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PMC0
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10.1038%2Fnri1105
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main
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Many neuronotropic viruses spread within the nervous system by axonal transport and move from neuron to neuron through connecting synapses>>1<<. If infection is induced experimentally by intracerebral inoculation, there might also be rapid spread through the cerebrospinal fluid2. Other viruses target the supporting glial cells of the CNS — oligodendrocytes, astrocytes and
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If infection is induced experimentally by intracerebral inoculation, there might also be rapid spread through the cerebrospinal fluid>>2<<. Other viruses target the supporting glial cells of the CNS — oligodendrocytes, astrocytes and microglia — alone or in addition to neurons. Oligodendrocytes form and maintain the myelin that surrounds the axons of neurons, and infection
n2:mentions
n3:9257852
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molecules that promote interaction with circulating leukocytes. Microglia, which express the CD200 receptor (CD200R), are kept in a quiescent state through interaction with electrically active, healthy neurons, which express CD200 (Ref. >>3<<), and by the local production of neurotrophins4. Production of the anti-inflammatory cytokine transforming growth factor-β (TGF-β) by astrocytes and meningeal cells suppresses activation of endothelial and other cells by cytokines5,6(Fig.
n2:mentions
n3:11099416
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3), and by the local production of neurotrophins>>4<<. Production of the anti-inflammatory cytokine transforming growth factor-β (TGF-β) by astrocytes and meningeal cells suppresses activation of endothelial and other cells by cytokines5,6(Fig.
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Production of the anti-inflammatory cytokine transforming growth factor-β (TGF-β) by astrocytes and meningeal cells suppresses activation of endothelial and other cells by cytokines>>5<<,6(Fig. 1a).Figure 1Changes that occur in the CNS during the immune response to a neuronal virus infection.
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Production of the anti-inflammatory cytokine transforming growth factor-β (TGF-β) by astrocytes and meningeal cells suppresses activation of endothelial and other cells by cytokines5,>>6<<(Fig. 1a).Figure 1Changes that occur in the CNS during the immune response to a neuronal virus infection.
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Rapid production of TYPE 1 INTERFERON (IFN) is important for host survival and, in most experimental virus infections of the CNS, susceptibility to fatal disease is markedly increased in mice that lack the receptor for IFN-α/β>>7<<,8,9,10. IFN-β, an immediate early IFN, which is produced by neurons as well as glial cells, is the main type 1 IFN that is produced by the CNS and is required for efficient induction of IFN-α in vitro11,12. Preferential production of
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Rapid production of TYPE 1 INTERFERON (IFN) is important for host survival and, in most experimental virus infections of the CNS, susceptibility to fatal disease is markedly increased in mice that lack the receptor for IFN-α/β7,>>8<<,9,10. IFN-β, an immediate early IFN, which is produced by neurons as well as glial cells, is the main type 1 IFN that is produced by the CNS and is required for efficient induction of IFN-α in vitro11,12. Preferential production of IFN-β
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Rapid production of TYPE 1 INTERFERON (IFN) is important for host survival and, in most experimental virus infections of the CNS, susceptibility to fatal disease is markedly increased in mice that lack the receptor for IFN-α/β7,8,>>9<<,10. IFN-β, an immediate early IFN, which is produced by neurons as well as glial cells, is the main type 1 IFN that is produced by the CNS and is required for efficient induction of IFN-α in vitro11,12. Preferential production of IFN-β
n2:mentions
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Rapid production of TYPE 1 INTERFERON (IFN) is important for host survival and, in most experimental virus infections of the CNS, susceptibility to fatal disease is markedly increased in mice that lack the receptor for IFN-α/β7,8,9,>>10<<. IFN-β, an immediate early IFN, which is produced by neurons as well as glial cells, is the main type 1 IFN that is produced by the CNS and is required for efficient induction of IFN-α in vitro11,12. Preferential production of IFN-β
n2:mentions
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IFN-β, an immediate early IFN, which is produced by neurons as well as glial cells, is the main type 1 IFN that is produced by the CNS and is required for efficient induction of IFN-α in vitro>>11<<,12. Preferential production of IFN-β rather than IFN-α in the CNS might be neuroprotective, as IFN-α has marked CNS toxicity compared with IFN-β13,14, and IFN-β might induce the production of neurotrophic factors by astrocytes15 and the
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n3:7933119
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IFN-β, an immediate early IFN, which is produced by neurons as well as glial cells, is the main type 1 IFN that is produced by the CNS and is required for efficient induction of IFN-α in vitro11,>>12<<. Preferential production of IFN-β rather than IFN-α in the CNS might be neuroprotective, as IFN-α has marked CNS toxicity compared with IFN-β13,14, and IFN-β might induce the production of neurotrophic factors by astrocytes15 and the
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n3:9501984
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Preferential production of IFN-β rather than IFN-α in the CNS might be neuroprotective, as IFN-α has marked CNS toxicity compared with IFN-β>>13<<,14, and IFN-β might induce the production of neurotrophic factors by astrocytes15 and the local production of the anti-inflammatory cytokine interleukin-10 (IL-10)16.
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n2:Context
rdf:value
Preferential production of IFN-β rather than IFN-α in the CNS might be neuroprotective, as IFN-α has marked CNS toxicity compared with IFN-β13,>>14<<, and IFN-β might induce the production of neurotrophic factors by astrocytes15 and the local production of the anti-inflammatory cytokine interleukin-10 (IL-10)16.
n2:mentions
n3:9345464
Subject Item
_:vb515155
rdf:type
n2:Context
rdf:value
Preferential production of IFN-β rather than IFN-α in the CNS might be neuroprotective, as IFN-α has marked CNS toxicity compared with IFN-β13,14, and IFN-β might induce the production of neurotrophic factors by astrocytes>>15<< and the local production of the anti-inflammatory cytokine interleukin-10 (IL-10)16.
n2:mentions
n3:9282915
Subject Item
_:vb515156
rdf:type
n2:Context
rdf:value
neuroprotective, as IFN-α has marked CNS toxicity compared with IFN-β13,14, and IFN-β might induce the production of neurotrophic factors by astrocytes15 and the local production of the anti-inflammatory cytokine interleukin-10 (IL-10)>>16<<. Rapid local production of type 1 IFN after infection slows virus spread and constrains virus replication before the induction of virus-specific adaptive immune responses.
n2:mentions
n3:11094104
Subject Item
_:vb515157
rdf:type
n2:Context
rdf:value
Virus inhibition of type 1 IFN production or activity — for example, by the leader protein of TMEV — increases the probability that the virus will not be cleared>>17<<.
n2:mentions
n3:11483724
Subject Item
_:vb515158
rdf:type
n2:Context
rdf:value
Damaged or stressed neurons signal to astrocytes and microglia in the vicinity of the neuronal cell body, even when the neuronal damage occurs to the axon at a distance>>18<<. The factors that are produced by neurons in response to injury — which includes injury caused by virus infection — are not completely catalogued, but they probably include the cytokines IFN-γ and IL-6, and the chemokines fractalkine
n2:mentions
n3:11596127
Subject Item
_:vb515159
rdf:type
n2:Context
rdf:value
injury caused by virus infection — are not completely catalogued, but they probably include the cytokines IFN-γ and IL-6, and the chemokines fractalkine (CX3CL1) and secondary lymphoid tissue chemokine (SLC, CCL21), in addition to IFN-β>>19<<,20,21. CX3CL1 is a membrane-bound chemokine that is constitutively expressed by subsets of neurons and is rapidly cleaved from the cell surface after neurotoxic damage22.
n2:mentions
n3:9396771
Subject Item
_:vb515160
rdf:type
n2:Context
rdf:value
caused by virus infection — are not completely catalogued, but they probably include the cytokines IFN-γ and IL-6, and the chemokines fractalkine (CX3CL1) and secondary lymphoid tissue chemokine (SLC, CCL21), in addition to IFN-β19,>>20<<,21. CX3CL1 is a membrane-bound chemokine that is constitutively expressed by subsets of neurons and is rapidly cleaved from the cell surface after neurotoxic damage22.
n2:mentions
n3:9724801
Subject Item
_:vb515161
rdf:type
n2:Context
rdf:value
caused by virus infection — are not completely catalogued, but they probably include the cytokines IFN-γ and IL-6, and the chemokines fractalkine (CX3CL1) and secondary lymphoid tissue chemokine (SLC, CCL21), in addition to IFN-β19,20,>>21<<. CX3CL1 is a membrane-bound chemokine that is constitutively expressed by subsets of neurons and is rapidly cleaved from the cell surface after neurotoxic damage22.
n2:mentions
n3:11907075
Subject Item
_:vb515162
rdf:type
n2:Context
rdf:value
CX3CL1 is a membrane-bound chemokine that is constitutively expressed by subsets of neurons and is rapidly cleaved from the cell surface after neurotoxic damage>>22<<. Macrophages and glial cells express the CX3CL1 receptor (CX3CR1) and so become activated in early phases of the response to neuronal infection23,24, in which they have an important role. Activated microglia and astrocytes can rapidly
n2:mentions
n3:10899174
Subject Item
_:vb515163
rdf:type
n2:Context
rdf:value
Macrophages and glial cells express the CX3CL1 receptor (CX3CR1) and so become activated in early phases of the response to neuronal infection>>23<<,24, in which they have an important role.
n2:mentions
n3:10805752
Subject Item
_:vb515164
rdf:type
n2:Context
rdf:value
Macrophages and glial cells express the CX3CL1 receptor (CX3CR1) and so become activated in early phases of the response to neuronal infection23,>>24<<, in which they have an important role.
n2:mentions
n3:10415068
Subject Item
_:vb515165
rdf:type
n2:Context
rdf:value
macrophage inflammatory protein-1β (MIP1β, CCL4), monocyte chemo attractant protein 1 (MCP1, CCL2), MCP3 (CCL7), regulated on activation normal T-cell expressed and secreted (RANTES, CCL5) and IFN-inducible 10 kDa protein (IP10, CXCL10)>>25<<,26,27,28,29,30. Production of these factors results in the upregulation of expression of MHC molecules on the cell surface of microglia and in increased expression of adhesion molecules by capillary endothelial cells.
n2:mentions
n3:9311871
Subject Item
_:vb515166
rdf:type
n2:Context
rdf:value
inflammatory protein-1β (MIP1β, CCL4), monocyte chemo attractant protein 1 (MCP1, CCL2), MCP3 (CCL7), regulated on activation normal T-cell expressed and secreted (RANTES, CCL5) and IFN-inducible 10 kDa protein (IP10, CXCL10)25,>>26<<,27,28,29,30. Production of these factors results in the upregulation of expression of MHC molecules on the cell surface of microglia and in increased expression of adhesion molecules by capillary endothelial cells.
n2:mentions
n3:9551936
Subject Item
_:vb515167
rdf:type
n2:Context
rdf:value
inflammatory protein-1β (MIP1β, CCL4), monocyte chemo attractant protein 1 (MCP1, CCL2), MCP3 (CCL7), regulated on activation normal T-cell expressed and secreted (RANTES, CCL5) and IFN-inducible 10 kDa protein (IP10, CXCL10)25,26,>>27<<,28,29,30. Production of these factors results in the upregulation of expression of MHC molecules on the cell surface of microglia and in increased expression of adhesion molecules by capillary endothelial cells.
n2:mentions
n3:8301132
Subject Item
_:vb515168
rdf:type
n2:Context
rdf:value
inflammatory protein-1β (MIP1β, CCL4), monocyte chemo attractant protein 1 (MCP1, CCL2), MCP3 (CCL7), regulated on activation normal T-cell expressed and secreted (RANTES, CCL5) and IFN-inducible 10 kDa protein (IP10, CXCL10)25,26,27,>>28<<,29,30. Production of these factors results in the upregulation of expression of MHC molecules on the cell surface of microglia and in increased expression of adhesion molecules by capillary endothelial cells.
n2:mentions
n3:9217050
Subject Item
_:vb515169
rdf:type
n2:Context
rdf:value
inflammatory protein-1β (MIP1β, CCL4), monocyte chemo attractant protein 1 (MCP1, CCL2), MCP3 (CCL7), regulated on activation normal T-cell expressed and secreted (RANTES, CCL5) and IFN-inducible 10 kDa protein (IP10, CXCL10)25,26,27,28,>>29<<,30. Production of these factors results in the upregulation of expression of MHC molecules on the cell surface of microglia and in increased expression of adhesion molecules by capillary endothelial cells.
n2:mentions
n3:11118358
Subject Item
_:vb515170
rdf:type
n2:Context
rdf:value
protein-1β (MIP1β, CCL4), monocyte chemo attractant protein 1 (MCP1, CCL2), MCP3 (CCL7), regulated on activation normal T-cell expressed and secreted (RANTES, CCL5) and IFN-inducible 10 kDa protein (IP10, CXCL10)25,26,27,28,29,>>30<<. Production of these factors results in the upregulation of expression of MHC molecules on the cell surface of microglia and in increased expression of adhesion molecules by capillary endothelial cells.
n2:mentions
n3:11031689
Subject Item
_:vb515171
rdf:type
n2:Context
rdf:value
Chemokines are transcytosed to the endothelial luminal surface and presented to circulating leukocytes on the tips of microvillous processes>>31<<.
n2:mentions
n3:12433694
Subject Item
_:vb515172
rdf:type
n2:Context
rdf:value
It is generally agreed that, although there are cells that can present antigen to primed T cells in the CNS, the activation of naive T cells and B cells occurs in secondary lymphoid tissues outside the CNS>>32<<,33,34. The parenchyma of the normal CNS does not contain dendritic cells, although they are present in the meninges and cerebro–spinal fluid (CSF)35. Some of the CSF and CNS interstitial fluid drains to the deep cervical lymph nodes
n2:mentions
n3:10580795
Subject Item
_:vb515173
rdf:type
n2:Context
rdf:value
It is generally agreed that, although there are cells that can present antigen to primed T cells in the CNS, the activation of naive T cells and B cells occurs in secondary lymphoid tissues outside the CNS32,>>33<<,34. The parenchyma of the normal CNS does not contain dendritic cells, although they are present in the meninges and cerebro–spinal fluid (CSF)35. Some of the CSF and CNS interstitial fluid drains to the deep cervical lymph nodes through
n2:mentions
n3:11160337
Subject Item
_:vb515174
rdf:type
n2:Context
rdf:value
It is generally agreed that, although there are cells that can present antigen to primed T cells in the CNS, the activation of naive T cells and B cells occurs in secondary lymphoid tissues outside the CNS32,33,>>34<<. The parenchyma of the normal CNS does not contain dendritic cells, although they are present in the meninges and cerebro–spinal fluid (CSF)35. Some of the CSF and CNS interstitial fluid drains to the deep cervical lymph nodes through the
n2:mentions
n3:9733812
Subject Item
_:vb515175
rdf:type
n2:Context
rdf:value
The parenchyma of the normal CNS does not contain dendritic cells, although they are present in the meninges and cerebro–spinal fluid (CSF)>>35<<. Some of the CSF and CNS interstitial fluid drains to the deep cervical lymph nodes through the nasal submucosa32,36. Experimental inoculation of virus directly into the CSF elicits a potent immune response, whereas inoculation directly
n2:mentions
n3:10098945
Subject Item
_:vb515176
rdf:type
n2:Context
rdf:value
Some of the CSF and CNS interstitial fluid drains to the deep cervical lymph nodes through the nasal submucosa>>32<<,36. Experimental inoculation of virus directly into the CSF elicits a potent immune response, whereas inoculation directly into the brain parenchyma elicits little or no response37. Responses that are initiated by inoculation of antigen
n2:mentions
n3:10580795
Subject Item
_:vb515177
rdf:type
n2:Context
rdf:value
Some of the CSF and CNS interstitial fluid drains to the deep cervical lymph nodes through the nasal submucosa32,>>36<<. Experimental inoculation of virus directly into the CSF elicits a potent immune response, whereas inoculation directly into the brain parenchyma elicits little or no response37. Responses that are initiated by inoculation of antigen into
n2:mentions
n3:1463583
Subject Item
_:vb515178
rdf:type
n2:Context
rdf:value
Experimental inoculation of virus directly into the CSF elicits a potent immune response, whereas inoculation directly into the brain parenchyma elicits little or no response>>37<<. Responses that are initiated by inoculation of antigen into the CSF tend to be characterized by marked antibody responses, priming of cytotoxic CD8+ T cells and no DELAYED-TYPE HYPERSENSITIVITY32. However, essentially all neurotropic
n2:mentions
n3:8985333
Subject Item
_:vb515179
rdf:type
n2:Context
rdf:value
Responses that are initiated by inoculation of antigen into the CSF tend to be characterized by marked antibody responses, priming of cytotoxic CD8+ T cells and no DELAYED-TYPE HYPERSENSITIVITY>>32<<. However, essentially all neurotropic viruses naturally initiate infection in the periphery either through the gastrointestinal tract, the respiratory tract or the bite of an infected mosquito or animal. Therefore, induction of the
n2:mentions
n3:10580795
Subject Item
_:vb515180
rdf:type
n2:Context
rdf:value
entry of circulating leukocytes into the CNS is generally restricted by the tight junctions and by the relative nonreactivity of cerebral capillary endothelial cells, which have limited endocytosis and expression of adhesion molecules>>38<<,39. However, several studies have shown that activated T cells routinely cross the blood–brain barrier as part of the normal immunological surveillance of all tissues39,40,41 (Fig.
n2:mentions
n3:10825349
Subject Item
_:vb515181
rdf:type
n2:Context
rdf:value
entry of circulating leukocytes into the CNS is generally restricted by the tight junctions and by the relative nonreactivity of cerebral capillary endothelial cells, which have limited endocytosis and expression of adhesion molecules38,>>39<<. However, several studies have shown that activated T cells routinely cross the blood–brain barrier as part of the normal immunological surveillance of all tissues39,40,41 (Fig.
n2:mentions
n3:8621923
Subject Item
_:vb515182
rdf:type
n2:Context
rdf:value
However, several studies have shown that activated T cells routinely cross the blood–brain barrier as part of the normal immunological surveillance of all tissues>>39<<,40,41 (Fig.
n2:mentions
n3:8621923
Subject Item
_:vb515183
rdf:type
n2:Context
rdf:value
However, several studies have shown that activated T cells routinely cross the blood–brain barrier as part of the normal immunological surveillance of all tissues39,40,>>41<< (Fig. 1a). Transendothelial migration of activated CD4+ T cells occurs, in part, through the interaction of P-selectin glycoprotein ligand, which is expressed by activated T cells, with P-selectin, which is expressed at low levels by
n2:mentions
n3:2033653
Subject Item
_:vb515184
rdf:type
n2:Context
rdf:value
of activated CD4+ T cells occurs, in part, through the interaction of P-selectin glycoprotein ligand, which is expressed by activated T cells, with P-selectin, which is expressed at low levels by normal cerebrovascular endothelial cells>>38<<,42. However, one side effect of the induction of a systemic immune response, even in the absence of infection in the CNS, is the upregulation of expression of adhesion molecules by CNS endothelial cells and increased surveillance of the
n2:mentions
n3:10825349
Subject Item
_:vb515185
rdf:type
n2:Context
rdf:value
activated CD4+ T cells occurs, in part, through the interaction of P-selectin glycoprotein ligand, which is expressed by activated T cells, with P-selectin, which is expressed at low levels by normal cerebrovascular endothelial cells38,>>42<<. However, one side effect of the induction of a systemic immune response, even in the absence of infection in the CNS, is the upregulation of expression of adhesion molecules by CNS endothelial cells and increased surveillance of the CNS
n2:mentions
n3:11823530
Subject Item
_:vb515186
rdf:type
n2:Context
rdf:value
of the induction of a systemic immune response, even in the absence of infection in the CNS, is the upregulation of expression of adhesion molecules by CNS endothelial cells and increased surveillance of the CNS by activated T cells>>43<<,44. Activated T cells that enter the CNS are not retained in the CNS in the absence of antigen, and either leave or die in situ (Fig.
n2:mentions
n3:11596120
Subject Item
_:vb515187
rdf:type
n2:Context
rdf:value
of the induction of a systemic immune response, even in the absence of infection in the CNS, is the upregulation of expression of adhesion molecules by CNS endothelial cells and increased surveillance of the CNS by activated T cells43,>>44<<. Activated T cells that enter the CNS are not retained in the CNS in the absence of antigen, and either leave or die in situ (Fig.
n2:mentions
n3:9399938
Subject Item
_:vb515188
rdf:type
n2:Context
rdf:value
However, activated T cells are retained in the CNS when the relevant antigen is present and associated with appropriate MHC molecules>>39<<. During inflammation, local production of chemokines and induction of expression of adhesion molecules by endothelial cells further enhance the entry of activated cells into the CNS45. Expression of intercellular adhesion molecule 1
n2:mentions
n3:8621923
Subject Item
_:vb515189
rdf:type
n2:Context
rdf:value
During inflammation, local production of chemokines and induction of expression of adhesion molecules by endothelial cells further enhance the entry of activated cells into the CNS>>45<<. Expression of intercellular adhesion molecule 1 (ICAM1, CD54) and vascular-cell adhesion molecule 1 (VCAM1) is upregulated by cerebral capillary endothelial cells during virus infection, and antibody specific for the integrin leukocyte
n2:mentions
n3:12209625
Subject Item
_:vb515190
rdf:type
n2:Context
rdf:value
during virus infection, and antibody specific for the integrin leukocyte function-associated antigen 1 (LFA1), which is the ligand for ICAM1, blocks lymphocyte entry into the brain during the acute inflammatory response to Sindbis virus>>39<<.
n2:mentions
n3:8621923
Subject Item
_:vb515191
rdf:type
n2:Context
rdf:value
MHC class I molecules are not expressed by cells in the normal CNS, but are abundantly expressed by endothelial, meningeal and microglial cells during the early phases of the immune response to most CNS virus infections>>46<<, and are occasionally expressed by astrocytes and oligodendrocytes47. However, neurons rarely express MHC class I molecules.
n2:mentions
n3:10846095
Subject Item
_:vb515192
rdf:type
n2:Context
rdf:value
the normal CNS, but are abundantly expressed by endothelial, meningeal and microglial cells during the early phases of the immune response to most CNS virus infections46, and are occasionally expressed by astrocytes and oligodendrocytes>>47<<. However, neurons rarely express MHC class I molecules. Neurons respond to IFN-β with induction of antiviral proteins, such as 2′–5′ oligoadenylate synthetase, but not with expression of MHC class I molecules48.
n2:mentions
n3:2831279
Subject Item
_:vb515193
rdf:type
n2:Context
rdf:value
Neurons respond to IFN-β with induction of antiviral proteins, such as 2′–5′ oligoadenylate synthetase, but not with expression of MHC class I molecules>>48<<. Suppression of expression of MHC class I molecules by neurons seems to be both transcriptional and post transcriptional. There is little translocation from the cytoplasm to the nucleus of nuclear factor-κB (NF-κB) — a key transcription
n2:mentions
n3:10321980
Subject Item
_:vb515194
rdf:type
n2:Context
rdf:value
Lack of NF-κB translocation in neurons seems to be due to a failure to induce degradation of the inhibitor of NF-κB (IκBα) in response to cytokines>>48<<,49. However, neurons that are damaged or electrically silenced by treatment with a neurotoxin can be induced to express MHC class I molecules by treatment with IFN-γ50. In vivo, the levels of mRNAs that encode MHC class I heavy chain,
n2:mentions
n3:10321980
Subject Item
_:vb515195
rdf:type
n2:Context
rdf:value
Lack of NF-κB translocation in neurons seems to be due to a failure to induce degradation of the inhibitor of NF-κB (IκBα) in response to cytokines48,>>49<<. However, neurons that are damaged or electrically silenced by treatment with a neurotoxin can be induced to express MHC class I molecules by treatment with IFN-γ50. In vivo, the levels of mRNAs that encode MHC class I heavy chain,
n2:mentions
n3:11555675
Subject Item
_:vb515196
rdf:type
n2:Context
rdf:value
However, neurons that are damaged or electrically silenced by treatment with a neurotoxin can be induced to express MHC class I molecules by treatment with IFN-γ>>50<<. In vivo, the levels of mRNAs that encode MHC class I heavy chain, β2-microglobulin (β2m), transporter associated with antigen processing 1 (TAP1) and TAP2 are increased in neurons during infection46. However, there is controversy over
n2:mentions
n3:9016879
Subject Item
_:vb515197
rdf:type
n2:Context
rdf:value
In vivo, the levels of mRNAs that encode MHC class I heavy chain, β2-microglobulin (β2m), transporter associated with antigen processing 1 (TAP1) and TAP2 are increased in neurons during infection>>46<<. However, there is controversy over whether mature MHC class I molecules are expressed on the neuronal cell surface46,51. The β2m molecules that are sometimes detected on the neuronal cell surface might be associated with expression of
n2:mentions
n3:10846095
Subject Item
_:vb515198
rdf:type
n2:Context
rdf:value
However, there is controversy over whether mature MHC class I molecules are expressed on the neuronal cell surface>>46<<,51. The β2m molecules that are sometimes detected on the neuronal cell surface might be associated with expression of non-classical rather than classical MHC class I molecules. Differences in sensitivity of detection might also account
n2:mentions
n3:10846095
Subject Item
_:vb515199
rdf:type
n2:Context
rdf:value
However, there is controversy over whether mature MHC class I molecules are expressed on the neuronal cell surface46,>>51<<. The β2m molecules that are sometimes detected on the neuronal cell surface might be associated with expression of non-classical rather than classical MHC class I molecules. Differences in sensitivity of detection might also account for
n2:mentions
n3:10400743
Subject Item
_:vb515200
rdf:type
n2:Context
rdf:value
of detection might also account for some of the controversy that surrounds this issue — when expression of MHC class I molecules by neurons is detected, it is at markedly lower levels than expression by glial cells in the same region>>51<<.
n2:mentions
n3:10400743
Subject Item
_:vb515201
rdf:type
n2:Context
rdf:value
MHC class II molecules are not constitutively expressed by cells in the brain parenchyma, but are induced quickly after initiation of infection and also by trafficking of activated T cells through the CNS>>52<<. The cells that express MHC class II molecules most abundantly are microglia and perivascular macrophages34.
n2:mentions
n3:9605131
Subject Item
_:vb515202
rdf:type
n2:Context
rdf:value
The cells that express MHC class II molecules most abundantly are microglia and perivascular macrophages>>34<<. Astrocytes can be induced to express MHC class II molecules in vitro, but this is rarely observed in vivo34.
n2:mentions
n3:9733812
Subject Item
_:vb515203
rdf:type
n2:Context
rdf:value
Astrocytes can be induced to express MHC class II molecules in vitro, but this is rarely observed in vivo>>34<<.
n2:mentions
n3:9733812
Subject Item
_:vb515204
rdf:type
n2:Context
rdf:value
Essentially all components of the cellular immune response are detected in the infiltrate: natural killer (NK) cells, antigen-specific CD4+ and CD8+ T cells, B cells and monocyte/macrophages>>53<<,54,55,56,57.
n2:mentions
n3:1675228
Subject Item
_:vb515205
rdf:type
n2:Context
rdf:value
Essentially all components of the cellular immune response are detected in the infiltrate: natural killer (NK) cells, antigen-specific CD4+ and CD8+ T cells, B cells and monocyte/macrophages53,>>54<<,55,56,57. NK cells are detected first, followed by CD8+ and CD4+ T cells, then B cells and monocytes/macrophages58. In addition, dendritic cells are detected in the CNS during inflammation and either enter the CNS from the circulation or
n2:mentions
n3:9817441
Subject Item
_:vb515206
rdf:type
n2:Context
rdf:value
Essentially all components of the cellular immune response are detected in the infiltrate: natural killer (NK) cells, antigen-specific CD4+ and CD8+ T cells, B cells and monocyte/macrophages53,54,>>55<<,56,57. NK cells are detected first, followed by CD8+ and CD4+ T cells, then B cells and monocytes/macrophages58. In addition, dendritic cells are detected in the CNS during inflammation and either enter the CNS from the circulation or
n2:mentions
n3:11222733
Subject Item
_:vb515207
rdf:type
n2:Context
rdf:value
Essentially all components of the cellular immune response are detected in the infiltrate: natural killer (NK) cells, antigen-specific CD4+ and CD8+ T cells, B cells and monocyte/macrophages53,54,55,>>56<<,57. NK cells are detected first, followed by CD8+ and CD4+ T cells, then B cells and monocytes/macrophages58. In addition, dendritic cells are detected in the CNS during inflammation and either enter the CNS from the circulation or
n2:mentions
n3:9510193
Subject Item
_:vb515208
rdf:type
n2:Context
rdf:value
Essentially all components of the cellular immune response are detected in the infiltrate: natural killer (NK) cells, antigen-specific CD4+ and CD8+ T cells, B cells and monocyte/macrophages53,54,55,56,>>57<<. NK cells are detected first, followed by CD8+ and CD4+ T cells, then B cells and monocytes/macrophages58. In addition, dendritic cells are detected in the CNS during inflammation and either enter the CNS from the circulation or develop
n2:mentions
n3:11884463
Subject Item
_:vb515209
rdf:type
n2:Context
rdf:value
NK cells are detected first, followed by CD8+ and CD4+ T cells, then B cells and monocytes/macrophages>>58<<. In addition, dendritic cells are detected in the CNS during inflammation and either enter the CNS from the circulation or develop from a subpopulation of activated microglia33,59. The number of inflammatory cells peaks within one to two
n2:mentions
n3:11417137
Subject Item
_:vb515210
rdf:type
n2:Context
rdf:value
In addition, dendritic cells are detected in the CNS during inflammation and either enter the CNS from the circulation or develop from a subpopulation of activated microglia>>33<<,59. The number of inflammatory cells peaks within one to two weeks after infection and then these cells are gradually eliminated, unless the virus persists60. Cytokines that are produced by these infiltrating cells include IFN-γ, IL-4 and
n2:mentions
n3:11160337
Subject Item
_:vb515211
rdf:type
n2:Context
rdf:value
In addition, dendritic cells are detected in the CNS during inflammation and either enter the CNS from the circulation or develop from a subpopulation of activated microglia33,>>59<<. The number of inflammatory cells peaks within one to two weeks after infection and then these cells are gradually eliminated, unless the virus persists60. Cytokines that are produced by these infiltrating cells include IFN-γ, IL-4 and
n2:mentions
n3:11929128
Subject Item
_:vb515212
rdf:type
n2:Context
rdf:value
The number of inflammatory cells peaks within one to two weeks after infection and then these cells are gradually eliminated, unless the virus persists>>60<<. Cytokines that are produced by these infiltrating cells include IFN-γ, IL-4 and IL-10, but rarely IL-2 (Refs 27–29,54,61,62). B cells in the CNS usually produce antibody, mainly of the immunoglobulin A and IgG subclasses63, although IgA
n2:mentions
n3:10933698
Subject Item
_:vb515213
rdf:type
n2:Context
rdf:value
Cytokines that are produced by these infiltrating cells include IFN-γ, IL-4 and IL-10, but rarely IL-2 (Refs >>27<<–29,54,61,62).
n2:mentions
n3:11118358 n3:9217050 n3:8301132
Subject Item
_:vb515214
rdf:type
n2:Context
rdf:value
Cytokines that are produced by these infiltrating cells include IFN-γ, IL-4 and IL-10, but rarely IL-2 (Refs 27–29,>>54<<,61,62). B cells in the CNS usually produce antibody, mainly of the immunoglobulin A and IgG subclasses63, although IgA is not produced in response to all infections.
n2:mentions
n3:9817441
Subject Item
_:vb515215
rdf:type
n2:Context
rdf:value
Cytokines that are produced by these infiltrating cells include IFN-γ, IL-4 and IL-10, but rarely IL-2 (Refs 27–29,54,>>61<<,62). B cells in the CNS usually produce antibody, mainly of the immunoglobulin A and IgG subclasses63, although IgA is not produced in response to all infections.
n2:mentions
n3:8057431
Subject Item
_:vb515216
rdf:type
n2:Context
rdf:value
Cytokines that are produced by these infiltrating cells include IFN-γ, IL-4 and IL-10, but rarely IL-2 (Refs 27–29,54,61,>>62<<). B cells in the CNS usually produce antibody, mainly of the immunoglobulin A and IgG subclasses63, although IgA is not produced in response to all infections.
n2:mentions
n3:9973422
Subject Item
_:vb515217
rdf:type
n2:Context
rdf:value
B cells in the CNS usually produce antibody, mainly of the immunoglobulin A and IgG subclasses>>63<<, although IgA is not produced in response to all infections.
n2:mentions
n3:7451970
Subject Item
_:vb515218
rdf:type
n2:Context
rdf:value
The immunoregulatory activity of brain gangliosides on T cells is due, in part, to suppression of NF-κB activity and phosphorylation of retinoblastoma protein>>64<<,65,66. T cells from strains of mice that are susceptible to TH1-mediated autoimmune disease of the CNS are less responsive to regulation by brain gangliosides than T cells from disease-resistant strains67.
n2:mentions
n3:8906807
Subject Item
_:vb515219
rdf:type
n2:Context
rdf:value
The immunoregulatory activity of brain gangliosides on T cells is due, in part, to suppression of NF-κB activity and phosphorylation of retinoblastoma protein64,>>65<<,66. T cells from strains of mice that are susceptible to TH1-mediated autoimmune disease of the CNS are less responsive to regulation by brain gangliosides than T cells from disease-resistant strains67.
n2:mentions
n3:9036980
Subject Item
_:vb515220
rdf:type
n2:Context
rdf:value
The immunoregulatory activity of brain gangliosides on T cells is due, in part, to suppression of NF-κB activity and phosphorylation of retinoblastoma protein64,65,>>66<<. T cells from strains of mice that are susceptible to TH1-mediated autoimmune disease of the CNS are less responsive to regulation by brain gangliosides than T cells from disease-resistant strains67.
n2:mentions
n3:8376939
Subject Item
_:vb515221
rdf:type
n2:Context
rdf:value
T cells from strains of mice that are susceptible to TH1-mediated autoimmune disease of the CNS are less responsive to regulation by brain gangliosides than T cells from disease-resistant strains>>67<<.
n2:mentions
n3:9743332
Subject Item
_:vb515222
rdf:type
n2:Context
rdf:value
However, ganglioside production by neurons does not inhibit the expression of MHC class II molecules by microglia>>4<<,68. Antigen presentation by microglia stimulates CD4+ T cells to produce IFN-γ, but not IL-2 and does not stimulate T-cell proliferation.
n2:mentions
n3:9576961
Subject Item
_:vb515223
rdf:type
n2:Context
rdf:value
However, ganglioside production by neurons does not inhibit the expression of MHC class II molecules by microglia4,>>68<<. Antigen presentation by microglia stimulates CD4+ T cells to produce IFN-γ, but not IL-2 and does not stimulate T-cell proliferation.
n2:mentions
n3:8466862
Subject Item
_:vb515224
rdf:type
n2:Context
rdf:value
By contrast, macrophages that infiltrate the CNS from the periphery, which tend to accumulate near vessels, can support complete T-cell activation and survival>>69<<. Activity of these macrophages might be controlled by the production of TGF-β in the CNS, which suppresses the development of T-cell cytotoxicity and delayed-type hypersensitivity, and which deactivates infiltrating antigen-presenting
n2:mentions
n3:8920862
Subject Item
_:vb515225
rdf:type
n2:Context
rdf:value
of these macrophages might be controlled by the production of TGF-β in the CNS, which suppresses the development of T-cell cytotoxicity and delayed-type hypersensitivity, and which deactivates infiltrating antigen-presenting cells (APCs)>>70<<,71.
n2:mentions
n3:9688327
Subject Item
_:vb515226
rdf:type
n2:Context
rdf:value
these macrophages might be controlled by the production of TGF-β in the CNS, which suppresses the development of T-cell cytotoxicity and delayed-type hypersensitivity, and which deactivates infiltrating antigen-presenting cells (APCs)70,>>71<<.
n2:mentions
n3:9669697
Subject Item
_:vb515227
rdf:type
n2:Context
rdf:value
Activated T cells that leave the perivascular area often undergo apoptosis after exposure to APCs in the parenchyma>>72<<,73. In addition, constitutive expression of FAS ligand (CD95L) by cells of the CNS induces apoptosis of FAS (CD95)-expressing activated T cells.
n2:mentions
n3:9736022
Subject Item
_:vb515228
rdf:type
n2:Context
rdf:value
Activated T cells that leave the perivascular area often undergo apoptosis after exposure to APCs in the parenchyma72,>>73<<. In addition, constitutive expression of FAS ligand (CD95L) by cells of the CNS induces apoptosis of FAS (CD95)-expressing activated T cells.
n2:mentions
n3:8800954
Subject Item
_:vb515229
rdf:type
n2:Context
rdf:value
Neuronal expression of CD95L might further protect neurons from T-cell cytotoxicity>>74<<,75,76.
n2:mentions
n3:11441070
Subject Item
_:vb515230
rdf:type
n2:Context
rdf:value
Neuronal expression of CD95L might further protect neurons from T-cell cytotoxicity74,>>75<<,76.
n2:mentions
n3:10885654
Subject Item
_:vb515231
rdf:type
n2:Context
rdf:value
Neuronal expression of CD95L might further protect neurons from T-cell cytotoxicity74,75,>>76<<.
n2:mentions
n3:11880149
Subject Item
_:vb515232
rdf:type
n2:Context
rdf:value
If the clearance process is not complete, then mechanisms for preventing resumption of virus replication must be in place to avoid progressive or relapsing disease>>58<<,60,77.
n2:mentions
n3:11417137
Subject Item
_:vb515233
rdf:type
n2:Context
rdf:value
If the clearance process is not complete, then mechanisms for preventing resumption of virus replication must be in place to avoid progressive or relapsing disease58,>>60<<,77.
n2:mentions
n3:10933698
Subject Item
_:vb515234
rdf:type
n2:Context
rdf:value
If the clearance process is not complete, then mechanisms for preventing resumption of virus replication must be in place to avoid progressive or relapsing disease58,60,>>77<<.
n2:mentions
n3:9584136
Subject Item
_:vb515235
rdf:type
n2:Context
rdf:value
Anti-viral antibodies are important for the clearance of rabies virus, Sindbis virus, MHV and TMEV from neurons>>78<<,79,80,81. B cells, including virus-specific antibody-secreting cells, infiltrate the brain as a part of the inflammatory response and are retained in the CNS57,82,83(Fig. 1c). Many of the specific antibodies that are associated with
n2:mentions
n3:2539504
Subject Item
_:vb515236
rdf:type
n2:Context
rdf:value
Anti-viral antibodies are important for the clearance of rabies virus, Sindbis virus, MHV and TMEV from neurons78,>>79<<,80,81. B cells, including virus-specific antibody-secreting cells, infiltrate the brain as a part of the inflammatory response and are retained in the CNS57,82,83(Fig. 1c). Many of the specific antibodies that are associated with
n2:mentions
n3:2535900
Subject Item
_:vb515237
rdf:type
n2:Context
rdf:value
Anti-viral antibodies are important for the clearance of rabies virus, Sindbis virus, MHV and TMEV from neurons78,79,>>80<<,81. B cells, including virus-specific antibody-secreting cells, infiltrate the brain as a part of the inflammatory response and are retained in the CNS57,82,83(Fig. 1c). Many of the specific antibodies that are associated with protection
n2:mentions
n3:1658936
Subject Item
_:vb515238
rdf:type
n2:Context
rdf:value
Anti-viral antibodies are important for the clearance of rabies virus, Sindbis virus, MHV and TMEV from neurons78,79,80,>>81<<. B cells, including virus-specific antibody-secreting cells, infiltrate the brain as a part of the inflammatory response and are retained in the CNS57,82,83(Fig. 1c). Many of the specific antibodies that are associated with protection or
n2:mentions
n3:1674967
Subject Item
_:vb515239
rdf:type
n2:Context
rdf:value
B cells, including virus-specific antibody-secreting cells, infiltrate the brain as a part of the inflammatory response and are retained in the CNS>>57<<,82,83(Fig.
n2:mentions
n3:11884463
Subject Item
_:vb515240
rdf:type
n2:Context
rdf:value
B cells, including virus-specific antibody-secreting cells, infiltrate the brain as a part of the inflammatory response and are retained in the CNS57,>>82<<,83(Fig. 1c). Many of the specific antibodies that are associated with protection or clearance are directed against virus surface proteins and inhibit intracellular virus replication in vitro80,84. The mechanism by which antibody specific
n2:mentions
n3:1334109
Subject Item
_:vb515241
rdf:type
n2:Context
rdf:value
B cells, including virus-specific antibody-secreting cells, infiltrate the brain as a part of the inflammatory response and are retained in the CNS57,82,>>83<<(Fig. 1c). Many of the specific antibodies that are associated with protection or clearance are directed against virus surface proteins and inhibit intracellular virus replication in vitro80,84. The mechanism by which antibody specific for
n2:mentions
n3:9817440
Subject Item
_:vb515242
rdf:type
n2:Context
rdf:value
Many of the specific antibodies that are associated with protection or clearance are directed against virus surface proteins and inhibit intracellular virus replication in vitro>>80<<,84. The mechanism by which antibody specific for a virus surface glycoprotein inhibits virus replication is not completely clear, but has been studied in greatest detail for Sindbis virus.
n2:mentions
n3:1658936
Subject Item
_:vb515243
rdf:type
n2:Context
rdf:value
Many of the specific antibodies that are associated with protection or clearance are directed against virus surface proteins and inhibit intracellular virus replication in vitro80,>>84<<. The mechanism by which antibody specific for a virus surface glycoprotein inhibits virus replication is not completely clear, but has been studied in greatest detail for Sindbis virus.
n2:mentions
n3:7853547
Subject Item
_:vb515244
rdf:type
n2:Context
rdf:value
The anti-viral effect of antibody specific for the Sindbis virus E2 glycoprotein does not require complement or phagocytic cells, but does require that the antibody be bivalent>>80<<,84. This implies that crosslinking of virus proteins on the surface of the infected cell is necessary for the control of intracellular virus replication.
n2:mentions
n3:1658936
Subject Item
_:vb515245
rdf:type
n2:Context
rdf:value
The anti-viral effect of antibody specific for the Sindbis virus E2 glycoprotein does not require complement or phagocytic cells, but does require that the antibody be bivalent80,>>84<<. This implies that crosslinking of virus proteins on the surface of the infected cell is necessary for the control of intracellular virus replication.
n2:mentions
n3:7853547
Subject Item
_:vb515246
rdf:type
n2:Context
rdf:value
Treatment of persistently infected cells with antibody leads to the restoration of sodium-pump function and synthesis of host proteins, both of which are inhibited by Sindbis-virus replication>>85<<. Type 1 IFN synergizes with antibody to further inhibit virus replication, and it is probable that this interaction is important for the clearance of virus in vivo86. Antibody is also necessary for the clearance of rabies virus87 and MHV
n2:mentions
n3:7474120
Subject Item
_:vb515247
rdf:type
n2:Context
rdf:value
Type 1 IFN synergizes with antibody to further inhibit virus replication, and it is probable that this interaction is important for the clearance of virus in vivo>>86<<. Antibody is also necessary for the clearance of rabies virus87 and MHV from neurons79. The 'clearance' of Sindbis virus and other viruses that infect neurons is incomplete, as virus RNA can be detected throughout the life of the
n2:mentions
n3:7474167
Subject Item
_:vb515248
rdf:type
n2:Context
rdf:value
Antibody is also necessary for the clearance of rabies virus>>87<< and MHV from neurons79.
n2:mentions
n3:9557653
Subject Item
_:vb515249
rdf:type
n2:Context
rdf:value
Antibody is also necessary for the clearance of rabies virus87 and MHV from neurons>>79<<. The 'clearance' of Sindbis virus and other viruses that infect neurons is incomplete, as virus RNA can be detected throughout the life of the animal82,88. Long-term secretion of antibody in the brains of mice that have recovered from
n2:mentions
n3:2535900
Subject Item
_:vb515250
rdf:type
n2:Context
rdf:value
The 'clearance' of Sindbis virus and other viruses that infect neurons is incomplete, as virus RNA can be detected throughout the life of the animal>>82<<,88. Long-term secretion of antibody in the brains of mice that have recovered from infection indicates that there is a continued requirement for immune-mediated control of virus replication57,82.
n2:mentions
n3:1334109
Subject Item
_:vb515251
rdf:type
n2:Context
rdf:value
The 'clearance' of Sindbis virus and other viruses that infect neurons is incomplete, as virus RNA can be detected throughout the life of the animal82,>>88<<. Long-term secretion of antibody in the brains of mice that have recovered from infection indicates that there is a continued requirement for immune-mediated control of virus replication57,82.
n2:mentions
n3:8995689
Subject Item
_:vb515252
rdf:type
n2:Context
rdf:value
Long-term secretion of antibody in the brains of mice that have recovered from infection indicates that there is a continued requirement for immune-mediated control of virus replication>>57<<,82.
n2:mentions
n3:11884463
Subject Item
_:vb515253
rdf:type
n2:Context
rdf:value
Long-term secretion of antibody in the brains of mice that have recovered from infection indicates that there is a continued requirement for immune-mediated control of virus replication57,>>82<<.
n2:mentions
n3:1334109
Subject Item
_:vb515254
rdf:type
n2:Context
rdf:value
Although studies of immunodeficient mice show that T cells are not necessary for the clearance of Sindbis virus from neurons, the level of intracellular virus RNA is reduced more rapidly when CD8+ T cells are present>>46<<. CD8+ T cells are also important for rapid control of rabies virus and herpes simplex virus replication in neurons87,89.
n2:mentions
n3:10846095
Subject Item
_:vb515255
rdf:type
n2:Context
rdf:value
CD8+ T cells are also important for rapid control of rabies virus and herpes simplex virus replication in neurons>>87<<,89. In general, CD8+ T cells can exert anti-viral effector functions either through a non-lytic cytokine-mediated pathway or by a cytotoxic pathway, which involves either the cytotoxic effector molecules perforin and granzymes or
n2:mentions
n3:9557653
Subject Item
_:vb515256
rdf:type
n2:Context
rdf:value
CD8+ T cells are also important for rapid control of rabies virus and herpes simplex virus replication in neurons87,>>89<<. In general, CD8+ T cells can exert anti-viral effector functions either through a non-lytic cytokine-mediated pathway or by a cytotoxic pathway, which involves either the cytotoxic effector molecules perforin and granzymes or CD95–CD95L.
n2:mentions
n3:11602757
Subject Item
_:vb515257
rdf:type
n2:Context
rdf:value
Neurons are severely restricted in their expression of MHC class I molecules, but MHC class I expression has been observed in damaged neurons that were exposed to IFN-γ and occasionally on virus-infected neurons>>51<<,90. CD8+ T-cell-mediated killing of neurons through a CD95–CD95L-dependent pathway has also been documented74,91. Although neuronal death mediated by CD8+ T cells has been reported in some virus infections of the CNS92, the main
n2:mentions
n3:10400743
Subject Item
_:vb515258
rdf:type
n2:Context
rdf:value
Neurons are severely restricted in their expression of MHC class I molecules, but MHC class I expression has been observed in damaged neurons that were exposed to IFN-γ and occasionally on virus-infected neurons51,>>90<<. CD8+ T-cell-mediated killing of neurons through a CD95–CD95L-dependent pathway has also been documented74,91. Although neuronal death mediated by CD8+ T cells has been reported in some virus infections of the CNS92, the main
n2:mentions
n3:7624779
Subject Item
_:vb515259
rdf:type
n2:Context
rdf:value
CD8+ T-cell-mediated killing of neurons through a CD95–CD95L-dependent pathway has also been documented>>74<<,91. Although neuronal death mediated by CD8+ T cells has been reported in some virus infections of the CNS92, the main contributions of CD8+ T cells to the clearance of virus from neurons are not mediated through cytolysis of infected
n2:mentions
n3:11441070
Subject Item
_:vb515260
rdf:type
n2:Context
rdf:value
CD8+ T-cell-mediated killing of neurons through a CD95–CD95L-dependent pathway has also been documented74,>>91<<. Although neuronal death mediated by CD8+ T cells has been reported in some virus infections of the CNS92, the main contributions of CD8+ T cells to the clearance of virus from neurons are not mediated through cytolysis of infected cells,
n2:mentions
n3:11169405
Subject Item
_:vb515261
rdf:type
n2:Context
rdf:value
Although neuronal death mediated by CD8+ T cells has been reported in some virus infections of the CNS>>92<<, the main contributions of CD8+ T cells to the clearance of virus from neurons are not mediated through cytolysis of infected cells, and do not require perforin or CD95 (Ref.
n2:mentions
n3:7912707
Subject Item
_:vb515262
rdf:type
n2:Context
rdf:value
cells has been reported in some virus infections of the CNS92, the main contributions of CD8+ T cells to the clearance of virus from neurons are not mediated through cytolysis of infected cells, and do not require perforin or CD95 (Ref. >>93<<).
n2:mentions
n3:11932415
Subject Item
_:vb515263
rdf:type
n2:Context
rdf:value
Recent studies of antibody-deficient mice have shown that both CD4+ and CD8+ T cells infiltrating the brain contribute to the clearance of Sindbis virus through the production of IFN-γ>>94<<. An important role for CD4+ and CD8+ T cells in virus clearance that is associated with the production of IFN-γ has also been shown for a neuronotropic strain of MHV61.
n2:mentions
n3:11452126
Subject Item
_:vb515264
rdf:type
n2:Context
rdf:value
An important role for CD4+ and CD8+ T cells in virus clearance that is associated with the production of IFN-γ has also been shown for a neuronotropic strain of MHV>>61<<. IFN-γ also has a role in the clearance of vesicular stomatitis virus, a virus that is related to rabies virus, from neurons, perhaps through the production of nitric oxide95. CD8+ T cells alone can clear LCMV from persistently infected
n2:mentions
n3:8057431
Subject Item
_:vb515265
rdf:type
n2:Context
rdf:value
IFN-γ also has a role in the clearance of vesicular stomatitis virus, a virus that is related to rabies virus, from neurons, perhaps through the production of nitric oxide>>95<<. CD8+ T cells alone can clear LCMV from persistently infected neurons after a prolonged period of time96 and these cells might produce anti-viral factors in addition to IFN-γ. For example, the cytotoxic effector molecule granzyme A seems
n2:mentions
n3:10366576
Subject Item
_:vb515266
rdf:type
n2:Context
rdf:value
CD8+ T cells alone can clear LCMV from persistently infected neurons after a prolonged period of time>>96<< and these cells might produce anti-viral factors in addition to IFN-γ.
n2:mentions
n3:3086743
Subject Item
_:vb515267
rdf:type
n2:Context
rdf:value
For example, the cytotoxic effector molecule granzyme A seems to have an additional role in restricting the replication of herpes simplex virus in neurons>>97<<. However, in contrast to its essential role in non-cytolytic clearance of hepatitis B virus from hepatocytes, there is no evidence of a role for TNF in clearing virus from neurons94,98.
n2:mentions
n3:10623769
Subject Item
_:vb515268
rdf:type
n2:Context
rdf:value
However, in contrast to its essential role in non-cytolytic clearance of hepatitis B virus from hepatocytes, there is no evidence of a role for TNF in clearing virus from neurons>>94<<,98.
n2:mentions
n3:11452126
Subject Item
_:vb515269
rdf:type
n2:Context
rdf:value
However, in contrast to its essential role in non-cytolytic clearance of hepatitis B virus from hepatocytes, there is no evidence of a role for TNF in clearing virus from neurons94,>>98<<.
n2:mentions
n3:9782311
Subject Item
_:vb515270
rdf:type
n2:Context
rdf:value
Motor neurons in the spinal cord show complete resolution of infection in the absence of antibody, whereas cortical neurons require antibody for the clearance of Sindbis virus>>94<<. Whether this is due to differences in the ability to bind IFN-γ or to differences in the ability to respond to IFN-γ is not known. The IFN-γ receptor is widely expressed by neurons and constitutive expression by neurons is markedly
n2:mentions
n3:11452126
Subject Item
_:vb515271
rdf:type
n2:Context
rdf:value
The IFN-γ receptor is widely expressed by neurons and constitutive expression by neurons is markedly higher than it is by glial cells>>99<<. It is interesting that some neurons can produce IFN-γ19, and the autocrine effects of IFN-γ might also have a role in virus clearance100. In all of these examples of virus clearance, no infectious virus can be recovered, but virus RNA in
n2:mentions
n3:10779704
Subject Item
_:vb515272
rdf:type
n2:Context
rdf:value
It is interesting that some neurons can produce IFN-γ>>19<<, and the autocrine effects of IFN-γ might also have a role in virus clearance100.
n2:mentions
n3:9396771
Subject Item
_:vb515273
rdf:type
n2:Context
rdf:value
It is interesting that some neurons can produce IFN-γ19, and the autocrine effects of IFN-γ might also have a role in virus clearance>>100<<. In all of these examples of virus clearance, no infectious virus can be recovered, but virus RNA in the CNS remains detectable by sensitive techniques — such as PCR after reverse transcription of RNA (RT-PCR) — and this RNA is presumed
n2:mentions
n3:1322463
Subject Item
_:vb515274
rdf:type
n2:Context
rdf:value
RNA in the CNS remains detectable by sensitive techniques — such as PCR after reverse transcription of RNA (RT-PCR) — and this RNA is presumed to reside in infected neurons that survived both infection and immune-mediated virus clearance>>77<<,82.
n2:mentions
n3:9584136
Subject Item
_:vb515275
rdf:type
n2:Context
rdf:value
in the CNS remains detectable by sensitive techniques — such as PCR after reverse transcription of RNA (RT-PCR) — and this RNA is presumed to reside in infected neurons that survived both infection and immune-mediated virus clearance77,>>82<<.
n2:mentions
n3:1334109
Subject Item
_:vb515276
rdf:type
n2:Context
rdf:value
Clearance of TMEV from strains of mice that do not develop persistent infection or demyelinating disease is dependent on a rapid CD8+ T-cell response>>101<<. In strains of mice that are susceptible to persistent infection, infectious virus is cleared from neurons, but virus is not completely cleared from oligodendrocytes, astrocytes and microglia, although it is reduced to low levels, and
n2:mentions
n3:2495323
Subject Item
_:vb515277
rdf:type
n2:Context
rdf:value
antibody can protect mice from fatal encephalomyelitis, in strains of mice that clear the infection, antibody is not detectable until after the virus is cleared, which indicates that antibody is not the main mechanism of clearance>>28<<,102,103. A reduction in virus replication can be mediated by CD4+ or CD8+ T cells104,105, and transfer of virus-specific CD4+ or CD8+ T cells can protect from fatal MHV-induced encephalitis104, which further indicates a role for T cells
n2:mentions
n3:9217050
Subject Item
_:vb515278
rdf:type
n2:Context
rdf:value
antibody can protect mice from fatal encephalomyelitis, in strains of mice that clear the infection, antibody is not detectable until after the virus is cleared, which indicates that antibody is not the main mechanism of clearance28,>>102<<,103. A reduction in virus replication can be mediated by CD4+ or CD8+ T cells104,105, and transfer of virus-specific CD4+ or CD8+ T cells can protect from fatal MHV-induced encephalitis104, which further indicates a role for T cells in
n2:mentions
n3:1655890
Subject Item
_:vb515279
rdf:type
n2:Context
rdf:value
can protect mice from fatal encephalomyelitis, in strains of mice that clear the infection, antibody is not detectable until after the virus is cleared, which indicates that antibody is not the main mechanism of clearance28,102,>>103<<. A reduction in virus replication can be mediated by CD4+ or CD8+ T cells104,105, and transfer of virus-specific CD4+ or CD8+ T cells can protect from fatal MHV-induced encephalitis104, which further indicates a role for T cells in
n2:mentions
n3:8985361
Subject Item
_:vb515280
rdf:type
n2:Context
rdf:value
A reduction in virus replication can be mediated by CD4+ or CD8+ T cells>>104<<,105, and transfer of virus-specific CD4+ or CD8+ T cells can protect from fatal MHV-induced encephalitis104, which further indicates a role for T cells in control of virus replication.
n2:mentions
n3:1705943
Subject Item
_:vb515281
rdf:type
n2:Context
rdf:value
A reduction in virus replication can be mediated by CD4+ or CD8+ T cells104,>>105<<, and transfer of virus-specific CD4+ or CD8+ T cells can protect from fatal MHV-induced encephalitis104, which further indicates a role for T cells in control of virus replication.
n2:mentions
n3:7815531
Subject Item
_:vb515282
rdf:type
n2:Context
rdf:value
A reduction in virus replication can be mediated by CD4+ or CD8+ T cells104,105, and transfer of virus-specific CD4+ or CD8+ T cells can protect from fatal MHV-induced encephalitis>>104<<, which further indicates a role for T cells in control of virus replication.
n2:mentions
n3:1705943
Subject Item
_:vb515283
rdf:type
n2:Context
rdf:value
The mechanism of T-cell-mediated clearance of MHV is glial-cell-type specific>>102<<. IFN-γ is particularly important for the clearance of MHV from oligodendrocytes, whereas perforin, and presumably CD8+ T-cell-mediated cytolysis, is important for the clearance of virus from astrocytes and microglia103,105,106.
n2:mentions
n3:1655890
Subject Item
_:vb515284
rdf:type
n2:Context
rdf:value
IFN-γ is particularly important for the clearance of MHV from oligodendrocytes, whereas perforin, and presumably CD8+ T-cell-mediated cytolysis, is important for the clearance of virus from astrocytes and microglia>>103<<,105,106. Clearance occurs in perforin-deficient mice but is delayed, indicating that many mechanisms are involved103. Auxiliary roles have been identified for CD4+ T cells, CD95 and IFN-γ in virus clearance98,107. CD4+ T-cell help is
n2:mentions
n3:8985361
Subject Item
_:vb515285
rdf:type
n2:Context
rdf:value
IFN-γ is particularly important for the clearance of MHV from oligodendrocytes, whereas perforin, and presumably CD8+ T-cell-mediated cytolysis, is important for the clearance of virus from astrocytes and microglia103,>>105<<,106. Clearance occurs in perforin-deficient mice but is delayed, indicating that many mechanisms are involved103. Auxiliary roles have been identified for CD4+ T cells, CD95 and IFN-γ in virus clearance98,107. CD4+ T-cell help is required
n2:mentions
n3:7815531
Subject Item
_:vb515286
rdf:type
n2:Context
rdf:value
IFN-γ is particularly important for the clearance of MHV from oligodendrocytes, whereas perforin, and presumably CD8+ T-cell-mediated cytolysis, is important for the clearance of virus from astrocytes and microglia103,105,>>106<<. Clearance occurs in perforin-deficient mice but is delayed, indicating that many mechanisms are involved103. Auxiliary roles have been identified for CD4+ T cells, CD95 and IFN-γ in virus clearance98,107. CD4+ T-cell help is required for
n2:mentions
n3:9973424
Subject Item
_:vb515287
rdf:type
n2:Context
rdf:value
Clearance occurs in perforin-deficient mice but is delayed, indicating that many mechanisms are involved>>103<<. Auxiliary roles have been identified for CD4+ T cells, CD95 and IFN-γ in virus clearance98,107. CD4+ T-cell help is required for the prevention of apoptosis and expression of the cytotoxic effector function of CD8+ T cells in the
n2:mentions
n3:8985361
Subject Item
_:vb515288
rdf:type
n2:Context
rdf:value
Auxiliary roles have been identified for CD4+ T cells, CD95 and IFN-γ in virus clearance>>98<<,107. CD4+ T-cell help is required for the prevention of apoptosis and expression of the cytotoxic effector function of CD8+ T cells in the MHV-infected CNS56. Virus RNA that persists can resume replication and, therefore, long-term local
n2:mentions
n3:9782311
Subject Item
_:vb515289
rdf:type
n2:Context
rdf:value
Auxiliary roles have been identified for CD4+ T cells, CD95 and IFN-γ in virus clearance98,>>107<<. CD4+ T-cell help is required for the prevention of apoptosis and expression of the cytotoxic effector function of CD8+ T cells in the MHV-infected CNS56. Virus RNA that persists can resume replication and, therefore, long-term local
n2:mentions
n3:10666278
Subject Item
_:vb515290
rdf:type
n2:Context
rdf:value
CD4+ T-cell help is required for the prevention of apoptosis and expression of the cytotoxic effector function of CD8+ T cells in the MHV-infected CNS>>56<<. Virus RNA that persists can resume replication and, therefore, long-term local immune control of virus replication is required60. IFN-γ-secreting CD8+ T cells that are specific for virus antigen persist in the CNS and might contribute to
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Virus RNA that persists can resume replication and, therefore, long-term local immune control of virus replication is required>>60<<. IFN-γ-secreting CD8+ T cells that are specific for virus antigen persist in the CNS and might contribute to the demyelinating process that follows resolution of the acute virus infection108,109,110. CD4+ T cells also persist in the CNS
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IFN-γ-secreting CD8+ T cells that are specific for virus antigen persist in the CNS and might contribute to the demyelinating process that follows resolution of the acute virus infection>>108<<,109,110. CD4+ T cells also persist in the CNS for several weeks60. Despite a lack of evidence for a role for antibody in the initial clearance of MHV, antibody-secreting B cells are essential for the prevention of renewed virus
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IFN-γ-secreting CD8+ T cells that are specific for virus antigen persist in the CNS and might contribute to the demyelinating process that follows resolution of the acute virus infection108,>>109<<,110. CD4+ T cells also persist in the CNS for several weeks60. Despite a lack of evidence for a role for antibody in the initial clearance of MHV, antibody-secreting B cells are essential for the prevention of renewed virus replication111.
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IFN-γ-secreting CD8+ T cells that are specific for virus antigen persist in the CNS and might contribute to the demyelinating process that follows resolution of the acute virus infection108,109,>>110<<. CD4+ T cells also persist in the CNS for several weeks60. Despite a lack of evidence for a role for antibody in the initial clearance of MHV, antibody-secreting B cells are essential for the prevention of renewed virus replication111.
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CD4+ T cells also persist in the CNS for several weeks>>60<<. Despite a lack of evidence for a role for antibody in the initial clearance of MHV, antibody-secreting B cells are essential for the prevention of renewed virus replication111.
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Despite a lack of evidence for a role for antibody in the initial clearance of MHV, antibody-secreting B cells are essential for the prevention of renewed virus replication>>111<<. T-cell numbers eventually decline, but antibody-secreting B cells persist57. Both B cells and the virus-specific antibody they produce seem to have independent roles in preventing the renewed replication of MHV in astrocytes and
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T-cell numbers eventually decline, but antibody-secreting B cells persist>>57<<. Both B cells and the virus-specific antibody they produce seem to have independent roles in preventing the renewed replication of MHV in astrocytes and oligodendrocytes111,112.
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Both B cells and the virus-specific antibody they produce seem to have independent roles in preventing the renewed replication of MHV in astrocytes and oligodendrocytes>>111<<,112. Although B cells can be cytotoxic in vitro, it is not clear whether this mechanism is relevant to the role of B cells in vivo.
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Both B cells and the virus-specific antibody they produce seem to have independent roles in preventing the renewed replication of MHV in astrocytes and oligodendrocytes111,>>112<<. Although B cells can be cytotoxic in vitro, it is not clear whether this mechanism is relevant to the role of B cells in vivo.
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