Evidence map›Paper›PMID 41690937›Full record

ArticleNature communications2026

Extrusion fountains are hallmarks of chromosome organization emerging upon zygotic genome activation.

Aleksandra Galitsyna, Sergey V Ulianov, Mariia Bazarevich, Nikolai S Bykov, Marina Veil, Meijiang Gao, Kristina Perevoschikova, Mikhail S Gelfand, Sergey V Razin, Leonid Mirny and 1 more

Abstract read
In one paragraph

Article in Nature communications, 2026. The graph could read no effect estimate from its abstract, so it casts no vote on the map. Cited by 28 papers.

0numbers the graph read from it
0cells of the map it votes in
28citing papers in PubMed
–field-weighted citation impact
1 · What the graph read from it

What it found

Each row is one number read from the abstract, on the scale the paper reported it, with its interval. Left of the dashed line favours the treatment, right favours the comparator. Under each row is the sentence it came from. New to these charts? A ten-minute tutorial.

The abstract states no effect estimate the extractor could read, or names no intervention and outcome on the map, so this paper lights no cell and moves no belief. It is still indexed, cited and linked below.

2 · The registry

The trial behind it

Trials whose registry record cites this paper, or whose number appears in the abstract. A trial that started after this paper was published is citing it as background, not reporting it.

Neither the registry nor the abstract names a trial number. If this is a trial report, that itself is worth knowing.

3 · Its place in the literature

Who cites it

28 citing papers in PubMed.

  1. Transcription and Three-Dimensional Genome Organization: Cause, Consequence, or Coordination?BioEssays : news and reviews in molecular, cellular and developmental biology · 2026
    Review
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  13. MIA-Jet: Multi-scale Identification Algorithm of Chromatin Jets.bioRxiv : the preprint server for biology · 2025
    Article
  14. Article
  15. Article
  16. Article
  17. Article
  18. Article
  19. Liquid condensates: a new barrier to loop extrusion?Cellular and molecular life sciences : CMLS · 2025
    Review
  20. Transcriptional machinery as an architect of genome structure.Current opinion in structural biology · 2024
    Review
4 · The record

Corrections and comments

5 · Who and what money

Authors and funding

11 authors.

Aleksandra Galitsyna *Institute for Medical Engineering and Science, Massachusetts Institute of Technology, Cambridge, MA, USA. galitsyn@mit.edu.ORCID http://orcid.org/0000-0001-8969-5694
Sergey V Ulianov *Institute of Gene Biology, Russian Academy of Sciences, Moscow, Russia.
Mariia BazarevichDepartment of Developmental Biology, University of Freiburg, Freiburg, Germany.
Nikolai S BykovIndependent researcher, Moscow, Russia.ORCID http://orcid.org/0000-0002-6321-487X
Marina VeilDepartment of Developmental Biology, University of Freiburg, Freiburg, Germany.
Meijiang GaoDepartment of Developmental Biology, University of Freiburg, Freiburg, Germany.
Kristina PerevoschikovaFaculty of Bioengineering and Bioinformatics, M.V. Lomonosov Moscow State University, Leninskiye Gory, 1, building 73, Moscow, Russia.ORCID http://orcid.org/0009-0007-7765-8045
Mikhail S GelfandIndependent researcher, P.O. Box 125476-22, Moscow, Russia.ORCID http://orcid.org/0000-0003-4181-0846
Sergey V RazinInstitute of Gene Biology, Russian Academy of Sciences, Moscow, Russia.ORCID http://orcid.org/0000-0003-1976-8661
Leonid MirnyInstitute for Medical Engineering and Science, Massachusetts Institute of Technology, Cambridge, MA, USA. leonid@mit.edu.ORCID http://orcid.org/0000-0002-0785-5410
Daria OnichtchoukDepartment of Developmental Biology, University of Freiburg, Freiburg, Germany. daria.onichtchouk@biologie.uni-freiburg.de.ORCID http://orcid.org/0000-0001-6497-1445

Funding

Deutsche Forschungsgemeinschaft (German Research Foundation) DFG-EXC2189Deutsche Forschungsgemeinschaft (German Research Foundation) DFG-ON86/5-1Russian Science Foundation (RSF) 21-64-00001-PRussian Science Foundation (RSF) 23-14-00136U.S. Department of Health & Human Services | National Institutes of Health (NIH) AI164728U.S. Department of Health & Human Services | National Institutes of Health (NIH) NS113929
6 · The paper itself

Abstract

The initiation of gene expression during development, known as zygotic genome activation (ZGA), is accompanied by massive changes in chromosome organization. However, the earliest events of chromosome folding and their functional roles remain unclear. Using Hi-C on zebrafish embryos, we discovered that chromosome folding begins early in development with the formation of fountains, distinct elements of chromosome organization. Emerging preferentially at enhancers, fountains show an initial accumulation of cohesin, which later redistributes to CTCF sites at TAD borders. Knockouts of pioneer transcription factors driving ZGA enhancers cause a specific loss of fountains, establishing a causal link between enhancer activation and fountain formation. Polymer simulations demonstrate that fountains may arise as sites of facilitated cohesin loading, requiring two-sided but desynchronized loop extrusion, potentially caused by cohesin collisions with obstacles or internal switching. Moreover, we detected cohesin-dependent fountain patterns at enhancers in mouse cells and found them reemerging with cohesin loading after mitosis. Altogether, fountains represent enhancer-specific elements of chromosome organization and suggest that chromosome folding during development and after cell division starts with facilitated cohesin loading. Observations in multiple systems further support facilitated loading at enhancers as a widespread phenomenon.

Indexed as

ChromosomesGenomeZygoteAnimalsCCCTC-Binding FactorCell Cycle ProteinsChromosomal Proteins, Non-HistoneCohesinsEnhancer Elements, GeneticGene Expression Regulation, DevelopmentalMiceTranscription FactorsZebrafishZebrafish ProteinsCCCTC-Binding FactorCell Cycle ProteinsChromosomal Proteins, Non-HistoneCohesinsTranscription FactorsZebrafish Proteins

Identifiers

PMID41690937
PMCPMC13018191

What OpenQuestion holds

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Registered trials

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Read under generation 80e0d062 · epoch 390. Bibliography from PubMed, PubMed Central and OpenAlex; grants from NIH RePORTER; trial links from ClinicalTrials.gov; estimates, votes and beliefs from the OpenQuestion graph.