Paper · 2026 · International Journal of Molecular Sciences

Physiological, Morphological, and Molecular Evaluation of Wheat Under Single (Drought, Salt, Heat) and Combined (Drought–Heat, Salt–Heat) Stress

Conghui Li; Xiaorui Guo; Lijuan Zhao; et al.

13
claims
30
verbatim quotes
0
citations
2
research areas
✓
Open access · cc-by

Abstract

L.), a key grain food crop worldwide, faces increasing threats from combined abiotic stresses exacerbated by climate change. However, the comprehensive effects of drought, salinity, and high-temperature pressure on wheat seedlings remain poorly understood. Using the cultivar "Yannong 1212", we conducted hydroponic experiments to investigate the physiological, morphological, antioxidant, osmoregulatory, membrane lipid peroxidation, and molecular responses of wheat seedlings to single and combined stresses, and then conducted multivariate statistical analyses. The results showed that drought or salt stress inhibited seed germination in a concentration-dependent manner. However, the combined stresses significantly inhibited germination and seedling growth, leading to leaf chlorosis, chlorophyll degradation, stomatal closure, and chloroplast damage. Physiologically, the combined effect of multiple stresses induced excessive ROS and MDA accumulation, promoted proline and soluble sugar synthesis, and triggered the dynamic responses of antioxidant enzymes. Drought stress increased SOD, POD, and CAT activities, whereas salt stress had the opposite effect, and combined stresses further increased SOD and POD activities, but reduced CAT activity. Additionally, stress-responsive genes were rapidly upregulated. Multivariate analyses confirmed that the combined stress of drought and heat was the most damaging. These findings explain the synergistic damage mechanisms of combined stresses, providing a theoretical basis for genetic improvement of wheat's stress tolerance.

Abstract from the paper’s OpenAlex record; open copy licensed cc-by. Read at the source ↗

The text the model read, with the source record’s inline markup (italics, sub- and superscripts) rendered and its character entities decoded; quote offsets refer to the raw text.

Claims 13

Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Drought inhibited germination in a concentration-dependent manner.

Treatment
droughte0
Outcome
seed germinatione0
Direction
inhibitede0
Qualifier
concentration-dependent mannere0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“drought or salt stress inhibited seed germination in a concentration-dependent manner”

✓ located verbatim in the source · abstract, characters 589–674 · show in text ↑
t0

“Wheat Under Single”

✓ located verbatim in the source · title, characters 58–76 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Salt stress inhibited germination in a concentration-dependent manner.

Treatment
salt stresse0
Outcome
seed germinatione0
Direction
inhibitede0
Qualifier
concentration-dependent mannere0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“drought or salt stress inhibited seed germination in a concentration-dependent manner”

✓ located verbatim in the source · abstract, characters 589–674 · show in text ↑
t0

“Wheat Under Single”

✓ located verbatim in the source · title, characters 58–76 · show in text ↑
Intervention effectobservedoriginal resultnegativesignificant (as stated)measured

Claim · model-written summary

Combined stresses significantly inhibited germination and seedling growth.

Treatment
combined stressese0
Outcome
germination and seedling growthe0
Direction
inhibitede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“the combined stresses significantly inhibited germination and seedling growth”

✓ located verbatim in the source · abstract, characters 685–762 · show in text ↑
t0

“Wheat Under Single”

✓ located verbatim in the source · title, characters 58–76 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Combined stresses caused chlorosis, chlorophyll degradation, stomatal closure and chloroplast damage.

Treatment
combined stressese1
Outcome
leaf chlorosis, chlorophyll degradation, stomatal closure, and chloroplast damagee0

Not reportedComparatorDirection

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“leading to leaf chlorosis, chlorophyll degradation, stomatal closure, and chloroplast damage”

✓ located verbatim in the source · abstract, characters 764–856 · show in text ↑
t0

“Wheat Under Single (Drought, Salt, Heat) and Combined (Drought–Heat, Salt–Heat) Stress”

✓ located verbatim in the source · title, characters 58–144 · show in text ↑
e1

“the combined stresses significantly inhibited germination and seedling growth”

✓ located verbatim in the source · abstract, characters 685–762 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Combined stresses induced excessive ROS and MDA accumulation.

Treatment
multiple stressese0
Outcome
ROS and MDA accumulatione0
Direction
excessivee0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“combined effect of multiple stresses induced excessive ROS and MDA accumulation”

✓ located verbatim in the source · abstract, characters 879–958 · show in text ↑
t0

“Wheat Under Single”

✓ located verbatim in the source · title, characters 58–76 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Combined stresses promoted proline and soluble sugar synthesis.

Treatment
combined stressese1
Outcome
proline and soluble sugar synthesise0
Direction
promotede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“promoted proline and soluble sugar synthesis”

✓ located verbatim in the source · abstract, characters 960–1004 · show in text ↑
t0

“Wheat Under Single (Drought, Salt, Heat) and Combined (Drought–Heat, Salt–Heat) Stress”

✓ located verbatim in the source · title, characters 58–144 · show in text ↑
e1

“the combined stresses significantly inhibited germination and seedling growth”

✓ located verbatim in the source · abstract, characters 685–762 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Combined stresses triggered dynamic antioxidant enzyme responses.

Treatment
combined stressese1
Outcome
dynamic responses of antioxidant enzymese0

Not reportedComparatorDirection

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“triggered the dynamic responses of antioxidant enzymes”

✓ located verbatim in the source · abstract, characters 1010–1064 · show in text ↑
t0

“Wheat Under Single (Drought, Salt, Heat) and Combined (Drought–Heat, Salt–Heat) Stress”

✓ located verbatim in the source · title, characters 58–144 · show in text ↑
e1

“the combined stresses significantly inhibited germination and seedling growth”

✓ located verbatim in the source · abstract, characters 685–762 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Drought increased SOD, POD and CAT activities.

Treatment
Drought stresse0
Outcome
SOD, POD, and CAT activitiese0
Direction
increasede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought stress increased SOD, POD, and CAT activities”

✓ located verbatim in the source · abstract, characters 1066–1119 · show in text ↑
t0

“Wheat Under Single”

✓ located verbatim in the source · title, characters 58–76 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Salt stress decreased SOD, POD and CAT activities.

Treatment
salt stresse0
Outcome
SOD, POD, and CAT activitiese0
Direction
decreaseinferrede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought stress increased SOD, POD, and CAT activities, whereas salt stress had the opposite effect”

✓ located verbatim in the source · abstract, characters 1066–1164 · show in text ↑
t0

“Wheat Under Single”

✓ located verbatim in the source · title, characters 58–76 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Combined stresses further increased SOD and POD activity.

Treatment
combined stressese0
Outcome
SOD and POD activitiese0
Direction
increasede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“combined stresses further increased SOD and POD activities”

✓ located verbatim in the source · abstract, characters 1170–1228 · show in text ↑
t0

“Wheat Under Single”

✓ located verbatim in the source · title, characters 58–76 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Combined stresses reduced CAT activity.

Treatment
combined stressese1
Outcome
CAT activitye0
Direction
reducede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“but reduced CAT activity”

✓ located verbatim in the source · abstract, characters 1230–1254 · show in text ↑
t0

“Wheat Under Single (Drought, Salt, Heat) and Combined (Drought–Heat, Salt–Heat) Stress”

✓ located verbatim in the source · title, characters 58–144 · show in text ↑
e1

“the combined stresses significantly inhibited germination and seedling growth”

✓ located verbatim in the source · abstract, characters 685–762 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Stress-responsive genes were rapidly upregulated.

Outcome
stress-responsive genese0
Direction
upregulatede0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“stress-responsive genes were rapidly upregulated”

✓ located verbatim in the source · abstract, characters 1270–1318 · show in text ↑
t0

“Wheat Under Single”

✓ located verbatim in the source · title, characters 58–76 · show in text ↑
Observationobservedoriginal resultnegativemeasured

Claim · model-written summary

Multivariate analysis identified combined drought and heat as most damaging.

Treatment
combined stress of drought and heate0
Outcome
most damaginge0
Direction
most damaginge0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Multivariate analyses confirmed that the combined stress of drought and heat was the most damaging”

✓ located verbatim in the source · abstract, characters 1320–1418 · show in text ↑
t0

“Wheat Under Single”

✓ located verbatim in the source · title, characters 58–76 · show in text ↑

salinity · physiology: antioxidant enzymes · Triticum aestivum (wheat)Results differ

48 papers: 21 positive, 27 negative or no effect

26 papers in the same direction as this paper · 21 in the opposite direction

  1. positive
  2. positive
  3. positive

44 papers more in this group

See all 48 papers →

drought · physiology: antioxidant enzymes · Triticum aestivum (wheat)Positive and negative results

79 papers: 49 positive, 30 negative or no effect

48 papers in the same direction as this paper · 30 in the opposite direction

  1. negative
  2. negative
  3. negative

75 papers more in this group

See all 79 papers →

salinity · growth · Triticum aestivum (wheat)Positive and negative results

377 papers: 49 positive, 328 negative or no effect

327 papers in the same direction as this paper · 49 in the opposite direction

  1. negative
  2. positive
  3. positive

373 papers more in this group

See all 377 papers →

drought · growth · Triticum aestivum (wheat)Positive and negative results

431 papers: 86 positive, 345 negative or no effect

344 papers in the same direction as this paper · 86 in the opposite direction

  1. negative
  2. positive
  3. positive

427 papers more in this group

See all 431 papers →

abiotic · physiology: oxidative · Triticum aestivum (wheat)Results agree

2 papers: 0 positive, 2 negative or no effect

1 paper in the same direction as this paper

  1. negative
See all 2 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.