{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,23]],"date-time":"2026-10-23T20:06:53Z","timestamp":1792786013103,"version":"4.5.0"},"reference-count":53,"publisher":"Oxford University Press (OUP)","license":[{"start":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T00:00:00Z","timestamp":1789430400000},"content-version":"am","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100004837","name":"MCIN","doi-asserted-by":"publisher","id":[{"id":"10.13039\/501100004837","id-type":"DOI","asserted-by":"publisher"}]},{"name":"Agencia Estatal de Investigaci\u00f3n","id":[{"id":"https:\/\/ror.org\/003x0zc53","id-type":"ROR","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100008530","name":"European Regional Development Fund","doi-asserted-by":"publisher","award":["PID2020-115856RB-I00"],"award-info":[{"award-number":["PID2020-115856RB-I00"]}],"id":[{"id":"10.13039\/501100008530","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100008530","name":"European Regional Development Fund","doi-asserted-by":"publisher","award":["PID2023-147220OB-I00"],"award-info":[{"award-number":["PID2023-147220OB-I00"]}],"id":[{"id":"10.13039\/501100008530","id-type":"DOI","asserted-by":"publisher"}]},{"name":"the Gobierno de Arag\u00f3n","award":["A09_17R, A09_23R"],"award-info":[{"award-number":["A09_17R, A09_23R"]}],"id":[{"id":"https:\/\/ror.org\/0425pg203","id-type":"ROR","asserted-by":"crossref"}]},{"name":"the Gobierno de Arag\u00f3n","award":["BES-2017-082913"],"award-info":[{"award-number":["BES-2017-082913"]}],"id":[{"id":"https:\/\/ror.org\/0425pg203","id-type":"ROR","asserted-by":"crossref"}]},{"name":"the Gobierno de Arag\u00f3n","award":["PRE2021-099952"],"award-info":[{"award-number":["PRE2021-099952"]}],"id":[{"id":"https:\/\/ror.org\/0425pg203","id-type":"ROR","asserted-by":"crossref"}]},{"name":"-Mexico","award":["288000\/471811"],"award-info":[{"award-number":["288000\/471811"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>Iron (Fe) deficiency is a widespread disorder limiting global soybean (Glycine max) production. Although root exudation is a key adaptive mechanism for Fe scarcity in species such as Arabidopsis, a detailed chemical characterization of soybean exudates is lacking. Here, we examined the accumulation and secretion of phenolic compounds in soybean roots and their correlation with intraspecific tolerance to Fe-deficiency chlorosis. Seven soybean genotypes with contrasting tolerance, derived from US breeding programs, were analyzed. Root exudates from Fe-deficient soybean plants solubilized ferric oxide. We identified and quantified 28 coumarin-type phenolic compounds, among which catechol methylsideretin was the predominant component. Although the qualitative coumarin profile was consistent across all genotypes, Fe-efficient lines secreted these compounds at higher concentrations or earlier during Fe deficiency than Fe-inefficient lines. The efficient genotype A7 showed coordinated up-regulation of coumarin biosynthesis and secretion, whereas this response was weaker in the Fe-inefficient genotype IsoClark. Catechol methylsideretin concentrations strongly correlated with the ability of root exudates to mobilize Fe from ferric oxide. The conserved phenolics profile, together with divergence from profiles reported in non-legume species, suggests lineage-specific adaptations and ecological roles beyond Fe mobilization. These results highlight genotype-dependent exudation as a determinant of soybean Fe-deficiency tolerance, with implications for breeding.<\/jats:p>","DOI":"10.1093\/jxb\/erag459","type":"journal-article","created":{"date-parts":[[2026,9,15]],"date-time":"2026-09-15T12:06:54Z","timestamp":1789474014000},"source":"Crossref","is-referenced-by-count":0,"title":["Redox-active di-\n                    <i>O<\/i>\n                    -methylated coumarins exudation contributes to genotype-dependent iron deficiency tolerance in soybean"],"prefix":"10.1093","author":[{"ORCID":"https:\/\/orcid.org\/0009-0004-2738-6692","authenticated-orcid":false,"given":"Francisco Jos\u00e9","family":"Jim\u00e9nez-Pastor","sequence":"first","affiliation":[{"name":"Plant Micronutrient Stress Physiology Group, Estaci\u00f3n Experimental de Aula Dei (EEAD), CSIC , Avenida de Monta\u00f1ana 1005, 50059 Zaragoza ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Edgar","family":"Garc\u00eda-Cruz","sequence":"additional","affiliation":[{"name":"Plant Micronutrient Stress Physiology Group, Estaci\u00f3n Experimental de Aula Dei (EEAD), CSIC , Avenida de Monta\u00f1ana 1005, 50059 Zaragoza ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0009-0275-0597","authenticated-orcid":false,"given":"Ra\u00fal","family":"Bouzada-D\u00edaz","sequence":"additional","affiliation":[{"name":"Plant Micronutrient Stress Physiology Group, Estaci\u00f3n Experimental de Aula Dei (EEAD), CSIC , Avenida de Monta\u00f1ana 1005, 50059 Zaragoza ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5470-5901","authenticated-orcid":false,"given":"Javier","family":"Abad\u00eda","sequence":"additional","affiliation":[{"name":"Plant Micronutrient Stress Physiology Group, Estaci\u00f3n Experimental de Aula Dei (EEAD), CSIC , Avenida de Monta\u00f1ana 1005, 50059 Zaragoza ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7967-5136","authenticated-orcid":false,"given":"Jorge","family":"Rodr\u00edguez-Celma","sequence":"additional","affiliation":[{"name":"Plant Micronutrient Stress Physiology Group, Estaci\u00f3n Experimental de Aula Dei (EEAD), CSIC , Avenida de Monta\u00f1ana 1005, 50059 Zaragoza ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4568-1201","authenticated-orcid":false,"given":"Ana","family":"\u00c1lvarez-Fern\u00e1ndez","sequence":"additional","affiliation":[{"name":"Plant Micronutrient Stress Physiology Group, Estaci\u00f3n Experimental de Aula Dei (EEAD), CSIC , Avenida de Monta\u00f1ana 1005, 50059 Zaragoza ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,9,15]]},"reference":[{"key":"2026101312113792300_erag459-B1","doi-asserted-by":"crossref","DOI":"10.3390\/ijms21114038","article-title":"Single and combined Fe and S deficiency differentially modulate root exudate composition in tomato: a double strategy for Fe acquisition?","volume":"21","author":"Astolfi","year":"2020","journal-title":"International Journal of Molecular Sciences"},{"key":"2026101312113792300_erag459-B2","first-page":"27","article-title":"USDA soybean genetic collection: isoline collection","volume":"18","author":"Bernard","year":"1991","journal-title":"Soybean Genetics Newsletter"},{"key":"2026101312113792300_erag459-B3","doi-asserted-by":"crossref","first-page":"196","DOI":"10.1111\/j.1399-3054.1983.tb00757.x","article-title":"Characterization of ferric reducing activity in roots of Fe-deficient Phaseolus vulgaris","volume":"59","author":"Bienfait","year":"1983","journal-title":"Physiologia Plantarum"},{"key":"2026101312113792300_erag459-B4","doi-asserted-by":"crossref","first-page":"887","DOI":"10.2174\/0929867053507315","article-title":"Simple coumarins and analogues in medicinal chemistry: occurrence, synthesis and biological activity","volume":"12","author":"Borges","year":"2005","journal-title":"Current Medicinal Chemistry"},{"key":"2026101312113792300_erag459-B5","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1002\/mas.20058","article-title":"Accurate mass measurement for the determination of elemental formula\u2014a tutorial","volume":"25","author":"Bristow","year":"2006","journal-title":"Mass Spectrometry Reviews"},{"key":"2026101312113792300_erag459-B6","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1097\/00010694-196102000-00008","article-title":"Iron chlorosis in soybeans as related to the genotype of rootstalk: chlorosis susceptibility and reductive capacity at the root","volume":"91","author":"Brown","year":"1961","journal-title":"Soil Science"},{"key":"2026101312113792300_erag459-B7","doi-asserted-by":"crossref","first-page":"311","DOI":"10.2134\/agronj1973.00021962006500020037x","article-title":"Reductants\u201d released by roots of Fe-deficient soybeans","volume":"65","author":"Brown","year":"1973","journal-title":"Agronomy Journal"},{"key":"2026101312113792300_erag459-B8","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1111\/nph.12471","article-title":"Involvement of the ABCG37 transporter in secretion of scopoletin and derivatives by Arabidopsis roots in response to iron deficiency","volume":"201","author":"Fourcroy","year":"2014","journal-title":"New Phytologist"},{"key":"2026101312113792300_erag459-B9","doi-asserted-by":"crossref","DOI":"10.1016\/j.plantsci.2020.110664","article-title":"Iron deficient Medicago scutellata grown in nutrient solution at high pH accumulates and secretes large amounts of flavins","volume":"303","author":"Gheshlaghi","year":"2021","journal-title":"Plant Science"},{"key":"2026101312113792300_erag459-B10","doi-asserted-by":"crossref","first-page":"2113","DOI":"10.1111\/nph.19098","article-title":"Forging a symbiosis: transition metal delivery in symbiotic nitrogen fixation","volume":"239","author":"Gonz\u00e1lez-Guerrero","year":"2023","journal-title":"New Phytologist"},{"key":"2026101312113792300_erag459-B11","doi-asserted-by":"crossref","first-page":"R19","DOI":"10.1186\/gb-2007-8-2-r19","article-title":"Qbase relative quantification framework and software for management and automated analysis of real-time quantitative PCR data","volume":"8","author":"Hellemans","year":"2007","journal-title":"Genome Biology"},{"key":"2026101312113792300_erag459-B12","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1007\/s11104-008-9668-3","article-title":"Global inputs of biological nitrogen fixation in agricultural 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Merr.) for iron deficiency tolerance","volume":"322","author":"Raj","year":"2019","journal-title":"Journal of Radioanalytical and Nuclear Chemistry"},{"key":"2026101312113792300_erag459-B28","doi-asserted-by":"crossref","first-page":"442","DOI":"10.1038\/s41589-018-0019-2","article-title":"Biosynthesis of redox-active metabolites in response to iron deficiency in plants","volume":"14","author":"Rajniak","year":"2018","journal-title":"Nature Chemical Biology"},{"key":"2026101312113792300_erag459-B29","doi-asserted-by":"crossref","first-page":"595","DOI":"10.1038\/s41477-024-01896-5","article-title":"The BRUTUS iron sensor and E3 ligase facilitates soybean root nodulation by monoubiquitination of NSP1","volume":"11","author":"Ren","year":"2025","journal-title":"Nature Plants"},{"key":"2026101312113792300_erag459-B30","doi-asserted-by":"crossref","first-page":"2062","DOI":"10.1111\/nph.17090","article-title":"Coumarin accumulation and trafficking in Arabidopsis thaliana: a complex and dynamic process","volume":"229","author":"Robe","year":"2021","journal-title":"New Phytologist"},{"key":"2026101312113792300_erag459-B31","doi-asserted-by":"crossref","DOI":"10.1016\/j.xplc.2025.101431","article-title":"Coumarin-facilitated iron transport: an IRT1-independent strategy for iron acquisition in Arabidopsis thaliana","volume":"6","author":"Robe","year":"2025","journal-title":"Plant Communications"},{"key":"2026101312113792300_erag459-B32","doi-asserted-by":"crossref","first-page":"694","DOI":"10.1038\/17800","article-title":"A ferric-chelate reductase for iron uptake from soils","volume":"397","author":"Robinson","year":"1999","journal-title":"Nature"},{"key":"2026101312113792300_erag459-B33","doi-asserted-by":"crossref","first-page":"2173","DOI":"10.1093\/pcp\/pcr149","article-title":"Characterization of flavins in roots of Fe-deficient strategy I plants, with a focus on Medicago truncatula","volume":"52","author":"Rodr\u00edguez-Celma","year":"2011","journal-title":"Plant & Cell Physiology"},{"key":"2026101312113792300_erag459-B34","doi-asserted-by":"crossref","first-page":"1473","DOI":"10.1104\/pp.113.220426","article-title":"Mutually exclusive alterations in secondary metabolism are critical for the uptake of insoluble iron compounds by Arabidopsis and Medicago truncatula","volume":"162","author":"Rodr\u00edguez-Celma","year":"2013","journal-title":"Plant Physiology"},{"key":"2026101312113792300_erag459-B35","doi-asserted-by":"crossref","first-page":"949","DOI":"10.1104\/pp.71.4.949","article-title":"Mechanism of iron uptake by peanut plants I. FeIII reduction, chelate slitting, and release of phenolics","volume":"71","author":"R\u00f6mheld","year":"1983","journal-title":"Plant Physiology"},{"key":"2026101312113792300_erag459-B36","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1104\/pp.80.1.175","article-title":"Evidence for a specific uptake system for iron phytosiderophores in roots of grasses","volume":"80","author":"R\u00f6mheld","year":"1986","journal-title":"Plant Physiology"},{"key":"2026101312113792300_erag459-B37","doi-asserted-by":"crossref","first-page":"1072","DOI":"10.1111\/j.1469-8137.2009.02908.x","article-title":"Dissecting iron deficiency-induced proton extrusion in Arabidopsis roots","volume":"183","author":"Santi","year":"2009","journal-title":"New Phytologist"},{"key":"2026101312113792300_erag459-B38","doi-asserted-by":"crossref","first-page":"160","DOI":"10.1104\/pp.113.228544","article-title":"Feruloyl-CoA 6\u2032-hydroxylase1-dependent coumarins mediate iron acquisition from alkaline substrates in Arabidopsis","volume":"164","author":"Schmid","year":"2014","journal-title":"Plant Physiology"},{"key":"2026101312113792300_erag459-B39","doi-asserted-by":"crossref","DOI":"10.1371\/journal.pone.0102444","article-title":"Metabolome analysis of Arabidopsis thaliana roots identifies a key metabolic pathway for iron acquisition","volume":"9","author":"Schmidt","year":"2014","journal-title":"PLoS One"},{"key":"2026101312113792300_erag459-B40","doi-asserted-by":"crossref","DOI":"10.3389\/fpls.2016.01711","article-title":"Accumulation and secretion of coumarinolignans and other coumarins by Arabidopsis thaliana roots in response to iron deficiency at high pH","volume":"7","author":"Sis\u00f3-Terraza","year":"2016","journal-title":"Frontiers in Plant Science"},{"key":"2026101312113792300_erag459-B41","doi-asserted-by":"crossref","first-page":"733","DOI":"10.1111\/nph.13633","article-title":"Flavins secreted by roots of iron-deficient Beta vulgaris enable mining of ferric oxide via reductive mechanisms","volume":"209","author":"Sis\u00f3-Terraza","year":"2016","journal-title":"New Phytologist"},{"key":"2026101312113792300_erag459-B42","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1007\/s11306-007-0082-2","article-title":"Proposed minimum reporting standards for chemical analysis. 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