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Using mutagenesis of endogenous Sidekick, we showed that its enrichment at apical tricellular junctions and its function in cell rearrangement require its fifth and sixth immunoglobulin domains, but not the first four, although these mediate homophilic adhesion of mammalian Sidekick homologues. The C-terminal PDZ-binding motif of Sidekick contributes to localizing both Sidekick and its intracellular binding partner Canoe to tricellular adherens junctions. We found that the PAK4 homologue Mushroom bodies tiny also binds to the cytoplasmic domain of Sidekick. Its kinase activity is necessary for Sidekick accumulation at tricellular junctions, and over-activity mislocalizes Sidekick along bicellular junctions. However, mutating predicted Mushroom bodies tiny phosphorylation sites in Sidekick itself did not affect its localization. Sidekick localizes to the dendrites of T4 and T5 neurons independently of its extracellular and PDZ-binding domains, and of Mushroom bodies tiny. Our findings reveal an important role for a PAK4 family member in establishing specialized structures at tricellular adherens junctions.<\/jats:p>","DOI":"10.1242\/dev.204777","type":"journal-article","created":{"date-parts":[[2025,9,1]],"date-time":"2025-09-01T13:45:52Z","timestamp":1756734352000},"source":"Crossref","is-referenced-by-count":4,"title":["Mushroom bodies tiny regulates Sidekick localization to tricellular adherens junctions"],"prefix":"10.1242","volume":"152","author":[{"ORCID":"https:\/\/orcid.org\/0009-0005-1915-5429","authenticated-orcid":false,"given":"Dhaval","family":"Gandhi","sequence":"first","affiliation":[{"name":"New York University Grossman School of Medicine 1 Department of Cell Biology , , 540 First Avenue, New York, NY 10016 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0002-8844-9648","authenticated-orcid":false,"given":"Brian P.","family":"Griffin","sequence":"additional","affiliation":[{"name":"New York University Grossman School of Medicine 1 Department of Cell Biology , , 540 First Avenue, New York, NY 10016 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5754-0970","authenticated-orcid":false,"given":"Maria E.","family":"Bustillo","sequence":"additional","affiliation":[{"name":"New York University Grossman School of Medicine 1 Department of Cell Biology , , 540 First Avenue, New York, NY 10016 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-5725-1973","authenticated-orcid":false,"given":"Fleur","family":"Chelemen","sequence":"additional","affiliation":[{"name":"Institut de Biologia Molecular de Barcelona, CSIC, Parc Cient\u00edfic de Barcelona 2 , Baldiri Reixac, 10-12, Barcelona 08028 ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0000-8442-1037","authenticated-orcid":false,"given":"Genie","family":"Jang","sequence":"additional","affiliation":[{"name":"New York University Grossman School of Medicine 1 Department of Cell Biology , , 540 First Avenue, New York, NY 10016 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4451-1609","authenticated-orcid":false,"given":"Ariel","family":"Hairston","sequence":"additional","affiliation":[{"name":"New York University Grossman School of Medicine 1 Department of Cell Biology , , 540 First Avenue, New York, NY 10016 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1309-4398","authenticated-orcid":false,"given":"Marta","family":"Llimargas","sequence":"additional","affiliation":[{"name":"Institut de Biologia Molecular de Barcelona, CSIC, Parc Cient\u00edfic de Barcelona 2 , Baldiri Reixac, 10-12, Barcelona 08028 ,","place":["Spain"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7453-107X","authenticated-orcid":false,"given":"Jessica E.","family":"Treisman","sequence":"additional","affiliation":[{"name":"New York University Grossman School of Medicine 1 Department of Cell Biology , , 540 First Avenue, New York, NY 10016 ,","place":["USA"]}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","published-online":{"date-parts":[[2025,9,18]]},"reference":[{"key":"2025091807531558600_DEV204777C1","doi-asserted-by":"publisher","first-page":"dev158246","DOI":"10.1242\/dev.158246","article-title":"Drosophila Sidekick is required in developing photoreceptors to enable visual motion detection","volume":"145","author":"Astigarraga","year":"2018","journal-title":"Development"},{"key":"2025091807531558600_DEV204777C2","doi-asserted-by":"publisher","first-page":"5315","DOI":"10.1038\/s41467-021-25011-w","article-title":"Proximity proteomics identifies PAK4 as a component of Afadin-Nectin junctions","volume":"12","author":"Baskaran","year":"2021","journal-title":"Nat. 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