The DOI matches its Crossref record; 43 of the 44 references it cites were found in a registry or an index.
CTCF-dependent enhancer-blocking by alternative chromatin loop formation
Open accessAbstract
The mechanism underlying enhancer-blocking by insulators is unclear. We explored the activity of human beta-globin HS5, the orthologue of the CTCF-dependent chicken HS4 insulator. An extra copy of HS5 placed between the beta-globin locus control region (LCR) and downstream genes on a transgene fulfills the classic predictions for an enhancer-blocker. Ectopic HS5 does not perturb the LCR but blocks gene activation by interfering with RNA pol II, activator and coactivator recruitment, and epigenetic modification at the downstream beta-globin gene. Underlying these effects, ectopic HS5 disrupts chromatin loop formation between beta-globin and the LCR, and instead forms a new loop with endogenous HS5 that topologically isolates the LCR. Both enhancer-blocking and insulator-loop formation depend on an intact CTCF site in ectopic HS5 and are sensitive to knock-down of the CTCF protein by siRNA. Thus, intrinsic looping activity of CTCF sites can nullify LCR function.
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“Transgene assays of the human β-globin locus then showed that two insulator sites binding CTCF could form a chromatin loop [332].”
From Transcriptional enhancers and their communication with gene promoters (2021) · cites this paper as [332]
In the citing paper
“In erythroid cells expressing globin genes, a large CTCF-CTCF loop encompasses the beta-globin genes and the locus control region (LCR), consistent with the organization of the locus in an insulated neighborhood.”
From Insulated Neighborhoods: Structural and Functional Units of Mammalian Gene Control (2016) · cites this paper as Hou et al., 2008
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Cites
High-Resolution Profiling of Histone Methylations in the Human Genome
Barski, A., Cuddapah, S., Cui, K., Roh, T.-Y., Schones, D. E., Wang, Z., Wei, G., Chepelev, I., & Zhao, K. (2007) · Cell · doi:10.1016/j.cell.2007.05.009 · Cited by 6,845
Capturing Chromosome Conformation
Dekker, J., Rippe, K., Dekker, M., & Kleckner, N. (2002) · Science · doi:10.1126/science.1067799 · Cited by 3,652
Looping and Interaction between Hypersensitive Sites in the Active β-globin Locus
Tolhuis, B., Palstra, R.-J., Splinter, E., Grosveld, F., & de Laat, W. (2002) · Molecular Cell · doi:10.1016/s1097-2765(02)00781-5 · Cited by 1,344
Cohesin mediates transcriptional insulation by CCCTC-binding factor
Wendt, K. S., Yoshida, K., Itoh, T., Bando, M., Koch, B., Schirghuber, E., Tsutsumi, S., Nagae, G., Ishihara, K., Mishiro, T., et al. (2008) · Nature · doi:10.1038/nature06634 · Cited by 1,193
The protein CTCF is required for the enhancer blocking activity of vertebrate insulators
Bell, A. C., West, A. G., & Felsenfeld, G. (1999) · Cell · doi:10.1016/s0092-8674(00)81967-4 · Cited by 1,101
Analysis of the Vertebrate Insulator Protein CTCF-Binding Sites in the Human Genome
Kim, T. H., Abdullaev, Z., Smith, A. D., Ching, K. A., Loukinov, D. I., Green, R. D., Zhang, M. Q., Lobanenkov, V. V., & Ren, B. (2007) · Cell · doi:10.1016/j.cell.2006.12.048 · Cited by 1,022
Cohesins Functionally Associate with CTCF on Mammalian Chromosome Arms
Parelho, V., Hadjur, S., Spivakov, M., Leleu, M., Sauer, S., Gregson, H. C., Jarmuz, A., Canzonetta, C., Webster, Z., Nesterova, T., et al. (2008) · Cell · doi:10.1016/j.cell.2008.01.011 · Cited by 900
CTCF mediates long-range chromatin looping and local histone modification in the β-globin locus
Splinter, E., Heath, H., Kooren, J., Palstra, R.-J., Klous, P., Grosveld, F., Galjart, N., & de Laat, W. (2006) · Genes & Development · doi:10.1101/gad.399506 · Cited by 677
Long-range chromatin regulatory interactions in vivo
Carter, D., Chakalova, L., Osborne, C. S., Dai, Y.-F., & Fraser, P. (2002) · Nature Genetics · doi:10.1038/ng1051 · Cited by 663
The β-globin nuclear compartment in development and erythroid differentiation
Palstra, R.-J., Tolhuis, B., Splinter, E., Nijmeijer, R., Grosveld, F., & de Laat, W. (2003) · Nature Genetics · doi:10.1038/ng1244 · Cited by 559
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Jia, Z., Li, J., Ge, X., Wu, Y., Guo, Y., & Wu, Q. (2020) · Figshare · doi:10.6084/m9.figshare.12211304
BLAST, FASTA, and String Algorithms
Xia, X. (2026) · Springer eBooks · doi:10.1007/978-3-032-18842-7_1
hnRNPK condensates facilitate enhancer–promoter looping and RNA polymerase II recruitment
Ye, R., Zhao, H., Chen, J., Wu, Q., Wang, D., Cai, Z., Wang, R., Li, J., Cheng, T., Zhang, M., et al. (2026) · Nature Genetics · doi:10.1038/s41588-026-02710-y
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Metadata
- Authors
- Hou, C., Zhao, H., Tanimoto, K., & Dean, A.
- Affiliations
- National Institutes of Health (US)
- University of Tsukuba (JP)
- Corresponding author
- Dean, Ann
- Venue
- Proceedings of the National Academy of Sciences
- Publisher
- National Academy of Sciences
- Published
- 2008-12-23
- Volume
- 105
- Issue
- 51
- Pages
- 20398-20403
- ISSN
- 0027-8424 (ISSN-L)
- 1091-6490
- PMC
- PMC2629272
- OpenAlex
- W2069172573
- Cited by
- 205
- References
- 44
- Citations by year
- 2026: 7 · 2025: 1 · 2024: 4 · 2023: 9 · 2022: 13 · 2021: 13 · 2020: 17 · 2019: 12 · 2018: 21 · 2017: 9 · 2016: 13 · 2015: 9 · 2014: 9 · 2013: 13 · 2012: 15
- FWCI
- 5.48
- Type
- journal article
- Language
- en
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- Proceedings of the National Academy of Sciences (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Funders
- National Institutes of Health
- Japan Society for the Promotion of Science (20678002)
- National Institute of Diabetes and Digestive and Kidney Diseases
- Topics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Genomics and Chromatin Dynamics
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Chromatin Remodeling and Cancer
- Life Sciences › Biochemistry, Genetics and Molecular Biology › Molecular Biology › Epigenetics and DNA Methylation
- Keywords
- CTCF, enhancer blocking, insulators, chromatin looping, locus control region, beta-globin, RNA polymerase II recruitment, coactivator recruitment, epigenetic modifications, topologically associating domains, chromatin topology
- MeSH terms
- CCCTC-Binding Factor, Chromatin, DNA-Binding Proteins, Gene Expression Regulation, Humans, Repressor Proteins, Transgenes, Locus Control Region, Insulator Elements, Epigenesis, Genetic, beta-Globins
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