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PMC0
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10.1046%2Fj.1365-2567.2000.00142.x
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introduction
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It has antimicrobial actions against various pathogens via its cytotoxic or cytostatic effects.>>1<<–5 Potent host defence against intruding microbes is also mediated by oxygen radicals and active oxygen species, including superoxide anion radical (O2–), hydrogen peroxide (H2O2), and hypochlorite anion (OCl–), produced from phagocytic
n2:mentions
n3:1711326 n3:8531884 n3:8386705 n3:9199469 n3:3280600
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also mediated by oxygen radicals and active oxygen species, including superoxide anion radical (O2–), hydrogen peroxide (H2O2), and hypochlorite anion (OCl–), produced from phagocytic cells such as neutrophils and activated macrophages.>>6<< It is now well accepted that the chemical and biological reactivities of NO produced in environments such as inflamed tissues are greatly affected by concomitantly formed oxygen radicals, particularly O2–, through formation of reactive
n2:mentions
n3:6250449
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biological reactivities of NO produced in environments such as inflamed tissues are greatly affected by concomitantly formed oxygen radicals, particularly O2–, through formation of reactive nitrogen oxides such as peroxynitrite (ONOO–).>>7<<–12 Although the importance of these reactive nitrogen and oxygen intermediate species has been documented for host defence reactions against bacteria and fungi,1–5 their role in the pathogenesis of virus infections is only partly
n2:mentions
n3:10906338 n3:10906340 n3:8944624 n3:8828915 n3:10617463 n3:2154753
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7–12 Although the importance of these reactive nitrogen and oxygen intermediate species has been documented for host defence reactions against bacteria and fungi,>>1<<–5 their role in the pathogenesis of virus infections is only partly understood.
n2:mentions
n3:1711326 n3:8531884 n3:8386705 n3:9199469 n3:3280600
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about a particular microbe. It is thus critical to evaluate the pathogenesis of virus infection as related to the emerging concept of free radicals that are generated as host-derived factors during interactions between viruses and hosts.>>13<<–17 In this review, the biological relevance of NO production is discussed in view of oxidative stress and immunomodulation of the host's responses caused by NO during virus infections.
n2:mentions
n3:9421208 n3:1656471
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induction of no biosynthesis and oxygen radicals in virus infection
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NO, mainly caused by inducible NO synthase (iNOS), which is usually expressed by inflammatory phagocytic cells and other types of cells (e.g. epithelial and neuronal cells), has a defence function against bacteria, fungi, and parasites.>>1<<–5,18 iNOS produces a much larger amount of NO for a longer time (i.e. 10–100 times more) than do the other two constitutive enzymes, neuronal NOS and endothelial NOS.
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mainly caused by inducible NO synthase (iNOS), which is usually expressed by inflammatory phagocytic cells and other types of cells (e.g. epithelial and neuronal cells), has a defence function against bacteria, fungi, and parasites.1–5,>>18<< iNOS produces a much larger amount of NO for a longer time (i.e. 10–100 times more) than do the other two constitutive enzymes, neuronal NOS and endothelial NOS.
n2:mentions
n3:9366554
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has a defence function against bacteria, fungi, and parasites.1–5,18 iNOS produces a much larger amount of NO for a longer time (i.e. 10–100 times more) than do the other two constitutive enzymes, neuronal NOS and endothelial NOS.>>19<<,20 Although NO seems to have a limited bactericidal effect,21–23 suppression or lack of NO production results in impaired clearance of some types of bacteria by the host.
n2:mentions
n3:7504210
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has a defence function against bacteria, fungi, and parasites.1–5,18 iNOS produces a much larger amount of NO for a longer time (i.e. 10–100 times more) than do the other two constitutive enzymes, neuronal NOS and endothelial NOS.19,>>20<< Although NO seems to have a limited bactericidal effect,21–23 suppression or lack of NO production results in impaired clearance of some types of bacteria by the host.
n2:mentions
n3:1373932
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19,20 Although NO seems to have a limited bactericidal effect,>>21<<–23 suppression or lack of NO production results in impaired clearance of some types of bacteria by the host.
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19,20 Although NO seems to have a limited bactericidal effect,21–23 suppression or lack of NO production results in impaired clearance of some types of bacteria by the host.>>5<<
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n3:9199469
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19,20 Although NO seems to have a limited bactericidal effect,21–23 suppression or lack of NO production results in impaired clearance of some types of bacteria by the host.5,>>18<<
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19,20 Although NO seems to have a limited bactericidal effect,21–23 suppression or lack of NO production results in impaired clearance of some types of bacteria by the host.5,18,>>24<<
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in rats and mice, including those with neuroviruses, such as Borna disease virus, herpes simplex virus type 1 (HSV-1), rabies virus, and pneumotropic and cardiotropic viruses, such as influenza virus, Sendai virus, and coxsackievirus.>>15<<,16,29–35 iNOS expression is also observed in human diseases caused by human immunodeficiency virus-1 (HIV-1) and hepatitis B virus (HBV).
n2:mentions
n3:9421208
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rats and mice, including those with neuroviruses, such as Borna disease virus, herpes simplex virus type 1 (HSV-1), rabies virus, and pneumotropic and cardiotropic viruses, such as influenza virus, Sendai virus, and coxsackievirus.15,16,>>29<<–35 iNOS expression is also observed in human diseases caused by human immunodeficiency virus-1 (HIV-1) and hepatitis B virus (HBV).
n2:mentions
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pneumotropic and cardiotropic viruses, such as influenza virus, Sendai virus, and coxsackievirus.15,16,29–35 iNOS expression is also observed in human diseases caused by human immunodeficiency virus-1 (HIV-1) and hepatitis B virus (HBV).>>36<<,37 It seems therefore that iNOS is ubiquitously expressed during host responses to viral replication in vivo (Fig.
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n3:7530762
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and cardiotropic viruses, such as influenza virus, Sendai virus, and coxsackievirus.15,16,29–35 iNOS expression is also observed in human diseases caused by human immunodeficiency virus-1 (HIV-1) and hepatitis B virus (HBV).36,>>37<< It seems therefore that iNOS is ubiquitously expressed during host responses to viral replication in vivo (Fig.
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n3:9525976
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Wild-type mice (C57BL/6, B6), iNOS heterozygotes (iNOS+/–), and mice deficient in iNOS (iNOS–/–) were inoculated with 2 × LD50 of influenza virus, and ESR was performed as described previously.>>34<<
n2:mentions
n3:8637894
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epithelial cells in lung tissues infected with influenza virus in mice; the high output of NO has been clearly identified and quantified by electron spin resonance (ESR) spin trapping with the use of a dithiocarbamate-iron complex.16,>>34<< NO-dithiocarbamate-iron adducts with a triplet hyperfine structure of g perpendicular 2·04 are generated (Fig.
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n3:8637894
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by pharmacological NOS inhibition with Nω-monomethyl-l-arginine (L-NMMA) or by genetic disruption of iNOS, indicating that excessive production of NO is because of localized iNOS expression in the area of the virus infection.16,>>34<< The time profile of iNOS induction in the lung correlates well with that of the pathological manifestations of the viral pneumonia, i.e. pulmonary consolidation characterized by extensive infiltration of macrophages and lymphocytes,
n2:mentions
n3:8637894
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Many pathological effects of NO are thought to be produced via its interaction with oxygen radicals, particularly O2–, producing ONOO–.>>7<<–12 In this context, we analysed O2– generation in virus-infected lung with a focus on xanthine oxidase (XO) as a potent generator of O2– in the influenza model.
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its interaction with oxygen radicals, particularly O2–, producing ONOO–.7–12 In this context, we analysed O2– generation in virus-infected lung with a focus on xanthine oxidase (XO) as a potent generator of O2– in the influenza model.>>38<< O2– production from XO in bronchoalveolar lavage fluid increases in a time-dependent manner with a time profile almost parallel to that of iNOS induction.
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rdf:type
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rdf:value
38 O2– production from XO in bronchoalveolar lavage fluid increases in a time-dependent manner with a time profile almost parallel to that of iNOS induction.>>15<<,16,34,38 To confirm the generation of NO and O2–, and their coupling reaction, in the local area of virus infection, ESR was used to measure NO-haemoglobin, which is formed de novo from the reaction of endogenous haemoglobin with NO
n2:mentions
n3:9421208
Subject Item
_:vb211486
rdf:type
n2:Context
rdf:value
38 O2– production from XO in bronchoalveolar lavage fluid increases in a time-dependent manner with a time profile almost parallel to that of iNOS induction.15,16,>>34<<,38 To confirm the generation of NO and O2–, and their coupling reaction, in the local area of virus infection, ESR was used to measure NO-haemoglobin, which is formed de novo from the reaction of endogenous haemoglobin with NO generated
n2:mentions
n3:8637894
Subject Item
_:vb211487
rdf:type
n2:Context
rdf:value
38 O2– production from XO in bronchoalveolar lavage fluid increases in a time-dependent manner with a time profile almost parallel to that of iNOS induction.15,16,34,>>38<< To confirm the generation of NO and O2–, and their coupling reaction, in the local area of virus infection, ESR was used to measure NO-haemoglobin, which is formed de novo from the reaction of endogenous haemoglobin with NO generated in
n2:mentions
n3:2155924
Subject Item
_:vb211488
rdf:type
n2:Context
rdf:value
in the NO-haemoglobin ESR signal in the lung is obtained by SOD treatment of virus-infected animals, possibly because an appreciable amount of NO is rescued from the quenching reaction with O2– because of elimination of O2– by SOD.>>34<< These results provide substantial evidence for ONOO– formation in virus-infected tissues.
n2:mentions
n3:8637894
Subject Item
_:vb211489
rdf:type
n5:Section
dc:title
mechanisms of inos induction and antiviral effects of no
n5:contains
_:vb211520 _:vb211521 _:vb211522 _:vb211523 _:vb211490 _:vb211491 _:vb211492 _:vb211493 _:vb211494 _:vb211495 _:vb211496 _:vb211497 _:vb211498 _:vb211499 _:vb211500 _:vb211501 _:vb211502 _:vb211503 _:vb211504 _:vb211505 _:vb211506 _:vb211507 _:vb211508 _:vb211509 _:vb211510 _:vb211511 _:vb211512 _:vb211513 _:vb211514 _:vb211515 _:vb211516 _:vb211517 _:vb211518 _:vb211519
Subject Item
_:vb211490
rdf:type
n2:Context
rdf:value
observation). Furthermore, the iNOS-inducing potential in bronchoalveolar lavage fluid in influenza virus pneumonia is attributable solely to IFN-γ, as revealed by an immunoadsorption study using a specific anti-IFN-γ antibody.>>34<< These results strongly support the suggestion that IFN-γ is a major cytokine inducing iNOS and NO overproduction in pathogenesis of virus infection.
n2:mentions
n3:8637894
Subject Item
_:vb211491
rdf:type
n2:Context
rdf:value
34 These results strongly support the suggestion that IFN-γ is a major cytokine inducing iNOS and NO overproduction in pathogenesis of virus infection.>>15<<
n2:mentions
n3:9421208
Subject Item
_:vb211492
rdf:type
n2:Context
rdf:value
34 These results strongly support the suggestion that IFN-γ is a major cytokine inducing iNOS and NO overproduction in pathogenesis of virus infection.15,16,>>31<<
n2:mentions
n3:7690156
Subject Item
_:vb211493
rdf:type
n2:Context
rdf:value
Many previous reports indicate that type 1 helper T cell (Th1) responses are important for viral clearance.>>39<<–41 However, IFN-γ, a Th1-dependent cytokine, seems to be inefficient in host defence against various viral pathogens including influenza virus, Sendai virus, and vaccinia virus.
n2:mentions
n3:7699320 n3:1919440 n3:2564415
Subject Item
_:vb211494
rdf:type
n2:Context
rdf:value
T cell (Th1) responses are important for viral clearance.39–41 However, IFN-γ, a Th1-dependent cytokine, seems to be inefficient in host defence against various viral pathogens including influenza virus, Sendai virus, and vaccinia virus.>>42<<–44 In addition, lack of an iNOS-dependent antiviral effect is also noted for the same virus infections, which was recently confirmed by a number of studies using iNOS-deficient (iNOS–/–) mice.
n2:mentions
n3:10605032 n3:8228818 n3:9032321
Subject Item
_:vb211495
rdf:type
n2:Context
rdf:value
42–44 In addition, lack of an iNOS-dependent antiviral effect is also noted for the same virus infections, which was recently confirmed by a number of studies using iNOS-deficient (iNOS–/–) mice.>>44<<
n2:mentions
n3:10605032 n3:10906226 n3:9782132 n3:10877838 n3:10580062
Subject Item
_:vb211496
rdf:type
n2:Context
rdf:value
Downregulation of iNOS expression is also reported for some cytokines, e.g. interleukin (IL)-4, IL-10, and transforming growth factor-β.>>49<<–51 In addition, these suppressor cytokines may reduce NO production indirectly via induction of arginase,52–54 which diminishes the supply of the substrate (l-arginine) for iNOS.
n2:mentions
n3:1371674 n3:7507968 n3:7688028
Subject Item
_:vb211497
rdf:type
n2:Context
rdf:value
49–51 In addition, these suppressor cytokines may reduce NO production indirectly via induction of arginase,>>52<<–54 which diminishes the supply of the substrate (l-arginine) for iNOS.
n2:mentions
n3:8898077 n3:7530004 n3:9013624
Subject Item
_:vb211498
rdf:type
n2:Context
rdf:value
In some viral diseases, viral replication or viral components directly induce iNOS without mediation by pro-inflammatory cytokines (Fig. 1a). iNOS expression in HIV-1 encephalitis is of particular interest in this regard.>>36<< An envelope glycoprotein of HIV, gp41, triggers iNOS expression in human astrocytes and murine cortical brain cells in culture.
n2:mentions
n3:7530762
Subject Item
_:vb211499
rdf:type
n2:Context
rdf:value
cytokines (Fig. 1a). iNOS expression in HIV-1 encephalitis is of particular interest in this regard.36 An envelope glycoprotein of HIV, gp41, triggers iNOS expression in human astrocytes and murine cortical brain cells in culture.>>55<<,56 Thus, NO produced by iNOS may contribute directly to the pathogenesis of HIV-associated dementia and cardiomyopathy as well.
n2:mentions
n3:9870939
Subject Item
_:vb211500
rdf:type
n2:Context
rdf:value
of HIV, gp41, triggers iNOS expression in human astrocytes and murine cortical brain cells in culture.55,56 Thus, NO produced by iNOS may contribute directly to the pathogenesis of HIV-associated dementia and cardiomyopathy as well.>>36<<,57,58 Similarly, the human paramyxovirus respiratory syncytial virus directly upregulates iNOS in human type 2 alveolar epithelial cells (A549 cells) through a pathway independent of pro-inflammatory cytokines.
n2:mentions
n3:7530762
Subject Item
_:vb211501
rdf:type
n2:Context
rdf:value
of HIV, gp41, triggers iNOS expression in human astrocytes and murine cortical brain cells in culture.55,56 Thus, NO produced by iNOS may contribute directly to the pathogenesis of HIV-associated dementia and cardiomyopathy as well.36,>>57<<,58 Similarly, the human paramyxovirus respiratory syncytial virus directly upregulates iNOS in human type 2 alveolar epithelial cells (A549 cells) through a pathway independent of pro-inflammatory cytokines.
n2:mentions
n3:10443886
Subject Item
_:vb211502
rdf:type
n2:Context
rdf:value
HIV, gp41, triggers iNOS expression in human astrocytes and murine cortical brain cells in culture.55,56 Thus, NO produced by iNOS may contribute directly to the pathogenesis of HIV-associated dementia and cardiomyopathy as well.36,57,>>58<< Similarly, the human paramyxovirus respiratory syncytial virus directly upregulates iNOS in human type 2 alveolar epithelial cells (A549 cells) through a pathway independent of pro-inflammatory cytokines.
n2:mentions
n3:10468523
Subject Item
_:vb211503
rdf:type
n2:Context
rdf:value
cardiomyopathy as well.36,57,58 Similarly, the human paramyxovirus respiratory syncytial virus directly upregulates iNOS in human type 2 alveolar epithelial cells (A549 cells) through a pathway independent of pro-inflammatory cytokines.>>59<< There are therefore two pathways for iNOS induction in virus infections: cytokine-dependent mechanisms, and direct upregulation by virus.
n2:mentions
n3:10410996
Subject Item
_:vb211504
rdf:type
n2:Context
rdf:value
NO has antimicrobial activity against bacteria, parasites, and fungi.>>1<<–5 Antiviral effects of NO are also known for some types of virus, most typically DNA viruses such as murine poxvirus (ectromelia virus, EV) and herpes viruses, including HSV and Epstein-Barr virus (EBV), and some RNA viruses such as
n2:mentions
n3:3280600 n3:1711326 n3:8531884 n3:8386705 n3:9199469
Subject Item
_:vb211505
rdf:type
n2:Context
rdf:value
effects of NO are also known for some types of virus, most typically DNA viruses such as murine poxvirus (ectromelia virus, EV) and herpes viruses, including HSV and Epstein-Barr virus (EBV), and some RNA viruses such as coxsackievirus.>>31<<,60–63 EBV reactivation appears to be inhibited by NO via suppression of an immediate early-transactivator gene.
n2:mentions
n3:7690156
Subject Item
_:vb211506
rdf:type
n2:Context
rdf:value
of NO are also known for some types of virus, most typically DNA viruses such as murine poxvirus (ectromelia virus, EV) and herpes viruses, including HSV and Epstein-Barr virus (EBV), and some RNA viruses such as coxsackievirus.31,>>60<<–63 EBV reactivation appears to be inhibited by NO via suppression of an immediate early-transactivator gene.
n2:mentions
n3:8390481 n3:7528106 n3:10366574 n3:10023767
Subject Item
_:vb211507
rdf:type
n2:Context
rdf:value
virus, EV) and herpes viruses, including HSV and Epstein-Barr virus (EBV), and some RNA viruses such as coxsackievirus.31,60–63 EBV reactivation appears to be inhibited by NO via suppression of an immediate early-transactivator gene.>>61<<,62 In contrast, coxsackievirus replication is suppressed by NO through inactivation of its viral cysteine protease by NO-dependent S-nitrosylation.
n2:mentions
n3:7528106
Subject Item
_:vb211508
rdf:type
n2:Context
rdf:value
virus, EV) and herpes viruses, including HSV and Epstein-Barr virus (EBV), and some RNA viruses such as coxsackievirus.31,60–63 EBV reactivation appears to be inhibited by NO via suppression of an immediate early-transactivator gene.61,>>62<< In contrast, coxsackievirus replication is suppressed by NO through inactivation of its viral cysteine protease by NO-dependent S-nitrosylation.
n2:mentions
n3:10366574
Subject Item
_:vb211509
rdf:type
n2:Context
rdf:value
to be inhibited by NO via suppression of an immediate early-transactivator gene.61,62 In contrast, coxsackievirus replication is suppressed by NO through inactivation of its viral cysteine protease by NO-dependent S-nitrosylation.>>63<< S-Nitrosylation of various proteins and sulfhydryl targets of pathogens is of great interest in view of the diverse functions of NO.
n2:mentions
n3:10023767
Subject Item
_:vb211510
rdf:type
n2:Context
rdf:value
suppressed by NO through inactivation of its viral cysteine protease by NO-dependent S-nitrosylation.63 S-Nitrosylation of various proteins and sulfhydryl targets of pathogens is of great interest in view of the diverse functions of NO.>>64<<–69 However, it remains ambiguous whether selective toxicity of NO for virus and virus-infected cells is brought about by NO-dependent S-nitrosylation.
n2:mentions
n3:10480920 n3:9039953 n3:1281928 n3:8248198
Subject Item
_:vb211511
rdf:type
n2:Context
rdf:value
shows redox regulation by S-nitrosylation of sulfhydryl-containing proteins involved in inter- and intracellular signalling pathways, including neurotransmission, transcription, and apoptosis involving a caspase (thiol protease) cascade.>>66<<,68–71 For example, NO and nitroso adducts of sulfhydryl compounds (nitrosothiols) have a potent antiapoptotic activity through S-nitrosylation or transnitrosylation reactions with caspases.
n2:mentions
n3:9039953
Subject Item
_:vb211512
rdf:type
n2:Context
rdf:value
redox regulation by S-nitrosylation of sulfhydryl-containing proteins involved in inter- and intracellular signalling pathways, including neurotransmission, transcription, and apoptosis involving a caspase (thiol protease) cascade.66,>>68<<–71 For example, NO and nitroso adducts of sulfhydryl compounds (nitrosothiols) have a potent antiapoptotic activity through S-nitrosylation or transnitrosylation reactions with caspases.
n2:mentions
n3:10213689 n3:9240442
Subject Item
_:vb211513
rdf:type
n2:Context
rdf:value
such as ortho- and paramyxovirus, murine vaccinia virus, coronavirus (mouse hepatitis virus, MHV), lymphocytic choriomeningitis virus (LCMV), murine encephalomyocarditis virus (EMCV), tick-born encephalitis virus (TBE-V), and others.>>44<<–48,72,73 This lack of antiviral activity of NO has been proven in murine pneumotropic virus infections caused by influenza and Sendai viruses in a series of our in vitro and in vivo studies (Akaike et al., unpublished observation).
n2:mentions
n3:10605032 n3:10906226 n3:9782132 n3:10877838 n3:10580062
Subject Item
_:vb211514
rdf:type
n2:Context
rdf:value
as ortho- and paramyxovirus, murine vaccinia virus, coronavirus (mouse hepatitis virus, MHV), lymphocytic choriomeningitis virus (LCMV), murine encephalomyocarditis virus (EMCV), tick-born encephalitis virus (TBE-V), and others.44–48,>>72<<,73 This lack of antiviral activity of NO has been proven in murine pneumotropic virus infections caused by influenza and Sendai viruses in a series of our in vitro and in vivo studies (Akaike et al., unpublished observation).
n2:mentions
n3:10438821
Subject Item
_:vb211515
rdf:type
n2:Context
rdf:value
44–48,72,>>73<< This lack of antiviral activity of NO has been proven in murine pneumotropic virus infections caused by influenza and Sendai viruses in a series of our in vitro and in vivo studies (Akaike et al., unpublished observation).
n2:mentions
n3:8623546
Subject Item
_:vb211516
rdf:type
n2:Context
rdf:value
This lack of antiviral activity of NO has been proven in murine pneumotropic virus infections caused by influenza and Sendai viruses in a series of our in vitro and in vivo studies (Akaike et al., unpublished observation).>>48<< Exposure of these viruses to biologically relevant concentrations of NO produces no appreciable reduction of viral growth in cultured cells in vitro.
n2:mentions
n3:10877838
Subject Item
_:vb211517
rdf:type
n2:Context
rdf:value
More important, antiviral host defence is not impaired by pharmacological interventions producing NOS inhibition or by genetic iNOS deficiency of in mice infected with either influenza virus or Sendai virus.>>34<<,48 Such NO inhibition and lack of NO biosynthesis, however, significantly reduce the pathological consequences of various virus infections, including viral pneumonia in mice caused by influenza virus, Sendai virus, and HSV-1;
n2:mentions
n3:8637894
Subject Item
_:vb211518
rdf:type
n2:Context
rdf:value
More important, antiviral host defence is not impaired by pharmacological interventions producing NOS inhibition or by genetic iNOS deficiency of in mice infected with either influenza virus or Sendai virus.34,>>48<< Such NO inhibition and lack of NO biosynthesis, however, significantly reduce the pathological consequences of various virus infections, including viral pneumonia in mice caused by influenza virus, Sendai virus, and HSV-1; HSV-1-induced
n2:mentions
n3:10877838
Subject Item
_:vb211519
rdf:type
n2:Context
rdf:value
viral pneumonia in mice caused by influenza virus, Sendai virus, and HSV-1; HSV-1-induced encephalitis in rats; EMCV-induced carditis and diabetes; and murine encephalitis induced by flavivirus (Murray Valley encephalitis virus; TBE-V).>>34<<,35,45,48,73–77 It is thus conceivable that NO is not entirely an antiviral molecule in various, if not all, virus infections.
n2:mentions
n3:8637894
Subject Item
_:vb211520
rdf:type
n2:Context
rdf:value
pneumonia in mice caused by influenza virus, Sendai virus, and HSV-1; HSV-1-induced encephalitis in rats; EMCV-induced carditis and diabetes; and murine encephalitis induced by flavivirus (Murray Valley encephalitis virus; TBE-V).34,>>35<<,45,48,73–77 It is thus conceivable that NO is not entirely an antiviral molecule in various, if not all, virus infections.
n2:mentions
n3:10191185
Subject Item
_:vb211521
rdf:type
n2:Context
rdf:value
pneumonia in mice caused by influenza virus, Sendai virus, and HSV-1; HSV-1-induced encephalitis in rats; EMCV-induced carditis and diabetes; and murine encephalitis induced by flavivirus (Murray Valley encephalitis virus; TBE-V).34,35,>>45<<,48,73–77 It is thus conceivable that NO is not entirely an antiviral molecule in various, if not all, virus infections.
n2:mentions
n3:9782132
Subject Item
_:vb211522
rdf:type
n2:Context
rdf:value
34,35,45,>>48<<,73–77 It is thus conceivable that NO is not entirely an antiviral molecule in various, if not all, virus infections.
n2:mentions
n3:10877838
Subject Item
_:vb211523
rdf:type
n2:Context
rdf:value
34,35,45,48,>>73<<–77 It is thus conceivable that NO is not entirely an antiviral molecule in various, if not all, virus infections.
n2:mentions
n3:10477536 n3:8623546 n3:9151890 n3:10482632 n3:10482607
Subject Item
_:vb211524
rdf:type
n5:Section
dc:title
effects of no on immunological responses during virus infection and the pathological consequences
n5:contains
_:vb211540 _:vb211541 _:vb211542 _:vb211543 _:vb211536 _:vb211537 _:vb211538 _:vb211539 _:vb211548 _:vb211549 _:vb211550 _:vb211544 _:vb211545 _:vb211546 _:vb211547 _:vb211525 _:vb211526 _:vb211527 _:vb211532 _:vb211533 _:vb211534 _:vb211535 _:vb211528 _:vb211529 _:vb211530 _:vb211531
Subject Item
_:vb211525
rdf:type
n2:Context
rdf:value
It has been suggested that NO affects the polarized Th1–Th2 response, causing a Th2-biased immunoregulatory balance, via a relatively specific suppressive effect on Th1 subpopulations.>>78<<–80 Such NO-induced immunomodulation occurs during virus infection in mice, as revealed by recent studies of HSV-1 and influenza virus infections.
n2:mentions
n3:9864946 n3:7539113 n3:8149966
Subject Item
_:vb211526
rdf:type
n2:Context
rdf:value
balance, via a relatively specific suppressive effect on Th1 subpopulations.78–80 Such NO-induced immunomodulation occurs during virus infection in mice, as revealed by recent studies of HSV-1 and influenza virus infections.>>45<<,81 These biased Th2 responses are most clearly demonstrated by using iNOS–/– mice, which show enhanced Th1 immune responses after these infections.
n2:mentions
n3:9782132
Subject Item
_:vb211527
rdf:type
n2:Context
rdf:value
balance, via a relatively specific suppressive effect on Th1 subpopulations.78–80 Such NO-induced immunomodulation occurs during virus infection in mice, as revealed by recent studies of HSV-1 and influenza virus infections.45,>>81<< These biased Th2 responses are most clearly demonstrated by using iNOS–/– mice, which show enhanced Th1 immune responses after these infections.
n2:mentions
n3:9568978
Subject Item
_:vb211528
rdf:type
n2:Context
rdf:value
in mice, as revealed by recent studies of HSV-1 and influenza virus infections.45,81 These biased Th2 responses are most clearly demonstrated by using iNOS–/– mice, which show enhanced Th1 immune responses after these infections.>>45<<,81 It is believed that Th1 cells produce IL-2 and IFN-γ, whereas production of IL-4 and IL-10 depends on Th2 cells.
n2:mentions
n3:9782132
Subject Item
_:vb211529
rdf:type
n2:Context
rdf:value
in mice, as revealed by recent studies of HSV-1 and influenza virus infections.45,81 These biased Th2 responses are most clearly demonstrated by using iNOS–/– mice, which show enhanced Th1 immune responses after these infections.45,>>81<< It is believed that Th1 cells produce IL-2 and IFN-γ, whereas production of IL-4 and IL-10 depends on Th2 cells.
n2:mentions
n3:9568978
Subject Item
_:vb211530
rdf:type
n2:Context
rdf:value
effect on the Th1 response in HSV-1 infection in the foot pad and the dorsal root ganglion in mice, as evidenced by increased IFN-γ production and reduced levels of IL-4, and a subsequent elevation of anti-HSV-1 antibody in iNOS–/– mice.>>81<< HSV-1 clearance is delayed in iNOS–/– mice compared with the heterozygote mice, possibly because of impairment of the direct anti-HSV-1 activity of NO.
n2:mentions
n3:9568978
Subject Item
_:vb211531
rdf:type
n2:Context
rdf:value
antiviral host defence in iNOS–/– mice against influenza virus is much greater than that in the wild-type mice, possibly because the NO-induced impairment of the Th1 response (particularly IFN-γ production) is restored in iNOS–/– mice.>>45<< The same group demonstrated that mouse poxvirus (EV) is susceptible to NO. Therefore, virus infection in iNOS–/– mice is significantly exacerbated, even though the Th1-dependent IFN-γ response is enhanced by the disrupted iNOS gene.
n2:mentions
n3:9782132
Subject Item
_:vb211532
rdf:type
n2:Context
rdf:value
The same group demonstrated that mouse poxvirus (EV) is susceptible to NO. Therefore, virus infection in iNOS–/– mice is significantly exacerbated, even though the Th1-dependent IFN-γ response is enhanced by the disrupted iNOS gene.>>82<< Similarly, Lowenstein's group reported that the antiviral defence is eliminated in coxsackievirus-infected iNOS–/– mice.
n2:mentions
n3:9696880
Subject Item
_:vb211533
rdf:type
n2:Context
rdf:value
is significantly exacerbated, even though the Th1-dependent IFN-γ response is enhanced by the disrupted iNOS gene.82 Similarly, Lowenstein's group reported that the antiviral defence is eliminated in coxsackievirus-infected iNOS–/– mice.>>83<<,84 However, in infections with vaccinia virus and corona virus (MHV), iNOS deficiency affects neither antiviral host defence nor pathology of viral diseases.
n2:mentions
n3:10553076
Subject Item
_:vb211534
rdf:type
n2:Context
rdf:value
significantly exacerbated, even though the Th1-dependent IFN-γ response is enhanced by the disrupted iNOS gene.82 Similarly, Lowenstein's group reported that the antiviral defence is eliminated in coxsackievirus-infected iNOS–/– mice.83,>>84<< However, in infections with vaccinia virus and corona virus (MHV), iNOS deficiency affects neither antiviral host defence nor pathology of viral diseases.
n2:mentions
n3:9482909
Subject Item
_:vb211535
rdf:type
n2:Context
rdf:value
the antiviral defence is eliminated in coxsackievirus-infected iNOS–/– mice.83,84 However, in infections with vaccinia virus and corona virus (MHV), iNOS deficiency affects neither antiviral host defence nor pathology of viral diseases.>>44<< Also, the antiviral immunological response against LCMV is unimpaired in mice lacking iNOS, although T-cell-mediated inflammation induced by LCMV is reduced.
n2:mentions
n3:10605032
Subject Item
_:vb211536
rdf:type
n2:Context
rdf:value
44 Also, the antiviral immunological response against LCMV is unimpaired in mice lacking iNOS, although T-cell-mediated inflammation induced by LCMV is reduced.>>46<<
n2:mentions
n3:10580062
Subject Item
_:vb211537
rdf:type
n2:Context
rdf:value
on the type of virus, Th cells, divided into two subsets (Th1 and Th2), protect hosts from intruding viral pathogens via virus-specific Th1 responses, potentiation of CD8+ cytotoxic T lymphocyte (CTL) activity, and B-cell proliferation.>>85<<,86 For non-cytopathic virus infections, CTLs rather than Th1–Th2 cells are important for antiviral host defenses.
n2:mentions
n3:8539616
Subject Item
_:vb211538
rdf:type
n2:Context
rdf:value
the type of virus, Th cells, divided into two subsets (Th1 and Th2), protect hosts from intruding viral pathogens via virus-specific Th1 responses, potentiation of CD8+ cytotoxic T lymphocyte (CTL) activity, and B-cell proliferation.85,>>86<< For non-cytopathic virus infections, CTLs rather than Th1–Th2 cells are important for antiviral host defenses.
n2:mentions
n3:103005
Subject Item
_:vb211539
rdf:type
n2:Context
rdf:value
via virus-specific Th1 responses, potentiation of CD8+ cytotoxic T lymphocyte (CTL) activity, and B-cell proliferation.85,86 For non-cytopathic virus infections, CTLs rather than Th1–Th2 cells are important for antiviral host defenses.>>85<<,87 In contrast, some types of viruses, such as influenza virus, can be eradicated without the help of CTLs.
n2:mentions
n3:8539616
Subject Item
_:vb211540
rdf:type
n2:Context
rdf:value
virus-specific Th1 responses, potentiation of CD8+ cytotoxic T lymphocyte (CTL) activity, and B-cell proliferation.85,86 For non-cytopathic virus infections, CTLs rather than Th1–Th2 cells are important for antiviral host defenses.85,>>87<< In contrast, some types of viruses, such as influenza virus, can be eradicated without the help of CTLs.
n2:mentions
n3:8499074
Subject Item
_:vb211541
rdf:type
n2:Context
rdf:value
For non-cytopathic virus infections, CTLs rather than Th1–Th2 cells are important for antiviral host defenses.85,87 In contrast, some types of viruses, such as influenza virus, can be eradicated without the help of CTLs.>>41<< A virus-specific Th1 response is more important for influenza antiviral defence than the Th2 responses, because Th2 cells exacerbate pathological lung reactions in influenza pneumonia.
n2:mentions
n3:1919440
Subject Item
_:vb211542
rdf:type
n2:Context
rdf:value
virus, can be eradicated without the help of CTLs.41 A virus-specific Th1 response is more important for influenza antiviral defence than the Th2 responses, because Th2 cells exacerbate pathological lung reactions in influenza pneumonia.>>88<< In this context, Karupiah et al. reported that NO impairs the anti-influenza virus response of the host by suppressing Th1-dependent IFN-γ induction and tipping the Th1–Th2 balance toward Th2 domination.
n2:mentions
n3:7931062
Subject Item
_:vb211543
rdf:type
n2:Context
rdf:value
88 In this context, Karupiah et al. reported that NO impairs the anti-influenza virus response of the host by suppressing Th1-dependent IFN-γ induction and tipping the Th1–Th2 balance toward Th2 domination.>>45<< However, it has now been demonstrated that IFN-γ, a Th1-dependent cytokine, is eventually inefficient in clearance of influenza virus from infectious foci,42 and even IL-4, induced by Th2 responses, possesses antiviral activity against
n2:mentions
n3:9782132
Subject Item
_:vb211544
rdf:type
n2:Context
rdf:value
45 However, it has now been demonstrated that IFN-γ, a Th1-dependent cytokine, is eventually inefficient in clearance of influenza virus from infectious foci,>>42<< and even IL-4, induced by Th2 responses, possesses antiviral activity against murine paramyxovirus (Sendai virus).
n2:mentions
n3:8228818
Subject Item
_:vb211545
rdf:type
n2:Context
rdf:value
that IFN-γ, a Th1-dependent cytokine, is eventually inefficient in clearance of influenza virus from infectious foci,42 and even IL-4, induced by Th2 responses, possesses antiviral activity against murine paramyxovirus (Sendai virus).>>89<< Our recent experiments using iNOS–/– mice indicate that clearance of virus from lungs infected with either influenza virus or Sendai virus is not affected by a lack of iNOS expression (Akaike et al., unpublished observation).
n2:mentions
n3:9032393
Subject Item
_:vb211546
rdf:type
n2:Context
rdf:value
virus).89 Our recent experiments using iNOS–/– mice indicate that clearance of virus from lungs infected with either influenza virus or Sendai virus is not affected by a lack of iNOS expression (Akaike et al., unpublished observation).>>48<< In fact, iNOS–/– mice recuperate from viral pneumonia much better than do wild-type animals, because of reduced levels of oxidative stress in virus-infected tissues.
n2:mentions
n3:10877838
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a lack of iNOS expression (Akaike et al., unpublished observation).48 In fact, iNOS–/– mice recuperate from viral pneumonia much better than do wild-type animals, because of reduced levels of oxidative stress in virus-infected tissues.>>48<< Therefore, not only NO-induced Th1 suppression, but also NO-induced oxidative injury may, be attributable to pathogenesis of infection with certain viruses that are resistant to the direct antiviral actions of NO.
n2:mentions
n3:10877838
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In addition, NO seems to have profound immunosuppressive and immunopathological effects, most typically in Mycobacterium avium and Salmonella typhimurium infections,>>90<<,91 which may be due to NO-induced cytotoxic effects on immune effector cells such as macrophages.
n2:mentions
n3:9144497
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In addition, NO seems to have profound immunosuppressive and immunopathological effects, most typically in Mycobacterium avium and Salmonella typhimurium infections,90,>>91<< which may be due to NO-induced cytotoxic effects on immune effector cells such as macrophages.
n2:mentions
n3:10899909
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Similar immunosuppression by NO is clearly demonstrated with vaccinia virus-infected murine macrophages, which show a loss of antiviral activity because of inhibition of IFN-α/β production by NO.>>72<<
n2:mentions
n3:10438821
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n5:Section
dc:title
no-induced oxidative stress in pathogenesis of virus infection
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NO itself is an inert radical and much less reactive compared with other naturally occurring oxygen and alkyl rad- icals.>>7<<–9,11,12 Of the complex chemistry of NO, the most important and biologically relevant reaction is formation of ONOO– via a very rapid radical coupling with O2– (NO + O2– → ONOO–:
n2:mentions
n3:2154753 n3:8944624 n3:8828915
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NO itself is an inert radical and much less reactive compared with other naturally occurring oxygen and alkyl rad- icals.7–9,>>11<<,12 Of the complex chemistry of NO, the most important and biologically relevant reaction is formation of ONOO– via a very rapid radical coupling with O2– (NO + O2– → ONOO–:
n2:mentions
n3:10906338
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NO itself is an inert radical and much less reactive compared with other naturally occurring oxygen and alkyl rad- icals.7–9,11,>>12<< Of the complex chemistry of NO, the most important and biologically relevant reaction is formation of ONOO– via a very rapid radical coupling with O2– (NO + O2– → ONOO–:
n2:mentions
n3:10906340
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oxygen and alkyl rad- icals.7–9,11,12 Of the complex chemistry of NO, the most important and biologically relevant reaction is formation of ONOO– via a very rapid radical coupling with O2– (NO + O2– → ONOO–: k = 6·7 × 109 M−1 s−1).>>7<<–9,11,12 Although NO can function as an antioxidant, particularly in lipid peroxidation,9 it also has indirect pro-oxidant activity after conversion to a strong oxidant and a potent nitrating agent (ONOO–) causing oxidative stress.
n2:mentions
n3:2154753 n3:8944624 n3:8828915
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oxygen and alkyl rad- icals.7–9,11,12 Of the complex chemistry of NO, the most important and biologically relevant reaction is formation of ONOO– via a very rapid radical coupling with O2– (NO + O2– → ONOO–: k = 6·7 × 109 M−1 s−1).7–9,>>11<<,12 Although NO can function as an antioxidant, particularly in lipid peroxidation,9 it also has indirect pro-oxidant activity after conversion to a strong oxidant and a potent nitrating agent (ONOO–) causing oxidative stress.
n2:mentions
n3:10906338
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and alkyl rad- icals.7–9,11,12 Of the complex chemistry of NO, the most important and biologically relevant reaction is formation of ONOO– via a very rapid radical coupling with O2– (NO + O2– → ONOO–: k = 6·7 × 109 M−1 s−1).7–9,11,>>12<< Although NO can function as an antioxidant, particularly in lipid peroxidation,9 it also has indirect pro-oxidant activity after conversion to a strong oxidant and a potent nitrating agent (ONOO–) causing oxidative stress.
n2:mentions
n3:10906340
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7–9,11,12 Although NO can function as an antioxidant, particularly in lipid peroxidation,>>9<< it also has indirect pro-oxidant activity after conversion to a strong oxidant and a potent nitrating agent (ONOO–) causing oxidative stress.
n2:mentions
n3:8828915
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s−1).7–9,11,12 Although NO can function as an antioxidant, particularly in lipid peroxidation,9 it also has indirect pro-oxidant activity after conversion to a strong oxidant and a potent nitrating agent (ONOO–) causing oxidative stress.>>8<< In addition, although NO and nitrosothiols show strong anti-apoptotic effects as described above,67–71 ONOO– induces apoptosis, possibly via mitochondrial damage leading to cytochrome c release.
n2:mentions
n3:8944624
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8 In addition, although NO and nitrosothiols show strong anti-apoptotic effects as described above,>>67<<–71 ONOO– induces apoptosis, possibly via mitochondrial damage leading to cytochrome c release.
n2:mentions
n3:10213689 n3:9240442 n3:10480920
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8 In addition, although NO and nitrosothiols show strong anti-apoptotic effects as described above,67–71 ONOO– induces apoptosis, possibly via mitochondrial damage leading to cytochrome c release.>>10<<,92 NO chemistry in biological systems is shown schematically in Fig. 2.
n2:mentions
n3:10617463
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8 In addition, although NO and nitrosothiols show strong anti-apoptotic effects as described above,67–71 ONOO– induces apoptosis, possibly via mitochondrial damage leading to cytochrome c release.10,>>92<< NO chemistry in biological systems is shown schematically in Fig. 2.
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analysis with antinitrotyrosine antibody shows positive staining in macrophages and neutrophils infiltrating the alveoli and interstitial tissues, as well as in inflammatory intra-alveolar exudate in virus-infected lung,>>34<< which provides indirect indication of ONOO– generation during virus infection.
n2:mentions
n3:8637894
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in NO-induced suppressive effects on macrophages, as described in earlier sections. In addition, we recently found that ONOO– activates matrix metalloproteinases (MMPs), which are involved in extracellular tissue damage and remodelling.>>93<< Accordingly, oxidative tissue injury in virus-infected lung may be mediated by ONOO–--induced MMP activation.
n2:mentions
n3:9186487
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MMP activation. In fact, remarkable improvements in pathological condition in the lung and in survival rate of virus-infected mice were observed with l-NMMA, and with the O2– scavenger SOD and the XO inhibitor allopurinol as well.>>34<<,38 Furthermore, a therapeutic effect on influenza pathogenesis was found with a selenium-containing organic compound, ebselen (unpublished observation), which shows potent ONOO–-scavenging action.
n2:mentions
n3:8637894
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MMP activation. In fact, remarkable improvements in pathological condition in the lung and in survival rate of virus-infected mice were observed with l-NMMA, and with the O2– scavenger SOD and the XO inhibitor allopurinol as well.34,>>38<< Furthermore, a therapeutic effect on influenza pathogenesis was found with a selenium-containing organic compound, ebselen (unpublished observation), which shows potent ONOO–-scavenging action.
n2:mentions
n3:2155924
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the XO inhibitor allopurinol as well.34,38 Furthermore, a therapeutic effect on influenza pathogenesis was found with a selenium-containing organic compound, ebselen (unpublished observation), which shows potent ONOO–-scavenging action.>>94<< The beneficial effects of these pharmacological interventions indicate that ONOO– could be an important molecular species in the pathogenesis of influenza virus-induced pneumonia in mice.
n2:mentions
n3:8924601
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The pathological effects of NO and O2– in virus infection mentioned above are in clear contrast to their beneficial antimicrobial effects in bacterial and fungal infections.>>1<<–5 Antibacterial host defence mediated by NO is typified in a murine salmonellosis model.
n2:mentions
n3:8531884 n3:1711326 n3:8386705 n3:3280600 n3:9199469
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of NO and O2– in virus infection mentioned above are in clear contrast to their beneficial antimicrobial effects in bacterial and fungal infections.1–5 Antibacterial host defence mediated by NO is typified in a murine salmonellosis model.>>5<< When Salmonella typhimurium is injected into mice, effective bacterial growth is observed in the liver.
n2:mentions
n3:9199469
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Treatment of murine salmonellosis with either l-NMMA or allopurinol results in a greater mortality rate as well as increased bacterial growth.>>5<<
n2:mentions
n3:9199469
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non-specific oxidative damage in virus-infected tissue, leading to various pathological events, because virus cannot be confined to limited areas by the non-specific host defence mediated by phagocytes and their mediators NO and O2–.>>15<<
n2:mentions
n3:9421208
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n5:Section
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no-induced viral mutation and its possible link to molecular evolution of viruses
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It was previously shown that human leucocytes producing O2–, but not leucocytes from patients with chronic granulomatous disease, are mutagenic for S. typhimurium TA100.>>95<< ONOO– has mutagenic effects on prokaryotic DNA, possibly via nitration of guanine residues of DNA.
n2:mentions
n3:6259738
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_:vb211574
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ONOO– has mutagenic effects on prokaryotic DNA, possibly via nitration of guanine residues of DNA. A typical base substitution caused by ONOO– is G to T transversion, which is an indirect result of depurination of nitroguanine in DNA.>>96<<,97 A recent study documented that a high output of NO induced mutations in an endogenous hypoxanthine–guanine phosphoribosyltransferase (hprt) gene of murine macrophages expressing iNOS.
n2:mentions
n3:7554052
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ONOO– has mutagenic effects on prokaryotic DNA, possibly via nitration of guanine residues of DNA. A typical base substitution caused by ONOO– is G to T transversion, which is an indirect result of depurination of nitroguanine in DNA.96,>>97<< A recent study documented that a high output of NO induced mutations in an endogenous hypoxanthine–guanine phosphoribosyltransferase (hprt) gene of murine macrophages expressing iNOS.
n2:mentions
n3:8569792
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96,97 A recent study documented that a high output of NO induced mutations in an endogenous hypoxanthine–guanine phosphoribosyltransferase (hprt) gene of murine macrophages expressing iNOS.>>98<<
n2:mentions
n3:9653179
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_:vb211577
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However, only a few reports explore a possible association between oxidative stress and viral mutation. Beck et al. showed that the pathogenicity of coxsackievirus B3 is strongly potentiated in vivo in mice fed a selenium-deficient diet.>>99<< More important, an avirulent strain of the virus is converted to a potent cardiotoxic variant during infection in selenium-depleted animals.
n2:mentions
n3:7585090
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_:vb211578
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system, e.g. low levels of glutathione peroxidase. Results of similar studies extended to animals deficient in vitamin E and glutathione peroxidase suggest that oxidative stress facilitates selection and generation of virulent mutants.>>100<< More specifically, the impaired immunological viral clearance related to oxidative stress may cause increased survival of heterogeneous mutants, resulting in selection of highly pathogenic variants of coxsackievirus.
n2:mentions
n3:9737717
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In addition, our recent study verifies for the first time that oxidative stress induced by a high output of NO accelerates mutation of RNA virus.>>48<< By using a recombinant RNA virus (Sendai) containing a marker gene (green fluorescent protein, GFP) for genetic mutation and iNOS knockout mice, we clearly showed that oxidative stress induced by NO in wild-type mice in vivo remarkably
n2:mentions
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