Paper · 2025 · Plants

Mixed Ammonium-Nitrate Nutrition Regulates Enzymes, Gene Expression, and Metabolic Pathways to Improve Nitrogen Uptake, Partitioning, and Utilization Efficiency in Rice

Xianting Fan; Chusheng Lu; Zaid Khan; et al.

18
claims
42
verbatim quotes
11
citations
4
research areas
✓
Open access · cc-by

Abstract

Ammonium and nitrate nitrogen are the two main forms of inorganic nitrogen (N) available to crops. However, it is not clear how mixtures of ammonium and nitrate N affect N uptake and partitioning in major rice cultivars in southern China. This study investigated the effects of different ammonium nitrogen and nitrate nitrogen mixture treatments (100:0, 75:25, 50:50, 25:75, and 0:100) on the growth, photosynthetic characteristics, nitrogen uptake, gene expression, and yield of different rice cultivars (Mei Xiang Zhan NO. 2: MXZ2; Nan Jing Xiang Zhan: NJXZ). Rice root biomass, tiller number, and yield were increased by 69.5%, 42.5%, and 46.8%, respectively, in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment. The nitrogen content in rice roots, stems, leaves, and grains increased by 69.5%, 64.0%, 65.5%, and 17.5%, respectively. In addition, compared with MXZ2, NJXZ had a greater proportion of N allocated to leaves and grains. Analysis of root enzyme activities revealed that the 75:25 ammonium-nitrate mixed nutrient treatment increased rice root glutamine synthetase activity by an average of 35.0% and glutamate synthetase activity by an average of 52.0%. Transcriptome analysis revealed that the 75:25 mixed ammonium-nitrate nutrient treatment upregulated the expression of genes related to the nitrogen metabolism transporter pathway. Weighted correlation network analysis revealed that some differentially expressed genes (HISX and RPAB5) regulated the activities of nitrogen-metabolizing enzymes in rice and some (SAT2, CYSKP, SYIM, CHI1, and XIP1) modulated amino acid synthesis; greater expression of these genes was detected in the 75:25 ammonium-nitrate mixed nutrient treatment. The expression characteristics of the above genes were further confirmed by RT‒qPCR. Interestingly, the expression levels of the above genes were significantly correlated with the glutamate synthase activity, photosynthetic rate, and root volume. It is noteworthy that increasing the expression of the aforementioned genes coupled with nitrogen uptake was observed in the three main rice cultivars. These results suggest that the 75:25 ammonium-nitrate mixture may have increased nitrogen-metabolizing enzyme activities and promoted nitrogen uptake through the upregulated expression of nitrogen metabolism-related genes, thereby increasing tiller number and improving rice yield.

Abstract from the paper’s OpenAlex record; open copy licensed cc-by. Read at the source ↗

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Claims 18

Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

The 75:25 mixture increased rice root biomass by 69.5% over 100:0.

Treatment
75:25 ammonium-nitrate mixed treatmente0e1
Outcome
Rice root biomasse0e1
Direction
increasede0e1
Value
69.5%e0e1
Comparator
100:0 ammonium-nitrate mixed treatmente0e1

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

“Rice root biomass, tiller number, and yield were increased by 69.5%, 42.5%, and 46.8%, respectively, in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 562–764 · show in text ↑
e1

“in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 663–764 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

The 75:25 mixture increased tiller number by 42.5%.

Treatment
75:25 ammonium-nitrate mixed treatmente0e1
Outcome
tiller numbere0e1
Direction
increasede0e1
Value
42.5%e0e1
Comparator
100:0 ammonium-nitrate mixed treatmente0e1

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

“Rice root biomass, tiller number, and yield were increased by 69.5%, 42.5%, and 46.8%, respectively, in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 562–764 · show in text ↑
e1

“in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 663–764 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

The 75:25 mixture increased yield by 46.8%.

Treatment
75:25 ammonium-nitrate mixed treatmente0e1
Outcome
yielde0e1
Direction
increasede0e1
Value
46.8%e0e1
Comparator
100:0 ammonium-nitrate mixed treatmente0e1

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

“Rice root biomass, tiller number, and yield were increased by 69.5%, 42.5%, and 46.8%, respectively, in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 562–764 · show in text ↑
e1

“in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 663–764 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Rice root nitrogen content increased by 69.5%.

Treatment
75:25 ammonium-nitrate mixed treatmente0e1
Outcome
nitrogen content in rice rootse0e1
Direction
increasede0e1
Value
69.5%e0e1
Comparator
100:0 ammonium-nitrate mixed treatmente0e1

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

“The nitrogen content in rice roots, stems, leaves, and grains increased by 69.5%, 64.0%, 65.5%, and 17.5%, respectively.”

✓ located verbatim in the source · abstract, characters 765–885 · show in text ↑
e1

“in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 663–764 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Rice stem nitrogen content increased by 64.0%.

Treatment
75:25 ammonium-nitrate mixed treatmente0e1
Outcome
stemse0e1
Direction
increasede0e1
Value
64.0%e0e1
Comparator
100:0 ammonium-nitrate mixed treatmente0e1

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

“The nitrogen content in rice roots, stems, leaves, and grains increased by 69.5%, 64.0%, 65.5%, and 17.5%, respectively.”

✓ located verbatim in the source · abstract, characters 765–885 · show in text ↑
e1

“in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 663–764 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Rice leaf nitrogen content increased by 65.5%.

Treatment
75:25 ammonium-nitrate mixed treatmente0e1
Outcome
leavese0e1
Direction
increasede0e1
Value
65.5%e0e1
Comparator
100:0 ammonium-nitrate mixed treatmente0e1

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

“The nitrogen content in rice roots, stems, leaves, and grains increased by 69.5%, 64.0%, 65.5%, and 17.5%, respectively.”

✓ located verbatim in the source · abstract, characters 765–885 · show in text ↑
e1

“in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 663–764 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Rice grain nitrogen content increased by 17.5%.

Treatment
75:25 ammonium-nitrate mixed treatmente0e1
Outcome
grainse0e1
Direction
increasede0e1
Value
17.5%e0e1
Comparator
100:0 ammonium-nitrate mixed treatmente0e1

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

“The nitrogen content in rice roots, stems, leaves, and grains increased by 69.5%, 64.0%, 65.5%, and 17.5%, respectively.”

✓ located verbatim in the source · abstract, characters 765–885 · show in text ↑
e1

“in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 663–764 · show in text ↑
Observationobservedoriginal resultpositivemeasured

Claim · model-written summary

NJXZ allocated a greater proportion of nitrogen to leaves and grains than MXZ2.

Material
NJXZe0e1
Outcome
proportion of N allocated to leaves and grainse0e1
Direction
greatere0e1
Comparator
MXZ2e0e1

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

“compared with MXZ2, NJXZ had a greater proportion of N allocated to leaves and grains.”

✓ located verbatim in the source · abstract, characters 899–985 · show in text ↑
e1

“in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 663–764 · show in text ↑
t0

“Rice”

✓ located verbatim in the source · title, characters 164–168 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

The 75:25 mixture increased root glutamine synthetase activity by an average of 35.0%.

Treatment
75:25 ammonium-nitrate mixed nutrient treatmente0e1
Gene
glutamine synthetasee0e1
Outcome
rice root glutamine synthetase activitye0e1
Direction
increasede0e1
Value
35.0%e0e1
Comparator
100:0 ammonium-nitrate mixed treatmente0e1

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

“the 75:25 ammonium-nitrate mixed nutrient treatment increased rice root glutamine synthetase activity by an average of 35.0% and glutamate synthetase activity by an average of 52.0%.”

✓ located verbatim in the source · abstract, characters 1035–1217 · show in text ↑
e1

“in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 663–764 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

The 75:25 mixture increased root glutamate synthetase activity by an average of 52.0%.

Treatment
75:25 ammonium-nitrate mixed nutrient treatmente0e1
Gene
glutamate synthetasee0e1
Outcome
glutamate synthetase activitye0e1
Direction
increasede0e1
Value
52.0%e0e1
Comparator
100:0 ammonium-nitrate mixed treatmente0e1

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

“the 75:25 ammonium-nitrate mixed nutrient treatment increased rice root glutamine synthetase activity by an average of 35.0% and glutamate synthetase activity by an average of 52.0%.”

✓ located verbatim in the source · abstract, characters 1035–1217 · show in text ↑
e1

“in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 663–764 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

The 75:25 mixture upregulated genes in the nitrogen metabolism transporter pathway.

Treatment
75:25 mixed ammonium-nitrate nutrient treatmente0e1t0
Outcome
expression of genes related to the nitrogen metabolism transporter pathwaye0e1t0
Direction
upregulatede0e1t0
Comparator
100:0 ammonium-nitrate mixed treatmente0e1t0

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

“the 75:25 mixed ammonium-nitrate nutrient treatment upregulated the expression of genes related to the nitrogen metabolism transporter pathway.”

✓ located verbatim in the source · abstract, characters 1255–1398 · show in text ↑
e1

“in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 663–764 · show in text ↑
t0

“Rice”

✓ located verbatim in the source · title, characters 164–168 · show in text ↑
Mechanismauthor interpretationoriginal resultmeasured

Claim · model-written summary

HISX and RPAB5 were inferred to regulate nitrogen-metabolizing enzyme activities.

Gene
HISX and RPAB5e0e1
Outcome
activities of nitrogen-metabolizing enzymese0e1

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

“some differentially expressed genes (HISX and RPAB5) regulated the activities of nitrogen-metabolizing enzymes in rice”

✓ located verbatim in the source · abstract, characters 1451–1569 · show in text ↑
e1

“in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 663–764 · show in text ↑
Mechanismauthor interpretationoriginal resultmeasured

Claim · model-written summary

SAT2, CYSKP, SYIM, CHI1, and XIP1 were inferred to modulate amino acid synthesis.

Gene
SAT2, CYSKP, SYIM, CHI1, and XIP1e0e1t0
Outcome
amino acid synthesise0e1t0

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

“some (SAT2, CYSKP, SYIM, CHI1, and XIP1) modulated amino acid synthesis”

✓ located verbatim in the source · abstract, characters 1574–1645 · show in text ↑
e1

“in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 663–764 · show in text ↑
t0

“Rice”

✓ located verbatim in the source · title, characters 164–168 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

The named genes had greater expression under the 75:25 mixture.

Treatment
75:25 ammonium-nitrate mixed nutrient treatmente0e1t0
Outcome
expression of these genese0e1t0
Direction
greatere0e1t0
Comparator
100:0 ammonium-nitrate mixed treatmente0e1t0

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

“greater expression of these genes was detected in the 75:25 ammonium-nitrate mixed nutrient treatment.”

✓ located verbatim in the source · abstract, characters 1647–1749 · show in text ↑
e1

“in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 663–764 · show in text ↑
t0

“Rice”

✓ located verbatim in the source · title, characters 164–168 · show in text ↑
Method / resourceobservedoriginal resultmeasured

Claim · model-written summary

RT-qPCR confirmed the expression characteristics of the named genes.

Outcome
expression characteristics of the above genese0e1t0
Comparator
RT‒qPCRe0e1t0

Not reportedValue

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

“The expression characteristics of the above genes were further confirmed by RT‒qPCR.”

✓ located verbatim in the source · abstract, characters 1750–1834 · show in text ↑
e1

“in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 663–764 · show in text ↑
t0

“Rice”

✓ located verbatim in the source · title, characters 164–168 · show in text ↑
Associationobservedoriginal resultsignificant (as stated)measured

Claim · model-written summary

Expression levels of the named genes correlated significantly with glutamate synthase activity, photosynthetic rate, and root volume.

Gene
above genese0e1t0
Outcome
glutamate synthase activity, photosynthetic rate, and root volumee0e1t0

Not reportedDirection

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

“the expression levels of the above genes were significantly correlated with the glutamate synthase activity, photosynthetic rate, and root volume.”

✓ located verbatim in the source · abstract, characters 1850–1996 · show in text ↑
e1

“in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 663–764 · show in text ↑
t0

“Rice”

✓ located verbatim in the source · title, characters 164–168 · show in text ↑
Associationobservedoriginal resultmeasured

Claim · model-written summary

Increased expression of the named genes accompanied nitrogen uptake in three main rice cultivars.

Gene
aforementioned genese0e1
Outcome
nitrogen uptakee0e1
Direction
increasinge0e1
Qualifier
three main rice cultivarse0e1

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

“increasing the expression of the aforementioned genes coupled with nitrogen uptake was observed in the three main rice cultivars.”

✓ located verbatim in the source · abstract, characters 2019–2148 · show in text ↑
e1

“in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 663–764 · show in text ↑
Mechanismauthor interpretationoriginal resultmeasured

Claim · model-written summary

The authors infer that the 75:25 mixture promotes nitrogen uptake through upregulated nitrogen-metabolism genes and increased enzyme activity, contributing to more tillers and yield.

Treatment
75:25 ammonium-nitrate mixturee0e1
Outcome
nitrogen uptakee0e1
Qualifier
through the upregulated expression of nitrogen metabolism-related genese0e1

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

“the 75:25 ammonium-nitrate mixture may have increased nitrogen-metabolizing enzyme activities and promoted nitrogen uptake through the upregulated expression of nitrogen metabolism-related genes, thereby increasing tiller number and improving rice yield.”

✓ located verbatim in the source · abstract, characters 2176–2430 · show in text ↑
e1

“in the 75:25 ammonium-nitrate mixed treatment compared to the 100:0 ammonium-nitrate mixed treatment.”

✓ located verbatim in the source · abstract, characters 663–764 · show in text ↑

nitrogen · physiology: enzyme activity · Oryza sativa (rice)Positive and negative results

36 papers: 28 positive, 8 negative or no effect

27 papers in the same direction as this paper · 7 in the opposite direction · 1 reporting no difference from an alternative

  1. negative
  2. no effect
  3. negative

32 papers more in this group

See all 36 papers →

nitrogen · molecular: expression · Oryza sativa (rice)Positive and negative results

45 papers: 29 positive, 16 negative or no effect

28 papers in the same direction as this paper · 16 in the opposite direction

  1. no effect
  2. negative
  3. negative

41 papers more in this group

See all 45 papers →

nitrogen · nutrient: content · Oryza sativa (rice)Positive and negative results

464 papers: 409 positive, 55 negative or no effect

408 papers in the same direction as this paper · 52 in the opposite direction · 3 reporting no difference from an alternative

  1. negative
  2. no effect
  3. negative

460 papers more in this group

See all 464 papers →

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

511 papers: 402 positive, 109 negative or no effect

401 papers in the same direction as this paper · 101 in the opposite direction · 8 reporting no difference from an alternative

  1. negative
  2. negative
  3. negative

507 papers more in this group

See all 511 papers →

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

2,045 papers: 1,803 positive, 242 negative or no effect

1,802 papers in the same direction as this paper · 193 in the opposite direction · 49 reporting no difference from an alternative

  1. negative
  2. negative
  3. negative

2,041 papers more in this group

See all 2,045 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.