{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,30]],"date-time":"2026-09-30T11:48:48Z","timestamp":1790768928578,"version":"4.1.0"},"reference-count":36,"publisher":"Rockefeller University Press","issue":"1","license":[{"start":{"date-parts":[[2019,12,24]],"date-time":"2019-12-24T00:00:00Z","timestamp":1577145600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/www.rupress.org\/terms\/"},{"start":{"date-parts":[[2020,7,6]],"date-time":"2020-07-06T00:00:00Z","timestamp":1593993600000},"content-version":"vor","delay-in-days":195,"URL":"https:\/\/creativecommons.org\/licenses\/by-nc-sa\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["GM031627"],"award-info":[{"award-number":["GM031627"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["GM118099"],"award-info":[{"award-number":["GM118099"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["GM118106"],"award-info":[{"award-number":["GM118106"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["S10OD020054"],"award-info":[{"award-number":["S10OD020054"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"publisher","award":["S10OD021741"],"award-info":[{"award-number":["S10OD021741"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000011","name":"Howard Hughes Medical Institute","doi-asserted-by":"publisher","id":[{"id":"10.13039\/100000011","id-type":"DOI","asserted-by":"publisher"}]}],"content-domain":{"domain":["rupress.org"],"crossmark-restriction":true},"short-container-title":[],"published-print":{"date-parts":[[2020,1,6]]},"abstract":"<jats:p>Cells use motile cilia to generate force in the extracellular space. The structure of a cilium can be classified into three subdomains: the intracellular basal body (BB) that templates cilium formation, the extracellular axoneme that generates force, and the transition zone (TZ) that bridges them. While the BB is composed of triplet microtubules (TMTs), the axoneme is composed of doublet microtubules (DMTs), meaning the cilium must convert between different microtubule geometries. Here, we performed electron cryotomography to define this conversion, and our reconstructions reveal identifying structural features of the BB, TZ, and axoneme. Each region is distinct in terms of microtubule number and geometry, microtubule inner proteins, and microtubule linkers. TMT to DMT conversion occurs within the BB, and microtubule geometry changes to axonemal by the end of the TZ, followed by the addition of axoneme-specific components essential for cilium motility. Our results provide the highest-resolution images of the motile cilium to date and reveal how BBs template axonemes.<\/jats:p>","DOI":"10.1083\/jcb.201907060","type":"journal-article","created":{"date-parts":[[2019,12,24]],"date-time":"2019-12-24T15:14:24Z","timestamp":1577200464000},"update-policy":"https:\/\/doi.org\/10.1083\/jcb.crossmarkpolicy","source":"Crossref","is-referenced-by-count":52,"title":["Electron cryotomography of intact motile cilia defines the basal body to axoneme transition"],"prefix":"10.1083","volume":"219","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-9045-7666","authenticated-orcid":true,"given":"Garrett A.","family":"Greenan","sequence":"first","affiliation":[{"name":"Department of Biochemistry and Biophysics, University of California, San Francisco, San Francisco, CA 1"},{"name":"Department of Cellular and Molecular Pharmacology, University of California, San Francisco, San Francisco, CA 2"},{"name":"The Howard Hughes Medical Institute, Chevy Chase, MD 3"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3460-2758","authenticated-orcid":true,"given":"Ronald D.","family":"Vale","sequence":"additional","affiliation":[{"name":"Department of Cellular and Molecular Pharmacology, University of California, San Francisco, San Francisco, CA 2"},{"name":"The Howard Hughes Medical Institute, Chevy Chase, MD 3"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3512-695X","authenticated-orcid":true,"given":"David A.","family":"Agard","sequence":"additional","affiliation":[{"name":"Department of Biochemistry and Biophysics, University of California, San Francisco, San Francisco, CA 1"},{"name":"The Howard Hughes Medical Institute, Chevy Chase, MD 3"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"291","published-online":{"date-parts":[[2019,12,24]]},"reference":[{"key":"2023072120333629600_bib1","doi-asserted-by":"publisher","first-page":"241","DOI":"10.1016\/S0040-8166(95)80044-1","article-title":"Flagellar structure in normal human spermatozoa and in spermatozoa that lack dynein arms","volume":"27","author":"Afzelius","year":"1995","journal-title":"Tissue Cell."},{"key":"2023072120333629600_bib2","doi-asserted-by":"publisher","first-page":"246","DOI":"10.1083\/jcb.54.2.246","article-title":"The three-dimensional structure of the basal body from the rhesus monkey oviduct","volume":"54","author":"Anderson","year":"1972","journal-title":"J. Cell Biol."},{"key":"2023072120333629600_bib3","doi-asserted-by":"publisher","first-page":"393","DOI":"10.1083\/jcb.60.2.393","article-title":"Isolation of ciliated or unciliated basal bodies from the rabbit oviduct","volume":"60","author":"Anderson","year":"1974","journal-title":"J. Cell Biol."},{"key":"2023072120333629600_bib4","doi-asserted-by":"publisher","first-page":"1743","DOI":"10.1016\/j.str.2015.06.026","article-title":"Advances in Single-Particle Electron Cryomicroscopy Structure Determination applied to Sub-tomogram Averaging","volume":"23","author":"Bharat","year":"2015","journal-title":"Structure."},{"key":"2023072120333629600_bib5","doi-asserted-by":"publisher","first-page":"261","DOI":"10.1016\/j.sbi.2013.02.003","article-title":"Structural biology in situ--the potential of subtomogram averaging","volume":"23","author":"Briggs","year":"2013","journal-title":"Curr. Opin. Struct. Biol."},{"key":"2023072120333629600_bib6","doi-asserted-by":"publisher","first-page":"913","DOI":"10.1083\/jcb.201201120","article-title":"Polarity and asymmetry in the arrangement of dynein and related structures in the Chlamydomonas axoneme","volume":"198","author":"Bui","year":"2012","journal-title":"J. Cell Biol."},{"key":"2023072120333629600_bib7","doi-asserted-by":"publisher","first-page":"56","DOI":"10.1006\/jsbi.1996.0010","article-title":"IVE (Image Visualization Environment): a software platform for all three-dimensional microscopy applications","volume":"116","author":"Chen","year":"1996","journal-title":"J. Struct. Biol."},{"key":"2023072120333629600_bib8","doi-asserted-by":"publisher","first-page":"927","DOI":"10.1083\/jcb.201006105","article-title":"CEP290 tethers flagellar transition zone microtubules to the membrane and regulates flagellar protein content","volume":"190","author":"Craige","year":"2010","journal-title":"J. Cell Biol."},{"key":"2023072120333629600_bib9","doi-asserted-by":"publisher","first-page":"249","DOI":"10.1042\/BC20100139","article-title":"Ultrastructure of cilia and flagella - back to the future!","volume":"103","author":"Fisch","year":"2011","journal-title":"Biol. Cell."},{"key":"2023072120333629600_bib10","doi-asserted-by":"publisher","first-page":"494","DOI":"10.1083\/jcb.53.2.494","article-title":"The ciliary necklace. A ciliary membrane specialization","volume":"53","author":"Gilula","year":"1972","journal-title":"J. Cell Biol."},{"key":"2023072120333629600_bib11","doi-asserted-by":"publisher","first-page":"243","DOI":"10.14348\/molcells.2017.0054","article-title":"The Ciliary Transition Zone: Finding the Pieces and Assembling the Gate","volume":"40","author":"Gon\u00e7alves","year":"2017","journal-title":"Mol. Cells."},{"key":"2023072120333629600_bib12","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.36851","article-title":"Insights into centriole geometry revealed by cryotomography of doublet and triplet centrioles","volume":"7","author":"Greenan","year":"2018","journal-title":"eLife."},{"key":"2023072120333629600_bib13","doi-asserted-by":"publisher","first-page":"553","DOI":"10.1126\/science.1222789","article-title":"Cartwheel architecture of Trichonympha basal body","volume":"337","author":"Guichard","year":"2012","journal-title":"Science."},{"key":"2023072120333629600_bib14","doi-asserted-by":"publisher","first-page":"921","DOI":"10.1083\/jcb.200908067","article-title":"The dynein regulatory complex is the nexin link and a major regulatory node in cilia and flagella","volume":"187","author":"Heuser","year":"2009","journal-title":"J. Cell Biol."},{"key":"2023072120333629600_bib15","doi-asserted-by":"publisher","first-page":"15035","DOI":"10.1038\/ncomms15035","article-title":"Subnanometre-resolution structure of the doublet microtubule reveals new classes of microtubule-associated proteins","volume":"8","author":"Ichikawa","year":"2017","journal-title":"Nat. Commun."},{"key":"2023072120333629600_bib16","doi-asserted-by":"publisher","DOI":"10.1101\/cshperspect.a028076","article-title":"Axoneme Structure from Motile Cilia","volume":"9","author":"Ishikawa","year":"2017","journal-title":"Cold Spring Harb. Perspect. Biol."},{"key":"2023072120333629600_bib17","doi-asserted-by":"publisher","first-page":"71","DOI":"10.1006\/jsbi.1996.0013","article-title":"Computer visualization of three-dimensional image data using IMOD","volume":"116","author":"Kremer","year":"1996","journal-title":"J. Struct. Biol."},{"key":"2023072120333629600_bib18","doi-asserted-by":"publisher","first-page":"552","DOI":"10.1038\/emboj.2011.460","article-title":"Three-dimensional structure of basal body triplet revealed by electron cryo-tomography","volume":"31","author":"Li","year":"2012","journal-title":"EMBO J."},{"key":"2023072120333629600_bib19","doi-asserted-by":"publisher","first-page":"5727","DOI":"10.1038\/ncomms6727","article-title":"Cryo-electron tomography reveals ciliary defects underlying human RSPH1 primary ciliary dyskinesia","volume":"5","author":"Lin","year":"2014","journal-title":"Nat. Commun."},{"key":"2023072120333629600_bib20","doi-asserted-by":"publisher","first-page":"126","DOI":"10.1016\/j.tcb.2016.08.002","article-title":"The Cilium: Cellular Antenna and Central Processing Unit","volume":"27","author":"Malicki","year":"2017","journal-title":"Trends Cell Biol."},{"key":"2023072120333629600_bib21","doi-asserted-by":"publisher","first-page":"15889","DOI":"10.1073\/pnas.0508274102","article-title":"3D structure of eukaryotic flagella in a quiescent state revealed by cryo-electron tomography","volume":"102","author":"Nicastro","year":"2005","journal-title":"Proc. Natl. Acad. Sci. USA."},{"key":"2023072120333629600_bib22","doi-asserted-by":"publisher","first-page":"944","DOI":"10.1126\/science.1128618","article-title":"The molecular architecture of axonemes revealed by cryoelectron tomography","volume":"313","author":"Nicastro","year":"2006","journal-title":"Science."},{"key":"2023072120333629600_bib23","doi-asserted-by":"publisher","first-page":"107","DOI":"10.1016\/1047-8477(92)90011-X","article-title":"Centrosome organization and centriole architecture: their sensitivity to divalent cations","volume":"108","author":"Paintrand","year":"1992","journal-title":"J. Struct. Biol."},{"key":"2023072120333629600_bib24","doi-asserted-by":"publisher","first-page":"1605","DOI":"10.1002\/jcc.20084","article-title":"UCSF Chimera--a visualization system for exploratory research and analysis","volume":"25","author":"Pettersen","year":"2004","journal-title":"J. Comput. Chem."},{"key":"2023072120333629600_bib25","doi-asserted-by":"publisher","first-page":"673","DOI":"10.1083\/jcb.201106125","article-title":"Cryoelectron tomography of radial spokes in cilia and flagella","volume":"195","author":"Pigino","year":"2011","journal-title":"J. Cell Biol."},{"key":"2023072120333629600_bib26","doi-asserted-by":"publisher","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":"2023072120333629600_bib27","doi-asserted-by":"publisher","first-page":"608","DOI":"10.1038\/embor.2012.73","article-title":"The base of the cilium: roles for transition fibres and the transition zone in ciliary formation, maintenance and compartmentalization","volume":"13","author":"Reiter","year":"2012","journal-title":"EMBO Rep."},{"key":"2023072120333629600_bib28","doi-asserted-by":"publisher","first-page":"372","DOI":"10.1126\/science.1139740","article-title":"Patched1 regulates hedgehog signaling at the primary cilium","volume":"317","author":"Rohatgi","year":"2007","journal-title":"Science."},{"key":"2023072120333629600_bib29","doi-asserted-by":"publisher","first-page":"377","DOI":"10.1146\/annurev.physiol.69.040705.141236","article-title":"Overview of structure and function of mammalian cilia","volume":"69","author":"Satir","year":"2007","journal-title":"Annu. Rev. Physiol."},{"key":"2023072120333629600_bib30","doi-asserted-by":"publisher","first-page":"1073","DOI":"10.1093\/biosci\/biu180","article-title":"A Structural Basis for How Motile Cilia Beat","volume":"64","author":"Satir","year":"2014","journal-title":"Bioscience."},{"key":"2023072120333629600_bib31","doi-asserted-by":"publisher","first-page":"499","DOI":"10.1242\/jcs.050377","article-title":"The primary cilium at a glance","volume":"123","author":"Satir","year":"2010","journal-title":"J. Cell Sci."},{"key":"2023072120333629600_bib32","doi-asserted-by":"publisher","first-page":"475","DOI":"10.1038\/nature04816","article-title":"Molecular architecture of axonemal microtubule doublets revealed by cryo-electron tomography","volume":"442","author":"Sui","year":"2006","journal-title":"Nature."},{"key":"2023072120333629600_bib33","doi-asserted-by":"publisher","first-page":"5","DOI":"10.1186\/s13630-016-0023-7","article-title":"Basal body structure and cell cycle-dependent biogenesis in Trypanosoma brucei","volume":"5","author":"Vaughan","year":"2016","journal-title":"Cilia."},{"key":"2023072120333629600_bib34","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.36979","article-title":"Systematic studies of all PIH proteins in zebrafish reveal their distinct roles in axonemal dynein assembly","volume":"7","author":"Yamaguchi","year":"2018","journal-title":"eLife."},{"key":"2023072120333629600_bib35","doi-asserted-by":"publisher","first-page":"14096","DOI":"10.1038\/srep14096","article-title":"Superresolution Pattern Recognition Reveals the Architectural Map of the Ciliary Transition Zone","volume":"5","author":"Yang","year":"2015","journal-title":"Sci. Rep."},{"key":"2023072120333629600_bib36","doi-asserted-by":"publisher","first-page":"2727","DOI":"10.1038\/s41598-018-21165-8","article-title":"A lumenal interrupted helix in human sperm tail microtubules","volume":"8","author":"Zabeo","year":"2018","journal-title":"Sci. Rep."}],"container-title":["Journal of Cell Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/rupress.org\/jcb\/article-pdf\/doi\/10.1083\/jcb.201907060\/1455649\/jcb_201907060.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/rupress.org\/jcb\/article-pdf\/doi\/10.1083\/jcb.201907060\/1455649\/jcb_201907060.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2023,7,21]],"date-time":"2023-07-21T20:37:47Z","timestamp":1689971867000},"score":1,"resource":{"primary":{"URL":"https:\/\/rupress.org\/jcb\/article\/219\/1\/e201907060\/133537\/Electron-cryotomography-of-intact-motile-cilia"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2019,12,24]]},"references-count":36,"journal-issue":{"issue":"1","published-print":{"date-parts":[[2020,1,6]]}},"URL":"https:\/\/doi.org\/10.1083\/jcb.201907060","relation":{},"ISSN":["0021-9525","1540-8140"],"issn-type":[{"value":"0021-9525","type":"print"},{"value":"1540-8140","type":"electronic"}],"subject":[],"published":{"date-parts":[[2019,12,24]]},"article-number":"e201907060"}}