Abstract
Systemic acquired resistance (SAR) is induced by pathogens and confers protection against a broad range of pathogens. Several SAR signals have been characterized, but the nature of the other unknown signalling by small metabolites in SAR remains unclear. Glutathione (GSH) has long been implicated in the defence reaction against biotic stress. However, the mechanism that GSH increases plant tolerance against virus infection is not entirely known. Here, a combination of a chemical, virus-induced gene-silencing-based genetics approach, and transgenic technology was undertaken to investigate the role of GSH in plant viral resistance in Nicotiana benthamiana. Tobacco mosaic virus (TMV) infection results in increasing the expression of GSH biosynthesis genes NbECS and NbGS, and GSH content. Silencing of NbECS or NbGS accelerated oxidative damage, increased accumulation of reactive oxygen species (ROS), compromised plant resistance to TMV, and suppressed the salicylic acid (SA)-mediated signalling pathway. Application of GSH or l-2-oxothiazolidine-4-carboxylic acid (a GSH activator) alleviated oxidative damage, decreased accumulation of ROS, elevated plant local and systemic resistance, enhanced the SA-mediated signalling pathway, and increased the expression of ROS scavenging-related genes. However, treatment with buthionine sulfoximine (a GSH inhibitor) accelerated oxidative damage, elevated ROS accumulation, compromised plant systemic resistance, suppressed the SA-mediated signalling pathway, and reduced the expression of ROS-regulating genes. Overexpression of NbECS reduced oxidative damage, decreased accumulation of ROS, increased resistance to TMV, activated the SA-mediated signalling pathway, and increased the expression of the ROS scavenging-related genes. We present molecular evidence suggesting GSH is essential for both local and systemic resistance of N. benthamiana to TMV through a differential modulation of SA and ROS.
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Claims 20
Claim · model-written summary
TMV infection increased the expression of GSH biosynthesis genes NbECS and NbGS and GSH content in N. benthamiana.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- Tobacco mosaic virus (TMV) infectione1
- Interactor
- Tobacco mosaic virus (TMV)
Tobacco mosaic viruse1pathogen - Gene
- NbECS and NbGSe1
- Outcome
- expression of GSH biosynthesis genes NbECS and NbGS, and GSH contente1
- Direction
- increasinge1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.


Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e1“Tobacco mosaic virus (TMV) infection results in increasing the expression of GSH biosynthesis genes NbECS and NbGS, and GSH content”
Claim · model-written summary
Silencing of NbECS or NbGS accelerated oxidative damage in N. benthamiana.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- Silencing of NbECS or NbGSe2
- Gene
- NbECS or NbGSe2
- Outcome
- oxidative damagee2
- Direction
- acceleratede2
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e2“Silencing of NbECS or NbGS accelerated oxidative damage, increased accumulation of reactive oxygen species (ROS), compromised plant resistance to TMV, and suppressed the salicylic acid (SA)-mediated signalling pathway”
Claim · model-written summary
Silencing of NbECS or NbGS increased ROS accumulation in N. benthamiana.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- Silencing of NbECS or NbGSe2
- Gene
- NbECS or NbGSe2
- Outcome
- accumulation of reactive oxygen species (ROS)e2
- Direction
- increasede2
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e2“Silencing of NbECS or NbGS accelerated oxidative damage, increased accumulation of reactive oxygen species (ROS), compromised plant resistance to TMV, and suppressed the salicylic acid (SA)-mediated signalling pathway”
Claim · model-written summary
Silencing of NbECS or NbGS compromised plant resistance to TMV.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- Silencing of NbECS or NbGSe2
- Gene
- NbECS or NbGSe2
- Outcome
- plant resistance to TMVe2
- Direction
- compromisede2
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e2“Silencing of NbECS or NbGS accelerated oxidative damage, increased accumulation of reactive oxygen species (ROS), compromised plant resistance to TMV, and suppressed the salicylic acid (SA)-mediated signalling pathway”
Claim · model-written summary
Silencing of NbECS or NbGS suppressed the SA-mediated signalling pathway.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- Silencing of NbECS or NbGSe2
- Gene
- NbECS or NbGSe2
- Outcome
- salicylic acid (SA)-mediated signalling pathwaye2
- Direction
- suppressede2
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e2“Silencing of NbECS or NbGS accelerated oxidative damage, increased accumulation of reactive oxygen species (ROS), compromised plant resistance to TMV, and suppressed the salicylic acid (SA)-mediated signalling pathway”
Claim · model-written summary
Application of GSH or its activator alleviated oxidative damage in N. benthamiana.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- Application of GSH or l-2-oxothiazolidine-4-carboxylic acid (a GSH activator)e3
- Compound
- GSHe3
- Outcome
- oxidative damagee3
- Direction
- alleviatede3
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e3“Application of GSH or l-2-oxothiazolidine-4-carboxylic acid (a GSH activator) alleviated oxidative damage, decreased accumulation of ROS, elevated plant local and systemic resistance, enhanced the SA-mediated signalling pathway, and increased the expression of ROS scavenging-related genes”
Claim · model-written summary
Application of GSH or its activator decreased ROS accumulation in N. benthamiana.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- Application of GSH or l-2-oxothiazolidine-4-carboxylic acid (a GSH activator)e3
- Compound
- GSHe3
- Outcome
- accumulation of ROSe3
- Direction
- decreasede3
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e3“Application of GSH or l-2-oxothiazolidine-4-carboxylic acid (a GSH activator) alleviated oxidative damage, decreased accumulation of ROS, elevated plant local and systemic resistance, enhanced the SA-mediated signalling pathway, and increased the expression of ROS scavenging-related genes”
Claim · model-written summary
Application of GSH or its activator elevated local and systemic resistance of N. benthamiana.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- Application of GSH or l-2-oxothiazolidine-4-carboxylic acid (a GSH activator)e3
- Compound
- GSHe3
- Outcome
- plant local and systemic resistancee3
- Direction
- elevatede3
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e3“Application of GSH or l-2-oxothiazolidine-4-carboxylic acid (a GSH activator) alleviated oxidative damage, decreased accumulation of ROS, elevated plant local and systemic resistance, enhanced the SA-mediated signalling pathway, and increased the expression of ROS scavenging-related genes”
Claim · model-written summary
Application of GSH or its activator enhanced the SA-mediated signalling pathway in N. benthamiana.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- Application of GSH or l-2-oxothiazolidine-4-carboxylic acid (a GSH activator)e3
- Compound
- GSHe3
- Outcome
- SA-mediated signalling pathwaye3
- Direction
- enhancede3
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e3“Application of GSH or l-2-oxothiazolidine-4-carboxylic acid (a GSH activator) alleviated oxidative damage, decreased accumulation of ROS, elevated plant local and systemic resistance, enhanced the SA-mediated signalling pathway, and increased the expression of ROS scavenging-related genes”
Claim · model-written summary
Application of GSH or its activator increased the expression of ROS scavenging-related genes in N. benthamiana.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- Application of GSH or l-2-oxothiazolidine-4-carboxylic acid (a GSH activator)e3
- Gene
- ROS scavenging-related genese3
- Compound
- GSHe3
- Outcome
- expression of ROS scavenging-related genese3
- Direction
- increasede3
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e3“Application of GSH or l-2-oxothiazolidine-4-carboxylic acid (a GSH activator) alleviated oxidative damage, decreased accumulation of ROS, elevated plant local and systemic resistance, enhanced the SA-mediated signalling pathway, and increased the expression of ROS scavenging-related genes”
Claim · model-written summary
Treatment with buthionine sulfoximine (a GSH inhibitor) accelerated oxidative damage in N. benthamiana.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- treatment with buthionine sulfoximine (a GSH inhibitor)e4
- Outcome
- oxidative damagee4
- Direction
- acceleratede4
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e4“However, treatment with buthionine sulfoximine (a GSH inhibitor) accelerated oxidative damage, elevated ROS accumulation, compromised plant systemic resistance, suppressed the SA-mediated signalling pathway, and reduced the expression of ROS-regulating genes”
Claim · model-written summary
Treatment with buthionine sulfoximine (a GSH inhibitor) elevated ROS accumulation in N. benthamiana.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- treatment with buthionine sulfoximine (a GSH inhibitor)e4
- Outcome
- ROS accumulatione4
- Direction
- elevatede4
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e4“However, treatment with buthionine sulfoximine (a GSH inhibitor) accelerated oxidative damage, elevated ROS accumulation, compromised plant systemic resistance, suppressed the SA-mediated signalling pathway, and reduced the expression of ROS-regulating genes”
Claim · model-written summary
Treatment with buthionine sulfoximine (a GSH inhibitor) compromised systemic resistance of N. benthamiana.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- treatment with buthionine sulfoximine (a GSH inhibitor)e4
- Outcome
- plant systemic resistancee4
- Direction
- compromisede4
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e4“However, treatment with buthionine sulfoximine (a GSH inhibitor) accelerated oxidative damage, elevated ROS accumulation, compromised plant systemic resistance, suppressed the SA-mediated signalling pathway, and reduced the expression of ROS-regulating genes”
Claim · model-written summary
Treatment with buthionine sulfoximine (a GSH inhibitor) suppressed the SA-mediated signalling pathway in N. benthamiana.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- treatment with buthionine sulfoximine (a GSH inhibitor)e4
- Outcome
- SA-mediated signalling pathwaye4
- Direction
- suppressede4
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e4“However, treatment with buthionine sulfoximine (a GSH inhibitor) accelerated oxidative damage, elevated ROS accumulation, compromised plant systemic resistance, suppressed the SA-mediated signalling pathway, and reduced the expression of ROS-regulating genes”
Claim · model-written summary
Treatment with buthionine sulfoximine (a GSH inhibitor) reduced the expression of ROS-regulating genes in N. benthamiana.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- treatment with buthionine sulfoximine (a GSH inhibitor)e4
- Gene
- ROS-regulating genese4
- Outcome
- expression of ROS-regulating genese4
- Direction
- reducede4
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e4“However, treatment with buthionine sulfoximine (a GSH inhibitor) accelerated oxidative damage, elevated ROS accumulation, compromised plant systemic resistance, suppressed the SA-mediated signalling pathway, and reduced the expression of ROS-regulating genes”
Claim · model-written summary
Overexpression of NbECS reduced oxidative damage in N. benthamiana.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- Overexpression of NbECSe5
- Gene
- NbECSe5
- Outcome
- oxidative damagee5
- Direction
- reducede5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e5“Overexpression of NbECS reduced oxidative damage, decreased accumulation of ROS, increased resistance to TMV, activated the SA-mediated signalling pathway, and increased the expression of the ROS scavenging-related genes”
Claim · model-written summary
Overexpression of NbECS decreased ROS accumulation in N. benthamiana.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- Overexpression of NbECSe5
- Gene
- NbECSe5
- Outcome
- accumulation of ROSe5
- Direction
- decreasede5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e5“Overexpression of NbECS reduced oxidative damage, decreased accumulation of ROS, increased resistance to TMV, activated the SA-mediated signalling pathway, and increased the expression of the ROS scavenging-related genes”
Claim · model-written summary
Overexpression of NbECS increased resistance of N. benthamiana to TMV.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- Overexpression of NbECSe5
- Gene
- NbECSe5
- Outcome
- resistance to TMVe5
- Direction
- increasede5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e5“Overexpression of NbECS reduced oxidative damage, decreased accumulation of ROS, increased resistance to TMV, activated the SA-mediated signalling pathway, and increased the expression of the ROS scavenging-related genes”
Claim · model-written summary
Overexpression of NbECS activated the SA-mediated signalling pathway in N. benthamiana.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- Overexpression of NbECSe5
- Gene
- NbECSe5
- Outcome
- SA-mediated signalling pathwaye5
- Direction
- activatede5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e5“Overexpression of NbECS reduced oxidative damage, decreased accumulation of ROS, increased resistance to TMV, activated the SA-mediated signalling pathway, and increased the expression of the ROS scavenging-related genes”
Claim · model-written summary
Overexpression of NbECS increased the expression of ROS scavenging-related genes in N. benthamiana.
- Taxon
- Nicotiana benthamiana
Nicotiana benthamianae0 - Treatment
- Overexpression of NbECSe5
- Gene
- ROS scavenging-related genese5
- Outcome
- expression of the ROS scavenging-related genese5
- Direction
- increasede5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“plant viral resistance in Nicotiana benthamiana”
e5“Overexpression of NbECS reduced oxidative damage, decreased accumulation of ROS, increased resistance to TMV, activated the SA-mediated signalling pathway, and increased the expression of the ROS scavenging-related genes”
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Claims are grouped by organism, treatment and what was measured or targeted. A paper counts as positive when any of its claims in the group is positive.