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However, the precise mechanism by which DMI3 activates this pathway remains unclear. Here, we show that phosphorylation of a triplet motif (S343S344T345) located at the C-terminus of the calmodulin-binding domain acts as a molecular switch. We identified T345 as a phosphorylation site that exhibits regulatory functions similar to those of S343 and S344. Phosphomimic mutations at any single residue of the 3 sites blocked DMI3-dependent nodulation, indicating that the kinase must initially remain nonphosphorylated at these positions. Conversely, simultaneous phosphorylation of at least 2 residues was required for symbiotic activity, revealing a transition from a phosphorylation-free state to a hyperphosphorylated status (simultaneous phosphorylation of at least 2 residues) at these sites. Phosphomimic mutations within the triplet enhanced DMI3 autophosphorylation (auto-P) but severely impaired its interaction with Ca2+\/CaM and IPD3. In addition, the physical interaction between DMI3 and IPD3 suppressed IPD3-triggered MtNIN transcription and nodulation, indicating that DMI3 also acts as a negative regulator by sequestering activated IPD3. Thus, we propose a model in which repeated auto-P at the triplet contributes to RNS activation by dissociating the CaM-DMI3-IPD3 complex, thereby releasing phosphorylated and activated IPD3 to initiate the downstream transcriptional cascade. Our findings uncover a dual role for DMI3\u2014as both a kinase and a scaffold\u2014and clarify how auto-P within the triplet motif licenses IPD3 to trigger RNS.<\/jats:p>","DOI":"10.1093\/plphys\/kiag347","type":"journal-article","created":{"date-parts":[[2026,6,4]],"date-time":"2026-06-04T11:44:14Z","timestamp":1780573454000},"source":"Crossref","is-referenced-by-count":0,"title":["Autophosphorylation at the C-terminus of the DMI3 calmodulin-binding domain dismantles CaM-DMI3-IPD3 to initiate root nodulation"],"prefix":"10.1093","volume":"202","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-5637-4445","authenticated-orcid":false,"given":"Yiting","family":"Zhou","sequence":"first","affiliation":[{"name":"College of Life and Environmental Sciences, Hangzhou Normal University , Hangzhou ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yuchen","family":"Qian","sequence":"additional","affiliation":[{"name":"College of Life and Environmental Sciences, Hangzhou Normal University , Hangzhou ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-0685-3409","authenticated-orcid":false,"given":"Yiang","family":"Wang","sequence":"additional","affiliation":[{"name":"College of Life and Environmental Sciences, Hangzhou Normal University , Hangzhou ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0008-1668-4511","authenticated-orcid":false,"given":"Xiang","family":"Li","sequence":"additional","affiliation":[{"name":"College of Life and Environmental Sciences, Hangzhou Normal University , Hangzhou ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yingying","family":"Huang","sequence":"additional","affiliation":[{"name":"College of Life and Environmental Sciences, Hangzhou Normal University , Hangzhou ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0008-8370-9635","authenticated-orcid":false,"given":"Liqiang","family":"Dai","sequence":"additional","affiliation":[{"name":"College of Life and Environmental Sciences, Hangzhou Normal University , Hangzhou ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0002-1173-8193","authenticated-orcid":false,"given":"Lihui","family":"Duan","sequence":"additional","affiliation":[{"name":"College of Life and Environmental Sciences, Hangzhou Normal University , Hangzhou ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-8384-488X","authenticated-orcid":false,"given":"Qian","family":"Guo","sequence":"additional","affiliation":[{"name":"College of Life and Environmental Sciences, Hangzhou Normal University , Hangzhou ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0007-8064-2984","authenticated-orcid":false,"given":"Kedi","family":"Jiang","sequence":"additional","affiliation":[{"name":"College of Life and Environmental Sciences, Hangzhou Normal University , Hangzhou ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Shang","family":"Wang","sequence":"additional","affiliation":[{"name":"College of Life and Environmental Sciences, Hangzhou Normal University , Hangzhou ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0412-5586","authenticated-orcid":false,"given":"Jinbo","family":"Shen","sequence":"additional","affiliation":[{"name":"State Key Laboratory of Subtropical Silviculture, Zhejiang A & F University , Hangzhou ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1251-5530","authenticated-orcid":false,"given":"Liqun","family":"Du","sequence":"additional","affiliation":[{"name":"College of Life and Environmental Sciences, Hangzhou Normal University , Hangzhou ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4722-0997","authenticated-orcid":false,"given":"Erxu","family":"Pi","sequence":"additional","affiliation":[{"name":"College of Life and Environmental Sciences, Hangzhou Normal University , Hangzhou ,","place":["China"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2026,6,9]]},"reference":[{"key":"2026101203574532900_kiag347-B1","doi-asserted-by":"publisher","first-page":"493","DOI":"10.1038\/s41586-024-07487-w","article-title":"Accurate structure prediction of biomolecular interactions with AlphaFold 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