Abstract
Heat stress decreases crop growth and yield worldwide. Spermidine (Spd) is a small aliphatic amine and acts as a ubiquitous regulator for plant growth, development and stress tolerance. Objectives of this study were to determine effects of exogenous Spd on changes in endogenous polyamine (PA) and γ-aminobutyric acid (GABA) metabolism, oxidative damage, senescence and heat shock protein (HSP) expression in white clover subjected to heat stress. Physiological and molecular methods, including colorimetric assay, high performance liquid chromatography and qRT-PCR, were applied. Results showed that exogenous Spd significantly alleviated heat-induced stress damage. Application of Spd not only increased endogenous putrescine, Spd, spermine and total PA accumulation, but also accelerated PA oxidation and improved glutamic acid decarboxylase activity, leading to GABA accumulation in leaves under heat stress. The Spd-pretreated white clover maintained a significantly higher chlorophyll (Chl) content than untreated plants under heat stress, which could be related to the roles of Spd in up-regulating genes encoding Chl synthesis (PBGD and Mg-CHT) and maintaining reduced Chl degradation (PaO and CHLASE) during heat stress. In addition, Spd up-regulated HSP70, HSP70B and HSP70-5 expression, which might function in stabilizing denatured proteins and helping proteins to folding correctly in white clover under high temperature stress. In summary, exogenous Spd treatment improves the heat tolerance of white clover by altering endogenous PA and GABA content and metabolism, enhancing the antioxidant system and HSP expression and slowing leaf senescence related to an increase in Chl biosynthesis and a decrease in Chl degradation during heat stress.
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Claims 11
Claim · model-written summary
Spermidine significantly alleviated heat stress damage.
- Taxon
- white clover
Trifolium repenst0 - Treatment
- exogenous Spde0
- Outcome
- heat-induced stress damagee0
- Direction
- alleviatede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Results showed that exogenous Spd significantly alleviated heat-induced stress damage.”
t0“white clover”
Claim · model-written summary
Spermidine increased endogenous putrescine, Spd, spermine and total polyamines under heat stress.
- Taxon
- white clover
Trifolium repenst0 - Treatment
- Application of Spde0
- Outcome
- endogenous putrescine, Spd, spermine and total PA accumulatione0
- Direction
- increasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Application of Spd not only increased endogenous putrescine, Spd, spermine and total PA accumulation, but also accelerated PA oxidation and improved glutamic acid decarboxylase activity, leading to GABA accumulation in leaves under heat stress.”
t0“white clover”
Claim · model-written summary
Spermidine accelerated polyamine oxidation.
- Taxon
- white clover
Trifolium repenst0 - Treatment
- Application of Spde0
- Outcome
- PA oxidatione0
- Direction
- acceleratede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Application of Spd not only increased endogenous putrescine, Spd, spermine and total PA accumulation, but also accelerated PA oxidation and improved glutamic acid decarboxylase activity, leading to GABA accumulation in leaves under heat stress.”
t0“white clover”
Claim · model-written summary
Spermidine improved glutamic acid decarboxylase activity.
- Taxon
- white clover
Trifolium repenst0 - Treatment
- Application of Spde0
- Outcome
- glutamic acid decarboxylase activitye0
- Direction
- improvede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Application of Spd not only increased endogenous putrescine, Spd, spermine and total PA accumulation, but also accelerated PA oxidation and improved glutamic acid decarboxylase activity, leading to GABA accumulation in leaves under heat stress.”
t0“white clover”
Claim · model-written summary
Spermidine treatment led to GABA accumulation in leaves.
- Taxon
- white clover
Trifolium repenst0 - Organ / stage
- leavese0
- Treatment
- Application of Spde0
- Outcome
- GABA accumulatione0
- Direction
- accumulatione0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Application of Spd not only increased endogenous putrescine, Spd, spermine and total PA accumulation, but also accelerated PA oxidation and improved glutamic acid decarboxylase activity, leading to GABA accumulation in leaves under heat stress.”
t0“white clover”
Claim · model-written summary
Spermidine-pretreated clover had higher chlorophyll content than untreated plants under heat stress.
- Taxon
- white clover
Trifolium repense0 - Treatment
- Spd-pretreatede0
- Outcome
- chlorophyll (Chl) contente0
- Direction
- highere0
- Comparator
- untreated plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The Spd-pretreated white clover maintained a significantly higher chlorophyll (Chl) content than untreated plants under heat stress, which could be related to the roles of Spd in up-regulating genes encoding Chl synthesis (PBGD and Mg-CHT) and maintaining reduced Chl degradation (PaO and CHLASE) during heat stress.”
Claim · model-written summary
Spermidine upregulated chlorophyll synthesis genes PBGD and Mg-CHT.
- Taxon
- white clover
Trifolium repense0 - Treatment
- Spd-pretreatede0
- Gene
- PBGD and Mg-CHTe0
- Outcome
- genes encoding Chl synthesise0
- Direction
- up-regulatinge0
- Comparator
- untreated plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The Spd-pretreated white clover maintained a significantly higher chlorophyll (Chl) content than untreated plants under heat stress, which could be related to the roles of Spd in up-regulating genes encoding Chl synthesis (PBGD and Mg-CHT) and maintaining reduced Chl degradation (PaO and CHLASE) during heat stress.”
Claim · model-written summary
Spermidine maintained reduced chlorophyll degradation.
- Taxon
- white clover
Trifolium repense0 - Treatment
- Spd-pretreatede0
- Gene
- PaO and CHLASEe0
- Outcome
- Chl degradatione0
- Direction
- reducede0
- Comparator
- untreated plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The Spd-pretreated white clover maintained a significantly higher chlorophyll (Chl) content than untreated plants under heat stress, which could be related to the roles of Spd in up-regulating genes encoding Chl synthesis (PBGD and Mg-CHT) and maintaining reduced Chl degradation (PaO and CHLASE) during heat stress.”
Claim · model-written summary
Spermidine upregulated HSP70, HSP70B and HSP70-5 expression.
- Taxon
- white clover
Trifolium repense0 - Treatment
- Spde0
- Outcome
- HSP70, HSP70B and HSP70-5 expressione0
- Direction
- up-regulatede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Spd up-regulated HSP70, HSP70B and HSP70-5 expression, which might function in stabilizing denatured proteins and helping proteins to folding correctly in white clover under high temperature stress.”
Claim · model-written summary
Authors proposed HSP upregulation could stabilize denatured proteins and aid folding.
- Taxon
- white clover
Trifolium repense0 - Gene
- HSP70, HSP70B and HSP70-5e0
- Outcome
- stabilizing denatured proteins and helping proteins to folding correctlye0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Spd up-regulated HSP70, HSP70B and HSP70-5 expression, which might function in stabilizing denatured proteins and helping proteins to folding correctly in white clover under high temperature stress.”
Claim · model-written summary
Authors concluded spermidine improved heat tolerance partly through antioxidant responses and delayed leaf senescence.
- Taxon
- white clover
Trifolium repense0 - Treatment
- exogenous Spd treatmente0
- Outcome
- heat tolerancee0
- Qualifier
- enhancing the antioxidant system and HSP expression and slowing leaf senescencee0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“exogenous Spd treatment improves the heat tolerance of white clover by altering endogenous PA and GABA content and metabolism, enhancing the antioxidant system and HSP expression and slowing leaf senescence related to an increase in Chl biosynthesis and a decrease in Chl degradation during heat stress.”
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spermidine · molecular: expression · Trifolium repens (white clover)Results agree
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See all 6 papers →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.