Signaling mechanisms that regulate actin-based motility processes in the nervous system

Journal of Neurochemistry

Volumn 83, Issue 3, 2002, Pages 490-503

  Meyer, Gary ^a   Feldman, Eva L ^a  

^a UNIVERSITY OF MICHIGAN MEDICAL SCHOOL   (United States)

Author keywords

Actin depolymerizing factor (ADF) cofilin; Actin related protein Arp2 3; Ena VASP; LIM kinase; Rho GTPase

Indexed keywords

ACTIN; ACTIN DEPOLYMERIZING FACTOR; COFILIN; GUANOSINE TRIPHOSPHATASE; PHOSPHOTRANSFERASE; PROTEIN CDC42; RAC PROTEIN; RHO FACTOR;

ACTIN POLYMERIZATION; CELL MIGRATION; CYTOSKELETON; ENZYME ACTIVITY; HYDROLYSIS; NERVE CELL; NERVOUS SYSTEM; NERVOUS SYSTEM DEVELOPMENT; NEURITE; NONHUMAN; PRIORITY JOURNAL; PROTEIN ANALYSIS; REGULATORY MECHANISM; REVIEW; SIGNAL TRANSDUCTION;

ACTINS; ANIMALS; CELL MOVEMENT; GROWTH CONES; HUMANS; NERVOUS SYSTEM; PSEUDOPODIA; RHO GTP-BINDING PROTEINS; SIGNAL TRANSDUCTION;

ENA;

EID: 0036828324     PISSN: 00223042     EISSN: None     Source Type: Journal    
DOI: 10.1046/j.1471-4159.2002.01185.x     Document Type: Review

References (129)

1. Adams A. E., Johnson D. I., Longnecker R. M., Sloat B. F. and Pringle J. R. (1990) CDC42 and CDC43, two additional genes involved in budding and the establishment of cell polarity in the yeast Saccharomyces cerevisiae. J. Cell Biol. 111, 131-142.
2. Aizawa H., Wakatsuki S., Ishii A., Moriyama K., Sasaki Y., Ohashi K., Sekine-Aizawa Y., Sehara-Fujisawa A., Mizuno K., Goshima Y. and Yahara I. (2001) Phosphorylation of cofilin by LIM-kinase is necessary for semaphorin 3A-induced growth cone collapse. Nat. Neurosci. 4, 367-373.
3. Altun-Gultekin Z. F. and Wagner J. A. (1996) Src, ras and rac mediate the migratory response elicited by NGF and PMA in PC cells. J. Neurosci. Res. 44, 308-327.
4. Amano M., Chihara K., Kimura K., Fukata Y., Nakamura N., Matsuura Y. and Kaibuchi K. (1997) Formation of actin stress fibers and focal adhesions enhanced by Rho-kinase. Science 275, 1308-1311.
5. Ambach A., Saunus J., Konstandin M., Wesselborg S., Meuer S. C. and Samstag Y. (2000) The serine phosphatases PP1 and PP2A associate with and activate the actin-binding protein cofilin in human T lymphocytes. Eur. J. Immunol. 30, 3422-3431.
6. Anderson P. S. and Lacal J. C. (1987) Expression of the Aplysia californica rho gene in Escherichia coli: Purification and characterization of its encoded p21 product. Mol. Cell. Biol. 7, 3620-3628.
7. Arber S., Barbayannis F. A., Hanser H., Schneider C., Stanyon C. A., Bernard O. and Caroni P. (1998) Regulation of actin dynamics through phosphorylation of cofilin by LIM-kinase. Nature 393, 805-809.
8. Bagrodia, S., Derijard B., Davis R. J. and Cerione R. A. (1995) Cdc42 and PAK-mediated signaling leads to jun kinase and p38 mitogen-activated protein kinase activation. J. Biol. Chem. 270, 27995-27998.
9. Bamburg J. R. (1999) Proteins of the ADF/cofilin family: Essential regulators of actin dynamics. Annu. Rev. Cell. Dev. Biol. 15, 185-230.
10. Bamburg J. R., Harris H. E. and Weeds A. G. (1980) Partial purification and characterization of an actin depolymerizing factor from brain. FEBS Lett. 121, 178-182.
11. Banzai Y., Miki H., Yamaguchi H. and Takenawa T. (2000) Essential role of neural Wiskott-Aldrich syndrome protein in neurite extension in PC12 cells and rat hippocampal primary culture cells. J. Biol. Chem. 275, 11987-11992.
12. Bashaw G. J., Hu H., Nobes C. D. and Goodman C. S. (2001) A novel Db1 family RhoGEF promotes Rho-dependent axon attraction to the central nervous system midline in Drosophila and overcomes Robo repulsion. J. Cell Biol. 155, 1117-1122.
13. Bateman J. and Van Vactor D. (2001) The Trio family of guanine-nucleotide-exchange factors: Regulators of axon guidance. J. Cell Sci. 114, 1973-1980.
14. Bateman J., Shu H. and Van Vactor D. (2000) The guanine nucleotide exchange factor trio mediates axonal development in the Drosophila embryo. Neuron 26, 93-106.
15. Bear J. E., Loureiro J. J., Libova I., Fassler R., Wehland J. and Gertler F. B. (2000) Negative regulation of fibroblast motility by Ena/VASP proteins. Cell 101, 717-728.
16. Bear J. E., Svitkina T. M., Krause M., Schafer D. A., Loureiro J. J., Strasser G. A., Maly I. V., Chaga O. Y., Cooper J. A., Borisy G. G. and Gertler F. B. (2002) Antagonism between Ena/VASP proteins and actin filament capping regulates fibroblast motility. Cell 109, 509-521.
17. Billuart P., Winter C. G., Maresh A., Zhao X. and Luo L. (2001) Regulating axon branch stability: The role of p190 RhoGAP in repressing a retraction signaling pathway. Cell 107, 195-207.
18. Bito H., Furuyashiki T., Ishihara H., Shibasaki Y., Ohashi K., Mizuno K., Maekawa M., Ishizaki T. and Narumiya S. (2000) A critical role for a Rho-associated kinase, p160ROCK, in determining axon outgrowth in mammalian CNS neurons. Neuron 26, 431-441.
19. Blanchoin L., Pollard T. D. and Mullins R. D. (2000) Interactions of ADF/cofilin, Arp2/3 complex, capping protein and profilin in remodeling of branched actin filament networks. Curr. Biol. 10, 1273-1282.
20. Bray J. J., Fernyhough P., Bamburg J. R. and Bray D. (1992) Actin depolymerizing factor is a component of slow axonal transport. J. Neurochem. 58, 2081-2087.
21. Brown M. D., Cornejo B. J., Kuhn T. B. and Bamburg J. R. (2000) Cdc42 stimulates neurite outgrowth and formation of growth cone filopodia and lamellipodia. J. Neurobiol. 43, 352-364.
22. Carlier M. F., Laurent V., Santolini J., Melki R., Didry D., Xia G. X., Hong Y., Chua N. H. and Pantaloni D. (1997) Actin depolymerizing factor (ADF/cofilin) enhances the rate of filament turnover: Implication in actin-based motility. J. Cell Biol. 136, 1307-1322.
23. Chan A. Y., Bailly M., Zebda N., Segall J. E. and Condeelis J. S. (2000) Role of cofilin in epidermal growth factor-stimulated actin polymerization and lamellipod protrusion. J. Cell Biol. 148, 531-542.
24. Cheng A. K. and Robertson E. J. (1995) The murine LIM-kinase gene (limk) encodes a novel serine threonine kinase expressed pre-dominantly in trophoblast giant cells and the developing nervous system. Mech. Dev. 52, 187-197.
25. Cheng H.-L., Steinway M., Russell J. W. and Feldman E. L. (2000) GTPases and phosphatidylinositol-3 kinase are critical for insulin-like growth factor-I mediated Schwann cell motility. J. Biol. Chem. 275, 27197-27204.
26. Dan C., Nath N., Liberto M. and Minden A. (2002) PAK5, a new brain-specific kinase, promotes neurite outgrowth in N1E-115 cells. Mol. Cell. Biol. 22, 567-577.
27. Daniels R. H., Hall P. S. and Bokoch G. M. (1998) Membrane targeting of p21-activated kinase 1 (PAK1) induces neurite outgrowth from PC12 cells. EMBO J. 17, 754-764.
28. Dharmawardhane S., Sanders L. C., Martin S. S., Daniels R. H. and Bokoch G. M. (1997) Localization of p21-activated kinase 1 (PAK1) to pinocytic vesicles and cortical actin structures in stimulated cells. J. Cell Biol. 138, 1265-1278.
29. Donnai D. and Karmiloff-Smith A. (2000) Williams syndrome: From genotype through to the cognitive phenotype. Am. J. Med. Genet. 97, 164-171.
30. Du Y., Weed S. A., Xiong W. C., Marshall T. D. and Parsons J. T. (1998) Identification of a novel cortactin SH3 domain-binding protein and its localization to growth cones of cultured neurons. Mol. Cell. Biol. 18, 5838-5851.
31. Edwards D. C., Sanders L. C., Bokoch G. M. and Gill G. N. (1999) Activation of LIM-kinase by Pak1 couples Rac/Cdc42 GTPase signalling to actin cytoskeletal dynamics. Nat. Cell. Biol. 1, 253-259.
32. Egile C., Loisel T. P., Laurent V., Li R., Pantaloni D., Sansonetti P. J. and Carlier M. F. (1999) Activation of the CDC42 effector N-WASP by the Shigella flexneri IcsA protein promotes actin nucleation by Arp2/3 complex and bacterial actin-based motility. J. Cell Biol. 146, 1319-1332.
33. Ehler E., van Leeuwen F., Collard J. G. and Salinas P. C. (1997) Expression of Tiam-1 in the developing brain suggests a role for the Tiam-1-Rac signaling pathway in cell migration and neurite outgrowth. Mol. Cell. Neurosci. 9, 1-12.
34. Frost J. A., Xu S., Hutchison M. R., Marcus S. and Cobb M. H. (1996) Actions of Rho family small G proteins and p21-activated protein kinases on mitogen-activated protein kinase family members. Mol. Cell. Biol. 16, 3707-3713.
35. Fukuoka M., Miki H. and Takenawa T. (1997) Identification of N-WASP homologs in human and rat brain. Gene 196, 43-48.
36. Goh K. L., Cai L., Cepko C. L. and Gertler F. B. (2002) Ena/VASP proteins regulate cortical neuronal positioning. Curr. Biol. 12, 565-569.
37. Goldberg D. J., Foley M. S., Tang D. and Grabham P. W. (2000) Recruitment of the Arp2/3 complex and mena for the stimulation of actin polymerization in growth cones by nerve growth factor. J. Neurosci. Res. 60, 458-467.
38. Hirose M., Ishizaki T., Watanabe N., Uehata M., Kranenburg O., Moolenaar W. H., Matsumura F., Maekawa M., Bito H. and Narumiya S. (1998) Molecular dissection of the Rho-associated protein kinase (p160ROCK)-regulated neurite remodeling in neuroblastoma N1E-115 cells. J. Cell Biol. 141, 1625-1636.
39. Hu H., Marton T. F. and Goodman C. S. (2001) Plexin B mediates axon guidance in Drosophila by simultaneously inhibiting active Rac and enhancing RhoA signaling. Neuron 32, 39-51.
40. Jin Z. and Strittmatter S. M. (1997) Rac1 mediates collapsin-1-induced growth cone collapse. J. Neurosci. 17, 6256-6263.
41. Johnson D. I. and Pringle J. R. (1990) Molecular characterization of CDC42, a Saccharomyces cerevisiae gene involved in the development of cell polarity. J. Cell Biol. 111, 143-152.
42. Katoh H., Aoki J., Ichikawa A. and Negishi M. (1998) p160 RhoA-binding kinase ROKalpha induces neurite retraction. J. Biol. Chem. 273, 2489-2492.
43. Katoh H., Yasui H., Yamaguchi Y., Aoki J., Fujita H., Mori K. and Negishi M. (2000) Small GTPase RhoG is a key regulator for neurite outgrowth in PC12 cells. Mol. Cell. Biol. 20, 7378-7387.
44. Kimura K., Ito M., Amano M., Chihara K., Fukata Y., Nakafuku M., Yamamori B., Feng J., Nakano T., Okawa K., Iwamatsu A. and Kaibuchi K. (1996) Regulation of myosin phosphatase by Rho and Rho-associated kinase (Rho-kinase). Science 273, 245-248.
45. Knaus U. G., Morris S., Dong H. J., Chernoff J. and Bokoch G. M. (1995) Regulation of human leukocyte p21-activated kinases through G protein-coupled receptors. Science 269, 221-223.
46. Kozma R., Sarner S., Ahmed S. and Lim L. (1997) Rho family GTPases and neuronal growth cone remodelling: Relationship between increased complexity induced by Cdc42Hs, Rac1, and acetylcholine and collapse induced by RhoA and lysophosphatidic acid. Mol. Cell. Biol. 17, 1201-1211.
47. Kranenburg O., Poland M., van Horck F. P., Drechsel D., Hall A. and Moolenaar W. H. (1999) Activation of RhoA by lysophosphatidic acid and Galpha12/13 subunits in neuronal cells: Induction of neurite retraction. Mol. Biol. Cell 10, 1851-1857.
48. Krugmann S., Jordens I., Gevaert K., Driessens M., Vandekerckhove J. and Hall A. (2001) Cdc42 induces filopodia by promoting the formation of an IRSp53: Mena complex. Curr. Biol. 11, 1645-1655.
49. Kuhn T. B., Brown M. D., Wilcox C. L., Raper J. A. and Bamburg J. R. (1999) Myelin and collapsin-1 induce motor neuron growth cone collapse through different pathways: Inhibition of collapse by opposing mutants of rac1. J. Neurosci. 19, 1965-1975.
50. Kuhn T. B., Meberg P. J., Brown M. D., Bernstein B. W., Minamide L. S., Jensen J. R., Okada K., Soda E. A. and Bamburg J. R. (2000) Regulating actin dynamics in neuronal growth cones by ADF/cofilin and rho family GTPases. J. Neurobiol. 44, 126-144.
51. PAK and the JNK/SAPK MAP kinase cascade. Cell 87, 519-529.
52. Lamoureux P., Altun-Gultekin Z. F., Lin C., Wagner J. A. and Heidemann S. R. (1997) Rac is required for growth cone function but not neurite assembly. J. Cell Sci. 110, 635-641.
53. Laurent V., Loisel T. P., Harbeck B., Wehman A., Grobe L., Jockusch B. M., Wehland J., Gertler F. B. and Carlier M. F. (1999) Role of proteins of the Ena/VASP family in actin-based motility of Listeria monocytogenes. J. Cell Biol. 144, 1245-1258.
54. Lee T., Winter C., Marticke S. S., Lee A. and Luo L. (2000) Essential roles of Drosophila RhoA in the regulation of neuroblast proliferation and dendritic but not axonal morphogenesis. Neuron 25, 307-316.
55. Leeuwen F. N., Kain H. E., Kammen R. A., Michiels F., Kranenburg O. W. and Collard J. G. (1997) The guanine nucleotide exchange factor Tiam1 affects neuronal morphology; opposing roles for the small GTPases Rac and Rho. J. Cell Biol. 139, 797-807.
56. Lehmann M., Fournier A., Selles-Navarro I., Dergham P., Sebok A., Leclerc N., Tigyi G. and McKerracher L. (1999) Inactivation of Rho signaling pathway promotes CNS axon regeneration. J. Neurosci. 19, 7537-7547.
57. Lena J. Y., Bamburg J. R., Rabie A. and Faivre-Sarrailh C. (1991) Actindepolymerizing factor (ADF) in the cerebellum of the developing rat: a quantitative and immunocytochemical study. J. Neurosci. Res. 30, 18-27.
58. Lewis A. K. and Bridgman P. C. (1992) Nerve growth cone lamellipodia contain two populations of actin filaments that differ in organization and polarity. J. Cell Biol. 119, 1219-1243.
59. Li Z., Van Aelst L. and Cline H. T. (2000) Rho GTPases regulate distinct aspects of dendritic arbor growth in Xenopus central neurons in vivo. Nat. Neurosci. 3, 217-225.
60. Loisel T. P., Boujemaa R., Pantaloni D. and Carlier M. F. (1999) Reconstitution of actin-based motility of Listeria and Shigella using pure proteins. Nature 401, 613-616.
61. Lundquist E. A., Reddien P. W., Hartwieg E., Horvitz H. R. and Bargmann C. I. (2001) Three C. elegans Rac proteins and several alternative Rac regulators control axon guidance, cell migration and apoptotic cell phagocytosis. Development 128, 4475-4488.
62. Ma L., Rohatgi R. and Kirschner M. W. (1998) The Arp2/3 complex mediates actin polymerization induced by the small GTP-binding protein Cdc42. Proc. Natl. Acad. Sci. USA 95, 15362-15367.
63. Machesky L. M., Atkinson S. J., Ampe C., Vandekerckhove J. and Pollard T. D. (1994) Purification of a cortical complex containing two unconventional actins from Acanthamoeba by affinity chromatography on profilin-agarose. J. Cell Biol. 127, 107-115.
64. Machesky L. M., Reeves E., Wientjes F., Mattheyse F. J., Grogan A., Totty N. F., Burlingame A. L., Hsuan J. J. and Segal A. W. (1997) Mammalian actin-related protein 2/3 complex localizes to regions of lamellipodial protrusion and is composed of evolutionarily conserved proteins. Biochem. J. 328, 105-112.
65. Machesky L. M., Mullins R. D., Higgs H. N., Kaiser D. A., Blanchoin L., May R. C., Hall M. E. and Pollard T. D. (1999) Scar, a WASp-related protein, activates nucleation of actin filaments by the Arp2/3 complex. Proc. Natl. Acad. Sci. USA 96, 3739-3744.
66. Madaule P. and Axel R. (1985) A novel ras-related gene family. Cell 41, 31-40.
67. Maekawa S., Nishida E., Ohta Y. and Sakai H. (1984) Isolation of low molecular weight actin-binding proteins from porcine brain. J. Biochem. (Tokyo) 95, 377-385.
68. Maekawa M., Ishizaki T., Boku S., Watanabe N., Fujita A., Iwamatsu A., Obinata T., Ohashi K., Mizuno K. and Narumiya S. (1999) Signaling from Rho to the actin cytoskeleton through protein kinases ROCK and LIM-kinase. Science 285, 895-898.
69. Manser E., Leung T., Salihunddin H., Zhao Z. S. and Lim L. (1994) A brain serine/threonine protein kinase activated by Cdc42 and Rac1. Nature 367, 40-46.
70. Marcus S., Polverino A., Chang E., Robbins D., Cobb M. H. and Wigler M. H. (1995) Shk1, a homolog of the Saccharomyces cerevisiae Ste20 and mammalian p65PAK protein kinases, is a component of a Ras/Cdc42 signaling module in the fission yeast Schizosaccharomyces pombe. Proc. Natl. Acad. Sci. USA 92, 6180-6184.
71. Matsui T., Amano M., Yamamoto T., Chihara K., Nakafuku M., Ito M., Nakano T., Okawa K., Iwamatsu A. and Kaibuchi K. (1996) Rho-associated kinase, a novel serine/threonine kinase, as a putative target for small GTP binding protein Rho. EMBO J. 15, 2208-2216.
72. Meberg P. J. and Bamburg J. R. (2000) Increase in neurite outgrowth mediated by overexpression of actin depolymerizing factor. J. Neurosci. 20, 2459-2469.
73. Meberg P. J., Ono S., Minamide L. S., Takahashi M. and Bamburg J. R. (1998) Actin depolymerizing factor and cofilin phosphorylation dynamics: Response to signals that regulate neurite extension. Cell. Motil. Cytoskeleton 39, 172-190.
74. Miki H., Miura K. and Takenawa T. (1996) N-WASP, a novel actindepolymerizing protein, regulates the cortical cytoskeletal rearrangement in a PIP2-dependent manner downstream of tyrosine kinases. EMBO J. 15, 5326-5335.
75. Miki H., Sasaki T., Takai Y. and Takenawa T. (1998a) Induction of filopodium formation by a WASP-related actin-depolymerizing protein N-WASP. Nature 391, 93-96.
76. Miki H., Suetsugu S. and Takenawa T. (1998b) WAVE, a novel WASP-family protein involved in actin reorganization induced by Rac. EMBO J. 17, 6932-6941.
77. Miki H., Yamaguchi H., Suetsugu S. and Takenawa T. (2000) IRSp53 is an essential intermediate between Rac and WAVE in the regulation of membrane ruffling. Nature 408, 732-735.
78. Minamide L. S., Striegl A. M., Boyle J. A., Meberg P. J. and Bamburg J. R. (2000) Neurodegenerative stimuli induce persistent ADF/cofilin-actin rods that disrupt distal neurite function. Nat. Cell. Biol. 2, 628-636.
79. Mizuno K., Okano I., Ohashi K., Nunoue K., Kuma K., Miyata T. and Nakamura T. (1994) Identification of a human cDNA encoding a novel protein kinase with two repeats of the LIM/double zinc finger motif. Oncogene 9, 1605-1612.
80. Mullins R. D., Heuser J. A. and Pollard T. D. (1998) The interaction of Arp2/3 complex with actin: Nucleation, high affinity pointed end capping, and formation of branching networks of filaments. Proc. Natl. Acad. Sci. USA 95, 6181-6186.
81. Nakagawa H., Miki H., Ito M., Ohashi K., Takenawa T. and Miyamoto S. (2001) N-WASP, WAVE and Mena play different roles in the organization of actin cytoskeleton in lamellipodia. J. Cell Sci. 114, 1555-1565.
82. Nakayama A. Y., Harms M. B. and Luo L. (2000) Small GTPases Rac and Rho in the maintenance of dendritic spines and branches in hippocampal pyramidal neurons. J. Neurosci. 20, 5329-5338.
83. Narumiya S., Ishizaki T. and Watanabe N. (1997) Rho effectors and reorganization of actin cytoskeleton. FEBS Lett. 410, 68-72.
84. Newsome T. P., Schmidt S., Dietzl G., Keleman K., Asling B., Debant A. and Dickson B. J. (2000) Trio combines with dock to regulate Pak activity during photoreceptor axon pathfinding in Drosophila. Cell 101, 283-294.
85. Nikolic M., Chou M. M., Lu W., Mayer B. J. and Tsai L. H. (1998) The p35/Cdk5 kinase is a neuron-specific Rac effector that inhibits Pak1 activity. Nature 395, 194-198.
86. Niwa R., Nagata-Ohashi K., Takeichi M., Mizuno K. and Uemura T. (2002) Control of actin reorganization by Slingshot, a family of phosphatases that dephosphorylate ADF/cofilin. Cell 108, 233-246.
87. Nobes C. D. and Hall A. (1995) Rho, rac, and cdc42 GTPases regulate the assembly of multimolecular focal complexes associated with stress fibers, lamellipodia, and filopodia. Cell 81, 53-62.
88. Ohashi K., Nagata K., Maekawa M., Ishizaki T., Narumiya S. and Mizuno K. (2000) Rho-associated kinase ROCK activates LIM-kinase 1 by phosphorylation at threonine 508 within the activation loop. J. Biol. Chem. 275, 3577-3582.
89. Olenik C., Barth H., Just I., Aktories K. and Meyer D. K. (1997) Gene expression of the small GTP-binding proteins RhoA, RhoB, Rac1, and Cdc42 in adult rat brain. Brain Res. Mol. Brain Res. 52, 263-269.
90. Olofsson B. (1999) Rho guanine dissociation inhibitors: Pivotal molecules in cellular signalling. Cell Signal. 11, 545-554.
91. Prehoda K. E., Scott J. A., Mullins R. D. and Lim W. A. (2000) Integration of multiple signals through cooperative regulation of the N-WASP-Arp2/3 complex. Science 290, 801-806.
92. Rashid T., Banerjee M. and Nikolic M. (2001) Phosphorylation of Pak1 by the p35/Cdk5 kinase affects neuronal morphology. J. Biol. Chem. 276, 49043-49052.
93. Ridley A. J. and Hall A. (1992) The small GTP-binding protein rho regulates the assembly of focal adhesions and actin stress fibers in response to growth factors. Cell 70, 389-399.
94. Ridley A. J., Paterson H. F., Johnston C. L., Diekmann D. and Hall A. (1992) The small GTP-binding protein rac regulates growth factor-induced membrane ruffling. Cell 70, 401-410.
95. Rohatgi R., Ma L., Miki H., Lopez M., Kirchhausen T., Takenawa T. and Kirschner M. W. (1999) The interaction between N-WASP and the Arp2/3 complex links Cdc42-dependent signals to actin assembly. Cell 97, 221-231.
96. Rosenblatt J., Agnew B. J., Abe H., Bamburg J. R. and Mitchison T. J. (1997) Xenopus actin depolymerizing factor/cofilin (XAC) is responsible for the turnover of actin filaments in Listeria monocytogenes tails. J. Cell Biol. 136, 1323-1332.
97. Sarner S., Kozma R., Ahmed S. and Lim L. (2000) Phosphatidylinositol 3-kinase, Cdc42, and Rac1 act downstream of Ras in integrin-dependent neurite outgrowth in N1E-115 neuroblastoma cells. Mol. Cell. Biol. 20, 158-172.
98. Sasaki T. and Takai Y. (1998) The Rho small G protein family-Rho GDI system as a temporal and spatial determinant for cytoskeletal control. Biochem. Biophys. Res. Commun. 245, 641-645.
99. Scheffzek K., Ahmadian M. R. and Wittinghofer A. (1998) GTPase-activating proteins: Helping hands to complement an active site. Trends Biochem. Sci. 23, 257-262.
100. Sebok A., Nusser N., Debreceni B., Guo Z., Santos M. F., Szeberenyi J. and Tigyi G. (1999) Different roles for RhoA during neurite initiation, elongation, and regeneration in PC12 cells. J. Neurochem. 73, 949-960.
101. Sells M. A., Knaus U. G., Bagrodia S., Ambrose D. M., Bokoch G. M. and Chernoff J. (1997) Human p21-activated kinase (Pak1) regulates actin organization in mammalian cells. Curr. Biol. 7, 202-210.
102. Shamah S. M., Lin M. Z., Goldberg J. L., Estrach S., Sahin M., Hu L., Bazalakova M., Neve R. L., Corfas G., Debant A. and Greenberg M. E. (2001) EphA receptors regulate growth cone dynamics through the novel guanine nucleotide exchange factor ephexin. Cell 105, 233-244.
103. Smith G. A., Theriot J. A. and Portnoy D. A. (1996) The tandem repeat domain in the Listeria monocytogenes ActA protein controls the rate of actin-based motility, the percentage of moving bacteria, and the localization of vasodilator-stimulated phosphoprotein and profilin. J. Cell Biol. 135, 647-660.
104. Sumi T., Matsumoto K., Takai Y. and Nakamura T. (1999) Cofilin phosphorylation and actin cytoskeletal dynamics regulated by rhoand Cdc42-activated LIM-kinase 2. J. Cell Biol. 147, 1519-1532.
105. Svitkina T. M. and Borisy G. G. (1999) Arp2/3 complex and actin depolymerizing factor/cofilin in dendritic organization and tread-milling of actin filament array in lamellipodia. J. Cell Biol. 145, 1009-1026.
106. Threadgill R., Bobb K. and Ghosh A. (1997) Regulation of dendritic growth and remodeling by Rho, Rac, and Cdc42. Neuron 19, 625-634.
107. 2+ signaling and Rho. J. Neurochem. 66, 537-548.
108. Toshima J., Toshima J. Y., Amano T., Yang N., Narumiya S. and Mizuno K. (2001a) Cofilin phosphorylation by protein kinase testicular protein kinase 1 and its role in integrin-mediated actin reorganization and focal adhesion formation. Mol. Biol. Cell 12, 1131-1145.
109. Toshima J., Toshima J. Y., Suzuki M., Noda T. and Mizuno K. (2001b) Cell-type-specific expression of a TESK1 promoter-linked lacZ gene in transgenic mice. Biochem. Biophys. Res. Commun. 286, 566-573.
110. Toshima J., Toshima J. Y., Takeuchi K., Mori R. and Mizuno K. (2001c) Cofilin phosphorylation and actin reorganization activities of testicular protein kinase 2 and its predominant expression in testicular Sertoli cells. J. Biol. Chem. 276, 31449-31458.
111. Uruno T., Liu J., Zhang P., Fan Y., Egile C., Li R., Mueller S. C. and Zhan X. (2001) Activation of Arp2/3 complex-mediated actin polymerization by cortactin. Nat. Cell. Biol. 3, 259-266.
112. Vetter I. R. and Wittinghofer A. (2001) The guanine nucleotide-binding switch in three dimensions. Science 294, 1299-1304.
113. Wahl S., Barth H., Ciossek T., Aktories K. and Mueller B. K. (2000) Ephrin-A5 induces collapse of growth cones by activating Rho and Rho kinase. J. Cell Biol. 149, 263-270.
114. Wang J. Y., Wigston D. J., Rees H. D., Levey A. I. and Falls D. L. (2000) LIM kinase 1 accumulates in presynaptic terminals during synapse maturation. J. Comp. Neurol. 416, 319-334.
115. Weaver A. M., Karginov A. V., Kinley A. W., Weed S. A., Li Y., Parsons J. T. and Cooper J. A. (2001) Cortactin promotes and stabilizes Arp2/3-induced actin filament network formation. Curr. Biol. 11, 370-374.
116. Welch M. D., DePace A. H., Verma S., Iwamatsu A. and Mitchison T. J. (1997) The human Arp2/3 complex is composed of evolutionarily conserved subunits and is localized to cellular regions of dynamic actin filament assembly. J. Cell Biol. 138, 375-384.
117. Welnhofer E. A., Zhao L. and Cohan C. S. (1997) Actin dynamics and organization during growth cone morphogenesis in Helisoma neurons. Cell. Motil. Cytoskeleton 37, 54-71.
118. Weston C. A., Anova L., Rialas C., Prives J. M. and Weeks B. S. (2000) Laminin-1 activates Cdc42 in the mechanism of laminin-1-mediated neurite outgrowth. Exp. Cell Res. 260, 374-378.
119. Winter C. G., Wang B., Ballew A., Royou A., Karess R., Axelrod J. D. and Luo L. (2001) Drosophila Rho-associated kinase (Drok) links frizzled-mediated planar cell polarity signaling to the actin cytoskeleton. Cell 105, 81-91.
120. Wong K., Ren X. R., Huang Y. Z., Xie Y., Liu G., Saito H., Tang H., Wen L., Brady-Kalnay S. M., Mei L., Wu J. Y., Xiong W. C. and Rao Y. (2001) Signal transduction in neuronal migration: Roles of GTPase activating proteins and the small GTPase Cdc42 in the Slit-Robo pathway. Cell 107, 209-221.
121. Yamamoto K., Kondo J., Hishida T., Teranishi Y. and Takai Y. (1988) Purification and characterization of a GTP-binding protein with a molecular weight of 20,000 in bovine brain membranes. Identification as the rho gene product. J. Biol. Chem. 263, 9926-9932.
122. Yamashita T., Tucker K. L. and Barde Y. A. (1999) Neurotrophin binding to the p75 receptor modulates Rho activity and axonal outgrowth. Neuron 24, 585-593.
123. Yang N., Higuchi O., Ohashi K., Nagata K., Wada A., Kangawa K., Nishida E. and Mizuno K. (1998) Cofilin phosphorylation by LIM-kinase 1 and its role in Rac-mediated actin reorganization. Nature 393, 809-812.
124. Yarar D., To W., Abo A. and Welch M. D. (1999) The Wiskott-Aldrich syndrome protein directs actin-based motility by stimulating actin nucleation with the Arp2/3 complex. Curr. Biol. 9, 555-558.
125. Zalcman G., Dorseuil O., Garcia-Ranea J. A., Gacon G. and Camonis J. (1999) RhoGAPs and RhoGDIs, (His) stories of two families. Prog. Mol. Subcell. Biol. 22, 85-113.
126. Zallen J. A., Cohen Y., Hudson A. M., Cooley L., Wieschaus E. and Schejter E. D. (2002) SCAR is a primary regulator of Arp2/3-dependent morphological events in Drosophila. J. Cell Biol. 156, 689-701.
127. Zebda N., Bernard O., Bailly M., Welti S., Lawrence D. S. and Condeelis J. S. (2000) Phosphorylation of ADF/cofilin abolishes EGF-induced actin nucleation at the leading edge and subsequent lamellipod extension. J. Cell Biol. 151, 1119-1128.
128. Zheng Y. (2001) Db1 family guanine nucleotide exchange factors. Trends Biochem. Sci. 26, 724-732.
129. Zhou F. Q. and Cohan C. S. (2001) Growth cone collapse through coincident loss of actin bundles and leading edge actin without actin depolymerization. J. Cell Biol. 153, 1071-1084.