Abstract
The aim of the present work was to investigate whether loss of germination ability and viability of sunflower ( Helianthus annuus L.) seeds during incubation at a high temperature (45°C) was related to changes in energy metabolism, loss of membrane integrity, and/or changes in lipid composition. Pre‐treatment of seeds at 45°C progressively reduced subsequent germination at the optimal temperature (25°C). Seeds did not germinate at 45°C and almost all of them were dead after 72 h of soaking at this high temperature. This loss of seed viability was associated with a large increase in leakage of K + and total electrolytes into the incubation medium, and with production of malondialdehyde in the embryonic axis and cotyledons, suggesting a loss of membrane integrity probably due to lipid peroxidation. ATP and ADP levels increased sharply during the first hours of imbibition at 45°C, remained high for about 24 h and then decreased. As a consequence, the energy charge followed a similar pattern. If the treatment at 45°C did not exceed 48 h, seeds recovered an apparently normal energy metabolism after transfer to 25°C, even though they lost their ability to germinate at this temperature. Therefore, energy metabolism at the whole embryo level cannot be considered as an indicator of germination ability. Incubation of seeds at 45°C resulted in an increase in triacylglycerols and diacylglycerols without a significant change in their fatty acid composition. It also induced a slight increase in phospholipid content with an increase in C 16:0 , C 18:0 and C 18:1 , but with no change in C 18:2 . In phospholipids, the C 18:2 /C 18:1 and (C 18:1 + C 18:2 )/ (C 16:0 + C 18:0 ) ratios thus declined during treatment at 45°C. The results obtained suggest that deterioration of sunflower seeds during incubation at a high temperature is mainly related to membrane damage and alteration of energy metabolism, and that accumulation of malondialdehyde, which is an index of lipid peroxidation, does not correspond to a decrease in total lipids and phospholipids nor to a significant change in fatty acid composition, except in PL in which the C 18:2 /C 18:1 and (C 18:1 + C 18:2 )/ (C 16:0 + C 18:0 ) ratios slightly declined.
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Claims 18
Claim · model-written summary
Pre-treatment of seeds at 45°C progressively reduced subsequent germination at 25°C.
- Taxon
- sunflower (Helianthus annuus)
Helianthus annuust0 - Treatment
- Pre‐treatment of seeds at 45°Ce1
- Outcome
- subsequent germinatione1
- Direction
- reducede1
- Qualifier
- progressivelye1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Decrease in sunflower (Helianthus annuus) seed viability”
e1“Pre‐treatment of seeds at 45°C progressively reduced subsequent germination at the optimal temperature (25°C)”
Claim · model-written summary
Seeds did not germinate at 45°C and almost all were dead after 72 h of soaking at high temperature.
- Taxon
- sunflower (Helianthus annuus)
Helianthus annuust0 - Treatment
- soaking at this high temperaturee2
- Outcome
- deade2
- Qualifier
- after 72 h of soakinge2
Not reportedComparatorDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Decrease in sunflower (Helianthus annuus) seed viability”
e2“Seeds did not germinate at 45°C and almost all of them were dead after 72 h of soaking at this high temperature”
Claim · model-written summary
Loss of seed viability was associated with a large increase in leakage of K+ and total electrolytes into the incubation medium.
- Taxon
- sunflower (Helianthus annuus)
Helianthus annuust0 - Outcome
- leakage of K + and total electrolytese3
- Direction
- increasee3
- Setting
- incubation mediume3
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Decrease in sunflower (Helianthus annuus) seed viability”
e3“This loss of seed viability was associated with a large increase in leakage of K + and total electrolytes into the incubation medium”
Claim · model-written summary
Malondialdehyde production was observed in the embryonic axis and cotyledons.
- Taxon
- sunflower (Helianthus annuus)
Helianthus annuust0 - Organ / stage
- embryonic axis and cotyledonse4
- Outcome
- production of malondialdehydee4
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Decrease in sunflower (Helianthus annuus) seed viability”
e4“with production of malondialdehyde in the embryonic axis and cotyledons, suggesting a loss of membrane integrity probably due to lipid peroxidation”
Claim · model-written summary
Loss of membrane integrity is suggested to be probably due to lipid peroxidation.
- Taxon
- sunflower (Helianthus annuus)
Helianthus annuust0 - Outcome
- loss of membrane integritye4
- Qualifier
- probably due to lipid peroxidatione4
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Decrease in sunflower (Helianthus annuus) seed viability”
e4“with production of malondialdehyde in the embryonic axis and cotyledons, suggesting a loss of membrane integrity probably due to lipid peroxidation”
Claim · model-written summary
ATP and ADP levels increased sharply during the first hours of imbibition at 45°C.
- Taxon
- sunflower (Helianthus annuus)
Helianthus annuust0 - Treatment
- imbibition at 45°Ce5
- Outcome
- ATP and ADP levelse5
- Direction
- increasede5
- Qualifier
- sharply during the first hours of imbibitione5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Decrease in sunflower (Helianthus annuus) seed viability”
e5“ATP and ADP levels increased sharply during the first hours of imbibition at 45°C, remained high for about 24 h and then decreased”
Claim · model-written summary
ATP and ADP levels remained high for about 24 h and then decreased during imbibition at 45°C.
- Taxon
- sunflower (Helianthus annuus)
Helianthus annuust0 - Treatment
- imbibition at 45°Ce5
- Outcome
- ATP and ADP levelse5
- Direction
- decreasede5
- Qualifier
- remained high for about 24 he5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Decrease in sunflower (Helianthus annuus) seed viability”
e5“ATP and ADP levels increased sharply during the first hours of imbibition at 45°C, remained high for about 24 h and then decreased”
Claim · model-written summary
The energy charge followed a similar pattern to the ATP and ADP levels.
- Taxon
- sunflower (Helianthus annuus)
Helianthus annuust0 - Outcome
- energy chargee6
- Qualifier
- followed a similar patterne6
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Decrease in sunflower (Helianthus annuus) seed viability”
e6“As a consequence, the energy charge followed a similar pattern”
Claim · model-written summary
Seeds recovered an apparently normal energy metabolism after transfer to 25°C when the 45°C treatment did not exceed 48 h.
- Taxon
- sunflower (Helianthus annuus)
Helianthus annuust0 - Outcome
- energy metabolisme7
- Qualifier
- did not exceed 48 he7
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Decrease in sunflower (Helianthus annuus) seed viability”
e7“If the treatment at 45°C did not exceed 48 h, seeds recovered an apparently normal energy metabolism after transfer to 25°C”
Claim · model-written summary
Energy metabolism at the whole embryo level cannot be considered an indicator of germination ability.
- Taxon
- sunflower (Helianthus annuus)
Helianthus annuust0 - Outcome
- energy metabolism at the whole embryo levele4
- Qualifier
- cannot be considered as an indicator of germination abilitye4
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Decrease in sunflower (Helianthus annuus) seed viability”
e4“Therefore, energy metabolism at the whole embryo level cannot be considered as an indicator of germination ability.”
Claim · model-written summary
Incubation at 45°C increased triacylglycerols and diacylglycerols.
- Taxon
- sunflower (Helianthus annuus)
Helianthus annuust0 - Treatment
- Incubation of seeds at 45°Ce5
- Outcome
- triacylglycerols and diacylglycerolse5
- Direction
- increasee5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Decrease in sunflower (Helianthus annuus) seed viability”
e5“Incubation of seeds at 45°C resulted in an increase in triacylglycerols and diacylglycerols without a significant change in their fatty acid composition”
Claim · model-written summary
Incubation at 45°C did not significantly change the fatty acid composition of total lipids.
- Taxon
- sunflower (Helianthus annuus)
Helianthus annuust0 - Treatment
- Incubation of seeds at 45°Ce5
- Outcome
- fatty acid compositione5
- Direction
- no_changeinferrede5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Decrease in sunflower (Helianthus annuus) seed viability”
e5“Incubation of seeds at 45°C resulted in an increase in triacylglycerols and diacylglycerols without a significant change in their fatty acid composition”
Claim · model-written summary
Incubation at 45°C induced a slight increase in phospholipid content.
- Taxon
- sunflower (Helianthus annuus)
Helianthus annuust0 - Treatment
- Incubation of seeds at 45°Ce5
- Outcome
- phospholipid contente6
- Direction
- increasee6
- Qualifier
- slighte6
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Decrease in sunflower (Helianthus annuus) seed viability”
e5“Incubation of seeds at 45°C resulted in an increase in triacylglycerols and diacylglycerols without a significant change in their fatty acid composition”
e6“It also induced a slight increase in phospholipid content with an increase in C 16:0 , C 18:0 and C 18:1 , but with no change in C 18:2”
Claim · model-written summary
Incubation at 45°C increased the phospholipid C 16:0, C 18:0 and C 18:1 fatty acids.
- Taxon
- sunflower (Helianthus annuus)
Helianthus annuust0 - Treatment
- Incubation of seeds at 45°Ce5
- Outcome
- C 16:0 , C 18:0 and C 18:1e6
- Direction
- increasee6
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Decrease in sunflower (Helianthus annuus) seed viability”
e5“Incubation of seeds at 45°C resulted in an increase in triacylglycerols and diacylglycerols without a significant change in their fatty acid composition”
e6“It also induced a slight increase in phospholipid content with an increase in C 16:0 , C 18:0 and C 18:1 , but with no change in C 18:2”
Claim · model-written summary
Incubation at 45°C did not change the phospholipid C 18:2 fatty acid.
- Taxon
- sunflower (Helianthus annuus)
Helianthus annuust0 - Treatment
- Incubation of seeds at 45°Ce5
- Outcome
- C 18:2e6
- Direction
- no changee6
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Decrease in sunflower (Helianthus annuus) seed viability”
e5“Incubation of seeds at 45°C resulted in an increase in triacylglycerols and diacylglycerols without a significant change in their fatty acid composition”
e6“It also induced a slight increase in phospholipid content with an increase in C 16:0 , C 18:0 and C 18:1 , but with no change in C 18:2”
Claim · model-written summary
In phospholipids, the C 18:2/C 18:1 and (C 18:1 + C 18:2)/(C 16:0 + C 18:0) ratios declined during treatment at 45°C.
- Taxon
- sunflower (Helianthus annuus)
Helianthus annuust0 - Treatment
- during treatment at 45°Ce7
- Outcome
- C 18:2 /C 18:1 and (C 18:1 + C 18:2 )/ (C 16:0 + C 18:0 ) ratiose7
- Direction
- declinede7
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Decrease in sunflower (Helianthus annuus) seed viability”
e7“In phospholipids, the C 18:2 /C 18:1 and (C 18:1 + C 18:2 )/ (C 16:0 + C 18:0 ) ratios thus declined during treatment at 45°C”
Claim · model-written summary
Deterioration of sunflower seeds during high-temperature incubation is mainly related to membrane damage and alteration of energy metabolism.
- Taxon
- sunflower (Helianthus annuus)
Helianthus annuust0 - Outcome
- deterioration of sunflower seedse4
- Qualifier
- mainly related to membrane damage and alteration of energy metabolisme4
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Decrease in sunflower (Helianthus annuus) seed viability”
e4“The results obtained suggest that deterioration of sunflower seeds during incubation at a high temperature is mainly related to membrane damage and alteration of energy metabolism”
Claim · model-written summary
Malondialdehyde accumulation did not correspond to a decrease in total lipids and phospholipids.
- Taxon
- sunflower (Helianthus annuus)
Helianthus annuust0 - Compound
- malondialdehydee5
- Outcome
- total lipids and phospholipidse5
- Direction
- decreasee5
- Qualifier
- does not corresponde5
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Decrease in sunflower (Helianthus annuus) seed viability”
e5“accumulation of malondialdehyde, which is an index of lipid peroxidation, does not correspond to a decrease in total lipids and phospholipids”