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Here, through a genome\u2010wide association study (GWAS) analysis of a diverse perennial ryegrass population, we identified\n                    <jats:italic>LpbZIP41<\/jats:italic>\n                    as a major heat tolerance quantitative trait locus (QTL) for heat tolerance. Natural variation at this locus comprises three haplotypes showing differential thermotolerance, with evidence of positive selection during breeding. LpbZIP41 localizes to the nucleus and exhibits rapid heat\u2010inducible expression. Functional validation via CRISPR\u2010Cas9\u2010mediated knockout confirmed that\n                    <jats:italic>LpbZIP41<\/jats:italic>\n                    is essential for heat stress. Overexpression of\n                    <jats:italic>LpbZIP41<\/jats:italic>\n                    in perennial ryegrass enhanced thermotolerance, as evidenced by improved survival, reduced lipid peroxidation, maintained membrane integrity, and global transcriptional reprogramming under heat stress. Mechanistically, we uncovered a transcriptional cascade that LpDREB2A directly activates\n                    <jats:italic>LpbZIP41<\/jats:italic>\n                    transcription by binding to dehydration\u2010responsive element (DRE) motifs in the promoter. LpbZIP41, in turn, regulates\n                    <jats:italic>LpNCED4<\/jats:italic>\n                    , a key abscisic acid (ABA) biosynthesis gene, and\n                    <jats:italic>Lp\u03b2CA2<\/jats:italic>\n                    , encoding a stress\u2010responsive carbonic anhydrase. Furthermore, LpbZIP66 functions as a negative regulator that antagonizes LpbZIP41 activity via heterodimerization. Collectively, our findings established an LpDREB2A\u2010LpbZIP41\/LpbZIP66 regulatory module as a central hub that integrates ABA biosynthesis and metabolic signals to confer heat stress tolerance in a cool\u2010season grass.\n                  <\/jats:p>","DOI":"10.1111\/tpj.71059","type":"journal-article","created":{"date-parts":[[2026,8,3]],"date-time":"2026-08-03T04:20:46Z","timestamp":1785730846000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["<scp>LpDREB2A<\/scp>\n                    \u2010\n                    <scp>LpbZIP41<\/scp>\n                    \/\n                    <scp>LpbZIP66<\/scp>\n                    module orchestrates heat stress tolerance by modulating\n                    <scp>ABA<\/scp>\n                    biosynthesis and carbonic anhydrase activity in perennial ryegrass"],"prefix":"10.1111","volume":"127","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5617-8719","authenticated-orcid":false,"given":"Zhengfu","family":"Fang","sequence":"first","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation &amp; Utilization of Horticultural Crops, College of Horticulture and Forestry Sciences Huazhong Agricultural University, Hubei Hongshan Laboratory  Wuhan 430070 Hubei China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jie","family":"Chen","sequence":"additional","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation &amp; Utilization of Horticultural Crops, College of Horticulture and Forestry Sciences Huazhong Agricultural University, Hubei Hongshan Laboratory  Wuhan 430070 Hubei China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Simin","family":"Wu","sequence":"additional","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation &amp; Utilization of Horticultural Crops, College of Horticulture and Forestry Sciences Huazhong Agricultural University, Hubei Hongshan Laboratory  Wuhan 430070 Hubei China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wensong","family":"Zhang","sequence":"additional","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation &amp; Utilization of Horticultural Crops, College of Horticulture and Forestry Sciences Huazhong Agricultural University, Hubei Hongshan Laboratory  Wuhan 430070 Hubei China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0001-7080-0957","authenticated-orcid":false,"given":"Jingya","family":"Lv","sequence":"additional","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation &amp; Utilization of Horticultural Crops, College of Horticulture and Forestry Sciences Huazhong Agricultural University, Hubei Hongshan Laboratory  Wuhan 430070 Hubei 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Huazhong Agricultural University, Hubei Hongshan Laboratory  Wuhan 430070 Hubei China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Weiliang","family":"Wang","sequence":"additional","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation &amp; Utilization of Horticultural Crops, College of Horticulture and Forestry Sciences Huazhong Agricultural University, Hubei Hongshan Laboratory  Wuhan 430070 Hubei China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Guomei","family":"Yin","sequence":"additional","affiliation":[{"name":"Inner Mongolia Academy of Agricultural and Animal Husbandry Sciences  Hohhot Inner Mongolia 750306 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4391-0883","authenticated-orcid":false,"given":"Yanping","family":"Wang","sequence":"additional","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation &amp; Utilization of Horticultural Crops, 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