Paper · 2025 · Industrial Crops and Products

Phytoremediation potential of Linum usitatissimum for copper-contaminated soil: Role of nitrogen to improve phytoextraction through altering physio-biochemical and molecular process

Atif Ur Rahman; Junde Hou; Dongmei Yin; et al.

11
claims
22
verbatim quotes
5
citations
2
research areas
✓
Open access · cc-by

Abstract

Flax ( Linum usitatissimun ) is a promising candidate for copper (Cu) phytoextraction from agricultural soils. However, the effects of nitrogen (N) fertilizer driving the efficiency of Cu phytoextraction remain poorly understood. This study aimed to evaluate the dynamics of different levels of N (0, 125, 250, 375 mg kg −1 soil) on flax growth and Cu uptake with Cu-contaminated soil (2137 mg kg −1 Cu) and in mixtures of Cu-contaminated soil and natural soil (1189 mg kg −1 Cu). Our results demonstrated that the application of N (250 mg kg −1 soil) alleviated Cu toxicity by increasing plant growth and biomass, including dry weight, plant height, stem length, number of pods and number of branches per plant by 153.7 %, 43.1 %, 52.2 %, and 220.8 %, respectively, compared to control. N application at Cu 0.5 N 2 significantly reduced malondialdehyde, proline, and peroxidase activity by 26 %, 66.8 %, and 60.1 %, respectively, while superoxide dismutase activity was maximally enhanced by 365.6 % under Cu 0.5 N 3 treatment. N application progressively modulated antioxidants response, indicating that N plays a protective role against Cu-induced oxidative stress. Furthermore, transcriptome analysis identified differentially expressed genes (DEGs) associated with signal transduction and the phenylpropanoid biosynthesis pathway, which play key roles in N-mediated Cu detoxification. The involvement of phytohormones, including auxin (Aux1, LAX, and SAUR), gibberellic acid (DELLA), and salicylic acid (PR1, TGA), suggests that N improves heavy metals (HMs) tolerance via hormone signal transduction. Notably, N fertilization activated the expression of genes encoding peroxidase (E1.11.1.7) and respiratory burst oxidase homolog (RBOH) in the phenylpropanoid biosynthesis and MAPK signaling pathways, respectively, thereby alleviating Cu-induced damage. These findings indicate that varying N levels enhance Cu accumulation while maintaining ionic balance, facilitated by the activity of key transporters such as ZIP, CTR1, and AMT. These findings provide deeper insights into the mechanisms through which N alleviates Cu phytotoxicity, laying the foundation for optimizing phytoremediation strategies. • Varying N levels influence physio-biochemical, and gene expression processes under Cu stress. • N mitigates Cu-induced growth inhibition in flax plants. • The signal transduction pathways are crucial in N-mediated alleviation of Cu toxicity. • N enhances Cu uptake by regulating key transporters.

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Claims 11

Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Nitrogen at 250 mg kg −1 soil increased flax dry weight by 153.7% relative to control.

Treatment
N (250 mg kg −1 soil)e1
Outcome
dry weighte1
Direction
increasinge1
Value
153.7 %e1
Comparator
controle1

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

Linum usitatissimum
Linum usitatissimum · TaxonH. Zell · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Flax ( Linum usitatissimun )”

✓ located verbatim in the source · abstract, characters 0–28 · show in text ↑
e1

“application of N (250 mg kg −1 soil) alleviated Cu toxicity by increasing plant growth and biomass, including dry weight, plant height, stem length, number of pods and number of branches per plant by 153.7 %, 43.1 %, 52.2 %, and 220.8 %, respectively, compared to control”

✓ located verbatim in the source · abstract, characters 514–785 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Nitrogen at 250 mg kg −1 soil increased flax plant height by 43.1% relative to control.

Treatment
N (250 mg kg −1 soil)e1
Outcome
plant heighte1
Direction
increasinge1
Value
43.1 %e1
Comparator
controle1

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

Linum usitatissimum
Linum usitatissimum · TaxonH. Zell · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Flax ( Linum usitatissimun )”

✓ located verbatim in the source · abstract, characters 0–28 · show in text ↑
e1

“application of N (250 mg kg −1 soil) alleviated Cu toxicity by increasing plant growth and biomass, including dry weight, plant height, stem length, number of pods and number of branches per plant by 153.7 %, 43.1 %, 52.2 %, and 220.8 %, respectively, compared to control”

✓ located verbatim in the source · abstract, characters 514–785 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Nitrogen at 250 mg kg −1 soil increased flax stem length by 52.2% relative to control.

Treatment
N (250 mg kg −1 soil)e1
Outcome
stem lengthe1
Direction
increasinge1
Value
52.2 %e1
Comparator
controle1

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

Linum usitatissimum
Linum usitatissimum · TaxonH. Zell · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Flax ( Linum usitatissimun )”

✓ located verbatim in the source · abstract, characters 0–28 · show in text ↑
e1

“application of N (250 mg kg −1 soil) alleviated Cu toxicity by increasing plant growth and biomass, including dry weight, plant height, stem length, number of pods and number of branches per plant by 153.7 %, 43.1 %, 52.2 %, and 220.8 %, respectively, compared to control”

✓ located verbatim in the source · abstract, characters 514–785 · show in text ↑
Intervention effectauthor interpretationoriginal resultpositivemeasured

Claim · model-written summary

Nitrogen at 250 mg kg −1 soil increased flax pod and branch number by 220.8% as reported.

Treatment
N (250 mg kg −1 soil)e1
Outcome
number of pods and number of branches per plante1
Direction
increasinge1
Value
220.8 %e1
Qualifier
respectivelye1
Comparator
controle1

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

Linum usitatissimum
Linum usitatissimum · TaxonH. Zell · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Flax ( Linum usitatissimun )”

✓ located verbatim in the source · abstract, characters 0–28 · show in text ↑
e1

“application of N (250 mg kg −1 soil) alleviated Cu toxicity by increasing plant growth and biomass, including dry weight, plant height, stem length, number of pods and number of branches per plant by 153.7 %, 43.1 %, 52.2 %, and 220.8 %, respectively, compared to control”

✓ located verbatim in the source · abstract, characters 514–785 · show in text ↑
Intervention effectobservedoriginal resultpositivesignificant (as stated)measured

Claim · model-written summary

Cu 0.5 N 2 nitrogen treatment reduced flax malondialdehyde by 26%.

Treatment
Cu 0.5 N 2e1
Outcome
malondialdehydee1
Direction
reducede1
Value
26 %e1

Not reportedComparator

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

Linum usitatissimum
Linum usitatissimum · TaxonH. Zell · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Flax ( Linum usitatissimun )”

✓ located verbatim in the source · abstract, characters 0–28 · show in text ↑
e1

“N application at Cu 0.5 N 2 significantly reduced malondialdehyde, proline, and peroxidase activity by 26 %, 66.8 %, and 60.1 %, respectively”

✓ located verbatim in the source · abstract, characters 787–928 · show in text ↑
Intervention effectobservedoriginal resultpositivesignificant (as stated)measured

Claim · model-written summary

Cu 0.5 N 2 nitrogen treatment reduced flax proline by 66.8%.

Treatment
Cu 0.5 N 2e1
Outcome
prolinee1
Direction
reducede1
Value
66.8 %e1

Not reportedComparator

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

Linum usitatissimum
Linum usitatissimum · TaxonH. Zell · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Flax ( Linum usitatissimun )”

✓ located verbatim in the source · abstract, characters 0–28 · show in text ↑
e1

“N application at Cu 0.5 N 2 significantly reduced malondialdehyde, proline, and peroxidase activity by 26 %, 66.8 %, and 60.1 %, respectively”

✓ located verbatim in the source · abstract, characters 787–928 · show in text ↑
Intervention effectobservedoriginal resultpositivesignificant (as stated)measured

Claim · model-written summary

Cu 0.5 N 2 nitrogen treatment reduced flax peroxidase activity by 60.1%.

Treatment
Cu 0.5 N 2e1
Outcome
peroxidase activitye1
Direction
reducede1
Value
60.1 %e1

Not reportedComparator

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

Linum usitatissimum
Linum usitatissimum · TaxonH. Zell · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Flax ( Linum usitatissimun )”

✓ located verbatim in the source · abstract, characters 0–28 · show in text ↑
e1

“N application at Cu 0.5 N 2 significantly reduced malondialdehyde, proline, and peroxidase activity by 26 %, 66.8 %, and 60.1 %, respectively”

✓ located verbatim in the source · abstract, characters 787–928 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Cu 0.5 N 3 treatment maximally enhanced superoxide dismutase activity by 365.6%.

Treatment
Cu 0.5 N 3 treatmente1
Outcome
superoxide dismutase activitye1
Direction
maximally enhancede1
Value
365.6 %e1

Not reportedComparator

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

Linum usitatissimum
Linum usitatissimum · TaxonH. Zell · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Flax ( Linum usitatissimun )”

✓ located verbatim in the source · abstract, characters 0–28 · show in text ↑
e1

“superoxide dismutase activity was maximally enhanced by 365.6 % under Cu 0.5 N 3 treatment”

✓ located verbatim in the source · abstract, characters 936–1026 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Transcriptomics identified differentially expressed genes in signal transduction and phenylpropanoid biosynthesis.

Outcome
differentially expressed genes (DEGs) associated with signal transduction and the phenylpropanoid biosynthesis pathwaye1

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

Linum usitatissimum
Linum usitatissimum · TaxonH. Zell · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Flax ( Linum usitatissimun )”

✓ located verbatim in the source · abstract, characters 0–28 · show in text ↑
e1

“transcriptome analysis identified differentially expressed genes (DEGs) associated with signal transduction and the phenylpropanoid biosynthesis pathway”

✓ located verbatim in the source · abstract, characters 1181–1333 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Nitrogen fertilization activated expression of genes encoding peroxidase and respiratory burst oxidase homolog.

Treatment
N fertilizatione1
Gene
peroxidase (E1.11.1.7) and respiratory burst oxidase homolog (RBOH)e1
Outcome
expression of genes encoding peroxidase (E1.11.1.7) and respiratory burst oxidase homolog (RBOH)e1
Direction
activatede1

Not reportedComparator

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

Linum usitatissimum
Linum usitatissimum · TaxonH. Zell · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Flax ( Linum usitatissimun )”

✓ located verbatim in the source · abstract, characters 0–28 · show in text ↑
e1

“N fertilization activated the expression of genes encoding peroxidase (E1.11.1.7) and respiratory burst oxidase homolog (RBOH) in the phenylpropanoid biosynthesis and MAPK signaling pathways, respectively”

✓ located verbatim in the source · abstract, characters 1615–1819 · show in text ↑
Intervention effectauthor interpretationoriginal resultpositivemeasured

Claim · model-written summary

Varying nitrogen levels enhanced copper accumulation while maintaining ionic balance.

Treatment
varying N levelse1
Gene
ZIP, CTR1, and AMTe1
Outcome
Cu accumulatione1
Direction
enhancee1

Not reportedComparator

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

Linum usitatissimum
Linum usitatissimum · TaxonH. Zell · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Flax ( Linum usitatissimun )”

✓ located verbatim in the source · abstract, characters 0–28 · show in text ↑
e1

“varying N levels enhance Cu accumulation while maintaining ionic balance, facilitated by the activity of key transporters such as ZIP, CTR1, and AMT”

✓ located verbatim in the source · abstract, characters 1889–2037 · show in text ↑

copper atom · physiology: oxidative · Linum usitatissimum (flax)Positive and negative results

2 papers: 1 positive, 1 negative or no effect

1 in the opposite direction

  1. negative
See all 2 papers →

nitrogen · nutrient: content · Linum usitatissimum (flax)Positive and negative results

9 papers: 8 positive, 1 negative or no effect

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

  1. no effect
  2. positive
  3. positive

5 papers more in this group

See all 9 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.