• We are sorry, but NCBI web applications do not support your browser and may not function properly. More information
Logo of geneticsGeneticsCurrent IssueInformation for AuthorsEditorial BoardSubscribeSubmit a Manuscript
Genetics. Jul 2000; 155(3): 1297–1311.
PMCID: PMC1461175

Deletion of an insulator element by the mutation facet-strawberry in Drosophila melanogaster.


Eukaryotic chromosomes are thought to be subdivided into a series of structurally and functionally independent units. Critical to this hypothesis is the identification of insulator or boundary elements that delimit chromosomal domains. The properties of a Notch mutation, facet-strawberry (fa(swb)), suggest that this small deletion disrupts such a boundary element. fa(swb) is located in the interband separating polytene band 3C7, which contains Notch, from the distal band 3C6. The fa(swb) mutation alters the structural organization of the chromosome by deleting the interband and fusing 3C7 with 3C6. Genetic studies also suggest that fa(swb) compromises the functional autonomy of Notch by allowing the locus to become sensitive to chromosomal position effects emanating from distal sequences. In the studies reported here, we show that a DNA fragment spanning the fa(swb) region can insulate reporter transgenes against chromosomal position effects and can block enhancer-promoter interactions. Moreover, we find that insulating activity is dependent on sequences deleted in fa(swb). These results provide evidence that the element defined by the fa(swb) mutation corresponds to an insulator.

Full Text

The Full Text of this article is available as a PDF (373K).

Selected References

These references are in PubMed. This may not be the complete list of references from this article.
  • Arkhipova IR. Promoter elements in Drosophila melanogaster revealed by sequence analysis. Genetics. 1995 Mar;139(3):1359–1369. [PMC free article] [PubMed]
  • Artavanis-Tsakonas S. The molecular biology of the Notch locus and the fine tuning of differentiation in Drosophila. Trends Genet. 1988 Apr;4(4):95–100. [PubMed]
  • Artavanis-Tsakonas S, Muskavitch MA, Yedvobnick B. Molecular cloning of Notch, a locus affecting neurogenesis in Drosophila melanogaster. Proc Natl Acad Sci U S A. 1983 Apr;80(7):1977–1981. [PMC free article] [PubMed]
  • Avramova Z, Tikhonov A. Are scs and scs' 'neutral' chromatin domain boundaries of the locus? Trends Genet. 1999 Apr;15(4):138–139. [PubMed]
  • Bell AC, Felsenfeld G. Stopped at the border: boundaries and insulators. Curr Opin Genet Dev. 1999 Apr;9(2):191–198. [PubMed]
  • Bell AC, West AG, Felsenfeld G. The protein CTCF is required for the enhancer blocking activity of vertebrate insulators. Cell. 1999 Aug 6;98(3):387–396. [PubMed]
  • Burke TW, Kadonaga JT. Drosophila TFIID binds to a conserved downstream basal promoter element that is present in many TATA-box-deficient promoters. Genes Dev. 1996 Mar 15;10(6):711–724. [PubMed]
  • Burke TW, Kadonaga JT. The downstream core promoter element, DPE, is conserved from Drosophila to humans and is recognized by TAFII60 of Drosophila. Genes Dev. 1997 Nov 15;11(22):3020–3031. [PMC free article] [PubMed]
  • Cai H, Levine M. Modulation of enhancer-promoter interactions by insulators in the Drosophila embryo. Nature. 1995 Aug 10;376(6540):533–536. [PubMed]
  • Chung JH, Whiteley M, Felsenfeld G. A 5' element of the chicken beta-globin domain serves as an insulator in human erythroid cells and protects against position effect in Drosophila. Cell. 1993 Aug 13;74(3):505–514. [PubMed]
  • Chung JH, Bell AC, Felsenfeld G. Characterization of the chicken beta-globin insulator. Proc Natl Acad Sci U S A. 1997 Jan 21;94(2):575–580. [PMC free article] [PubMed]
  • Dillon N, Trimborn T, Strouboulis J, Fraser P, Grosveld F. The effect of distance on long-range chromatin interactions. Mol Cell. 1997 Dec;1(1):131–139. [PubMed]
  • Dorsett D. Distance-independent inactivation of an enhancer by the suppressor of Hairy-wing DNA-binding protein of Drosophila. Genetics. 1993 Aug;134(4):1135–1144. [PMC free article] [PubMed]
  • Galloni M, Gyurkovics H, Schedl P, Karch F. The bluetail transposon: evidence for independent cis-regulatory domains and domain boundaries in the bithorax complex. EMBO J. 1993 Mar;12(3):1087–1097. [PMC free article] [PubMed]
  • Gaszner M, Vazquez J, Schedl P. The Zw5 protein, a component of the scs chromatin domain boundary, is able to block enhancer-promoter interaction. Genes Dev. 1999 Aug 15;13(16):2098–2107. [PMC free article] [PubMed]
  • Gerasimova TI, Corces VG. Polycomb and trithorax group proteins mediate the function of a chromatin insulator. Cell. 1998 Feb 20;92(4):511–521. [PubMed]
  • Gerasimova TI, Corces VG. Domains and boundaries in chromosomes. Results Probl Cell Differ. 1999;25:229–249. [PubMed]
  • Gerasimova TI, Gdula DA, Gerasimov DV, Simonova O, Corces VG. A Drosophila protein that imparts directionality on a chromatin insulator is an enhancer of position-effect variegation. Cell. 1995 Aug 25;82(4):587–597. [PubMed]
  • Geyer PK. The role of insulator elements in defining domains of gene expression. Curr Opin Genet Dev. 1997 Apr;7(2):242–248. [PubMed]
  • Geyer PK, Corces VG. DNA position-specific repression of transcription by a Drosophila zinc finger protein. Genes Dev. 1992 Oct;6(10):1865–1873. [PubMed]
  • Ramos RG, Grimwade BG, Wharton KA, Scottgale TN, Artavanis-Tsakonas S. Physical and functional definition of the Drosophila Notch locus by P element transformation. Genetics. 1989 Oct;123(2):337–348. [PMC free article] [PubMed]
  • Glover DM, Leibowitz MH, McLean DA, Parry H. Mutations in aurora prevent centrosome separation leading to the formation of monopolar spindles. Cell. 1995 Apr 7;81(1):95–105. [PubMed]
  • Rykowski MC, Parmelee SJ, Agard DA, Sedat JW. Precise determination of the molecular limits of a polytene chromosome band: regulatory sequences for the Notch gene are in the interband. Cell. 1988 Aug 12;54(4):461–472. [PubMed]
  • GREEN MM. Back mutation in Drosophila melanogaster. II. Data on additional yellow and white mutants. Genetics. 1962 Apr;47:483–488. [PMC free article] [PubMed]
  • Scott KS, Geyer PK. Effects of the su(Hw) insulator protein on the expression of the divergently transcribed Drosophila yolk protein genes. EMBO J. 1995 Dec 15;14(24):6258–6267. [PMC free article] [PubMed]
  • Greenwald I. Structure/function studies of lin-12/Notch proteins. Curr Opin Genet Dev. 1994 Aug;4(4):556–562. [PubMed]
  • Smith PA, Corces VG. The suppressor of Hairy-wing binding region is required for gypsy mutagenesis. Mol Gen Genet. 1992 May;233(1-2):65–70. [PubMed]
  • Gyurkovics H, Gausz J, Kummer J, Karch F. A new homeotic mutation in the Drosophila bithorax complex removes a boundary separating two domains of regulation. EMBO J. 1990 Aug;9(8):2579–2585. [PMC free article] [PubMed]
  • Smith PA, Corces VG. The suppressor of Hairy-wing protein regulates the tissue-specific expression of the Drosophila gypsy retrotransposon. Genetics. 1995 Jan;139(1):215–228. [PMC free article] [PubMed]
  • Hagstrom K, Muller M, Schedl P. Fab-7 functions as a chromatin domain boundary to ensure proper segment specification by the Drosophila bithorax complex. Genes Dev. 1996 Dec 15;10(24):3202–3215. [PubMed]
  • Spradling AC, Rubin GM. Transposition of cloned P elements into Drosophila germ line chromosomes. Science. 1982 Oct 22;218(4570):341–347. [PubMed]
  • Zannad F, Vaur L, Dutrey-Dupagne C, Genès N, Chatellier G, Elkik F, Ménard J. Assessment of drug efficacy using home self-blood pressure measurement: the SMART study. Self Measurement for the Assessment of the Response to Trandolapril. J Hum Hypertens. 1996 Jun;10(6):341–347. [PubMed]
  • Tomlinson A, Kimmel BE, Rubin GM. rough, a Drosophila homeobox gene required in photoreceptors R2 and R5 for inductive interactions in the developing eye. Cell. 1988 Dec 2;55(5):771–784. [PubMed]
  • Hart CM, Zhao K, Laemmli UK. The scs' boundary element: characterization of boundary element-associated factors. Mol Cell Biol. 1997 Feb;17(2):999–1009. [PMC free article] [PubMed]
  • Hiromi Y, Gehring WJ. Regulation and function of the Drosophila segmentation gene fushi tarazu. Cell. 1987 Sep 11;50(6):963–974. [PubMed]
  • Udvardy A, Maine E, Schedl P. The 87A7 chromomere. Identification of novel chromatin structures flanking the heat shock locus that may define the boundaries of higher order domains. J Mol Biol. 1985 Sep 20;185(2):341–358. [PubMed]
  • Holdridge C, Dorsett D. Repression of hsp70 heat shock gene transcription by the suppressor of hairy-wing protein of Drosophila melanogaster. Mol Cell Biol. 1991 Apr;11(4):1894–1900. [PMC free article] [PubMed]
  • Vazquez J, Schedl P. Sequences required for enhancer blocking activity of scs are located within two nuclease-hypersensitive regions. EMBO J. 1994 Dec 15;13(24):5984–5993. [PMC free article] [PubMed]
  • Kellum R, Schedl P. A group of scs elements function as domain boundaries in an enhancer-blocking assay. Mol Cell Biol. 1992 May;12(5):2424–2431. [PMC free article] [PubMed]
  • Welshons WJ, Keppy DO. Intragenic deletions and salivary band relationships in Drosophila. Genetics. 1975 May;80(1):143–155. [PMC free article] [PubMed]
  • Kellum R, Schedl P. A position-effect assay for boundaries of higher order chromosomal domains. Cell. 1991 Mar 8;64(5):941–950. [PubMed]
  • Keppy DO, Welshons WJ. The cytogenetics of a recessive visible mutant associated with a deficiency adjacent to the notch locus in Drosophila melanogaster. Genetics. 1977 Mar;85(3):497–506. [PMC free article] [PubMed]
  • Welshons WJ, Welshons HJ. Enhancement and Suppression of a Euchromatic Position Effect at NOTCH in Drosophila. Genetics. 1986 Jun;113(2):337–354. [PMC free article] [PubMed]
  • Kidd S, Lockett TJ, Young MW. The Notch locus of Drosophila melanogaster. Cell. 1983 Sep;34(2):421–433. [PubMed]
  • Wharton KA, Johansen KM, Xu T, Artavanis-Tsakonas S. Nucleotide sequence from the neurogenic locus notch implies a gene product that shares homology with proteins containing EGF-like repeats. Cell. 1985 Dec;43(3 Pt 2):567–581. [PubMed]
  • Mihaly J, Hogga I, Gausz J, Gyurkovics H, Karch F. In situ dissection of the Fab-7 region of the bithorax complex into a chromatin domain boundary and a Polycomb-response element. Development. 1997 May;124(9):1809–1820. [PubMed]
  • Worcel A, Gargiulo G, Jessee B, Udvardy A, Louis C, Schedl P. Chromatin fine structure of the histone gene complex of Drosophila melanogaster. Nucleic Acids Res. 1983 Jan 25;11(2):421–439. [PMC free article] [PubMed]
  • Mihaly J, Hogga I, Barges S, Galloni M, Mishra RK, Hagstrom K, Müller M, Schedl P, Sipos L, Gausz J, et al. Chromatin domain boundaries in the Bithorax complex. Cell Mol Life Sci. 1998 Jan;54(1):60–70. [PubMed]
  • Zhao K, Hart CM, Laemmli UK. Visualization of chromosomal domains with boundary element-associated factor BEAF-32. Cell. 1995 Jun 16;81(6):879–889. [PubMed]
  • Ohtsuki S, Levine M. GAGA mediates the enhancer blocking activity of the eve promoter in the Drosophila embryo. Genes Dev. 1998 Nov 1;12(21):3325–3330. [PMC free article] [PubMed]
  • Zhou J, Barolo S, Szymanski P, Levine M. The Fab-7 element of the bithorax complex attenuates enhancer-promoter interactions in the Drosophila embryo. Genes Dev. 1996 Dec 15;10(24):3195–3201. [PubMed]
  • Peifer M, Bender W. Sequences of the gypsy transposon of Drosophila necessary for its effects on adjacent genes. Proc Natl Acad Sci U S A. 1988 Dec;85(24):9650–9654. [PMC free article] [PubMed]

Articles from Genetics are provided here courtesy of Genetics Society of America


Related citations in PubMed

See reviews...See all...

Cited by other articles in PMC

See all...


Recent Activity

Your browsing activity is empty.

Activity recording is turned off.

Turn recording back on

See more...