Abstract
Abstract Thermotolerance is improved by heat stress (HS) acclimation, and the thermotolerance level is “remembered” by plants. However, the underlying signalling mechanisms remain largely unknown. Here, we showed NADPH oxidase‐mediated H 2 O 2 (NADPH‐H 2 O 2 ), and chloroplast‐H 2 O 2 promoted the sustained expression of HS‐responsive genes and programmed cell death (PCD) genes, respectively, during recovery after HS acclimation. When spraying the NADPH oxidase inhibitor, diphenylene iodonium, after HS acclimation, the NADPH‐H 2 O 2 level significantly decreased, resulting in a decrease in the expression of HS‐responsive genes and the loss of maintenance of acquired thermotolerance (MAT). In contrast, compared with HS acclimation, NADPH‐H 2 O 2 declined but chloroplast‐H 2 O 2 further enhanced during recovery after HS over‐acclimation, resulting in the reduced expression of HS‐responsive genes and substantial production of PCD. Notably, the further inhibition of NADPH‐H 2 O 2 after HS over‐acclimation also inhibited chloroplast‐H 2 O 2 , alleviating the severe PCD and surpassing the MAT of HS over‐acclimation treatment. Due to the change in subcellular H 2 O 2 after HS acclimation, the tomato seedlings maintained a constant H 2 O 2 level during recovery, resulting in stable and lower total H 2 O 2 levels during a tester HS challenge conducted after recovery. We conclude that tomato seedlings increase their MAT by enhancing NADPH‐H 2 O 2 content and controlling chloroplast‐H 2 O 2 production during recovery, which enhances the expression of HS‐responsive genes and balances PCD levels, respectively.
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Claims 15
Claim · model-written summary
NADPH‐H 2 O 2 promoted sustained expression of HS-responsive genes.
- Taxon
- tomato seedlings
Solanum lycopersicume7 - Compound
- NADPH‐H 2 O 2e0
- Outcome
- sustained expression of HS‐responsive genese0
- Qualifier
- during recovery after HS acclimatione0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e7“the tomato seedlings maintained a constant H 2 O 2 level”
e0“NADPH oxidase‐mediated H 2 O 2 (NADPH‐H 2 O 2 ), and chloroplast‐H 2 O 2 promoted the sustained expression of HS‐responsive genes and programmed cell death (PCD) genes, respectively, during recovery after HS acclimation.”
Claim · model-written summary
Chloroplast H2O2 promoted PCD gene expression.
- Taxon
- tomato seedlings
Solanum lycopersicume7 - Compound
- chloroplast‐H 2 O 2e0
- Outcome
- programmed cell death (PCD) genese0
- Qualifier
- during recovery after HS acclimatione0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e7“the tomato seedlings maintained a constant H 2 O 2 level”
e0“NADPH oxidase‐mediated H 2 O 2 (NADPH‐H 2 O 2 ), and chloroplast‐H 2 O 2 promoted the sustained expression of HS‐responsive genes and programmed cell death (PCD) genes, respectively, during recovery after HS acclimation.”
Claim · model-written summary
Diphenylene iodonium after acclimation caused decreased NADPH‐H 2 O 2 level.
- Taxon
- tomato seedlings
Solanum lycopersicume7 - Treatment
- spraying the NADPH oxidase inhibitor, diphenylene iodoniume0
- Outcome
- NADPH‐H 2 O 2 levele0
- Direction
- decreasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e7“the tomato seedlings maintained a constant H 2 O 2 level”
e0“When spraying the NADPH oxidase inhibitor, diphenylene iodonium, after HS acclimation, the NADPH‐H 2 O 2 level significantly decreased, resulting in a decrease in the expression of HS‐responsive genes and the loss of maintenance of acquired thermotolerance (MAT).”
Claim · model-written summary
Diphenylene iodonium after acclimation caused decrease expression of HS‐responsive genes.
- Taxon
- tomato seedlings
Solanum lycopersicume7 - Treatment
- spraying the NADPH oxidase inhibitor, diphenylene iodoniume0
- Outcome
- expression of HS‐responsive genese0
- Direction
- decreasee0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e7“the tomato seedlings maintained a constant H 2 O 2 level”
e0“When spraying the NADPH oxidase inhibitor, diphenylene iodonium, after HS acclimation, the NADPH‐H 2 O 2 level significantly decreased, resulting in a decrease in the expression of HS‐responsive genes and the loss of maintenance of acquired thermotolerance (MAT).”
Claim · model-written summary
Diphenylene iodonium after acclimation caused loss maintenance of acquired thermotolerance (MAT).
- Taxon
- tomato seedlings
Solanum lycopersicume7 - Treatment
- spraying the NADPH oxidase inhibitor, diphenylene iodoniume0
- Outcome
- maintenance of acquired thermotolerance (MAT)e0
- Direction
- losse0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e7“the tomato seedlings maintained a constant H 2 O 2 level”
e0“When spraying the NADPH oxidase inhibitor, diphenylene iodonium, after HS acclimation, the NADPH‐H 2 O 2 level significantly decreased, resulting in a decrease in the expression of HS‐responsive genes and the loss of maintenance of acquired thermotolerance (MAT).”
Claim · model-written summary
After over-acclimation, NADPH‐H 2 O 2 was declined relative to acclimation.
- Taxon
- tomato seedlings
Solanum lycopersicume7 - Treatment
- HS over‐acclimatione0
- Outcome
- NADPH‐H 2 O 2e0
- Direction
- declinede0
- Comparator
- HS acclimatione0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e7“the tomato seedlings maintained a constant H 2 O 2 level”
e0“compared with HS acclimation, NADPH‐H 2 O 2 declined but chloroplast‐H 2 O 2 further enhanced during recovery after HS over‐acclimation, resulting in the reduced expression of HS‐responsive genes and substantial production of PCD.”
Claim · model-written summary
After over-acclimation, chloroplast‐H 2 O 2 was enhanced relative to acclimation.
- Taxon
- tomato seedlings
Solanum lycopersicume7 - Treatment
- HS over‐acclimatione0
- Outcome
- chloroplast‐H 2 O 2e0
- Direction
- enhancede0
- Comparator
- HS acclimatione0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e7“the tomato seedlings maintained a constant H 2 O 2 level”
e0“compared with HS acclimation, NADPH‐H 2 O 2 declined but chloroplast‐H 2 O 2 further enhanced during recovery after HS over‐acclimation, resulting in the reduced expression of HS‐responsive genes and substantial production of PCD.”
Claim · model-written summary
After over-acclimation, expression of HS‐responsive genes was reduced relative to acclimation.
- Taxon
- tomato seedlings
Solanum lycopersicume7 - Treatment
- HS over‐acclimatione0
- Outcome
- expression of HS‐responsive genese0
- Direction
- reducede0
- Comparator
- HS acclimatione0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e7“the tomato seedlings maintained a constant H 2 O 2 level”
e0“compared with HS acclimation, NADPH‐H 2 O 2 declined but chloroplast‐H 2 O 2 further enhanced during recovery after HS over‐acclimation, resulting in the reduced expression of HS‐responsive genes and substantial production of PCD.”
Claim · model-written summary
After over-acclimation, production of PCD was substantial relative to acclimation.
- Taxon
- tomato seedlings
Solanum lycopersicume7 - Treatment
- HS over‐acclimatione0
- Outcome
- production of PCDe0
- Direction
- substantiale0
- Comparator
- HS acclimatione0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e7“the tomato seedlings maintained a constant H 2 O 2 level”
e0“compared with HS acclimation, NADPH‐H 2 O 2 declined but chloroplast‐H 2 O 2 further enhanced during recovery after HS over‐acclimation, resulting in the reduced expression of HS‐responsive genes and substantial production of PCD.”
Claim · model-written summary
Further NADPH H2O2 inhibition after over-acclimation inhibited chloroplast‐H 2 O 2.
- Taxon
- tomato seedlings
Solanum lycopersicume7 - Treatment
- further inhibition of NADPH‐H 2 O 2e0
- Outcome
- chloroplast‐H 2 O 2e0
- Direction
- inhibitede0
- Comparator
- HS over‐acclimation treatmente0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e7“the tomato seedlings maintained a constant H 2 O 2 level”
e0“the further inhibition of NADPH‐H 2 O 2 after HS over‐acclimation also inhibited chloroplast‐H 2 O 2 , alleviating the severe PCD and surpassing the MAT of HS over‐acclimation treatment.”
Claim · model-written summary
Further NADPH H2O2 inhibition after over-acclimation alleviating severe PCD.
- Taxon
- tomato seedlings
Solanum lycopersicume7 - Treatment
- further inhibition of NADPH‐H 2 O 2e0
- Outcome
- severe PCDe0
- Direction
- alleviatinge0
- Comparator
- HS over‐acclimation treatmente0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e7“the tomato seedlings maintained a constant H 2 O 2 level”
e0“the further inhibition of NADPH‐H 2 O 2 after HS over‐acclimation also inhibited chloroplast‐H 2 O 2 , alleviating the severe PCD and surpassing the MAT of HS over‐acclimation treatment.”
Claim · model-written summary
Further NADPH H2O2 inhibition after over-acclimation surpassing MAT.
- Taxon
- tomato seedlings
Solanum lycopersicume7 - Treatment
- further inhibition of NADPH‐H 2 O 2e0
- Outcome
- MATe0
- Direction
- surpassinge0
- Comparator
- HS over‐acclimation treatmente0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e7“the tomato seedlings maintained a constant H 2 O 2 level”
e0“the further inhibition of NADPH‐H 2 O 2 after HS over‐acclimation also inhibited chloroplast‐H 2 O 2 , alleviating the severe PCD and surpassing the MAT of HS over‐acclimation treatment.”
Claim · model-written summary
Tomato seedlings maintained constant H2O2 during recovery.
- Taxon
- tomato seedlings
Solanum lycopersicume0 - Outcome
- H 2 O 2 levele0
- Direction
- constante0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“the tomato seedlings maintained a constant H 2 O 2 level during recovery, resulting in stable and lower total H 2 O 2 levels during a tester HS challenge conducted after recovery.”
Claim · model-written summary
Total H2O2 was stable and lower during a later heat challenge.
- Taxon
- tomato seedlings
Solanum lycopersicume0 - Outcome
- total H 2 O 2 levelse0
- Direction
- lowere0
- Qualifier
- during a tester HS challenge conducted after recoverye0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“the tomato seedlings maintained a constant H 2 O 2 level during recovery, resulting in stable and lower total H 2 O 2 levels during a tester HS challenge conducted after recovery.”
Claim · model-written summary
NADPH H2O2 enhancement and chloroplast H2O2 control increased tomato MAT.
- Taxon
- tomato seedlings
Solanum lycopersicume0 - Compound
- NADPH‐H 2 O 2e0
- Outcome
- MATe0
- Direction
- increasee0
- Qualifier
- controlling chloroplast‐H 2 O 2 production during recoverye0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“tomato seedlings increase their MAT by enhancing NADPH‐H 2 O 2 content and controlling chloroplast‐H 2 O 2 production during recovery, which enhances the expression of HS‐responsive genes and balances PCD levels, respectively.”
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