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Although plants have evolved sophisticated mechanisms to enhance Fe uptake, transport, and utilization, key components of the underlying regulatory network remain unclear. Here, we identify the NAM, ATAF1\/2, and CUP\u2010SHAPED COTYLEDON 2 (NAC) transcription factor ANAC017 as an essential regulator of Fe homeostasis in\n                    <jats:italic>Arabidopsis<\/jats:italic>\n                    .\n                    <jats:italic>ANAC017<\/jats:italic>\n                    expression is strongly induced under Fe deficiency, and loss of\n                    <jats:italic>ANAC017<\/jats:italic>\n                    function leads to reduced Fe accumulation in both roots and shoots, impaired root elongation, and pronounced leaf chlorosis. Notably,\n                    <jats:italic>anac017<\/jats:italic>\n                    mutants show excessive Fe retention in the cell wall\u2014particularly within the hemicellulose fraction\u2014under Fe\u2010deficient conditions, suggesting that ANAC017 contributes to the mobilization and reutilization of apoplastic Fe. Mechanistically, ANAC017 functions within the canonical Fe signaling cascade. We show that the core Fe regulator, the BASIC HELIX\u2010LOOP\u2010HELIX (bHLH) transcription factor bHLH121\/URI, directly activates\n                    <jats:italic>ANAC017<\/jats:italic>\n                    transcription, and overexpression of\n                    <jats:italic>ANAC017<\/jats:italic>\n                    partially rescues the Fe\u2010sensitive phenotype of the\n                    <jats:italic>bhlh121\/uri<\/jats:italic>\n                    mutant. Further experiments revealed that the transcription factor ANAC017 binds to the promoter of\n                    <jats:italic>MAT1<\/jats:italic>\n                    \/\n                    <jats:italic>SAM1<\/jats:italic>\n                    (encoding\n                    <jats:italic>S<\/jats:italic>\n                    \u2010adenosylmethionine synthetase), potentially modulating its expression. In conclusion, this study reveals that ANAC017 functions downstream of the bHLH121\/URI pathway to regulate Fe homeostasis via two separate mechanisms: facilitating cell wall Fe reutilization and mediating\n                    <jats:italic>S<\/jats:italic>\n                    \u2010adenosylmethionine synthetase metabolism by regulating\n                    <jats:italic>MAT1\/SAM1<\/jats:italic>\n                    .\n                  <\/jats:p>","DOI":"10.1111\/tpj.71157","type":"journal-article","created":{"date-parts":[[2026,10,15]],"date-time":"2026-10-15T05:55:27Z","timestamp":1792043727000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["A\n                    <scp>bHLH121<\/scp>\n                    \/\n                    <scp>URI<\/scp>\n                    \u2013\n                    <scp>ANAC017<\/scp>\n                    \u2013\n                    <scp>MAT1<\/scp>\n                    \/\n                    <scp>SAM1<\/scp>\n                    regulatory cascade controls iron homeostasis in\n                    <i>Arabidopsis<\/i>"],"prefix":"10.1111","volume":"128","author":[{"given":"Chunyan","family":"Tu","sequence":"first","affiliation":[{"name":"State Key Laboratory of Soil and Sustainable Agriculture, Institute of Soil Science Chinese Academy of Sciences  Nanjing 210008 China"},{"name":"College of Land Resources and Environment Jiangxi Agricultural University  Nanchang Jiangxi 330045 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5100-7585","authenticated-orcid":false,"given":"Qi","family":"Wu","sequence":"additional","affiliation":[{"name":"Tobacco Research Institute Chinese Academy of Agricultural Sciences  Qingdao 266101 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7785-2266","authenticated-orcid":false,"given":"Fangwei","family":"Yu","sequence":"additional","affiliation":[{"name":"Institute of Vegetable Crops Jiangsu Academy of Agricultural Sciences  Nanjing 210014 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Renfang","family":"Shen","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Soil and Sustainable Agriculture, Institute of Soil Science Chinese Academy of Sciences  Nanjing 210008 China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2052-2212","authenticated-orcid":false,"given":"Xiaofang","family":"Zhu","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Soil and Sustainable Agriculture, Institute of Soil Science Chinese Academy of Sciences  Nanjing 210008 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