Paper · 2026 · Scientific Reports

Drought tolerance mechanisms across C3 and C3–C4 intermediate photosynthetic types revealed by physiological and gene expression profiling

Rabab H. M. Mohamed; Reem Badr; Amani Abdel-Latif; et al.

14
claims
23
verbatim quotes
0
citations
2
research areas
✓
Open access · cc-by

Abstract

Abstract Abiotic stress, particularly drought, significantly reduces crop yields and threatens global agricultural sustainability. This study investigated drought and recovery responses in four plant species with contrasting photosynthetic types: Triticum aestivum (C3), Helianthus annuus (C3), Chenopodium album (intermediate-C4), and Alternanthera brasiliana (C4-like). Drought markedly reduced plant fresh biomass (up to 80% in H. annuus ) and relative water content, particularly in C. album . Oxidative damage intensified, with H. annuus showing the greatest increase in hydrogen peroxide (258%) and C. album exhibiting the highest malondialdehyde accumulation (284%). Antioxidant enzymes were strongly activated; catalase activity increased dramatically in C. album (837%) and H. annuus (630%). Proline levels increased sharply, particularly in T. aestivum and C. album , indicating enhanced osmotic adjustment. Carotenoid content also rose significantly in H. annuus (141%), suggesting photoprotective adaptation. Gene expression analysis revealed upregulation of TaP5CS in T. aestivum , correlating with proline accumulation, and CaHSP26 in C. album , potentially stabilizing photosystem II. Principal component analysis identified catalase activity, root-to-shoot ratio, hydrogen peroxide, and proline as major contributors to drought response variance. These findings highlight species-specific strategies for drought tolerance and recovery, with C3 species showing strong enzymatic and osmotic adjustments, and intermediate-C4 and C4-like species exhibiting greater tissue integrity and ROS balance. This comparative framework provides valuable insights for developing drought-resilient crops.

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 14

Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Drought reduced fresh biomass by up to 80% in Helianthus annuus.

Taxon
H. annuusHelianthus annuuse0
Treatment
Droughte0
Outcome
plant fresh biomasse0
Direction
reducede0
Value
80%e0

Not reportedComparator

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

Helianthus annuus
Helianthus annuus · TaxonJon Sullivan · Public domain · Wikimedia Commons ↗

Verbatim

e0

“Drought markedly reduced plant fresh biomass (up to 80% in H. annuus ) and relative water content, particularly in C. album”

✓ located verbatim in the source · abstract, characters 372–495 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Drought reduced relative water content, particularly in Chenopodium album.

Taxon
C. albumChenopodium albume0
Treatment
Droughte0
Outcome
relative water contente0
Direction
reducede0

Not reportedComparator

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

Chenopodium album
Chenopodium album · TaxonOlivier Pichard · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought markedly reduced plant fresh biomass (up to 80% in H. annuus ) and relative water content, particularly in C. album”

✓ located verbatim in the source · abstract, characters 372–495 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Hydrogen peroxide increased most in H. annuus under drought, by 258%.

Taxon
H. annuusHelianthus annuuse1
Treatment
Droughte0
Outcome
hydrogen peroxidee1
Direction
greatest increasee1
Value
258%e1

Not reportedComparator

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

Helianthus annuus
Helianthus annuus · TaxonJon Sullivan · Public domain · Wikimedia Commons ↗

Verbatim

e0

“Drought markedly reduced”

✓ located verbatim in the source · abstract, characters 372–396 · show in text ↑
e1

“H. annuus showing the greatest increase in hydrogen peroxide (258%)”

✓ located verbatim in the source · abstract, characters 533–600 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Malondialdehyde accumulation was highest in C. album under drought, at 284%.

Taxon
C. albumChenopodium albume1
Treatment
Droughte0
Outcome
malondialdehyde accumulatione1
Direction
higheste1
Value
284%e1

Not reportedComparator

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

Chenopodium album
Chenopodium album · TaxonOlivier Pichard · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought markedly reduced”

✓ located verbatim in the source · abstract, characters 372–396 · show in text ↑
e1

“C. album exhibiting the highest malondialdehyde accumulation (284%)”

✓ located verbatim in the source · abstract, characters 605–672 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Catalase activity increased by 837% in C. album under drought.

Taxon
C. albumChenopodium albume1
Treatment
Droughte0
Outcome
catalase activitye1
Direction
increasede1
Value
837%e1

Not reportedComparator

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

Chenopodium album
Chenopodium album · TaxonOlivier Pichard · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought markedly reduced”

✓ located verbatim in the source · abstract, characters 372–396 · show in text ↑
e1

“catalase activity increased dramatically in C. album (837%) and H. annuus (630%)”

✓ located verbatim in the source · abstract, characters 719–799 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Catalase activity increased by 630% in H. annuus under drought.

Taxon
H. annuusHelianthus annuuse1
Treatment
Droughte0
Outcome
catalase activitye1
Direction
increasede1
Value
630%e1

Not reportedComparator

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

Helianthus annuus
Helianthus annuus · TaxonJon Sullivan · Public domain · Wikimedia Commons ↗

Verbatim

e0

“Drought markedly reduced”

✓ located verbatim in the source · abstract, characters 372–396 · show in text ↑
e1

“catalase activity increased dramatically in C. album (837%) and H. annuus (630%)”

✓ located verbatim in the source · abstract, characters 719–799 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Proline levels increased sharply under drought, particularly in T. aestivum, indicating enhanced osmotic adjustment.

Taxon
T. aestivumTriticum aestivume1
Treatment
Droughte0
Outcome
Proline levelse1
Direction
increased sharplye1

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 markedly reduced”

✓ located verbatim in the source · abstract, characters 372–396 · show in text ↑
e1

“Proline levels increased sharply, particularly in T. aestivum and C. album”

✓ located verbatim in the source · abstract, characters 801–875 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Proline levels increased sharply under drought, particularly in C. album.

Taxon
C. albumChenopodium albume1
Treatment
Droughte0
Outcome
Proline levelse1
Direction
increased sharplye1

Not reportedComparator

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

Chenopodium album
Chenopodium album · TaxonOlivier Pichard · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought markedly reduced”

✓ located verbatim in the source · abstract, characters 372–396 · show in text ↑
e1

“Proline levels increased sharply, particularly in T. aestivum and C. album”

✓ located verbatim in the source · abstract, characters 801–875 · show in text ↑
Intervention effectobservedoriginal resultpositivesignificant (as stated)measured

Claim · model-written summary

Carotenoid content rose significantly in H. annuus by 141% under drought.

Taxon
H. annuusHelianthus annuuse1
Treatment
Droughte0
Outcome
Carotenoid contente1
Direction
rosee1
Value
141%e1

Not reportedComparator

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

Helianthus annuus
Helianthus annuus · TaxonJon Sullivan · Public domain · Wikimedia Commons ↗

Verbatim

e0

“Drought markedly reduced”

✓ located verbatim in the source · abstract, characters 372–396 · show in text ↑
e1

“Carotenoid content also rose significantly in H. annuus (141%)”

✓ located verbatim in the source · abstract, characters 918–980 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

TaP5CS was upregulated in T. aestivum under drought, correlating with proline accumulation.

Taxon
T. aestivumTriticum aestivume1
Treatment
Droughte0
Gene
TaP5CSe1
Outcome
upregulation of TaP5CSe1
Direction
upregulatione1
Qualifier
correlating with proline accumulatione1

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 markedly reduced”

✓ located verbatim in the source · abstract, characters 372–396 · show in text ↑
e1

“upregulation of TaP5CS in T. aestivum , correlating with proline accumulation”

✓ located verbatim in the source · abstract, characters 1055–1132 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

CaHSP26 was upregulated in C. album under drought, potentially stabilizing photosystem II.

Taxon
C. albumChenopodium albume1
Treatment
Droughte0
Gene
CaHSP26e1
Outcome
CaHSP26e1
Qualifier
potentially stabilizing photosystem IIe1

Not reportedComparatorDirection

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

Chenopodium album
Chenopodium album · TaxonOlivier Pichard · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought markedly reduced”

✓ located verbatim in the source · abstract, characters 372–396 · show in text ↑
e1

“and CaHSP26 in C. album , potentially stabilizing photosystem II”

✓ located verbatim in the source · abstract, characters 1134–1198 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Principal component analysis identified catalase activity, root-to-shoot ratio, hydrogen peroxide and proline as major contributors to drought response variance.

Outcome
major contributors to drought response variancee0

Not reportedTaxon

Verbatim

e0

“Principal component analysis identified catalase activity, root-to-shoot ratio, hydrogen peroxide, and proline as major contributors to drought response variance”

✓ located verbatim in the source · abstract, characters 1200–1361 · show in text ↑
Observationauthor interpretationoriginal resultmeasured

Claim · model-written summary

C3 species showed strong enzymatic and osmotic adjustments under drought.

Taxon
C3 speciese0
Outcome
strong enzymatic and osmotic adjustmentse0

Verbatim

e0

“C3 species showing strong enzymatic and osmotic adjustments”

✓ located verbatim in the source · abstract, characters 1457–1516 · show in text ↑
Observationauthor interpretationoriginal resultmeasured

Claim · model-written summary

Intermediate-C4 and C4-like species showed greater tissue integrity and ROS balance under drought.

Taxon
intermediate-C4 and C4-like speciese0
Outcome
greater tissue integrity and ROS balancee0

Verbatim

e0

“intermediate-C4 and C4-like species exhibiting greater tissue integrity and ROS balance”

✓ located verbatim in the source · abstract, characters 1522–1609 · show in text ↑

drought · physiology: antioxidant enzymes · Helianthus annuus (common sunflower)Results differ

9 papers: 5 positive, 4 negative or no effect

4 papers in the same direction as this paper · 4 in the opposite direction

  1. positive
  2. negative
  3. negative

5 papers more in this group

See all 9 papers →

drought · physiology: oxidative · Helianthus annuus (common sunflower)Results differ

16 papers: 3 positive, 13 negative or no effect

12 papers in the same direction as this paper · 3 in the opposite direction

  1. negative
  2. positive
  3. positive

12 papers more in this group

See all 16 papers →

drought · growth · Helianthus annuus (common sunflower)Results differ

59 papers: 9 positive, 50 negative or no effect

49 papers in the same direction as this paper · 9 in the opposite direction

  1. negative
  2. positive
  3. positive

55 papers more in this group

See all 59 papers →

drought · physiology: oxidative · Chenopodium album (common lambsquarters)Positive and negative results

2 papers: 1 positive, 1 negative or no effect

1 in the opposite direction

  1. positive
See all 2 papers →

drought · physiology: pigments · Helianthus annuus (common sunflower)Positive and negative results

19 papers: 1 positive, 18 negative or no effect

18 in the opposite direction

  1. negative
  2. negative
  3. negative

15 papers more in this group

See all 19 papers →

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

75 papers: 48 positive, 27 negative or no effect

47 papers in the same direction as this paper · 27 in the opposite direction

  1. negative
  2. negative
  3. negative

71 papers more in this group

See all 75 papers →

drought · molecular: expression · Triticum aestivum (wheat)Positive and negative results

98 papers: 74 positive, 24 negative or no effect

73 papers in the same direction as this paper · 24 in the opposite direction

  1. negative
  2. negative
  3. negative

94 papers more in this group

See all 98 papers →

drought · physiology: water · Chenopodium album (common lambsquarters)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 →

8 of 9 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.