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Previous studies lack insights into chloroplast-specific Cd detoxification mechanisms and metabolic regulators governing metal homeostasis. This study investigated contrasting Cd responses between two wheat genotypes: Cd-tolerant \u201cH97\u201d and Cd-sensitive \u201cL193\u201d. Compared to Cd-compromised \u201cL193\u201d, \u201cH97\u201d manifested through mitigated leaf chlorosis, optimized root architecture and photosynthetic capability, reduced reactive oxygen species accumulation, and lower grain Cd content. Moreover, \u201cH97\u201d exhibited significantly lower Cd accumulation within chloroplasts. Comparative transcriptomics and functional analyses identified TaHMA1-7B, a chloroplast-localized heavy metal ATPase, as a pivotal chloroplast Cd exporter, and its overexpression significantly enhanced rice tolerance against Cd toxicity. Metabolomic profiling coupled with exogenous application revealed that indole-3-carboxylic acid (ICA) conferred dose-dependent protection against Cd toxicity. Mechanistically, ICA significantly reduced leaf Cd concentrations and induced TaHMA1-7B expression. Collectively, our findings establish a chloroplast-centric detoxification pathway in which ICA potentially activates TaHMA1-7B-mediated Cd export from chloroplasts, thereby enhancing Cd tolerance.<\/jats:p>","DOI":"10.1021\/acs.jafc.6c09893","type":"journal-article","created":{"date-parts":[[2026,9,18]],"date-time":"2026-09-18T14:48:18Z","timestamp":1789742898000},"page":"30445-30462","source":"Crossref","is-referenced-by-count":0,"title":["Integrated Transcriptomics,\nMetabolomics, and Functional\nAnalyses Uncover\n                    <i>TaHma1-7b<\/i>\n                    and Indole-3-carboxylic\nAcid as Key Regulators of Wheat Cadmium Detoxification"],"prefix":"10.1021","volume":"74","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5730-3737","authenticated-orcid":true,"given":"Yingpeng","family":"Hua","sequence":"first","affiliation":[{"name":"Zhengzhou University , , ,","place":["Zhengzhou, China, 450001"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xuan","family":"Wang","sequence":"additional","affiliation":[{"name":"Zhengzhou University , , ,","place":["Zhengzhou, China, 450001"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rui","family":"Cui","sequence":"additional","affiliation":[{"name":"Zhengzhou University , , ,","place":["Zhengzhou, China, 450001"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yingna","family":"Feng","sequence":"additional","affiliation":[{"name":"Zhengzhou University , , ,","place":["Zhengzhou, China, 450001"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Hao","family":"Li","sequence":"additional","affiliation":[{"name":"Henan Academy of Agricultural Sciences , , ,","place":["Zhengzhou, China, 450002"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wenxu","family":"Li","sequence":"additional","affiliation":[{"name":"Henan Academy of Agricultural Sciences , , ,","place":["Zhengzhou, China, 450002"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jinna","family":"Hou","sequence":"additional","affiliation":[{"name":"Henan Academy of Agricultural Sciences , , ,","place":["Zhengzhou, China, 450002"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yunhong","family":"Zhang","sequence":"additional","affiliation":[{"name":"Henan Academy of Agricultural Sciences , , ,","place":["Zhengzhou, China, 450002"]},{"name":"Ministry of Agriculture and Rural Affairs of the People\u2019s Republic of China , , ,","place":["Xinxiang, China, 453514"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhensheng","family":"Lei","sequence":"additional","affiliation":[{"name":"Henan Academy of Agricultural Sciences , , ,","place":["Zhengzhou, China, 450002"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ting","family":"Zhou","sequence":"additional","affiliation":[{"name":"Zhengzhou University , , ,","place":["Zhengzhou, China, 450001"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Zhengfu","family":"Zhou","sequence":"additional","affiliation":[{"name":"Henan Academy of Agricultural Sciences , , ,","place":["Zhengzhou, China, 450002"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,9,18]]},"reference":[{"issue":"4","key":"2026093009081261700_cit1","doi-asserted-by":"publisher","first-page":"44","DOI":"10.1007\/s44307-024-00052-6","article-title":"Defense guard: strategies of plants in the fight against cadmium\nstress","volume":"2","author":"Zhang","year":"2024","journal-title":"Adv. Biotechnol."},{"issue":"1","key":"2026093009081261700_cit2","doi-asserted-by":"publisher","first-page":"1","DOI":"10.1146\/annurev-arplant-062923-021424","article-title":"Metal transport\nsystems in plants","volume":"75","author":"Huang","year":"2024","journal-title":"Annu. Rev. Plant Biol."},{"issue":"5","key":"2026093009081261700_cit3","doi-asserted-by":"publisher","DOI":"10.1111\/ppl.14544","article-title":"Accelerating wheat improvement through\ntrait characterization: advances and perspectives","volume":"176","author":"Roychowdhury","year":"2024","journal-title":"Physiol. 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Agric. Food Chem."},{"issue":"3","key":"2026093009081261700_cit26","doi-asserted-by":"publisher","first-page":"54","DOI":"10.1007\/s00122-024-04560-w","article-title":"Pectin demethylation-mediated\ncell wall Na+ retention positively regulates salt stress\ntolerance in oilseed\nrape","volume":"137","author":"Zhou","year":"2024","journal-title":"Theor. Appl. Genet."},{"key":"2026093009081261700_cit27","doi-asserted-by":"publisher","first-page":"205","DOI":"10.1007\/s11104-017-3204-2","article-title":"Genome-scale\nmRNA\ntranscriptomic insights into the responses of oilseed rape (Brassica napus L.) to varying boron availabilities","volume":"416","author":"Hua","year":"2017","journal-title":"Plant Soil"},{"issue":"1","key":"2026093009081261700_cit28","doi-asserted-by":"publisher","first-page":"245","DOI":"10.1104\/pp.121.1.245","article-title":"Cytokinins in tobacco\nand wheat chloroplasts.\nOccurrence and changes due to light\/dark treatment","volume":"121","author":"Benkova\u0301","year":"1999","journal-title":"Plant Physiol."},{"key":"2026093009081261700_cit29","doi-asserted-by":"publisher","first-page":"402","DOI":"10.1006\/meth.2001.1262","article-title":"Analysis\nof relative gene expression data using real-time\nquantitative PCR and the 2-\u0394\u0394CT method","volume":"25","author":"Livak","year":"2001","journal-title":"Methods"},{"issue":"6","key":"2026093009081261700_cit30","doi-asserted-by":"publisher","first-page":"2796","DOI":"10.1038\/nprot.2006.469","article-title":"A protocol for Agrobacterium-mediated\ntransformation in rice","volume":"1","author":"Nishimura","year":"2006","journal-title":"Nat. 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Seedlings\nto Cadmium Chloride stress and Freeze\u2013Thaw Cycles through Protocatechuic\nAcid Application","volume":"236","author":"Ali","year":"2025","journal-title":"Water Air Soil Pollut."},{"key":"2026093009081261700_cit35","first-page":"F4.3.1","article-title":"Chlorophylls\nand carotenoids: measurement and characterization by UV\u2013VIS\nspectroscopy","volume-title":"Current protocols in food\nanalytical chemistry","author":"Lichtenthaler","year":"2001"},{"issue":"4","key":"2026093009081261700_cit36","doi-asserted-by":"publisher","first-page":"1112","DOI":"10.1111\/pce.13456","article-title":"SpHMA1 is a chloroplast cadmium exporter protecting photochemical\nreactions in the Cd hyperaccumulator Sedum plumbizincicola","volume":"42","author":"Zhao","year":"2019","journal-title":"Plant, Cell Environ."},{"key":"2026093009081261700_cit37","doi-asserted-by":"publisher","first-page":"132480","DOI":"10.1016\/j.jhazmat.2023.132480","article-title":"CaHMA1 promotes\nCd accumulation in pepper fruit","volume":"460","author":"Xu","year":"2023","journal-title":"J. Hazard.\nMater."},{"key":"2026093009081261700_cit38","doi-asserted-by":"publisher","first-page":"136766","DOI":"10.1016\/j.jhazmat.2024.136766","article-title":"Chloroplast-localized\ntransporter OsNTP1 mediates cadmium transport from root to shoot and\nsugar metabolism in rice","volume":"484","author":"Wang","year":"2025","journal-title":"J. Hazard. Mater."},{"key":"2026093009081261700_cit39","doi-asserted-by":"publisher","first-page":"224","DOI":"10.1186\/1471-2229-12-224","article-title":"Auxin as\na player in the biocontrol of Fusarium head blight disease of barley\nand its potential as a disease control agent","volume":"12","author":"Petti","year":"2012","journal-title":"BMC Plant Biol."},{"issue":"3","key":"2026093009081261700_cit40","doi-asserted-by":"publisher","first-page":"518","DOI":"10.1111\/tpj.14045","article-title":"Starch degradation,\nabscisic acid and vesicular trafficking are important elements in\ncallose priming by indole-3-carboxylic acid in response to Plectosphaerella cucumerina infection","volume":"96","author":"Gamir","year":"2018","journal-title":"Plant J."}],"container-title":["Journal of Agricultural\nand Food Chemistry"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/pubs.acs.org\/doi\/pdf\/10.1021\/acs.jafc.6c09893","content-type":"application\/pdf","content-version":"vor","intended-application":"unspecified"},{"URL":"https:\/\/pubs.acs.org\/jafcau\/article-pdf\/74\/38\/30445\/68557357\/acs.jafc.6c09893.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/pubs.acs.org\/jafcau\/article-pdf\/74\/38\/30445\/68557357\/acs.jafc.6c09893.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T13:14:46Z","timestamp":1790774086000},"score":1,"resource":{"primary":{"URL":"https:\/\/pubs.acs.org\/jafcau\/article\/74\/38\/30445\/5432858\/Integrated-Transcriptomics-Metabolomics-and"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,9,18]]},"references-count":40,"journal-issue":{"issue":"38","published-online":{"date-parts":[[2026,9,18]]},"published-print":{"date-parts":[[2026,9,30]]}},"URL":"https:\/\/doi.org\/10.1021\/acs.jafc.6c09893","relation":{},"ISSN":["0021-8561","1520-5118"],"issn-type":[{"value":"0021-8561","type":"print"},{"value":"1520-5118","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,9,18]]}}}