Paper · 2025 · Agriculture

Brassinosteroids Alleviate Ethylene-Induced Copper Oxide Nanoparticle Toxicity and Ultrastructural and Stomatal Damage in Rice Seedlings

Wardah Azhar; Abdul Salam; Ali Raza Khan; et al.

15
claims
36
verbatim quotes
2
citations
2
research areas
✓
Open access · cc-by

Abstract

Nanoparticle contamination has been associated with adverse impacts on crop productivity. Thus, effective approaches are necessary to ameliorate NP-induced phytotoxicity. The present study aimed to investigate the efficacy of brassinosteroids and ethylene in regulating CuO NPs toxicity in rice seedlings. Therefore, we comprehensively evaluated the crosstalk of 24-Epibrassinolide and ethylene in regulating CuO NP-induced phytotoxicity at the physiological, cellular ultrastructural, and biochemical levels. The results of the study illustrated that exposure to CuO NPs at 450 mg/L displayed a significant decline in growth attributes and induced toxic effects in rice seedlings. Furthermore, the exogenous application of ethylene biosynthesis precursor 1-aminocyclopropane-1-carboxylic acid (ACC) at 20 µM with 450 mg/L of CuO NPs significantly enhanced the reactive oxygen species (ROS) accumulation that led to the stimulation of ultrastructural and stomatal damage and reduced antioxidant enzyme activities (CAT and APX) in rice tissues. On the contrary, it was noticed that 24-Epibrassinolide (BR) at 0.01 µM improved plant biomass and growth, restored cellular ultrastructure, and enhanced antioxidant enzyme activities (CAT and APX) under exposure to 450 mg/L of CuO NPs. In addition, brassinosteroids reduced ROS accumulation and the toxic effects of 450 mg/L of CuO NPs on guard cells and the stomatal aperture of rice seedlings. Interestingly, when 0.01 µM of brassinosteroids, 20 µM of ACC, and 450 mg/L of CuO NPs were applied together, BRs and ethylene showed antagonistic crosstalk under CuO NP stress via partially reducing the ethylene-induced CuO NP toxicity on plant growth, cellular ultrastructure, stomatal aperture, and guard cell and antioxidant enzyme activities (CAT and APX) in rice seedlings. BR supplementation with ACC and CuO NPs notably diminished ACC-induced CuO NPs’ toxic effects on all of the mentioned attributes in rice seedlings. This study uncovered the interesting crosstalk of two main phytohormones under CuO NPs stress, providing basic knowledge to improve crop yield and productivity in CuO NPs-contaminated areas.

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 15

Intervention effectobservedoriginal resultnegativesignificant (as stated)measured

Claim · model-written summary

Exposure to 450 mg/L CuO NPs significantly reduced rice seedling growth attributes.

Taxon
rice seedlingsOryza sativae0
Treatment
CuO NPs at 450 mg/Le0
Outcome
growth attributese0
Direction
declinee0

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“exposure to CuO NPs at 450 mg/L displayed a significant decline in growth attributes and induced toxic effects in rice seedlings”

✓ located verbatim in the source · abstract, characters 552–680 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Exposure to 450 mg/L CuO NPs caused toxic effects in rice seedlings.

Taxon
rice seedlingsOryza sativae0
Treatment
CuO NPs at 450 mg/Le0
Outcome
toxic effectse0
Direction
inducede0

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“exposure to CuO NPs at 450 mg/L displayed a significant decline in growth attributes and induced toxic effects in rice seedlings”

✓ located verbatim in the source · abstract, characters 552–680 · show in text ↑
Intervention effectobservedoriginal resultnegativesignificant (as stated)measured

Claim · model-written summary

ACC with CuO NPs significantly increased reactive oxygen species accumulation in rice tissue.

Taxon
rice seedlingsOryza sativat0
Treatment
ACC) at 20 µM with 450 mg/L of CuO NPse0
Outcome
reactive oxygen species (ROS) accumulatione0
Direction
enhancede0

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“ACC) at 20 µM with 450 mg/L of CuO NPs significantly enhanced the reactive oxygen species (ROS) accumulation”

✓ located verbatim in the source · abstract, characters 795–903 · show in text ↑
t0

“Rice Seedlings”

✓ located verbatim in the source · title, characters 122–136 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

ACC with CuO NPs stimulated rice ultrastructural damage.

Taxon
rice seedlingsOryza sativat0
Treatment
ACC) at 20 µM with 450 mg/L of CuO NPse1
Outcome
ultrastructural and stomatal damagee0
Direction
stimulatione0

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“that led to the stimulation of ultrastructural and stomatal damage”

✓ located verbatim in the source · abstract, characters 904–970 · show in text ↑
e1

“ACC) at 20 µM with 450 mg/L of CuO NPs”

✓ located verbatim in the source · abstract, characters 795–833 · show in text ↑
t0

“Rice Seedlings”

✓ located verbatim in the source · title, characters 122–136 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

ACC with CuO NPs stimulated rice stomatal damage.

Taxon
rice seedlingsOryza sativat0
Treatment
ACC) at 20 µM with 450 mg/L of CuO NPse1
Outcome
stomatal damagee0
Direction
stimulatione0

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“that led to the stimulation of ultrastructural and stomatal damage”

✓ located verbatim in the source · abstract, characters 904–970 · show in text ↑
e1

“ACC) at 20 µM with 450 mg/L of CuO NPs”

✓ located verbatim in the source · abstract, characters 795–833 · show in text ↑
t0

“Rice Seedlings”

✓ located verbatim in the source · title, characters 122–136 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

ACC with CuO NPs reduced CAT and APX antioxidant enzyme activities in rice tissues.

Treatment
ACC) at 20 µM with 450 mg/L of CuO NPse1
Gene
CAT and APXe0
Outcome
antioxidant enzyme activities (CAT and APX)e0
Direction
reducede0

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“reduced antioxidant enzyme activities (CAT and APX) in rice tissues”

✓ located verbatim in the source · abstract, characters 975–1042 · show in text ↑
e1

“ACC) at 20 µM with 450 mg/L of CuO NPs”

✓ located verbatim in the source · abstract, characters 795–833 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

BR at 0.01 µM improved rice biomass under 450 mg/L CuO NPs.

Taxon
rice seedlingsOryza sativat0
Treatment
24-Epibrassinolide (BR) at 0.01 µMe0
Outcome
plant biomasse0
Direction
improvede0
Qualifier
under exposure to 450 mg/L of CuO NPse1

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“24-Epibrassinolide (BR) at 0.01 µM improved plant biomass and growth”

✓ located verbatim in the source · abstract, characters 1081–1149 · show in text ↑
e1

“under exposure to 450 mg/L of CuO NPs”

✓ located verbatim in the source · abstract, characters 1242–1279 · show in text ↑
t0

“Rice Seedlings”

✓ located verbatim in the source · title, characters 122–136 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

BR at 0.01 µM improved rice growth under 450 mg/L CuO NPs.

Taxon
rice seedlingsOryza sativat0
Treatment
24-Epibrassinolide (BR) at 0.01 µMe0
Outcome
growthe0
Direction
improvede0
Qualifier
under exposure to 450 mg/L of CuO NPse1

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“24-Epibrassinolide (BR) at 0.01 µM improved plant biomass and growth”

✓ located verbatim in the source · abstract, characters 1081–1149 · show in text ↑
e1

“under exposure to 450 mg/L of CuO NPs”

✓ located verbatim in the source · abstract, characters 1242–1279 · show in text ↑
t0

“Rice Seedlings”

✓ located verbatim in the source · title, characters 122–136 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

BR restored cellular ultrastructure under CuO NP exposure.

Taxon
rice seedlingsOryza sativat0
Treatment
24-Epibrassinolide (BR) at 0.01 µMe1
Outcome
cellular ultrastructuree0
Direction
restorede0
Qualifier
under exposure to 450 mg/L of CuO NPse2

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“restored cellular ultrastructure”

✓ located verbatim in the source · abstract, characters 1151–1183 · show in text ↑
e1

“24-Epibrassinolide (BR) at 0.01 µM”

✓ located verbatim in the source · abstract, characters 1081–1115 · show in text ↑
e2

“under exposure to 450 mg/L of CuO NPs”

✓ located verbatim in the source · abstract, characters 1242–1279 · show in text ↑
t0

“Rice Seedlings”

✓ located verbatim in the source · title, characters 122–136 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

BR enhanced CAT and APX antioxidant enzyme activities under CuO NP exposure.

Taxon
rice seedlingsOryza sativat0
Treatment
24-Epibrassinolide (BR) at 0.01 µMe1
Gene
CAT and APXe0
Outcome
antioxidant enzyme activities (CAT and APX)e0
Direction
enhancede0
Qualifier
under exposure to 450 mg/L of CuO NPse0

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“enhanced antioxidant enzyme activities (CAT and APX) under exposure to 450 mg/L of CuO NPs”

✓ located verbatim in the source · abstract, characters 1189–1279 · show in text ↑
e1

“24-Epibrassinolide (BR) at 0.01 µM”

✓ located verbatim in the source · abstract, characters 1081–1115 · show in text ↑
t0

“Rice Seedlings”

✓ located verbatim in the source · title, characters 122–136 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Brassinosteroids reduced ROS accumulation in rice seedlings exposed to CuO NPs.

Taxon
rice seedlingsOryza sativat0
Treatment
brassinosteroidse0
Outcome
ROS accumulatione0
Direction
reducede0
Qualifier
450 mg/L of CuO NPse1

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“brassinosteroids reduced ROS accumulation”

✓ located verbatim in the source · abstract, characters 1294–1335 · show in text ↑
e1

“exposure to 450 mg/L of CuO NPs”

✓ located verbatim in the source · abstract, characters 1248–1279 · show in text ↑
t0

“Rice Seedlings”

✓ located verbatim in the source · title, characters 122–136 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Brassinosteroids reduced CuO NP toxic effects on rice guard cells.

Taxon
rice seedlingsOryza sativae0
Organ / stage
guard cellse0
Treatment
brassinosteroidse1
Outcome
toxic effects of 450 mg/L of CuO NPs on guard cellse0
Direction
reducede0

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“reduced ROS accumulation and the toxic effects of 450 mg/L of CuO NPs on guard cells and the stomatal aperture of rice seedlings”

✓ located verbatim in the source · abstract, characters 1311–1439 · show in text ↑
e1

“brassinosteroids reduced ROS accumulation”

✓ located verbatim in the source · abstract, characters 1294–1335 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Brassinosteroids reduced CuO NP toxic effects on the rice stomatal aperture.

Taxon
rice seedlingsOryza sativae0
Organ / stage
stomatal aperturee0
Treatment
brassinosteroidse1
Outcome
toxic effects of 450 mg/L of CuO NPs on guard cells and the stomatal aperturee0
Direction
reducede0

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“reduced ROS accumulation and the toxic effects of 450 mg/L of CuO NPs on guard cells and the stomatal aperture of rice seedlings”

✓ located verbatim in the source · abstract, characters 1311–1439 · show in text ↑
e1

“brassinosteroids reduced ROS accumulation”

✓ located verbatim in the source · abstract, characters 1294–1335 · show in text ↑
Intervention effectauthor interpretationoriginal resultpositivemeasured

Claim · model-written summary

Brassinosteroid and ACC cotreatment partially reduced ethylene-induced CuO NP toxicity in rice growth.

Taxon
rice seedlingsOryza sativat0
Treatment
0.01 µM of brassinosteroids, 20 µM of ACC, and 450 mg/L of CuO NPse1
Outcome
ethylene-induced CuO NP toxicity on plant growthe0
Direction
partially reducinge0

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“BRs and ethylene showed antagonistic crosstalk under CuO NP stress via partially reducing the ethylene-induced CuO NP toxicity on plant growth”

✓ located verbatim in the source · abstract, characters 1551–1693 · show in text ↑
e1

“0.01 µM of brassinosteroids, 20 µM of ACC, and 450 mg/L of CuO NPs”

✓ located verbatim in the source · abstract, characters 1461–1527 · show in text ↑
t0

“Rice Seedlings”

✓ located verbatim in the source · title, characters 122–136 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Brassinosteroid and ACC cotreatment diminished CuO NP toxicity affecting rice ultrastructure and stomata.

Taxon
rice seedlingsOryza sativae0
Treatment
BR supplementation with ACC and CuO NPse0
Outcome
ACC-induced CuO NPs’ toxic effectse0
Direction
diminishede0
Qualifier
all of the mentioned attributese0

Not reportedComparator

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

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“BR supplementation with ACC and CuO NPs notably diminished ACC-induced CuO NPs’ toxic effects on all of the mentioned attributes in rice seedlings”

✓ located verbatim in the source · abstract, characters 1821–1967 · show in text ↑

brassinolide · growth · Oryza sativa (rice)Results differ

32 papers: 27 positive, 5 negative or no effect

26 papers in the same direction as this paper · 5 in the opposite direction

  1. negative
  2. negative
  3. negative

28 papers more in this group

See all 32 papers →

1-aminocyclopropanecarboxylic acid · molecular: metabolites · Oryza sativa (rice)Positive and negative results

4 papers: 3 positive, 1 negative or no effect

2 papers in the same direction as this paper · 1 in the opposite direction

  1. no effect
  2. positive
  3. positive
See all 4 papers →

1-aminocyclopropanecarboxylic acid · physiology: hormones · Oryza sativa (rice)Positive and negative results

5 papers: 3 positive, 2 negative or no effect

2 papers in the same direction as this paper · 2 in the opposite direction

  1. no effect
  2. negative
  3. positive

1 paper more in this group

See all 5 papers →

brassinolide · molecular: metabolites · Oryza sativa (rice)Positive and negative results

9 papers: 6 positive, 3 negative or no effect

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

  1. negative
  2. no effect
  3. negative

5 papers more in this group

See all 9 papers →

brassinolide · soil: biology · Oryza sativa (rice)Results agree

2 papers: 2 positive, 0 negative or no effect

1 paper in the same direction as this paper

  1. positive
See all 2 papers →

24-epi-brassinolide · soil: biology · Oryza sativa (rice)Results agree

3 papers: 3 positive, 0 negative or no effect

2 papers in the same direction as this paper

  1. positive
  2. positive
See all 3 papers →

24-epi-brassinolide · growth · Oryza sativa (rice)Results agree

10 papers: 10 positive, 0 negative or no effect

9 papers in the same direction as this paper

  1. positive
  2. positive
  3. positive

6 papers more in this group

See all 10 papers →

nanoparticle · pest · Oryza sativa (rice)Positive and negative results

4 papers: 3 positive, 1 negative or no effect

3 in the opposite direction

  1. positive
  2. positive
  3. positive
See all 4 papers →

8 of 16 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.