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To reveal structural transitions underlying heterochromatin formation in maturing mouse rod photoreceptors, we apply cryo-electron microscopy (cryo-EM) tomography, AI-assisted denoising, and molecular modeling. We find that chromatin isolated from immature retina cells contains many closely apposed nucleosomes with extremely short or absent nucleosome linkers, which are inconsistent with the typical two-start zigzag chromatin folding. In mature retina cells, the fraction of short-linker nucleosomes is much lower, supporting stronger chromatin compaction. By cryo-EM-assisted nucleosome interaction capture, we observe that chromatin in immature retina is enriched with i\u00a0\u00b1\u00a01 interactions, while chromatin in mature retina contains predominantly i\u00a0\u00b1\u00a02 interactions typical of the two-start zigzag. By mesoscale modeling and computational simulation, we clarify that the unusually short linkers typical of immature retina are sufficient to inhibit the two-start zigzag and chromatin compaction by the interference of very short linkers with linker DNA stems. We propose that this short linker composition renders nucleosome arrays more open in immature retina and that, as the linker DNA length increases in mature retina, chromatin becomes globally condensed via tight zigzag folding. This mechanism may be broadly utilized to introduce higher chromatin folding entropy for epigenomic plasticity.<\/jats:p>","DOI":"10.1093\/nar\/gkaf457","type":"journal-article","created":{"date-parts":[[2025,6,4]],"date-time":"2025-06-04T13:00:39Z","timestamp":1749042039000},"source":"Crossref","is-referenced-by-count":5,"title":["Compromised two-start zigzag chromatin folding in immature mouse retina cells driven by irregularly spaced nucleosomes with short DNA linkers"],"prefix":"10.1093","volume":"53","author":[{"given":"Brianna","family":"Kable","sequence":"first","affiliation":[{"name":"Penn State University College of Medicine Department Biochemistry & Molecular Biology, , 500 University Drive , Hershey , PA 17033,\u00a0","place":["United States"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Stephanie","family":"Portillo-Ledesma","sequence":"additional","affiliation":[{"name":"New York University Department of Chemistry and Simons Center for Computational 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