Gene Symbol: e(y)2
Description: enhancer of yellow 2
Alias: CG15191, Dmel\CG15191, ENY2, Sus1, enhancer of yellow 2, CG15191-PA, e(y)2-PA
Species: fruit fly
Products:     e(y)2

Top Publications

  1. Georgiev P. Identification of mutations in three genes that interact with zeste in the control of white gene expression in Drosophila melanogaster. Genetics. 1994;138:733-9 pubmed
    ..The products of these three genes may represent, together with zeste, a group of proteins involved in the organization of long-distance interactions between DNA sequences...
  2. Kopytova D, Orlova A, Krasnov A, Gurskiy D, Nikolenko J, Nabirochkina E, et al. Multifunctional factor ENY2 is associated with the THO complex and promotes its recruitment onto nascent mRNA. Genes Dev. 2010;24:86-96 pubmed publisher
    Metazoan E(y)2/ENY2 is a multifunctional protein important for transcription activation and mRNA export, being a component of SAGA/TFTC and the mRNA export complex AMEX...
  3. Kurshakova M, Maksimenko O, Golovnin A, Pulina M, Georgieva S, Georgiev P, et al. Evolutionarily conserved E(y)2/Sus1 protein is essential for the barrier activity of Su(Hw)-dependent insulators in Drosophila. Mol Cell. 2007;27:332-8 pubmed
    ..Here we demonstrate that an evolutionarily conserved protein, E(y)2/Sus1, is recruited to the Su(Hw) insulators via binding to the zinc-finger domain of Su(Hw)...
  4. Gurskiy D, Orlova A, Vorobyeva N, Nabirochkina E, Krasnov A, Shidlovskii Y, et al. The DUBm subunit Sgf11 is required for mRNA export and interacts with Cbp80 in Drosophila. Nucleic Acids Res. 2012;40:10689-700 pubmed publisher
    ..Drosophila DUBm consists of Sgf11, ENY2 and Nonstop proteins. We show that Sgf11 has other DUBm-independent functions...
  5. Rodriguez Navarro S. Insights into SAGA function during gene expression. EMBO Rep. 2009;10:843-50 pubmed publisher
    ..I focus particularly on the new components of human SAGA, which was recently discovered and confirms the conservation of the SAGA complex throughout evolution. ..
  6. Zhao Y, Lang G, Ito S, Bonnet J, Metzger E, Sawatsubashi S, et al. A TFTC/STAGA module mediates histone H2A and H2B deubiquitination, coactivates nuclear receptors, and counteracts heterochromatin silencing. Mol Cell. 2008;29:92-101 pubmed publisher
    ..We have identified three additional subunits of TFTC/STAGA (ATXN7L3, USP22, and ENY2) that form the deubiquitination module...
  7. Kurshakova M, Krasnov A, Kopytova D, Shidlovskii Y, Nikolenko J, Nabirochkina E, et al. SAGA and a novel Drosophila export complex anchor efficient transcription and mRNA export to NPC. EMBO J. 2007;26:4956-65 pubmed
    ..Thus, E(y)2 and Xmas-2 together with SAGA/TFTC function in the anchoring of a subset of transcription sites to the NPCs to achieve efficient transcription and mRNA export. ..
  8. Georgieva S, Nabirochkina E, Dilworth F, Eickhoff H, Becker P, Tora L, et al. The novel transcription factor e(y)2 interacts with TAF(II)40 and potentiates transcription activation on chromatin templates. Mol Cell Biol. 2001;21:5223-31 pubmed
  9. Xie G, Yu Z, Jia D, Jiao R, Deng W. E(y)1/TAF9 mediates the transcriptional output of Notch signaling in Drosophila. J Cell Sci. 2014;127:3830-9 pubmed publisher

More Information


  1. Georgiev P, Kiselev S, Simonova O, Gerasimova T. A novel transposition system in Drosophila melanogaster depending on the Stalker mobile genetic element. EMBO J. 1990;9:2037-44 pubmed
    ..Many known and novel mutations have been obtained. Comparison of their genetic localization with Stalker distribution suggests that the majority of them have been induced by the Stalker insertion. ..
  2. Orlova A, Kopytova D, Krasnov A, Nabirochkina E, Ilyin Y, Georgieva S, et al. Transcriptional factor ENY2 promotes recruitment of the THO complex to the hsp70 gene of Drosophila melanogaster. Dokl Biochem Biophys. 2010;434:227-31 pubmed publisher
  3. Kyrchanova O, Maksimenko O, Stakhov V, Ivlieva T, Parshikov A, Studitsky V, et al. Effective blocking of the white enhancer requires cooperation between two main mechanisms suggested for the insulator function. PLoS Genet. 2013;9:e1003606 pubmed publisher
  4. Georgieva S, Nabirochkina E, Pustovoĭtov M, Krasnov A, Soldatov A. [Molecular characteristics of the new evolutionary-conserved nuclear protein e(y)2]. Genetika. 2001;37:18-23 pubmed
    Molecular structural and function analyses of the Drosophila melanogaster enhancer of yellow 2 (e(y)2) gene showed that its product acts as a transcription factor and is one of the basic elements of the eukaryotic transcription system...
  5. Bezborodova E, Kulikov A, Georgiev P. A new family of genes which, when mutated, suppress the inhibitory effect of the mod(mdg4)1u1 mutation on y2 expression in Drosophila melanogaster. Mol Gen Genet. 1997;257:83-90 pubmed
    ..Most of Su(mg) mutations do not affect the viability or fertility of homozygous flies. We speculate that the Su(mg) genes represent a new family of redundant regulatory genes in Drosophila melanogaster. ..
  6. Soldatov A, Nabirochkina E, Dzitoeva S, Matiunina L, Georgiev P. [Molecular cloning of regions of localization of the e(y)1 and e(y)2 genes in Drosophila melanogaster]. Genetika. 1996;32:1714-6 pubmed
    ..Molecular cloning of the e(y)1 and e(y)2 gene regions is described. The results of cloning were confirmed by experiments on the rescue of the mutant phenotype. ..
  7. Elagin V, Medvedeva N, Georgiev P. [Mutations affecting intrachromosomal recombination and conversion in the binary y(2ns)sc(me) system in Drosophila melanogaster]. Genetika. 1994;30:330-6 pubmed
    ..Some of the mutations under study also modify the phenotypic manifestation of the y2nsscme allele. ..
  8. Georgieva S, Nabirochkina E, Ladygina N, Georgiev P, Soldatov A. [Nuclear protein e(y)2 from Drosophila melanogaster participates in transcription control]. Genetika. 2001;37:24-8 pubmed
    ..Genetic analysis showed that the C-terminal amino acid residues of transcription factor TAFII40 are important for its interaction with e(y)2. ..
  9. Belen kaia T, Georgiev P. [Role of the "enhancer of yellow" genes in the regulation of expression of the "yellow" gene in Drosophila melanogaster]. Genetika. 1999;35:432-7 pubmed
    ..When introduced at +2490 from the transcription initiation site, the su(Hw) insulator becomes able to block interactions between the bristle enhancer and yellow promoter, not separating them from each other. ..
  10. Kozitsina M, Abramova N, Georgiev P. [Interaction between the enhancers of yellow and the yellow gene in Drosophila melanogaster]. Genetika. 1992;28:59-67 pubmed
    ..At the same time, the interaction of here described gene e(y)7 with the yellow gene is likely to occur at the level of protein products. ..
  11. Gurskiy D, Nabirochkina E, Kopytova D, Nikolenko Y, Ilyin Y, Georgieva S, et al. ENY2 protein forms a part of the THO complex of Drosophila melanogaster. Dokl Biochem Biophys. 2010;433:212-5 pubmed publisher
  12. Krasnov A, Kurshakova M, Ramensky V, Mardanov P, Nabirochkina E, Georgieva S. A retrocopy of a gene can functionally displace the source gene in evolution. Nucleic Acids Res. 2005;33:6654-61 pubmed publisher
    ..This must have resulted from retroposition into a transcriptionally favorable region of the genome...
  13. Mohan R, Dialynas G, Weake V, Liu J, Martin Brown S, Florens L, et al. Loss of Drosophila Ataxin-7, a SAGA subunit, reduces H2B ubiquitination and leads to neural and retinal degeneration. Genes Dev. 2014;28:259-72 pubmed publisher
    ..When we examined the consequences of reduced Ataxin-7 in vivo, we found that flies exhibited pronounced neural and retinal degeneration, impaired movement, and early lethality...
  14. Melnikova L, Kulikov A, Georgiev P. Interactions between cut wing mutations and mutations in zeste, and the enhancer of yellow and Polycomb group genes of Drosophila melanogaster. Mol Gen Genet. 1996;252:230-6 pubmed
    ..The results are discussed in terms of the role of long-distance interactions and DNA compaction in transcriptional control of the cut locus. ..
  15. Georgiev P, Gerasimova T. Novel genes influencing the expression of the yellow locus and mdg4 (gypsy) in Drosophila melanogaster. Mol Gen Genet. 1989;220:121-6 pubmed
    ..All e(y)n genes are located in different regions of the X chromosome. One may speculate that e(y)n genes are involved in trans-regulation of the yellow locus and possibly of some other loci. ..
  16. Gurskiĭ D, Orlova A, Kopytova D, Krasnov A, Nabirochkina E, Georgieva S, et al. [Multifunctional factor ENY2 integrates the various stages of gene expression]. Genetika. 2010;46:1700-3 pubmed
    The ENY2/Sus1 is a multifunctional transcription factor which couples transcription with mRNA export. It is the component of the SAGA/TFTC and TREX-2/AMEX protein complexes...
  17. Bonchuk A, Georgiev P, Maksimenko O. CTCF and Sgfl1 proteins form alternative complexes with ENY2 proteins. Dokl Biochem Biophys. 2016;468:180-2 pubmed publisher
    b>ENY2 is a multifunctional protein, a component of the SAGA complex (protein complex involved in the regulation of transcription, possessing histone-acetyltransferase and H2B-histone-deubiquitinylating activities [1]) and TREX-2 (complex ..
  18. Vorobyeva N, Mazina M, Golovnin A, Kopytova D, Gurskiy D, Nabirochkina E, et al. Insulator protein Su(Hw) recruits SAGA and Brahma complexes and constitutes part of Origin Recognition Complex-binding sites in the Drosophila genome. Nucleic Acids Res. 2013;41:5717-30 pubmed publisher
    ..Su(Hw) is the first example of such a protein. ..
  19. Maksimenko O, Kyrchanova O, Bonchuk A, Stakhov V, Parshikov A, Georgiev P. Highly conserved ENY2/Sus1 protein binds to Drosophila CTCF and is required for barrier activity. Epigenetics. 2014;9:1261-70 pubmed publisher
    ..Here we demonstrate that the evolutionarily conserved protein ENY2 is recruited to the zinc-finger domain of dCTCF and is required for the barrier activity of dCTCF-dependent ..
  20. Kopytova D, Popova V, Kurshakova M, Shidlovskii Y, Nabirochkina E, Brechalov A, et al. ORC interacts with THSC/TREX-2 and its subunits promote Nxf1 association with mRNP and mRNA export in Drosophila. Nucleic Acids Res. 2016;44:4920-33 pubmed publisher
    ..The knockdown of Orc5, Orc3 and several other ORC subunits led to an accumulation of mRNA in the nucleus, suggesting that ORC participates in the regulation of the mRNP export. ..
  21. Brechalov A, Gurskiĭ D, Georgieva S, Shidlovskiĭ I. [Functioning of macromolecular complexes at successive stages of gene expression as self-coordinated molecular machines]. Biofizika. 2011;56:831-9 pubmed
    The results of systemic investigations devoted to the role of the proteins SAYP and ENY2 from Drosophila melanogaster in the regulation of different stages of gene expression are presented...
  22. Erokhin M, Parshikov A, Georgiev P, Chetverina D. E(y)2/Sus1 is required for blocking PRE silencing by the Wari insulator in Drosophila melanogaster. Chromosoma. 2010;119:243-53 pubmed publisher
    ..Thus, the E(y)2 protein in different Drosophila insulators serves to protect gene expression from silencing. ..
  23. Kurshakova m, Kopytova D, Nabirochkina E, Nikolenko I, Shidlovskiĭ I, Georgieva S, et al. [Conservative E(y)2/Sus1 protein is the member of SAGA complex and new nuclear pore-associated complex in Drosophila]. Genetika. 2009;45:1332-40 pubmed
    ..Thus, E(y)2 and Xmas-2 together with SAGA/TFTC functioned in the anchoring of a subset of transcription sites to the NPCs to achieve efficient transcription and mRNA export. ..
  24. Lee C, Reichman T, Baik T, Mathews M. MLE functions as a transcriptional regulator of the roX2 gene. J Biol Chem. 2004;279:47740-5 pubmed
    ..Our observations suggest a novel mechanism by which roX2 RNA is regulated at the transcriptional level. ..
  25. Gause M, Georgieva S, Georgiev P. Phenotypic reversion of the gypsy-induced mutation scD1 of Drosophila melanogaster by replicative transposition of a sc enhancer to the yellow gene and by mutations in the enhancer of yellow and zeste loci. Mol Gen Genet. 1996;253:370-6 pubmed
    ..In addition, we found that mutations in the e(y)1, e(y)3 and zeste genes may interfere with transcriptional insulation by the su(Hw)-binding region in the scD1 allele. ..
  26. Kurshakova m, Nabirochkina E, Georgieva S. [E(y)2, the novel component of SAGA/TFTC complex in eucaryotes participates in export of mRNP from nucleus and couples transcription with nuclear pore]. Mol Biol (Mosk). 2009;43:253-63 pubmed
    ..The new protein E(y)2 which plays an important role in these processes is described. ..
  27. Kurshakova m, Kopytova D, Nabirochkina E, Soshnikova N, Georgieva S, Krasnov A. [Conservative E(y)2/Sus1 protein interacts with the Su(Hw)-dependent insulators in Drosophila]. Genetika. 2009;45:330-5 pubmed
    ..In this study it was demonstrated that novel, evolutionary conservative transcription factor E(y)2/Sus1 interacted with the Su(Hw) zinc-finger domain and was present in the protein complex, associated with the Su(Hw)-..