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A9 also links the regulation of seed maturation with that of early seedling greening through its effects on various light receptors. Transcriptomic analyses of transgenic tobacco seeds suggested that A9 might affect chromatin remodeling. Here, using formaldehyde\u2010assisted isolation of regulatory elements (FAIRE) and micrococcal nuclease digestion, we analyzed the A9 effects on chromatin accessibility shortly after seed imbibition in the proximal promoter region of developmentally relevant genes, including HY5 and PHYA. A9, expressed from a seed\u2010specific promoter, enhanced chromatin accessibility in the analyzed regions. Converse, promoter\u2010specific, effects were observed upon loss\u2010of\u2010function of tobacco A9 (NtA9) in transgenic seeds. Furthermore, a memory effect was observed, as the induced chromatin accessibility persisted for up to 4\u2009days after seed imbibition, when A9 was no longer detected. The A9\u2010induced chromatin effects involved labile\/unstable nucleosomes placed at proximal promoter locations where A9 induced substantial nucleosomal depletion. Specific inhibitors of BRAHMA\u2010like ATPase subunits of SWI\/SNF (SWItch\/Sucrose Non\u2010Fermentable) chromatin\u2010remodeling complexes and histone deacetylase (HDAC) impaired A9\u2010induced memory. Thus, SWI\/SNF remodeling and HDAC activity mechanistically contribute to the A9\u2010induced memory. Furthermore, SWI\/SNF inhibition specifically reduced the HY5 and PHYA promoter accessibility in both transgenic and non\u2010transgenic seeds. Our results identify HSFA9 as a potential master, short\u2010term, \u2018epigenetic\u2019 regulator that operates in seeds in anticipation of seedling establishment. The new, A9\u2010induced, somatic memory effect reported here may facilitate early seedling greening and stress tolerance during the seed\u2010to\u2010seedling developmental transition.<\/jats:p>","DOI":"10.1111\/tpj.70261","type":"journal-article","created":{"date-parts":[[2025,6,6]],"date-time":"2025-06-06T16:59:31Z","timestamp":1749229171000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Sunflower seed\u2010specific <scp>HSFA9<\/scp> induces persistent chromatin accessibility during seed\u2010to\u2010seedling developmental transition"],"prefix":"10.1111","volume":"122","author":[{"given":"Pilar","family":"Prieto\u2010Dapena","sequence":"first","affiliation":[{"name":"Instituto de Recursos Naturales y Agrobiolog\u00eda de Sevilla Consejo Superior de Investigaciones Cient\u00edficas (IRNAS\u2010CSIC)  Sevilla Spain"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Ra\u00fal","family":"Carranco","sequence":"additional","affiliation":[{"name":"Instituto de Recursos Naturales y Agrobiolog\u00eda de Sevilla Consejo Superior de Investigaciones Cient\u00edficas (IRNAS\u2010CSIC)  Sevilla Spain"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Concepci\u00f3n","family":"Almoguera","sequence":"additional","affiliation":[{"name":"Instituto de Recursos Naturales y Agrobiolog\u00eda de Sevilla Consejo Superior de Investigaciones Cient\u00edficas (IRNAS\u2010CSIC)  Sevilla Spain"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2583-4453","authenticated-orcid":false,"given":"Juan","family":"Jordano","sequence":"additional","affiliation":[{"name":"Instituto de Recursos Naturales y Agrobiolog\u00eda de Sevilla Consejo Superior de Investigaciones Cient\u00edficas (IRNAS\u2010CSIC)  Sevilla Spain"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"311","published-online":{"date-parts":[[2025,6,6]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.pbi.2021.102091"},{"key":"e_1_2_9_3_1","doi-asserted-by":"publisher","DOI":"10.3390\/plants9030335"},{"key":"e_1_2_9_4_1","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M207330200"},{"key":"e_1_2_9_5_1","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkw1273"},{"key":"e_1_2_9_6_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/eraa098"},{"key":"e_1_2_9_7_1","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.106.048272"},{"key":"e_1_2_9_8_1","doi-asserted-by":"publisher","DOI":"10.1146\/annurev\u2010arplant\u2010102820\u2010093218"},{"key":"e_1_2_9_9_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1365\u2010313X.2004.02169.x"},{"key":"e_1_2_9_10_1","doi-asserted-by":"publisher","DOI":"10.1111\/pce.13365"},{"key":"e_1_2_9_11_1","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.17061"},{"key":"e_1_2_9_12_1","doi-asserted-by":"publisher","DOI":"10.1038\/nsmb1333"},{"key":"e_1_2_9_13_1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.1014856107"},{"key":"e_1_2_9_14_1","doi-asserted-by":"publisher","DOI":"10.1111\/tpj.15901"},{"key":"e_1_2_9_15_1","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.107.056457"},{"key":"e_1_2_9_16_1","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.109.066845"},{"key":"e_1_2_9_17_1","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.17.00655"},{"key":"e_1_2_9_18_1","doi-asserted-by":"publisher","DOI":"10.1105\/tpc.13.3.599"},{"key":"e_1_2_9_19_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/eraa335"},{"key":"e_1_2_9_20_1","doi-asserted-by":"publisher","DOI":"10.1093\/nar\/gkac826"},{"key":"e_1_2_9_21_1","doi-asserted-by":"publisher","DOI":"10.3389\/fpls.2022.865361"},{"key":"e_1_2_9_22_1","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms1732"},{"key":"e_1_2_9_23_1","doi-asserted-by":"publisher","DOI":"10.1074\/jbc.M114.579474"},{"key":"e_1_2_9_24_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1399\u20103054.2011.01476.x"},{"key":"e_1_2_9_25_1","doi-asserted-by":"publisher","DOI":"10.1093\/jxb\/erp304"},{"key":"e_1_2_9_26_1","doi-asserted-by":"publisher","DOI":"10.1111\/pce.13373"},{"key":"e_1_2_9_27_1","doi-asserted-by":"publisher","DOI":"10.1038\/s41467\u2010021\u201023786\u20106"},{"key":"e_1_2_9_28_1","doi-asserted-by":"publisher","DOI":"10.1093\/plcell\/koad111"},{"key":"e_1_2_9_29_1","doi-asserted-by":"publisher","DOI":"10.1111\/j.1469\u20108137.2011.03818.x"},{"key":"e_1_2_9_30_1","doi-asserted-by":"publisher","DOI":"10.1016\/j.molp.2016.07.002"},{"key":"e_1_2_9_31_1","unstructured":"Gasper M. 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