Extracellular regulators of axonal growth in the adult central nervous system

Philosophical Transactions of the Royal Society B: Biological Sciences

Volumn 361, Issue 1473, 2006, Pages 1593-1610

  Liu, Betty P ^a   Cafferty, William B J ^a   Budel, Stephane O ^a   Strittmatter, Stephen M ^a  

^a YALE UNIVERSITY SCHOOL OF MEDICINE   (United States)

Author keywords

Axon regeneration; Chondroitin sulphate proteoglycans; Myelin; Nogo; Nogo receptor; Spinal cord injury

Indexed keywords

ADULT; DEVELOPMENTAL STABILITY; GROWTH; INJURY; MAMMAL; NERVOUS SYSTEM; PROTEIN;

MAMMALIA;

EID: 33749028999     PISSN: 09628436     EISSN: None     Source Type: Journal    
DOI: 10.1098/rstb.2006.1891     Document Type: Review

References (215)

1. Ahmed, Z., Dent, R. G., Suggate, E. L., Barrett, L. B., Seabright, R. J., Berry, M. & Logan, A. 2005 Disinhibition of neurotrophin-induced dorsal root ganglion cell neurite outgrowth on CNS myelin by siRNA-mediated knockdown of NgR, p75NTR and Rho-A. Mol. Cell. Neurosci. 28, 509-523. (doi:10.1016/j.mcn. 2004.11.002)
2. Asher, R. A. et al. 2000 Neurocan is upregulated in injured brain and in cytokine-treated astrocytes. J. Neurosci. 20, 2427-2438.
3. Bandtlow, C. E. & Zimmermann, D. R. 2000 Proteoglycans in the developing brain: new conceptual insights for old proteins. Physiol. Rev. 80, 1267-1290.
4. Bandtlow, C., Zachleder, T. & Schwab, M. E. 1990 Oligodendrocytes arrest neurite growth by contact inhibition. J. Neurosci. 10, 3837-3848.
5. Bareyre, F. M., Haudenschild, B. & Schwab, M. E. 2002 Long-lasting sprouting and gene expression changes induced by the monoclonal antibody IN-1 in the adult spinal cord. J. Neurosci. 22, 7097-7110.
6. Bareyre, F. M., Kerschensteiner, M., Raineteau, O., Mettenleiter, T. C., Weinmann, O. & Schwab, M. E. 2004 The injured spinal cord spontaneously forms a new intraspinal circuit in adult rats. Nat. Neurosci. 7, 269-277. (doi:10.1038/nn1195)
7. Barton, W. A., Liu, B. P., Tzvetkova, D., Jeffrey, P. D., Fournier, A. E., Sah, D., Cate, R., Strittmatter, S. M. & Nikolov, D. B. 2003 Structure and axon outgrowth inhibitor binding of the Nogo-66 receptor and related proteins. EMBO J. 22, 3291-3302. (doi:10.1093/emboj/cdg325)
8. Bartsch, U. et al. 1995 Lack of evidence that myelin-associated glycoprotein is a major inhibitor of axonal regeneration in the CNS. Neuron 15, 1375-1381. (doi:10.1016/0896-6273(95)90015-2)
9. Benson, M. D., Romero, M. I., Lush, M. E., Lu, Q. R., Henkemeyer, M. & Parada, L. F. 2005 Ephrin-B3 is a myelin-based inhibitor of neurite outgrowth. Proc. Natl Acad. Sci. USA 102, 10 694-10 699. (doi:10.1073/pnas. 0504021102)
10. Berry, M., Maxwell, W. L., Logan, A., Mathewson, A., McConnell, P., Ashhurst, D. E. & Thomas, G. H. 1983 Deposition of scar tissue in the central nervous system. Acta Neurochir. 32(Suppl. (Wien)), 31-53.
11. Biffiger, K., Bartsch, S., Montag, D., Aguzzi, A., Schachner, M. & Bartsch, U. 2000 Severe hypomyelination of the murine CNS in the absence of myelin-associated glycoprotein and fyn tyrosine kinase. J. Neurosci. 20, 7430-7437.
12. Bjartmar, C., Yin, X. & Trapp, B. D. 1999 Axonal pathology in myelin disorders. J. Neurocytol. 28, 383-395. (doi:10.1023/A:1007010205037)
13. Bomze, H. M., Bulsara, K. R., Iskandar, B. J., Caroni, P. & Skene, J. H. 2001 Spinal axon regeneration evoked by replacing two growth cone proteins in adult neurons. Nat. Neurosci. 4, 38-43. (doi:10.1038/82881)
14. Borisoff, J. F., Chan, C. C., Hiebert, G. W., Oschipok, L., Robertson, G. S., Zamboni, R., Steeves, J. D. & Tetzlaff, W. 2003 Suppression of Rho-kinase activity promotes axonal growth on inhibitory CNS substrates. Mol. Cell. Neurosci. 22, 405-416. (doi:10.1016/S1044-7431(02)00032-5)
15. Bradbury, E. J., Moon, L. D., Popat, R. J., King, V. R., Bennett, G. S., Patel, P. N., Fawcett, J. W. & McMahon, S. B. 2002 Chondroitinase ABC promotes functional recovery after spinal cord injury. Nature 416, 636-640. (doi:10.1038/416636a)
16. Brosamle, C., Huber, A. B., Fiedler, M., Skerra, A. & Schwab, M. E. 2000 Regeneration of lesioned corticospinal tract fibers in the adult rat induced by a recombinant, humanized IN-1 antibody fragment. J. Neurosci. 20, 8061-8068.
17. Buffo, A., Zagrebelsky, M., Huber, A. B., Skerra, A., Schwab, M. E., Strata, P. & Rossi, F. 2000 Application of neutralizing antibodies against NI-35/250 myelin-associated neurite growth inhibitory proteins to the adult rat cerebellum induces sprouting of uninjured Purkinje cell axons. J. Neurosci. 20, 2275-2286.
18. Bundesen, L. Q., Scheel, T. A., Bregman, B. S. & Kromer, L. F. 2003 Ephrin-B2 and EphB2 regulation of astrocyte-meningeal fibroblast interactions in response to spinal cord lesions in adult rats. J. Neurosci. 23, 7789-7800.
19. Burg, M. A., Tillet, E., Timpl, R. & Stallcup, W. B. 1996 Binding of the NG2 proteoglycan to type VI collagen and other extracellular matrix molecules. J. Biol. Chem. 271, 26 110-26 116. (doi:10.1074/jbc.271.42.26110)
20. Caroni, P. & Schwab, M. E. 1988 Two membrane protein fractions from rat central myelin with inhibitory properties for neurite growth and fibroblast spreading. J. Cell Biol. 106, 1281-1288. (doi:10.1083/jcb.106.4.1281)
21. Carulli, D., Laabs, T., Geller, H. M. & Fawcett, J. W. 2005 Chondroitin sulfate proteoglycans in neural development and regeneration. Curr. Opin. Neurobiol. 15, 116-120. (doi:10.1016/j.conb.2005.03.018)
22. Chau, C. H., Shum, D. K., Li, H., Pei, J., Lui, Y. Y., Wirthlin, L., Chan, Y. S. & Xu, X. M. 2004 Chondroitinase ABC enhances axonal regrowth through Schwann cell-seeded guidance channels after spinal cord injury. FASEB J. 18, 194-196.
23. Chen, M. S., Huber, A. B., van der Haar, M. E., Frank, M., Schnell, L., Spillmann, A. A., Christ, F. & Schwab, M. E. 2000 Nogo-A is a myelin-associated neurite outgrowth inhibitor and an antigen for monoclonal antibody IN-1. Nature 403, 434-439. (doi:10.1038/35000219)
24. Chen, Z. J., Ughrin, Y. & Levine, J. M. 2002 Inhibition of axon growth by oligodendrocyte precursor cells. Mol. Cell. Neurosci. 20, 125-139. (doi:10.1006/mcne.2002.1102)
25. David, S. & Aguayo, A. J. 1981 Axonal elongation into peripheral nervous system 'bridges' after central nervous system injury in adult rats. Science 214, 931-933.
26. Davies, S. J., Fitch, M. T., Memberg, S. P., Hall, A. K., Raisman, G. & Silver, J. 1997 Regeneration of adult axons in white matter tracts of the central nervous system. Nature 390, 680-683.
27. Davies, S. J., Goucher, D. R., Doller, C. & Silver, J. 1999 Robust regeneration of adult sensory axons in degenerating white matter of the adult rat spinal cord. J. Neurosci. 19, 5810-5822.
28. Davies, J. E., Tang, X., Denning, J. W., Archibald, S. J. & Davies, S. J. 2004 Decorin suppresses neurocan, brevican, phosphacan and NG2 expression and promotes axon growth across adult rat spinal cord injuries. Eur. J. Neurosci. 19, 1226-1242. (doi:10.1111/j.1460-9568.2004.03184.x)
29. de Castro Jr, R., Tajrishi, R., Claros, J. & Stallcup, W. B. 2005 Differential responses of spinal axons to transection: influence of the NG2 proteoglycan. Exp. Neurol. 192, 299-309. (doi:10.1016/j.expneurol.2004.11.027)
30. De Winter, F., Holtmaat, A. J. & Verhaagen, J. 2002 Neuropilin and class 3 semaphorins in nervous system regeneration. Adv. Exp. Med. Biol. 515, 115-139.
31. Deguchi, K. et al. 2005 Expression of neurocan after transient middle cerebral artery occlusion in adult rat brain. Brain Res. 1037, 194-199. (doi:10.1016/j.brainres.2004.12.016)
32. Dergham, P., Ellezam, B., Essagian, C., Avedissian, H., Lubell, W. D. & McKerracher, L. 2002 Rho signaling pathway targeted to promote spinal cord repair. J. Neurosci. 22, 6570-6577.
33. Domeniconi, M. et al. 2002 Myelin-associated glycoprotein interacts with the Nogo66 receptor to inhibit neurite outgrowth. Neuron 35, 283-290. (doi:10.1016/S0896-6273(02)00770-5)
34. Domeniconi, M., Zampieri, N., Spencer, T., Hilaire, M., Mellado, W., Chao, M. V. & Filbin, M. T. 2005 MAG induces regulated intramembrane proteolysis of the p75 neurotrophin receptor to inhibit neurite outgrowth. Neuron 46, 849-855. (doi:10.1016/j.neuron.2005.05.029)
35. Domowicz, M., Li, H., Hennig, A., Henry, J., Vertel, B. M. & Schwartz, N. B. 1995 The biochemically and immunologically distinct CSPG of notochord is a product of the aggrecan gene. Dev. Biol. 171, 655-664. (doi:10.1006/dbio.1995.1312)
36. Dou, C. L. & Levine, J. M. 1994 Inhibition of neurite growth by the NG2 chondroitin sulfate proteoglycan. J. Neurosci. 14, 7616-7628.
37. Dou, C. L. & Levine, J. M. 1997 Identification of a neuronal cell surface receptor for a growth inhibitory chondroitin sulfate proteoglycan (NG2). J. Neurochem. 68, 1021-1030.
38. Dours-Zimmermann, M. T. & Zimmermann, D. R. 1994 A novel glycosaminoglycan attachment domain identified in two alternative splice variants of human versican. J. Biol. Chem. 269, 32 992-32 998.
39. Emerick, A. J., Neafsey, E. J., Schwab, M. E. & Kartje, G. L. 2003 Functional reorganization of the motor cortex in adult rats after cortical lesion and treatment with monoclonal antibody IN-1. J. Neurosci. 23, 4826-4830.
40. Faissner, A., Clement, A., Lochter, A., Streit, A., Mandl, C. & Schachner, M. 1994 Isolation of a neural chondroitin sulfate proteoglycan with neurite outgrowth promoting properties. J. Cell Biol. 126, 783-799. (doi:10.1083/jcb.126.3.783)
41. Fidler, P. S. et al. 1999 Comparing astrocytic cell lines that are inhibitory or permissive for axon growth: the major axon-inhibitory proteoglycan is NG2. J. Neurosci. 19, 8778-8788.
42. Fiedler, M., Horn, C., Bandtlow, C., Schwab, M. E. & Skerra, A. 2002 An engineered IN-1 F(ab) fragment with improved affinity for the Nogo-A axonal growth inhibitor permits immunochemical detection and shows enhanced neutralizing activity. Protein Eng. 15, 931-941. (doi:10.1093/protein/15.11.931)
43. Fischer, D., He, Z. & Benowitz, L. I. 2004 Counteracting the Nogo receptor enhances optic nerve regeneration if retinal ganglion cells are in an active growth state. J. Neurosci. 24, 1646-1651. (doi:10.1523/JNEUROSCI.5119-03. 2004)
44. Fitch, M. T. & Silver, J. 1997 Glial cell extracellular matrix: boundaries for axon growth in development and regeneration. Cell Tissue Res. 290, 379-384. (doi:10.1007/s004410050944)
45. Fitch, M. T., Doller, C., Combs, C. K., Landreth, G. E. & Silver, J. 1999 Cellular and molecular mechanisms of glial scarring and progressive cavitation: in vivo and in vitro analysis of inflammation-induced secondary injury after CNS trauma. J. Neurosci. 19, 8182-8198.
46. Fouad, K., Klusman, I. & Schwab, M. E. 2004 Regenerating corticospinal fibers in the Marmoset (Callitrix jacchus) after spinal cord lesion and treatment with the anti-Nogo-A antibody IN-1. Eur. J. Neurosci. 20, 2479-2482. (doi:10.1111/j.1460-9568.2004.03716.x)
47. Fouad, K., Schnell, L., Bunge, M. B., Schwab, M. E., Liebscher, T. & Pearse, D. D. 2005 Combining Schwann cell bridges and olfactory-ensheathing glia grafts with chondroitinase promotes locomotor recovery after complete transection of the spinal cord. J. Neurosci. 25, 1169-1178. (doi:10.1523/ JNEUROSCI.3562-04.2005)
48. Fournier, A. E., GrandPre, T. & Strittmatter, S. M. 2001 Identification of a receptor mediating Nogo-66 inhibition of axonal regeneration. Nature 409, 341-346. (doi:10.1038/35053072)
49. Fournier, A. E., Gould, G. C., Liu, B. P. & Strittmatter, S. M. 2002 Truncated soluble Nogo receptor binds Nogo-66 and blocks inhibition of axon growth by myelin. J. Neurosci. 22, 8876-8883.
50. Fournier, A. E., Takizawa, B. T. & Strittmatter, S. M. 2003 Rho kinase inhibition enhances axonal regeneration in the injured CNS. J. Neurosci. 23, 1416-1423.
51. Fox, K. & Wong, R. O. 2005 A comparison of experience-dependent plasticity in the visual and somatosensory systems. Neuron 48, 465-477. (doi:10.1016/j.neuron.2005.10.013)
52. Friedlander, D. R., Milev, P., Karthikeyan, L., Margolis, R. K., Margolis, R. U. & Grumet, M. 1994 The neuronal chondroitin sulfate proteoglycan neurocan binds to the neural cell adhesion molecules Ng-CAM/L1/NILE and N-CAM, and inhibits neuronal adhesion and neurite outgrowth. J. Cell Biol. 125, 669-680. (doi:10.1083/jcb.125.3.669)
53. Fruttiger, M., Montag, D., Schachner, M. & Martini, R. 1995 Crucial role for the myelin-associated glycoprotein in the maintenance of axon-myelin integrity. Eur. J. Neurosci. 7, 511-515. (doi:10.1111/j.1460-9568.1995.tb00347. x)
54. Fujisawa, H. 2004 Discovery of semaphorin receptors, neuropilin and plexin, and their functions in neural development. J. Neurobiol. 59, 24-33. (doi:10.1002/neu.10337)
55. Fujita, N., Kemper, A., Dupree, J., Nakayasu, H., Bartsch, U., Schachner, M., Maeda, N., Suzuki, K. & Popko, B. 1998 The cytoplasmic domain of the large myelin-associated glycoprotein isoform is needed for proper CNS but not peripheral nervous system myelination. J. Neurosci. 18, 1970-1978.
56. Geng, J., Shin, M. E., Gilbert, P. M., Collins, R. N. & Burd, C. G. 2005 Saccharomyces cerevisiae Rab-GDI displacement factor ortholog Yip3p forms distinct complexes with the Ypt1 Rab GTPase and the reticulon Rtn1p. Eukaryot Cell 4, 1166-1174. (doi:10.1128/EC.4.7.1166-1174.2005)
57. Goldberg, J. L. & Barres, B. A. 1998 Neuronal regeneration: extending axons from bench to brain. Curr. Biol. 8, R310-R312. (doi:10.1016/S0960- 9822(98)70195-2)
58. Goldshmit, Y., Galea, M. P., Wise, G., Bartlett, P. F. & Turnley, A. M. 2004 Axonal regeneration and lack of astrocytic gliosis in EphA4-deficient mice. J. Neurosci. 24, 10 064-10 073. (doi:10.1523/JNEUROSCI.2981-04.2004)
59. GrandPre, T., Li, S. & Strittmatter, S. M. 2002 Nogo-66 receptor antagonist peptide promotes axonal regeneration. Nature 417, 547-551.
60. GrandPre, T., Nakamura, F., Vartanian, T. & Strittmatter, S. M. 2000 Identification of the Nogo inhibitor of axon regeneration as a Reticulon protein. Nature 403, 439-444. (doi:10.1038/35000226)
61. Grimpe, B. & Silver, J. 2004 A novel DNA enzyme reduces glycosaminoglycan chains in the glial scar and allows microtransplanted dorsal root ganglia axons to regenerate beyond lesions in the spinal cord. J. Neurosci. 24, 1393-1397. (doi:10.1523/JNEUROSCI.4986-03.2004)
62. Grumet, M., Flaccus, A. & Margolis, R. U. 1993 Functional characterization of chondroitin sulfate proteoglycans of brain: interactions with neurons and neural cell adhesion molecules. J. Cell Biol. 120, 815-824. (doi:10.1083/jcb.120.3.815)
63. Grumet, M., Milev, P., Sakurai, T., Karthikeyan, L., Bourdon, M., Margolis, R. K. & Margolis, R. U. 1994 Interactions with tenascin and differential effects on cell adhesion of neurocan and phosphacan, two major chondroitin sulfate proteoglycans of nervous tissue. J. Biol. Chem. 269, 12 142-12 146.
64. Habib, A. A., Marton, L. S., Allwardt, B., Gulcher, J. R., Mikol, D. D., Hognason, T., Chattopadhyay, N. & Stefansson, K. 1998 Expression of the oligodendrocyte-myelin glycoprotein by neurons in the mouse central nervous system. J. Neurochem. 70, 1704-1711.
65. Hasegawa, Y., Fujitani, M., Hata, K., Tohyama, M., Yamagishi, S. & Yamashita, T. 2004 Promotion of axon regeneration by myelin-associated glycoprotein and Nogo through divergent signals downstream of Gi/G. J. Neurosci. 24, 6826-6832. (doi:10.1523/JNEUROSCI.1856-04.2004)
66. He, Z. & Tessier-Lavigne, M. 1997 Neuropilin is a receptor for the axonal chemorepellent Semaphorin III. Cell 90, 739-751. (doi:10.1016/S0092- 8674(00)80534-6)
67. He, X. L., Bazan, J. F., McDermott, G., Park, J. B., Wang, K., Tessier-Lavigne, M., He, Z. & Garcia, K. C. 2003 Structure of the Nogo receptor ectodomain: a recognition module implicated in myelin inhibition. Neuron 38, 177-185. (doi:10.1016/S0896-6273(03)00232-0)
68. Hensch, T. K. 2005 Critical period mechanisms in developing visual cortex. Curr. Top. Dev. Biol. 69, 215-237.
69. Hensch, T. K. & Fagiolini, M. 2005 Excitatory-inhibitory balance and critical period plasticity in developing visual cortex. Prog. Brain Res. 147, 115-124.
70. Holtmaat, A. J., Trachtenberg, J. T., Wilbrecht, L., Shepherd, G. M., Zhang, X., Knott, G. W. & Svoboda, K. 2005 Transient and persistent dendritic spines in the neocortex in vivo. Neuron 45, 279-291. (doi:10.1016/j.neuron.2005.01.003)
71. Hu, F. & Strittmatter, S. M. 2004 Regulating axon growth within the postnatal central nervous system. Semin. Perinatol. 28, 371-378. (doi:10.1053/j.semperi.2004.10.001)
72. Hu, F., Liu, B. P., Budel, S., Liao, J., Chin, J., Fournier, A. & Strittmatter, S. M. 2005 Nogo-A interacts with the Nogo-66 receptor through multiple sites to create an isoform-selective subnanomolar agonist. J. Neurosci. 25, 5298-5304. (doi:10.1523/JNEUROSCI.5235-04.2005)
73. Huang, J. K. et al. 2005 Glial membranes at the Node of Ranvier prevent neurite outgrowth. Science 310, 1813-1817.
74. Huizinga, E. G., Tsuji, S., Romijn, R. A., Schiphorst, M. E., de Groot, P. G., Sixma, J. J. & Gros, P. 2002 Structures of glycoprotein Ibalpha and its complex with von Willebrand factor Al domain. Science 297, 1176-1179. (doi:10.1126/science.107355)
75. Hunt, D., Mason, M. R., Campbell, G., Coffin, R. & Anderson, P. N. 2002 Nogo receptor mRNA expression in intact and regenerating CNS neurons. Mol. Cell. Neurosci. 20, 537-552. (doi:10.1006/mcne.2002.1153)
76. Ji, B., Li, M., Budel, S., Pepinsky, R. B., Walus, L., Engber, T. M., Strittmatter, S. M. & Relton, J. K. 2005 Effect of combined treatment with methylprednisolone and soluble Nogo-66 receptor after rat spinal cord injury. Eur. J. Neurosci. 22, 587-594. (doi:10.1111/j.1460-9568.2005.04241.x)
77. Jin, Z. & Strittmatter, S. M. 1997 Rac1 mediates collapsin-1-induced growth cone collapse. J. Neurosci. 17, 6256-6263.
78. Jones, L. L., Yamaguchi, Y., Stallcup, W. B. & Tuszynski, M. H. 2002 NG2 is a major chondroitin sulfate proteoglycan produced after spinal cord injury and is expressed by macrophages and oligodendrocyte progenitors. J. Neurosci. 22, 2792-2803.
79. Jones, L. L., Sajed, D. & Tuszynski, M. H. 2003 Axonal regeneration through regions of chondroitin sulfate proteoglycan deposition after spinal cord injury: a balance of permissiveness and inhibition. J. Neurosci. 23, 9276-9288.
80. Kalil, K. & Reh, T. 1982 A light and electron microscopic study of regrowing pyramidal tract fibers. J. Comp. Neurol. 211, 265-275. (doi:10.1002/cne.902110305)
81. Kim, J. E., Bonilla, I. E., Qiu, D. & Strittmatter, S. M. 2003a Nogo-C is sufficient to delay nerve regeneration. Mol. Cell. Neurosci. 23, 451-459. (doi:10.1016/S1044-7431(03)00076-9)
82. Kim, J. E., Li, S., GrandPre, T., Qiu, D. & Strittmatter, S. M. 2003b Axon regeneration in young adult mice lacking Nogo-A/B. Neuron 38, 187-199. (doi:10.1016/S0896-6273(03)00147-8)
83. Kim, J. E., Liu, B. P., Park, J. H. & Strittmatter, S. M. 2004 Nogo-66 receptor prevents raphespinal and rubrospinal axon regeneration and limits functional recovery from spinal cord injury. Neuron 44, 439-451. (doi:10.1016/j.neuron.2004.10.015)
84. Klein, R. 2004 Eph/ephrin signaling in morphogenesis, neural development and plasticity. Curr. Opin. Cell Biol. 16, 580-589. (doi:10.1016/j.ceb.2004.07. 002)
85. Kolodkin, A. L., Levengood, D. V., Rowe, E. G., Tai, Y. X., Giger, R. J. & Ginty, D. D. 1997 Neuropilin is a semaphorin III receptor. Cell 90, 753-762. (doi:10.1016/S0092-8674(00)80535-8)
86. Koprivica, V. et al. 2005 EGFR activation mediates inhibition of axon regeneration by myelin and chondroitin sulfate proteoglycans. Science 310, 106-110. (doi:10.1126/science.1115462)
87. Krueger Jr, R. C., Hennig, A. K. & Schwartz, N. B. 1992 Two immunologically and developmentally distinct chondroitin sulfate proteolglycans in embryonic chick brain. J. Biol. Chem. 267, 12 149-12 161.
88. Krusius, T., Gehlsen, K. R. & Ruoslahti, E. 1987 A fibroblast chondroitin sulfate proteoglycan core protein contains lectin-like and growth factor-like sequences. J. Biol. Chem. 262, 13 120-13 125.
89. Kullander, K., Mather, N. K., Diella, F., Dottori, M., Boyd, A. W. & Klein, R. 2001 Kinase-dependent and kinase-independent functions of EphA4 receptors in major axon tract formation in vivo. Neuron 29, 73-84. (doi:10.1016/S0896-6273(01)00181-7)
90. Kursula, P., Lehto, V. P. & Heape, A. M. 2000 S100beta inhibits the phosphorylation of the L-MAG cytoplasmic domain by PKA. Brain Res. Mol. Brain Res. 76, 407-410. (doi:10.1016/S0169-328X(00)00018-8)
91. Lassmann, H., Bartsch, U., Montag, D. & Schachner, M. 1997 Dying-back oligodendrogliopathy: a late sequel of myelin-associated glycoprotein deficiency. Glia 19, 104-110. (doi:10.1002/(SICI)1098-1136(199702)19:2〈104: :AID-GLIA2〉3.0.CO;2-0)
92. Lauren, J., Airaksinen, M. S., Saarma, M. & Timmusk, T. 2003 Two novel mammalian Nogo receptor homologs differentially expressed in the central and peripheral nervous systems. Mol. Cell. Neurosci. 24, 581-594. (doi:10.1016/S1044-7431(03)00199-4)
93. Lee, J. K., Kim, J. E., Sivula, M. & Strittmatter, S. M. 2004 Nogo receptor antagonism promotes stroke recovery by enhancing axonal plasticity. J. Neurosci. 24, 6209-6217. (doi:10.1523/JNEUROSCI.1643-04.2004)
94. Lehmann, M., Fournier, A., Selles-Navarro, I., Dergham, P., Sebok, A., Leclerc, N., Tigyi, G. & McKerracher, L. 1999 Inactivation of Rho signaling pathway promotes CNS axon regeneration. J. Neurosci. 19, 7537-7547.
95. Leung, K. M., Margolis, R. U. & Chan, S. O. 2004 Expression of phosphacan and neurocan during early development of mouse retinofugal pathway. Brain Res. Dev. Brain Res. 152, 1-10. (doi:10.1016/j.devbrainres.2004.05.010)
96. Levine, J. M. 1994 Increased expression of the NG2 chondroitin-sulfate proteoglycan after brain injury. J. Neurosci. 14, 4716-4730.
97. Levine, J. M. & Card, J. P. 1987 Light and electron microscopic localization of a cell surface antigen (NG2) in the rat cerebellum: association with smooth protoplasmic astrocytes. J. Neurosci. 7, 2711-2720.
98. Li, S. & Strittmatter, S. M. 2003 Delayed systemic Nogo-66 receptor antagonist promotes recovery from spinal cord injury. J. Neurosci. 23, 4219-4227.
99. Li, C., Tropak, M. B., Gerlai, R., Clapoff, S., Abramow-Newerly, W., Trapp, B., Peterson, A. & Roder, J. 1994 Myelination in the absence of myelin-associated glycoprotein. Nature 369, 747-750. (doi:10.1038/369747a0)
100. Li, H., Domowicz, M., Hennig, A. & Schwartz, N. B. 1996 S103L reactive chondroitin sulfate proteoglycan (aggrecan) mRNA expressed in developing chick brain and cartilage is encoded by a single gene. Brain Res. Mol. Brain Res. 36, 309-321. (doi:10.1016/0169-328X(95)00269-X)
101. Li, S. et al. 2004a Blockade of Nogo-66, myelin-associated glycoprotein, and oligodendrocyte myelin glycoprotein by soluble Nogo-66 receptor promotes axonal sprouting and recovery after spinal injury. J. Neurosci. 24, 10 511-10 520. (doi:10.1523/JNEUROSCI.2828-04.2004)
102. Li, W. et al. 2004b A neutralizing anti-Nogo66 receptor monoclonal antibody reverses inhibition of neurite outgrowth by central nervous system myelin. J. Biol. Chem. 279, 43 780-43 788. (doi:10.1074/jbc.M401803200)
103. Li, S., Kim, J. E., Budel, S., Hampton, T. G. & Strittmatter, S. M. 2005 Transgenic inhibition of Nogo-66 receptor function allows axonal sprouting and improved locomotion after spinal injury. Mol. Cell. Neurosci. 29, 26-39.
104. Liu, B. P., Fournier, A., GrandPre, T. & Strittmatter, S. M. 2002 Myelin-associated glycoprotein as a functional ligand for the Nogo-66 receptor. [comment]. Science 297, 1190-1193. (doi:10.1126/science.1073031)
105. MacDermid, V. E., McPhail, L. T., Tsang, B., Rosenthal, A., Davies, A. & Ramer, M. S. 2004 A soluble Nogo receptor differentially affects plasticity of spinally projecting axons. Eur. J. Neurosci. 20, 2567-2579. (doi:10.1111/j.1460-9568.2004.03715.x)
106. Maeda, N., Hamanaka, H., Shintani, T., Nishiwaki, T. & Noda, M. 1994 Multiple receptor-like protein tyrosine phosphatases in the form of chondroitin sulfate proteoglycan. FEBS Lett. 354, 67-70. (doi:10.1016/0014-5793(94)01093-5)
107. Manitt, C., Colicos, M. A., Thompson, K. M., Rousselle, E., Peterson, A. C. & Kennedy, T. E. 2001 Widespread expression of netrin-1 by neurons and oligodendrocytes in the adult mammalian spinal cord. J. Neurosci. 21, 3911-3922.
108. Manitt, C., Thompson, K. M. & Kennedy, T. E. 2004 Developmental shift in expression of netrin receptors in the rat spinal cord: predominance of UNC-5 homologues in adulthood. J. Neurosci. Res 77, 690-700. (doi:10.1002/jnr.20199)
109. Margolis, R. U. & Margolis, R. K. 1997 Chondroitin sulfate proteoglycans as mediators of axon growth and pathfinding. Cell Tissue Res. 290, 343-348. (doi:10.1007/s004410050939)
110. Matsui, F., Watanabe, E. & Oohira, A. 1994 Immunological identification of two proteoglycan fragments derived from neurocan, a brain-specific chondroitin sulfate proteoglycan. Neurochem. Int. 25, 425-431. (doi:10.1016/0197-0186(94)90018-3)
111. Matsunaga, E., Tauszig-Delamasure, S., Monnier, P. P., Mueller, B. K., Strittmatter, S. M., Mehlen, P. & Chedotal, A. 2004 RGM and its receptor neogenin regulate neuronal survival. Nat. Cell Biol. 6, 749-755.
112. Maurel, P., Rauch, U., Flad, M., Margolis, R. K. & Margolis, R. U. 1994 Phosphacan, a chondroitin sulfate proteoglycan of brain that interacts with neurons and neural cell-adhesion molecules, is an extracellular variant of a receptor-type protein tyrosine phosphatase. Proc. Natl. Acad. Sci. USA 91, 2512-2516. (doi:10.1073/pnas.91.7.2512)
113. McGee, A. W. & Strittmatter, S. M. 2003 The Nogo-66 receptor: focusing myelin inhibition of axon regeneration. Trends Neurosci. 26, 193-198. (doi:10.1016/S0166-2236(03)00062-6)
114. McGee, A. W., Yang, Y., Fischer, Q. S., Daw, N. W. & Strittmatter, S. M. 2005 Experience-driven plasticity of visual cortex limited by myelin and Nogo receptor. Science 309, 2222-2226. (doi:10.1126/science.1114362)
115. McKeon, R. J., Schreiber, R. C., Rudge, J. S. & Silver, J. 1991 Reduction of neurite outgrowth in a model of glial scarring following CNS injury is correlated with the expression of inhibitory molecules on reactive astrocytes. J. Neurosci. 11, 3398-3411.
116. McKeon, R. J., Hoke, A. & Silver, J. 1995 Injury-induced proteoglycans inhibit the potential for laminin-mediated axon growth on astrocytic scars. Exp. Neurol. 136, 32-43. (doi:10.1006/exnr.1995.1081)
117. McKerracher, L., David, S., Jackson, D. L., Kottis, V., Dunn, R. J. & Braun, P. E. 1994 Identification of myelin-associated glycoprotein as a major myelin-derived inhibitor of neunte growth. Neuron 13, 805-811. (doi:10.1016/0896-6273(94)90247-X)
118. Merkler, D., Metz, G. A., Raineteau, O., Dietz, V., Schwab, M. E. & Fouad, K. 2001 Locomotor recovery in spinal cord-injured rats treated with an antibody neutralizing the myelin-associated neurite growth inhibitor Nogo-A. J. Neurosci. 21, 3665-3673.
119. Mi, S. et al. 2004 LINGO-1 is a component of the Nogo-66 receptor/p75 signaling complex. Nat. Neurosci. 7, 221-228. (doi:10.1038/nn1188)
120. Milev, P., Friedlander, D. R., Sakurai, T., Karthikeyan, L., Flad, M., Margolis, R. K., Grumet, M. & Margolis, R. U. 1994 Interactions of the chondroitin sulfate proteoglycan phosphacan, the extracellular domain of a receptor-type protein tyrosine phosphatase, with neurons, glia, and neural cell adhesion molecules. J. Cell Biol. 127, 1703-1715. (doi:10.1083/jcb.127.6.1703)
121. Milev, P., Maurel, P., Haring, M., Margolis, R. K. & Margolis, R. U. 1996 TAG-1/axonin-1 is a high-affinity ligand of neurocan, phosphacan/protein- tyrosine phosphatase-zeta/beta, and N-CAM. J. Biol. Chem. 271, 15 716-15 723. (doi:10.1074/jbc.271.26.15716)
122. Milev, P., Maurel, P., Chiba, A., Mevissen, M., Popp, S., Yamaguchi, Y., Margolis, R. K. & Margolis, R. U. 1998 Differential regulation of expression of hyaluronan-binding proteoglycans in developing brain: aggrecan, versican, neurocan, and brevican. Biochem. Biophys. Res. Commun. 247, 207-212. (doi:10.1006/bbrc.1998.8759)
123. Mingorance, A., Fontana, X., Soriano, E. & Del Rio, J. A. 2005 Overexpression of myelin-associated glycoprotein after axotomy of the perforant pathway. Mol. Cell. Neurosci. 29, 471-483. (doi:10.1016/j.mcn.2005.03.016)
124. Monnier, P. P. et al. 2002 RGM is a repulsive guidance molecule for retinal axons. Nature 419, 392-395. (doi:10.1038/nature01041)
125. Monnier, P. P., Sierra, A., Schwab, J. M., Henke-Fahle, S. & Mueller, B. K. 2003 The Rho/ROCK pathway mediates neurite growth-inhibitory activity associated with the chondroitin sulfate proteoglycans of the CNS glial scar. Mol. Cell. Neurosci. 22, 319-330. (doi:10.1016/S1044-7431(02)00035-0)
126. Montag, D. et al. 1994 Mice deficient for the myelin-associated glycoprotein show subtle abnormalities in myelin. Neuron 13, 229-246. (doi:10.1016/0896-6273(94)90472-3)
127. Moon, L. D., Asher, R. A., Rhodes, K. E. & Fawcett, J. W. 2001 Regeneration of CNS axons back to their target following treatment of adult rat brain with chondroitinase ABC. Nat. Neurosci. 4, 465-466.
128. Moreau-Fauvarque, C. et al. 2003 The transmembrane semaphorin Sema4D/CD100, an inhibitor of axonal growth, is expressed on oligodendrocytes and upregulated after CNS lesion. J. Neurosci. 23, 9229-9239.
129. Mukhopadhyay, G., Doherty, P., Walsh, F. S., Crocker, P. R. & Filbin, M. T. 1994 A novel role for myelin-associated glycoprotein as an inhibitor of axonal regeneration. Neuron 13, 757-767. (doi:10.1016/0896-6273(94)90042-6)
130. Nakamura, F., Kalb, R. G. & Strittmatter, S. M. 2000 Molecular basis of semaphorin-mediated axon guidance. J. Neurobiol. 44, 219-229. (doi:10.1002/1097-4695(200008)44:2〈219::AID-NEU11〉3.0.CO;2-W)
131. Niederost, B. P., Zimmermann, D. R., Schwab, M. E. & Bandtlow, C. E. 1999 Bovine CNS myelin contains neurite growth-inhibitory activity associated with chondroitin sulfate proteoglycans. J. Neurosci. 19, 8979-8989.
132. Nishiyama, A., Dahlin, K. J., Prince, J. T., Johnstone, S. R. & Stallcup, W. B. 1991 The primary structure of NG2, a novel membrane-spanning proteoglycan. J. Cell Biol. 114, 359-371. (doi:10.1083/jcb.114.2.359)
133. Oakley, R. A. & Tosney, K. W. 1991 Peanut agglutinin and chondroitin-6-sulfate are molecular markers for tissues that act as barriers to axon advance in the avian embryo. Dev. Biol. 147, 187-206. (doi:10.1016/S0012- 1606(05)80017-X)
134. Oertle, T. et al. 2003 Nogo-A inhibits neurite outgrowth and cell spreading with three discrete regions. J. Neurosci. 23, 5393-5406.
135. Oohira, A., Matsui, F., Watanabe, E., Kushima, Y. & Maeda, N. 1994 Developmentally regulated expression of a brain specific species of chondroitin sulfate proteoglycan, neurocan, identified with a monoclonal antibody IG2 in the rat cerebrum. Neuroscience 60, 145-157. (doi:10.1016/0306-4522(94)90210-0)
136. Oster, S. F., Bodeker, M. O., He, F. & Sretavan, D. W. 2003 Invariant Sema5A inhibition serves an ensheathing function during optic nerve development. Development 130, 775-784. (doi:10.1242/dev.00299)
137. Papadopoulos, C. M., Tsai, S. Y., Alsbiei, T., O'Brien, T. E., Schwab, M. E. & Kartje, G. L. 2002 Functional recovery and neuroanatomical plasticity following middle cerebral artery occlusion and IN-1 antibody treatment in the adult rat. Ann. Neurol. 51, 433-441. (doi:10.1002/ana.10144)
138. Park, J. B., Yiu, G., Kaneko, S., Wang, J., Chang, J., He, X. L., Garcia, K. C. & He, Z. 2005 ATNF receptor family member, TROY, is a coreceptor with Nogo receptor in mediating the inhibitory activity of myelin inhibitors. Neuron 45, 345-351. (doi:10.1016/j.neuron.2004.12.040)
139. Paulsson, M., Morgelin, M., Wiedemann, H., Beardmore-Gray, M., Dunham, D., Hardingham, T., Heinegard, D., Timpl, R. & Engel, J. 1987 Extended and globular protein domains in cartilage proteoglycans. Biochem. J. 245, 763-772.
140. Perissinotto, D. et al. 2000 Avian neural crest cell migration is diversely regulated by the two major hyaluronan-binding proteoglycans PG-M/versican and aggrecan. Development 127, 2823-2842.
141. Pignot, V. et al. 2003 Characterization of two novel proteins, NgRH1 and NgRH2, structurally and biochemically homologous to the Nogo-66 receptor. J. Neurochem. 85, 717-728.
142. Pizzorusso, T., Medini, P., Berardi, N., Chierzi, S., Fawcett, J. W. & Maffei, L. 2002 Reactivation of ocular dominance plasticity in the adult visual cortex. Science 298, 1248-1251. (doi:10.1126/science.1072699)
143. Popp, S., Maurel, P., Andersen, J. S. & Margolis, R. U. 2004 Developmental changes of aggrecan, versican and neurocan in the retina and optic nerve. Exp. Eye Res. 79, 351-356. (doi:10.1016/j.exer.2004.05.016)
144. Pot, C. et al. 2002 Nogo-A expressed in Schwann cells impairs axonal regeneration after peripheral nerve injury. J. Cell Biol. 159, 29-35. (doi:10.1083/jcb.200206068)
145. Prinjha, R., Moore, S. E., Vinson, M., Blake, S., Morrow, R., Christie, G., Michalovich, D., Simmons, D. L. & Walsh, F. S. 2000 Inhibitor of neurite outgrowth in humans. Nature 403, 383-384. (doi:10.1038/35000287)
146. Quarles, R. H., Everly, J. L. & Brady, R. O. 1972 Demonstration of a glycoprotein which is associated with a purified myelin fraction from rat brain. Biochem. Biophys. Res. Commun. 47, 491-497. (doi:10.1016/0006-291X(72)90741-3)
147. Quarles, R. H., Everly, J. L. & Brady, R. O. 1973 Myelin-associated glycoprotein: a developmental change. Brain Res. 58, 506-509. (doi:10.1016/0006-8993(73)90022-X)
148. Raff, M. C., Miller, R. H. & Noble, M. 1983 A glial progenitor cell that develops in vitro into an astrocyte or an oligodendrocyte depending on culture medium. Nature 303, 390-396. (doi:10.1038/303390a0)
149. Raff, M. C., Durand, B. & Gao, F. B. 1998 Cell number control and timing in animal development: the oligodendrocyte cell lineage. Int. J. Dev. Biol. 42, 263-267.
150. Raineteau, O., Z'Graggen, W. J., Thallmair, M. & Schwab, M. E. 1999
151. Raineteau, O., Fouad, K., Noth, P., Thallmair, Mr & Schwab, M. E. 2001 Functional switch between motor tracts in the presence of the mAb IN-1 in the adult rat. Proc. Natl Acad. Sci. USA 98, 6929-6934. (doi:10.1073/pnas. 111165498)
152. Rajagopalan, S., Deitinghoff, L., Davis, D., Conrad, S., Skutella, T., Chedotal, A., Mueller, B. K. & Strittmatter, S. M. 2004 Neogenin mediates the action of repulsive guidance molecule. Nat. Cell Biol. 6, 756-762.
153. Rauch, U., Karthikeyan, L., Maurel, P., Margolis, R. U. & Margolis, R. K. 1992 Cloning and primary structure of neurocan, a developmentally regulated, aggregating chondroitin sulfate proteoglycan of brain. J. Biol. Chem. 267, 19 536-19 547.
154. Rhodes, K. E., Moon, L. D. & Fawcett, J. W. 2003 Inhibiting cell proliferation during formation of the glial scar: effects on axon regeneration in the CNS. Neuroscience 120, 41-56. (doi:10.1016/S0306-4522(03)00285-9)
155. Richardson, P. M., McGuinness, U. M. & Aguayo, A. J. 1982 Peripheral nerve autografts to the rat spinal cord: studies with axonal tracing methods. Brain Res. 237, 147-162. (doi:10.1016/0006-8993(82)90563-7)
156. Sango, K., Oohira, A., Ajiki, K., Tokashiki, A., Horie, M. & Kawano, H. 2003 Phosphacan and neurocan are repulsive substrata for adhesion and neurite extension of adult rat dorsal root ganglion neurons in vitro. Exp. Neurol. 182, 1-11. (doi:10.1016/S0014-4886(03)00090-6)
157. Savio, T. & Schwab, M. E. 1989 Rat CNS white matter, but not gray matter, is nonpermissive for neuronal cell adhesion and fiber outgrowth. J. Neurosci. 9, 1126-1133.
158. Schachner, M. & Bartsch, U. 2000 Multiple functions of the myelin-associated glycoprotein MAG (siglec-4a) in formation and maintenance of myelin. Glia 29, 154-165. (doi:10.1002/(SICI)1098-1136(20000115)29:2〈154:: AID-GLIA9〉3.0.CO;2-3)
159. Schafer, M., Fruttiger, M., Montag, D., Schachner, M. & Martini, R. 1996 Disruption of the gene for the myelin-associated glycoprotein improves axonal regrowth along myelin in C57BL/Wlds mice. Neuron 16, 1107-1113. (doi:10.1016/S0896-6273(00)80137-3)
160. Schimmele, B. & Pluckthun, A. 2005 Identification of a functional epitope of the Nogo receptor by a combinatorial approach using ribosome display. J. Mol. Biol. 352, 229-241. (doi:10.1016/j.jmb.2005.06.073)
161. Schmalfeldt, M., Dours-Zimmermann, M. T., Winterhalter, K. H. & Zimmermann, D. R. 1998 Versican V2 is a major extracellular matrix component of the mature bovine brain. J. Biol. Chem. 273, 15 758-15 764. (doi:10.1074/jbc. 273.25.15758)
162. Schnell, L. & Schwab, M. E. 1990 Axonal regeneration in the rat spinal cord produced by an antibody against myelin-associated neurite growth inhibitors. Nature 343, 269-272. (doi:10.1038/343269a0)
163. Schreyer, D. J. & Jones, E. G. 1983 Growing corticospinal axons by-pass lesions of neonatal rat spinal cord. Neuroscience 9, 31-40. (doi:10.1016/0306-4522(83)90044-1)
164. Schreyer, D. J. & Skene, J. H. 1993 Injury-associated induction of GAP-43 expression displays axon branch specificity in rat dorsal root ganglion neurons. J. Neurobiol. 24, 959-970. (doi:10.1002/neu.480240709)
165. Schwab, M. E. & Thoenen, H. 1985 Dissociated neurons regenerate into sciatic but not optic nerve explants in culture irrespective of neurotrophic factors. J. Neurosci. 5, 2415-2423.
166. Schwab, M. E. & Caroni, P. 1988 Oligodendrocytes and CNS myelin are nonpermissive substrates for neurite growth and fibroblast spreading in vitro. J. Neurosci. 8, 2381-2393.
167. Schweigreiter, R. et al. 2004 Versican V2 and the central inhibitory domain of Nogo-A inhibit neurite growth via p75NTR/NgR-independent pathways that converge at RhoA. Mol. Cell. Neurosci. 27, 163-174. (doi:10.1016/j.mcn.2004.06. 004)
168. Seymour, A. B. et al. 2005 Delayed treatment with monoclonal antibody IN-1 1 week after stroke results in recovery of function and corticorubral plasticity in adult rats. J. Cereb. Blood Flow Metab. 25, 1366-1375. (doi:10.1038/sj.jcbfm.9600134)
169. Shao, Z. et al. 2005 TAJ/TROY, an orphan TNF receptor family member, binds Nogo-66 receptor 1 and regulates axonal regeneration. Neuron 45, 353-359. (doi:10.1016/j.neuron.2004.12.050)
170. Silver, J. & Miller, J. H. 2004 Regeneration beyond the glial scar. Nat. Rev. Neurosci. 5, 146-156. (doi:10.1038/nrn1326)
171. Simonen, M. et al. 2003 Systemic deletion of the myelin-associated outgrowth inhibitor Nogo-A improves regenerative and plastic responses after spinal cord injury. Neuron 38, 201-211. (doi:10.1016/S0896-6273(03)00226-5)
172. Sivasankaran, R., Pei, J., Wang, K. C., Zhang, Y. P., Shields, C. B., Xu, X. M. & He, Z. 2004 PKC mediates inhibitory effects of myelin and chondroitin sulfate proteoglycans on axonal regeneration. Nat. Neurosci. 7, 261-268. (doi:10.1038/nn1193)
173. Skene, J. H. 1989 Axonal growth-associated proteins. Annu. Rev. Neurosci. 12, 127-156. (doi:10.1146/annurev.ne.12.030189.001015)
174. Smith-Thomas, L. C., Fok-Seang, J., Stevens, J., Du, J. S., Muir, E., Faissner, A., Geller, H. M., Rogers, J. H. & Fawcett, J. W. 1994 An inhibitor of neurite outgrowth produced by astrocytes. J. Cell Sci. 107, 1687-1695.
175. Smith-Thomas, L. C., Stevens, J., Fok-Seang, J., Faissner, A., Rogers, J. H. & Fawcett, J. W. 1995 Increased axon regeneration in astrocytes grown in the presence of proteoglycan synthesis inhibitors. J. Cell Sci. 108, 1307-1315.
176. Snow, D. M., Lemmon, V., Carrino, D. A., Caplan, A. I. & Silver, J. 1990 Sulfated proteoglycans in astroglial barriers inhibit neurite outgrowth in vitro. Exp. Neurol. 109, 111-130. (doi:10.1016/S0014-4886(05)80013-5)
177. Spillmann, A. A., Bandtlow, C. E., Lottspeich, F., Keller, F. & Schwab, M. E. 1998 Identification and characterization of a bovine neurite growth inhibitor (bNI-220). J. Biol. Chem. 273, 19 283-19 293. (doi:10.1074/jbc.273.30.19283)
178. Stallcup, W. B. & Beasley, L. 1987 Bipotential glial precursor cells of the optic nerve express the NG2 proteoglycan. J. Neurosci. 7, 2737-2744.
179. Stallcup, W. B. & Dahlin-Huppe, K. 2001 Chondroitin sulfate and cytoplasmic domain-dependent membrane targeting of the NG2 proteoglycan promotes retraction fiber formation and cell polarization. J. Cell Sci. 114, 2315-2325.
180. Steiner, P., Kulangara, K., Sarria, J. C., Glauser, L., Regazzi, R. & Hirling, H. 2004 Reticulon 1-C/neuroendocrine-specific protein-C interacts with SNARE proteins. J. Neurochem. 89, 569-580. (doi:10.1111/j.1471-4159.2004.02345. x)
181. Streit, A., Nolte, C., Rasony, T. & Schachner, M. 1993 Interaction of astrochondrin with extracellular matrix components and its involvement in astrocyte process formation and cerebellar granule cell migration. J. Cell Biol. 120, 799-814. (doi:10.1083/jcb.120.3.799)
182. Taha, S. A. & Stryker, M. P. 2005 Molecular substrates of plasticity in the developing visual cortex. Prog. Brain Res. 147, 103-114.
183. Takahashi, T., Fournier, A., Nakamura, F., Wang, L. H., Murakami, Y., Kalb, R. G., Fujisawa, H. & Strittmatter, S. M. 1999 Plexin-neuropilin-1 complexes form functional semaphorin-3A receptors. Cell 99, 59-69. (doi:10.1016/S0092-8674(00)80062-8)
184. Tamagnone, L. et al. 1999 Plexins are a large family of receptors for transmembrane, secreted, and GPI-anchored semaphorins in vertebrates. Cell 99, 71-80. (doi:10.1016/S0092-8674(00)80063-X)
185. Tang, S., Woodhall, R. W., Shen, Y. J., deBellard, M. E., Saffell, J. L., Doherty, P., Walsh, F. S. & Filbin, M. T. 1997 Soluble myelin-associated glycoprotein (MAG) found in vivo inhibits axonal regeneration. Mol. Cell. Neurosci. 9, 333-346. (doi:10.1006/mcne.1997.0633)
186. Tang, X., Davies, J. E. & Davies, S. J. 2003 Changes in distribution, cell associations, and protein expression levels of NG2, neurocan, phosphacan, brevican, versican V2, and tenascin-C during acute to chronic maturation of spinal cord scar tissue. J. Neurosci. Res 71, 427-444. (doi:10.1002/jnr.10523)
187. Teng, F. Y. & Tang, B. L. 2005 Why do Nogo/Nogo-66 receptor gene knockouts result in inferior regeneration compared to treatment with neutralizing agents? J. Neurochem. 94, 865-874. (doi:10.1111/j.1471-4159.2005. 03238.x)
188. Tessier-Lavigne, M. & Goodman, C. S. 1996 The molecular biology of axon guidance. Science 274, 1123-1133. (doi:10.1126/science.274.5290.1123)
189. Tropak, M. B., Johnson, P. W., Dunn, R. J. & Roder, J. C. 1988 Differential splicing of MAG transcripts during CNS and PNS development. Brain Res. 464, 143-155.
190. Ughrin, Y. M., Chen, Z. J. & Levine, J. M. 2003 Multiple regions of the NG2 proteoglycan inhibit neurite growth and induce growth cone collapse. J. Neurosci. 23, 175-186.
191. Umemori, H., Sato, S., Yagi, T., Aizawa, S. & Yamamoto, T. 1994 Initial events of myelination involve Fyn tyrosine kinase signalling. Nature 367, 572-576. (doi:10.1038/367572a0)
192. Venkatesh, K., Chivatakarn, O., Lee, H., Joshi, P. S., Kantor, D. B., Newman, B. A., Mage, R., Rader, C. & Giger, R. J. 2005 The Nogo-66 receptor homolog NgR2 is a sialic acid-dependent receptor selective for myelin-associated glycoprotein. J. Neurosci. 25, 808-822. (doi:10.1523/JNEUROSCI.4464-04.2005)
193. Vinson, M., Strijbos, P. J., Rowles, A., Facci, L., Moore, S. E., Simmons, D. L. & Walsh, F. S. 2001 Myelin-associated glycoprotein interacts with ganglioside GT1b. A mechanism for neurite outgrowth inhibition. J. Biol. Chem. 276, 20 280-20 285. (doi:10.1074/jbc.M100345200)
194. Vourc'h, P. & Andres, C. 2004 Oligodendrocyte myelin glycoprotein (OMgp): evolution, structure and function. Brain Res. Brain Res. Rev. 45, 115-124. (doi:10.1016/j.brainresrev.2004.01.003)
195. Wang, K. C., Kim, J. A., Sivasankaran, R., Segal, R. & He, Z. 2002a P75 interacts with the Nogo receptor as a co-receptor for Nogo, MAG and OMgp. Nature 420, 74-78. (doi:10.1038/nature01176)
196. Wang, K. C., Koprivica, V., Kim, J. A., Sivasankaran, R., Guo, Y., Neve, R. L. & He, Z. 2002b Oligodendrocyte-myelin glycoprotein is a Nogo receptor ligand that inhibits neurite outgrowth. Nature 417, 941-944. (doi:10.1038/nature00867)
197. Wang, X., Chun, S. J., Treloar, H., Vartanian, T., Greer, C. A. & Strittmatter, S. M. 2002c Localization of Nogo-A and Nogo-66 receptor proteins at sites of axon-myelin and synaptic contact. J. Neurosci. 22, 5505-5515.
198. Weiss, M. D., Hammer, J. & Quarles, R. H. 2000 Oligodendrocytes in aging mice lacking myelin-associated glycoprotein are dystrophic but not apoptotic. J. Neurosci. Res 62, 772-780. (doi:10.1002/1097-4547(20001215)62: 6〈772::AID-JNR3〉3.0.CO;2-X)
199. Wenk, C. A., Thallmair, M., Kartje, G. L. & Schwab, M. E. 1999 Increased corticofugal plasticity after unilateral cortical lesions combined with neutralization of the IN-1 antigen in adult rats. J. Comp. Neurol. 410, 143-157. (doi:10.1002/(SICI)1096-9861(19990719)410:1〈143::AID-CNE12〉3. 0.CO;2-)
200. Winberg, M. L., Noordermeer, J. N., Tamagnone, L., Comoglio, P. M., Spriggs, M. K., Tessier-Lavigne, M. & Goodman, C. S. 1998 Plexin A is a neuronal semaphorin receptor that controls axon guidance. Cell 95, 903-916. (doi:10.1016/S0092-8674(00)81715-8)
201. Winton, M. J., Dubreuil, C. I., Lasko, D., Leclerc, N. & McKerracher, L. 2002 Characterization of new cell permeable C3-like proteins that inactivate Rho and stimulate neurite outgrowth on inhibitory substrates. J. Biol. Chem. 277, 32 820-32 829. (doi:10.1074/jbc.M201195200)
202. Wong, S. T., Henley, J. R., Kanning, K. C., Huang, K. H., Bothwell, M. & Poo, M. M. 2002 A p75(NTR) and Nogo receptor complex mediates repulsive signaling by myelin-associated glycoprotein. Nat. Neurosci. 5, 1302-1308. (doi:10.1038/nn975)
203. Woolf, C. J. 2003 No Nogo: now where to go? Neuron 38, 153-156. (doi:10.1016/S0896-6273(03)00233-2)
204. Yaginuma, H., Homma, S., Kunzi, R. & Oppenheim, R. W. 1991 Pathfinding by growth cones of commissural interneurons in the chick embryo spinal cord: a light and electron microscopic study. J. Comp. Neurol. 304, 78-102. (doi:10.1002/cne.903040107)
205. Yamada, H., Watanabe, K., Shimonaka, M. & Yamaguchi, Y. 1994 Molecular cloning of brevican, a novel brain proteoglycan of the aggrecan/versican family. J. Biol. Chem. 269, 10 119-10 126.
206. Yamada, H., Fredette, B., Shitara, K., Hagihara, K., Miura, R., Ranscht, B., Stallcup, W. B. & Yamaguchi, Y. 1997 The brain chondroitin sulfate proteoglycan brevican associates with astrocytes ensheathing cerebellar glomeruli and inhibits neurite outgrowth from granule neurons. J. Neurosci. 17, 7784-7795.
207. Yamagata, M. & Sanes, J. R. 2005 Versican in the developing brain: lamina-specific expression in interneuronal subsets and role in presynaptic maturation. J. Neurosci. 25, 8457-8467. (doi:10.1523/JNEUROSCI.1976-05.2005)
208. Yamashita, T. & Tohyama, M. 2003 The p75 receptor acts as a displacement factor that releases Rho from Rho-GDI. Nat. Neurosci. 6, 461-467.
209. Yang, Y., Fischer, Q. S., Zhang, Y., Baumgartel, K., Mansuy, I. M. & Daw, N. W. 2005 Reversible blockade of experience-dependent plasticity by calcineurin in mouse visual cortex. Nat. Neurosci. 8, 791-796. (doi:10.1038/nn1464)
210. Yick, L. W., Wu, W., So, K. F., Yip, H. K. & Shum, D. K. 2000 Chondroitinase ABC promotes axonal regeneration of Clarke's neurons after spinal cord injury. Neuroreport 11, 1063-1067.
211. Yick, L. W., Cheung, P. T., So, K. F. & Wu, W. 2003 Axonal regeneration of Clarke's neurons beyond the spinal cord injury scar after treatment with chondroitinase ABC. Exp. Neurol. 182, 160-168. (doi:10.1016/S0014-4886(02)00052-3)
212. Yu, T. W. & Bargmann, C. I. 2001 Dynamic regulation of axon guidance. Nat. Neurosci. 4(Suppl.), 1169-1176. (doi:10.1038/nn748)
213. Zheng, B., Ho, C., Li, S., Keirstead, H., Steward, O. & Tessier-Lavigne, M. 2003 Lack of enhanced spinal regeneration in Nogo-deficient mice. Neuron 38, 213-224. (doi:10.1016/S0896-6273(03)00225-3)
214. Zheng, B., Atwal, J., Ho, C., Case, L., He, X. L., Garcia, K. C., Steward, O. & Tessier-Lavigne, M. 2005 Genetic deletion of the Nogo receptor does not reduce neurite inhibition in vitro or promote corticospinal tract regeneration in vivo. Proc. Natl Acad. Sci. USA 102, 1205-1210. (doi:10.1073/pnas.0409026102)
215. Zhou, X. H. et al. 2001 Neurocan is dispensable for brain development. Mol. Cell. Biol. 21, 5970-5978. (doi:10. 1128/MCB.21.17.5970-5978.2001)