Gene Symbol: E(bx)
Description: Enhancer of bithorax
Alias: CG10894, CG17135, CG32346, CG32478, CG7022, Dmel\CG32346, E(BX), E(Bx), Ebx, En-bx, NURF, NURF 215, NURF-215, NURF-301, NURF215, NURF301, NuRF301, Nurf 301, Nurf-215, Nurf301, dNURF, dNURF301, e(bx), l(3)122, l(3)ry122, nurf301, p215, p301, enhancer of bithorax, CG32346-PA, CG32346-PB, CG32346-PC, CG32346-PE, CG32346-PF, CG32346-PG, CG32346-PH, CG32346-PI, CG32346-PJ, E(bx)-PA, E(bx)-PB, E(bx)-PC, E(bx)-PE, E(bx)-PF, E(bx)-PG, E(bx)-PH, E(bx)-PI, E(bx)-PJ, nucleosome remodeling factor, nucleosome remodeling factor - 215kD
Species: fruit fly

Top Publications

  1. 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
    ..Here we use a sensitized system to examine the function of roX genes on the X chromosome. In mutants that lack the NURF nucleosome remodeling complex, the male polytene X chromosome is severely distorted, appearing decondensed...
  2. Kirilly D, Wong J, Lim E, Wang Y, Zhang H, Wang C, et al. Intrinsic epigenetic factors cooperate with the steroid hormone ecdysone to govern dendrite pruning in Drosophila. Neuron. 2011;72:86-100 pubmed publisher
    ..Thus, specific intrinsic epigenetic factors cooperate with steroid hormones to activate selective transcriptional programs, thereby initiating neuronal remodeling. ..
  3. Carré C, Ciurciu A, Komonyi O, Jacquier C, Fagegaltier D, Pidoux J, et al. The Drosophila NURF remodelling and the ATAC histone acetylase complexes functionally interact and are required for global chromosome organization. EMBO Rep. 2008;9:187-92 pubmed
    ..Ada2a induce a decondensation of the male X chromosome, similar to that induced by mutations in the Iswi and Nurf301 subunits of the NURF nucleosome remodelling complex...
  4. Ferreira R, Eberharter A, Bonaldi T, Chioda M, Imhof A, Becker P. Site-specific acetylation of ISWI by GCN5. BMC Mol Biol. 2007;8:73 pubmed
    ..The acetylated form of ISWI represents a minor species presumably associated with the nucleosome remodelling factor NURF. Acetylation of histone H3 and ISWI by GCN5 is explained by the sequence similarity between the histone and ISWI ..
  5. Stephens G, Xiao H, Lankenau D, Wu C, Elgin S. Heterochromatin protein 2 interacts with Nap-1 and NURF: a link between heterochromatin-induced gene silencing and the chromatin remodeling machinery in Drosophila. Biochemistry. 2006;45:14990-9 pubmed
    ..Upon probing fractions in which HP2 and Nap-1 are both present, we find that the nucleosome remodeling factor (NURF), an ISWI-dependent chromatin remodeling complex, is also present...
  6. Wysocka J, Swigut T, Xiao H, Milne T, Kwon S, Landry J, et al. A PHD finger of NURF couples histone H3 lysine 4 trimethylation with chromatin remodelling. Nature. 2006;442:86-90 pubmed
  7. Varga Weisz P, Becker P. Regulation of higher-order chromatin structures by nucleosome-remodelling factors. Curr Opin Genet Dev. 2006;16:151-6 pubmed
    ..Recent observations support roles for nucleosome-remodelling factors at the supra-nucleosomal level. ..
  8. Kang J, Hamiche A, Wu C. GAL4 directs nucleosome sliding induced by NURF. EMBO J. 2002;21:1406-13 pubmed
    The Drosophila nucleosome remodeling factor (NURF) is an imitation switch (ISWI)-containing chromatin remodeling complex that can catalyze nucleosome repositioning at promoter regions to regulate access by the transcription machinery...
  9. Tsukiyama T, Wu C. Purification and properties of an ATP-dependent nucleosome remodeling factor. Cell. 1995;83:1011-20 pubmed
    We report the purification of an ATP-dependent nucleosome remodeling factor (NURF) from Drosophila embryo extracts...

More Information


  1. Feller C, Prestel M, Hartmann H, Straub T, Söding J, Becker P. The MOF-containing NSL complex associates globally with housekeeping genes, but activates only a defined subset. Nucleic Acids Res. 2012;40:1509-22 pubmed publisher
    ..and in particular housekeeping genes, at which it colocalizes with the chromatin remodeler NURF (nucleosome remodeling factor) and the histone methyltransferase Trithorax...
  2. Kugler S, Nagel A. A novel Pzg-NURF complex regulates Notch target gene activity. Mol Biol Cell. 2010;21:3443-8 pubmed publisher
    ..Loss of any NURF component including the NURF-specific subunit Nurf 301 impedes binding of Putzig to Notch target genes, suggesting that NURF recruits Putzig to these sites...
  3. Li M, Belozerov V, Cai H. Modulation of chromatin boundary activities by nucleosome-remodeling activities in Drosophila melanogaster. Mol Cell Biol. 2010;30:1067-76 pubmed publisher
    ..We found that knockdown of NURF301 and ISWI, key components of the nucleosome-remodeling factor (NURF), synergistically disrupted the enhancer-..
  4. Corona D, Tamkun J. Multiple roles for ISWI in transcription, chromosome organization and DNA replication. Biochim Biophys Acta. 2004;1677:113-9 pubmed
    ..Here we discuss recent progress toward understanding the biological functions of ISWI, with an emphasis on its roles in transcription, chromosome organization and DNA replication. ..
    ..in vitro requires the concerted action of the GAGA transcription factor and NURF, an ATP-dependent nucleosome remodeling factor. NURF is composed of four subunits and is biochemically distinct from the SWI2/SNF2 multiprotein ..
  6. Xiao H, Sandaltzopoulos R, Wang H, Hamiche A, Ranallo R, Lee K, et al. Dual functions of largest NURF subunit NURF301 in nucleosome sliding and transcription factor interactions. Mol Cell. 2001;8:531-43 pubmed
    ..We have cloned cDNA encoding the largest NURF subunit, revealing a 301 kDa polypeptide (NURF301) that shares structural motifs with ACF1...
  7. 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. ..
  8. Petesch S, Lis J. Rapid, transcription-independent loss of nucleosomes over a large chromatin domain at Hsp70 loci. Cell. 2008;134:74-84 pubmed publisher
  9. Hamiche A, Sandaltzopoulos R, Gdula D, Wu C. ATP-dependent histone octamer sliding mediated by the chromatin remodeling complex NURF. Cell. 1999;97:833-42 pubmed
    Drosophila NURF is an ATP-dependent chromatin remodeling complex that contains ISWI, a member of the SWI2/SNF2 family of ATPases...
  10. Badenhorst P, Voas M, Rebay I, Wu C. Biological functions of the ISWI chromatin remodeling complex NURF. Genes Dev. 2002;16:3186-98 pubmed
    The nucleosome remodeling factor (NURF) is one of several ISWI-containing protein complexes that catalyze ATP-dependent nucleosome sliding and facilitate transcription of chromatin in vitro...
  11. Kwon S, Xiao H, Wu C, Badenhorst P. Alternative splicing of NURF301 generates distinct NURF chromatin remodeling complexes with altered modified histone binding specificities. PLoS Genet. 2009;5:e1000574 pubmed publisher
    ..By sliding nucleosomes, NURF can alter chromatin structure and regulate transcription. NURF301/BPTF is the only NURF-specific subunit of NURF and is instrumental in recruiting the complex to target genes...
  12. Song H, Spichiger Haeusermann C, Basler K. The ISWI-containing NURF complex regulates the output of the canonical Wingless pathway. EMBO Rep. 2009;10:1140-6 pubmed publisher
  13. Alkhatib S, Landry J. The nucleosome remodeling factor. FEBS Lett. 2011;585:3197-207 pubmed publisher
    An essential component of the chromatin remodeling machinery is NURF (Nucleosome Remodeling Factor), the founding member of the ISWI family of chromatin remodeling complexes...
  14. Ables E, Drummond Barbosa D. The steroid hormone ecdysone functions with intrinsic chromatin remodeling factors to control female germline stem cells in Drosophila. Cell Stem Cell. 2010;7:581-92 pubmed publisher
    ..the chromatin remodeling factors ISWI, an intrinsic epigenetic factor required for GSC fate and activity, and Nurf301, the largest subunit of the ISWI-containing NURF chromatin remodeling complex...
  15. Lucchesi J, Kuroda M. Dosage compensation in Drosophila. Cold Spring Harb Perspect Biol. 2015;7: pubmed publisher
    ..Interestingly, two noncoding RNAs are key for MSL assembly and spreading to active genes along the length of the X chromosome. ..
  16. Nakayama T, Shimojima T, Hirose S. The PBAP remodeling complex is required for histone H3.3 replacement at chromatin boundaries and for boundary functions. Development. 2012;139:4582-90 pubmed publisher
    ..3 replacement to execute boundary functions. Our results provide new insight into the function of the trithorax group during development. ..
  17. Fry C, Peterson C. Chromatin remodeling enzymes: who's on first?. Curr Biol. 2001;11:R185-97 pubmed
    ..Here we review recent developments concerning the role of chromatin remodeling enzymes in gene regulation, and propose several models to explain how different chromatin remodeling activities can be functionally coupled. ..
  18. Tsai S, Chang Y, Swamy K, Chiang R, Huang D. GAGA factor, a positive regulator of global gene expression, modulates transcriptional pausing and organization of upstream nucleosomes. Epigenetics Chromatin. 2016;9:32 pubmed publisher
  19. Tyler J, Kadonaga J. The "dark side" of chromatin remodeling: repressive effects on transcription. Cell. 1999;99:443-6 pubmed
  20. Kugler S, Gehring E, Wallkamm V, Krüger V, Nagel A. The Putzig-NURF nucleosome remodeling complex is required for ecdysone receptor signaling and innate immunity in Drosophila melanogaster. Genetics. 2011;188:127-39 pubmed publisher
    ..This function of Pzg entails an association with the nucleosome remodeling factor complex NURF, which directly binds the ecdysone receptor EcR and coregulates targets of the EcR via the ..
  21. Peterson C. Multiple SWItches to turn on chromatin?. Curr Opin Genet Dev. 1996;6:171-5 pubmed
  22. Soruco M, Larschan E. A new player in X identification: the CLAMP protein is a key factor in Drosophila dosage compensation. Chromosome Res. 2014;22:505-15 pubmed publisher
    ..Understanding how the newly identified CLAMP protein is involved in X targeting in the fly could provide key insights into how the X chromosome is initially identified across species. ..
  23. Burgio G, La Rocca G, Sala A, Arancio W, Di Gesù D, Collesano M, et al. Genetic identification of a network of factors that functionally interact with the nucleosome remodeling ATPase ISWI. PLoS Genet. 2008;4:e1000089 pubmed publisher
    ..Consistent with these findings, the acetylation of histone H4 is altered when ISWI activity is perturbed in vivo. These findings suggest that ISWI associates with the Sin3A/Rpd3 complex to support its function in vivo. ..
  24. Lusser A, Kadonaga J. Chromatin remodeling by ATP-dependent molecular machines. Bioessays. 2003;25:1192-200 pubmed
    ..In this review, we provide an overview of chromatin-remodeling enzymes and discuss two possible mechanisms by which these factors might act to reorganize nucleosome structure. ..
  25. Di Croce L, Koop R, Beato M. Rapid purification of intact minichromosomes over a glycerol cushion. Nucleic Acids Res. 1999;27:e11 pubmed
    ..As centrifugation through glycerol yields chromatin free of ATP and several characterized chromatin remodeling complexes, this method should be useful for structural and functional studies in vitro. ..
  26. Jacobson S, Pillus L. Modifying chromatin and concepts of cancer. Curr Opin Genet Dev. 1999;9:175-84 pubmed
    ..Understanding the function of these factors will be key to defining mechanisms of transcriptional regulation and understanding cancers linked to aberrant chromatin modification. ..
  27. Belles X, Piulachs M. Ecdysone signalling and ovarian development in insects: from stem cells to ovarian follicle formation. Biochim Biophys Acta. 2015;1849:181-6 pubmed publisher
    ..This article is part of a Special Issue entitled: Nuclear receptors in animal development. ..
  28. Koe C, Li S, Rossi F, Wong J, Wang Y, Zhang Z, et al. The Brm-HDAC3-Erm repressor complex suppresses dedifferentiation in Drosophila type II neuroblast lineages. elife. 2014;3:e01906 pubmed publisher
    ..Thus, the Brm-HDAC3-Erm repressor complex suppresses dedifferentiation of INPs back into type II neuroblasts. DOI: http://dx.doi.org/10.7554/eLife.01906.001. ..
  29. Acharya P, Negre N, Johnston J, Wei Y, White K, Henry R, et al. Evidence for autoregulation and cell signaling pathway regulation from genome-wide binding of the Drosophila retinoblastoma protein. G3 (Bethesda). 2012;2:1459-72 pubmed publisher
    ..Many of these targeted genes are bound by Rbf1 homologs in human cells, indicating that a conserved role of RB proteins may be to adjust the set point of interlinked signaling networks essential for growth and development. ..
    ..We therefore suggest that nucleosome positioning by RSSs provides a previously unanticipated level of protection and regulation of V(D)J recombination. ..
  31. Reinberg D, Orphanides G, Ebright R, Akoulitchev S, Carcamo J, Cho H, et al. The RNA polymerase II general transcription factors: past, present, and future. Cold Spring Harb Symp Quant Biol. 1998;63:83-103 pubmed
  32. Schwanbeck R, Xiao H, Wu C. Spatial contacts and nucleosome step movements induced by the NURF chromatin remodeling complex. J Biol Chem. 2004;279:39933-41 pubmed
    The nucleosome remodeling factor NURF is a four-subunit, ISWI-containing chromatin remodeling complex that catalyzes nucleosome sliding in an ATP-dependent fashion, thereby modulating the accessibility of the DNA...
  33. Spradling A, Rubin G. Transposition of cloned P elements into Drosophila germ line chromosomes. Science. 1982;218:341-7 pubmed
    ..Thus P elements can efficiently and selectively transpose from extrachromosomal DNA to the DNA of germ line chromosomes in Drosophila embryos. These observations provide the basis for efficient DNA-mediated gene transfer in Drosophila. ..
  35. Cherry C, MATUNIS E. Epigenetic regulation of stem cell maintenance in the Drosophila testis via the nucleosome-remodeling factor NURF. Cell Stem Cell. 2010;6:557-67 pubmed publisher
    ..Here we show that epigenetic regulation via the nucleosome-remodeling factor (NURF) complex ensures GSC and CPC maintenance by positively regulating JAK-STAT signaling, thereby preventing premature ..
  36. Follmer N, Wani A, Francis N. A polycomb group protein is retained at specific sites on chromatin in mitosis. PLoS Genet. 2012;8:e1003135 pubmed publisher
  37. Bohla D, Herold M, Panzer I, Buxa M, Ali T, Demmers J, et al. A functional insulator screen identifies NURF and dREAM components to be required for enhancer-blocking. PLoS ONE. 2014;9:e107765 pubmed publisher
  38. Shen X, Xiao H, Ranallo R, Wu W, Wu C. Modulation of ATP-dependent chromatin-remodeling complexes by inositol polyphosphates. Science. 2003;299:112-4 pubmed
    ..Inositol hexakisphosphate (IP6) inhibits nucleosome mobilization by NURF, ISW2, and INO80 complexes...
  39. 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...
  40. Neely K, Workman J. The complexity of chromatin remodeling and its links to cancer. Biochim Biophys Acta. 2002;1603:19-29 pubmed
    ..Their subunit composition and brief overview of their functional properties will be discussed. Also, current links between human chromatin-remodeling complexes and cell growth and cancer will be presented. ..
  41. Achary B, Campbell K, Co I, Gilmour D. RNAi screen in Drosophila larvae identifies histone deacetylase 3 as a positive regulator of the hsp70 heat shock gene expression during heat shock. Biochim Biophys Acta. 2014;1839:355-63 pubmed publisher
    ..We validated the screen by showing that the depletion of HSF, CycT, Cdk9, Nurf 301, or ELL prevented the full induction of hsp70 by heat shock...
  42. Li G, Zhou L. Genome-wide identification of chromatin transitional regions reveals diverse mechanisms defining the boundary of facultative heterochromatin. PLoS ONE. 2013;8:e67156 pubmed publisher
    ..However, boundary-associated insulator binding sites are distinctively flanked by nucleosome destabilizing sequences, which correlates with significantly decreased nucleosome density and increased binding of co-factors. ..
  43. Boyer L, Logie C, Bonte E, Becker P, Wade P, Wolffe A, et al. Functional delineation of three groups of the ATP-dependent family of chromatin remodeling enzymes. J Biol Chem. 2000;275:18864-70 pubmed
    ..We compare six different remodeling complexes: ySWI/SNF, yRSC, hSWI/SNF, xMi-2, dCHRAC, and dNURF. We find that each complex uses similar amounts of ATP to remodel nucleosomal arrays at nearly identical rates...
  44. Demakov S, Vatolina T, Babenko V, Semeshin V, Belyaeva E, Zhimulev I. Protein composition of interband regions in polytene and cell line chromosomes of Drosophila melanogaster. BMC Genomics. 2011;12:566 pubmed publisher
  45. Guschin D, Wolffe A. SWItched-on mobility. Curr Biol. 1999;9:R742-6 pubmed
    Recent studies have shown that two nucleosome-remodeling complexes, NURF and CHRAC, open chromatin for transcription and replication by using their common catalytic subunit, the nucleosomal ATPase ISWI, to increase the mobility of ..
  46. Kingston R, Bunker C, Imbalzano A. Repression and activation by multiprotein complexes that alter chromatin structure. Genes Dev. 1996;10:905-20 pubmed
    ..This implies a regulation of the activity of these complexes in response to developmental cues and further implies that the work to fully understand these complexes will occupy a generation of scientists. ..
  47. Shimojima T, Okada M, Nakayama T, Ueda H, Okawa K, Iwamatsu A, et al. Drosophila FACT contributes to Hox gene expression through physical and functional interactions with GAGA factor. Genes Dev. 2003;17:1605-16 pubmed
    ..These data illustrate a crucial role of FACT in the modulation of chromatin structure for the regulation of gene expression. ..
  48. Peterson C. ATP-dependent chromatin remodeling: going mobile. FEBS Lett. 2000;476:68-72 pubmed
    ..Here the biochemical properties of these enzymes are reviewed and the manner in which ATP-driven nucleosome movements might account for many of these diverse activities is discussed. ..
  49. Kessler R, Hausmann G, Basler K. The PHD domain is required to link Drosophila Pygopus to Legless/beta-catenin and not to histone H3. Mech Dev. 2009;126:752-9 pubmed publisher
    ..Our results reveal an interesting evolutionary dichotomy in Pygo structure-function, as well as evidence underpinning the chain of adaptors model. ..
  50. Wu C, Tsukiyama T, Gdula D, Georgel P, Martinez Balbas M, Mizuguchi G, et al. ATP-dependent remodeling of chromatin. Cold Spring Harb Symp Quant Biol. 1998;63:525-34 pubmed
  51. Englund C, Birve A, Falileeva L, Grabbe C, Palmer R. Miple1 and miple2 encode a family of MK/PTN homologues in Drosophila melanogaster. Dev Genes Evol. 2006;216:10-8 pubmed publisher
    ..The identification of homologues of the MK/PTN family in this genetically tractable model organism should allow an analysis of their function during complex developmental processes...
  52. Jones K, Kadonaga J. Exploring the transcription-chromatin interface. Genes Dev. 2000;14:1992-6 pubmed
  53. Shi S, Calhoun H, Xia F, Li J, Le L, Li W. JAK signaling globally counteracts heterochromatic gene silencing. Nat Genet. 2006;38:1071-6 pubmed
    ..These results demonstrate that the JAK/STAT pathway regulates cellular epigenetic status and that globally disrupting heterochromatin-mediated tumor suppression is essential for tumorigenesis induced by JAK overactivation. ..
  54. Kwon S, Grisan V, Jang B, Herbert J, Badenhorst P. Genome-Wide Mapping Targets of the Metazoan Chromatin Remodeling Factor NURF Reveals Nucleosome Remodeling at Enhancers, Core Promoters and Gene Insulators. PLoS Genet. 2016;12:e1005969 pubmed publisher
    ..Previous biochemical and genetic analysis of the specificity-subunit of Drosophila NURF (Nurf301/Enhancer of Bithorax (E(bx)) has defined NURF as a critical regulator of homeotic, heat-shock and steroid-responsive gene ..
  55. Mills A. Throwing the cancer switch: reciprocal roles of polycomb and trithorax proteins. Nat Rev Cancer. 2010;10:669-82 pubmed publisher
    ..Recent work highlights the dynamic interplay between these opposing classes of proteins, providing new avenues for understanding how these epigenetic regulators function in tumorigenesis. ..
  56. Mizuguchi G, Tsukiyama T, Wisniewski J, Wu C. Role of nucleosome remodeling factor NURF in transcriptional activation of chromatin. Mol Cell. 1997;1:141-50 pubmed
  57. Jung A, Hollmann M, Schäfer M. The fatty acid elongase NOA is necessary for viability and has a somatic role in Drosophila sperm development. J Cell Sci. 2007;120:2924-34 pubmed
    ..An RNAi construct selectively driven in cyst cells leads to male sterility, demonstrating the necessity of noa function for male germline development and the interaction of the somatic cyst cells with the developing sperm. ..