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This study integrated expression genome\u2010wide association study (eGWAS) across 115\n                    <jats:italic>Arabidopsis thaliana<\/jats:italic>\n                    ecotypes with differential methylome profiling to identify these regulators. Focusing on the memory genes\n                    <jats:italic>LKR<\/jats:italic>\n                    ,\n                    <jats:italic>HIS1\u20103<\/jats:italic>\n                    , and\n                    <jats:italic>DREB1A<\/jats:italic>\n                    , eGWAS identified signaling and epigenetic loci involved in ABA\/JA responses and DNA methylation. Methylome profiling by whole\u2010genome bisulfite sequencing of ecotypes contrasting in drought tolerance, and in the superinduction of memory genes revealed significantly greater methylation variation during the second drought (D2) than during the first (D1), highlighting the role of epigenetic reprogramming in memory maintenance. Functional validation using T\u2010DNA mutants demonstrated specific modulation of the D2\/D1 induction ratio without affecting initial drought responses. Mutants of\n                    <jats:italic>LKR<\/jats:italic>\n                    eGWAS\u2010delineated genes AT1G56660, AT2G19120, AT4G16490, and\n                    <jats:italic>DEG3<\/jats:italic>\n                    , those of\n                    <jats:italic>HIS1\u20103<\/jats:italic>\n                    eGWAS genes AT1G14220, AT2G24960, AT3G10845, AT3G19340,\n                    <jats:italic>CNGC10<\/jats:italic>\n                    ,\n                    <jats:italic>EMB2770<\/jats:italic>\n                    ,\n                    <jats:italic>GRF7<\/jats:italic>\n                    , and\n                    <jats:italic>RPP2A<\/jats:italic>\n                    , and\n                    <jats:italic>DREB1A<\/jats:italic>\n                    eGWAS genes AT1G67000, AT3G61610, AT5G62110,\n                    <jats:italic>HK2<\/jats:italic>\n                    ,\n                    <jats:italic>JMJ12<\/jats:italic>\n                    , and\n                    <jats:italic>LUP5<\/jats:italic>\n                    abolished respective memory gene induction. The eGWAS and methylome approaches converged on DNA repair, chromatin modification, vesicular transport, and proteostasis as core memory hubs. These findings reveal a genetic\u2010epigenetic interplay that coordinates transcriptional memory, priming plants for rapid reactivation of stress pathways during recurrent drought.\n                  <\/jats:p>","DOI":"10.1111\/pce.70914","type":"journal-article","created":{"date-parts":[[2026,9,23]],"date-time":"2026-09-23T05:50:30Z","timestamp":1790142630000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Expression GWAS and Differential Methylome Profiling Reveal Upstream Regulators of Drought Memory Genes in\n                    <i>Arabidopsis thaliana<\/i>"],"prefix":"10.1111","author":[{"given":"Debankona","family":"Marik","sequence":"first","affiliation":[{"name":"Department of Bioscience and Bioengineering Indian Institute of Technology Jodhpur Jodhpur Rajasthan India"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-2831-2382","authenticated-orcid":false,"given":"Rishabh","family":"Kumar","sequence":"additional","affiliation":[{"name":"Department of Bioscience and Bioengineering Indian Institute of Technology Jodhpur Jodhpur Rajasthan India"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-6745-749X","authenticated-orcid":false,"given":"Ayan","family":"Sadhukhan","sequence":"additional","affiliation":[{"name":"Department of Bioscience and Bioengineering Indian Institute of Technology Jodhpur Jodhpur Rajasthan India"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"311","published-online":{"date-parts":[[2026,9,22]]},"reference":[{"key":"e_1_2_9_2_1","doi-asserted-by":"crossref","first-page":"34","DOI":"10.1016\/j.envexpbot.2017.08.004","article-title":"Analysis of DNA Methylation Patterns Associated With Drought Stress Response in Faba Bean (Vicia faba L.) 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