Abstract
Drought and heat stress are two major constraints that limit chickpea (Cicer arietinum L.) yield, particularly during seed filling. The present study aimed (i) to assess the individual and combined effects of drought and heat stress on oxidative metabolism during seed filling, and (ii) to determine any genetic variation in oxidative metabolism among genotypes differing in drought and heat tolerance and sensitivity. The plants were raised in outdoor conditions with two different times of sowing, one in November (normal-sown, temperatures <32°C-20°C (day–night) during seed filling), and the other in February (late-sown, temperatures >32°C-20°C (day–night) during seed filling). Plants were regularly irrigated to prevent any water shortage until the water treatments were applied. At both sowing times, the drought treatment was applied during seed filling (at ~75% podding) by withholding water from half of the pots until the relative leaf water content (RLWC) of leaves on the top three branches reached 42–45%, whereas leaves in the fully irrigated control plants were maintained at RLWC 85–90%. Drought-stressed plants were then rewatered and maintained under fully irrigated conditions until maturity. Several biochemical parameters were measured on the leaves and seeds at the end of the stress treatments, and seed yield and aboveground biomass were measured at maturity. Individual and combined stresses damaged membranes, and decreased PSII function and leaf chlorophyll content, more so under the combined stress treatment. The levels of oxidative molecules (malondialdehyde (MDA) and H2O2) markedly increased compared with the control plants in all stress treatments, especially across genotypes in the combined heat + drought stress treatment (increases in leaves: MDA 5.4–8.4-fold and H2O2 5.1–7.1-fold; in seeds: MDA 1.9–3.3-fold and H2O2 3.8–7.9-fold). The enzymatic and non-enzymatic antioxidants related to oxidative metabolism increased under individual stress treatments but decreased in the combined heat + drought stress treatment. Leaves had higher oxidative damage than seeds, and this likely inhibited their photosynthetic efficiency. Yields were reduced more by drought stress than by heat stress, with the lowest yields in the combined heat + drought stress treatment. Heat- and drought-tolerant genotypes suffered less damage and had higher yields than the heat- and drought-sensitive genotypes under the individual and combined stress treatments, suggesting partial cross-tolerance in these genotypes. A drought-tolerant genotype ICC8950 produced more seed yield under the combined heat + drought stress than other genotypes, and this was associated with low oxidative damage in leaves and seeds.
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Claims 17
Claim · model-written summary
Individual and combined heat and drought stresses damaged membranes in chickpea.
- Taxon
- chickpea
Cicer arietinume2 - Treatment
- Individual and combined stressese1
- Outcome
- membranese1
Not reportedComparatorDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Individual and combined stresses damaged membranes, and decreased PSII function and leaf chlorophyll content, more so under the combined stress treatment”
e2“Effects of individual and combined heat and drought stress during seed filling on the oxidative metabolism and yield of chickpea (Cicer arietinum) genotypes”
Claim · model-written summary
Individual and combined stresses decreased PSII function and leaf chlorophyll content in chickpea, more under the combined stress.
- Taxon
- chickpea
Cicer arietinume2 - Treatment
- Individual and combined stressese1
- Outcome
- PSII function and leaf chlorophyll contente1
- Direction
- decreasede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Individual and combined stresses damaged membranes, and decreased PSII function and leaf chlorophyll content, more so under the combined stress treatment”
e2“Effects of individual and combined heat and drought stress during seed filling on the oxidative metabolism and yield of chickpea (Cicer arietinum) genotypes”
Claim · model-written summary
Leaf malondialdehyde increased 5.4–8.4-fold under heat and drought stress relative to control plants.
- Taxon
- chickpea
Cicer arietinume4 - Organ / stage
- leavese2
- Treatment
- combined heat + drought stress treatmente1
- Outcome
- malondialdehyde (MDA)e1
- Direction
- increasede1
- Value
- 5.4–8.4-folde2
- Comparator
- control plantse1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“The levels of oxidative molecules (malondialdehyde (MDA) and H2O2) markedly increased compared with the control plants in all stress treatments, especially across genotypes in the combined heat + drought stress treatment”
e2“increases in leaves: MDA 5.4–8.4-fold and H2O2 5.1–7.1-fold”
e4“Effects of individual and combined heat and drought stress during seed filling on the oxidative metabolism and yield of chickpea (Cicer arietinum) genotypes”
Claim · model-written summary
Leaf hydrogen peroxide increased 5.1–7.1-fold under heat and drought stress relative to control plants.
- Taxon
- chickpea
Cicer arietinume4 - Organ / stage
- leavese2
- Treatment
- combined heat + drought stress treatmente1
- Outcome
- H2O2e1
- Direction
- increasede1
- Value
- 5.1–7.1-folde2
- Comparator
- control plantse1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“The levels of oxidative molecules (malondialdehyde (MDA) and H2O2) markedly increased compared with the control plants in all stress treatments, especially across genotypes in the combined heat + drought stress treatment”
e2“increases in leaves: MDA 5.4–8.4-fold and H2O2 5.1–7.1-fold”
e4“Effects of individual and combined heat and drought stress during seed filling on the oxidative metabolism and yield of chickpea (Cicer arietinum) genotypes”
Claim · model-written summary
Seed malondialdehyde increased 1.9–3.3-fold under heat and drought stress relative to control plants.
- Taxon
- chickpea
Cicer arietinume4 - Organ / stage
- seedse2
- Treatment
- combined heat + drought stress treatmente1
- Outcome
- malondialdehyde (MDA)e1
- Direction
- increasede1
- Value
- 1.9–3.3-folde2
- Comparator
- control plantse1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“The levels of oxidative molecules (malondialdehyde (MDA) and H2O2) markedly increased compared with the control plants in all stress treatments, especially across genotypes in the combined heat + drought stress treatment”
e2“in seeds: MDA 1.9–3.3-fold and H2O2 3.8–7.9-fold”
e4“Effects of individual and combined heat and drought stress during seed filling on the oxidative metabolism and yield of chickpea (Cicer arietinum) genotypes”
Claim · model-written summary
Seed hydrogen peroxide increased 3.8–7.9-fold under heat and drought stress relative to control plants.
- Taxon
- chickpea
Cicer arietinume4 - Organ / stage
- seedse2
- Treatment
- combined heat + drought stress treatmente1
- Outcome
- H2O2e1
- Direction
- increasede1
- Value
- 3.8–7.9-folde2
- Comparator
- control plantse1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“The levels of oxidative molecules (malondialdehyde (MDA) and H2O2) markedly increased compared with the control plants in all stress treatments, especially across genotypes in the combined heat + drought stress treatment”
e2“in seeds: MDA 1.9–3.3-fold and H2O2 3.8–7.9-fold”
e4“Effects of individual and combined heat and drought stress during seed filling on the oxidative metabolism and yield of chickpea (Cicer arietinum) genotypes”
Claim · model-written summary
Antioxidants related to oxidative metabolism increased in chickpea under individual stress treatments.
- Taxon
- chickpea
Cicer arietinume2 - Treatment
- individual stress treatmentse1
- Outcome
- enzymatic and non-enzymatic antioxidantse1
- Direction
- increasede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“The enzymatic and non-enzymatic antioxidants related to oxidative metabolism increased under individual stress treatments”
e2“Effects of individual and combined heat and drought stress during seed filling on the oxidative metabolism and yield of chickpea (Cicer arietinum) genotypes”
Claim · model-written summary
Antioxidants related to oxidative metabolism decreased in chickpea under the combined heat + drought stress.
- Taxon
- chickpea
Cicer arietinume2 - Treatment
- combined heat + drought stress treatmente1
- Outcome
- antioxidantse3
- Direction
- decreasede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“but decreased in the combined heat + drought stress treatment”
e2“Effects of individual and combined heat and drought stress during seed filling on the oxidative metabolism and yield of chickpea (Cicer arietinum) genotypes”
e3“The enzymatic and non-enzymatic antioxidants related to oxidative metabolism increased under individual stress treatments”
Claim · model-written summary
Leaves had higher oxidative damage than seeds in chickpea.
- Taxon
- chickpea
Cicer arietinume2 - Organ / stage
- Leavese1
- Outcome
- oxidative damagee1
- Direction
- highere1
- Comparator
- seedse1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Leaves had higher oxidative damage than seeds, and this likely inhibited their photosynthetic efficiency”
e2“Effects of individual and combined heat and drought stress during seed filling on the oxidative metabolism and yield of chickpea (Cicer arietinum) genotypes”
Claim · model-written summary
Oxidative damage in leaves likely inhibited their photosynthetic efficiency in chickpea.
- Taxon
- chickpea
Cicer arietinume2 - Outcome
- photosynthetic efficiencye1
- Qualifier
- oxidative damagee1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Leaves had higher oxidative damage than seeds, and this likely inhibited their photosynthetic efficiency”
e2“Effects of individual and combined heat and drought stress during seed filling on the oxidative metabolism and yield of chickpea (Cicer arietinum) genotypes”
Claim · model-written summary
Yields were reduced more by drought stress than by heat stress in chickpea.
- Taxon
- chickpea
Cicer arietinume2 - Treatment
- drought stresse1
- Outcome
- Yieldse1
- Direction
- reducede1
- Comparator
- heat stresse1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Yields were reduced more by drought stress than by heat stress”
e2“Effects of individual and combined heat and drought stress during seed filling on the oxidative metabolism and yield of chickpea (Cicer arietinum) genotypes”
Claim · model-written summary
The lowest chickpea yields occurred under the combined heat + drought stress.
- Taxon
- chickpea
Cicer arietinume2 - Treatment
- combined heat + drought stress treatmente1
- Outcome
- yieldse1
- Direction
- loweste1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“with the lowest yields in the combined heat + drought stress treatment”
e2“Effects of individual and combined heat and drought stress during seed filling on the oxidative metabolism and yield of chickpea (Cicer arietinum) genotypes”
Claim · model-written summary
Heat- and drought-tolerant chickpea genotypes suffered less damage than sensitive genotypes under individual and combined stress.
- Taxon
- chickpea
Cicer arietinume2 - Material
- heat- and drought-tolerant genotypese1
- Treatment
- individual and combined stress treatmentse1
- Outcome
- damagee1
- Direction
- lesse1
- Comparator
- heat- and drought-sensitive genotypese1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Heat- and drought-tolerant genotypes suffered less damage and had higher yields than the heat- and drought-sensitive genotypes under the individual and combined stress treatments”
e2“Effects of individual and combined heat and drought stress during seed filling on the oxidative metabolism and yield of chickpea (Cicer arietinum) genotypes”
Claim · model-written summary
Heat- and drought-tolerant chickpea genotypes had higher yields than sensitive genotypes under individual and combined stress.
- Taxon
- chickpea
Cicer arietinume2 - Material
- heat- and drought-tolerant genotypese1
- Treatment
- individual and combined stress treatmentse1
- Outcome
- yieldse1
- Direction
- highere1
- Comparator
- heat- and drought-sensitive genotypese1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Heat- and drought-tolerant genotypes suffered less damage and had higher yields than the heat- and drought-sensitive genotypes under the individual and combined stress treatments”
e2“Effects of individual and combined heat and drought stress during seed filling on the oxidative metabolism and yield of chickpea (Cicer arietinum) genotypes”
Claim · model-written summary
The tolerant chickpea genotypes suggest partial cross-tolerance between heat and drought.
- Taxon
- chickpea
Cicer arietinume2 - Material
- these genotypese1
- Outcome
- partial cross-tolerancee1
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“suggesting partial cross-tolerance in these genotypes”
e2“Effects of individual and combined heat and drought stress during seed filling on the oxidative metabolism and yield of chickpea (Cicer arietinum) genotypes”
Claim · model-written summary
Chickpea genotype ICC8950 produced more seed yield under combined heat and drought stress than other genotypes.
- Taxon
- chickpea
Cicer arietinume2 - Material
- ICC8950e1
- Treatment
- combined heat + drought stresse1
- Outcome
- seed yielde1
- Direction
- moree1
- Comparator
- other genotypese1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“A drought-tolerant genotype ICC8950 produced more seed yield under the combined heat + drought stress than other genotypes”
e2“Effects of individual and combined heat and drought stress during seed filling on the oxidative metabolism and yield of chickpea (Cicer arietinum) genotypes”
Claim · model-written summary
Higher seed yield of ICC8950 was associated with low oxidative damage in leaves and seeds.
- Taxon
- chickpea
Cicer arietinume2 - Material
- ICC8950e3
- Organ / stage
- leaves and seedse1
- Outcome
- oxidative damagee1
- Direction
- lowe1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“this was associated with low oxidative damage in leaves and seeds”
e2“Effects of individual and combined heat and drought stress during seed filling on the oxidative metabolism and yield of chickpea (Cicer arietinum) genotypes”
e3“A drought-tolerant genotype ICC8950 produced more seed yield under the combined heat + drought stress than other genotypes”
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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.