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Despite this agricultural importance, the molecular basis underlying boron homeostasis in plants remains unclear. To identify molecular players involved in boron homeostasis in maize (Zea mays L.), we measured boron levels in the Goodman-Buckler association panel and performed genome-wide association studies. These analyses identified a benzoxazinless (bx) gene, bx3, involved in the biosynthesis of benzoxazinoids, such as 2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3-one (DIMBOA), which are major defense compounds in maize. Genes involved in DIMBOA biosynthesis are all located in close proximity in the genome, and benzoxazinoid biosynthesis mutants, including bx3, are all DIMBOA deficient. We determined that leaves of the bx3 mutant have a greater boron concentration than those of B73 control plants, which corresponded with enhanced leaf tip necrosis, a phenotype associated with boron toxicity. By contrast, other DIMBOA-deficient maize mutants did not show altered boron levels or the leaf tip necrosis phenotype, suggesting that boron is not associated with DIMBOA. Instead, our analyses suggest that the accumulation of boron is linked to the benzoxazinoid intermediates indolin-2-one (ION) and 3-hydroxy-ION. Therefore, our results connect boron homeostasis to the benzoxazinoid plant defense pathway through bx3 and specific intermediates, rendering the benzoxazinoid biosynthesis pathway a potential target for crop improvement under inadequate boron conditions.<\/jats:p>","DOI":"10.1093\/plphys\/kiae611","type":"journal-article","created":{"date-parts":[[2024,11,8]],"date-time":"2024-11-08T15:14:38Z","timestamp":1731078878000},"source":"Crossref","is-referenced-by-count":3,"title":["Association of the benzoxazinoid pathway with boron homeostasis in maize"],"prefix":"10.1093","volume":"197","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7405-8326","authenticated-orcid":false,"given":"Liuyang","family":"Chu","sequence":"first","affiliation":[{"name":"Institute of Crop Science and Resource Conservation, Crop Functional Genomics, University of Bonn , Friedrich-Ebert-Allee 144, Bonn 53113 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Freising 85354 ,","place":["Germany"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1025-9484","authenticated-orcid":false,"given":"Henning","family":"Jessen","sequence":"additional","affiliation":[{"name":"Institute of Organic Chemistry, University of Freiburg , Albertstr. 21, Freiburg im Breisgau 79104 ,","place":["Germany"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9022-4515","authenticated-orcid":false,"given":"Gabriel","family":"Schaaf","sequence":"additional","affiliation":[{"name":"Institute of Crop Science and Resource Conservation, Plant Nutrition, University of Bonn , Karl-Robert-Kreiten Stra\u00dfe 13, Bonn 53115 ,","place":["Germany"]}],"role":[{"role":"author","vocabulary":"crossref"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5155-0884","authenticated-orcid":false,"given":"Frank","family":"Hochholdinger","sequence":"additional","affiliation":[{"name":"Institute of Crop 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