Paper · 2026 · Plant Physiology and Biochemistry

Cyanobacterial priming enhances lignin biosynthesis for cell wall macromolecular reinforcement: A mechanism for mitigating In2O3 nanoparticle Stress in the legume alfalfa and the grass rye

Seham M. Hamed; Uğur Tan; Marwa Yousry A. Mohamed; et al.

14
claims
26
verbatim quotes
0
citations
2
research areas
✓
Open access · cc-by-nc-nd

Abstract

Exposure to indium oxide nanoparticles (In 2 O 3 -NPs) presents a critical challenge as an emerging soil contaminant that severely impairs plant growth and metabolic health. In this study, In 2 O 3-NP exposure reduced biomass by 58% in alfalfa, driven by excessive indium (In) accumulation, disrupted mineral (P and Fe) balance, and inhibited photosynthesis. This physiological decline led to a significant depletion of sugars and nitrogen metabolites, particularly in the more sensitive alfalfa model. To mitigate these toxic effects, we utilized cyanobacterial priming (CP) with Anabaena laxa to investigate how species-specific metabolic and physiological responses are shaped in alfalfa (legume) and rye (grass). CP treatment differentially mitigated toxicity by reducing In uptake, maintaining nutrient homeostasis, and restoring photosynthetic efficiency. This resulted in improved biomass, with alfalfa showing the most significant recovery. Mechanistically, CP enhanced sugar and nitrogen metabolism, providing the necessary precursors for the accumulation of protective secondary metabolites, such as phenolics and flavonoids, which reduced oxidative damage. Furthermore, CP induced a structural adaptation through the activation of the phenylpropanoid–lignin biosynthetic pathway. In alfalfa, elevated levels of cinnamic and coumaric acids were linked to increased activities of key enzymes, phenylalanine ammonia-lyase (PAL), caffeic acid O-methyltransferase (COMT), cinnamyl alcohol dehydrogenase (CAD), and cinnamoyl-CoA reductase (CCR). This resulted in a substantial 243.6% increase in lignin in alfalfa, compared to a 115.1% rise in rye. This study demonstrates that while rye relies on inherent physiological tolerance, alfalfa exhibits superior metabolic plasticity when primed with CP. These findings prove that alfalfa and rye utilize distinct survival strategies at the biochemical level; while rye leverages its natural resilience, alfalfa undergoes a complete metabolic reorganization to rescue its growth and survive nanoparticle stress. • In 2 O 3 -NPs reduce growth by inhibiting photosynthesis and inducing oxidative stress. • CP differentially restores plant growth, mineral homeostasis, and metabolic balance in alfalfa and rye. • CP facilitates metabolic reallocation of carbon and nitrogen, promoting biosynthesis of phenolics and lignin. • Activation of the phenylpropanoid pathway at the enzymatic level drives lignification and cell wall reinforcement. • Alfalfa demonstrates enhanced plasticity, resulting in increased synthesis of antioxidant and cell wall lignin.

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Claims 14

Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Nanoparticle exposure reduced alfalfa biomass by 58%.

Taxon
alfalfaMedicago sativae0
Treatment
In 2 O 3-NP exposuree0
Outcome
biomasse0
Direction
reducede0
Value
58%e0

Not reportedComparator

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

Medicago sativa
Medicago sativa · TaxonDrepanostoma · CC BY 4.0 · iNaturalist ↗

Verbatim

e0

“In 2 O 3-NP exposure reduced biomass by 58% in alfalfa”

✓ located verbatim in the source · abstract, characters 189–243 · show in text ↑
Intervention effectauthor interpretationoriginal resultnegativemeasured

Claim · model-written summary

Nanoparticle exposure caused excessive indium accumulation and disrupted phosphorus and iron balance in alfalfa.

Taxon
alfalfaMedicago sativae0
Outcome
excessive indium (In) accumulation, disrupted mineral (P and Fe) balancee0

Not reportedTreatmentComparatorDirection

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

Medicago sativa
Medicago sativa · TaxonDrepanostoma · CC BY 4.0 · iNaturalist ↗

Verbatim

e0

“in alfalfa, driven by excessive indium (In) accumulation, disrupted mineral (P and Fe) balance”

✓ located verbatim in the source · abstract, characters 233–327 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Nanoparticle exposure inhibited photosynthesis in alfalfa.

Taxon
alfalfaMedicago sativae0
Treatment
In 2 O 3-NP exposuree0
Outcome
photosynthesise0
Direction
inhibitede0

Not reportedComparator

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

Medicago sativa
Medicago sativa · TaxonDrepanostoma · CC BY 4.0 · iNaturalist ↗

Verbatim

e0

“In 2 O 3-NP exposure reduced biomass by 58% in alfalfa, driven by excessive indium (In) accumulation, disrupted mineral (P and Fe) balance, and inhibited photosynthesis”

✓ located verbatim in the source · abstract, characters 189–357 · show in text ↑
Observationobservedoriginal resultnegativesignificant (as stated)measured

Claim · model-written summary

Sugars and nitrogen metabolites were significantly depleted, particularly in alfalfa.

Taxon
alfalfaMedicago sativae0
Outcome
sugars and nitrogen metabolitese0
Direction
depletione0

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

Medicago sativa
Medicago sativa · TaxonDrepanostoma · CC BY 4.0 · iNaturalist ↗

Verbatim

e0

“significant depletion of sugars and nitrogen metabolites, particularly in the more sensitive alfalfa model”

✓ located verbatim in the source · abstract, characters 395–501 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Priming with Anabaena laxa reduced indium uptake in alfalfa and rye.

Taxon
alfalfaMedicago sativae2
Treatment
cyanobacterial priming (CP) with Anabaena laxae0
Interactor
Anabaena laxaAnabaena laxae0symbiont
Outcome
In uptakee1
Direction
reducinge1

Not reportedComparator

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

Medicago sativa
Medicago sativa · TaxonDrepanostoma · CC BY 4.0 · iNaturalist ↗

Verbatim

e0

“cyanobacterial priming (CP) with Anabaena laxa”

✓ located verbatim in the source · abstract, characters 548–594 · show in text ↑
e1

“CP treatment differentially mitigated toxicity by reducing In uptake”

✓ located verbatim in the source · abstract, characters 717–785 · show in text ↑
e2

“alfalfa (legume) and rye (grass)”

✓ located verbatim in the source · abstract, characters 683–715 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Priming maintained nutrient homeostasis in alfalfa.

Taxon
alfalfaMedicago sativae1
Treatment
cyanobacterial priming (CP) with Anabaena laxae2
Interactor
Anabaena laxaAnabaena laxae2symbiont
Outcome
nutrient homeostasise0
Direction
maintaininge0

Not reportedComparator

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

Medicago sativa
Medicago sativa · TaxonDrepanostoma · CC BY 4.0 · iNaturalist ↗

Verbatim

e0

“CP treatment differentially mitigated toxicity by reducing In uptake, maintaining nutrient homeostasis, and restoring photosynthetic efficiency”

✓ located verbatim in the source · abstract, characters 717–860 · show in text ↑
e1

“alfalfa (legume) and rye (grass)”

✓ located verbatim in the source · abstract, characters 683–715 · show in text ↑
e2

“cyanobacterial priming (CP) with Anabaena laxa”

✓ located verbatim in the source · abstract, characters 548–594 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Priming restored photosynthetic efficiency in alfalfa.

Taxon
alfalfaMedicago sativae1
Treatment
cyanobacterial priming (CP) with Anabaena laxae2
Interactor
Anabaena laxaAnabaena laxae2symbiont
Outcome
photosynthetic efficiencye0
Direction
restoringe0

Not reportedComparator

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

Medicago sativa
Medicago sativa · TaxonDrepanostoma · CC BY 4.0 · iNaturalist ↗

Verbatim

e0

“restoring photosynthetic efficiency”

✓ located verbatim in the source · abstract, characters 825–860 · show in text ↑
e1

“alfalfa (legume) and rye (grass)”

✓ located verbatim in the source · abstract, characters 683–715 · show in text ↑
e2

“cyanobacterial priming (CP) with Anabaena laxa”

✓ located verbatim in the source · abstract, characters 548–594 · show in text ↑
Intervention effectobservedoriginal resultpositivesignificant (as stated)measured

Claim · model-written summary

Priming improved biomass, with alfalfa showing the most significant recovery.

Taxon
alfalfaMedicago sativae0
Treatment
cyanobacterial priming (CP) with Anabaena laxae1
Interactor
Anabaena laxaAnabaena laxae1symbiont
Outcome
biomasse0
Direction
improvede0

Not reportedComparator

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

Medicago sativa
Medicago sativa · TaxonDrepanostoma · CC BY 4.0 · iNaturalist ↗

Verbatim

e0

“This resulted in improved biomass, with alfalfa showing the most significant recovery”

✓ located verbatim in the source · abstract, characters 862–947 · show in text ↑
e1

“cyanobacterial priming (CP) with Anabaena laxa”

✓ located verbatim in the source · abstract, characters 548–594 · show in text ↑
Mechanismauthor interpretationoriginal resultpositivemeasured

Claim · model-written summary

Priming enhanced sugar and nitrogen metabolism, supplying precursors for protective secondary metabolites.

Taxon
alfalfaMedicago sativae1
Outcome
accumulation of protective secondary metabolitese0
Qualifier
sugar and nitrogen metabolisme0

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

Medicago sativa
Medicago sativa · TaxonDrepanostoma · CC BY 4.0 · iNaturalist ↗

Verbatim

e0

“CP enhanced sugar and nitrogen metabolism, providing the necessary precursors for the accumulation of protective secondary metabolites, such as phenolics and flavonoids”

✓ located verbatim in the source · abstract, characters 966–1134 · show in text ↑
e1

“alfalfa (legume) and rye (grass)”

✓ located verbatim in the source · abstract, characters 683–715 · show in text ↑
Mechanismauthor interpretationoriginal resultpositivemeasured

Claim · model-written summary

Protective secondary metabolites reduced oxidative damage under priming.

Taxon
alfalfaMedicago sativae1
Compound
phenolics and flavonoidse0
Outcome
oxidative damagee0
Direction
reducede0

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

Medicago sativa
Medicago sativa · TaxonDrepanostoma · CC BY 4.0 · iNaturalist ↗

Verbatim

e0

“protective secondary metabolites, such as phenolics and flavonoids, which reduced oxidative damage”

✓ located verbatim in the source · abstract, characters 1068–1166 · show in text ↑
e1

“alfalfa (legume) and rye (grass)”

✓ located verbatim in the source · abstract, characters 683–715 · show in text ↑
Mechanismauthor interpretationoriginal resultpositivemeasured

Claim · model-written summary

Priming activated the phenylpropanoid–lignin biosynthetic pathway.

Taxon
alfalfaMedicago sativae1
Outcome
activation of the phenylpropanoid–lignin biosynthetic pathwaye0
Qualifier
structural adaptatione0

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

Medicago sativa
Medicago sativa · TaxonDrepanostoma · CC BY 4.0 · iNaturalist ↗

Verbatim

e0

“CP induced a structural adaptation through the activation of the phenylpropanoid–lignin biosynthetic pathway”

✓ located verbatim in the source · abstract, characters 1181–1289 · show in text ↑
e1

“alfalfa (legume) and rye (grass)”

✓ located verbatim in the source · abstract, characters 683–715 · show in text ↑
Associationobservedoriginal resultpositivemeasured

Claim · model-written summary

In alfalfa, elevated cinnamic and coumaric acids were linked to increased PAL, COMT, CAD, and CCR activity.

Taxon
alfalfaMedicago sativae0
Gene
phenylalanine ammonia-lyase (PAL), caffeic acid O-methyltransferase (COMT), cinnamyl alcohol dehydrogenase (CAD), and cinnamoyl-CoA reductase (CCR)e0
Compound
cinnamic and coumaric acidse0
Outcome
activities of key enzymese0
Direction
increasede0

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

Medicago sativa
Medicago sativa · TaxonDrepanostoma · CC BY 4.0 · iNaturalist ↗

Verbatim

e0

“In alfalfa, elevated levels of cinnamic and coumaric acids were linked to increased activities of key enzymes, phenylalanine ammonia-lyase (PAL), caffeic acid O-methyltransferase (COMT), cinnamyl alcohol dehydrogenase (CAD), and cinnamoyl-CoA reductase (CCR)”

✓ located verbatim in the source · abstract, characters 1291–1549 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Priming increased lignin in alfalfa by 243.6%.

Taxon
alfalfaMedicago sativae0
Treatment
cyanobacterial priming (CP) with Anabaena laxae1
Interactor
Anabaena laxaAnabaena laxae1symbiont
Outcome
lignine0
Direction
increasee0
Value
243.6%e0
Comparator
ryee0

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

Medicago sativa
Medicago sativa · TaxonDrepanostoma · CC BY 4.0 · iNaturalist ↗

Verbatim

e0

“243.6% increase in lignin in alfalfa, compared to a 115.1% rise in rye”

✓ located verbatim in the source · abstract, characters 1582–1652 · show in text ↑
e1

“cyanobacterial priming (CP) with Anabaena laxa”

✓ located verbatim in the source · abstract, characters 548–594 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Priming increased lignin in rye by 115.1%.

Treatment
cyanobacterial priming (CP) with Anabaena laxae1
Interactor
Anabaena laxaAnabaena laxae1symbiont
Outcome
lignine0
Direction
risee0
Value
115.1%e0
Comparator
alfalfae0

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

Secale cereale
Secale cereale · TaxonLSDSL · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“243.6% increase in lignin in alfalfa, compared to a 115.1% rise in rye”

✓ located verbatim in the source · abstract, characters 1582–1652 · show in text ↑
e1

“cyanobacterial priming (CP) with Anabaena laxa”

✓ located verbatim in the source · abstract, characters 548–594 · show in text ↑

ozone · growth · Medicago sativa (alfalfa)Positive and negative results

6 papers: 1 positive, 5 negative or no effect

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

  1. positive
  2. no effect
  3. no effect

2 papers more in this group

See all 6 papers →

seed treatment · growth · Medicago sativa (alfalfa)Positive and negative results

34 papers: 23 positive, 11 negative or no effect

22 papers in the same direction as this paper · 11 in the opposite direction

  1. negative
  2. no effect
  3. no effect

30 papers more in this group

See all 34 papers →

seed treatment · physiology: photosynthesis · Medicago sativa (alfalfa)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 →

seed treatment · molecular: metabolites · Medicago sativa (alfalfa)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 →

seed treatment · nutrient: content · Medicago sativa (alfalfa)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 →

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.