Abstract
High temperatures occurring during grain filling are known to affect wheat grain yield and quality considerably. In this paper we report the results of experiments carried out with two cultivars of bread wheat ( Triticum aestivum L.) and two cultivars of durum wheat ( Triticum durum Desf.). The plants, cultivated in pots, were subjected to 13 heat treatments (temperature up to 40°C) differing in duration and timing and starting 7 days after anthesis. Heat treatments were applied by temporary transfer of the pots to a glasshouse where the temperature rose to 40°C as a consequence of solar radiation for periods ranging from 5 to 30 days. The applied heat shocks substantially affected dry matter and protein accumulation in the different parts of the plant. Early heat shock (5 days with a total of 18 h of temperature in the range 35–40°C) caused a small reduction of kernel mass and no effect on protein per kernel; the damage was greater in the central and in the final stage of grain filling. Plants subjected to a progressive increase of temperature, or to an early heat shock, acquired thermotolerance to further heat shocks. Continuous exposure to very high temperatures from 27 days after pollination to maturity did not negatively affect grain yield and it facilitated the remobilisation of nitrogen from vegetative to reproductive organs. Rheological properties were severely affected by heat shocks at all stages of grain filling: 5 days of heat shock were sufficient to reduce mixing tolerance by 40–60%. These variations in rheological properties were accompanied by modification of the level of protein aggregation: soluble polymeric proteins and low molecular weight gliadins progressively increased according to the intensity of the stress, while insoluble polymeric proteins decreased. Our experiments, carried out in conditions close to the Mediterranean climate, indicate that the occurrence of very high temperature in the range 35–40°C during grain filling substantially affects dry matter and protein accumulation in the different parts of the plant. The formation of the complex protein aggregates responsible for positive dough mixing properties is significantly reduced by very high temperature. When heat shock came late in grain filling, grain yield and protein concentration were not negatively affected but a ‘dough weakening’ effect, which may reduce the commercial value of the production, is to be expected.
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Claims 13
Claim · model-written summary
Heat shocks affected dry matter and protein accumulation.
- Taxon
- Wheat
Triticum aestivumt0 - Treatment
- heat shockse0
- Outcome
- dry matter and protein accumulatione0
Not reportedComparatorDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Wheat”
e0“The applied heat shocks substantially affected dry matter and protein accumulation in the different parts of the plant.”
Claim · model-written summary
Early heat shock slightly reduced kernel mass.
- Taxon
- Wheat
Triticum aestivumt0 - Treatment
- Early heat shock (5 days with a total of 18 h of temperature in the range 35–40°C)e0
- Outcome
- kernel masse0
- Direction
- reductione0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Wheat”
e0“Early heat shock (5 days with a total of 18 h of temperature in the range 35–40°C) caused a small reduction of kernel mass”
Claim · model-written summary
Early heat shock had no effect on protein per kernel.
- Taxon
- Wheat
Triticum aestivumt0 - Outcome
- protein per kernele0
- Direction
- no effecte0
Not reportedTreatmentComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Wheat”
e0“no effect on protein per kernel”
Claim · model-written summary
Damage was greater for shocks in central and final grain filling.
- Taxon
- Wheat
Triticum aestivumt0 - Treatment
- central and in the final stage of grain fillinge0
- Outcome
- damagee0
- Direction
- greatere0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Wheat”
e0“the damage was greater in the central and in the final stage of grain filling”
Claim · model-written summary
Progressively heated plants acquired thermotolerance to later heat shocks.
- Taxon
- Wheat
Triticum aestivumt0 - Treatment
- progressive increase of temperaturee0
- Outcome
- thermotolerance to further heat shockse0
Not reportedComparatorDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Wheat”
e0“Plants subjected to a progressive increase of temperature, or to an early heat shock, acquired thermotolerance to further heat shocks.”
Claim · model-written summary
Plants given an early heat shock acquired thermotolerance to later shocks.
- Taxon
- Wheat
Triticum aestivumt0 - Treatment
- early heat shocke0
- Outcome
- thermotolerance to further heat shockse0
Not reportedComparatorDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Wheat”
e0“Plants subjected to a progressive increase of temperature, or to an early heat shock, acquired thermotolerance to further heat shocks.”
Claim · model-written summary
Continuous late high temperature did not negatively affect grain yield.
- Taxon
- Wheat
Triticum aestivumt0 - Treatment
- Continuous exposure to very high temperatures from 27 days after pollination to maturitye0
- Outcome
- grain yielde0
- Direction
- did not negatively affecte0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Wheat”
e0“Continuous exposure to very high temperatures from 27 days after pollination to maturity did not negatively affect grain yield”
Claim · model-written summary
Continuous late high temperature facilitated nitrogen remobilization.
- Taxon
- Wheat
Triticum aestivumt0 - Outcome
- remobilisation of nitrogene0
- Direction
- facilitatede0
Not reportedTreatmentComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Wheat”
e0“it facilitated the remobilisation of nitrogen from vegetative to reproductive organs”
Claim · model-written summary
Five days of heat shock reduced mixing tolerance by 40–60%.
- Taxon
- Wheat
Triticum aestivumt0 - Treatment
- 5 days of heat shocke0
- Outcome
- mixing tolerancee0
- Direction
- reducee0
- Value
- 40–60%e0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Wheat”
e0“5 days of heat shock were sufficient to reduce mixing tolerance by 40–60%”
Claim · model-written summary
Soluble polymeric proteins and low molecular weight gliadins increased with stress intensity.
- Taxon
- Wheat
Triticum aestivumt0 - Treatment
- intensity of the stresse0
- Outcome
- soluble polymeric proteins and low molecular weight gliadinse0
- Direction
- increasede0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Wheat”
e0“soluble polymeric proteins and low molecular weight gliadins progressively increased according to the intensity of the stress”
Claim · model-written summary
Insoluble polymeric proteins decreased with heat stress intensity.
- Taxon
- Wheat
Triticum aestivumt0 - Outcome
- insoluble polymeric proteinse0
- Direction
- decreasede0
Not reportedTreatment
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Wheat”
e0“while insoluble polymeric proteins decreased”
Claim · model-written summary
Very high temperature significantly reduced formation of complex protein aggregates.
- Taxon
- Wheat
Triticum aestivumt0 - Treatment
- very high temperaturee0
- Outcome
- formation of the complex protein aggregatese0
- Direction
- reducede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Wheat”
e0“The formation of the complex protein aggregates responsible for positive dough mixing properties is significantly reduced by very high temperature.”
Claim · model-written summary
Late heat shock did not negatively affect protein concentration.
- Taxon
- Wheat
Triticum aestivumt0 - Treatment
- heat shock came late in grain fillinge0
- Outcome
- protein concentratione0
- Direction
- not negatively affectede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Wheat”
e0“When heat shock came late in grain filling, grain yield and protein concentration were not negatively affected”
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