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George B Witman PhD

TitleProfessor Emeritus
Endowed TitleGeorge F. Booth Chair in the Basic Sciences
InstitutionUniversity of Massachusetts Medical School
DepartmentRadiology
AddressUniversity of Massachusetts Medical School
55 Lake Avenue North
Worcester MA 01655
Phone508-856-4038
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    Other Positions
    InstitutionUMMS - School of Medicine
    DepartmentRadiology

    InstitutionUMMS - Graduate School of Biomedical Sciences
    DepartmentCell Biology

    InstitutionUMMS - Graduate School of Biomedical Sciences
    DepartmentInterdisciplinary Graduate Program


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    University of California, Riverside, Riverside, CA, United StatesBAZoology
    Yale University, New Haven, CT, United StatesPH DBiology

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    Cilia and Flagella, Molecular Motors, Sensory Transduction, Proteomics, Molecular Basis for Diseases Involving Cilia and Flagella


    Our research is concerned with the biology of cilia and flagella, including the non-motile primary cilia that are present on most cells in our bodies and function as cell antennae, receiving signals from the environment and transmitting these to the cell body. Our findings have important implications for human development and male infertility, and for diseases of the lung, kidney, and eye, all of which contain cilia. Such diseases are known as "ciliopathies."


    In many of these studies we are using the unicellular Chlamydomonas, a model flagellated organism amenable to biochemical, genetic, and molecular genetic approaches. We recently completed a proteomic analysis of the Chlamydomonas flagellum. This has resulted in a virtual "gold mine" of data that has and will continue to form the basis for many exciting projects. Because the proteins of cilia and flagella have been highly conserved throughout evolution, the human homologues of most of these proteins are readily identified. This opens the door to understanding the functions of many previously uncharacterized ciliary proteins. We currently are investigating the functions of several proteins whose homologues in humans or mice are known to cause disease, including blindness (Leber congenital amaurosis), cystic kidney disease, hydrocephalus, and syndromic ciliopathies such as Bardet-Biedl syndrome and primary ciliary dyskinesia. Typically, we explore the functions of these proteins in Chlamydomonas and then in the mouse to be sure that what we learn from Chlamydomonas is applicable to mammals.


    In addition, many members of this laboratory are participating in a large-scale project to generate and identify insertional Chlamydomonas mutants for all the genes encoding flagellar proteins. Chlamydomonas cells are transformed with a selectable marker that integrates at random into the genome, disrupting any gene at the site of insertion. Using PCR, we can then readily determine the genomic sequence flanking the insert, and thus identify the mutated gene. The mutant can then be characterized structurally and biochemically to understand the function of the mutated gene.


    Finally, we are studying a process called "intraflagellar transport" (IFT), which involves the active movement of multi-subunit protein particles from the base to the tip of the cilium or flagellum, and back to the base again (Fig. 1). These particles carry cargo necessary for assembly and maintenance of the cilium or flagellum, and also transport signals from the cilium or flagellum to the cell body and vice versa (Fig. 2). We are characterizing the motors responsible for this transport, the individual polypeptides that make up the IFT particles, and the proteins and protein complexes that interact with the IFT particle and generally function as cargo adaptors. These studies are providing new insights into a process that is essential for the assembly of almost all cilia and flagella.


    Witman Figure 1 a


    Figure 1. The intraflagellar transport (IFT) machinery. During IFT, linear arrays of IFT particles (yellow) are transported towards the 'plus' (distal) ends of the flagellar outer doublet microtubules (blue) by kinesin-II (pink), and towards the 'minus' (proximal) ends of the microtubules by cytoplasmic dynein 1b (green). The IFT particles, which are composed of at least 19 different proteins, are believed to be carrying precursors that are necessary for the assembly of the flagellar axoneme. The IFT particles are linked to the flagellar membrane (grey lines), and there is evidence that their cargo also includes membrane proteins.


    Witman Figure 2


    Figure 2. IFT and targeting of proteins to the flagellar compartment. Flagellar membrane proteins are carried by vesicles from the Golgi apparatus to the base of the flagellum, where they fuse with the plasma membrane of the cell. In this figure, proteins destined for the flagellar membrane are sorted into specific vesicles that are then targeted to the base of the flagellum. This sorting and targeting appears to be aided by one or more IFT-particle proteins that cycle from the base of the flagellum back through the endomembrane system, where they become associated with the proteins that are destined for the flagellar membrane. Once the vesicle is exocytosed, the IFT-particle proteins, with attached flagellar membrane proteins, become incorporated into IFT particles and are moved through the flagellar pore (involving outer doublet-membrane links in the flagellar transition zone) into the flagellar compartment.




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    Publications listed below are automatically derived from MEDLINE/PubMed and other sources, which might result in incorrect or missing publications. Faculty can login to make corrections and additions.
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    1. Van De Weghe JC, Harris JA, Kubo T, Witman GB, Lechtreck KF. Diffusion rather than IFT likely provides most of the tubulin required for axonemal assembly. J Cell Sci. 2020 Aug 14. PMID: 32801124.
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    2. Picariello T, Hou Y, Kubo T, McNeill NA, Yanagisawa HA, Oda T, Witman GB. TIM, a targeted insertional mutagenesis method utilizing CRISPR/Cas9 in Chlamydomonas reinhardtii. PLoS One. 2020; 15(5):e0232594. PMID: 32401787.
      View in: PubMed
    3. Zhao L, Hou Y, McNeill NA, Witman GB. The unity and diversity of the ciliary central apparatus. Philos Trans R Soc Lond B Biol Sci. 2020 Feb 17; 375(1792):20190164. PMID: 31884923.
      View in: PubMed
    4. Fu G, Zhao L, Dymek E, Hou Y, Song K, Phan N, Shang Z, Smith EF, Witman GB, Nicastro D. Structural organization of the C1a-e-c supercomplex within the ciliary central apparatus. J Cell Biol. 2019 Oct 31. PMID: 31672705.
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    5. Zhao L, Hou Y, Picariello T, Craige B, Witman GB. Proteome of the central apparatus of a ciliary axoneme. J Cell Biol. 2019 May 15. PMID: 31092556.
      View in: PubMed
    6. Picariello T, Brown JM, Hou Y, Swank G, Cochran DA, King OD, Lechtreck K, Pazour GJ, Witman GB. A global analysis of IFT-A function reveals specialization for transport of membrane-associated proteins into cilia. J Cell Sci. 2019 Jan 18. PMID: 30659111.
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    7. Kubo T, Hou Y, Cochran DA, Witman GB, Oda T. A microtubule-dynein tethering complex regulates the axonemal inner dynein f (I1). Mol Biol Cell. 2018 May 01; 29(9):1060-1074. PMID: 29540525.
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    8. Hou Y, Witman GB. The N-terminus of IFT46 mediates intraflagellar transport of outer arm dynein and its cargo-adaptor ODA16. Mol Biol Cell. 2017 Sep 01; 28(18):2420-2433. PMID: 28701346.
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    9. Wingfield JL, Mengoni I, Bomberger H, Jiang YY, Walsh JD, Brown JM, Picariello T, Cochran DA, Zhu B, Pan J, Eggenschwiler J, Gaertig J, Witman GB, Kner P, Lechtreck K. IFT trains in different stages of assembly queue at the ciliary base for consecutive release into the cilium. Elife. 2017 May 31; 6. PMID: 28562242.
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    10. Brown JM, Mosley M, Montes-Berrueta D, Hou Y, Yang F, Scarbrough C, Witman GB, Wirschell M. Characterization of a new oda3 allele, oda3-6, defective in assembly of the outer dynein arm-docking complex in Chlamydomonas reinhardtii. PLoS One. 2017; 12(3):e0173842. PMID: 28291812.
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    11. Kubo T, Brown JM, Bellve K, Craige B, Craft JM, Fogarty K, Lechtreck KF, Witman GB. Together, the IFT81 and IFT74 N-termini form the main module for intraflagellar transport of tubulin. J Cell Sci. 2016 May 15; 129(10):2106-19. PMID: 27068536.
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    12. Schmidts M, Hou Y, Cortés CR, Mans DA, Huber C, Boldt K, Patel M, van Reeuwijk J, Plaza JM, van Beersum SE, Yap ZM, Letteboer SJ, Taylor SP, Herridge W, Johnson CA, Scambler PJ, Ueffing M, Kayserili H, Krakow D, King SM. Corrigendum: TCTEX1D2 mutations underlie Jeune asphyxiating thoracic dystrophy with impaired retrograde intraflagellar transport. Nat Commun. 2016; 7:11270. PMID: 27021811.
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    13. San Agustin JT, Pazour GJ, Witman GB. Intraflagellar transport is essential for mammalian spermiogenesis but is absent in mature sperm. Mol Biol Cell. 2015 Dec 01; 26(24):4358-72. PMID: 26424803.
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    14. Yang TT, Su J, Wang WJ, Craige B, Witman GB, Tsou MF, Liao JC. Superresolution Pattern Recognition Reveals the Architectural Map of the Ciliary Transition Zone. Sci Rep. 2015 Sep 14; 5:14096. PMID: 26365165.
      View in: PubMed
    15. McKenzie CW, Craige B, Kroeger TV, Finn R, Wyatt TA, Sisson JH, Pavlik JA, Strittmatter L, Hendricks GM, Witman GB, Lee L. CFAP54 is required for proper ciliary motility and assembly of the central pair apparatus in mice. Mol Biol Cell. 2015 Sep 15; 26(18):3140-9. PMID: 26224312.
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    16. Kubo T, Hirono M, Aikawa T, Kamiya R, Witman GB. Reduced tubulin polyglutamylation suppresses flagellar shortness in Chlamydomonas. Mol Biol Cell. 2015 Aug 1; 26(15):2810-22. PMID: 26085508.
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    17. Awata J, Song K, Lin J, King SM, Sanderson MJ, Nicastro D, Witman GB. DRC3 connects the N-DRC to dynein g to regulate flagellar waveform. Mol Biol Cell. 2015 Aug 1; 26(15):2788-800. PMID: 26063732.
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    18. Schmidts M, Hou Y, Cortés CR, Mans DA, Huber C, Boldt K, Patel M, van Reeuwijk J, Plaza JM, van Beersum SE, Yap ZM, Letteboer SJ, Taylor SP, Herridge W, Johnson CA, Scambler PJ, Ueffing M, Kayserili H, Krakow D, King SM. TCTEX1D2 mutations underlie Jeune asphyxiating thoracic dystrophy with impaired retrograde intraflagellar transport. Nat Commun. 2015 Jun 05; 6:7074. PMID: 26044572.
      View in: PubMed
    19. Brown JM, Cochran DA, Craige B, Kubo T, Witman GB. Assembly of IFT trains at the ciliary base depends on IFT74. Curr Biol. 2015 Jun 15; 25(12):1583-93. PMID: 26051893.
      View in: PubMed
    20. Damerla RR, Cui C, Gabriel GC, Liu X, Craige B, Gibbs BC, Francis R, Li Y, Chatterjee B, San Agustin JT, Eguether T, Subramanian R, Witman GB, Michaud JL, Pazour GJ, Lo CW. Novel Jbts17 mutant mouse model of Joubert syndrome with cilia transition zone defects and cerebellar and other ciliopathy related anomalies. Hum Mol Genet. 2015 Jul 15; 24(14):3994-4005. PMID: 25877302.
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    21. Hou Y, Witman GB. Dynein and intraflagellar transport. Exp Cell Res. 2015 May 15; 334(1):26-34. PMID: 25725253.
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    22. Song K, Awata J, Tritschler D, Bower R, Witman GB, Porter ME, Nicastro D. In situ localization of N and C termini of subunits of the flagellar nexin-dynein regulatory complex (N-DRC) using SNAP tag and cryo-electron tomography. J Biol Chem. 2015 Feb 27; 290(9):5341-53. PMID: 25564608.
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    23. Brown JM, Witman GB. Cilia and Diseases. Bioscience. 2014 Dec 1; 64(12):1126-1137. PMID: 25960570.
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    24. Craige B, Witman GB. Flipping a phosphate switch on kinesin-II to turn IFT around. Dev Cell. 2014 Sep 8; 30(5):492-3. PMID: 25203204.
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    25. Awata J, Takada S, Standley C, Lechtreck KF, Bellvé KD, Pazour GJ, Fogarty KE, Witman GB. NPHP4 controls ciliary trafficking of membrane proteins and large soluble proteins at the transition zone. J Cell Sci. 2014 Nov 1; 127(Pt 21):4714-27. PMID: 25150219.
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    26. Johnson EA, Rice SL, Preimesberger MR, Nye DB, Gilevicius L, Wenke BB, Brown JM, Witman GB, Lecomte JT. Characterization of THB1, a Chlamydomonas reinhardtii truncated hemoglobin: linkage to nitrogen metabolism and identification of lysine as the distal heme ligand. Biochemistry. 2014 Jul 22; 53(28):4573-89. PMID: 24964018.
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    27. Blaby IK, Blaby-Haas CE, Tourasse N, Hom EF, Lopez D, Aksoy M, Grossman A, Umen J, Dutcher S, Porter M, King S, Witman GB, Stanke M, Harris EH, Goodstein D, Grimwood J, Schmutz J, Vallon O, Merchant SS, Prochnik S. The Chlamydomonas genome project: a decade on. Trends Plant Sci. 2014 Oct; 19(10):672-80. PMID: 24950814.
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    28. Owa M, Furuta A, Usukura J, Arisaka F, King SM, Witman GB, Kamiya R, Wakabayashi K. Cooperative binding of the outer arm-docking complex underlies the regular arrangement of outer arm dynein in the axoneme. Proc Natl Acad Sci U S A. 2014 Jul 1; 111(26):9461-6. PMID: 24979786.
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    29. Lechtreck KF, Gould TJ, Witman GB. Flagellar central pair assembly in Chlamydomonas reinhardtii. Cilia. 2013 Nov 27; 2(1):15. PMID: 24283352.
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    30. Craige B, Brown JM, Witman GB. Isolation of Chlamydomonas flagella. Curr Protoc Cell Biol. 2013 Jun; Chapter 3:Unit 3.41.1-9. PMID: 23728744.
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    31. Lechtreck KF, Brown JM, Sampaio JL, Craft JM, Shevchenko A, Evans JE, Witman GB. Cycling of the signaling protein phospholipase D through cilia requires the BBSome only for the export phase. J Cell Biol. 2013 Apr 15; 201(2):249-61. PMID: 23589493.
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    32. Ludington WB, Wemmer KA, Lechtreck KF, Witman GB, Marshall WF. Avalanche-like behavior in ciliary import. Proc Natl Acad Sci U S A. 2013 Mar 5; 110(10):3925-30. PMID: 23431147.
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    33. Engel BD, Ishikawa H, Wemmer KA, Geimer S, Wakabayashi K, Hirono M, Craige B, Pazour GJ, Witman GB, Kamiya R, Marshall WF. The role of retrograde intraflagellar transport in flagellar assembly, maintenance, and function. J Cell Biol. 2012 Oct 1; 199(1):151-67. PMID: 23027906.
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    34. Brown JM, Dipetrillo CG, Smith EF, Witman GB. A FAP46 mutant provides new insights into the function and assembly of the C1d complex of the ciliary central apparatus. J Cell Sci. 2012 Aug 15; 125(Pt 16):3904-13. PMID: 22573824.
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    35. Hom EF, Witman GB, Harris EH, Dutcher SK, Kamiya R, Mitchell DR, Pazour GJ, Porter ME, Sale WS, Wirschell M, Yagi T, King SM. A unified taxonomy for ciliary dyneins. Cytoskeleton (Hoboken). 2011 Oct; 68(10):555-65. PMID: 21953912.
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    36. Yang Y, Cochran DA, Gargano MD, King I, Samhat NK, Burger BP, Sabourin KR, Hou Y, Awata J, Parry DA, Marshall WF, Witman GB, Lu X. Regulation of flagellar motility by the conserved flagellar protein CG34110/Ccdc135/FAP50. Mol Biol Cell. 2011 Apr; 22(7):976-87. PMID: 21289096.
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    37. Craige B, Tsao CC, Diener DR, Hou Y, Lechtreck KF, Rosenbaum JL, Witman GB. CEP290 tethers flagellar transition zone microtubules to the membrane and regulates flagellar protein content. J Cell Biol. 2010 Sep 6; 190(5):927-40. PMID: 20819941.
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    38. Lechtreck KF, Johnson EC, Sakai T, Cochran D, Ballif BA, Rush J, Pazour GJ, Ikebe M, Witman GB. The Chlamydomonas reinhardtii BBSome is an IFT cargo required for export of specific signaling proteins from flagella. J Cell Biol. 2009 Dec 28; 187(7):1117-32. PMID: 20038682.
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    39. Engel BD, Lechtreck KF, Sakai T, Ikebe M, Witman GB, Marshall WF. Total internal reflection fluorescence (TIRF) microscopy of Chlamydomonas flagella. Methods Cell Biol. 2009; 93:157-77. PMID: 20409817.
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    40. Lechtreck KF, Sanderson MJ, Witman GB. High-speed digital imaging of ependymal cilia in the murine brain. Methods Cell Biol. 2009; 91:255-64. PMID: 20409790.
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    41. Lechtreck KF, Luro S, Awata J, Witman GB. HA-tagging of putative flagellar proteins in Chlamydomonas reinhardtii identifies a novel protein of intraflagellar transport complex B. Cell Motil Cytoskeleton. 2009 Aug; 66(8):469-82. PMID: 19382199.
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    42. Wirschell M, Yang C, Yang P, Fox L, Yanagisawa HA, Kamiya R, Witman GB, Porter ME, Sale WS. IC97 is a novel intermediate chain of I1 dynein that interacts with tubulin and regulates interdoublet sliding. Mol Biol Cell. 2009 Jul; 20(13):3044-54. PMID: 19420136.
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    43. Lechtreck KF, Delmotte P, Robinson ML, Sanderson MJ, Witman GB. Mutations in Hydin impair ciliary motility in mice. J Cell Biol. 2008 Feb 11; 180(3):633-43. PMID: 18250199.
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    44. Huang K, Diener DR, Mitchell A, Pazour GJ, Witman GB, Rosenbaum JL. Function and dynamics of PKD2 in Chlamydomonas reinhardtii flagella. J Cell Biol. 2007 Nov 5; 179(3):501-14. PMID: 17984324.
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    45. Merchant SS, Prochnik SE, Vallon O, Harris EH, Karpowicz SJ, Witman GB, Terry A, Salamov A, Fritz-Laylin LK, Maréchal-Drouard L, Marshall WF, Qu LH, Nelson DR, Sanderfoot AA, Spalding MH, Kapitonov VV, Ren Q, Ferris P, Lindquist E, Shapiro H, Lucas SM, Grimwood J, Schmutz J, Cardol P, Cerutti H, Chanfreau G, Chen CL, Cognat V, Croft MT, Dent R, Dutcher S, Fernández E, Fukuzawa H, González-Ballester D, González-Halphen D, Hallmann A, Hanikenne M, Hippler M, Inwood W, Jabbari K, Kalanon M, Kuras R, Lefebvre PA, Lemaire SD, Lobanov AV, Lohr M, Manuell A, Meier I, Mets L, Mittag M, Mittelmeier T, Moroney JV, Moseley J, Napoli C, Nedelcu AM, Niyogi K, Novoselov SV, Paulsen IT, Pazour G, Purton S, Ral JP, Riaño-Pachón DM, Riekhof W, Rymarquis L, Schroda M, Stern D, Umen J, Willows R, Wilson N, Zimmer SL, Allmer J, Balk J, Bisova K, Chen CJ, Elias M, Gendler K, Hauser C, Lamb MR, Ledford H, Long JC, Minagawa J, Page MD, Pan J, Pootakham W, Roje S, Rose A, Stahlberg E, Terauchi AM, Yang P, Ball S, Bowler C, Dieckmann CL, Gladyshev VN, Green P, Jorgensen R, Mayfield S, Mueller-Roeber B, Rajamani S, Sayre RT, Brokstein P, Dubchak I, Goodstein D, Hornick L, Huang YW, Jhaveri J, Luo Y, Martínez D, Ngau WC, Otillar B, Poliakov A, Porter A, Szajkowski L, Werner G, Zhou K, Grigoriev IV, Rokhsar DS, Grossman AR. The Chlamydomonas genome reveals the evolution of key animal and plant functions. Science. 2007 Oct 12; 318(5848):245-50. PMID: 17932292.
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    46. Hou Y, Qin H, Follit JA, Pazour GJ, Rosenbaum JL, Witman GB. Functional analysis of an individual IFT protein: IFT46 is required for transport of outer dynein arms into flagella. J Cell Biol. 2007 Feb 26; 176(5):653-65. PMID: 17312020.
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    47. Lechtreck KF, Witman GB. Chlamydomonas reinhardtii hydin is a central pair protein required for flagellar motility. J Cell Biol. 2007 Feb 12; 176(4):473-82. PMID: 17296796.
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    48. Yang P, Diener DR, Yang C, Kohno T, Pazour GJ, Dienes JM, Agrin NS, King SM, Sale WS, Kamiya R, Rosenbaum JL, Witman GB. Radial spoke proteins of Chlamydomonas flagella. J Cell Sci. 2006 Mar 15; 119(Pt 6):1165-74. PMID: 16507594.
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    49. Pfister KK, Fisher EM, Gibbons IR, Hays TS, Holzbaur EL, McIntosh JR, Porter ME, Schroer TA, Vaughan KT, Witman GB, King SM, Vallee RB. Cytoplasmic dynein nomenclature. J Cell Biol. 2005 Nov 7; 171(3):411-3. PMID: 16260502.
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    50. DiBella LM, Gorbatyuk O, Sakato M, Wakabayashi K, Patel-King RS, Pazour GJ, Witman GB, King SM. Differential light chain assembly influences outer arm dynein motor function. Mol Biol Cell. 2005 Dec; 16(12):5661-74. PMID: 16195342.
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    51. Pazour GJ, Agrin N, Leszyk J, Witman GB. Proteomic analysis of a eukaryotic cilium. J Cell Biol. 2005 Jul 4; 170(1):103-13. PMID: 15998802.
      View in: PubMed
    52. Wang H, San Agustin JT, Witman GB, Kilpatrick DL. Novel role for a sterol response element binding protein in directing spermatogenic cell-specific gene expression. Mol Cell Biol. 2004 Dec; 24(24):10681-8. PMID: 15572673.
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    53. Jurczyk A, Gromley A, Redick S, San Agustin J, Witman G, Pazour GJ, Peters DJ, Doxsey S. Pericentrin forms a complex with intraflagellar transport proteins and polycystin-2 and is required for primary cilia assembly. J Cell Biol. 2004 Aug 30; 166(5):637-43. PMID: 15337773.
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    54. Hou Y, Pazour GJ, Witman GB. A dynein light intermediate chain, D1bLIC, is required for retrograde intraflagellar transport. Mol Biol Cell. 2004 Oct; 15(10):4382-94. PMID: 15269286.
      View in: PubMed
    55. Wirschell M, Pazour G, Yoda A, Hirono M, Kamiya R, Witman GB. Oda5p, a novel axonemal protein required for assembly of the outer dynein arm and an associated adenylate kinase. Mol Biol Cell. 2004 Jun; 15(6):2729-41. PMID: 15064350.
      View in: PubMed
    56. Wang S, Luo Y, Wilson PD, Witman GB, Zhou J. The autosomal recessive polycystic kidney disease protein is localized to primary cilia, with concentration in the basal body area. J Am Soc Nephrol. 2004 Mar; 15(3):592-602. PMID: 14978161.
      View in: PubMed
    57. Witman GB. Cell motility: deaf Drosophila keep the beat. Curr Biol. 2003 Oct 14; 13(20):R796-8. PMID: 14561418.
      View in: PubMed
    58. Casey DM, Yagi T, Kamiya R, Witman GB. DC3, the smallest subunit of the Chlamydomonas flagellar outer dynein arm-docking complex, is a redox-sensitive calcium-binding protein. J Biol Chem. 2003 Oct 24; 278(43):42652-9. PMID: 12920131.
      View in: PubMed
    59. Casey DM, Inaba K, Pazour GJ, Takada S, Wakabayashi K, Wilkerson CG, Kamiya R, Witman GB. DC3, the 21-kDa subunit of the outer dynein arm-docking complex (ODA-DC), is a novel EF-hand protein important for assembly of both the outer arm and the ODA-DC. Mol Biol Cell. 2003 Sep; 14(9):3650-63. PMID: 12972554.
      View in: PubMed
    60. Pazour GJ, Witman GB. The vertebrate primary cilium is a sensory organelle. Curr Opin Cell Biol. 2003 Feb; 15(1):105-10. PMID: 12517711.
      View in: PubMed
    61. Rosenbaum JL, Witman GB. Intraflagellar transport. Nat Rev Mol Cell Biol. 2002 Nov; 3(11):813-25. PMID: 12415299.
      View in: PubMed
    62. Pazour GJ, San Agustin JT, Follit JA, Rosenbaum JL, Witman GB. Polycystin-2 localizes to kidney cilia and the ciliary level is elevated in orpk mice with polycystic kidney disease. Curr Biol. 2002 Jun 4; 12(11):R378-80. PMID: 12062067.
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    63. Pazour GJ, Baker SA, Deane JA, Cole DG, Dickert BL, Rosenbaum JL, Witman GB, Besharse JC. The intraflagellar transport protein, IFT88, is essential for vertebrate photoreceptor assembly and maintenance. J Cell Biol. 2002 Apr 1; 157(1):103-13. PMID: 11916979.
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    64. Takada S, Wilkerson CG, Wakabayashi K, Kamiya R, Witman GB. The outer dynein arm-docking complex: composition and characterization of a subunit (oda1) necessary for outer arm assembly. Mol Biol Cell. 2002 Mar; 13(3):1015-29. PMID: 11907279.
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