Gene Symbol: esc
Description: extra sexcombs
Alias: CG14941, Dmel\CG14941, ESC, Esc, L41867, dESC, extra sexcombs, CG14941-PA, esc-PA, extra Sex combs, extra sex comb
Species: fruit fly
Products:     esc

Top Publications

  1. Mollaaghababa R, Sipos L, Tiong S, Papoulas O, Armstrong J, Tamkun J, et al. Mutations in Drosophila heat shock cognate 4 are enhancers of Polycomb. Proc Natl Acad Sci U S A. 2001;98:3958-63 pubmed
    ..HSC4 might be needed for the proper folding of a component of the Polycomb repression complex, or it may be a functional member of that complex...
  2. Wang L, Brown J, Cao R, Zhang Y, Kassis J, Jones R. Hierarchical recruitment of polycomb group silencing complexes. Mol Cell. 2004;14:637-46 pubmed
  3. Simon J, Chiang A, Bender W. Ten different Polycomb group genes are required for spatial control of the abdA and AbdB homeotic products. Development. 1992;114:493-505 pubmed
    ..Embryos containing mutations in the genes Polycomb (Pc), extra sex combs (esc), Enhancer of zeste [E(z)], polyhomeotic (ph), Sex comb on midleg (Scm), Polycomb-like (Pcl), Sex comb ..
  4. Klymenko T, Papp B, Fischle W, Kocher T, Schelder M, Fritsch C, et al. A Polycomb group protein complex with sequence-specific DNA-binding and selective methyl-lysine-binding activities. Genes Dev. 2006;20:1110-22 pubmed
    ..We propose that PRE-tethered PhoRC selectively interacts with methylated histones in the chromatin flanking PREs to maintain a Polycomb-repressed chromatin state. ..
  5. Cao R, Wang L, Wang H, Xia L, Erdjument Bromage H, Tempst P, et al. Role of histone H3 lysine 27 methylation in Polycomb-group silencing. Science. 2002;298:1039-43 pubmed
    ..we report the purification and characterization of an EED-EZH2 complex, the human counterpart of the Drosophila ESC-E(Z) complex. We demonstrate that the complex specifically methylates nucleosomal histone H3 at lysine 27 (H3-K27)...
  6. Tie F, Furuyama T, Prasad Sinha J, Jane E, Harte P. The Drosophila Polycomb Group proteins ESC and E(Z) are present in a complex containing the histone-binding protein p55 and the histone deacetylase RPD3. Development. 2001;128:275-86 pubmed
    ..Among PcG genes, esc is unique in being critically required for establishment of PcG-mediated silencing during early embryogenesis, but ..
  7. Nekrasov M, Wild B, Muller J. Nucleosome binding and histone methyltransferase activity of Drosophila PRC2. EMBO Rep. 2005;6:348-53 pubmed
    ..E(z) exists in a multiprotein complex called Polycomb repressive complex 2 (PRC2) that also contains Su(z)12, Esc and Nurf55...
  8. Moazed D, O Farrell P. Maintenance of the engrailed expression pattern by Polycomb group genes in Drosophila. Development. 1992;116:805-10 pubmed
    ..Additionally, in contrast to the strong ectopic expression of homeotic genes observed in extra sex combs- (esc-) mutant embryos, the en expression pattern is nearly normal in esc- embryos...
  9. O Connell S, Wang L, Robert S, Jones C, Saint R, Jones R. Polycomblike PHD fingers mediate conserved interaction with enhancer of zeste protein. J Biol Chem. 2001;276:43065-73 pubmed
    ..E(Z)) is a Drosophila PcG protein that previously has been shown to bind directly to another PcG protein, Extra Sex Combs (ESC), and is present along with ESC in a 600-kDa complex in Drosophila embryos...

More Information


  1. Tolhuis B, de Wit E, Muijrers I, Teunissen H, Talhout W, van Steensel B, et al. Genome-wide profiling of PRC1 and PRC2 Polycomb chromatin binding in Drosophila melanogaster. Nat Genet. 2006;38:694-9 pubmed
    ..We used a genome-wide approach to map binding patterns of PcG proteins (Pc, esc and Sce) in Drosophila melanogaster Kc cells...
  2. Huang D, Chang Y, Yang C, Pan I, King B. pipsqueak encodes a factor essential for sequence-specific targeting of a polycomb group protein complex. Mol Cell Biol. 2002;22:6261-71 pubmed
    ..The functional relevance of psq to the silencing mechanism is strongly supported by its synergistic interactions with a subset of Pc-G that cause misexpression of homeotic genes. ..
  3. 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
    ..In the early embryo it is found in a complex that includes ESC and is recruited to Polycomb Response Elements...
  4. Tie F, Prasad Sinha J, Birve A, Rasmuson Lestander A, Harte P. A 1-megadalton ESC/E(Z) complex from Drosophila that contains polycomblike and RPD3. Mol Cell Biol. 2003;23:3352-62 pubmed
    ..The PcG proteins ESC and E(Z) are present in a prominent 600-kDa complex as well as in a number of higher-molecular-mass complexes...
  5. Muller J, Hart C, Francis N, Vargas M, Sengupta A, Wild B, et al. Histone methyltransferase activity of a Drosophila Polycomb group repressor complex. Cell. 2002;111:197-208 pubmed
    ..The Extra Sex Combs (ESC) and Enhancer of Zeste [E(Z)] proteins are partners in an essential PcG complex, but its full ..
  6. Campbell R, Sinclair D, Couling M, Brock H. Genetic interactions and dosage effects of Polycomb group genes of Drosophila. Mol Gen Genet. 1995;246:291-300 pubmed
    ..We show genetic interactions among extra sex combs (esc) and Asx, Enhancer of Pc, Pcl, Enhancer of zeste E(z), and super sex combs and reassess the idea that ..
  7. Poux S, Melfi R, Pirrotta V. Establishment of Polycomb silencing requires a transient interaction between PC and ESC. Genes Dev. 2001;15:2509-14 pubmed
    Two distinct types of Polycomb complexes have been identified in flies and in vertebrates, one containing ESC and one containing PC...
  8. Soto M, Chou T, Bender W. Comparison of germline mosaics of genes in the Polycomb group of Drosophila melanogaster. Genetics. 1995;140:231-43 pubmed
    ..Psc shows unusual effects on two of the targets, Ultrabithorax and abdominal-A. These results show that the PcG genes do not act only in a common complex or pathway; they must have some independent functions. ..
  9. Ketel C, Andersen E, Vargas M, Suh J, Strome S, Simon J. Subunit contributions to histone methyltransferase activities of fly and worm polycomb group complexes. Mol Cell Biol. 2005;25:6857-68 pubmed
    The ESC-E(Z) complex of Drosophila melanogaster Polycomb group (PcG) repressors is a histone H3 methyltransferase (HMTase)...
  10. Scheuermann J, de Ayala Alonso A, Oktaba K, Ly Hartig N, McGinty R, Fraterman S, et al. Histone H2A deubiquitinase activity of the Polycomb repressive complex PR-DUB. Nature. 2010;465:243-7 pubmed publisher
    ..Polycomb gene silencing may thus entail a dynamic balance between H2A ubiquitination by PRC1 and dRAF, and H2A deubiquitination by PR-DUB. ..
  11. Gorfinkiel N, Fanti L, Melgar T, García E, Pimpinelli S, Guerrero I, et al. The Drosophila Polycomb group gene Sex combs extra encodes the ortholog of mammalian Ring1 proteins. Mech Dev. 2004;121:449-62 pubmed
    ..We also show that Ring/Sce interacts directly with PcG proteins, as it occurs in mammals. ..
  12. Mishra R, Yamagishi T, Vasanthi D, Ohtsuka C, Kondo T. Involvement of polycomb-group genes in establishing HoxD temporal colinearity. Genesis. 2007;45:570-6 pubmed
    ..repressive activity in Drosophila depends on some members of the Polycomb-group (PcG) genes, for example, extra sex combs. We also showed direct association of these factors with the repressive fragment, both in transgenic flies ..
  13. Joshi P, Carrington E, Wang L, Ketel C, Miller E, Jones R, et al. Dominant alleles identify SET domain residues required for histone methyltransferase of Polycomb repressive complex 2. J Biol Chem. 2008;283:27757-66 pubmed publisher
    ..These results identify critical SET domain residues needed for PRC2 enzyme function, and they also emphasize functional inputs from outside the SET domain. ..
  14. Struhl G. Role of the esc+ gene product in ensuring the selective expression of segment-specific homeotic genes in Drosophila. J Embryol Exp Morphol. 1983;76:297-331 pubmed
    The product of the extra sex combs+ (esc+) gene is required during embryogenesis for the correct determination of segments in Drosophila. If this product is absent, most segments develop like the normal eighth abdominal segment...
  15. Pirrotta V. PcG complexes and chromatin silencing. Curr Opin Genet Dev. 1997;7:249-58 pubmed
    ..How such a complex interferes with transcriptional activity is not entirely clear but current evidence suggests that they compete with the chromatin structure required for the binding of activators. ..
  16. 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. ..
  17. Rai A, Vargas M, Wang L, Andersen E, Miller E, Simon J. Elements of the polycomb repressor SU(Z)12 needed for histone H3-K27 methylation, the interface with E(Z), and in vivo function. Mol Cell Biol. 2013;33:4844-56 pubmed publisher
    ..In Drosophila melanogaster, PRC2 contains four core subunits: E(Z), SU(Z)12, ESC, and NURF55. E(Z) bears a SET domain that houses the enzyme active site...
  18. 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...
  19. Di Croce L, Helin K. Transcriptional regulation by Polycomb group proteins. Nat Struct Mol Biol. 2013;20:1147-55 pubmed publisher
    ..Here we discuss our current knowledge of the PRC complexes, how they are targeted to chromatin and how the high diversity of the PcG proteins allows these complexes to influence cell identity. ..
  20. Anderson A, Karandikar U, Pepple K, Chen Z, Bergmann A, Mardon G. The enhancer of trithorax and polycomb gene Caf1/p55 is essential for cell survival and patterning in Drosophila development. Development. 2011;138:1957-66 pubmed publisher
    ..More importantly, our mutant phenotypes confirm that Caf1-mediated silencing is vital to Drosophila development. These studies underscore the importance of Caf1 and its mammalian homologs in development and disease. ..
  21. Margueron R, Justin N, Ohno K, Sharpe M, Son J, Drury W, et al. Role of the polycomb protein EED in the propagation of repressive histone marks. Nature. 2009;461:762-7 pubmed publisher
    ..These findings suggest a model for the propagation of the H3K27me3 mark that accounts for the maintenance of repressive chromatin domains and for the transmission of a histone modification from mother to daughter cells. ..
  22. Wang L, Ding L, Jones C, Jones R. Drosophila Enhancer of zeste protein interacts with dSAP18. Gene. 2002;285:119-25 pubmed
    ..These studies provide support for a model in which dSAP18 contributes to the activities of multiple protein complexes, and potentially may mediate interactions between distinct proteins and/or protein complexes. ..
  23. Grossniklaus U, Paro R. Transcriptional silencing by polycomb-group proteins. Cold Spring Harb Perspect Biol. 2014;6:a019331 pubmed publisher
    ..PcG silencing is medically relevant as it is often observed in human disorders, including cancer, and tissue regeneration, which involve the reprogramming of PcG-controlled target genes. ..
  24. Struhl G, Brower D. Early role of the esc+ gene product in the determination of segments in Drosophila. Cell. 1982;31:285-92 pubmed
    Recent studies of the esc+ gene suggest that its product acts principally in the initiation, but not the maintenance, of segmental determination...
  25. Gould A. Functions of mammalian Polycomb group and trithorax group related genes. Curr Opin Genet Dev. 1997;7:488-94 pubmed
    ..Initial indications suggest the existence of a large mammalian PcG and trxG family, with a potential to encode multiple specialised functions in cell fate and cell-cycle control. ..
  26. Kurzhals R, Tie F, Stratton C, Harte P. Drosophila ESC-like can substitute for ESC and becomes required for Polycomb silencing if ESC is absent. Dev Biol. 2008;313:293-306 pubmed
    The Drosophila esc-like gene (escl) encodes a protein very similar to ESC. Like ESC, ESCL binds directly to the E(Z) histone methyltransferase via its WD region...
  27. Saget O, Forquignon F, Santamaria P, Randsholt N. Needs and targets for the multi sex combs gene product in Drosophila melanogaster. Genetics. 1998;149:1823-38 pubmed
    ..We discuss these data in view of the numerous functions of PcG genes during development. ..
  28. Lanzuolo C, Orlando V. Memories from the polycomb group proteins. Annu Rev Genet. 2012;46:561-89 pubmed publisher
    ..In this review, we focus on the multiple modes of action of the PcG complexes and describe their biological roles. ..
  29. Stepanik V, Harte P. A mutation in the E(Z) methyltransferase that increases trimethylation of histone H3 lysine 27 and causes inappropriate silencing of active Polycomb target genes. Dev Biol. 2012;364:249-58 pubmed publisher
  30. Tie F, Furuyama T, Harte P. The Drosophila Polycomb Group proteins ESC and E(Z) bind directly to each other and co-localize at multiple chromosomal sites. Development. 1998;125:3483-96 pubmed
    The Polycomb Group gene esc encodes an evolutionarily conserved protein required for transcriptional silencing of the homeotic genes...
  31. Kang S, Kim S, Kim S, Jeon S. Temporal and spatial expression of homeotic genes is important for segment-specific neuroblast 6-4 lineage formation in Drosophila. Mol Cells. 2006;21:436-42 pubmed
    ..Our results indicate that temporal and spa-tial expression of the homeotic genes is important to determine segmental-specificity of NB6-4 daughter cells along the anterior-posterior (A/P) axis. ..
  32. Herz H, Mohan M, Garrett A, Miller C, Casto D, Zhang Y, et al. Polycomb repressive complex 2-dependent and -independent functions of Jarid2 in transcriptional regulation in Drosophila. Mol Cell Biol. 2012;32:1683-93 pubmed publisher
    ..Identification of Jarid2 as a conserved PRC2 interactor in flies provides an opportunity to begin to probe some of its novel functions in Drosophila development. ..
  33. Sultana H, Verma S, Mishra R. A BEAF dependent chromatin domain boundary separates myoglianin and eyeless genes of Drosophila melanogaster. Nucleic Acids Res. 2011;39:3543-57 pubmed publisher
    ..We also report that BEAF and GAF are the two major proteins responsible for the ME boundary function. Our studies demonstrate a rational approach to search for potential boundaries in genomic regions that are well annotated. ..
  34. Gould A, Lai R, Green M, White R. Blocking cell division does not remove the requirement for Polycomb function in Drosophila embryogenesis. Development. 1990;110:1319-25 pubmed
    ..Thus, even in the absence of DNA replication, Pc is required to maintain spatially restricted patterns of homeotic gene expression. The role of the Pc gene product in the stable repression of homeotic gene transcription is discussed. ..
  35. Muller J. Transcriptional silencing by the Polycomb protein in Drosophila embryos. EMBO J. 1995;14:1209-20 pubmed
    ..The extent of DNA over which the Pc-G machinery acts is quite selective, as silencing established on one enhancer does not necessarily 'spread' to a juxtaposed synthetic enhancer...
  36. Nekrasov M, Klymenko T, Fraterman S, Papp B, Oktaba K, Kocher T, et al. Pcl-PRC2 is needed to generate high levels of H3-K27 trimethylation at Polycomb target genes. EMBO J. 2007;26:4078-88 pubmed
    ..Pcl-PRC2 is thus the HMTase that generates the high levels of H3-K27 trimethylation in Polycomb target genes that are needed to maintain a Polycomb-repressed chromatin state. ..
  37. Steffen P, Ringrose L. What are memories made of? How Polycomb and Trithorax proteins mediate epigenetic memory. Nat Rev Mol Cell Biol. 2014;15:340-56 pubmed publisher
    ..Several recent studies have advanced our understanding of the molecular mechanisms of this epigenetic memory during DNA replication and mitosis. ..
  38. Dietz K, Di Stefano L, Maher R, Zhu H, MacDonald M, Gusella J, et al. The Drosophila Huntington's disease gene ortholog dhtt influences chromatin regulation during development. Hum Mol Genet. 2015;24:330-45 pubmed publisher
    ..These findings suggest that a fundamental aspect of huntingtin function in heterochromatin/euchromatin organization is evolutionarily conserved across phyla. ..
  39. Frei E, Bopp D, Burri M, Baumgartner S, Edstrom J, Noll M. Isolation and structural analysis of the extra sex combs gene of Drosophila. Cold Spring Harb Symp Quant Biol. 1985;50:127-34 pubmed
  40. Calvo Martín J, Librado P, Aguadé M, Papaceit M, Segarra C. Adaptive selection and coevolution at the proteins of the Polycomb repressive complexes in Drosophila. Heredity (Edinb). 2016;116:213-23 pubmed publisher
    ..These clusters include well-established binary interactions among PcG proteins as well as new putative interactions. ..
  41. Wutz A. RNAs templating chromatin structure for dosage compensation in animals. Bioessays. 2003;25:434-42 pubmed
    ..They do not appear to act in a sequence-specific manner but might provide scaffolds for co-operative binding of chromatin-associated complexes that enable spreading of chromatin modifications. ..
  42. Yuan W, Wu T, Fu H, Dai C, Wu H, Liu N, et al. Dense chromatin activates Polycomb repressive complex 2 to regulate H3 lysine 27 methylation. Science. 2012;337:971-5 pubmed publisher
    ..We propose that PRC2 can sense the chromatin environment to exert its role in the maintenance of transcriptional states. ..
  43. Furuyama T, Tie F, Harte P. Polycomb group proteins ESC and E(Z) are present in multiple distinct complexes that undergo dynamic changes during development. Genesis. 2003;35:114-24 pubmed
    ..distinct Polycomb Group complexes have been identified, a 2-MDa PRC1 complex and a 600-kDa complex containing the ESC and E(Z) proteins together with the histone deacetylase RPD3 and the histone-binding protein p55...
  44. Pirrotta V. Polycombing the genome: PcG, trxG, and chromatin silencing. Cell. 1998;93:333-6 pubmed
  45. Holmes A, Weedmark K, Gloor G. Mutations in the extra sex combs and Enhancer of Polycomb genes increase homologous recombination in somatic cells of Drosophila melanogaster. Genetics. 2006;172:2367-77 pubmed
    We found that heterozygous mutant alleles of E(Pc) and esc increased homologous recombination from an allelic template in somatic cells in a P-element-induced double-strand break repair assay...
  46. Tie F, Stratton C, Kurzhals R, Harte P. The N terminus of Drosophila ESC binds directly to histone H3 and is required for E(Z)-dependent trimethylation of H3 lysine 27. Mol Cell Biol. 2007;27:2014-26 pubmed
    ..The 600-kDa E(Z)/ESC complex, also known as Polycomb repressive complex 2 (PRC2), specifically methylates histone H3 lysine 27 (H3 K27) ..
  47. McKeon J, Slade E, Sinclair D, Cheng N, Couling M, Brock H. Mutations in some Polycomb group genes of Drosophila interfere with regulation of segmentation genes. Mol Gen Genet. 1994;244:474-83 pubmed
    ..Surprisingly, flies carrying duplications of extra sex combs (esc), that were heterozygous for mutations of even-skipped (eve), were extremely subvital...
  48. Sathe S, Harte P. The extra sex combs protein is highly conserved between Drosophila virilis and Drosophila melanogaster. Mech Dev. 1995;52:225-32 pubmed
    b>Extra sex combs (esc) is one of the Polycomb Group genes, whose products are required for long term maintenance of the spatially restricted domains of homeotic gene expression initially established by the products of the segmentation ..
  49. Simon J, Bornemann D, Lunde K, Schwartz C. The extra sex combs product contains WD40 repeats and its time of action implies a role distinct from other Polycomb group products. Mech Dev. 1995;53:197-208 pubmed
    The extra sex combs (esc) gene product is a transcriptional repressor of homeotic genes...
  50. Roseman R, Morgan K, Mallin D, Roberson R, Parnell T, Bornemann D, et al. Long-range repression by multiple polycomb group (PcG) proteins targeted by fusion to a defined DNA-binding domain in Drosophila. Genetics. 2001;158:291-307 pubmed
    ..Three PcG fusion proteins, ZnF-PC, ZnF-SCM, and ZnF-ESC, were studied, since biochemical analyses place these PcG proteins in distinct complexes...
  51. Salvaing J, Lopez A, Boivin A, Deutsch J, Peronnet F. The Drosophila Corto protein interacts with Polycomb-group proteins and the GAGA factor. Nucleic Acids Res. 2003;31:2873-82 pubmed
    In Drosophila, PcG complexes provide heritable transcriptional silencing of target genes. Among them, the ESC/E(Z) complex is thought to play a role in the initiation of silencing whereas other complexes such as the PRC1 complex are ..
  52. Chopra V, Hendrix D, Core L, Tsui C, Lis J, Levine M. The polycomb group mutant esc leads to augmented levels of paused Pol II in the Drosophila embryo. Mol Cell. 2011;42:837-44 pubmed publisher
    ..and GRO-Seq assays were used to determine the genome-wide distributions of Pol II, H3K27me3, and H3K4me3 in extra sex combs (esc) mutant embryos...
  53. van Lohuizen M. The trithorax-group and polycomb-group chromatin modifiers: implications for disease. Curr Opin Genet Dev. 1999;9:355-61 pubmed
    ..This is illustrated by recent mechanistic insights that link Polycomb repression with histone deacetylases and the identification of new target genes that provide a connection to cell cycle control and tumorigenesis. ..
  54. Tie F, Banerjee R, Stratton C, Prasad Sinha J, Stepanik V, Zlobin A, et al. CBP-mediated acetylation of histone H3 lysine 27 antagonizes Drosophila Polycomb silencing. Development. 2009;136:3131-41 pubmed publisher
    ..We propose that TRX-dependent acetylation of H3K27 by CBP prevents H3K27me3 at Polycomb target genes and constitutes a key part of the molecular mechanism by which TRX antagonizes or prevents Polycomb silencing. ..
  55. Cheng N, Sinclair D, Campbell R, Brock H. Interactions of polyhomeotic with Polycomb group genes of Drosophila melanogaster. Genetics. 1994;138:1151-62 pubmed
    ..ph2 was not lethal with extra sex combs, E(z), super sex combs (sxc) or l(4)102EFc heterozygotes, but did cause earlier lethality in embryos ..
  56. Atchison L, Ghias A, Wilkinson F, Bonini N, Atchison M. Transcription factor YY1 functions as a PcG protein in vivo. EMBO J. 2003;22:1347-58 pubmed
    ..Finally, we found that YY1, as well as Polycomb, required the co-repressor protein CtBP for repression in vivo. These results provide a mechanism for recruitment of vertebrate PcG complexes to DNA and demonstrate new functions for YY1. ..
  57. Dorsett D, Kassis J. Checks and balances between cohesin and polycomb in gene silencing and transcription. Curr Biol. 2014;24:R535-9 pubmed publisher
  58. Dura J, Brock H, Santamaria P. Polyhomeotic: a gene of Drosophila melanogaster required for correct expression of segmental identity. Mol Gen Genet. 1985;198:213-20 pubmed interacts with three other similar mutations: Polycomb (Pc), Polycomblike (Pcl), and extra sex comb (esc), and acts as a dominant enhancer of Pc. The expression of ph depends on the ANT-C and BX-C dosage...
  59. Struhl G, White R. Regulation of the Ultrabithorax gene of Drosophila by other bithorax complex genes. Cell. 1985;43:507-19 pubmed
    ..Using mutations in the abd-A, Abd-B, and esc (extra sex combs) genes we show that these distinct patterns result in large part from regulation of Ubx gene expression by ..
  60. Birchler J, Bhadra M, Bhadra U. Making noise about silence: repression of repeated genes in animals. Curr Opin Genet Dev. 2000;10:211-6 pubmed
    ..The silencing mechanisms potentially act as a protection against high expression of transposons and viruses. ..
  61. Remillieux Leschelle N, Santamaria P, Randsholt N. Regulation of larval hematopoiesis in Drosophila melanogaster: a role for the multi sex combs gene. Genetics. 2002;162:1259-74 pubmed
    ..We propose that mxc provides one of the regulatory inputs in larval hematopoiesis that control normal rates of plasmatocyte and crystal lineage proliferation as well as normal rates and timing of hemocyte differentiation. ..
  62. Bienz M, Muller J. Transcriptional silencing of homeotic genes in Drosophila. Bioessays. 1995;17:775-84 pubmed
    ..We propose that this complex consists of specific non-histone proteins, the so-called Polycomb group proteins, and that it is anchored at SIL-M elements and at the promoter. ..
  63. Ptak S, Petrov D. How intron splicing affects the deletion and insertion profile in Drosophila melanogaster. Genetics. 2002;162:1233-44 pubmed
    ..Altogether we demonstrate that constraints in introns may explain much of the difference in the pattern of deletions and insertions observed in Drosophila introns and pseudogenes. ..
  64. Ali J, Bender W. Cross-regulation among the polycomb group genes in Drosophila melanogaster. Mol Cell Biol. 2004;24:7737-47 pubmed
    ..proteins found in two evolutionarily conserved chromatin complexes, Polycomb repressive complex 1 (PRC1) and the ESC-E(Z) complex. The majority of the group, lacking clear biochemical functions, might be indirect regulators...
  65. Sathe S, Harte P. The Drosophila extra sex combs protein contains WD motifs essential for its function as a repressor of homeotic genes. Mech Dev. 1995;52:77-87 pubmed
    b>Extra sex combs is a member of the Polycomb Group genes, whose products are required for stable long term transcriptional repression of the homeotic genes of the Bithorax and Antennapedia complexes...
  66. Carroll S, Laymon R, McCutcheon M, Riley P, Scott M. The localization and regulation of Antennapedia protein expression in Drosophila embryos. Cell. 1986;47:113-22 pubmed
    ..The distributions of the Antp and the Ultrabithorax (Ubx) proteins in doubly-labeled embryos suggest that the Ubx protein may be one direct negative regulator of Antp gene expression. ..
  67. Furuyama T, Banerjee R, Breen T, Harte P. SIR2 is required for polycomb silencing and is associated with an E(Z) histone methyltransferase complex. Curr Biol. 2004;14:1812-21 pubmed
    ..SIR2 is involved in the epigenetic inheritance of silent chromatin states mediated by the Drosophila Polycomb group proteins and is physically associated with a complex containing the E(Z) histone methyltransferase. ..
  68. Kalb R, Latwiel S, Baymaz H, Jansen P, Müller C, Vermeulen M, et al. Histone H2A monoubiquitination promotes histone H3 methylation in Polycomb repression. Nat Struct Mol Biol. 2014;21:569-71 pubmed publisher
    ..Jarid2, Aebp2 and H2Aub thus constitute components of a positive feedback loop establishing H3K27me3 chromatin domains. ..
  69. Cao R, Zhang Y. The functions of E(Z)/EZH2-mediated methylation of lysine 27 in histone H3. Curr Opin Genet Dev. 2004;14:155-64 pubmed
    ..Drosophila and mammalian cells indicate that PcG proteins function in at least two distinct protein complexes: the ESC-E(Z) or EED-EZH2 complex, and the PRC1 complex...
  70. Ng J, Li R, Morgan K, Simon J. Evolutionary conservation and predicted structure of the Drosophila extra sex combs repressor protein. Mol Cell Biol. 1997;17:6663-72 pubmed
    The Drosophila extra sex combs (esc) protein, a member of the Polycomb group (PcG), is a transcriptional repressor of homeotic genes...
  71. Lopez A, Higuet D, Rosset R, Deutsch J, Peronnet F. corto genetically interacts with Pc-G and trx-G genes and maintains the anterior boundary of Ultrabithorax expression in Drosophila larvae. Mol Genet Genomics. 2001;266:572-83 pubmed
    ..Our results provide evidence that corto maintains the anterior border of Ubx expression in third-instar larvae. We suggest that this regulation is accomplished through an interaction with the products of the Pc-G and trx-G genes. ..
  72. Gebuhr T, Bultman S, Magnuson T. Pc-G/trx-G and the SWI/SNF connection: developmental gene regulation through chromatin remodeling. Genesis. 2000;26:189-97 pubmed
    ..genesis 26:189-197, 2000. ..
  73. Pindyurin A, Moorman C, de Wit E, Belyakin S, Belyaeva E, Christophides G, et al. SUUR joins separate subsets of PcG, HP1 and B-type lamin targets in Drosophila. J Cell Sci. 2007;120:2344-51 pubmed
    ..Distinct subsets of SUUR targets are associated with PcG proteins (Pc and Esc; Polycomb and Extra sexcombs), heterochromatic proteins [HP1 and SU(VAR)3-9] and B-type lamin...
  74. Fitzgerald D, Bender W. Polycomb group repression reduces DNA accessibility. Mol Cell Biol. 2001;21:6585-97 pubmed
    ..In contrast to SIR silencing, however, episomes excised from Polycomb-repressed chromosomal sites do not show an altered superhelix density. ..
  75. Mishra K, Chopra V, Srinivasan A, Mishra R. Trl-GAGA directly interacts with lola like and both are part of the repressive complex of Polycomb group of genes. Mech Dev. 2003;120:681-9 pubmed
    ..These observations suggest a possible mechanism of how Trl-GAGA plays a role in maintaining the repressed state of target genes involving lolal, which may function as a mediator to recruit PcG complexes. ..
  76. Mason Suares H, Tie F, Yan C, Harte P. Polycomb silencing of the Drosophila 4E-BP gene regulates imaginal disc cell growth. Dev Biol. 2013;380:111-24 pubmed publisher
    ..These include mutations in E(z), Su(z)12, esc and escl, which encode Polycomb repressive complex 2 (PRC2) subunits...
  77. Casanova J, Sánchez Herrero E, Morata G. Developmental analysis of a hybrid gene composed of parts of the Ubx and abd-A genes of Drosophila. EMBO J. 1988;7:1097-105 pubmed
    ..of conventional bithorax genes: it can be spatially derepressed in the absence of trans-acting genes like extra Sex combs or Polycomb and in turn represses other homeotics like Sex combs reduced...
  78. Bengani H, Mendiratta S, Maini J, Vasanthi D, Sultana H, Ghasemi M, et al. Identification and Validation of a Putative Polycomb Responsive Element in the Human Genome. PLoS ONE. 2013;8:e67217 pubmed publisher
    ..Taken together, our results strongly suggest that PRE-PIK3C2B functions as a site of interaction for polycomb proteins. ..
  79. Sinclair D, Clegg N, Antonchuk J, Milne T, Stankunas K, Ruse C, et al. Enhancer of Polycomb is a suppressor of position-effect variegation in Drosophila melanogaster. Genetics. 1998;148:211-20 pubmed
    ..However, E(Pc) appears to be an important link between the two groups. We discuss why Asx might act as an enhancer of PEV. ..
  80. Choi S, Oh C, Kim S, Kim Y, Jeon S. Effects of Polycomb group mutations on the expression of Ultrabithorax in the Drosophila visceral mesoderm. Mol Cells. 2000;10:156-61 pubmed
    ..Mutant embryos of the genes extra sex combs (esc), Polycomb (Pc), additional sex combs (Asx) and pleiohomeotic (pho) were examined...
  81. Maeda R, Karch F. The ABC of the BX-C: the bithorax complex explained. Development. 2006;133:1413-22 pubmed
    ..This primer attempts to distill and explain our current knowledge about this classic, complex locus. ..
  82. Tie F, Siebold A, Harte P. The N-terminus of Drosophila ESC mediates its phosphorylation and dimerization. Biochem Biophys Res Commun. 2005;332:622-32 pubmed
    The ESC protein, like other Polycomb Group proteins, is required for heritable silencing of the homeotic genes. ESC is phosphorylated in vivo, but the region of ESC that is phosphorylated and its consequences are not known...
  83. Paro R. Imprinting a determined state into the chromatin of Drosophila. Trends Genet. 1990;6:416-21 pubmed
  84. Hannah Alava A. Developmental Genetics of the Posterior Legs in Drosophila Melanogaster. Genetics. 1958;43:878-905 pubmed