Gene Symbol: Su(var)3-9
Description: Suppressor of variegation 3-9
Alias: CG43664, CG6476, Dmel\CG43664, Dmel_CG6476, E(var)3-2, SU(VAR)3-9, SUV39, SUV39H1, Su(VAR)3-9, Su(Var)3-9, Su(var), Su(var) 3-9, Su(var)3-902, Su(var)306, Su(var)310, Su(var)328, Su-var(3)9, Su-var(3)902, SuVar3-9, Suv 3-9, Suvar(3)9, Suvar3-9, Suvar39, caa56376 Suvar39 Dm, dSU(VAR)3-9, dSu(var)3-9, ptn, su(var)3-9, suppressor of variegation 3-9, CG43664-PA, Su(var)3-9-PA, enhancer of variegation 3-2, histone methyltransferase, pitkin, suppressor of variegation 306, suppressor of variegation 310, suppressor of variegation 328
Species: fruit fly

Top Publications

  1. Lu X, Wontakal S, Kavi H, Kim B, Guzzardo P, Emelyanov A, et al. Drosophila H1 regulates the genetic activity of heterochromatin by recruitment of Su(var)3-9. Science. 2013;340:78-81 pubmed publisher
    ..We propose that H1 plays a key role in silencing by tethering Su(var)3-9 to heterochromatin. The tethering function of H1 adds to its established role as a regulator of chromatin compaction and accessibility. ..
  2. Alvarez Venegas R, Avramova Z. SET-domain proteins of the Su(var)3-9, E(z) and trithorax families. Gene. 2002;285:25-37 pubmed
    ..The architecture of the entire protein supported the distribution pattern built upon SET-domain similarity. Parallel cladistic and protein-architecture analyses outlined two plausible criteria for predicting function...
  3. Riddle N, Minoda A, Kharchenko P, Alekseyenko A, Schwartz Y, Tolstorukov M, et al. Plasticity in patterns of histone modifications and chromosomal proteins in Drosophila heterochromatin. Genome Res. 2011;21:147-63 pubmed publisher
    ..This comprehensive analysis provides a foundation for future studies of gene activity and chromosomal functions that are influenced by or dependent upon heterochromatin...
  4. Eskeland R, Czermin B, Boeke J, Bonaldi T, Regula J, Imhof A. The N-terminus of Drosophila SU(VAR)3-9 mediates dimerization and regulates its methyltransferase activity. Biochemistry. 2004;43:3740-9 pubmed
    In most eukaryotes, the histone methyltransferase SU(VAR)3-9 and its orthologues play a major role in the function of centromeric heterochromatin...
  5. Lemos B, Branco A, Hartl D. Epigenetic effects of polymorphic Y chromosomes modulate chromatin components, immune response, and sexual conflict. Proc Natl Acad Sci U S A. 2010;107:15826-31 pubmed publisher
  6. Grewal S, Elgin S. Transcription and RNA interference in the formation of heterochromatin. Nature. 2007;447:399-406 pubmed
    ..Silencing of chromatin might involve trans-acting sources of the crucial small RNAs that carry out RNA interference, but in some cases, transcription of the region to be silenced seems to be required--an apparent contradiction. ..
  7. Rudolph T, Yonezawa M, Lein S, Heidrich K, Kubicek S, Schafer C, et al. Heterochromatin formation in Drosophila is initiated through active removal of H3K4 methylation by the LSD1 homolog SU(VAR)3-3. Mol Cell. 2007;26:103-15 pubmed
  8. Dillon S, Zhang X, Trievel R, Cheng X. The SET-domain protein superfamily: protein lysine methyltransferases. Genome Biol. 2005;6:227 pubmed
    ..The SET-domain protein methyltransferase superfamily includes all but one of the proteins known to methylate histones on lysine. Histone methylation is important in the regulation of chromatin and gene expression. ..
  9. Zhang Y, Lin N, Carroll P, Chan G, Guan B, Xiao H, et al. Epigenetic blocking of an enhancer region controls irradiation-induced proapoptotic gene expression in Drosophila embryos. Dev Cell. 2008;14:481-93 pubmed publisher
    ..Thus, direct epigenetic regulation of two proapoptotic genes controls cellular sensitivity to cytotoxic stimuli. ..

More Information


  1. Brehm A, Tufteland K, Aasland R, Becker P. The many colours of chromodomains. Bioessays. 2004;26:133-40 pubmed
  2. Boeke J, Regnard C, Cai W, Johansen J, Johansen K, Becker P, et al. Phosphorylation of SU(VAR)3-9 by the chromosomal kinase JIL-1. PLoS ONE. 2010;5:e10042 pubmed publisher
    The histone methyltransferase SU(VAR)3-9 plays an important role in the formation of heterochromatin within the eukaryotic nucleus...
  3. Haynes K, Gracheva E, Elgin S. A Distinct type of heterochromatin within Drosophila melanogaster chromosome 4. Genetics. 2007;175:1539-42 pubmed
    ..distal arm chromosome 4 heterochromatin, distinguishing these two heterochromatin types. ..
  4. Mis J, Ner S, Grigliatti T. Identification of three histone methyltransferases in Drosophila: dG9a is a suppressor of PEV and is required for gene silencing. Mol Genet Genomics. 2006;275:513-26 pubmed
    ..The combined Su(var)3-9 and dG9a mutations have severe developmental defects suggesting an overlapping role for dG9a and Su(var)3-9 in the packaging of heterochromatin and gene silencing via a K9H3 methylation pathway. ..
  5. Stassen M, Bailey D, Nelson S, Chinwalla V, Harte P. The Drosophila trithorax proteins contain a novel variant of the nuclear receptor type DNA binding domain and an ancient conserved motif found in other chromosomal proteins. Mech Dev. 1995;52:209-23 pubmed
    ..This pattern is transient and evolves into a broader expression domain encompassing the entire germ band during the extended germ band stage. ..
  6. Czermin B, Melfi R, McCabe D, Seitz V, Imhof A, Pirrotta V. Drosophila enhancer of Zeste/ESC complexes have a histone H3 methyltransferase activity that marks chromosomal Polycomb sites. Cell. 2002;111:185-96 pubmed
    ..Histone H3 methylated in vitro by the E(Z)/ESC complex binds specifically to Polycomb protein. ..
  7. Naumann K, Fischer A, Hofmann I, Krauss V, Phalke S, Irmler K, et al. Pivotal role of AtSUVH2 in heterochromatic histone methylation and gene silencing in Arabidopsis. EMBO J. 2005;24:1418-29 pubmed
    ..Gene silencing by SUVH2 depends on MET1 and DDM1, but not CMT3. In Arabidopsis, SUVH2 with its histone H3K9 and H4K20 methylation activity has a central role in heterochromatic gene silencing. ..
  8. Huang X, Yin H, Sweeney S, Raha D, Snyder M, Lin H. A major epigenetic programming mechanism guided by piRNAs. Dev Cell. 2013;24:502-16 pubmed publisher
    ..Piwi deficiency drastically changed the epigenetic landscape and polymerase II profile throughout the genome, revealing the Piwi-piRNA mechanism as a major epigenetic programming mechanism in Drosophila. ..
  9. Tschiersch B, Hofmann A, Krauss V, Dorn R, Korge G, Reuter G. The protein encoded by the Drosophila position-effect variegation suppressor gene Su(var)3-9 combines domains of antagonistic regulators of homeotic gene complexes. EMBO J. 1994;13:3822-31 pubmed
  10. Phalke S, Nickel O, Walluscheck D, Hortig F, Onorati M, Reuter G. Retrotransposon silencing and telomere integrity in somatic cells of Drosophila depends on the cytosine-5 methyltransferase DNMT2. Nat Genet. 2009;41:696-702 pubmed publisher
    ..Together, these results demonstrate a previously unappreciated role of DNA methylation in retrotransposon silencing and telomere integrity in Drosophila. ..
  11. Seum C, Bontron S, Reo E, Delattre M, Spierer P. Drosophila G9a is a nonessential gene. Genetics. 2007;177:1955-7 pubmed
    ..Here, we characterize the Drosophila homolog of G9a, dG9a. We generated a dG9a deletion allele by homologous recombination. Analysis of this allele revealed that, in contrast to recent findings, dG9a is not required for fly viability. ..
  12. Hearn M, Hedrick A, Grigliatti T, Wakimoto B. The effect of modifiers of position-effect variegation on the variegation of heterochromatic genes of Drosophila melanogaster. Genetics. 1991;128:785-97 pubmed
    ..We conclude that these heterochromatic genes have fundamentally different regulatory requirements compared to those typical of euchromatic genes. ..
  13. Danzer J, Wallrath L. Mechanisms of HP1-mediated gene silencing in Drosophila. Development. 2004;131:3571-80 pubmed
    ..Silencing was minimally affected at 1.9 kb, but eliminated at 3.7 kb, suggesting that HP1-mediated silencing can operate in a SU(VAR)3-9-independent and -dependent manner. ..
  14. Figueiredo M, Philip P, Stenberg P, Larsson J. HP1a recruitment to promoters is independent of H3K9 methylation in Drosophila melanogaster. PLoS Genet. 2012;8:e1003061 pubmed publisher
    ..Our results support a hypothesis in which HP1a nucleates with high affinity independently of H3K9me in promoters of active genes and then spreads via H3K9 methylation and transient looping contacts with those H3K9me target sites...
  15. Sims R, Nishioka K, Reinberg D. Histone lysine methylation: a signature for chromatin function. Trends Genet. 2003;19:629-39 pubmed
  16. Ner S, Harrington M, Grigliatti T. A role for the Drosophila SU(VAR)3-9 protein in chromatin organization at the histone gene cluster and in suppression of position-effect variegation. Genetics. 2002;162:1763-74 pubmed
    ..The chromosomal deficiency for the HIS-C is also a suppressor of PEV. In contrast to what might be expected, we show that hemizygosity for the HIS-C locus leads to a substantial increase in the histone transcripts. ..
  17. Peng J, Karpen G. H3K9 methylation and RNA interference regulate nucleolar organization and repeated DNA stability. Nat Cell Biol. 2007;9:25-35 pubmed
    ..These results suggest a mechanism for how local chromatin structure can regulate genome stability, and the organization of chromosomal elements and nuclear organelles. ..
  18. Zhang W, Deng H, Bao X, Lerach S, Girton J, Johansen J, et al. The JIL-1 histone H3S10 kinase regulates dimethyl H3K9 modifications and heterochromatic spreading in Drosophila. Development. 2006;133:229-35 pubmed
    ..Based on these findings, we propose a model where JIL-1 kinase activity functions to maintain euchromatic regions by antagonizing Su(var)3-9-mediated heterochromatization. ..
  19. Czermin B, Schotta G, Hülsmann B, Brehm A, Becker P, Reuter G, et al. Physical and functional association of SU(VAR)3-9 and HDAC1 in Drosophila. EMBO Rep. 2001;2:915-9 pubmed immunoaffinity purification that two activities important for chromatin-mediated gene silencing, the histone methyltransferase SU(VAR)3-9 and the histone deacetylase HDAC1, associate in vivo...
  20. Haynes K, Caudy A, Collins L, Elgin S. Element 1360 and RNAi components contribute to HP1-dependent silencing of a pericentric reporter. Curr Biol. 2006;16:2222-7 pubmed
    ..Silencing of the 1360, hsp70-white reporter is sensitive to mutations in RNAi components. Our results implicate 1360 as a target for sequence-specific heterochromatic silencing through an RNAi-dependent mechanism. ..
  21. Johansson A, Stenberg P, Bernhardsson C, Larsson J. Painting of fourth and chromosome-wide regulation of the 4th chromosome in Drosophila melanogaster. EMBO J. 2007;26:2307-16 pubmed
    ..We propose a balancing mechanism involving POF and HP1 that provides a feedback system for fine-tuning expression status of genes on the 4th chromosome. ..
  22. Krauss V, Reuter G. Two genes become one: the genes encoding heterochromatin protein Su(var)3-9 and translation initiation factor subunit eIF-2gamma are joined to a dicistronic unit in holometabolic insects. Genetics. 2000;156:1157-67 pubmed
  23. Schotta G, Ebert A, Krauss V, Fischer A, Hoffmann J, Rea S, et al. Central role of Drosophila SU(VAR)3-9 in histone H3-K9 methylation and heterochromatic gene silencing. EMBO J. 2002;21:1121-31 pubmed
    ..Like its mammalian and Schizosaccharomyces pombe homologues, Su(var) 3-9 encodes a histone methyltransferase (HMTase), which selectively methylates histone H3 at lysine 9 (H3-K9)...
  24. Cryderman D, Grade S, Li Y, Fanti L, Pimpinelli S, Wallrath L. Role of Drosophila HP1 in euchromatic gene expression. Dev Dyn. 2005;232:767-74 pubmed
    ..Collectively, these data demonstrate multiple mechanisms for HP1 localization within euchromatin and show that some genes associated with HP1 are not affected by alterations in Su(var)3-9 dosage. ..
  25. Bushey D, Locke J. Mutations in Su(var)205 and Su(var)3-7 suppress P-element-dependent silencing in Drosophila melanogaster. Genetics. 2004;168:1395-411 pubmed
    ..Studying w(+) variegation from P[lacW]ci(Dplac) provides a model for the interactions that can enhance heterochromatic silencing at single P-element inserts. ..
  26. Ebert A, Schotta G, Lein S, Kubicek S, Krauss V, Jenuwein T, et al. Su(var) genes regulate the balance between euchromatin and heterochromatin in Drosophila. Genes Dev. 2004;18:2973-83 pubmed
    ..A hyperactive Su(var)3-9 mutant, pitkin(D), displays extensive H3-K9 di- and trimethylation within but also outside pericentric heterochromatin...
  27. Perrini B, Piacentini L, Fanti L, Altieri F, Chichiarelli S, Berloco M, et al. HP1 controls telomere capping, telomere elongation, and telomere silencing by two different mechanisms in Drosophila. Mol Cell. 2004;15:467-76 pubmed
    ..Here, we tested this model, and we found that the capping function of HP1 is due to its direct binding to telomeric DNA, while the silencing of telomeric sequences and telomere elongation is due to its interaction with H3-MeK9. ..
  28. Wang C, Girton J, Johansen J, Johansen K. A balance between euchromatic (JIL-1) and heterochromatic [SU(var)2-5 and SU(var)3-9] factors regulates position-effect variegation in Drosophila. Genetics. 2011;188:745-8 pubmed publisher
  29. Sentmanat M, Elgin S. Ectopic assembly of heterochromatin in Drosophila melanogaster triggered by transposable elements. Proc Natl Acad Sci U S A. 2012;109:14104-9 pubmed publisher
  30. Yoon J, Lee K, Park J, Yu K, Paik S, Kang Y. dSETDB1 and SU(VAR)3-9 sequentially function during germline-stem cell differentiation in Drosophila melanogaster. PLoS ONE. 2008;3:e2234 pubmed publisher
    ..These results indicate that dSETDB1 and SU(VAR)3-9 act together with distinct roles during oogenesis, with dsetdb1 being of particular importance due to its GSC-specific function and more severe mutant phenotype. ..
  31. Csink A, Linsk R, Birchler J. The Lighten up (Lip) gene of Drosophila melanogaster, a modifier of retroelement expression, position effect variegation and white locus insertion alleles. Genetics. 1994;138:153-63 pubmed
    ..Additionally, Lip modifies the total transcript abundance of both the blood and copia retrotransposons, having an inverse effect on the steady state level of blood transcripts, while showing a non-additive effect on copia RNA. ..
  32. Westphal T, Reuter G. Recombinogenic effects of suppressors of position-effect variegation in Drosophila. Genetics. 2002;160:609-21 pubmed
    ..Our data suggest that crossing-over suppression by heterochromatin is controlled at chromatin structure as well as illustrate the possible effects of heterochromatin on total crossing-over frequencies in the genome. ..
  33. de Wit E, Greil F, van Steensel B. Genome-wide HP1 binding in Drosophila: developmental plasticity and genomic targeting signals. Genome Res. 2005;15:1265-73 pubmed
    ..These results provide insights into the mechanisms of HP1 targeting in the natural genomic context. ..
  34. Seum C, Reo E, Peng H, Rauscher F, Spierer P, Bontron S. Drosophila SETDB1 is required for chromosome 4 silencing. PLoS Genet. 2007;3:e76 pubmed
    ..This study defines DmSETDB1 as a H3K9 methyltransferase that specifically targets euchromatin and the autosomal Chromosome 4 and shows that it is an essential factor for Chromosome 4 silencing...
  35. Brower Toland B, Riddle N, Jiang H, Huisinga K, Elgin S. Multiple SET methyltransferases are required to maintain normal heterochromatin domains in the genome of Drosophila melanogaster. Genetics. 2009;181:1303-19 pubmed publisher
    ..In addition, the genetic interactions between dSETDB1 and Su(var)3-9 mutations emphasize the importance of maintaining the activities of these histone methyltransferases in balance for normal genome function. ..
  36. Peng J, Karpen G. Heterochromatic genome stability requires regulators of histone H3 K9 methylation. PLoS Genet. 2009;5:e1000435 pubmed publisher
    ..Similar effects of lower magnitude were observed in animals that lack the RNA interference pathway component Dcr2. These results suggest that the H3K9 methylation and RNAi pathways ensure heterochromatin stability. ..
  37. Wustmann G, Szidonya J, Taubert H, Reuter G. The genetics of position-effect variegation modifying loci in Drosophila melanogaster. Mol Gen Genet. 1989;217:520-7 pubmed
    ..Most of these loci proved not to display significant triplo-effects (35). The group of haplo-abnormal loci with a triplo-effect may represent genes which play an important role in heterochromatin packaging. ..
  38. Andreyeva E, Kolesnikova T, Demakova O, Mendez Lago M, Pokholkova G, Belyaeva E, et al. High-resolution analysis of Drosophila heterochromatin organization using SuUR Su(var)3-9 double mutants. Proc Natl Acad Sci U S A. 2007;104:12819-24 pubmed
    ..This region is enriched in H3diMetK9 and H4triMetK20 but is devoid of other proteins analyzed. ..
  39. Fauvarque M, Dura J. polyhomeotic regulatory sequences induce developmental regulator-dependent variegation and targeted P-element insertions in Drosophila. Genes Dev. 1993;7:1508-20 pubmed
    ..We suggest that a multimeric complex containing PH and PC proteins, at a minimum, causes a local and clonally inherited heterochromatinization, which maintains the repressed state of transcription of the homeotic genes. ..
  40. Swaminathan J, Baxter E, Corces V. The role of histone H2Av variant replacement and histone H4 acetylation in the establishment of Drosophila heterochromatin. Genes Dev. 2005;19:65-76 pubmed
  41. Lu X, Wontakal S, Emelyanov A, Morcillo P, Konev A, Fyodorov D, et al. Linker histone H1 is essential for Drosophila development, the establishment of pericentric heterochromatin, and a normal polytene chromosome structure. Genes Dev. 2009;23:452-65 pubmed publisher
    ..Thus, linker histone H1 is essential in Drosophila and plays a fundamental role in the architecture and activity of chromosomes in vivo. ..
  42. Greil F, de Wit E, Bussemaker H, van Steensel B. HP1 controls genomic targeting of four novel heterochromatin proteins in Drosophila. EMBO J. 2007;26:741-51 pubmed
    ..These results indicate that HP1 acts as a docking platform for several mediator proteins that contribute to heterochromatin function. ..
  43. Paredes S, Maggert K. Ribosomal DNA contributes to global chromatin regulation. Proc Natl Acad Sci U S A. 2009;106:17829-34 pubmed publisher
    ..We propose that the rDNA contributes to a balance between heterochromatin and euchromatin in the nucleus, and alterations in rDNA--induced or natural--affect this balance. ..
  44. Hwang K, Eissenberg J, Worman H. Transcriptional repression of euchromatic genes by Drosophila heterochromatin protein 1 and histone modifiers. Proc Natl Acad Sci U S A. 2001;98:11423-7 pubmed
    ..These data provide genetic evidence that an HP1-family protein represses the expression of euchromatic genes in a metazoan, and that histone modifiers cooperate with HP1 in euchromatic gene repression. ..
  45. Fagegaltier D, Bouge A, Berry B, Poisot E, Sismeiro O, Coppee J, et al. The endogenous siRNA pathway is involved in heterochromatin formation in Drosophila. Proc Natl Acad Sci U S A. 2009;106:21258-63 pubmed publisher
    ..Similar effects were observed in dcr2, r2d2, and ago2 mutants. Our findings provide evidence that a nuclear pool of TE-derived endo-siRNAs is involved in heterochromatin formation in somatic tissues in Drosophila. ..
  46. Aagaard L, Laible G, Selenko P, Schmid M, Dorn R, Schotta G, et al. Functional mammalian homologues of the Drosophila PEV-modifier Su(var)3-9 encode centromere-associated proteins which complex with the heterochromatin component M31. EMBO J. 1999;18:1923-38 pubmed
    ..Based on sequence information from the SET domain, we have isolated human (SUV39H1) and mouse (Suv39h1) homologues of the dominant Drosophila modifier of position-effect-variegation (PEV) Su(var)3-..
  47. Delattre M, Spierer A, Jaquet Y, Spierer P. Increased expression of Drosophila Su(var)3-7 triggers Su(var)3-9-dependent heterochromatin formation. J Cell Sci. 2004;117:6239-47 pubmed
    ..Moreover they show that Su(var)3-7 expression is sufficient to induce Su(var)3-9-dependent ectopic heterochromatinisation and suggest a functional link between Su(var)3-7 and the histone-methyltransferase Su(var)3-9. ..
  48. Kharchenko P, Alekseyenko A, Schwartz Y, Minoda A, Riddle N, Ernst J, et al. Comprehensive analysis of the chromatin landscape in Drosophila melanogaster. Nature. 2011;471:480-5 pubmed publisher
    ..This systematic profiling and integrative analysis of chromatin signatures provides insights into how genomic elements are regulated, and will serve as a resource for future experimental investigations of genome structure and function. ..
  49. Csink A, Henikoff S. Genetic modification of heterochromatic association and nuclear organization in Drosophila. Nature. 1996;381:529-31 pubmed
    ..This suggests that heterochromatin and proteins involved in its formation provide a structural framework for the interphase nucleus. ..
  50. Gracheva E, Dus M, Elgin S. Drosophila RISC component VIG and its homolog Vig2 impact heterochromatin formation. PLoS ONE. 2009;4:e6182 pubmed publisher
  51. Moshkovich N, Lei E. HP1 recruitment in the absence of argonaute proteins in Drosophila. PLoS Genet. 2010;6:e1000880 pubmed publisher
    ..Taken together, these results indicate that heterochromatin forms independently of endo-siRNA and piRNA pathways. ..
  52. Xu N, Emelyanov A, Fyodorov D, Skoultchi A. Drosophila linker histone H1 coordinates STAT-dependent organization of heterochromatin and suppresses tumorigenesis caused by hyperactive JAK-STAT signaling. Epigenetics Chromatin. 2014;7:16 pubmed publisher
    ..H1 serves as a molecular reservoir for STAT92E in chromatin, enabling H1 to act as a tumor suppressor and oppose an oncogenic mutation in the JAK-STAT signaling pathway. ..
  53. Pickersgill H, Kalverda B, de Wit E, Talhout W, Fornerod M, van Steensel B. Characterization of the Drosophila melanogaster genome at the nuclear lamina. Nat Genet. 2006;38:1005-14 pubmed
    ..This genome-wide analysis gives clear insight into the nature and dynamic behavior of the genome at the nuclear lamina, and implies that intergenic DNA functions in the global organization of chromatin in the nucleus. ..
  54. Deng H, Cai W, Wang C, Lerach S, Delattre M, Girton J, et al. JIL-1 and Su(var)3-7 interact genetically and counteract each other's effect on position-effect variegation in Drosophila. Genetics. 2010;185:1183-92 pubmed publisher
    ..These observations suggest a model where Su(var)3-7 functions as an effector downstream of Su(var)3-9 and H3K9 dimethylation in heterochromatic spreading and gene silencing that is normally counteracted by JIL-1 kinase activity. ..
  55. Demakova O, Pokholkova G, Kolesnikova T, Demakov S, Andreyeva E, Belyaeva E, et al. The SU(VAR)3-9/HP1 complex differentially regulates the compaction state and degree of underreplication of X chromosome pericentric heterochromatin in Drosophila melanogaster. Genetics. 2007;175:609-20 pubmed
    ..Focusing on study of X chromosome heterochromatin, we demonstrate that loss of either SU(VAR)3-9 histone methyltransferase activity or HP1 protein differentially affects the compaction of different pericentric regions...
  56. Scaria G, Ramsay G, Katzen A. Two components of the Myb complex, DMyb and Mip130, are specifically associated with euchromatin and degraded during prometaphase throughout development. Mech Dev. 2008;125:646-61 pubmed publisher
    ..We conclude that cell cycle specific degradation of DMyb and Mip130 is likely to be utilized as a key regulatory mechanism in down-regulating their levels and the activity of the Myb complex. ..
  57. 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. ..
  58. Bai X, Larschan E, Kwon S, Badenhorst P, Kuroda M. Regional control of chromatin organization by noncoding roX RNAs and the NURF remodeling complex in Drosophila melanogaster. Genetics. 2007;176:1491-9 pubmed
    ..Together, these results demonstrate the importance of a local balance between modifying activities that promote and antagonize chromatin compaction within defined chromatin domains in higher organisms. ..