Symmons, O. et al. The Shh topological domain facilitates the action of remote enhancers by reducing the effects of genomic distances. Dev. Cell 39, 529–543 (2016).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Dekker, C., Haering, C. H., Peters, J. M. & Rowland, B. D. How do molecular motors fold the genome? Science 382, 646–648 (2023).

Article 
CAS 
PubMed 

Google Scholar
 

Gabriele, M. et al. Dynamics of CTCF- and cohesin-mediated chromatin looping revealed by live-cell imaging. Science 376, 496–501 (2022).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Kane, L. et al. Cohesin is required for long-range enhancer action at the Shh locus. Nat. Struct. Mol. Biol. 29, 891–897 (2022).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Gómez Acuña, L. I., Flyamer, I., Boyle, S., Friman, E. T. & Bickmore, W. A. Transcription decouples estrogen-dependent changes in enhancer-promoter contact frequencies and spatial proximity. PLoS Genet. 20, e1011277 (2024).

Article 
PubMed 
PubMed Central 

Google Scholar
 

Alexander, J. M. et al. Live-cell imaging reveals enhancer dependent Sox2 transcription in the absence of enhancer proximity. eLife 8, e41769 (2019).

Article 
PubMed 
PubMed Central 

Google Scholar
 

Calderon, L. et al. Cohesin-dependence of neuronal gene expression relates to chromatin loop length. eLife. 11, e76539 (2022).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Rinzema, N. J. et al. Building regulatory landscapes reveals that an enhancer can recruit cohesin to create contact domains, engage CTCF sites and activate distant genes. Nat. Struct. Mol. Biol. 29, 563–574 (2022).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Karr, J. P., Ferrie, J. J., Tjian, R. & Darzacq, X. The transcription factor activity gradient (TAG) model: contemplating a contact-independent mechanism for enhancer-promoter communication. Genes Dev. 36, 7–16 (2022).

Article 
CAS 
PubMed 
PubMed Central 

Google Scholar
 

Williamson, I. et al. Bystander activation across a TAD boundary supports a cohesin-dependent transcription cluster model for enhancer function. Genes Dev. https://doi.org/10.1101/gad.352648.125 (2025).

Article 
PubMed 
PubMed Central 

Google Scholar