Gene Symbol: pbl
Description: pebble
Alias: 0083/20, 0293/09, 0542/03, CG8114, Dmel\CG8114, ECT2, Ect-2, GEF, PBL, Pbl, Pbl/Ect2, anon-WO0172774.22, anon-WO0172774.25, l(3)09645, pebble, CG8114-PA, CG8114-PB, CG8114-PC, CG8114-PD, CG8114-PE, CG8114-PF, pbl-PA, pbl-PB, pbl-PC, pbl-PD, pbl-PE, pbl-PF
Species: fruit fly
Products:     pbl

Top Publications

  1. Hime G, Saint R. Zygotic expression of the pebble locus is required for cytokinesis during the postblastoderm mitoses of Drosophila. Development. 1992;114:165-71 pubmed
    Mutations at the pebble locus of Drosophila melanogaster result in embryonic lethality. Examination of homozygous mutant embryos at the end of embryogenesis revealed the presence of fewer and larger cells which contained enlarged nuclei...
  2. Schumacher S, Gryzik T, Tannebaum S, Müller H. The RhoGEF Pebble is required for cell shape changes during cell migration triggered by the Drosophila FGF receptor Heartless. Development. 2004;131:2631-40 pubmed
    ..In a genetic screen we identified pebble (pbl) as a novel gene required for mesoderm migration...
  3. Dean S, Rogers S, Stuurman N, Vale R, Spudich J. Distinct pathways control recruitment and maintenance of myosin II at the cleavage furrow during cytokinesis. Proc Natl Acad Sci U S A. 2005;102:13473-8 pubmed
    ..Three genes in the rho1 signaling pathway, pebble (a Rho guanidine nucleotide exchange factor), rho1, and rho kinase, are required for the initial recruitment of ..
  4. Koizumi K, Higashida H, Yoo S, Islam M, Ivanov A, Guo V, et al. RNA interference screen to identify genes required for Drosophila embryonic nervous system development. Proc Natl Acad Sci U S A. 2007;104:5626-31 pubmed
    ..The total number of dsRNAs that we have tested for an RNAi-induced phenotype affecting the embryonic nervous system, including our previous study, is 7,312, which corresponds to approximately 50% of the genes in the Drosophila genome. ..
  5. Hickson G, Echard A, O Farrell P. Rho-kinase controls cell shape changes during cytokinesis. Curr Biol. 2006;16:359-70 pubmed
    ..sensitivity of cell elongation to Rok depletion was highlighted by RNAi to other genes in the Rho pathway, such as pebble, racGAP50C, and diaphanous, which had profound effects on furrowing but lesser effects on elongation...
  6. Prokopenko S, Brumby A, O KEEFE L, Prior L, He Y, Saint R, et al. A putative exchange factor for Rho1 GTPase is required for initiation of cytokinesis in Drosophila. Genes Dev. 1999;13:2301-14 pubmed
    ..Here we show that in Drosophila, Pebble (Pbl), a putative exchange factor for Rho GTPases (RhoGEF), is required for the formation of the contractile ring ..
  7. Smallhorn M, Murray M, Saint R. The epithelial-mesenchymal transition of the Drosophila mesoderm requires the Rho GTP exchange factor Pebble. Development. 2004;131:2641-51 pubmed
    Drosophila pebble (pbl) encodes a Rho-family GTP exchange factor (GEF) required for cytokinesis...
  8. Echard A, O Farrell P. The degradation of two mitotic cyclins contributes to the timing of cytokinesis. Curr Biol. 2003;13:373-83 pubmed
    ..Reduced function of Pebble, a Rho GEF required for cytokinesis, also delayed and slowed furrow invagination, but incomplete furrows were ..
  9. Somma M, Fasulo B, Cenci G, Cundari E, Gatti M. Molecular dissection of cytokinesis by RNA interference in Drosophila cultured cells. Mol Biol Cell. 2002;13:2448-60 pubmed
    ..We show that double-stranded RNAs for anillin, acGAP, pavarotti, rho1, pebble, spaghetti squash, syntaxin1A, and twinstar all disrupt cytokinesis in S2 tissue culture cells, causing gene-..

More Information


  1. Zavortink M, Contreras N, Addy T, Bejsovec A, Saint R. Tum/RacGAP50C provides a critical link between anaphase microtubules and the assembly of the contractile ring in Drosophila melanogaster. J Cell Sci. 2005;118:5381-92 pubmed
    ..embryos we show that cells lacking Tum do not form furrows and fail to localise the key cytokinetic components Pebble (a RhoGEF), Aurora B kinase, Diaphanous, Pav-KLP and Anillin...
  2. Prokopenko S, Saint R, Bellen H. Untying the Gordian knot of cytokinesis. Role of small G proteins and their regulators. J Cell Biol. 2000;148:843-8 pubmed
  3. Carreno S, Kouranti I, Glusman E, Fuller M, Echard A, Payre F. Moesin and its activating kinase Slik are required for cortical stability and microtubule organization in mitotic cells. J Cell Biol. 2008;180:739-46 pubmed publisher
    ..We propose that the spatiotemporal control of moesin activation at the mitotic cortex provides localized cues to coordinate cortical contractility and microtubule interactions during cell division. ..
  4. Lehner C. The pebble gene is required for cytokinesis in Drosophila. J Cell Sci. 1992;103 ( Pt 4):1021-30 pubmed
    ..However, cytokinesis never occurs in somatic cells of embryos homozygous or transheterozygous for mutations in the pebble gene...
  5. Prokopenko S, Saint R, Bellen H. Tissue distribution of PEBBLE RNA and pebble protein during Drosophila embryonic development. Mech Dev. 2000;90:269-73 pubmed
    b>pebble (pbl) is required for cytokinesis during postblastoderm mitoses (Hime, G., Saint, R., 1992. Zygotic expression of the pebble locus is required for cytokinesis during the postblastoderm mitoses of Drosophila...
  6. D Avino P, Savoian M, Glover D. Cleavage furrow formation and ingression during animal cytokinesis: a microtubule legacy. J Cell Sci. 2005;118:1549-58 pubmed
    ..The same mechanism might be conserved in other organisms. ..
  7. Eggert U, Kiger A, Richter C, Perlman Z, Perrimon N, Mitchison T, et al. Parallel chemical genetic and genome-wide RNAi screens identify cytokinesis inhibitors and targets. PLoS Biol. 2004;2:e379 pubmed
    ..Our study shows that parallel RNA interference and small molecule screening is a generally useful approach to identifying active small molecules and their target pathways...
  8. Giansanti M, Farkas R, Bonaccorsi S, Lindsley D, Wakimoto B, Fuller M, et al. Genetic dissection of meiotic cytokinesis in Drosophila males. Mol Biol Cell. 2004;15:2509-22 pubmed
    ..These include 16 novel loci and three genes, diaphanous, four wheel drive, and pebble, already known to be involved in Drosophila cytokinesis...
  9. Ebrahimi S, Fraval H, Murray M, Saint R, Gregory S. Polo kinase interacts with RacGAP50C and is required to localize the cytokinesis initiation complex. J Biol Chem. 2010;285:28667-73 pubmed publisher
    ..between a microtubule-associated kinesin-like protein (KLP), a member of the RacGAP family, and the RhoGEF Pebble. Recently, the activity of the mammalian Polo kinase ortholog Plk1 has been implicated in the formation of this ..
  10. Somers W, Saint R. A RhoGEF and Rho family GTPase-activating protein complex links the contractile ring to cortical microtubules at the onset of cytokinesis. Dev Cell. 2003;4:29-39 pubmed
    ..We show that an interaction between the Rho GTP exchange factor, Pebble, and the Rho family GTPase-activating protein, RacGAP50C, connects the contractile ring to cortical microtubules ..
  11. van Impel A, Schumacher S, Draga M, Herz H, Grosshans J, Müller H. Regulation of the Rac GTPase pathway by the multifunctional Rho GEF Pebble is essential for mesoderm migration in the Drosophila gastrula. Development. 2009;136:813-22 pubmed publisher
    The Drosophila guanine nucleotide exchange factor Pebble (Pbl) is essential for cytokinesis and cell migration during gastrulation...
  12. Albertson R, Cao J, Hsieh T, Sullivan W. Vesicles and actin are targeted to the cleavage furrow via furrow microtubules and the central spindle. J Cell Biol. 2008;181:777-90 pubmed publisher
    ..We find that Rho-specific guanine nucleotide exchange factor mutants, pebble (pbl), severely disrupt this F-actin-associated vesicle transport...
  13. Burgio G, La Rocca G, Sala A, Arancio W, Di Gesù D, Collesano M, et al. Genetic identification of a network of factors that functionally interact with the nucleosome remodeling ATPase ISWI. PLoS Genet. 2008;4:e1000089 pubmed publisher
    ..Consistent with these findings, the acetylation of histone H4 is altered when ISWI activity is perturbed in vivo. These findings suggest that ISWI associates with the Sin3A/Rpd3 complex to support its function in vivo. ..
  14. Giansanti M, Bonaccorsi S, Kurek R, Farkas R, Dimitri P, Fuller M, et al. The class I PITP giotto is required for Drosophila cytokinesis. Curr Biol. 2006;16:195-201 pubmed
    ..This change in membrane composition would ultimately facilitate vesicle fusion, allowing membrane addition to the furrow and/or targeted delivery of proteins required for cytokinesis. ..
  15. Nakamura M, Matsumoto K, Iwamoto Y, Muguruma T, Nakazawa N, Hatori R, et al. Reduced cell number in the hindgut epithelium disrupts hindgut left-right asymmetry in a mutant of pebble, encoding a RhoGEF, in Drosophila embryos. Mech Dev. 2013;130:169-80 pubmed publisher
    ..From this screen, we isolated pebble (pbl), which encodes a homolog of a mammalian RhoGEF, Ect2...
  16. Das A, Shah S, Fan B, Paik D, DiSanto D, Hinman A, et al. Spindle Assembly and Chromosome Segregation Requires Central Spindle Proteins in Drosophila Oocytes. Genetics. 2016;202:61-75 pubmed publisher
    ..In combination our results support a model where the meiotic central spindle and associated proteins are essential for acentrosomal chromosome segregation. ..
  17. Tolwinski N. New advances in signaling and pattern formation. Fly (Austin). 2007;1:116-7 pubmed
  18. Ng J, Luo L. Rho GTPases regulate axon growth through convergent and divergent signaling pathways. Neuron. 2004;44:779-93 pubmed
    ..These genetic analyses reveal convergent and divergent pathways from Rho GTPases to the cytoskeleton during axon growth in vivo and suggest that different developmental outcomes could be achieved by biases in pathway selection...
  19. Menant A, Karess R. Inducing "cytokinesis" without mitosis in unfertilized Drosophila eggs. Cell Cycle. 2012;11:2856-63 pubmed publisher
  20. ROBINETT C, Giansanti M, Gatti M, Fuller M. TRAPPII is required for cleavage furrow ingression and localization of Rab11 in dividing male meiotic cells of Drosophila. J Cell Sci. 2009;122:4526-34 pubmed publisher
    ..These results suggest that Bru cooperates with Rab11 and PI4Kbeta to regulate the efficiency of membrane addition to the cleavage furrow, thus promoting cytokinesis in Drosophila male meiotic cells. ..
  21. Murray M, Southall T, Liu W, Fraval H, Lorensuhewa N, Brand A, et al. Snail-dependent repression of the RhoGEF pebble is required for gastrulation consistency in Drosophila melanogaster. Dev Genes Evol. 2012;222:361-8 pubmed publisher
    The Rho GTP exchange factor, Pebble (Pbl), long recognised as an essential activator of Rho during cytokinesis, also regulates mesoderm migration at gastrulation...
  22. Weigmann K, Lehner C. Cell fate specification by even-skipped expression in the Drosophila nervous system is coupled to cell cycle progression. Development. 1995;121:3713-21 pubmed
    ..during the first NB division is not required for even-skipped expression as revealed by observations in pebble mutant embryos...
  23. Akiyama Oda Y, Hotta Y, Tsukita S, Oda H. Distinct mechanisms triggering glial differentiation in Drosophila thoracic and abdominal neuroblasts 6-4. Dev Biol. 2000;222:429-39 pubmed
    ..These results showed that NB6-4T, but not NB6-4A, requires cell cycle progression for acquisition of glial fate, suggesting that distinct mechanisms trigger glial differentiation in the different lineages. ..
  24. Murray M, Ng M, Fraval H, Tan J, Liu W, Smallhorn M, et al. Regulation of Drosophila mesoderm migration by phosphoinositides and the PH domain of the Rho GTP exchange factor Pebble. Dev Biol. 2012;372:17-27 pubmed publisher
    The Drosophila RhoGEF Pebble (Pbl) is required for cytokinesis and migration of mesodermal cells...
  25. Duan G, Saint R, Helliwell C, Behm C, Wang M, Waterhouse P, et al. C. elegans RNA-dependent RNA polymerases rrf-1 and ego-1 silence Drosophila transgenes by differing mechanisms. Cell Mol Life Sci. 2013;70:1469-81 pubmed publisher
  26. Bateman J, Larschan E, D Souza R, Marshall L, Dempsey K, Johnson J, et al. A genome-wide screen identifies genes that affect somatic homolog pairing in Drosophila. G3 (Bethesda). 2012;2:731-40 pubmed publisher
    ..Consistent with our results in cultured cells, a classical mutation in one of the strongest candidate genes, pebble (pbl), causes a decrease in somatic homolog pairing in developing embryos...
  27. Yamamoto R, Axton J, Yamamoto Y, Saunders R, Glover D, Henderson D. The Drosophila mus101 gene, which links DNA repair, replication and condensation of heterochromatin in mitosis, encodes a protein with seven BRCA1 C-terminus domains. Genetics. 2000;156:711-21 pubmed
    ..Mus101 also shares sequence similarity with the fission yeast Rad4/Cut5 protein required for repair, replication, and checkpoint control, suggesting that the two proteins may be functional homologs. ..
  28. Grosskortenhaus R, Pearson B, Marusich A, Doe C. Regulation of temporal identity transitions in Drosophila neuroblasts. Dev Cell. 2005;8:193-202 pubmed
    ..We conclude that two distinct "timers" regulate neuroblast gene expression: a hunchback --> Kruppel timer requiring cytokinesis, and a Kruppel --> pdm1 --> castor timer which is cell cycle independent. ..
  29. Allada R, White N, So W, Hall J, Rosbash M. A mutant Drosophila homolog of mammalian Clock disrupts circadian rhythms and transcription of period and timeless. Cell. 1998;93:791-804 pubmed
    ..The remarkable sequence conservation strongly supports common clock components present in the common ancestor of Drosophila and mammals. ..
  30. Vervoort M, Merritt D, Ghysen A, Dambly Chaudiere C. Genetic basis of the formation and identity of type I and type II neurons in Drosophila embryos. Development. 1997;124:2819-28 pubmed
  31. Kitazawa D, Matsuo T, Kaizuka K, Miyauchi C, Hayashi D, Inoue Y. Orbit/CLASP is required for myosin accumulation at the cleavage furrow in Drosophila male meiosis. PLoS ONE. 2014;9:e93669 pubmed publisher
    ..Orbit was also necessary for the maintenance of the CR. Our data suggest that Orbit plays an essential role as a connector between MTs and the CR in Drosophila male meiosis. ..
  32. Pandey R, Heidmann S, Lehner C. Epithelial re-organization and dynamics of progression through mitosis in Drosophila separase complex mutants. J Cell Sci. 2005;118:733-42 pubmed
  33. Kotadia S, Montembault E, Sullivan W, Royou A. Cell elongation is an adaptive response for clearing long chromatid arms from the cleavage plane. J Cell Biol. 2012;199:745-53 pubmed publisher
    ..Rho guanine-nucleotide exchange factor (Pebble)-depleted cells failed to elongate during segregation of long chromatids...
  34. Reiter L, Seagroves T, Bowers M, Bier E. Expression of the Rho-GEF Pbl/ECT2 is regulated by the UBE3A E3 ubiquitin ligase. Hum Mol Genet. 2006;15:2825-35 pubmed
    ..One protein responsive to UBE3A was the Rho-GEF pebble (pbl). Here, we present three lines of evidence suggesting that UBE3A regulates Pbl...
  35. Choi K, Benzer S. Rotation of photoreceptor clusters in the developing Drosophila eye requires the nemo gene. Cell. 1994;78:125-36 pubmed
    ..In another mutant, roulette, excessive rotation through varying angles occurs in many ommatidia. This defect is suppressed by nmo, indicating that nmo acts upstream in a rotation-regulating pathway. ..
  36. CUKIER H, Perez A, Collins A, Zhou Z, Zoghbi H, Botas J. Genetic modifiers of MeCP2 function in Drosophila. PLoS Genet. 2008;4:e1000179 pubmed publisher
    ..Additional sex combs, corto, osa, Sex combs on midleg, and trithorax), the kinase tricornered, the UBE3A target pebble, and Drosophila homologues of the MeCP2 physical interactors Sin3a, REST, and N-CoR...
  37. Cui X, Doe C. The role of the cell cycle and cytokinesis in regulating neuroblast sublineage gene expression in the Drosophila CNS. Development. 1995;121:3233-43 pubmed
    ..cytokinesis and cell cycle progression (string mutants) and in embryos lacking only neuroblast cytokinesis (pebble mutants)...
  38. Rosa A, Vlassaks E, Pichaud F, Baum B. Ect2/Pbl acts via Rho and polarity proteins to direct the assembly of an isotropic actomyosin cortex upon mitotic entry. Dev Cell. 2015;32:604-16 pubmed publisher
    ..Here, we delineate a pathway downstream of the RhoGEF Pbl/Ect2 that directs this process in a model epithelium...
  39. Castoreno A, Smurnyy Y, Torres A, Vokes M, Jones T, Carpenter A, et al. Small molecules discovered in a pathway screen target the Rho pathway in cytokinesis. Nat Chem Biol. 2010;6:457-63 pubmed publisher
    ..Rhodblocks differentially perturb Rho pathway proteins in cells and can be used to dissect the mechanism of the Rho pathway during cytokinesis. ..
  40. Del Castillo U, Lu W, Winding M, Lakonishok M, Gelfand V. Pavarotti/MKLP1 regulates microtubule sliding and neurite outgrowth in Drosophila neurons. Curr Biol. 2015;25:200-5 pubmed publisher
    ..These data suggest an important parallel between long microtubule-microtubule sliding in anaphase B and sliding of interphase microtubules during neurite formation. ..
  41. Miller K, Kiehart D. Fly division. J Cell Biol. 1995;131:1-5 pubmed
  42. Gregory S, Lorensuhewa N, Saint R. Signalling through the RhoGEF Pebble in Drosophila. IUBMB Life. 2010;62:290-5 pubmed publisher
    ..Here we review the role of the Drosophila RhoGEF, Pebble (the Drosophila ortholog of mammalian ECT2)...
  43. Taniguchi K, Kokuryo A, Imano T, Minami R, Nakagoshi H, Adachi Yamada T. Isoform-specific functions of Mud/NuMA mediate binucleation of Drosophila male accessory gland cells. BMC Dev Biol. 2014;14:46 pubmed publisher
    ..We also propose that Mud(S) is a key regulator triggering cytokinesis skipping in binucleation processes. ..
  44. Jankovics F, Henn L, Bujna Ã, Vilmos P, Kiss N, Erdélyi M. A functional genomic screen combined with time-lapse microscopy uncovers a novel set of genes involved in dorsal closure of Drosophila embryos. PLoS ONE. 2011;6:e22229 pubmed publisher
    ..One of the novel dorsal closure genes, the small GTPase activator pebble (pbl), was selected for detailed analysis...
  45. Axelsson E, Sandmann T, Horn T, Boutros M, Huber W, Fischer B. Extracting quantitative genetic interaction phenotypes from matrix combinatorial RNAi. BMC Bioinformatics. 2011;12:342 pubmed publisher
    ..The reliable extraction of presence, absence, and strength of interactions provides insights into molecular mechanisms. ..
  46. Shandala T, Gregory S, Dalton H, Smallhorn M, Saint R. Citron kinase is an essential effector of the Pbl-activated Rho signalling pathway in Drosophila melanogaster. Development. 2004;131:5053-63 pubmed
    b>Pebble (Pbl)-activated RhoA signalling is essential for cytokinesis in Drosophila melanogaster. Here we report that the Drosophila citron gene encodes an essential effector kinase of Pbl-RhoA signalling in vivo...
  47. Sechi S, Frappaolo A, Belloni G, Giansanti M. The roles of the oncoprotein GOLPH3 in contractile ring assembly and membrane trafficking during cytokinesis. Biochem Soc Trans. 2015;43:117-21 pubmed publisher
    ..Because cytokinesis failures have been linked with pre-malignant disease and cancer, the novel connection between GOLPH3 and cytokinesis imposes new fields of investigation in cancer biology and therapy. ..
  48. Nakamura M, Verboon J, Parkhurst S. Prepatterning by RhoGEFs governs Rho GTPase spatiotemporal dynamics during wound repair. J Cell Biol. 2017;216:3959-3969 pubmed publisher
    ..Thus, Annexin proteins may link membrane resealing to cytoskeletal remodeling processes in single cell wound repair. ..
  49. Crest J, Concha Moore K, Sullivan W. RhoGEF and positioning of rappaport-like furrows in the early Drosophila embryo. Curr Biol. 2012;22:2037-41 pubmed publisher
    ..Unlike the zygotic, central spindle-localized RhoGEF (Pebble), RhoGEF2 localizes to metaphase furrows, a function distinct from RhoGEF/Pebble and likely due to the absence of ..
  50. Brumby A, Goulding K, Schlosser T, Loi S, Galea R, Khoo P, et al. Identification of novel Ras-cooperating oncogenes in Drosophila melanogaster: a RhoGEF/Rho-family/JNK pathway is a central driver of tumorigenesis. Genetics. 2011;188:105-25 pubmed publisher
    ..Ras(ACT)-cooperating genes identified were Rac1 Rho1, RhoGEF2, pbl, rib, and east, which encode cell morphology regulators...
  51. Montembault E, Zhang W, Przewloka M, Archambault V, Sevin E, Laue E, et al. Nessun Dorma, a novel centralspindlin partner, is required for cytokinesis in Drosophila spermatocytes. J Cell Biol. 2010;191:1351-65 pubmed publisher
    ..Our findings indicate that Nesd is a novel carbohydrate-binding protein that functions together with centralspindlin in late cytokinesis, thus highlighting the importance of glycosylation in this process...
  52. Slack C, Somers W, Sousa Nunes R, Chia W, Overton P. A mosaic genetic screen for novel mutations affecting Drosophila neuroblast divisions. BMC Genet. 2006;7:33 pubmed
    ..Taken together, these results demonstrate the value of mosaic screens in the identification of genes involved in neuroblast division. ..
  53. Subramanian A, Prokop A, Yamamoto M, Sugimura K, Uemura T, Betschinger J, et al. Shortstop recruits EB1/APC1 and promotes microtubule assembly at the muscle-tendon junction. Curr Biol. 2003;13:1086-95 pubmed
    ..This is achieved by Shot association with the cytoplasmic faces of the basal hemiadherens junction and with the EB1/APC1 complex. ..
  54. Gonzalez C, Tavosanis G, Mollinari C. Centrosomes and microtubule organisation during Drosophila development. J Cell Sci. 1998;111 ( Pt 18):2697-706 pubmed
    ..In this review we present some of the many examples which illustrate how centrosomes and microtubule organisation changes through development in Drosophila and discuss some of its implications. ..
  55. Saint R, Patterson B. Zygotic transcription and cell proliferation during Drosophila embryogenesis. Genetica. 1993;90:157-63 pubmed
    ..Genes that comprise the first class are involved in the developmental control of the cell cycle, while those of the second class identify components of the cell cycle machinery. ..
  56. Greer E, Chao A, Bejsovec A. Pebble/ECT2 RhoGEF negatively regulates the Wingless/Wnt signaling pathway. Development. 2013;140:4937-46 pubmed publisher
    ..mechanism for regulating the Wnt pathway: activity of a Rho guanine nucleotide exchange factor (GEF) encoded by pebble (pbl) in Drosophila and ECT2 in humans...
  57. Schotman H, Karhinen L, Rabouille C. Integrins mediate their unconventional, mechanical-stress-induced secretion via RhoA and PINCH in Drosophila. J Cell Sci. 2009;122:2662-72 pubmed publisher
  58. Broihier H, Skeath J. Drosophila homeodomain protein dHb9 directs neuronal fate via crossrepressive and cell-nonautonomous mechanisms. Neuron. 2002;35:39-50 pubmed
    ..This mutually antagonistic relationship bears similarity to the crossrepressive relationships between pairs of HD proteins that pattern the vertebrate neural tube. ..
  59. Bowman A, Kamal A, Ritchings B, Philp A, McGrail M, Gindhart J, et al. Kinesin-dependent axonal transport is mediated by the sunday driver (SYD) protein. Cell. 2000;103:583-94 pubmed
    ..We propose that SYD mediates the axonal transport of at least one class of vesicles by interacting directly with KLC. ..