Abstract
• Spermine and ethylene enhance antioxidant defense and upregulates psbA, psbB , and psbC . • Spermine enhances Rubisco activity, nitrogen metabolism and p-NUE. • Controlled ethylene levels are essential for optimal heat stress protection. • Spermine inhibits ACS/ACO expression, hence reducing ethylene overproduction. Global warming-induced heat stress (HS) limits wheat productivity by impairing photosynthesis and nitrogen metabolism. We evaluated the effect of spermine (Spm) and ethephon (ETH; an ethylene donor) in HS tolerance across eight cultivars, and selected PBW343 (heat-tolerant) and PBW17 (heat-sensitive) for a detailed study. Under HS, ROS accumulation and lipid peroxidation increased by 145.6%, while photosynthetic rate and photosynthetic nitrogen use efficiency (p-NUE) declined by 39.3% and 35%, respectively. Spm (100 μM) reduced ROS and lipid peroxidation by 33.9%, enhanced antioxidant enzymes (SOD, APX, GR), upregulated psbA, psbB, and psbC, and suppressed ACS/ACO expression, decreasing ethylene emission by 23.8%. HS increased oxidative stress by increasing H 2 O 2 and MDA levels, but ETH (200 µL L⁻¹) and Spm applied together most effectively reduced oxidative stress under control and heat stress conditions. Combining Spm with the ethylene inhibitor aminoethoxyvinylglycine (AVG; 50 μL L⁻¹) under HS reduced GR activity and photosynthesis, confirming that balanced ethylene signaling is essential for thermoprotection. Our findings reveal that Spm enhances HS tolerance by redirecting S-adenosylmethionine metabolism toward polyamine biosynthesis, stabilizing membranes, detoxifying ROS, and sustaining photosynthesis. This supports the Spm ethylene pathway modulation to improve wheat resilience to HS.
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Claims 13
Claim · model-written summary
Heat stress increased wheat ROS accumulation and lipid peroxidation by 145.6%.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- HSe0
- Outcome
- ROS accumulation and lipid peroxidatione0
- Direction
- increasede0
- Value
- 145.6%e0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Under HS, ROS accumulation and lipid peroxidation increased by 145.6%”
t0“wheat (Triticum aestivum L.)”
Claim · model-written summary
Heat stress reduced wheat photosynthetic rate by 39.3%.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- HSe0
- Outcome
- photosynthetic ratee0
- Direction
- declinede0
- Value
- 39.3%e0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Under HS, ROS accumulation and lipid peroxidation increased by 145.6%, while photosynthetic rate and photosynthetic nitrogen use efficiency (p-NUE) declined by 39.3% and 35%, respectively”
t0“wheat (Triticum aestivum L.)”
Claim · model-written summary
Heat stress reduced wheat photosynthetic nitrogen use efficiency by 35%.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- HSe1
- Outcome
- photosynthetic nitrogen use efficiency (p-NUE)e0
- Direction
- declinede0
- Value
- 35%e0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“photosynthetic nitrogen use efficiency (p-NUE) declined by 39.3% and 35%, respectively”
e1“Under HS, ROS accumulation and lipid peroxidation increased by 145.6%, while photosynthetic rate and photosynthetic nitrogen use efficiency (p-NUE) declined by 39.3% and 35%, respectively”
t0“wheat (Triticum aestivum L.)”
Claim · model-written summary
Spermine reduced wheat ROS and lipid peroxidation by 33.9%.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- Spm (100 μM)e0
- Outcome
- ROS and lipid peroxidatione0
- Direction
- reducede0
- Value
- 33.9%e0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Spm (100 μM) reduced ROS and lipid peroxidation by 33.9%”
t0“wheat (Triticum aestivum L.)”
Claim · model-written summary
Spermine enhanced wheat antioxidant enzymes SOD, APX, and GR.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- Spm (100 μM)e0
- Gene
- SOD, APX, GRe0
- Outcome
- antioxidant enzymes (SOD, APX, GR)e0
- Direction
- enhancede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Spm (100 μM) reduced ROS and lipid peroxidation by 33.9%, enhanced antioxidant enzymes (SOD, APX, GR)”
t0“wheat (Triticum aestivum L.)”
Claim · model-written summary
Spermine upregulated wheat psbA, psbB, and psbC.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- Spm (100 μM)e1
- Gene
- psbA, psbB, and psbCe0
- Outcome
- psbA, psbB, and psbCe0
- Direction
- upregulatede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“upregulated psbA, psbB, and psbC”
e1“Spm (100 μM) reduced ROS and lipid peroxidation by 33.9%, enhanced antioxidant enzymes (SOD, APX, GR), upregulated psbA, psbB, and psbC”
t0“wheat (Triticum aestivum L.)”
Claim · model-written summary
Spermine suppressed wheat ACS/ACO expression.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- Spm (100 μM)e1
- Gene
- ACS/ACOe0
- Outcome
- ACS/ACO expressione0
- Direction
- suppressede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“suppressed ACS/ACO expression”
e1“Spm (100 μM) reduced ROS and lipid peroxidation by 33.9%, enhanced antioxidant enzymes (SOD, APX, GR), upregulated psbA, psbB, and psbC, and suppressed ACS/ACO expression”
t0“wheat (Triticum aestivum L.)”
Claim · model-written summary
Spermine decreased wheat ethylene emission by 23.8%.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- Spm (100 μM)e1
- Outcome
- ethylene emissione0
- Direction
- decreasinge0
- Value
- 23.8%e0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“decreasing ethylene emission by 23.8%”
e1“Spm (100 μM) reduced ROS and lipid peroxidation by 33.9%, enhanced antioxidant enzymes (SOD, APX, GR), upregulated psbA, psbB, and psbC, and suppressed ACS/ACO expression, decreasing ethylene emission by 23.8%”
t0“wheat (Triticum aestivum L.)”
Claim · model-written summary
Heat stress increased wheat H 2 O 2 and MDA levels.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- HSe0
- Outcome
- H 2 O 2 and MDA levelse0
- Direction
- increasinge0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“HS increased oxidative stress by increasing H 2 O 2 and MDA levels”
t0“wheat (Triticum aestivum L.)”
Claim · model-written summary
Combined ethephon and spermine most effectively reduced wheat oxidative stress under control and heat stress conditions.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- ETH (200 µL L⁻¹) and Spm applied togethere0
- Outcome
- oxidative stresse0
- Direction
- reducede0
- Qualifier
- under control and heat stress conditionse0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“ETH (200 µL L⁻¹) and Spm applied together most effectively reduced oxidative stress under control and heat stress conditions”
t0“wheat (Triticum aestivum L.)”
Claim · model-written summary
Spermine with AVG under heat stress reduced wheat GR activity.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- Combining Spm with the ethylene inhibitor aminoethoxyvinylglycine (AVG; 50 μL L⁻¹) under HSe0
- Gene
- GRe0
- Outcome
- GR activitye0
- Direction
- reducede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Combining Spm with the ethylene inhibitor aminoethoxyvinylglycine (AVG; 50 μL L⁻¹) under HS reduced GR activity and photosynthesis”
t0“wheat (Triticum aestivum L.)”
Claim · model-written summary
Spermine with AVG under heat stress reduced wheat photosynthesis.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- Combining Spm with the ethylene inhibitor aminoethoxyvinylglycine (AVG; 50 μL L⁻¹) under HSe0
- Outcome
- photosynthesise0
- Direction
- reducede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Combining Spm with the ethylene inhibitor aminoethoxyvinylglycine (AVG; 50 μL L⁻¹) under HS reduced GR activity and photosynthesis”
t0“wheat (Triticum aestivum L.)”
Claim · model-written summary
The authors attribute spermine-enhanced heat tolerance to redirected S-adenosylmethionine metabolism, membrane stabilization, ROS detoxification, and sustained photosynthesis.
- Taxon
- wheat
Triticum aestivumt0 - Outcome
- HS tolerancee0
- Qualifier
- redirecting S-adenosylmethionine metabolism toward polyamine biosynthesis, stabilizing membranes, detoxifying ROS, and sustaining photosynthesise0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Spm enhances HS tolerance by redirecting S-adenosylmethionine metabolism toward polyamine biosynthesis, stabilizing membranes, detoxifying ROS, and sustaining photosynthesis”
t0“wheat (Triticum aestivum L.)”
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See all 28 papers →8 of 10 groups shown: named compounds and organisms first, then groups whose results differ (checked claim by claim), then those with positive and negative results, then the smaller groups.
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.