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Phenotypic analysis revealed that the\n                    <jats:italic>zik4<\/jats:italic>\n                    mutant had stronger tolerance to phosphate starvation than that to the wild type. However, the kinase domain was still expressed in\n                    <jats:italic>zik4<\/jats:italic>\n                    . The CRISPR\/Cas9 system was used to construct two independent knockout mutants\n                    <jats:italic>zik4-c3<\/jats:italic>\n                    and\n                    <jats:italic>zik4-c15<\/jats:italic>\n                    by targeting two different sites in the AtZIK4 kinase domain. Physiological measurements showed that the phosphate content in\n                    <jats:italic>zik4-c3<\/jats:italic>\n                    and\n                    <jats:italic>zik4-c15<\/jats:italic>\n                    was lower than that in the wild type. Expression pattern analysis revealed that ZIK4 was induced to express under phosphate starvation. Quantitative real-time PCR results showed that the expression levels of multiple key phosphate uptake-related genes were\u00a0significantly up-regulated in the\n                    <jats:italic>zik4<\/jats:italic>\n                    mutant. These results indicate that ZIK4, as a phosphate starvation-induced MAPKKK, promotes phosphate accumulation in Arabidopsis seedlings by upregulating the expression levels of phosphate uptake-related genes. These findings not only elucidate the regulatory role of AtZIK4 kinase during abiotic stress and expand the low-phosphate (LP) signaling cascade in\n                    <jats:italic>Arabidopsis thaliana<\/jats:italic>\n                    , but also providing promising genetic targets to enhance crop performance under phosphorus-limited conditions.\n                  <\/jats:p>","DOI":"10.1515\/biol-2025-1361","type":"journal-article","created":{"date-parts":[[2026,8,26]],"date-time":"2026-08-26T16:03:30Z","timestamp":1787760210000},"source":"Crossref","is-referenced-by-count":0,"title":["AtZIK4, a MAPK kinase kinase, positively regulates phosphate uptake gene expression to modulate phosphate starvation tolerance in\n                    <i>Arabidopsis<\/i>"],"prefix":"10.1515","volume":"21","author":[{"given":"Hongying","family":"Deng","sequence":"first","affiliation":[{"name":"Hebei Center for Industrial Energy-Saving and Pollution Control Research, Hebei Vocational University of Technology and Engineering , Xingtai , Hebei , 054000 , China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Qian","family":"Yang","sequence":"additional","affiliation":[{"name":"Hebei Center for Industrial Energy-Saving and Pollution Control Research, Hebei Vocational University of Technology and Engineering , Xingtai , Hebei , 054000 , China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shirong","family":"Zhao","sequence":"additional","affiliation":[{"name":"Hebei Center for Industrial Energy-Saving and Pollution Control Research, Hebei Vocational University of Technology and Engineering , Xingtai , Hebei , 054000 , China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xuyang","family":"Lei","sequence":"additional","affiliation":[{"name":"Hebei Center for Industrial Energy-Saving and Pollution Control Research, Hebei Vocational University of Technology and Engineering , Xingtai , Hebei , 054000 , China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Liye","family":"Zhu","sequence":"additional","affiliation":[{"name":"School of Medicine, Nankai University , Tianjin , 300071 , China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"374","published-online":{"date-parts":[[2026,8,27]]},"reference":[{"key":"2026082616032695642_j_biol-2025-1361_ref_001","doi-asserted-by":"crossref","unstructured":"Jia, X, Wang, L, Nussaume, L, Yi, K. 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