Abstract
Reduced glutathione (γ-glutamyl-cysteinyl-glycine, GSH), the primary non-protein sulfhydryl group in organisms, plays a pivotal role in the plant salt stress response. This study aimed to explore the impact of GSH on the photosynthetic apparatus, and carbon assimilation in tomato plants under salt stress, and then investigate the role of nitric oxide (NO) in this process. The investigation involved foliar application of 5 mM GSH, 0.1% (w/v) hemoglobin (Hb, a nitric oxide scavenger), and GSH+Hb on the endogenous NO levels, rapid chlorophyll fluorescence, enzyme activities, and gene expression related to the Calvin cycle in tomato seedlings ( Solanum lycopersicum L. cv. ‘Zhongshu No. 4’) subjected short-term salt stress (100 mM NaCl) for 24, 48 and 72 hours. GSH treatment notably boosted nitrate reductase (NR) and NO synthase (NOS) activities, elevating endogenous NO signaling in salt-stressed tomato seedling leaves. It also mitigated chlorophyll fluorescence (OJIP) curve distortion and damage to the oxygen-evolving complex (OEC) induced by salt stress. Furthermore, GSH improved photosystem II (PSII) electron transfer efficiency, reduced Q A - accumulation, and countered salt stress effects on photosystem I (PSI) redox properties, enhancing the light energy absorption index (PI abs ). Additionally, GSH enhanced key enzyme activities in the Calvin cycle and upregulated their genes. Exogenous GSH optimized PSII energy utilization via endogenous NO, safeguarded the photosynthetic reaction center, improved photochemical and energy efficiency, and boosted carbon assimilation, ultimately enhancing net photosynthetic efficiency (P n ) in salt-stressed tomato seedling leaves. Conversely, Hb hindered P n reduction and NO signaling under salt stress and weakened the positive effects of GSH on NO levels, photosynthetic apparatus, and carbon assimilation in tomato plants. Thus, the positive regulation of photosynthesis in tomato seedlings under salt stress by GSH requires the involvement of NO.
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Claims 17
Claim · model-written summary
GSH boosted nitrate reductase and NO synthase activities in salt-stressed tomato.
- Taxon
- Solanum lycopersicum L
Solanum lycopersicumt0 - Treatment
- GSHe0
- Outcome
- nitrate reductase (NR) and NO synthase (NOS) activitiese0
- Direction
- boostede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“GSH treatment notably boosted nitrate reductase (NR) and NO synthase (NOS) activities, elevating endogenous NO signaling in salt-stressed tomato seedling leaves.”
t0“Solanum lycopersicum L”
Claim · model-written summary
GSH elevated endogenous NO signaling in salt-stressed tomato.
- Taxon
- Solanum lycopersicum L
Solanum lycopersicumt0 - Treatment
- GSHe0
- Outcome
- endogenous NO signalinge0
- Direction
- elevatinge0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“GSH treatment notably boosted nitrate reductase (NR) and NO synthase (NOS) activities, elevating endogenous NO signaling in salt-stressed tomato seedling leaves.”
t0“Solanum lycopersicum L”
Claim · model-written summary
GSH mitigated chlorophyll fluorescence (OJIP) curve distortion.
- Taxon
- Solanum lycopersicum L
Solanum lycopersicumt0 - Treatment
- GSHe0
- Outcome
- chlorophyll fluorescence (OJIP) curve distortione0
- Direction
- mitigatede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“GSH treatment notably boosted nitrate reductase (NR) and NO synthase (NOS) activities, elevating endogenous NO signaling in salt-stressed tomato seedling leaves. It also mitigated chlorophyll fluorescence (OJIP) curve distortion and damage to the oxygen-evolving complex (OEC) induced by salt stress.”
t0“Solanum lycopersicum L”
Claim · model-written summary
GSH mitigated damage to the oxygen-evolving complex (OEC).
- Taxon
- Solanum lycopersicum L
Solanum lycopersicumt0 - Treatment
- GSHe0
- Outcome
- damage to the oxygen-evolving complex (OEC)e0
- Direction
- mitigatede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“GSH treatment notably boosted nitrate reductase (NR) and NO synthase (NOS) activities, elevating endogenous NO signaling in salt-stressed tomato seedling leaves. It also mitigated chlorophyll fluorescence (OJIP) curve distortion and damage to the oxygen-evolving complex (OEC) induced by salt stress.”
t0“Solanum lycopersicum L”
Claim · model-written summary
GSH improved photosystem II (PSII) electron transfer efficiency.
- Taxon
- Solanum lycopersicum L
Solanum lycopersicumt0 - Treatment
- GSHe0
- Outcome
- photosystem II (PSII) electron transfer efficiencye0
- Direction
- improvede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“GSH improved photosystem II (PSII) electron transfer efficiency, reduced Q A - accumulation, and countered salt stress effects on photosystem I (PSI) redox properties, enhancing the light energy absorption index (PI abs ).”
t0“Solanum lycopersicum L”
Claim · model-written summary
GSH reduced Q A - accumulation.
- Taxon
- Solanum lycopersicum L
Solanum lycopersicumt0 - Treatment
- GSHe0
- Outcome
- Q A - accumulatione0
- Direction
- reducede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“GSH improved photosystem II (PSII) electron transfer efficiency, reduced Q A - accumulation, and countered salt stress effects on photosystem I (PSI) redox properties, enhancing the light energy absorption index (PI abs ).”
t0“Solanum lycopersicum L”
Claim · model-written summary
GSH countered photosystem I (PSI) redox properties.
- Taxon
- Solanum lycopersicum L
Solanum lycopersicumt0 - Treatment
- GSHe0
- Outcome
- photosystem I (PSI) redox propertiese0
- Direction
- counterede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“GSH improved photosystem II (PSII) electron transfer efficiency, reduced Q A - accumulation, and countered salt stress effects on photosystem I (PSI) redox properties, enhancing the light energy absorption index (PI abs ).”
t0“Solanum lycopersicum L”
Claim · model-written summary
GSH enhancing light energy absorption index (PI abs ).
- Taxon
- Solanum lycopersicum L
Solanum lycopersicumt0 - Treatment
- GSHe0
- Outcome
- light energy absorption index (PI abs )e0
- Direction
- enhancinge0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“GSH improved photosystem II (PSII) electron transfer efficiency, reduced Q A - accumulation, and countered salt stress effects on photosystem I (PSI) redox properties, enhancing the light energy absorption index (PI abs ).”
t0“Solanum lycopersicum L”
Claim · model-written summary
GSH enhanced Calvin cycle enzyme activities.
- Taxon
- Solanum lycopersicum L
Solanum lycopersicumt0 - Treatment
- GSHe0
- Outcome
- key enzyme activities in the Calvin cyclee0
- Direction
- enhancede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Additionally, GSH enhanced key enzyme activities in the Calvin cycle and upregulated their genes.”
t0“Solanum lycopersicum L”
Claim · model-written summary
GSH upregulated genes for Calvin cycle enzymes.
- Taxon
- Solanum lycopersicum L
Solanum lycopersicumt0 - Treatment
- GSHe0
- Outcome
- their genese0
- Direction
- upregulatede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Additionally, GSH enhanced key enzyme activities in the Calvin cycle and upregulated their genes.”
t0“Solanum lycopersicum L”
Claim · model-written summary
Exogenous GSH optimized PSII energy utilization.
- Taxon
- Solanum lycopersicum L
Solanum lycopersicumt0 - Treatment
- Exogenous GSHe0
- Outcome
- PSII energy utilizatione0
- Direction
- optimizede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Exogenous GSH optimized PSII energy utilization via endogenous NO, safeguarded the photosynthetic reaction center, improved photochemical and energy efficiency, and boosted carbon assimilation, ultimately enhancing net photosynthetic efficiency (P n ) in salt-stressed tomato seedling leaves.”
t0“Solanum lycopersicum L”
Claim · model-written summary
Exogenous GSH safeguarded photosynthetic reaction center.
- Taxon
- Solanum lycopersicum L
Solanum lycopersicumt0 - Treatment
- Exogenous GSHe0
- Outcome
- photosynthetic reaction centere0
- Direction
- safeguardede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Exogenous GSH optimized PSII energy utilization via endogenous NO, safeguarded the photosynthetic reaction center, improved photochemical and energy efficiency, and boosted carbon assimilation, ultimately enhancing net photosynthetic efficiency (P n ) in salt-stressed tomato seedling leaves.”
t0“Solanum lycopersicum L”
Claim · model-written summary
Exogenous GSH improved photochemical and energy efficiency.
- Taxon
- Solanum lycopersicum L
Solanum lycopersicumt0 - Treatment
- Exogenous GSHe0
- Outcome
- photochemical and energy efficiencye0
- Direction
- improvede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Exogenous GSH optimized PSII energy utilization via endogenous NO, safeguarded the photosynthetic reaction center, improved photochemical and energy efficiency, and boosted carbon assimilation, ultimately enhancing net photosynthetic efficiency (P n ) in salt-stressed tomato seedling leaves.”
t0“Solanum lycopersicum L”
Claim · model-written summary
Exogenous GSH boosted carbon assimilation.
- Taxon
- Solanum lycopersicum L
Solanum lycopersicumt0 - Treatment
- Exogenous GSHe0
- Outcome
- carbon assimilatione0
- Direction
- boostede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Exogenous GSH optimized PSII energy utilization via endogenous NO, safeguarded the photosynthetic reaction center, improved photochemical and energy efficiency, and boosted carbon assimilation, ultimately enhancing net photosynthetic efficiency (P n ) in salt-stressed tomato seedling leaves.”
t0“Solanum lycopersicum L”
Claim · model-written summary
Exogenous GSH enhancing net photosynthetic efficiency (P n ).
- Taxon
- Solanum lycopersicum L
Solanum lycopersicumt0 - Treatment
- Exogenous GSHe0
- Outcome
- net photosynthetic efficiency (P n )e0
- Direction
- enhancinge0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Exogenous GSH optimized PSII energy utilization via endogenous NO, safeguarded the photosynthetic reaction center, improved photochemical and energy efficiency, and boosted carbon assimilation, ultimately enhancing net photosynthetic efficiency (P n ) in salt-stressed tomato seedling leaves.”
t0“Solanum lycopersicum L”
Claim · model-written summary
Hb weakened GSH effects on NO, photosynthetic apparatus, and carbon assimilation.
- Taxon
- tomato plants
Solanum lycopersicume0 - Treatment
- Hbe0
- Outcome
- NO levels, photosynthetic apparatus, and carbon assimilatione0
- Direction
- weakenede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Conversely, Hb hindered P n reduction and NO signaling under salt stress and weakened the positive effects of GSH on NO levels, photosynthetic apparatus, and carbon assimilation in tomato plants.”
Claim · model-written summary
GSH regulation of photosynthesis under salt stress requires NO.
- Taxon
- tomato seedlings
Solanum lycopersicume0 - Treatment
- GSHe0
- Compound
- NOe0
- Outcome
- positive regulation of photosynthesise0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Thus, the positive regulation of photosynthesis in tomato seedlings under salt stress by GSH requires the involvement of NO.”
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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.