{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,6]],"date-time":"2026-07-06T15:13:21Z","timestamp":1783350801867,"version":"3.54.6"},"reference-count":220,"publisher":"MDPI AG","issue":"8","license":[{"start":{"date-parts":[[2019,7,29]],"date-time":"2019-07-29T00:00:00Z","timestamp":1564358400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100002426","name":"Fondazione Telethon","doi-asserted-by":"publisher","award":["GGP16003"],"award-info":[{"award-number":["GGP16003"]}],"id":[{"id":"10.13039\/501100002426","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100005010","name":"Associazione Italiana per la Ricerca sul Cancro","doi-asserted-by":"publisher","award":["IG 2017-20148"],"award-info":[{"award-number":["IG 2017-20148"]}],"id":[{"id":"10.13039\/501100005010","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Cells"],"abstract":"<jats:p>The primary cilium has gone from being a vestigial organelle to a crucial signaling hub of growing interest given the association between a group of human disorders, collectively known as ciliopathies, and defects in its structure or function. In recent years many ciliogenesis proteins have been observed at extraciliary sites in cells and likely perform cilium-independent functions ranging from regulation of the cytoskeleton to vesicular trafficking. Perhaps the most striking example is the non-ciliated T lymphocyte, in which components of the ciliary machinery are repurposed for the assembly and function of the immunological synapse even in the absence of a primary cilium. Furthermore, the specialization traits described at the immunological synapse are similar to those seen in the primary cilium. Here, we review common regulators and features shared by the immunological synapse and the primary cilium that document the remarkable homology between these structures.<\/jats:p>","DOI":"10.3390\/cells8080789","type":"journal-article","created":{"date-parts":[[2019,7,29]],"date-time":"2019-07-29T11:20:18Z","timestamp":1564399218000},"page":"789","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":49,"title":["A Ciliary View of the Immunological Synapse"],"prefix":"10.3390","volume":"8","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-4853-2298","authenticated-orcid":false,"given":"Chiara","family":"Cassioli","sequence":"first","affiliation":[{"name":"Department of Life Sciences, University of Siena, 53100 Siena, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Cosima T.","family":"Baldari","sequence":"additional","affiliation":[{"name":"Department of Life Sciences, University of Siena, 53100 Siena, Italy"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2019,7,29]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"1023","DOI":"10.1158\/2326-6066.CIR-14-0161","article-title":"The immunological synapse","volume":"2","author":"Dustin","year":"2014","journal-title":"Cancer Immunol. Res."},{"key":"ref_2","doi-asserted-by":"crossref","unstructured":"Ortega-Carrion, A., and Vicente-Manzanares, M. (2016). Concerning immune synapses: A spatiotemporal timeline. F1000Research, 5.","DOI":"10.12688\/f1000research.7796.1"},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1111\/imr.12017","article-title":"How the immune system talks to itself: The varied role of synapses","volume":"251","author":"Xie","year":"2013","journal-title":"Immunol. Rev."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"a002311","DOI":"10.1101\/cshperspect.a002311","article-title":"Understanding the structure and function of the immunological synapse","volume":"2","author":"Dustin","year":"2010","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"ref_5","doi-asserted-by":"crossref","first-page":"a002873","DOI":"10.1101\/cshperspect.a002873","article-title":"Modular design of immunological synapses and kinapses","volume":"1","author":"Dustin","year":"2009","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"911","DOI":"10.1038\/ni836","article-title":"Staging and resetting T cell activation in SMACs","volume":"3","author":"Freiberg","year":"2002","journal-title":"Nat. Immunol."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"82","DOI":"10.1038\/25764","article-title":"Three-dimensional segregation of supramolecular activation clusters in T cells","volume":"395","author":"Monks","year":"1998","journal-title":"Nature"},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"691","DOI":"10.1016\/S1074-7613(01)00231-X","article-title":"Exclusion of CD43 from the immunological synapse is mediated by phosphorylation-regulated relocation of the cytoskeletal adaptor moesin","volume":"15","author":"Delon","year":"2001","journal-title":"Immunity"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"10138","DOI":"10.1073\/pnas.97.18.10138","article-title":"A supramolecular basis for CD45 tyrosine phosphatase regulation in sustained T cell activation","volume":"97","author":"Johnson","year":"2000","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_10","first-page":"213","article-title":"Centriole formation and ciliogenesis","volume":"11","author":"Sorokin","year":"1968","journal-title":"Aspen Emphysema Conf."},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"709","DOI":"10.1083\/jcb.151.3.709","article-title":"Chlamydomonas IFT88 and its mouse homologue, polycystic kidney disease gene tg737, are required for assembly of cilia and flagella","volume":"151","author":"Pazour","year":"2000","journal-title":"J. Cell Biol."},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"533","DOI":"10.1038\/nrm.2017.60","article-title":"Genes and molecular pathways underpinning ciliopathies","volume":"18","author":"Reiter","year":"2017","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"693","DOI":"10.1083\/jcb.201411070","article-title":"Centrin2 regulates CP110 removal in primary cilium formation","volume":"208","author":"Prosser","year":"2015","journal-title":"J. Cell Biol."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"3239","DOI":"10.1016\/j.cub.2015.10.028","article-title":"Mother Centriole Distal Appendages Mediate Centrosome Docking at the Immunological Synapse and Reveal Mechanistic Parallels with Ciliogenesis","volume":"25","author":"Stinchcombe","year":"2015","journal-title":"Curr. Biol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1098\/rspb.1976.0086","article-title":"The mechanism of T cell mediated cytotoxicity. III. Changes in target cell susceptibility during the cell cycle","volume":"194","author":"Sanderson","year":"1976","journal-title":"Proc. R. Soc. Lond. B Biol. Sci."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"3713","DOI":"10.1083\/jcb.201610039","article-title":"Microtubule stabilization drives 3D centrosome migration to initiate primary ciliogenesis","volume":"216","author":"Pitaval","year":"2017","journal-title":"J. Cell Biol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"399","DOI":"10.1083\/jcb.201002027","article-title":"The immunological synapse: A focal point for endocytosis and exocytosis","volume":"189","author":"Griffiths","year":"2010","journal-title":"J. Cell Biol."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1002\/cm.970050302","article-title":"Analysis of the morphology and function of primary cilia in connective tissues: A cellular cybernetic probe?","volume":"5","author":"Poole","year":"1985","journal-title":"Cell Motil."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"2762","DOI":"10.4049\/jimmunol.133.5.2762","article-title":"Reorientation of the microtubule-organizing center and the Golgi apparatus in cloned cytotoxic lymphocytes triggered by binding to lysable target cells","volume":"133","author":"Kupfer","year":"1984","journal-title":"J. Immunol."},{"key":"ref_20","doi-asserted-by":"crossref","unstructured":"Follit, J.A., San Agustin, J.T., Xu, F., Jonassen, J.A., Samtani, R., Lo, C.W., and Pazour, G.J. (2008). The Golgin GMAP210\/TRIP11 anchors IFT20 to the Golgi complex. PLoS Genet., 4.","DOI":"10.1371\/journal.pgen.1000315"},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"1332","DOI":"10.1038\/ncb1977","article-title":"Intraflagellar transport is required for polarized recycling of the TCR\/CD3 complex to the immune synapse","volume":"11","author":"Finetti","year":"2009","journal-title":"Nat. Cell Biol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"1110","DOI":"10.1242\/jcs.200006","article-title":"The T cell IFT20 interactome reveals new players in immune synapse assembly","volume":"130","author":"Galgano","year":"2017","journal-title":"J. Cell Sci."},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"241","DOI":"10.1016\/j.tcb.2016.10.005","article-title":"Mechanical Communication at the Immunological Synapse","volume":"27","author":"Basu","year":"2017","journal-title":"Trends Cell Biol."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"298","DOI":"10.1016\/j.coi.2006.03.011","article-title":"Immune synapses in T-cell activation","volume":"18","author":"Cemerski","year":"2006","journal-title":"Curr. Opin. Immunol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"2658","DOI":"10.1038\/s41467-018-05077-9","article-title":"Priming of dendritic cells by DNA-containing extracellular vesicles from activated T cells through antigen-driven contacts","volume":"9","author":"Torralba","year":"2018","journal-title":"Nat. Commun."},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"629","DOI":"10.1126\/science.1124534","article-title":"The primary cilium as the cell\u2019s antenna: Signaling at a sensory organelle","volume":"313","author":"Singla","year":"2006","journal-title":"Science"},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"8","DOI":"10.3389\/fcell.2018.00008","article-title":"Signaling through the Primary Cilium","volume":"6","author":"Wheway","year":"2018","journal-title":"Front. Cell Dev. Biol."},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"117","DOI":"10.1016\/j.immuni.2006.04.010","article-title":"T cell receptor-proximal signals are sustained in peripheral microclusters and terminated in the central supramolecular activation cluster","volume":"25","author":"Varma","year":"2006","journal-title":"Immunity"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"531","DOI":"10.1016\/j.immuni.2010.04.005","article-title":"Essential role of ubiquitin and TSG101 protein in formation and function of the central supramolecular activation cluster","volume":"32","author":"Vardhana","year":"2010","journal-title":"Immunity"},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"110","DOI":"10.3389\/fcell.2019.00110","article-title":"Orchestration of Immunological Synapse Assembly by Vesicular Trafficking","volume":"7","author":"Onnis","year":"2019","journal-title":"Front. Cell Dev. Biol."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"6709","DOI":"10.4049\/jimmunol.172.11.6709","article-title":"Recruitment of transferrin receptor to immunological synapse in response to TCR engagement","volume":"172","author":"Batista","year":"2004","journal-title":"J. Immunol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"577","DOI":"10.1016\/S1074-7613(04)00106-2","article-title":"Activation-induced polarized recycling targets T cell antigen receptors to the immunological synapse; involvement of SNARE complexes","volume":"20","author":"Das","year":"2004","journal-title":"Immunity"},{"key":"ref_33","first-page":"1924","article-title":"Specific recycling receptors are targeted to the immune synapse by the intraflagellar transport system","volume":"127","author":"Finetti","year":"2014","journal-title":"J. Cell Sci."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"1009","DOI":"10.1242\/jcs.00968","article-title":"Dynamic recruitment of the adaptor protein LAT: LAT exists in two distinct intracellular pools and controls its own recruitment","volume":"117","author":"Bonello","year":"2004","journal-title":"J. Cell Sci."},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"809","DOI":"10.1016\/S1074-7613(02)00481-8","article-title":"Dynamics of p56lck translocation to the T cell immunological synapse following agonist and antagonist stimulation","volume":"17","author":"Ehrlich","year":"2002","journal-title":"Immunity"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"1387","DOI":"10.1038\/ncb2888","article-title":"The roles of evolutionarily conserved functional modules in cilia-related trafficking","volume":"15","author":"Sung","year":"2013","journal-title":"Nat. Cell Biol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1016\/j.ceb.2012.10.011","article-title":"The ciliary pocket","volume":"25","author":"Benmerah","year":"2013","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"784","DOI":"10.1016\/j.tibs.2016.06.002","article-title":"Endocytic Control of Cellular Signaling at the Primary Cilium","volume":"41","author":"Pedersen","year":"2016","journal-title":"Trends Biochem. Sci."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"1880","DOI":"10.12688\/f1000research.11944.1","article-title":"Transcellular communication at the immunological synapse: A vesicular traffic-mediated mutual exchange","volume":"6","author":"Finetti","year":"2017","journal-title":"F1000Research"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"449","DOI":"10.1007\/s10571-016-0345-4","article-title":"Ciliary Extracellular Vesicles: Txt Msg Organelles","volume":"36","author":"Wang","year":"2016","journal-title":"Cell Mol. Neurobiol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"2823","DOI":"10.1091\/mbc.E15-01-0009","article-title":"Myristoylated CIL-7 regulates ciliary extracellular vesicle biogenesis","volume":"26","author":"Maguire","year":"2015","journal-title":"Mol. Biol. Cell"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"519","DOI":"10.1016\/j.cub.2014.01.002","article-title":"C. elegans ciliated sensory neurons release extracellular vesicles that function in animal communication","volume":"24","author":"Wang","year":"2014","journal-title":"Curr. Biol."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"252.e214","DOI":"10.1016\/j.cell.2016.11.036","article-title":"An Actin Network Dispatches Ciliary GPCRs into Extracellular Vesicles to Modulate Signaling","volume":"168","author":"Nager","year":"2017","journal-title":"Cell"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"118","DOI":"10.1038\/nature12951","article-title":"Polarized release of T-cell-receptor-enriched microvesicles at the immunological synapse","volume":"507","author":"Choudhuri","year":"2014","journal-title":"Nature"},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1274","DOI":"10.4049\/jimmunol.163.3.1274","article-title":"Activated human T cells release bioactive Fas ligand and APO2 ligand in microvesicles","volume":"163","author":"Anel","year":"1999","journal-title":"J. Immunol."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"282","DOI":"10.1038\/ncomms1285","article-title":"Unidirectional transfer of microRNA-loaded exosomes from T cells to antigen-presenting cells","volume":"2","author":"Mittelbrunn","year":"2011","journal-title":"Nat. Commun."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"1521","DOI":"10.1007\/s00018-017-2740-5","article-title":"Primary cilia proteins: Ciliary and extraciliary sites and functions","volume":"75","author":"Hua","year":"2018","journal-title":"Cell Mol. Life Sci."},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"4140","DOI":"10.1038\/emboj.2012.242","article-title":"End-binding protein 1 controls signal propagation from the T cell receptor","volume":"31","author":"Baixauli","year":"2012","journal-title":"EMBO J."},{"key":"ref_49","doi-asserted-by":"crossref","first-page":"506","DOI":"10.1016\/j.ceb.2013.04.011","article-title":"Assembling a primary cilium","volume":"25","author":"Kim","year":"2013","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"76","DOI":"10.1080\/21541248.2017.1353847","article-title":"Rab GTPases in cilium formation and function","volume":"9","author":"Blacque","year":"2018","journal-title":"Small GTPases"},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"2864","DOI":"10.1038\/s41467-019-10891-w","article-title":"The golgin GMAP210 organizes the transfer of signaling vesicles to the immune synapse: Parallel between vesicular trafficking at the immune synapse and at the cilium","volume":"10","author":"Zucchetti","year":"2019","journal-title":"Nat. Commun."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"779","DOI":"10.1083\/jcb.201301004","article-title":"Centrosome repositioning in T cells is biphasic and driven by microtubule end-on capture-shrinkage","volume":"202","author":"Yi","year":"2013","journal-title":"J. Cell Biol."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"389","DOI":"10.1016\/S1074-7613(02)00421-1","article-title":"In the immune synapse, ZAP-70 controls T cell polarization and recruitment of signaling proteins but not formation of the synaptic pattern","volume":"17","author":"Blanchard","year":"2002","journal-title":"Immunity"},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"860","DOI":"10.4049\/jimmunol.171.2.860","article-title":"Linker for activation of T cells, zeta-associated protein-70, and Src homology 2 domain-containing leukocyte protein-76 are required for TCR-induced microtubule-organizing center polarization","volume":"171","author":"Kuhne","year":"2003","journal-title":"J. Immunol."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"861","DOI":"10.1083\/jcb.140.4.861","article-title":"Cytoskeletal polarization of T cells is regulated by an immunoreceptor tyrosine-based activation motif-dependent mechanism","volume":"140","author":"Shapiro","year":"1998","journal-title":"J. Cell Biol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"4201","DOI":"10.4049\/jimmunol.176.7.4201","article-title":"Role of Fyn in the rearrangement of tubulin cytoskeleton induced through TCR","volume":"176","author":"Sancho","year":"2006","journal-title":"J. Immunol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"663","DOI":"10.1083\/jcb.201008140","article-title":"Centrosome docking at the immunological synapse is controlled by Lck signaling","volume":"192","author":"Tsun","year":"2011","journal-title":"J. Cell Biol."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"647","DOI":"10.1038\/ni.2033","article-title":"A cascade of protein kinase C isozymes promotes cytoskeletal polarization in T cells","volume":"12","author":"Quann","year":"2011","journal-title":"Nat. Immunol."},{"key":"ref_59","doi-asserted-by":"crossref","first-page":"627","DOI":"10.1038\/ni.1734","article-title":"Localized diacylglycerol drives the polarization of the microtubule-organizing center in T cells","volume":"10","author":"Quann","year":"2009","journal-title":"Nat. Immunol."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"14883","DOI":"10.1073\/pnas.0600914103","article-title":"Recruitment of dynein to the Jurkat immunological synapse","volume":"103","author":"Combs","year":"2006","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"1815","DOI":"10.15252\/embj.201488147","article-title":"NuMA interacts with phosphoinositides and links the mitotic spindle with the plasma membrane","volume":"33","author":"Kotak","year":"2014","journal-title":"EMBO J."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"447","DOI":"10.1016\/S0092-8674(00)81365-3","article-title":"A complex of NuMA and cytoplasmic dynein is essential for mitotic spindle assembly","volume":"87","author":"Merdes","year":"1996","journal-title":"Cell"},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"2301","DOI":"10.1038\/emboj.2010.127","article-title":"Ezrin tunes T-cell activation by controlling Dlg1 and microtubule positioning at the immunological synapse","volume":"29","author":"Lasserre","year":"2010","journal-title":"EMBO J."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"5421","DOI":"10.4049\/jimmunol.1200242","article-title":"Stathmin regulates microtubule dynamics and microtubule organizing center polarization in activated T cells","volume":"188","author":"Filbert","year":"2012","journal-title":"J. Immunol."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"781","DOI":"10.1083\/jcb.201106025","article-title":"Casein kinase I delta controls centrosome positioning during T cell activation","volume":"195","author":"Zyss","year":"2011","journal-title":"J. Cell Biol."},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"1629","DOI":"10.1091\/mbc.e13-10-0598","article-title":"Casein kinase 1delta functions at the centrosome and Golgi to promote ciliogenesis","volume":"25","author":"Greer","year":"2014","journal-title":"Mol. Biol. Cell"},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"2539","DOI":"10.1242\/jcs.085852","article-title":"EB1 and EB3 promote cilia biogenesis by several centrosome-related mechanisms","volume":"124","author":"Schroder","year":"2011","journal-title":"J. Cell Sci."},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"483","DOI":"10.1006\/cbir.1997.0177","article-title":"Confocal analysis of primary cilia structure and colocalization with the Golgi apparatus in chondrocytes and aortic smooth muscle cells","volume":"21","author":"Poole","year":"1997","journal-title":"Cell Biol. Int."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"1733","DOI":"10.1242\/bio.014787","article-title":"The primary cilium is a self-adaptable, integrating nexus for mechanical stimuli and cellular signaling","volume":"4","author":"Nguyen","year":"2015","journal-title":"Biol. Open"},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"1351","DOI":"10.1016\/j.cell.2007.04.035","article-title":"HEF1-dependent Aurora A activation induces disassembly of the primary cilium","volume":"129","author":"Pugacheva","year":"2007","journal-title":"Cell"},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"21517","DOI":"10.1073\/pnas.1013728107","article-title":"The major alpha-tubulin K40 acetyltransferase alphaTAT1 promotes rapid ciliogenesis and efficient mechanosensation","volume":"107","author":"Shida","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"1305","DOI":"10.1242\/jcs.180885","article-title":"HDAC6 regulates the dynamics of lytic granules in cytotoxic T lymphocytes","volume":"129","author":"Iborra","year":"2016","journal-title":"J. Cell Sci."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"417","DOI":"10.1016\/S1074-7613(04)00078-0","article-title":"HDAC6 deacetylase activity links the tubulin cytoskeleton with immune synapse organization","volume":"20","author":"Serrador","year":"2004","journal-title":"Immunity"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"1025","DOI":"10.1083\/jcb.201202137","article-title":"INF2 promotes the formation of detyrosinated microtubules necessary for centrosome reorientation in T cells","volume":"198","author":"Anton","year":"2012","journal-title":"J. Cell Biol."},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"1217","DOI":"10.1242\/jcs.199042","article-title":"Microtubule-associated protein-4 controls nanovesicle dynamics and T cell activation","volume":"130","year":"2017","journal-title":"J. Cell Sci."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"951","DOI":"10.1083\/jcb.200801014","article-title":"MTOC translocation modulates IS formation and controls sustained T cell signaling","volume":"182","author":"Cabrero","year":"2008","journal-title":"J. Cell Biol."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"919","DOI":"10.1016\/j.immuni.2011.05.012","article-title":"Dynein-driven transport of T cell receptor microclusters regulates immune synapse formation and T cell activation","volume":"34","author":"Yokosuka","year":"2011","journal-title":"Immunity"},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"17447","DOI":"10.1038\/s41598-018-35593-z","article-title":"CLIP-170 is essential for MTOC repositioning during T cell activation by regulating dynein localisation on the cell surface","volume":"8","author":"Lim","year":"2018","journal-title":"Sci. Rep."},{"key":"ref_79","doi-asserted-by":"crossref","first-page":"1175","DOI":"10.15252\/embj.201593071","article-title":"Tyrosination of alpha-tubulin controls the initiation of processive dynein-dynactin motility","volume":"35","author":"McKenney","year":"2016","journal-title":"EMBO J."},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"321","DOI":"10.1083\/jcb.200707181","article-title":"Cep164, a novel centriole appendage protein required for primary cilium formation","volume":"179","author":"Graser","year":"2007","journal-title":"J. Cell Biol."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"517","DOI":"10.1038\/ncb1251","article-title":"Odf2-deficient mother centrioles lack distal\/subdistal appendages and the ability to generate primary cilia","volume":"7","author":"Ishikawa","year":"2005","journal-title":"Nat. Cell Biol."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"5987","DOI":"10.1073\/pnas.1220927110","article-title":"CCDC41 is required for ciliary vesicle docking to the mother centriole","volume":"110","author":"Joo","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"437","DOI":"10.1016\/0014-4835(88)90054-1","article-title":"Immunogold localization of actin in developing photoreceptor cilia of normal and rds mutant mice","volume":"47","author":"Chaitin","year":"1988","journal-title":"Exp. Eye Res."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"1521","DOI":"10.15252\/embr.201643846","article-title":"The ciliary membrane-associated proteome reveals actin-binding proteins as key components of cilia","volume":"18","author":"Kohli","year":"2017","journal-title":"EMBO Rep."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1083\/jcb.201004003","article-title":"Cell shape and contractility regulate ciliogenesis in cell cycle-arrested cells","volume":"191","author":"Pitaval","year":"2010","journal-title":"J. Cell Biol."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"1048","DOI":"10.1038\/nature08895","article-title":"Functional genomic screen for modulators of ciliogenesis and cilium length","volume":"464","author":"Kim","year":"2010","journal-title":"Nature"},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"2716","DOI":"10.1242\/jcs.043794","article-title":"Nesprin-2 interacts with meckelin and mediates ciliogenesis via remodelling of the actin cytoskeleton","volume":"122","author":"Dawe","year":"2009","journal-title":"J. Cell Sci."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1038\/s41556-017-0018-7","article-title":"Myosin-Va is required for preciliary vesicle transportation to the mother centriole during ciliogenesis","volume":"20","author":"Wu","year":"2018","journal-title":"Nat. Cell Biol."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"12917","DOI":"10.1038\/srep12917","article-title":"Deacetylation of alpha-tubulin and cortactin is required for HDAC6 to trigger ciliary disassembly","volume":"5","author":"Ran","year":"2015","journal-title":"Sci. Rep."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"264.e215","DOI":"10.1016\/j.cell.2016.12.032","article-title":"Dynamic Remodeling of Membrane Composition Drives Cell Cycle through Primary Cilia Excision","volume":"168","author":"Phua","year":"2017","journal-title":"Cell"},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"3255","DOI":"10.1083\/jcb.201703196","article-title":"Actin polymerization controls cilia-mediated signaling","volume":"217","author":"Drummond","year":"2018","journal-title":"J. Cell Biol."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"23","DOI":"10.1038\/76877","article-title":"The immunological synapse and the actin cytoskeleton: Molecular hardware for T cell signaling","volume":"1","author":"Dustin","year":"2000","journal-title":"Nat. Immunol."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"864","DOI":"10.1016\/j.immuni.2015.04.013","article-title":"Actin depletion initiates events leading to granule secretion at the immunological synapse","volume":"42","author":"Ritter","year":"2015","journal-title":"Immunity"},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"E6585","DOI":"10.1073\/pnas.1710751114","article-title":"Cortical actin recovery at the immunological synapse leads to termination of lytic granule secretion in cytotoxic T lymphocytes","volume":"114","author":"Ritter","year":"2017","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_95","first-page":"427.e424","article-title":"PIP5 Kinases Regulate Membrane Phosphoinositide and Actin Composition for Targeted Granule Secretion by Cytotoxic Lymphocytes","volume":"49","author":"Frazer","year":"2018","journal-title":"Immunity"},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"775","DOI":"10.1083\/jcb.201201018","article-title":"F-actin polymerization and retrograde flow drive sustained PLCgamma1 signaling during T cell activation","volume":"197","author":"Babich","year":"2012","journal-title":"J. Cell Biol."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"834","DOI":"10.1091\/mbc.e11-08-0731","article-title":"Actin retrograde flow and actomyosin II arc contraction drive receptor cluster dynamics at the immunological synapse in Jurkat T cells","volume":"23","author":"Yi","year":"2012","journal-title":"Mol. Biol. Cell"},{"key":"ref_98","doi-asserted-by":"crossref","first-page":"68","DOI":"10.3389\/fimmu.2016.00068","article-title":"Action and Traction: Cytoskeletal Control of Receptor Triggering at the Immunological Synapse","volume":"7","author":"Comrie","year":"2016","journal-title":"Front. Immunol."},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"1658","DOI":"10.4049\/jimmunol.173.3.1658","article-title":"Differential roles for Wiskott-Aldrich syndrome protein in immune synapse formation and IL-2 production","volume":"173","author":"Cannon","year":"2004","journal-title":"J. Immunol."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1016\/j.cub.2005.11.036","article-title":"The WAVE2 complex regulates actin cytoskeletal reorganization and CRAC-mediated calcium entry during T cell activation","volume":"16","author":"Nolz","year":"2006","journal-title":"Curr. Biol."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"5986","DOI":"10.1128\/MCB.00136-07","article-title":"WAVE2 regulates high-affinity integrin binding by recruiting vinculin and talin to the immunological synapse","volume":"27","author":"Nolz","year":"2007","journal-title":"Mol. Cell Biol."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"eaal3118","DOI":"10.1126\/science.aal3118","article-title":"Visualizing dynamic microvillar search and stabilization during ligand detection by T cells","volume":"356","author":"Cai","year":"2017","journal-title":"Science"},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"e04953","DOI":"10.7554\/eLife.04953","article-title":"Actin foci facilitate activation of the phospholipase C-gamma in primary T lymphocytes via the WASP pathway","volume":"4","author":"Kumari","year":"2015","journal-title":"eLife"},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"383","DOI":"10.1083\/jcb.201603080","article-title":"Formin-generated actomyosin arcs propel T cell receptor microcluster movement at the immune synapse","volume":"215","author":"Murugesan","year":"2016","journal-title":"J. Cell Biol."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"177","DOI":"10.1016\/j.immuni.2007.01.008","article-title":"Formins regulate the actin-related protein 2\/3 complex-independent polarization of the centrosome to the immunological synapse","volume":"26","author":"Gomez","year":"2007","journal-title":"Immunity"},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"1517","DOI":"10.1073\/pnas.1424829112","article-title":"Force-dependent transition in the T-cell receptor beta-subunit allosterically regulates peptide discrimination and pMHC bond lifetime","volume":"112","author":"Das","year":"2015","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"E8204","DOI":"10.1073\/pnas.1703559114","article-title":"Mechanosensing drives acuity of alphabeta T-cell recognition","volume":"114","author":"Feng","year":"2017","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"31028","DOI":"10.1074\/jbc.M109.052712","article-title":"The alphabeta T cell receptor is an anisotropic mechanosensor","volume":"284","author":"Kim","year":"2009","journal-title":"J. Biol. Chem."},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1016\/j.cell.2014.02.053","article-title":"Accumulation of dynamic catch bonds between TCR and agonist peptide-MHC triggers T cell signaling","volume":"157","author":"Liu","year":"2014","journal-title":"Cell"},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"990","DOI":"10.1038\/35050128","article-title":"Co-assembly of polycystin-1 and -2 produces unique cation-permeable currents","volume":"408","author":"Hanaoka","year":"2000","journal-title":"Nature"},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"4141","DOI":"10.1242\/jcs.087007","article-title":"Septins at a glance","volume":"124","author":"Beise","year":"2011","journal-title":"J. Cell Sci."},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1038\/nrm3284","article-title":"Septins: The fourth component of the cytoskeleton","volume":"13","author":"Mostowy","year":"2012","journal-title":"Nat. Rev. Mol. Cell Biol."},{"key":"ref_113","first-page":"1476","article-title":"Sept7b is essential for pronephric function and development of left-right asymmetry in zebrafish embryogenesis","volume":"127","author":"Dash","year":"2014","journal-title":"J. Cell Sci."},{"key":"ref_114","first-page":"2583","article-title":"Septins 2, 7 and 9 and MAP4 colocalize along the axoneme in the primary cilium and control ciliary length","volume":"126","author":"Ghossoub","year":"2013","journal-title":"J. Cell Sci."},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"436","DOI":"10.1126\/science.1191054","article-title":"A septin diffusion barrier at the base of the primary cilium maintains ciliary membrane protein distribution","volume":"329","author":"Hu","year":"2010","journal-title":"Science"},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"1337","DOI":"10.1126\/science.1191184","article-title":"Planar cell polarity acts through septins to control collective cell movement and ciliogenesis","volume":"329","author":"Kim","year":"2010","journal-title":"Science"},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"315","DOI":"10.1038\/ni.3330","article-title":"A septin requirement differentiates autonomous and contact-facilitated T cell proliferation","volume":"17","author":"Mujal","year":"2016","journal-title":"Nat. Immunol."},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"5843","DOI":"10.1038\/ncomms6843","article-title":"Translocation between PI(4,5)P2-poor and PI(4,5)P2-rich microdomains during store depletion determines STIM1 conformation and Orai1 gating","volume":"5","author":"Maleth","year":"2014","journal-title":"Nat. Commun."},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"238","DOI":"10.1038\/nature12229","article-title":"An siRNA screen for NFAT activation identifies septins as coordinators of store-operated Ca2+ entry","volume":"499","author":"Sharma","year":"2013","journal-title":"Nature"},{"key":"ref_120","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1038\/emboj.2008.267","article-title":"Ciliary targeting motif VxPx directs assembly of a trafficking module through Arf4","volume":"28","author":"Mazelova","year":"2009","journal-title":"EMBO J."},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"4057","DOI":"10.1038\/emboj.2012.253","article-title":"The Arf GAP ASAP1 provides a platform to regulate Arf4- and Rab11-Rab8-mediated ciliary receptor targeting","volume":"31","author":"Wang","year":"2012","journal-title":"EMBO J."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"6346","DOI":"10.1073\/pnas.1002401107","article-title":"Coordination of Rab8 and Rab11 in primary ciliogenesis","volume":"107","author":"Knodler","year":"2010","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"1201","DOI":"10.1016\/j.cell.2007.03.053","article-title":"A core complex of BBS proteins cooperates with the GTPase Rab8 to promote ciliary membrane biogenesis","volume":"129","author":"Nachury","year":"2007","journal-title":"Cell"},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"2759","DOI":"10.1073\/pnas.1018823108","article-title":"Primary cilia membrane assembly is initiated by Rab11 and transport protein particle II (TRAPPII) complex-dependent trafficking of Rabin8 to the centrosome","volume":"108","author":"Westlake","year":"2011","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"363","DOI":"10.1083\/jcb.200703047","article-title":"Functional dissection of Rab GTPases involved in primary cilium formation","volume":"178","author":"Yoshimura","year":"2007","journal-title":"J. Cell Biol."},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"2522","DOI":"10.1091\/mbc.e08-07-0772","article-title":"The exocyst protein Sec10 is necessary for primary ciliogenesis and cystogenesis in vitro","volume":"20","author":"Zuo","year":"2009","journal-title":"Mol. Biol. Cell"},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"3781","DOI":"10.1091\/mbc.e06-02-0133","article-title":"The intraflagellar transport protein IFT20 is associated with the Golgi complex and is required for cilia assembly","volume":"17","author":"Follit","year":"2006","journal-title":"Mol. Biol. Cell"},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"921","DOI":"10.1091\/mbc.e10-09-0792","article-title":"IFT20 is required for opsin trafficking and photoreceptor outer segment development","volume":"22","author":"Keady","year":"2011","journal-title":"Mol. Biol. Cell"},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"2131","DOI":"10.1083\/jcb.201611138","article-title":"BLOC-1 is required for selective membrane protein trafficking from endosomes to primary cilia","volume":"216","author":"Monis","year":"2017","journal-title":"J. Cell Biol."},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"365","DOI":"10.1083\/jcb.200907126","article-title":"Lateral transport of Smoothened from the plasma membrane to the membrane of the cilium","volume":"187","author":"Milenkovic","year":"2009","journal-title":"J. Cell Biol."},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"2347","DOI":"10.1101\/gad.173443.111","article-title":"An ARL3-UNC119-RP2 GTPase cycle targets myristoylated NPHP3 to the primary cilium","volume":"25","author":"Wright","year":"2011","journal-title":"Genes Dev."},{"key":"ref_132","doi-asserted-by":"crossref","first-page":"2415","DOI":"10.1084\/jem.20130150","article-title":"Regulated vesicle fusion generates signaling nanoterritories that control T cell activation at the immunological synapse","volume":"210","author":"Soares","year":"2013","journal-title":"J. Exp. Med."},{"key":"ref_133","first-page":"2541","article-title":"The small GTPase Rab8 interacts with VAMP-3 to regulate the delivery of recycling T-cell receptors to the immune synapse","volume":"128","author":"Finetti","year":"2015","journal-title":"J. Cell Sci."},{"key":"ref_134","doi-asserted-by":"crossref","first-page":"1160","DOI":"10.15252\/embj.201593274","article-title":"Rac1-Rab11-FIP3 regulatory hub coordinates vesicle traffic with actin remodeling and T-cell activation","volume":"35","author":"Bouchet","year":"2016","journal-title":"EMBO J."},{"key":"ref_135","doi-asserted-by":"crossref","first-page":"2967","DOI":"10.4049\/jimmunol.1600671","article-title":"Rab11-FIP3 Regulation of Lck Endosomal Traffic Controls TCR Signal Transduction","volume":"198","author":"Bouchet","year":"2017","journal-title":"J. Immunol."},{"key":"ref_136","doi-asserted-by":"crossref","first-page":"1687","DOI":"10.1038\/cdd.2015.17","article-title":"The small GTPase Rab29 is a common regulator of immune synapse assembly and ciliogenesis","volume":"22","author":"Onnis","year":"2015","journal-title":"Cell Death Differ."},{"key":"ref_137","doi-asserted-by":"crossref","first-page":"18323","DOI":"10.1074\/jbc.M800056200","article-title":"Rab35 and its GAP EPI64C in T cells regulate receptor recycling and immunological synapse formation","volume":"283","author":"Dong","year":"2008","journal-title":"J. Biol. Chem."},{"key":"ref_138","doi-asserted-by":"crossref","first-page":"122.e124","DOI":"10.1016\/j.devcel.2018.08.012","article-title":"The Ciliary Machinery Is Repurposed for T Cell Immune Synapse Trafficking of LCK","volume":"47","author":"Stephen","year":"2018","journal-title":"Dev. Cell"},{"key":"ref_139","doi-asserted-by":"crossref","first-page":"451","DOI":"10.1016\/j.cub.2012.01.060","article-title":"Endocytosis genes facilitate protein and membrane transport in C. elegans sensory cilia","volume":"22","author":"Kaplan","year":"2012","journal-title":"Curr. Biol."},{"key":"ref_140","doi-asserted-by":"crossref","first-page":"3289","DOI":"10.1091\/mbc.e11-01-0082","article-title":"A conserved signal and GTPase complex are required for the ciliary transport of polycystin-1","volume":"22","author":"Ward","year":"2011","journal-title":"Mol. Biol. Cell"},{"key":"ref_141","doi-asserted-by":"crossref","first-page":"F495","DOI":"10.1152\/ajprenal.00198.2010","article-title":"Rab10 associates with primary cilia and the exocyst complex in renal epithelial cells","volume":"299","author":"Babbey","year":"2010","journal-title":"Am. J. Physiol. Renal. Physiol."},{"key":"ref_142","doi-asserted-by":"crossref","first-page":"711","DOI":"10.1038\/ncb3370","article-title":"Cilium assembly and disassembly","volume":"18","author":"Sanchez","year":"2016","journal-title":"Nat. Cell Biol."},{"key":"ref_143","doi-asserted-by":"crossref","first-page":"1375","DOI":"10.1242\/jcs.162925","article-title":"The Arf and Rab11 effector FIP3 acts synergistically with ASAP1 to direct Rabin8 in ciliary receptor targeting","volume":"128","author":"Wang","year":"2015","journal-title":"J. Cell Sci."},{"key":"ref_144","doi-asserted-by":"crossref","first-page":"1460","DOI":"10.1242\/jcs.058883","article-title":"Differential role of Rab proteins in ciliary trafficking: Rab23 regulates smoothened levels","volume":"123","author":"Boehlke","year":"2010","journal-title":"J. Cell Sci."},{"key":"ref_145","doi-asserted-by":"crossref","first-page":"2","DOI":"10.1016\/j.visres.2012.07.016","article-title":"Arl3 and RP2 mediated assembly and traffic of membrane associated cilia proteins","volume":"75","author":"Schwarz","year":"2012","journal-title":"Vis. Res."},{"key":"ref_146","doi-asserted-by":"crossref","first-page":"1208","DOI":"10.1016\/j.cell.2010.05.015","article-title":"The conserved Bardet-Biedl syndrome proteins assemble a coat that traffics membrane proteins to cilia","volume":"141","author":"Jin","year":"2010","journal-title":"Cell"},{"key":"ref_147","doi-asserted-by":"crossref","first-page":"1358","DOI":"10.1093\/hmg\/ddq012","article-title":"The retinitis pigmentosa protein RP2 links pericentriolar vesicle transport between the Golgi and the primary cilium","volume":"19","author":"Evans","year":"2010","journal-title":"Hum. Mol. Genet."},{"key":"ref_148","doi-asserted-by":"crossref","first-page":"437","DOI":"10.1038\/ncb1706","article-title":"Elipsa is an early determinant of ciliogenesis that links the IFT particle to membrane-associated small GTPase Rab8","volume":"10","author":"Omori","year":"2008","journal-title":"Nat. Cell Biol."},{"key":"ref_149","doi-asserted-by":"crossref","first-page":"21354","DOI":"10.1073\/pnas.1218272110","article-title":"Arl13b regulates endocytic recycling traffic","volume":"109","author":"Barral","year":"2012","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_150","doi-asserted-by":"crossref","first-page":"953","DOI":"10.1083\/jcb.200908133","article-title":"Joubert syndrome Arl13b functions at ciliary membranes and stabilizes protein transport in Caenorhabditis elegans","volume":"188","author":"Cevik","year":"2010","journal-title":"J. Cell Biol."},{"key":"ref_151","doi-asserted-by":"crossref","first-page":"712","DOI":"10.1016\/j.devcel.2009.09.010","article-title":"The Arp2\/3 activator WASH controls the fission of endosomes through a large multiprotein complex","volume":"17","author":"Derivery","year":"2009","journal-title":"Dev. Cell"},{"key":"ref_152","doi-asserted-by":"crossref","first-page":"699","DOI":"10.1016\/j.devcel.2009.09.009","article-title":"A FAM21-containing WASH complex regulates retromer-dependent sorting","volume":"17","author":"Gomez","year":"2009","journal-title":"Dev. Cell"},{"key":"ref_153","doi-asserted-by":"crossref","first-page":"6345","DOI":"10.4049\/jimmunol.1003771","article-title":"MAL protein controls protein sorting at the supramolecular activation cluster of human T lymphocytes","volume":"186","author":"Anton","year":"2011","journal-title":"J. Immunol."},{"key":"ref_154","doi-asserted-by":"crossref","first-page":"820","DOI":"10.1242\/jcs.078832","article-title":"Endosomal clathrin drives actin accumulation at the immunological synapse","volume":"124","author":"Mittelbrunn","year":"2011","journal-title":"J. Cell Sci."},{"key":"ref_155","doi-asserted-by":"crossref","first-page":"559","DOI":"10.1002\/embj.201386022","article-title":"beta-Arrestin-1 mediates the TCR-triggered re-routing of distal receptors to the immunological synapse by a PKC-mediated mechanism","volume":"33","author":"Calleja","year":"2014","journal-title":"EMBO J."},{"key":"ref_156","doi-asserted-by":"crossref","first-page":"3966","DOI":"10.1242\/jcs.073908","article-title":"The AP-1 clathrin adaptor facilitates cilium formation and functions with RAB-8 in C. elegans ciliary membrane transport","volume":"123","author":"Kaplan","year":"2010","journal-title":"J. Cell Sci."},{"key":"ref_157","doi-asserted-by":"crossref","first-page":"C862","DOI":"10.1152\/ajpcell.00338.2011","article-title":"VIP17\/MAL expression modulates epithelial cyst formation and ciliogenesis","volume":"303","author":"Takiar","year":"2012","journal-title":"Am. J. Physiol. Cell Physiol."},{"key":"ref_158","doi-asserted-by":"crossref","first-page":"1134","DOI":"10.1016\/j.cub.2007.05.055","article-title":"EB1 is required for primary cilia assembly in fibroblasts","volume":"17","author":"Schroder","year":"2007","journal-title":"Curr. Biol."},{"key":"ref_159","doi-asserted-by":"crossref","first-page":"4555","DOI":"10.4049\/jimmunol.1402734","article-title":"SNX17 affects T cell activation by regulating TCR and integrin recycling","volume":"194","author":"Osborne","year":"2015","journal-title":"J. Immunol."},{"key":"ref_160","doi-asserted-by":"crossref","first-page":"723","DOI":"10.1038\/ni.2609","article-title":"VAMP7 controls T cell activation by regulating the recruitment and phosphorylation of vesicular Lat at TCR-activation sites","volume":"14","author":"Larghi","year":"2013","journal-title":"Nat. Immunol."},{"key":"ref_161","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1038\/cr.2011.134","article-title":"A SNX10\/V-ATPase pathway regulates ciliogenesis in vitro and in vivo","volume":"22","author":"Chen","year":"2012","journal-title":"Cell Res."},{"key":"ref_162","doi-asserted-by":"crossref","unstructured":"Szalinski, C.M., Labilloy, A., Bruns, J.R., and Weisz, O.A. (2014). VAMP7 modulates ciliary biogenesis in kidney cells. PLoS ONE, 9.","DOI":"10.1371\/journal.pone.0086425"},{"key":"ref_163","doi-asserted-by":"crossref","first-page":"889","DOI":"10.1242\/jcs.023861","article-title":"Intraflagellar transport at a glance","volume":"122","author":"Hao","year":"2009","journal-title":"J. Cell Sci."},{"key":"ref_164","doi-asserted-by":"crossref","first-page":"124","DOI":"10.1016\/j.ceb.2018.03.004","article-title":"The molecular machines that traffic signaling receptors into and out of cilia","volume":"51","author":"Nachury","year":"2018","journal-title":"Curr. Opin. Cell Biol."},{"key":"ref_165","doi-asserted-by":"crossref","first-page":"6943","DOI":"10.1073\/pnas.1221011110","article-title":"Evolution of modular intraflagellar transport from a coatomer-like progenitor","volume":"110","author":"Townsend","year":"2013","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_166","doi-asserted-by":"crossref","first-page":"1505","DOI":"10.1016\/j.celrep.2016.10.018","article-title":"Role for the IFT-A Complex in Selective Transport to the Primary Cilium","volume":"17","author":"Fu","year":"2016","journal-title":"Cell Rep."},{"key":"ref_167","doi-asserted-by":"crossref","first-page":"210","DOI":"10.1016\/j.cell.2012.12.026","article-title":"The ciliary G-protein-coupled receptor Gpr161 negatively regulates the Sonic hedgehog pathway via cAMP signaling","volume":"152","author":"Mukhopadhyay","year":"2013","journal-title":"Cell"},{"key":"ref_168","doi-asserted-by":"crossref","first-page":"4242","DOI":"10.1073\/pnas.0711027105","article-title":"Bardet-Biedl syndrome proteins are required for the localization of G protein-coupled receptors to primary cilia","volume":"105","author":"Berbari","year":"2008","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_169","doi-asserted-by":"crossref","unstructured":"Seo, S., Zhang, Q., Bugge, K., Breslow, D.K., Searby, C.C., Nachury, M.V., and Sheffield, V.C. (2011). A novel protein LZTFL1 regulates ciliary trafficking of the BBSome and Smoothened. PLoS Genet., 7.","DOI":"10.1371\/journal.pgen.1002358"},{"key":"ref_170","doi-asserted-by":"crossref","first-page":"5441","DOI":"10.1093\/hmg\/ddu267","article-title":"Bardet-Biedl syndrome proteins 1 and 3 regulate the ciliary trafficking of polycystic kidney disease 1 protein","volume":"23","author":"Su","year":"2014","journal-title":"Hum. Mol. Genet."},{"key":"ref_171","doi-asserted-by":"crossref","first-page":"1847","DOI":"10.1083\/jcb.201709041","article-title":"BBSome trains remove activated GPCRs from cilia by enabling passage through the transition zone","volume":"217","author":"Ye","year":"2018","journal-title":"J. Cell Biol."},{"key":"ref_172","doi-asserted-by":"crossref","unstructured":"Robichaux, M.A., Potter, V.L., Zhang, Z., He, F., Schmid, M.F., and Wensel, T.G. (2017). Defining the Layers of a Sensory Cilium with STORM and Cryo-Electron Nanoscopies. bioRxiv, 198655.","DOI":"10.1101\/198655"},{"key":"ref_173","doi-asserted-by":"crossref","first-page":"191","DOI":"10.1002\/bies.20369","article-title":"Evolution of intraflagellar transport from coated vesicles and autogenous origin of the eukaryotic cilium","volume":"28","author":"Jekely","year":"2006","journal-title":"Bioessays"},{"key":"ref_174","doi-asserted-by":"crossref","first-page":"194","DOI":"10.1038\/nature12639","article-title":"Functional interaction between autophagy and ciliogenesis","volume":"502","author":"Pampliega","year":"2013","journal-title":"Nature"},{"key":"ref_175","doi-asserted-by":"crossref","first-page":"386","DOI":"10.1073\/pnas.1513601113","article-title":"IFT20 controls LAT recruitment to the immune synapse and T-cell activation in vivo","volume":"113","author":"Vivar","year":"2016","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_176","doi-asserted-by":"crossref","first-page":"901","DOI":"10.1083\/jcb.149.4.901","article-title":"Distinct membrane domains on endosomes in the recycling pathway visualized by multicolor imaging of Rab4, Rab5, and Rab11","volume":"149","author":"Sonnichsen","year":"2000","journal-title":"J. Cell Biol."},{"key":"ref_177","doi-asserted-by":"crossref","first-page":"913","DOI":"10.1083\/jcb.135.4.913","article-title":"Rab11 regulates recycling through the pericentriolar recycling endosome","volume":"135","author":"Ullrich","year":"1996","journal-title":"J. Cell Biol."},{"key":"ref_178","doi-asserted-by":"crossref","first-page":"729","DOI":"10.1016\/0092-8674(92)90307-X","article-title":"The small GTP-binding protein rab4 controls an early sorting event on the endocytic pathway","volume":"70","author":"Hull","year":"1992","journal-title":"Cell"},{"key":"ref_179","doi-asserted-by":"crossref","first-page":"1245","DOI":"10.1084\/jem.20162042","article-title":"Rab6-dependent retrograde traffic of LAT controls immune synapse formation and T cell activation","volume":"215","author":"Carpier","year":"2018","journal-title":"J. Exp. Med."},{"key":"ref_180","doi-asserted-by":"crossref","first-page":"279","DOI":"10.1016\/j.devcel.2014.09.011","article-title":"IFT27 links the BBSome to IFT for maintenance of the ciliary signaling compartment","volume":"31","author":"Eguether","year":"2014","journal-title":"Dev. Cell"},{"key":"ref_181","doi-asserted-by":"crossref","first-page":"940","DOI":"10.1016\/j.devcel.2012.04.009","article-title":"IFT25 links the signal-dependent movement of Hedgehog components to intraflagellar transport","volume":"22","author":"Keady","year":"2012","journal-title":"Dev. Cell"},{"key":"ref_182","doi-asserted-by":"crossref","first-page":"2194","DOI":"10.1242\/dev.115261","article-title":"Intraflagellar transport 27 is essential for hedgehog signaling but dispensable for ciliogenesis during hair follicle morphogenesis","volume":"142","author":"Yang","year":"2015","journal-title":"Development"},{"key":"ref_183","doi-asserted-by":"crossref","first-page":"338","DOI":"10.1016\/j.devcel.2015.06.016","article-title":"Modulation of Ciliary Phosphoinositide Content Regulates Trafficking and Sonic Hedgehog Signaling Output","volume":"34","author":"Chavez","year":"2015","journal-title":"Dev. Cell"},{"key":"ref_184","doi-asserted-by":"crossref","first-page":"400","DOI":"10.1016\/j.devcel.2015.08.001","article-title":"Phosphoinositides Regulate Ciliary Protein Trafficking to Modulate Hedgehog Signaling","volume":"34","author":"Phua","year":"2015","journal-title":"Dev. Cell"},{"key":"ref_185","doi-asserted-by":"crossref","first-page":"18556","DOI":"10.1073\/pnas.0608291103","article-title":"FAPP2, cilium formation, and compartmentalization of the apical membrane in polarized Madin-Darby canine kidney (MDCK) cells","volume":"103","author":"Vieira","year":"2006","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_186","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1016\/j.bbrc.2018.12.109","article-title":"TULP3 is required for localization of membrane-associated proteins ARL13B and INPP5E to primary cilia","volume":"509","author":"Han","year":"2019","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_187","doi-asserted-by":"crossref","first-page":"2180","DOI":"10.1101\/gad.1966210","article-title":"TULP3 bridges the IFT-A complex and membrane phosphoinositides to promote trafficking of G protein-coupled receptors into primary cilia","volume":"24","author":"Mukhopadhyay","year":"2010","journal-title":"Genes Dev."},{"key":"ref_188","doi-asserted-by":"crossref","first-page":"1032","DOI":"10.1038\/ng.423","article-title":"Mutations in INPP5E, encoding inositol polyphosphate-5-phosphatase E, link phosphatidyl inositol signaling to the ciliopathies","volume":"41","author":"Bielas","year":"2009","journal-title":"Nat. Genet."},{"key":"ref_189","doi-asserted-by":"crossref","first-page":"529","DOI":"10.1038\/nrneph.2009.124","article-title":"Inherited cerebrorenal syndromes","volume":"5","author":"Schurman","year":"2009","journal-title":"Nat. Rev. Nephrol."},{"key":"ref_190","doi-asserted-by":"crossref","first-page":"3447","DOI":"10.1242\/jcs.200857","article-title":"Loss of OCRL increases ciliary PI(4,5)P2 in Lowe oculocerebrorenal syndrome","volume":"130","author":"Prosseda","year":"2017","journal-title":"J. Cell Sci."},{"key":"ref_191","doi-asserted-by":"crossref","first-page":"4966","DOI":"10.1038\/s41598-018-23109-8","article-title":"Phosphoinositides regulate the TCR\/CD3 complex membrane dynamics and activation","volume":"8","author":"Ruminski","year":"2018","journal-title":"Sci. Rep."},{"key":"ref_192","doi-asserted-by":"crossref","first-page":"701","DOI":"10.1083\/jcb.200909025","article-title":"An electrostatic switch displaces phosphatidylinositol phosphate kinases from the membrane during phagocytosis","volume":"187","author":"Fairn","year":"2009","journal-title":"J. Cell Biol."},{"key":"ref_193","doi-asserted-by":"crossref","first-page":"e1600925","DOI":"10.1126\/sciadv.1600925","article-title":"The activation loop of PIP5K functions as a membrane sensor essential for lipid substrate processing","volume":"2","author":"Liu","year":"2016","journal-title":"Sci. Adv."},{"key":"ref_194","doi-asserted-by":"crossref","first-page":"502","DOI":"10.3389\/fimmu.2017.00502","article-title":"ISA-2011B, a Phosphatidylinositol 4-Phosphate 5-Kinase alpha Inhibitor, Impairs CD28-Dependent Costimulatory and Pro-inflammatory Signals in Human T Lymphocytes","volume":"8","author":"Kunkl","year":"2017","journal-title":"Front. Immunol."},{"key":"ref_195","doi-asserted-by":"crossref","first-page":"1955","DOI":"10.4049\/jimmunol.1501788","article-title":"Phosphatidylinositol 4-Phosphate 5-Kinase beta Controls Recruitment of Lipid Rafts into the Immunological Synapse","volume":"196","author":"Kallikourdis","year":"2016","journal-title":"J. Immunol."},{"key":"ref_196","doi-asserted-by":"crossref","first-page":"389","DOI":"10.1083\/jcb.134.2.389","article-title":"Coordinated regulation of platelet actin filament barbed ends by gelsolin and capping protein","volume":"134","author":"Barkalow","year":"1996","journal-title":"J. Cell Biol."},{"key":"ref_197","doi-asserted-by":"crossref","first-page":"311","DOI":"10.1016\/S0960-9822(00)00384-5","article-title":"Phosphatidylinositol 4,5-bisphosphate induces actin-based movement of raft-enriched vesicles through WASP-Arp2\/3","volume":"10","author":"Rozelle","year":"2000","journal-title":"Curr. Biol."},{"key":"ref_198","doi-asserted-by":"crossref","unstructured":"Szatmari, D., Xue, B., Kannan, B., Burtnick, L.D., Bugyi, B., Nyitrai, M., and Robinson, R.C. (2018). ATP competes with PIP2 for binding to gelsolin. PLoS ONE, 13.","DOI":"10.1371\/journal.pone.0201826"},{"key":"ref_199","doi-asserted-by":"crossref","first-page":"2327","DOI":"10.1016\/j.bpj.2010.01.046","article-title":"ADF\/cofilin binds phosphoinositides in a multivalent manner to act as a PIP(2)-density sensor","volume":"98","author":"Zhao","year":"2010","journal-title":"Biophys. J."},{"key":"ref_200","doi-asserted-by":"crossref","first-page":"700","DOI":"10.3389\/fimmu.2019.00700","article-title":"Phospholipids: Pulling Back the Actin Curtain for Granule Delivery to the Immune Synapse","volume":"10","author":"Griffiths","year":"2019","journal-title":"Front. Immunol."},{"key":"ref_201","doi-asserted-by":"crossref","first-page":"307","DOI":"10.1016\/j.bbrc.2018.07.155","article-title":"Regulation of actin assembly by PI(4,5)P2 and other inositol phospholipids: An update on possible mechanisms","volume":"506","author":"Janmey","year":"2018","journal-title":"Biochem. Biophys. Res. Commun."},{"key":"ref_202","doi-asserted-by":"crossref","first-page":"ra82","DOI":"10.1126\/scisignal.2005287","article-title":"Diacylglycerol kinase alpha establishes T cell polarity by shaping diacylglycerol accumulation at the immunological synapse","volume":"7","author":"Chauveau","year":"2014","journal-title":"Sci. Signal."},{"key":"ref_203","doi-asserted-by":"crossref","first-page":"555","DOI":"10.1091\/mbc.e12-06-0437","article-title":"The PCP effector Fuzzy controls cilial assembly and signaling by recruiting Rab8 and Dishevelled to the primary cilium","volume":"24","author":"Zilber","year":"2013","journal-title":"Mol. Biol. Cell"},{"key":"ref_204","doi-asserted-by":"crossref","first-page":"1451","DOI":"10.1016\/j.cub.2004.08.025","article-title":"Polarity proteins control ciliogenesis via kinesin motor interactions","volume":"14","author":"Fan","year":"2004","journal-title":"Curr. Biol."},{"key":"ref_205","doi-asserted-by":"crossref","first-page":"5649","DOI":"10.4049\/jimmunol.179.9.5649","article-title":"Cutting edge: Atypical PKCs regulate T lymphocyte polarity and scanning behavior","volume":"179","author":"Real","year":"2007","journal-title":"J. Immunol."},{"key":"ref_206","doi-asserted-by":"crossref","first-page":"737","DOI":"10.1016\/j.immuni.2005.04.009","article-title":"A network of PDZ-containing proteins regulates T cell polarity and morphology during migration and immunological synapse formation","volume":"22","author":"Oliaro","year":"2005","journal-title":"Immunity"},{"key":"ref_207","doi-asserted-by":"crossref","first-page":"367","DOI":"10.4049\/jimmunol.0903627","article-title":"Asymmetric cell division of T cells upon antigen presentation uses multiple conserved mechanisms","volume":"185","author":"Oliaro","year":"2010","journal-title":"J. Immunol."},{"key":"ref_208","doi-asserted-by":"crossref","first-page":"365","DOI":"10.1038\/ni.2842","article-title":"Early specification of CD8+ T lymphocyte fates during adaptive immunity revealed by single-cell gene-expression analyses","volume":"15","author":"Arsenio","year":"2014","journal-title":"Nat. Immunol."},{"key":"ref_209","doi-asserted-by":"crossref","first-page":"492","DOI":"10.1016\/j.immuni.2011.03.017","article-title":"Asymmetric proteasome segregation as a mechanism for unequal partitioning of the transcription factor T-bet during T lymphocyte division","volume":"34","author":"Chang","year":"2011","journal-title":"Immunity"},{"key":"ref_210","doi-asserted-by":"crossref","first-page":"1687","DOI":"10.1126\/science.1139393","article-title":"Asymmetric T lymphocyte division in the initiation of adaptive immune responses","volume":"315","author":"Chang","year":"2007","journal-title":"Science"},{"key":"ref_211","doi-asserted-by":"crossref","first-page":"2249","DOI":"10.4049\/jimmunol.1401652","article-title":"Regulation of asymmetric division and CD8+ T lymphocyte fate specification by protein kinase Czeta and protein kinase Clambda\/iota","volume":"194","author":"Metz","year":"2015","journal-title":"J. Immunol."},{"key":"ref_212","doi-asserted-by":"crossref","first-page":"9866","DOI":"10.1073\/pnas.0501202102","article-title":"Control of T helper 2 cell function and allergic airway inflammation by PKCzeta","volume":"102","author":"Martin","year":"2005","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_213","doi-asserted-by":"crossref","first-page":"1099","DOI":"10.1073\/pnas.0805907106","article-title":"Loss of PKC lambda\/iota impairs Th2 establishment and allergic airway inflammation in vivo","volume":"106","author":"Yang","year":"2009","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_214","doi-asserted-by":"crossref","first-page":"181","DOI":"10.1016\/j.celrep.2017.09.020","article-title":"Adenomatous Polyposis Coli Defines Treg Differentiation and Anti-inflammatory Function through Microtubule-Mediated NFAT Localization","volume":"21","author":"Burton","year":"2017","journal-title":"Cell Rep."},{"key":"ref_215","doi-asserted-by":"crossref","first-page":"421","DOI":"10.1038\/nri.2016.54","article-title":"Origins of the cytolytic synapse","volume":"16","author":"Asano","year":"2016","journal-title":"Nat. Rev. Immunol."},{"key":"ref_216","doi-asserted-by":"crossref","first-page":"E2589","DOI":"10.1073\/pnas.1519458113","article-title":"Canonical and noncanonical intraflagellar transport regulates craniofacial skeletal development","volume":"113","author":"Noda","year":"2016","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_217","doi-asserted-by":"crossref","unstructured":"Finetti, F., Cassioli, C., Cianfanelli, V., Onnis, A., Paccagnini, E., Kabanova, A., and Baldari, C.T. (2019). The intraflagellar transport protein IFT20 controls lysosome biogenesis by regulating the post-Golgi transport of acid hydrolases. Cell Death Differ.","DOI":"10.1038\/s41418-019-0357-y"},{"key":"ref_218","doi-asserted-by":"crossref","first-page":"1427","DOI":"10.1242\/dev.074666","article-title":"Switching on cilia: Transcriptional networks regulating ciliogenesis","volume":"141","author":"Choksi","year":"2014","journal-title":"Development"},{"key":"ref_219","doi-asserted-by":"crossref","unstructured":"Viau, A., Bienaime, F., Lukas, K., Todkar, A.P., Knoll, M., Yakulov, T.A., Hofherr, A., Kretz, O., Helmstadter, M., and Reichardt, W. (2018). Cilia-localized LKB1 regulates chemokine signaling, macrophage recruitment, and tissue homeostasis in the kidney. EMBO J., 37.","DOI":"10.15252\/embj.201798615"},{"key":"ref_220","doi-asserted-by":"crossref","first-page":"1399","DOI":"10.1182\/blood-2011-04-350686","article-title":"A mutation in the human Uncoordinated 119 gene impairs TCR signaling and is associated with CD4 lymphopenia","volume":"119","author":"Gorska","year":"2012","journal-title":"Blood"}],"container-title":["Cells"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/2073-4409\/8\/8\/789\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T13:11:02Z","timestamp":1760188262000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/2073-4409\/8\/8\/789"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,7,29]]},"references-count":220,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2019,8]]}},"alternative-id":["cells8080789"],"URL":"https:\/\/doi.org\/10.3390\/cells8080789","relation":{},"ISSN":["2073-4409"],"issn-type":[{"value":"2073-4409","type":"electronic"}],"subject":[],"published":{"date-parts":[[2019,7,29]]}}}