Paper · 2026 · Agronomy

Nitrogen Fertilizer Affects Culm Lodging Resistance by Regulating Phenylpropanoid Metabolism in Rice

Fei Weng; Yi Wang; Qingkui Li; et al.

11
claims
23
verbatim quotes
0
citations
3
research areas
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Open access · cc-by

Abstract

Excessive nitrogen (N) fertilization is widely used to increase rice yield, but it often leads to lodging by weakening culm strength. This study aimed to elucidate the structural and molecular mechanisms underlying nitrogen-induced changes in culm lodging resistance in rice. Field and pot experiments with two nitrogen levels were conducted using a randomized design with three biological replicates to evaluate the effects of high nitrogen application on culm mechanical properties, secondary cell wall development, and associated metabolic pathways. Mechanical measurements and microscopic analysis revealed that high nitrogen significantly reduced culm rigidity and impaired sclerenchyma development. To investigate the underlying mechanisms, integrated transcriptomic and proteomic analyses were performed on developing internodes. Differentially expressed genes and proteins were predominantly enriched in carbohydrate metabolism and phenylpropanoid biosynthesis pathways. Notably, key enzymes involved in lignin biosynthesis were consistently downregulated at the protein level under high-nitrogen conditions. In contrast, genes and proteins related to cellulose and hemicellulose biosynthesis exhibited transient inhibition at early stages followed by recovery or upregulation at later stages. Consistent with these findings, histochemical staining and quantitative assays demonstrated a significant reduction (14–16%) in lignin content in the fourth internode, whereas cellulose content showed no substantial change. Furthermore, lignin biosynthetic genes (OsCAD2, Os4CL3, and OsCOMT) were persistently suppressed during critical stages of secondary wall formation, while cellulose synthase genes (OsCESA4, OsCESA7, and OsCESA9) displayed more variable and less sustained expression patterns. Collectively, these results demonstrate that excessive nitrogen application weakens rice culms primarily by inhibiting lignin accumulation rather than cellulose deposition. The preferential suppression of the phenylpropanoid pathway and disruption of secondary cell wall formation provide a mechanistic basis for nitrogen-induced lodging susceptibility in rice.

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Claims 11

Intervention effectobservedoriginal resultnegativesignificant (as stated)measured

Claim · model-written summary

High nitrogen significantly reduced culm rigidity in rice.

Treatment
high nitrogene1
Outcome
culm rigiditye1
Direction
reducede1

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

“changes in culm lodging resistance in rice”

✓ located verbatim in the source · abstract, characters 232–274 · show in text ↑
e1

“high nitrogen significantly reduced culm rigidity and impaired sclerenchyma development”

✓ located verbatim in the source · abstract, characters 616–703 · show in text ↑
Intervention effectobservedoriginal resultnegativesignificant (as stated)measured

Claim · model-written summary

High nitrogen significantly impaired sclerenchyma development in rice culms.

Organ / stage
sclerenchymae1
Treatment
high nitrogene1
Outcome
sclerenchyma developmente1
Direction
impairede1

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

“changes in culm lodging resistance in rice”

✓ located verbatim in the source · abstract, characters 232–274 · show in text ↑
e1

“high nitrogen significantly reduced culm rigidity and impaired sclerenchyma development”

✓ located verbatim in the source · abstract, characters 616–703 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Differentially expressed genes and proteins in developing internodes were predominantly enriched in carbohydrate metabolism and phenylpropanoid biosynthesis pathways.

Organ / stage
developing internodese4
Outcome
carbohydrate metabolism and phenylpropanoid biosynthesis pathwayse3
Qualifier
predominantly enrichede3

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

“changes in culm lodging resistance in rice”

✓ located verbatim in the source · abstract, characters 232–274 · show in text ↑
e3

“Differentially expressed genes and proteins were predominantly enriched in carbohydrate metabolism and phenylpropanoid biosynthesis pathways”

✓ located verbatim in the source · abstract, characters 837–977 · show in text ↑
e4

“integrated transcriptomic and proteomic analyses were performed on developing internodes”

✓ located verbatim in the source · abstract, characters 747–835 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

High nitrogen consistently downregulated key lignin biosynthesis enzymes at the protein level.

Treatment
high-nitrogen conditionse5
Outcome
key enzymes involved in lignin biosynthesise5
Direction
downregulatede5
Qualifier
at the protein levele5

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

“changes in culm lodging resistance in rice”

✓ located verbatim in the source · abstract, characters 232–274 · show in text ↑
e5

“key enzymes involved in lignin biosynthesis were consistently downregulated at the protein level under high-nitrogen conditions”

✓ located verbatim in the source · abstract, characters 988–1115 · show in text ↑
Associationobservedoriginal resultmixedmeasured

Claim · model-written summary

Cellulose and hemicellulose biosynthesis genes and proteins showed transient inhibition at early stages.

Organ / stage
early stagese6
Outcome
genes and proteins related to cellulose and hemicellulose biosynthesise6
Direction
transient inhibitione6

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

“changes in culm lodging resistance in rice”

✓ located verbatim in the source · abstract, characters 232–274 · show in text ↑
e6

“genes and proteins related to cellulose and hemicellulose biosynthesis exhibited transient inhibition at early stages”

✓ located verbatim in the source · abstract, characters 1130–1247 · show in text ↑
Associationobservedoriginal resultnegativesignificant (as stated)measured

Claim · model-written summary

High nitrogen reduced lignin content by 14–16% in the fourth internode of rice.

Organ / stage
fourth internodee7
Outcome
lignin contente7
Direction
reductione7
Value
14–16%e7

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

“changes in culm lodging resistance in rice”

✓ located verbatim in the source · abstract, characters 232–274 · show in text ↑
e7

“histochemical staining and quantitative assays demonstrated a significant reduction (14–16%) in lignin content in the fourth internode, whereas cellulose content showed no substantial change”

✓ located verbatim in the source · abstract, characters 1334–1524 · show in text ↑
Associationobservedoriginal resultno effectnot significant (as stated)measured

Claim · model-written summary

Cellulose content in the fourth internode of rice showed no substantial change.

Organ / stage
fourth internodee7
Outcome
cellulose contente7
Direction
no substantial changee7

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

“changes in culm lodging resistance in rice”

✓ located verbatim in the source · abstract, characters 232–274 · show in text ↑
e7

“histochemical staining and quantitative assays demonstrated a significant reduction (14–16%) in lignin content in the fourth internode, whereas cellulose content showed no substantial change”

✓ located verbatim in the source · abstract, characters 1334–1524 · show in text ↑
Associationobservedoriginal resultnegativemeasured

Claim · model-written summary

Lignin biosynthetic genes OsCAD2, Os4CL3 and OsCOMT were persistently suppressed during critical stages of secondary wall formation in rice.

Organ / stage
secondary wall formatione1
Gene
OsCAD2, Os4CL3, and OsCOMTe1
Outcome
lignin biosynthetic genese1
Direction
suppressede1

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

“changes in culm lodging resistance in rice”

✓ located verbatim in the source · abstract, characters 232–274 · show in text ↑
e1

“lignin biosynthetic genes (OsCAD2, Os4CL3, and OsCOMT) were persistently suppressed during critical stages of secondary wall formation”

✓ located verbatim in the source · abstract, characters 1539–1673 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Cellulose synthase genes OsCESA4, OsCESA7 and OsCESA9 showed more variable and less sustained expression in rice.

Gene
OsCESA4, OsCESA7, and OsCESA9e2
Outcome
expression patternse2
Qualifier
more variable and less sustainede2

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

“changes in culm lodging resistance in rice”

✓ located verbatim in the source · abstract, characters 232–274 · show in text ↑
e2

“cellulose synthase genes (OsCESA4, OsCESA7, and OsCESA9) displayed more variable and less sustained expression patterns”

✓ located verbatim in the source · abstract, characters 1681–1800 · show in text ↑
Mechanismauthor interpretationoriginal resultnegativemeasured

Claim · model-written summary

Excessive nitrogen weakens rice culms mainly by inhibiting lignin accumulation rather than cellulose deposition.

Treatment
excessive nitrogen applicatione3
Outcome
lignin accumulatione3

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

“changes in culm lodging resistance in rice”

✓ located verbatim in the source · abstract, characters 232–274 · show in text ↑
e3

“excessive nitrogen application weakens rice culms primarily by inhibiting lignin accumulation rather than cellulose deposition”

✓ located verbatim in the source · abstract, characters 1847–1973 · show in text ↑
Mechanismauthor interpretationoriginal resultnegativemeasured

Claim · model-written summary

Suppression of the phenylpropanoid pathway and disruption of secondary cell wall formation provide a mechanistic basis for nitrogen-induced lodging susceptibility in rice.

Outcome
lodging susceptibilitye4
Qualifier
suppression of the phenylpropanoid pathwaye4

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

“changes in culm lodging resistance in rice”

✓ located verbatim in the source · abstract, characters 232–274 · show in text ↑
e4

“The preferential suppression of the phenylpropanoid pathway and disruption of secondary cell wall formation provide a mechanistic basis for nitrogen-induced lodging susceptibility in rice”

✓ located verbatim in the source · abstract, characters 1975–2162 · show in text ↑

nitrogen · growth · Oryza sativa (rice)Positive and negative results

511 papers: 402 positive, 109 negative or no effect

108 papers in the same direction as this paper · 402 in the opposite direction

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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.