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Therefore, creating diverse fruit morphologies and elucidating their regulatory networks are major objectives in tomato breeding. Currently, although multiple genes regulating fruit shape (such as\n                    <jats:italic>SUN<\/jats:italic>\n                    ,\n                    <jats:italic>OVATE<\/jats:italic>\n                    ,\n                    <jats:italic>FAS<\/jats:italic>\n                    , etc.) have been identified, the molecular network governing tomato fruit shape remains incomplete.\n                    <jats:italic>SHAGGY\u2010like kinases<\/jats:italic>\n                    (\n                    <jats:italic>SKs<\/jats:italic>\n                    )\n                    <jats:italic>\/Glycogen synthase kinase3<\/jats:italic>\n                    (\n                    <jats:italic>GSK3<\/jats:italic>\n                    ) are evolutionarily conserved regulators of plant development and have been implicated in diverse processes. However, the biological functions of this family in tomato fruit development remain to be systematically elucidated. In this study, we performed systematic CRISPR\/Cas9 mutagenesis of all nine members of the tomato\n                    <jats:italic>SHAGGY\u2010like kinases<\/jats:italic>\n                    family (\n                    <jats:italic>SlSKs<\/jats:italic>\n                    ), generating higher order mutants across different clades. Phenotypic screening revealed that\n                    <jats:italic>SlSKs<\/jats:italic>\n                    clade II (\n                    <jats:italic>SlSK2s<\/jats:italic>\n                    ) members (\n                    <jats:italic>SlSK21<\/jats:italic>\n                    \/\n                    <jats:italic>SlSK22<\/jats:italic>\n                    \/\n                    <jats:italic>SlSK23<\/jats:italic>\n                    ) redundantly regulate fruit shape by acting as negative regulators, controlling cell expansion in the columella and modulating ovary wall cell division patterns during ovary development. Proteomics analysis revealed a significant reduction of the microtubule\u2010associated protein SlMAP65\u20101 in the\n                    <jats:italic>slsk2s<\/jats:italic>\n                    mutant lines. Our results further confirmed that SlMAP65\u20101 interacts with SlSK21, SlSK22, and SlSK23, respectively,\n                    <jats:italic>in planta<\/jats:italic>\n                    . Phosphorylation assays demonstrated that SlSK23 directly phosphorylates SlMAP65\u20101 at Ser504 and Ser532, thereby enhancing its protein stability. These results shed new light on the molecular mechanism by which\n                    <jats:italic>SlSK2s<\/jats:italic>\n                    modulate fruit shape in tomato.\n                  <\/jats:p>","DOI":"10.1111\/tpj.70918","type":"journal-article","created":{"date-parts":[[2026,5,18]],"date-time":"2026-05-18T06:00:43Z","timestamp":1779084043000},"update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Clade\n                    <scp>II SHAGGY<\/scp>\n                    \u2010like kinases regulate fruit shape in tomato by modulating\n                    <scp>SlMAP65<\/scp>\n                    \u20101 stability"],"prefix":"10.1111","volume":"126","author":[{"given":"Rong","family":"Huang","sequence":"first","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation &amp; Utilization of Horticultural Crops Huazhong Agricultural University  Wuhan 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Innovation &amp; Utilization of Horticultural Crops Huazhong Agricultural University  Wuhan China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Wenran","family":"Su","sequence":"additional","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation &amp; Utilization of Horticultural Crops Huazhong Agricultural University  Wuhan China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jiajun","family":"Ran","sequence":"additional","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation &amp; Utilization of Horticultural Crops Huazhong Agricultural University  Wuhan China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lei","family":"Song","sequence":"additional","affiliation":[{"name":"National Key Laboratory for Germplasm Innovation &amp; Utilization of Horticultural Crops Huazhong Agricultural University  Wuhan 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