axon guidance. Cell. 2000;101:239-42 pubmed
  43. Iyengar B, Roote J, Campos A. The tamas gene, identified as a mutation that disrupts larval behavior in Drosophila melanogaster, codes for the mitochondrial DNA polymerase catalytic subunit (DNApol-gamma125). Genetics. 1999;153:1809-24 pubmed
    ..Sequencing of mutant alleles demonstrated that tamas codes for the mitochondrial DNA polymerase catalytic subunit (DNApol-gamma125). ..
  44. Kim S, Kim J, Malik S, Son W, Kwon K, Kim C. Negative regulation of EGFR/MAPK pathway by Pumilio in Drosophila melanogaster. PLoS ONE. 2012;7:e34016 pubmed publisher
    ..Taken together, our evidence suggests that Pum functions as a negative regulator of EGFR signaling by directly targeting components of the pathway in Drosophila. ..
  45. Pawson T, Olivier P, Rozakis Adcock M, McGlade J, Henkemeyer M. Proteins with SH2 and SH3 domains couple receptor tyrosine kinases to intracellular signalling pathways. Philos Trans R Soc Lond B Biol Sci. 1993;340:279-85 pubmed
    ..Evidence suggesting that SH2 and SH3 domains act synergistically in stimulation of the Ras pathway is discussed. ..
  46. Lu X, Chou T, Williams N, Roberts T, Perrimon N. Control of cell fate determination by p21ras/Ras1, an essential component of torso signaling in Drosophila. Genes Dev. 1993;7:621-32 pubmed
    ..Furthermore, we have analyzed the maternal-effect phenotype of Son of sevenless (Sos), a positive regulator of Ras1, and showed that embryos derived from germ cells lacking Sos+ activity ..
  47. Cherniack A, Klarlund J, Czech M. Phosphorylation of the Ras nucleotide exchange factor son of sevenless by mitogen-activated protein kinase. J Biol Chem. 1994;269:4717-20 pubmed
    b>Son of sevenless-1 and -2 (Sos-1 and -2) are guanosine nucleotide exchange factors implicated in the activation of Ras by both the insulin and epidermal growth factor signal transduction pathways...
  48. Park J, Al Ramahi I, Tan Q, Mollema N, Diaz Garcia J, Gallego Flores T, et al. RAS-MAPK-MSK1 pathway modulates ataxin 1 protein levels and toxicity in SCA1. Nature. 2013;498:325-331 pubmed publisher
    ..Collectively, these data reveal new therapeutic entry points for SCA1 and provide a proof-of-principle for tackling other classes of intractable neurodegenerative diseases. ..
  49. Firth L, Baker N. Extracellular signals responsible for spatially regulated proliferation in the differentiating Drosophila eye. Dev Cell. 2005;8:541-51 pubmed
    ..These studies identify the specific extracellular signals that pattern the retinal cell cycles and show how differentiation can be uncoupled from cell cycle exit. ..
  50. Hafen E, Basler K. Specification of cell fate in the developing eye of Drosophila. Dev Suppl. 1991;1:123-30 pubmed
    ..Two components of this cascade--Sos and sina--have been identified genetically. sina encodes a nuclear protein whose expression is not limited to R7...
  51. The I, Hannigan G, Cowley G, Reginald S, Zhong Y, Gusella J, et al. Rescue of a Drosophila NF1 mutant phenotype by protein kinase A. Science. 1997;276:791-4 pubmed
    ..Thus, NF1 and PKA appear to interact in a pathway that controls the overall growth of Drosophila. ..
  52. Harrison S, Rudolph J, Spencer M, Wes P, Montell C, Andres D, et al. Activated RIC, a small GTPase, genetically interacts with the Ras pathway and calmodulin during Drosophila development. Dev Dyn. 2005;232:817-26 pubmed
    ..RIC could be mitigated by a reduction in dosage of several genes in the Drosophila Ras cascade, including Son of sevenless (Sos), Dsor (MEK), rolled (MAPK), and Ras itself...
  53. Cabernard C, Affolter M. Distinct roles for two receptor tyrosine kinases in epithelial branching morphogenesis in Drosophila. Dev Cell. 2005;9:831-42 pubmed
    ..Ras and the Mapk pathway are also needed for Egf-regulated cell division/survival, but Pointed is dispensable. ..
  54. Simon M, Bowtell D, Dodson G, Laverty T, Rubin G. Ras1 and a putative guanine nucleotide exchange factor perform crucial steps in signaling by the sevenless protein tyrosine kinase. Cell. 1991;67:701-16 pubmed
    ..These results suggest that the stimulation of ras protein activity is a key element in the signaling by sevenless and Ellipse and that this stimulation may be achieved by activating the exchange of GTP for bound GDP by the ras protein. ..
  55. Hsouna A, VanBerkum M. Abelson tyrosine kinase and Calmodulin interact synergistically to transduce midline guidance cues in the Drosophila embryonic CNS. Int J Dev Neurosci. 2008;26:345-54 pubmed publisher
  56. Végh M, Basler K. A genetic screen for hedgehog targets involved in the maintenance of the Drosophila anteroposterior compartment boundary. Genetics. 2003;163:1427-38 pubmed
    ..Among these, we identified a class of patched (ptc) alleles exhibiting a novel phenotype. These results demonstrate the value of our setup in the identification of genes involved in distinct wing-patterning processes. ..
  57. Coleman H, Labrador J, Chance R, Bashaw G. The Adam family metalloprotease Kuzbanian regulates the cleavage of the roundabout receptor to control axon repulsion at the midline. Development. 2010;137:2417-26 pubmed publisher
    ..Finally, Kuz activity is required for Robo to recruit its downstream signaling partner, Son of sevenless (Sos)...
  58. Charng W, Yamamoto S, Bellen H. Shared mechanisms between Drosophila peripheral nervous system development and human neurodegenerative diseases. Curr Opin Neurobiol. 2014;27:158-64 pubmed publisher
    ..In this review, we summarize and discuss how studies of Drosophila PNS and NMJ development have provided guidance in experimental approaches for these diseases. ..
  59. Dorn R, Szidonya J, Korge G, Sehnert M, Taubert H, Archoukieh E, et al. P transposon-induced dominant enhancer mutations of position-effect variegation in Drosophila melanogaster. Genetics. 1993;133:279-90 pubmed
    ..First studies on the developmentally regulated gene expression of PEV enhancer genes were performed by beta-galactosidase staining in P[lArB] induced mutations. ..
  60. Chou T, Noll E, Perrimon N. Autosomal P[ovoD1] dominant female-sterile insertions in Drosophila and their use in generating germ-line chimeras. Development. 1993;119:1359-69 pubmed
    ..Specifically, we show that the Gap1 gene, which encodes a Drosophila homologue of mammalian GTPase-activating protein, is required in somatic follicle cells for embryonic dorsoventral polarity determination. ..
  61. Kramer S, West S, Hiromi Y. Cell fate control in the Drosophila retina by the orphan receptor seven-up: its role in the decisions mediated by the ras signaling pathway. Development. 1995;121:1361-72 pubmed
    ..However, a decrease in ras signaling results in enhancement of the phenotype caused by the ro-svp transgene. We discuss the relationship between decisions controlled by seven-up and those controlled by ras signaling. ..
  62. Rubin G. Signal transduction and the fate of the R7 photoreceptor in Drosophila. Trends Genet. 1991;7:372-7 pubmed
    ..Studies of the mechanisms governing the specificity of signal transmission and reception suggest that the sevenless signal directs a bipotential cell towards a neuronal rather than a cone cell fate. ..
  63. Silver S, Chen F, Doyon L, Zink A, Rebay I. New class of Son-of-sevenless (Sos) alleles highlights the complexities of Sos function. Genesis. 2004;39:263-72 pubmed
    The guanine nucleotide exchange factor (GEF) Son-of-sevenless (Sos) encodes a complex multidomain protein best known for its role in activating the small GTPase RAS in response to receptor tyrosine kinase (RTK) stimulation...
  64. Woodruff R, Ashburner M. The genetics of a small autosomal region of Drosophila melanogaster containing the structural gene for alcohol dehydrogenase. II. Lethal mutations in the region. Genetics. 1979;92:133-49 pubmed
  65. Biggs W, Zavitz K, Dickson B, van der Straten A, Brunner D, Hafen E, et al. The Drosophila rolled locus encodes a MAP kinase required in the sevenless signal transduction pathway. EMBO J. 1994;13:1628-35 pubmed
    ..of the signal transduction pathway activated by Sev in the R7 precursor include proteins encoded by the gap1, drk, Sos, ras1 and raf loci...
  66. Okabe M, Okano H. Two-step induction of chordotonal organ precursors in Drosophila embryogenesis. Development. 1997;124:1045-53 pubmed
    ..Three of these argos-expressing cells differentiate into the three remaining COPs and they prevent neighboring cells from becoming extra COPs. ..
  67. Simon M, Dodson G, Rubin G. An SH3-SH2-SH3 protein is required for p21Ras1 activation and binds to sevenless and Sos proteins in vitro. Cell. 1993;73:169-77 pubmed
    ..of p21Ras1 but not for any subsequent events, and that this protein can bind in vitro to sevenless and to Son of sevenless (Sos), a putative guanine nucleotide exchange factor for p21Ras1...
  68. Venkatesh T. Neuronal development in the Drosophila retina. J Neurobiol. 1993;24:740-56 pubmed
    ..Recent genetic and molecular studies have provided important clues regarding the nature of the molecules involved in cellular signalling and neuronal differentiation. ..
  69. Butler S, Ray S, Hiromi Y. klingon, a novel member of the Drosophila immunoglobulin superfamily, is required for the development of the R7 photoreceptor neuron. Development. 1997;124:781-92 pubmed
    ..Genetic analysis has revealed that klingon is an essential gene that participates in the development of the R7 neuron. Ectopic expression of klingon in all neurons in a sevenless background can alter the position of the R8 rhabdomere. ..
  70. Fritz J, VanBerkum M. Regulation of rho family GTPases is required to prevent axons from crossing the midline. Dev Biol. 2002;252:46-58 pubmed
    ..Homozygous loss of Son-of-Sevenless (Sos) also enhances the ctDrac phenotype and causes errors in embryos expressing either dnRho or dnDrac...
  71. Hou X, Chou T, Melnick M, Perrimon N. The torso receptor tyrosine kinase can activate Raf in a Ras-independent pathway. Cell. 1995;81:63-71 pubmed
    ..This result is supported by analysis of D-Raf activation in the absence of either the exchange factor Son of sevenless (Sos) or the adaptor protein drk (Grb2), as well as by the phenotype of a D-Raf mutation that abolishes ..
  72. Lu X, Perkins L, Perrimon N. The torso pathway in Drosophila: a model system to study receptor tyrosine kinase signal transduction. Dev Suppl. 1993;:47-56 pubmed
    ..a signal transduction pathway that is mediated sequentially by the guanine nucleotide releasing factor son of sevenless (Sos), the p21Ras1 GTPase, the serine/threonine kinase D-raf and the tyrosine/threonine kinase MAPKK (Dsor1)...
  73. Williams J, Su H, Bernards A, Field J, Sehgal A. A circadian output in Drosophila mediated by neurofibromatosis-1 and Ras/MAPK. Science. 2001;293:2251-6 pubmed
  74. Ruan W, Srinivasan A, Lin S, Kara k, Barker P. Eiger-induced cell death relies on Rac1-dependent endocytosis. Cell Death Dis. 2016;7:e2181 pubmed publisher
    ..These findings reveal novel regulatory mechanisms that allow Rac1 to contribute to Egr-induced JNK activation and cell death. ..
  75. van Dam T, Rehmann H, Bos J, Snel B. Phylogeny of the CDC25 homology domain reveals rapid differentiation of Ras pathways between early animals and fungi. Cell Signal. 2009;21:1579-85 pubmed publisher
    ..Finally, a comparison of the domain composition of the Ras guanine nucleotide exchange factors shows that domain addition and diversification occurred both prior to and after the fungal-animal split. ..
  76. Yan H, Chin M, Horvath E, Kane E, Pfleger C. Impairment of ubiquitylation by mutation in Drosophila E1 promotes both cell-autonomous and non-cell-autonomous Ras-ERK activation in vivo. J Cell Sci. 2009;122:1461-70 pubmed publisher dramatically suppressed by removing one copy of Ras, removal of the more upstream components Egfr, Grb2 or sos shows no suppression...
  77. Mousset S, Brazier L, Cariou M, Chartois F, Depaulis F, Veuille M. Evidence of a high rate of selective sweeps in African Drosophila melanogaster. Genetics. 2003;163:599-609 pubmed
    ..Polymorphism data from five loci from intervening regions (dpp, Acp26Ab, Acp29AB, GH10711, and Sos) did not show the characteristic deviation from neutrality caused by local selective sweeps...
  78. Taguchi A, Sawamoto K, Okano H. Mutations modulating the Argos-regulated signaling pathway in Drosophila eye development. Genetics. 2000;154:1639-48 pubmed
    ..In addition to the known genes, we identified six mutations of novel genes. Genetic characterization of these mutants suggested that they have distinct roles in cell differentiation and/or survival regulated by the EGF receptor pathway. ..
  79. Bhadra U, Pal Bhadra M, Birchler J. A trans-acting modifier causing extensive overexpression of genes in Drosophila melanogaster. Mol Gen Genet. 1997;254:621-34 pubmed
    ..Regulatory genes such as Ufo may contribute to the wide fluctuations in gene expression between tissues and stages of development...
  80. MacDougall L, Waterfield M. Receptor signalling: to sevenless, a daughter. Curr Biol. 1996;6:1250-3 pubmed
    ..But the discovery of a substrate for the Sevenless receptor indicates that the concept of a single, linear pathway may be an oversimplification. ..
  81. Simon M. Signal transduction during the development of the Drosophila R7 photoreceptor. Dev Biol. 1994;166:431-42 pubmed
  82. Malartre M, Ayaz D, Amador F, Martín Bermudo M. The guanine exchange factor vav controls axon growth and guidance during Drosophila development. J Neurosci. 2010;30:2257-67 pubmed publisher
    ..Thus, we propose that vav acts in coordination with other GEFs to regulate axon growth and guidance during development by linking guidance signals to the cytoskeleton via the modulation of Rac activity. ..
  83. McNeill H, Downward J. Apoptosis: Ras to the rescue in the fly eye. Curr Biol. 1999;9:R176-9 pubmed
    ..Recent results on the way Ras and downstream pathways can influence cell-death induction suggest the regulatory pathways in these distinct organisms might be more similar than was at first sight apparent. ..
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