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However, these gibberellin (GA)\u2010insensitive genes require higher water and fertilizer inputs and impair seedling emergence and early vigor, prompting the search for alternative GA\u2010sensitive dwarfing genes to breed environmentally friendly wheat varieties for sustainable wheat production. Here, we reported the map\u2010based cloning of\n                    <jats:italic>Rht4<\/jats:italic>\n                    and functional characterization of its potential causal gene\n                    <jats:italic>TaBSK3\u20102B<\/jats:italic>\n                    , which encodes a plasma membrane\u2010localized serine\/threonine\u2010protein kinase involved in the brassinosteroid (BR) signaling pathway. Loss function of\n                    <jats:italic>TaBSK3\u20102B<\/jats:italic>\n                    attenuated BR responses and inhibited stem elongation by reducing cell expansion, which was accompanied by decreased levels of the bioactive gibberellins GA\n                    <jats:sub>1<\/jats:sub>\n                    and GA\n                    <jats:sub>4<\/jats:sub>\n                    . Under controlled conditions,\n                    <jats:italic>TaBSK3\u20102B<\/jats:italic>\n                    positively regulates wheat drought tolerance by integrating transcriptional level changes of key drought\u2010related genes and physiological adjustments under water deficit. Furthermore, we identified a natural haplotype\n                    <jats:italic>TaBSK3\u20102B<\/jats:italic>\n                    <jats:sup>\n                      <jats:italic>H2<\/jats:italic>\n                    <\/jats:sup>\n                    which was associated with reduced plant height and improved yield\u2010related traits, with the potential to mitigate the yield penalty associated with the original\n                    <jats:italic>Rht4<\/jats:italic>\n                    allele, a major limitation on its application in wheat breeding. The frequency of\n                    <jats:italic>TaBSK3\u20102B<\/jats:italic>\n                    <jats:sup>\n                      <jats:italic>H2<\/jats:italic>\n                    <\/jats:sup>\n                    is dramatically higher in cultivars compared to landraces, indicating a strong artificial selection of\n                    <jats:italic>TaBSK3\u20102B<\/jats:italic>\n                    <jats:sup>\n                      <jats:italic>H2<\/jats:italic>\n                    <\/jats:sup>\n                    during semi\u2010dwarf wheat breeding. Our study not only provides new insights into the synergistic improvement of plant height and grain yield by fine\u2010tuning BR signaling but also offers potential genetic resources for breeding drought\u2010adapted semi\u2010dwarf wheat varieties.\n                  <\/jats:p>","DOI":"10.1111\/tpj.71118","type":"journal-article","created":{"date-parts":[[2026,9,17]],"date-time":"2026-09-17T08:15:23Z","timestamp":1789632923000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Deletion of\n                    <i>TaBSK3\u20102B<\/i>\n                    attenuates\n                    <scp>BR<\/scp>\n                    signaling to regulate the\n                    <i>Rht4<\/i>\n                    semi\u2010dwarf phenotype and drought tolerance in 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