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Among tested phytohormones, salicylic acid (SA) promoted seedling growth, folate accumulation, and antioxidant capacity. Composite coatings were fabricated using chitosan of varying molecular weights; films formulated with 0.3% medium molecular weight chitosan and 0.1% SA (SA-MCS) exhibited moderate water vapor permeability and sustained release, creating a favorable microenvironment for germination. SA-MCS coatings significantly increased total folates from 569.87 to 702.75 \u03bcg 100 g\u20131 DW. Mechanistically, controlled SA delivery upregulated GTP cyclohydrolase 1 (GCH1) and aminodeoxychorismate synthase (ADCS), accelerating metabolic flux into the pterin and para-aminobenzoic acid (pABA) branches. Multiomics analysis revealed coordinated mobilization of carbon units from histidine catabolism and glutamate into one-carbon metabolism, providing structural backbones and methyl groups for folate enrichment. These findings demonstrate SA-CS coatings as an efficient, biodegradable seed delivery platform for eco-friendly folate biofortification.<\/jats:p>","DOI":"10.1021\/acs.jafc.6c01087","type":"journal-article","created":{"date-parts":[[2026,6,16]],"date-time":"2026-06-16T15:24:44Z","timestamp":1781623484000},"page":"24292-24308","source":"Crossref","is-referenced-by-count":0,"title":["Bioactive Seed\nCoating with Chitosan-Salicylic Acid\nNetworks Activates Folate Metabolism and Nutritional Fortification\nin Wheat Seedlings"],"prefix":"10.1021","volume":"74","author":[{"given":"Muhammad","family":"Bilal","sequence":"first","affiliation":[{"name":"Nanjing Agricultural University , , , ,","place":["Nanjing, Jiangsu, People\u2019s Republic of China, 210095"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Bowen","family":"Qin","sequence":"additional","affiliation":[{"name":"Nanjing Agricultural University , , , ,","place":["Nanjing, Jiangsu, People\u2019s Republic of China, 210095"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yiwei","family":"Wu","sequence":"additional","affiliation":[{"name":"Nanjing Agricultural University , , , ,","place":["Nanjing, Jiangsu, People\u2019s Republic of China, 210095"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-8925-7173","authenticated-orcid":true,"given":"Dandan","family":"Li","sequence":"additional","affiliation":[{"name":"Nanjing Agricultural University , , , ,","place":["Nanjing, Jiangsu, People\u2019s Republic of China, 210095"]},{"name":"The Sanya Institute of Nanjing Agricultural University , ,","place":["Sanya, People\u2019s Republic of China, 572024"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chong","family":"Xie","sequence":"additional","affiliation":[{"name":"Nanjing Agricultural University , , , ,","place":["Nanjing, Jiangsu, People\u2019s Republic of China, 210095"]},{"name":"The Sanya Institute of Nanjing Agricultural University , ,","place":["Sanya, People\u2019s Republic of China, 572024"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0191-6315","authenticated-orcid":true,"given":"Runqiang","family":"Yang","sequence":"additional","affiliation":[{"name":"Nanjing Agricultural University , , , ,","place":["Nanjing, Jiangsu, People\u2019s Republic of China, 210095"]},{"name":"The Sanya Institute of Nanjing Agricultural University , ,","place":["Sanya, People\u2019s Republic of China, 572024"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Dong","family":"Jiang","sequence":"additional","affiliation":[{"name":"Nanjing Agricultural University , , , ,","place":["Nanjing, Jiangsu, People\u2019s Republic of China, 210095"]},{"name":"Nanjing Agricultural University , , , ,","place":["Nanjing, Jiangsu, People\u2019s Republic of China, 210095"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2338-9851","authenticated-orcid":true,"given":"Pei","family":"Wang","sequence":"additional","affiliation":[{"name":"Nanjing Agricultural University , , , ,","place":["Nanjing, Jiangsu, People\u2019s Republic of China, 210095"]},{"name":"The Sanya Institute of Nanjing Agricultural University , ,","place":["Sanya, People\u2019s Republic of China, 572024"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"316","published-online":{"date-parts":[[2026,6,16]]},"reference":[{"issue":"5","key":"2026081208091213800_cit1","doi-asserted-by":"publisher","first-page":"693","DOI":"10.1080\/14735903.2021.1975437","article-title":"Continuing cereals research for sustainable\nhealth and well-being","volume":"20","author":"Poole","year":"2022","journal-title":"Int. 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Cereal Sci."},{"key":"2026081208091213800_cit22","doi-asserted-by":"publisher","first-page":"102306","DOI":"10.1016\/j.fochx.2025.102306","article-title":"Insight into the effect\nof wheatgrass powder on steamed bread properties: Impacts on gluten\npolymerization and starch gelatinization behavior","volume":"26","author":"Bilal","year":"2025","journal-title":"Food Chem. X."},{"issue":"3","key":"2026081208091213800_cit23","doi-asserted-by":"publisher","first-page":"426","DOI":"10.1111\/j.1365-313X.2006.02685.x","article-title":"Evidence for folate-salvage reactions\nin plants","volume":"46","author":"Orsomando","year":"2006","journal-title":"Plant J."},{"issue":"23","key":"2026081208091213800_cit24","doi-asserted-by":"publisher","first-page":"20731","DOI":"10.1074\/jbc.M302894200","article-title":"The folate precursor\np-aminobenzoate\nis reversibly converted to its glucose ester in the plant cytosol","volume":"278","author":"Quinlivan","year":"2003","journal-title":"J. Bio. 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