{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,8]],"date-time":"2026-05-08T05:10:51Z","timestamp":1778217051918,"version":"3.51.4"},"reference-count":71,"publisher":"Frontiers Media SA","license":[{"start":{"date-parts":[[2026,5,8]],"date-time":"2026-05-08T00:00:00Z","timestamp":1778198400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"content-domain":{"domain":["frontiersin.org"],"crossmark-restriction":true},"short-container-title":["Front. Plant Sci."],"abstract":"<jats:sec>\n                    <jats:title>Introduction<\/jats:title>\n                    <jats:p>Basic helix-loop-helix (bHLH) transcription factors play pivotal roles in plant adaptation to metallic elements, yet the mechanisms by which leguminous bHLHs regulate iron (Fe) homeostasis and cadmium (Cd) stress adaptation remain inadequately understood.<\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Methods<\/jats:title>\n                    <jats:p>\n                      MsbHLH60 and its promoter was cloned from alfalfa. The expression of\n                      <jats:italic>MsbHLH60<\/jats:italic>\n                      under iron deficiency (\u2013Fe), Cd, and Cd accompanied by Fe deficiency (Cd(\u2013Fe)) was analyzed by qRT-PCR and GUS assay. MsbHLH60 was subcellular localized. Overexpressing\n                      <jats:italic>MsbHLH60<\/jats:italic>\n                      Arabidopsis was generated and subjected to phenotypic and physiological assays under \u2013Fe, Cd and Cd(\u2013Fe) stress. The interaction was studied through Y2H, Y1H and transient co-expression.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Results<\/jats:title>\n                    <jats:p>\n                      MsbHLH60 belongs to bHLH transcription factor family and was localized to nucleus and cytoplasm.\n                      <jats:italic>MsbHLH60<\/jats:italic>\n                      was strongly expressed in the young tissues of alfalfa, and was significantly upregulated in roots by \u2013Fe or Cd stress, especially in the stele. However,\n                      <jats:italic>MsbHLH60<\/jats:italic>\n                      was not induced by Cd(\u2013Fe). Overexpression of\n                      <jats:italic>MsbHLH60<\/jats:italic>\n                      enhanced Arabidopsis tolerance to \u2013Fe and Cd stress.\n                      <jats:italic>MsbHLH60<\/jats:italic>\n                      mediated Cd tolerance was notably attenuated by \u2013Fe. MsbHLH60 could interact with MsFIT and enhanced the activate effect on\n                      <jats:italic>MsIRT1<\/jats:italic>\n                      expression.\n                    <\/jats:p>\n                  <\/jats:sec>\n                  <jats:sec>\n                    <jats:title>Discussion<\/jats:title>\n                    <jats:p>\n                      The results suggested that\n                      <jats:italic>MsbHLH60<\/jats:italic>\n                      might serve as an auxiliary regulator in the Fe regulation network and might orchestrate plant adaptive response to \u2013Fe and Cd by modulating metal transport and distribution and enhancing antioxidant capacity. This study provides novel insights into the intersecting mechanisms of nutrient homeostasis and metal stress tolerance in plants, laying a theoretical foundation and genetic resources for crop improvement.\n                    <\/jats:p>\n                  <\/jats:sec>","DOI":"10.3389\/fpls.2026.1821873","type":"journal-article","created":{"date-parts":[[2026,5,8]],"date-time":"2026-05-08T04:42:09Z","timestamp":1778215329000},"update-policy":"https:\/\/doi.org\/10.3389\/crossmark-policy","source":"Crossref","is-referenced-by-count":0,"title":["Iron status modulates cadmium tolerance: mechanistic insights from MsbHLH60-mediated regulation of the FIT\/IRT1 module in alfalfa"],"prefix":"10.3389","volume":"17","author":[{"given":"Xin","family":"Liu","sequence":"first","affiliation":[{"name":"Heilongjiang Provincial Key Laboratory of Molecular Cell Genetics and Genetic Breeding, College of Life Science and Technology, Harbin Normal University","place":["Harbin, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Menghan","family":"Chang","sequence":"additional","affiliation":[{"name":"Heilongjiang Provincial Key Laboratory of Molecular Cell Genetics and Genetic Breeding, College of Life Science and Technology, Harbin Normal University","place":["Harbin, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Wenzhe","family":"Du","sequence":"additional","affiliation":[{"name":"Heilongjiang Provincial Key Laboratory of Molecular Cell Genetics and Genetic Breeding, College of Life Science and Technology, Harbin Normal University","place":["Harbin, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Gemeng","family":"Yu","sequence":"additional","affiliation":[{"name":"Heilongjiang Provincial Key Laboratory of Molecular Cell Genetics and Genetic Breeding, College of Life Science and Technology, Harbin Normal University","place":["Harbin, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yuzhe","family":"Yang","sequence":"additional","affiliation":[{"name":"Heilongjiang Provincial Key Laboratory of Molecular Cell Genetics and Genetic Breeding, College of Life Science and Technology, Harbin Normal University","place":["Harbin, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Liang","family":"Si","sequence":"additional","affiliation":[{"name":"Heilongjiang Provincial Key Laboratory of Molecular Cell Genetics and Genetic Breeding, College of Life Science and Technology, Harbin Normal University","place":["Harbin, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Changhong","family":"Guo","sequence":"additional","affiliation":[{"name":"Heilongjiang Provincial Key Laboratory of Molecular Cell Genetics and Genetic Breeding, College of Life Science and Technology, Harbin Normal University","place":["Harbin, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Yingdong","family":"Bi","sequence":"additional","affiliation":[{"name":"Institute of Crops Tillage and Cultivation, Heilongjiang Academy of Agricultural Sciences","place":["Harbin, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Donglin","family":"Guo","sequence":"additional","affiliation":[{"name":"Heilongjiang Provincial Key Laboratory of Molecular Cell Genetics and Genetic Breeding, College of Life Science and Technology, Harbin Normal University","place":["Harbin, China"]}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"1965","published-online":{"date-parts":[[2026,5,8]]},"reference":[{"key":"B1","doi-asserted-by":"publisher","first-page":"528","DOI":"10.1104\/pp.114.246124","article-title":"Radial transport of nutrients: The plant root as a polarized epithelium","volume":"166","author":"Barberon","year":"2014","journal-title":"Plant Physiol."},{"key":"B2","doi-asserted-by":"publisher","first-page":"1733","DOI":"10.1016\/j.molp.2023.09.010","article-title":"TBtools-II: A \u201cone for all, all for one\u201d bioinformatics platform for biological big-data mining","volume":"16","author":"Chen","year":"2023","journal-title":"Mol. 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