Conserved CTCF Insulator Elements Flank the Mouse and Human β-Globin Loci

REFERENCES

• Ashe, H. L., J. Monks, M. Wijgerde, P. Fraser, and N. J. Proudfoot. 1998. Intergenic transcription and transinduction of the human β-globin locus. Genes Dev. 11: 2494–2509.
   Google Scholar
• Bell, A. C., A. G. West, and G. Felsenfeld. 1999. The protein CTCF is required for the enhancer-blocking activity of vertebrate insulators. Cell 98: 387–396.
   PubMed Web of Science ®Google Scholar
• Bell, A. C., A. G. West, and G. Felsenfeld. 2001. Insulators and boundaries: versatile regulatory elements in the eukaryotic genome. Science 291: 447–450.
   PubMedGoogle Scholar
• Bender, M. A., A. Reik, J. Close, A. Telling, E. Epner, S. Fiering, R. Hardison, and M. Groudine. 1998. Description and targeted deletion of 5′ hypersensitive site 5 and 6 of the mouse β-globin locus control region. Blood 92: 4394–4403.
   PubMedGoogle Scholar
• Bender, M. A., M. Bulger, J. Close, and M. Groudine. 2000. β-Globin gene switching and DNase I sensitivity of the endogenous β-globin locus in mice do not require the locus control region. Mol. Cell 5: 387–393.
   PubMed Web of Science ®Google Scholar
• Bulger, M., J. H. von Doorninck, N. Saitoh, A. Telling, C. Farrell, M. A. Bender, G. Felsenfeld, R. Axel, and M. Groudine. 1999. Conservation of sequence and structure flanking the mouse and human β-globin loci: the β-globin genes are embedded within an array of odorant receptor genes. Proc. Natl. Acad. Sci. USA 96: 5129–5134.
   PubMedGoogle Scholar
• Bulger, M., M. A. Bender, J. H. van Doorninck, B. Wertman, C. M. Farrell, G. Felsenfeld, M. Groudine, and R. C. Hardison. 2000. Comparative structural and functional analysis of the olfactory receptor genes flanking the human and mouse β-globin gene clusters. Proc. Natl. Acad. Sci. USA 97: 14560–14565.
   PubMedGoogle Scholar
• Burcin, M., R. Arnold, M. Lutz, B. Kaiser, D. Runge, F. Lottspeich, G. N. Filippova, V. V. Lobanenkov, and R. Renkawitz. 1997. Negative protein 1, which is required for function of the chicken lysozyme gene silencer in conjunction with hormone receptors, is identical to the multivalent zinc finger repressor CTCF. Mol. Cell. Biol. 17: 1281–1288.
   PubMed Web of Science ®Google Scholar
• Chung, J. H., M. Whitely, and G. Felsenfeld. 1993. A 5′ element of the chicken β-globin domain serves as an insulator in human erythroid cells and protects against position effect in Drosophila. Cell 74: 505–514.
   PubMed Web of Science ®Google Scholar
• Chung, J. H., A. C. Bell, and G. Felsenfeld. 1997. Characterization of the chicken β-globin insulator. Proc. Natl. Acad. Sci. USA 94: 575–580.
   PubMed Web of Science ®Google Scholar
• Dhar, V., A. Nandi, C. L. Schildkraut, and A. I. Skoultchi. 1990. Erythroid-specific nuclease-hypersensitive sites flanking the human β-globin domain. Mol. Cell. Biol. 10: 4324–4333.
   PubMedGoogle Scholar
• Engel, J. D., and K. Tanimoto. 2000. Looping, linking, and chromatin activity: new insights into β-globin locus regulation. Cell 100: 499–502.
   PubMedGoogle Scholar
• Epner, E., A. Reik, D. Cimbora, A. Telling, M. A. Bender, S. Fiering, T. Enver, D. I. K. Martin, M. Kennedy, G. Keller, and M. Groudine. 1998. The β-globin LCR is not necessary for an open chromatin structure or developmentally regulated transcription of the native mouse β-globin locus. Mol. Cell 2: 447–455.
   PubMedGoogle Scholar
• Farrell, C. M., A. Grinberg, S. P. Huang, D. Chen, J. G. Pichel, H. Westphal, and G. Felsenfeld. 2000. A large upstream region is not necessary for gene expression or hypersensitive site formation at the mouse β-globin locus. Proc. Natl. Acad. Sci. USA 97: 14554–14559.
   PubMedGoogle Scholar
• Filippova, G. N., S. Fagerlie, E. M. Klenova, C. Myers, Y. Dehner, G. Goodwin, P. E. Neiman, S. J. Collins, and V. V. Lobanenkov. 1996. An exceptionally conserved transcriptional repressor, CTCF, employs different combinations of zinc fingers to bind diverged promoter sequences of avian and mammalian c-myc oncogenes. Mol. Cell. Biol. 16: 2802–2813.
   PubMed Web of Science ®Google Scholar
• Fleenor, D. E., and R. E. Kaufman. 1993. Characterization of the DNase I hypersensitive site 3′ of the human β globin gene domain. Blood 81: 2781–2790.
   PubMed Web of Science ®Google Scholar
• Forget, B. G. 1998. Molecular basis of hereditary persistence of fetal hemoglobin. Ann. N. Y. Acad. Sci. 850: 38–44.
   PubMed Web of Science ®Google Scholar
• Forsberg, C. M., and E. H. Bresnick. 2001. Histone acetylation beyond promoters: long-range acetylation patterns in the chromatin world. Bioessays 23: 820–830.
   PubMedGoogle Scholar
• Gerasimova, T. I., K. Byrd, and V. G. Corces. 2000. A chromatin insulator determines the nuclear localization of DNA. Mol. Cell 6: 1025–1035.
   PubMedGoogle Scholar
• Gribnau, J., K. Diderich, S. Pruzina, R. Calzolari, and P. Fraser. 2000. Intergenic transcription and developmental remodeling of chromatin subdomains in the human β-globin locus. Mol. Cell 5: 377–386.
   PubMedGoogle Scholar
• Hardison, R., J. L. Slightom, D. L. Gumucio, M. Goodman, N. Stojanovic, and W. Miller. 1997. Locus control regions of mammalian β-globin gene clusters: combining phylogenetic analyses and experimental results to gain functional insights. Gene 205: 73–94.
   PubMed Web of Science ®Google Scholar
• Hebbes, T. R., A. L. Clayton, A. W. Thorne, and C. Crane-Robinson. 1994. Core histone hyperacetylation co-maps with generalized DNase I sensitivity in the chicken β-globin chromosomal domain. EMBO J. 13: 1823–1830.
   PubMed Web of Science ®Google Scholar
• Higgs, D. R. 1998. Do LCRs open chromatin domains? Cell 95: 299–302.
   PubMedGoogle Scholar
• Jackson, J. D., H. Petrykowska, S. Philipsen, W. Miller, and R. Hardison. 1996. Role of DNA sequences outside the cores of DNase hypersensitive sites (HSs) in functions of the β-globin locus control region: domain opening and synergism between HS2 and HS3. J. Biol. Chem. 271: 11871–11878.
   PubMed Web of Science ®Google Scholar
• Jarman, A. P., and D. R. Higgs. 1988. Nuclear scaffold attachment sites in the human globin gene complexes. EMBO J. 7: 3337–3344.
   PubMedGoogle Scholar
• Li, Q., and G. Stamatoyannopoulos. 1994. Hypersensitive site 5 of the human β locus control region functions as a chromatin insulator. Blood 84: 1399–1401.
   PubMed Web of Science ®Google Scholar
• Li, Q., M. Zhang, Z. Duan, and G. Stamatoyannopoulos. 1999. Structural analysis and mapping of DNase I hypersensitivity of HS5 of the β-globin locus control region. Genomics 61: 183–193.
   PubMedGoogle Scholar
• Litt, M. D., M. Simpson, F. Recillas-Targa, M.-N. Prioleau, and G. Felsenfeld. 2001. Transitions in histone acetylation reveal boundaries of three separately regulated neighboring loci. EMBO J. 20: 2224–2235.
   PubMedGoogle Scholar
• Long, Q., C. Bengra, C. Li, F. Kutlar, and D. Tuan. 1998. A long terminal repeat of the human endogenous retrovirus ERV-9 is located in the 5′ boundary area of the human β-globin locus control region. Genomics 54: 542–555.
   PubMedGoogle Scholar
• Ohlsson, R., R. Renkawitz, and V. Lobanenkov. 2001. CTCF is a uniquely versatile transcription regulator linked to epigenetics and disease. Trends Gen. 17: 520–527.
   PubMed Web of Science ®Google Scholar
• Plant, K. E., S. J. E. Routledge, and N. J. Proudfoot. 2001. Intergenic transcription in the human β-globin gene cluster. Mol. Cell. Biol. 21: 6507–6514.
   PubMedGoogle Scholar
• Prioleau, M.-N., P. Nony, M. Simpson, and G. Felsenfeld. 1999. An insulator element and condensed chromatin region separate the chicken β-globin locus from an independently regulated erythroid-specific folate receptor gene. EMBO J. 18: 4035–4048.
   PubMedGoogle Scholar
• Ramchandran, R., C. Bengra, B. Whitney, K. Lanclos, and D. Tuan. 2000. A (GATA)7 motif located in the 5′ boundary area of the human β-globin locus control region exhibits silencer activity in erythroid cells. Am. J. Hematol. 65: 14–24.
   PubMedGoogle Scholar
• Reik, A., A. Telling, G. Zitnik, D. Cimbora, E. Epner, and M. Groudine. 1998. The locus control region is necessary for gene expression in the human β-globin locus but not the maintenance of an open chromatin structure in erythroid cells. Mol. Cell. Biol. 18: 5992–6000.
   PubMed Web of Science ®Google Scholar
• Reitman, M., and G. Felsenfeld. 1990. Developmental regulation of topoisomerase II sites and DNase I-hypersensitive sites in the chicken β-globin locus. Mol. Cell. Biol. 10: 2774–2786.
   PubMedGoogle Scholar
• Saitoh, N., A. C. Bell, F. Recillas-Targa, A. G. West, M. Simpson, M. Pikaart, and G. Felsenfeld. 1999. Structural and functional conservation at the boundaries of the chicken β-globin domain. EMBO J. 19: 2315–2322.
   Google Scholar
• Schübeler, D., C. Francastel, D. M. Cimbora, A. Reik, D. I. K. Martin, and M. Groudine. 2000. Nuclear localization and histone acetylation: a pathway for chromatin opening and transcriptional activation of the human β-globin locus. Genes Dev. 14: 940–950.
   PubMed Web of Science ®Google Scholar
• Staines, D. M., and J. O. Thomas. 1999. A sequence with homology to human HPFH-linked enhancer elements and to a family of G-protein linked membrane receptor genes is located downstream of the chicken β-globin locus. Gene 234: 345–352.
   PubMedGoogle Scholar
• Stamatoyannopoulos, G., and F. Grosveld. 2001. Hemoglobin switching, p. 135–182. In G. Stamatoyannopoulos, P. W. Majerus, R. M. Perlmutter, and H. Varmus (ed.), The molecular basis of blood diseases, 3rd ed. W. B. Saunders Company, Philadelphia, Pa.
   Google Scholar
• Tanimoto, K., Q. Liu, J. Bungert, and J. D. Engel. 1999. Effects of altered gene order or orientation of the locus control region on human β-globin gene expression in mice. Nature 398: 344–348.
   PubMedGoogle Scholar
• Tuan, D., W. Solomon, Q. Li, and I. M. London. 1985. The “beta-like-globin” gene domain in human erythroid cells. Proc. Natl. Acad. Sci. USA 82: 6384–6388.
   PubMed Web of Science ®Google Scholar
• Yu, J., J. H. Bock, J. L. Slightom, and B. Villeponteau. 1994. A 5′ β-globin matrix-attachment region and the polyoma enhancer together confer position-independent transcription. Gene 139: 139–145.
   PubMed Web of Science ®Google Scholar
• Zafarana, G., S. Raguz, S. Pruzina, F. Grosveld, and D. Meijer. 1995. The regulation of human β-globin gene expression: the analysis of hypersensitive site 5 (HS5) in the LCR, p. 39–44. In G. Stammatoyannopoulos (ed.), Molecular biology of hemoglobin switching. Intercept, Andover, Mass.
   Google Scholar