Lame ducks or fierce creatures? - The role of oligodendrocytes in multiple sclerosis

Journal of Molecular Neuroscience

Volumn 35, Issue 1, 2008, Pages 91-100

  Zeis, T ^a   Schaeren Wiemers, N ^a  

^a UNIVERSITY HOSPITAL BASEL   (Switzerland)

Author keywords

Inflammation; Innate immunity; Multiple sclerosis; Normal appearing white matter; Oligodendrocyte; Oligodendrocyte pathology

Indexed keywords

HEAT SHOCK PROTEIN 32; HEAT SHOCK PROTEIN 70; NERVE GROWTH FACTOR; SOMATOMEDIN; STAT6 PROTEIN; TRANSFORMING GROWTH FACTOR BETA; ANTIBODY; SIGNAL PEPTIDE;

ASTROCYTE; B LYMPHOCYTE; CD4+ T LYMPHOCYTE; CD8+ T LYMPHOCYTE; CELL ACTIVATION; CELL DAMAGE; CELL DEATH; CELL SURVIVAL; HUMAN; IMMUNE RESPONSE; INFLAMMATION; MACROPHAGE; MICROGLIA; MULTIPLE SCLEROSIS; NONHUMAN; OLIGODENDROGLIA; OXIDATIVE STRESS; PATHOGENESIS; REVIEW; ANIMAL; CYTOLOGY; DEMYELINATING DISEASE; IMMUNOLOGY; METABOLISM; MYELIN SHEATH; PATHOLOGY; PATHOPHYSIOLOGY; PHYSIOLOGY; T LYMPHOCYTE;

ANAS;

ANIMALS; ANTIBODIES; ASTROCYTES; B-LYMPHOCYTES; DEMYELINATING DISEASES; HUMANS; INTERCELLULAR SIGNALING PEPTIDES AND PROTEINS; MACROPHAGES; MICROGLIA; MULTIPLE SCLEROSIS; MYELIN SHEATH; OLIGODENDROGLIA; T-LYMPHOCYTES;

EID: 45949112049     PISSN: 08958696     EISSN: None     Source Type: Journal    
DOI: 10.1007/s12031-008-9042-1     Document Type: Review

References (108)

1. F. Aboul-Enein H. Rauschka B. Kornek C. Stadelmann A. Stefferl W. Bruck 2003 Preferential loss of myelin-associated glycoprotein reflects hypoxia-like white matter damage in stroke and inflammatory brain diseases Journal of Neuropathology and Experimental Neurology 62 25 33
2. H. Babbe A. Roers A. Waisman H. Lassmann N. Goebels R. Hohlfeld 2000 Clonal expansions of CD8(+) T cells dominate the T cell infiltrate in active multiple sclerosis lesions as shown by micromanipulation and single cell polymerase chain reaction Journal of Experimental Medicine 192 393 404
3. R. Balabanov K. Strand R. Goswami E. McMahon W. Begolka S. D. Miller 2007 Interferon-gamma-oligodendrocyte interactions in the regulation of experimental autoimmune encephalomyelitis Journal of Neuroscience 27 2013 2024
4. M. H. Barnett J. W. Prineas 2004 Relapsing and remitting multiple sclerosis: Pathology of the newly forming lesion Annals of Neurology 55 458 468
5. B. Becher P. A. Barker T. Owens J. P. Antel 1998 CD95-CD95L: Can the brain learn from the immune system? Trends in Neurosciences 21 114 117
6. B. Becher A. Prat J. P. Antel 2000 Brain-immune connection: Immuno-regulatory properties of CNS-resident cells Glia 29 293 304
7. T. Berger S. Weerth K. Kojima C. Linington H. Wekerle H. Lassmann 1997 Experimental autoimmune encephalomyelitis: The antigen specificity of T lymphocytes determines the topography of lesions in the central and peripheral nervous system Laboratory Investigation 76 355 364
8. M. Bergeron J. M. Gidday A. Y. Yu G. L. Semenza D. M. Ferriero F. R. Sharp 2000 Role of hypoxia-inducible factor-1 in hypoxia-induced ischemic tolerance in neonatal rat brain Annals of Neurology 48 285 296
9. K. Bergsteindottir A. Brennan K. R. Jessen R. Mirsky 1992 In the presence of dexamethasone, gamma interferon induces rat oligodendrocytes to express major histocompatibility complex class II molecules Proceedings of the National Academy of Sciences of the United States of America 89 9054 9058
10. M. Bernaudin Y. Tang M. Reilly E. Petit F. R. Sharp 2002 Brain genomic response following hypoxia and re-oxygenation in the neonatal rat. Identification of genes that might contribute to hypoxia-induced ischemic tolerance Journal of Biological Chemistry 277 39728 39738
11. B. Brune J. Zhou 2007 Hypoxia-inducible factor-1alpha under the control of nitric oxide Methods in Enzymology 435 463 478
12. M. Buntinx E. Gielen P. Van Hummelen J. Raus M. Ameloot P. Steels 2004 Cytokine-induced cell death in human oligodendroglial cell lines. II: Alterations in gene expression induced by interferon-gamma and tumor necrosis factor-alpha Journal of Neuroscience Research 76 846 861
13. B. Cannella S. Gaupp K. M. Omari C. S. Raine 2007 Multiple sclerosis: Death receptor expression and oligodendrocyte apoptosis in established lesions Journal of Neuroimmunology 188 128 137
14. B. Cannella D. Pitt E. Capello C. S. Raine 2000 Insulin-like growth factor-1 fails to enhance central nervous system myelin repair during autoimmune demyelination American Journal of Pathology 157 933 943
15. B. Cannella C. S. Raine 2004 Multiple sclerosis: Cytokine receptors on oligodendrocytes predict innate regulation Annals of Neurology 55 46 57
16. P. Casaccia-Bonnefil C. Gu G. Khursigara M. V. Chao 1999 p75 neurotrophin receptor as a modulator of survival and death decisions Microscopy Research and Technique 45 217 224
17. T. Chitnis N. Najafian C. Benou A. D. Salama M. J. Grusby M. H. Sayegh 2001 Effect of targeted disruption of STAT4 and STAT6 on the induction of experimental autoimmune encephalomyelitis Journal of Clinical Investigation 108 739 747
18. E. S. Christians L. J. Yan I. J. Benjamin 2002 Heat shock factor 1 and heat shock proteins: Critical partners in protection against acute cell injury Critical Care Medicine 30 S43 S50
19. Compston, A., McDonald, I., Noseworthy, J., Lassmann, H., Miller, D., Smith, K., et al. (2006). McAlpine's multiple sclerosis. New York: Churchill Livingstone.
20. J. R. Connor S. L. Menzies 1995 Cellular management of iron in the brain Journal of the Neurological Sciences 134 Suppl 33 44
21. S. D. D'Souza K. A. Alinauskas J. P. Antel 1996 Ciliary neurotrophic factor selectively protects human oligodendrocytes from tumor necrosis factor-mediated injury Journal of Neuroscience Research 43 289 298
22. S. D. D'Souza J. P. Antel 1994 Freedman MS. Cytokine induction of heat shock protein expression in human oligodendrocytes: An interleukin-1-mediated mechanism Journal of Neuroimmunology 50 17 24
23. A. da Cunha J. A. Jefferson R. W. Jackson L. Vitkovic 1993 Glial cell-specific mechanisms of TGF-beta 1 induction by IL-1 in cerebral cortex Journal of Neuroimmunology 42 71 85
24. C. J. De Groot S. R. Ruuls J. W. Theeuwes C. D. Dijkstra P. Van der Valk 1997 Immunocytochemical characterization of the expression of inducible and constitutive isoforms of nitric oxide synthase in demyelinating multiple sclerosis lesions Journal of Neuropathology and Experimental Neurology 56 10 20
25. Y. Du C. F. Dreyfus 2002 Oligodendrocytes as providers of growth factors Journal of Neuroscience Research 68 647 654
26. M. S. Freedman T. C. Ruijs L. K. Selin J. P. Antel 1991 Peripheral blood gamma-delta T cells lyse fresh human brain-derived oligodendrocytes Annals of Neurology 30 794 800
27. C. P. Genain B. Cannella S. L. Hauser C. S. Raine 1999 Identification of autoantibodies associated with myelin damage in multiple sclerosis Natural Medicines 5 170 175
28. C. L. Gibson T. C. Coughlan S. P. Murphy 2005 Glial nitric oxide and ischemia Glia 50 417 426
29. R. Gold H. P. Hartung K. V. Toyka 2000 Animal models for autoimmune demyelinating disorders of the nervous system Molecular Medicine Today 6 88 91
30. R. Gold C. Linington H. Lassmann 2006 Understanding pathogenesis and therapy of multiple sclerosis via animal models: 70 years of merits and culprits in experimental autoimmune encephalomyelitis research Brain 129 1953 1971
31. U. Graumann R. Reynolds A. J. Steck N. Schaeren-Wiemers 2003 Molecular changes in normal appearing white matter in multiple sclerosis are characteristic of neuroprotective mechanisms against hypoxic insult Brain Pathology 13 554 573
32. Y. Grenier T. C. Ruijs Y. Robitaille A. Olivier J. P. Antel 1989 Immunohistochemical studies of adult human glial cells Journal of Neuroimmunology 21 103 115
33. M. Greter F. L. Heppner M. P. Lemos B. M. Odermatt N. Goebels T. Laufer 2005 Dendritic cells permit immune invasion of the CNS in an animal model of multiple sclerosis Natural Medicines 11 328 334
34. M. Griot-Wenk C. Griot H. Pfister M. Vandevelde 1991 Antibody-dependent cellular cytotoxicity in antimyelin antibody-induced oligodendrocyte damage in vitro Journal of Neuroimmunology 33 145 155
35. I. Gutcher E. Urich K. Wolter M. Prinz B. Becher 2006 Interleukin 18-independent engagement of interleukin 18 receptor-alpha is required for autoimmune inflammation Nature Immunology 7 946 953
36. J. M. Gutteridge B. Halliwell 1989 Iron toxicity and oxygen radicals Baillieres Clinics in Haematology 2 195 256
37. K. E. Hill L. V. Zollinger H. E. Watt N. G. Carlson J. W. Rose 2004 Inducible nitric oxide synthase in chronic active multiple sclerosis plaques: Distribution, cellular expression and association with myelin damage Journal of Neuroimmunology 151 171 179
38. R. Hoftberger F. Aboul-Enein W. Brueck C. Lucchinetti M. Rodriguez M. Schmidbauer 2004 Expression of major histocompatibility complex class I molecules on the different cell types in multiple sclerosis lesions Brain Pathology 14 43 50
39. E. S. Huseby D. Liggitt T. Brabb B. Schnabel C. Ohlen J. Goverman 2001 A pathogenic role for myelin-specific CD8(+) T cells in a model for multiple sclerosis Journal of Experimental Medicine 194 669 676
40. Y. Iwakura H. Ishigame 2006 The IL-23/IL-17 axis in inflammation Journal of Clinical Investigation 116 1218 1222
41. C. Jack J. Antel W. Bruck T. Kuhlmann 2007 Contrasting potential of nitric oxide and peroxynitrite to mediate oligodendrocyte injury in multiple sclerosis Glia 55 926 934
42. A. Jurewicz W. E. Biddison J. P. Antel 1998 MHC class I-restricted lysis of human oligodendrocytes by myelin basic protein peptide-specific CD8 T lymphocytes Journal of Immunology 160 3056 3059
43. A. Jurewicz M. Matysiak K. Tybor L. Kilianek C. S. Raine K. Selmaj 2005 Tumour necrosis factor-induced death of adult human oligodendrocytes is mediated by apoptosis inducing factor Brain 128 2675 2688
44. B. H. Juurlink S. K. Thorburne L. Hertz 1998 Peroxide-scavenging deficit underlies oligodendrocyte susceptibility to oxidative stress Glia 22 371 378
45. M. H. Kaplan U. Schindler S. T. Smiley M. J. Grusby 1996 Stat6 is required for mediating responses to IL-4 and for development of Th2 cells Immunity 4 313 319
46. P. G. Kennedy I. Steiner 1994 On the possible viral aetiology of multiple sclerosis QJM 87 523 528
47. S. U. Kim 1985 Antigen expression by glial cells grown in culture Journal of Neuroimmunology 8 255 282
48. K. Kreymborg R. Etzensperger L. Dumoutier S. Haak A. Rebollo T. Buch 2007 IL-22 is expressed by Th17 cells in an IL-23-dependent fashion, but not required for the development of autoimmune encephalomyelitis Journal of Immunology 179 8098 8104
49. C. L. Langrish Y. Chen W. M. Blumenschein J. Mattson B. Basham J. D. Sedgwick 2005 IL-23 drives a pathogenic T cell population that induces autoimmune inflammation Journal of Experimental Medicine 201 233 240
50. H. Lassmann 2003 Hypoxia-like tissue injury as a component of multiple sclerosis lesions Journal of the Neurological Sciences 206 187 191
51. H. Lassmann R. M. Ransohoff 2004 The CD4-Th1 model for multiple sclerosis: A critical [correction of crucial] re-appraisal Trends in Immunology 25 132 137
52. S. C. Lee C. S. Raine 1989 Multiple sclerosis: Oligodendrocytes in active lesions do not express class II major histocompatibility complex molecules Journal of Neuroimmunology 25 261 266
53. J. Li O. Baud T. Vartanian J. J. Volpe P. A. Rosenberg 2005 Peroxynitrite generated by inducible nitric oxide synthase and NADPH oxidase mediates microglial toxicity to oligodendrocytes Proceedings of the National Academy of Sciences of the United States of America 102 9936 9941
54. H. Li M. I. Lebedeva A. S. Llera B. A. Fields M. B. Brenner R. A. Mariuzza 1998 Structure of the Vdelta domain of a human gammadelta T-cell antigen receptor Nature 391 502 506
55. W. Li Y. Maeda X. Ming S. Cook J. Chapin W. Husar 2002 Apoptotic death following Fas activation in human oligodendrocyte hybrid cultures Journal of Neuroscience Research 69 189 196
56. W. Lin S. L. Bailey H. Ho H. P. Harding D. Ron S. D. Miller 2007 The integrated stress response prevents demyelination by protecting oligodendrocytes against immune-mediated damage Journal of Clinical Investigation 117 448 456
57. C. Linington M. Bradl H. Lassmann C. Brunner K. Vass 1988 Augmentation of demyelination in rat acute allergic encephalomyelitis by circulating mouse monoclonal antibodies directed against a myelin/oligodendrocyte glycoprotein American Journal of Pathology 130 443 454
58. X. Liu D. L. Yao H. Webster 1995 Insulin-like growth factor I treatment reduces clinical deficits and lesion severity in acute demyelinating experimental autoimmune encephalomyelitis Multiple Sclerosis 1 2 9
59. C. Lucchinetti W. Bruck J. Parisi B. Scheithauer M. Rodriguez H. Lassmann 2000 Heterogeneity of multiple sclerosis lesions: Implications for the pathogenesis of demyelination Annals of Neurology 47 707 717
60. S. K. Ludwin 1997 The pathobiology of the oligodendrocyte Journal of Neuropathology and Experimental Neurology 56 111 124
61. R. Magliozzi O. Howell A. Vora B. Serafini R. Nicholas M. Puopolo 2007 Meningeal B-cell follicles in secondary progressive multiple sclerosis associate with early onset of disease and severe cortical pathology Brain 130 1089 1104
62. M. J. McGeachy K. S. Bak-Jensen Y. Chen C. M. Tato W. Blumenschein T. McClanahan 2007 TGF-beta and IL-6 drive the production of IL-17 and IL-10 by T cells and restrain T(H)-17 cell-mediated pathology Nature Immunology 8 1390 1397
63. R. D. McKinnon G. Piras J. A. Ida Jr. M. Dubois-Dalcq 1993 A role for TGF-beta in oligodendrocyte differentiation Journal of Cell Biology 121 1397 1407
64. R. J. Mead S. K. Singhrao J. W. Neal H. Lassmann B. P. Morgan 2002 The membrane attack complex of complement causes severe demyelination associated with acute axonal injury Journal of Immunology 168 458 465
65. J. E. Merrill L. J. Ignarro M. P. Sherman J. Melinek T. E. Lane 1993 Microglial cell cytotoxicity of oligodendrocytes is mediated through nitric oxide Journal of Immunology 151 2132 2141
66. J. E. Merrill N. J. Scolding 1999 Mechanisms of damage to myelin and oligodendrocytes and their relevance to disease Neuropathology & Applied Neurobiology 25 435 458
67. A. Micera F. Properzi V. Triaca L. Aloe 2000 Nerve growth factor antibody exacerbates neuropathological signs of experimental allergic encephalomyelitis in adult lewis rats Journal of Neuroimmunology 104 116 123
68. B. Mitrovic L. J. Ignarro H. V. Vinters M. A. Akers I. Schmid C. Uittenbogaart 1995 Nitric oxide induces necrotic but not apoptotic cell death in oligodendrocytes Neuroscience 65 531 539
69. S. Nataf P. Naveilhan L. Sindji F. Darcy P. Brachet C. N. Montero-Menei 1998 Low affinity NGF receptor expression in the central nervous system during experimental allergic encephalomyelitis Journal of Neuroscience Research 52 83 92
70. J. Newcombe A. Uddin R. Dove B. Patel L. Turski Y. Nishizawa 2007 Glutamate receptor expression in multiple sclerosis lesions Brain Pathology 18 1 52 61
71. K. C. O'Connor H. Appel L. Bregoli M. E. Call I. Catz J. A. Chan 2005 Antibodies from inflamed central nervous system tissue recognize myelin oligodendrocyte glycoprotein Journal of Immunology 175 1974 1982
72. B. Oderfeld-Nowak M. Zaremba A. W. Lipkowski B. Kwiatkowska-Patzer V. Triaca L. Aloe 2003 High-affinity NGF receptor in the rat spinal cord during acute and chronic phases of experimental autoimmune encephalomyelitis: A possible functional significance Archives Italiennes de Biologie 141 103 116
73. B. Oderfeld-Nowak M. Zaremba A. Micera L. Aloe 2001 The upregulation of nerve growth factor receptors in reactive astrocytes of rat spinal cord during experimental autoimmune encephalomyelitis Neuroscience Letters 308 165 168
74. A. Odyniec M. Szczepanik M. P. Mycko M. Stasiolek C. S. Raine K. W. Selmaj 2004 Gammadelta T cells enhance the expression of experimental autoimmune encephalomyelitis by promoting antigen presentation and IL-12 production Journal of Immunology 173 682 694
75. K. Ozawa G. Suchanek H. Breitschopf W. Bruck H. Budka K. Jellinger 1994 Patterns of oligodendroglia pathology in multiple sclerosis Brain 117 Pt 6 1311 1322
76. J. Parkin B. Cohen 2001 An overview of the immune system Lancet 357 1777 1789
77. B. M. Pratt J. M. McPherson 1997 TGF-beta in the central nervous system: Potential roles in ischemic injury and neurodegenerative diseases Cytokine and Growth Factor Reviews 8 267 292
78. C. S. Raine 1997 The Norton lecture: A review of the oligodendrocyte in the multiple sclerosis lesion Journal of Neuroimmunology 77 135 152
79. C. S. Raine B. Bonetti B. Cannella 1998 Multiple sclerosis: Expression of molecules of the tumor necrosis factor ligand and receptor families in relationship to the demyelinated plaque Revista de Neurología (Paris) 154 577 585
80. A. J. Rajan Y. L. Gao C. S. Raine C. F. Brosnan 1996 A pathogenic role for gamma delta T cells in relapsing-remitting experimental allergic encephalomyelitis in the SJL mouse Journal of Immunology 157 941 949
81. E. J. Redford R. Kapoor K. J. Smith 1997 Nitric oxide donors reversibly block axonal conduction: Demyelinated axons are especially susceptible Brain 120 Pt 12 2149 2157
82. J. M. Redwine M. J. Buchmeier C. F. Evans 2001 In vivo expression of major histocompatibility complex molecules on oligodendrocytes and neurons during viral infection American Journal of Pathology 159 1219 1224
83. M. Reindl C. Linington U. Brehm R. Egg E. Dilitz F. Deisenhammer 1999 Antibodies against the myelin oligodendrocyte glycoprotein and the myelin basic protein in multiple sclerosis and other neurological diseases: A comparative study Brain 122 Pt 11 2047 2056
84. A. J. Roskams J. R. Connor 1994 Iron, transferrin, and ferritin in the rat brain during development and aging Journal of Neurochemistry 63 709 716
85. T. C. Ruijs M. S. Freedman Y. G. Grenier A. Olivier J. P. Antel 1990 Human oligodendrocytes are susceptible to cytolysis by major histocompatibility complex class I-restricted lymphocytes Journal of Neuroimmunology 27 89 97
86. N. J. Scolding D. A. Compston 1991 Oligodendrocyte-macrophage interactions in vitro triggered by specific antibodies Immunology 72 127 132
87. K. W. Selmaj C. S. Raine 1988 Tumor necrosis factor mediates myelin and oligodendrocyte damage in vitro Annals of Neurology 23 339 346
88. G. Semenza 2002 Signal transduction to hypoxia-inducible factor 1 Biochemical Pharmacology 64 993 998
89. A. Setzu J. D. Lathia C. Zhao K. Wells M. S. Rao C. Ffrench-Constant 2006 Inflammation stimulates myelination by transplanted oligodendrocyte precursor cells Glia 54 297 303
90. F. R. Sharp M. Bergeron M. Bernaudin 2001 Hypoxia-inducible factor in brain Advances in Experimental Medicine and Biology 502 273 291
91. + T cells persist as clonal expansions in the cerebrospinal fluid and blood Proceedings of the National Academy of Sciences of the United States of America 101 2428 2433
92. K. J. Smith R. Kapoor P. A. Felts 1999 Demyelination: The role of reactive oxygen and nitrogen species Brain Pathology 9 69 92
93. K. J. Smith H. Lassmann 2002 The role of nitric oxide in multiple sclerosis Lancet Neurology 1 232 241
94. M. Sospedra R. Martin 2005 Immunology of multiple sclerosis Annual Review of Immunology 23 683 747
95. C. Stadelmann S. Ludwin T. Tabira A. Guseo C. F. Lucchinetti L. Leel-Ossy 2005 Tissue preconditioning may explain concentric lesions in Balo's type of multiple sclerosis Brain 128 979 987
96. T. Stahnke C. Stadelmann A. Netzler W. Bruck C. Richter-Landsberg 2007 Differential upregulation of heme oxygenase-1 (HSP32) in glial cells after oxidative stress and in demyelinating disorders Journal of Molecular Neuroscience 32 25 37
97. + T cells are encephalitogenic and produce severe disease in C57BL/6 mice Journal of Immunology 166 7579 7587
98. S. K. Thorburne B. H. Juurlink 1996 Low glutathione and high iron govern the susceptibility of oligodendroglial precursors to oxidative stress Journal of Neurochemistry 67 1014 1022
99. R. C. van der Veen L. J. Roberts 1999 Contrasting roles for nitric oxide and peroxynitrite in the peroxidation of myelin lipids Journal of Neuroimmunology 95 1 7
100. P. Villoslada S. L. Hauser I. Bartke J. Unger N. Heald D. Rosenberg 2000 Human nerve growth factor protects common marmosets against autoimmune encephalomyelitis by switching the balance of T helper cell type 1 and 2 cytokines within the central nervous system Journal of Experimental Medicine 191 1799 1806
101. C. T. Weaver L. E. Harrington P. R. Mangan M. Gavrieli K. M. Murphy 2006 Th17: An effector CD4 T cell lineage with regulatory T cell ties Immunity 24 677 688
102. D. O. Willenborg M. A. Staykova W. B. Cowden 1999 Our shifting understanding of the role of nitric oxide in autoimmune encephalomyelitis: A review Journal of Neuroimmunology 100 21 35
103. A. Williams G. Piaton C. Lubetzki 2007 Astrocytes-friends or foes in multiple sclerosis? Glia 55 1300 1312
104. K. W. Wucherpfennig J. Newcombe H. Li C. Keddy M. L. Cuzner D. A. Hafler 1992 Gamma delta T-cell receptor repertoire in acute multiple sclerosis lesions Proceedings of the National Academy of Sciences of the United States of America 89 4588 4592
105. P. Ye G. Kollias D A. J. Ercole 2007 Insulin-like growth factor-I ameliorates demyelination induced by tumor necrosis factor-alpha in transgenic mice Journal of Neuroscience Research 85 712 722
106. S. O. Yoon P. Casaccia-Bonnefil B. Carter M. V. Chao 1998 Competitive signaling between TrkA and p75 nerve growth factor receptors determines cell survival Journal of Neuroscience 18 3273 3281
107. T. Zeis U. Graumann R. Reynolds N. Schaeren-Wiemers 2008 Normal-appearing white matter in multiple sclerosis is in a subtle balance between inflammation and neuroprotection Brain 131 288 303
108. J. Zhang V. L. Dawson T. M. Dawson S. H. Snyder 1994 Nitric oxide activation of poly(ADP-ribose) synthetase in neurotoxicity Science 263 687 689