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Using a model of transcriptional regulation, we investigated co-regulation between p53 and hormone receptors in different cellular contexts and highlight an enrichment in mammary and muscle cell types. Sequence, structural and epigenomic evidence demonstrate a broadly distributed functional 73-bp p53-estrogen receptor grammar spanning nucleosome entry\/exit to dyad, while p53-androgen receptor grammar spans 146 bp. At a functional breast cancer enhancer regulating Cyclin D, varied spacing led to lower predicted p53 binding. Genome-wide analysis found a major role of Alu sequences in distributing this grammar, in particular in mTORC1 pathway genes, with an evolutionary association of primate body-size sexual dimorphism and muscle aging.<\/jats:p>","DOI":"10.64898\/2026.09.26.754600","type":"posted-content","created":{"date-parts":[[2026,9,28]],"date-time":"2026-09-28T14:51:04Z","timestamp":1790607064000},"source":"Crossref","is-referenced-by-count":0,"title":["Nucleosome-scale p53\u2013hormone receptor grammar distributed by Alu elements"],"prefix":"10.64898","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-9835-5018","authenticated-orcid":false,"given":"Xi","family":"Fu","sequence":"first","affiliation":[{"name":"Program of Mathematical Genomics, Department of Systems Biology, Columbia University Irving Medical Center, New York N.Y"},{"name":"Herbert Irving Comprehensive Cancer Center, Columbia University Irving Medical Center, New York N.Y"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qingyuan","family":"Cai","sequence":"additional","affiliation":[{"name":"Program of Mathematical Genomics, Department of Systems Biology, Columbia University Irving Medical Center, New York N.Y"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Pranay","family":"Satya","sequence":"additional","affiliation":[{"name":"Program of Mathematical Genomics, Department of Systems Biology, Columbia University Irving Medical Center, New York N.Y"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Raul","family":"Rabadan","sequence":"additional","affiliation":[{"name":"Program of Mathematical Genomics, Department of Systems Biology, Columbia University Irving Medical Center, New York N.Y"},{"name":"Herbert Irving Comprehensive Cancer Center, Columbia University Irving Medical Center, New York N.Y"},{"name":"Department of Biomedical Informatics, Columbia University Irving Medical Center, New York N.Y"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Arnold J.","family":"Levine","sequence":"additional","affiliation":[{"name":"Simons Center for Systems Biology, Institute for Advanced Study, Princeton, N.J"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"54368","reference":[{"key":"2026100109151088000_2026.09.26.754600v1.1","doi-asserted-by":"publisher","DOI":"10.1038\/s41568-020-0262-1"},{"key":"2026100109151088000_2026.09.26.754600v1.2","first-page":"7261","article-title":"Estrogens and progesterone promote persistent CCND1 gene activation during G1 by inducing transcriptional derepression via c-jun\/c-fos\/estrogen receptor (progesterone receptor) complex assembly to a distal regulatory element and recruitment of cyclin D1 to its own gene promoter","volume":"24","year":"2004","journal-title":"Mol. 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