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Culm strength, mainly determined by lignin biosynthesis and secondary cell wall (SCW) formation, is core to lodging resistance. We identified an ethylene response factor (ERF) transcription factor STRONG3 via genome\u2010wide association study (GWAS) of 405 rice accessions based on lignin content.<\/jats:p>\n                      <\/jats:list-item>\n                      <jats:list-item>\n                        <jats:p>\n                          Functional validation demonstrated that\n                          <jats:italic>STRONG3<\/jats:italic>\n                          positively regulates lignin accumulation, culm diameter, and mechanical strength by promoting sclerenchyma tissue development, vascular bundle enlargement, and SCW thickening.\n                        <\/jats:p>\n                      <\/jats:list-item>\n                      <jats:list-item>\n                        <jats:p>\n                          Molecular analyses revealed that STRONG3 directly binds to the GCC\u2010box motif in the promoter of\n                          <jats:italic>4CL4<\/jats:italic>\n                          , a rate\u2010limiting enzyme in the phenylpropanoid pathway, to activate its transcription. Genetic complementation confirmed that\n                          <jats:italic>4CL4<\/jats:italic>\n                          acts downstream of\n                          <jats:italic>STRONG3<\/jats:italic>\n                          to mediate lignin biosynthesis and culm development. Haplotype analysis identified subspecies\u2010specific favorable alleles:\n                          <jats:italic>STRONG3<\/jats:italic>\n                          <jats:sup>\n                            <jats:italic>G<\/jats:italic>\n                          <\/jats:sup>\n                          in\n                          <jats:italic>Geng<\/jats:italic>\n                          rice, caused by a single nucleotide polymorphism (4349512) leading to an Ala289Thr substitution in the activation domain that enhances STRONG3's transcriptional activation of\n                          <jats:italic>4CL4<\/jats:italic>\n                          , and\n                          <jats:italic>4CL4<\/jats:italic>\n                          <jats:sup>TC<\/jats:sup>\n                          in\n                          <jats:italic>Xian<\/jats:italic>\n                          rice. Combinatorial analysis showed these alleles synergistically improve culm traits in their respective subspecies.\n                        <\/jats:p>\n                      <\/jats:list-item>\n                      <jats:list-item>\n                        <jats:p>\n                          Overexpression of\n                          <jats:italic>STRONG3<\/jats:italic>\n                          enhanced lodging resistance without compromising grain yield. Our findings uncover a critical regulatory pathway for rice culm development and provide valuable genetic resources and molecular markers for breeding high\u2010yield, lodging\u2010resistant rice varieties.\n                        <\/jats:p>\n                      <\/jats:list-item>\n                    <\/jats:list>\n                  <\/jats:p>","DOI":"10.1111\/nph.71574","type":"journal-article","created":{"date-parts":[[2026,10,8]],"date-time":"2026-10-08T02:51:22Z","timestamp":1791427882000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["<i>STRONG3<\/i>\n                    promotes lignin biosynthesis and culm strength in rice"],"prefix":"10.1111","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-7070-798X","authenticated-orcid":false,"given":"Yong","family":"Zhao","sequence":"first","affiliation":[{"name":"Frontiers Science Center for Molecular Design Breeding, Key Laboratory of Crop Heterosis and 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Agricultural University  100193 Beijing China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2955-6307","authenticated-orcid":false,"given":"Lujia","family":"Han","sequence":"additional","affiliation":[{"name":"College of Engineering China Agricultural University  100083 Beijing China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8842-3432","authenticated-orcid":false,"given":"Wensheng","family":"Wang","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Crop Gene Resources and Breeding, Institute of Crop Sciences Chinese Academy of Agricultural Sciences  100081 Beijing China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3186-1132","authenticated-orcid":false,"given":"Zichao","family":"Li","sequence":"additional","affiliation":[{"name":"Frontiers Science Center for Molecular Design Breeding, Key Laboratory of Crop 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