Paper · 2026 · Agriculture Ecosystems & Environment

Long-term nitrogen fertilizer improves nitrogen use efficiency and wheat yield by regulating the interactions of rhizosphere soil-root-microbes

Xiaoyan Zhang; Xiaojie Han; Qingcheng Li; et al.

12
claims
36
verbatim quotes
2
citations
3
research areas
✓
Open access · cc-by-nc-nd

Abstract

Nitrogen (N) fertilizer is widely used in agroecosystems to maintain and improve soil fertility while affecting soil microbial communities. The ability of root to absorb nutrients is critically mediated by these soil microbial communities, ultimately determining crop growth and yield formation. Thus, comprehensively understanding the rhizosphere root-soil-microbe interaction mechanism is critical for synergistically regulating nitrogen use efficiency (NUE) and yield. Here, a 13-year field experiment in a wheat-maize rotation system was conducted to explore the effects of N fertilizer on soil physicochemical properties, root parameters and rhizosphere microorganisms. Compared with N0 (0 kg ha −1 ), moderate N application (180–240 kg ha −1 ) optimized soil nutrient availability, as evidenced by increasing available nitrogen (AN) and ammonium-N (NH 4 + -N) by 17.42 %–27.33 % and 12.24 %–6 2.49 %, respectively, and enhanced root morphology, with root vessel diameter (RVD), root length density (RLD), and root weight density (RWD) rising by 5.12 %–21.57 %, 14.17 %–65.77 %, and 3.46 %–62.10 %, respectively. Consequently, N uptake and metabolic enzyme activities were significantly boosted. These root modifications, in turn, reshaped the rhizosphere microbiome by enriching beneficial bacterial and fungal taxa, Pseudomonadota and Basidiomycota increased by an average of 20.93 % and 73.28 %, facilitating efficient N cycling and organic matter decomposition. Structural equation model (SEM) revealed that soil N and root enzyme activity mediated microbial community compositions and diversity, which collectively enhanced both yield and NUE. Overall, our findings highlight the critical role of root-microbe feedback in driving N efficiency and propose that optimizing N inputs can sustainably enhance crop productivity by leveraging soil-plant-microbial synergies. • A 13-year experiment clarified the synergistic regulation of wheat yield and NUE by rhizosphere soil-root-microbe interaction . • Optimal N fertilization enhanced soil nutrient availability and root traits, improved root N metabolic enzyme activities. • Rhizosphere microbiomes promote N cycling and organic matter decomposition via beneficial taxa. • SEM revealed soil and root traits mediated microbial community characteristics, collectively enhancing wheat yield and NUE.

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Claims 12

Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Moderate N application increased available soil nitrogen by 17.42%–27.33% relative to N0.

Treatment
moderate N application (180–240 kg ha −1 )e0e1t0
Outcome
available nitrogen (AN)e0e1t0
Direction
increasinge0e1t0
Value
17.42 %–27.33 %e0e1t0
Comparator
N0e0e1t0

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

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Compared with N0 (0 kg ha −1 ), moderate N application (180–240 kg ha −1 ) optimized soil nutrient availability, as evidenced by increasing available nitrogen (AN) and ammonium-N (NH 4 + -N) by 17.42 %–27.33 % and 12.24 %–6 2.49 %, respectively, and enhanced root morphology, with root vessel diameter (RVD), root length density (RLD), and root weight density (RWD) rising by 5.12 %–21.57 %, 14.17 %–65.77 %, and 3.46 %–62.10 %, respectively.”

✓ located verbatim in the source · abstract, characters 675–1117 · show in text ↑
e1

“Compared with N0 (0 kg ha −1 ), moderate N application (180–240 kg ha −1 ) optimized soil nutrient availability, as evidenced by increasing available nitrogen (AN) and ammonium-N (NH 4 + -N) by 17.42 %–27.33 % and 12.24 %–6 2.49 %, respectively, and enhanced root morphology, with root vessel diameter (RVD), root length density (RLD), and root weight density (RWD) rising by 5.12 %–21.57 %, 14.17 %–65.77 %, and 3.46 %–62.10 %, respectively.”

✓ located verbatim in the source · abstract, characters 675–1117 · show in text ↑
t0

“wheat”

✓ located verbatim in the source · title, characters 67–72 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Moderate N application increased ammonium-N by 12.24%–62.49% relative to N0.

Treatment
moderate N application (180–240 kg ha −1 )e0e1t0
Outcome
ammonium-N (NH 4 + -N)e0e1t0
Direction
increasinge0e1t0
Value
12.24 %–6 2.49 %e0e1t0
Comparator
N0e0e1t0

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

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Compared with N0 (0 kg ha −1 ), moderate N application (180–240 kg ha −1 ) optimized soil nutrient availability, as evidenced by increasing available nitrogen (AN) and ammonium-N (NH 4 + -N) by 17.42 %–27.33 % and 12.24 %–6 2.49 %, respectively, and enhanced root morphology, with root vessel diameter (RVD), root length density (RLD), and root weight density (RWD) rising by 5.12 %–21.57 %, 14.17 %–65.77 %, and 3.46 %–62.10 %, respectively.”

✓ located verbatim in the source · abstract, characters 675–1117 · show in text ↑
e1

“Compared with N0 (0 kg ha −1 ), moderate N application (180–240 kg ha −1 ) optimized soil nutrient availability, as evidenced by increasing available nitrogen (AN) and ammonium-N (NH 4 + -N) by 17.42 %–27.33 % and 12.24 %–6 2.49 %, respectively, and enhanced root morphology, with root vessel diameter (RVD), root length density (RLD), and root weight density (RWD) rising by 5.12 %–21.57 %, 14.17 %–65.77 %, and 3.46 %–62.10 %, respectively.”

✓ located verbatim in the source · abstract, characters 675–1117 · show in text ↑
t0

“wheat”

✓ located verbatim in the source · title, characters 67–72 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Moderate N application increased root vessel diameter by 5.12%–21.57%.

Treatment
moderate N application (180–240 kg ha −1 )e0e1t0
Outcome
root vessel diameter (RVD)e0e1t0
Direction
risinge0e1t0
Value
5.12 %–21.57 %e0e1t0
Comparator
N0e0e1t0

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

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“root vessel diameter (RVD), root length density (RLD), and root weight density (RWD) rising by 5.12 %–21.57 %, 14.17 %–65.77 %, and 3.46 %–62.10 %, respectively.”

✓ located verbatim in the source · abstract, characters 956–1117 · show in text ↑
e1

“Compared with N0 (0 kg ha −1 ), moderate N application (180–240 kg ha −1 ) optimized soil nutrient availability, as evidenced by increasing available nitrogen (AN) and ammonium-N (NH 4 + -N) by 17.42 %–27.33 % and 12.24 %–6 2.49 %, respectively, and enhanced root morphology, with root vessel diameter (RVD), root length density (RLD), and root weight density (RWD) rising by 5.12 %–21.57 %, 14.17 %–65.77 %, and 3.46 %–62.10 %, respectively.”

✓ located verbatim in the source · abstract, characters 675–1117 · show in text ↑
t0

“wheat”

✓ located verbatim in the source · title, characters 67–72 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Moderate N application increased root length density by 14.17%–65.77%.

Treatment
moderate N application (180–240 kg ha −1 )e0e1t0
Outcome
root length density (RLD)e0e1t0
Direction
risinge0e1t0
Value
14.17 %–65.77 %e0e1t0
Comparator
N0e0e1t0

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

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“root vessel diameter (RVD), root length density (RLD), and root weight density (RWD) rising by 5.12 %–21.57 %, 14.17 %–65.77 %, and 3.46 %–62.10 %, respectively.”

✓ located verbatim in the source · abstract, characters 956–1117 · show in text ↑
e1

“Compared with N0 (0 kg ha −1 ), moderate N application (180–240 kg ha −1 ) optimized soil nutrient availability, as evidenced by increasing available nitrogen (AN) and ammonium-N (NH 4 + -N) by 17.42 %–27.33 % and 12.24 %–6 2.49 %, respectively, and enhanced root morphology, with root vessel diameter (RVD), root length density (RLD), and root weight density (RWD) rising by 5.12 %–21.57 %, 14.17 %–65.77 %, and 3.46 %–62.10 %, respectively.”

✓ located verbatim in the source · abstract, characters 675–1117 · show in text ↑
t0

“wheat”

✓ located verbatim in the source · title, characters 67–72 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Moderate N application increased root weight density by 3.46%–62.10%.

Treatment
moderate N application (180–240 kg ha −1 )e0e1t0
Outcome
root weight density (RWD)e0e1t0
Direction
risinge0e1t0
Value
3.46 %–62.10 %e0e1t0
Comparator
N0e0e1t0

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

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“root vessel diameter (RVD), root length density (RLD), and root weight density (RWD) rising by 5.12 %–21.57 %, 14.17 %–65.77 %, and 3.46 %–62.10 %, respectively.”

✓ located verbatim in the source · abstract, characters 956–1117 · show in text ↑
e1

“Compared with N0 (0 kg ha −1 ), moderate N application (180–240 kg ha −1 ) optimized soil nutrient availability, as evidenced by increasing available nitrogen (AN) and ammonium-N (NH 4 + -N) by 17.42 %–27.33 % and 12.24 %–6 2.49 %, respectively, and enhanced root morphology, with root vessel diameter (RVD), root length density (RLD), and root weight density (RWD) rising by 5.12 %–21.57 %, 14.17 %–65.77 %, and 3.46 %–62.10 %, respectively.”

✓ located verbatim in the source · abstract, characters 675–1117 · show in text ↑
t0

“wheat”

✓ located verbatim in the source · title, characters 67–72 · show in text ↑
Intervention effectobservedoriginal resultpositivesignificant (as stated)measured

Claim · model-written summary

Moderate N application significantly boosted N uptake and metabolic enzyme activities.

Treatment
moderate N application (180–240 kg ha −1 )e0e1t0
Outcome
N uptake and metabolic enzyme activitiese0e1t0
Direction
boostede0e1t0
Comparator
N0e0e1t0

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

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Consequently, N uptake and metabolic enzyme activities were significantly boosted.”

✓ located verbatim in the source · abstract, characters 1118–1200 · show in text ↑
e1

“Compared with N0 (0 kg ha −1 ), moderate N application (180–240 kg ha −1 ) optimized soil nutrient availability, as evidenced by increasing available nitrogen (AN) and ammonium-N (NH 4 + -N) by 17.42 %–27.33 % and 12.24 %–6 2.49 %, respectively, and enhanced root morphology, with root vessel diameter (RVD), root length density (RLD), and root weight density (RWD) rising by 5.12 %–21.57 %, 14.17 %–65.77 %, and 3.46 %–62.10 %, respectively.”

✓ located verbatim in the source · abstract, characters 675–1117 · show in text ↑
t0

“wheat”

✓ located verbatim in the source · title, characters 67–72 · show in text ↑
Associationobservedoriginal resultpositivemeasured

Claim · model-written summary

Root changes enriched beneficial bacterial and fungal taxa in the rhizosphere.

Organ / stage
roote0e1t0
Interactor
beneficial bacterial and fungal taxae0e1t0symbiont
Outcome
beneficial bacterial and fungal taxae0e1t0
Direction
enrichinge0e1t0

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

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“These root modifications, in turn, reshaped the rhizosphere microbiome by enriching beneficial bacterial and fungal taxa, Pseudomonadota and Basidiomycota increased by an average of 20.93 % and 73.28 %, facilitating efficient N cycling and organic matter decomposition.”

✓ located verbatim in the source · abstract, characters 1201–1470 · show in text ↑
e1

“Compared with N0 (0 kg ha −1 ), moderate N application (180–240 kg ha −1 ) optimized soil nutrient availability, as evidenced by increasing available nitrogen (AN) and ammonium-N (NH 4 + -N) by 17.42 %–27.33 % and 12.24 %–6 2.49 %, respectively, and enhanced root morphology, with root vessel diameter (RVD), root length density (RLD), and root weight density (RWD) rising by 5.12 %–21.57 %, 14.17 %–65.77 %, and 3.46 %–62.10 %, respectively.”

✓ located verbatim in the source · abstract, characters 675–1117 · show in text ↑
t0

“wheat”

✓ located verbatim in the source · title, characters 67–72 · show in text ↑
Associationobservedoriginal resultpositivemeasured

Claim · model-written summary

Pseudomonadota increased by an average of 20.93% following root changes.

Organ / stage
roote0e1t0
Interactor
PseudomonadotaPseudomonadotae0e1t0symbiont
Outcome
Pseudomonadotae0e1t0
Direction
increasede0e1t0
Value
20.93 %e0e1t0

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

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Pseudomonadota and Basidiomycota increased by an average of 20.93 % and 73.28 %”

✓ located verbatim in the source · abstract, characters 1323–1402 · show in text ↑
e1

“Compared with N0 (0 kg ha −1 ), moderate N application (180–240 kg ha −1 ) optimized soil nutrient availability, as evidenced by increasing available nitrogen (AN) and ammonium-N (NH 4 + -N) by 17.42 %–27.33 % and 12.24 %–6 2.49 %, respectively, and enhanced root morphology, with root vessel diameter (RVD), root length density (RLD), and root weight density (RWD) rising by 5.12 %–21.57 %, 14.17 %–65.77 %, and 3.46 %–62.10 %, respectively.”

✓ located verbatim in the source · abstract, characters 675–1117 · show in text ↑
t0

“wheat”

✓ located verbatim in the source · title, characters 67–72 · show in text ↑
Associationobservedoriginal resultpositivemeasured

Claim · model-written summary

Basidiomycota increased by an average of 73.28% following root changes.

Organ / stage
roote0e1t0
Interactor
BasidiomycotaBasidiomycotae0e1t0symbiont
Outcome
Basidiomycotae0e1t0
Direction
increasede0e1t0
Value
73.28 %e0e1t0

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

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Pseudomonadota and Basidiomycota increased by an average of 20.93 % and 73.28 %”

✓ located verbatim in the source · abstract, characters 1323–1402 · show in text ↑
e1

“Compared with N0 (0 kg ha −1 ), moderate N application (180–240 kg ha −1 ) optimized soil nutrient availability, as evidenced by increasing available nitrogen (AN) and ammonium-N (NH 4 + -N) by 17.42 %–27.33 % and 12.24 %–6 2.49 %, respectively, and enhanced root morphology, with root vessel diameter (RVD), root length density (RLD), and root weight density (RWD) rising by 5.12 %–21.57 %, 14.17 %–65.77 %, and 3.46 %–62.10 %, respectively.”

✓ located verbatim in the source · abstract, characters 675–1117 · show in text ↑
t0

“wheat”

✓ located verbatim in the source · title, characters 67–72 · show in text ↑
Associationobservedoriginal resultsimulated

Claim · model-written summary

The SEM linked soil N and root enzyme activity to microbial community composition and diversity.

Organ / stage
roote0e1t0
Compound
soil Ne0e1t0
Outcome
microbial community compositions and diversitye0e1t0
Qualifier
root enzyme activitye0e1t0

Not reportedDirection

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

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Structural equation model (SEM) revealed that soil N and root enzyme activity mediated microbial community compositions and diversity, which collectively enhanced both yield and NUE.”

✓ located verbatim in the source · abstract, characters 1471–1653 · show in text ↑
e1

“Compared with N0 (0 kg ha −1 ), moderate N application (180–240 kg ha −1 ) optimized soil nutrient availability, as evidenced by increasing available nitrogen (AN) and ammonium-N (NH 4 + -N) by 17.42 %–27.33 % and 12.24 %–6 2.49 %, respectively, and enhanced root morphology, with root vessel diameter (RVD), root length density (RLD), and root weight density (RWD) rising by 5.12 %–21.57 %, 14.17 %–65.77 %, and 3.46 %–62.10 %, respectively.”

✓ located verbatim in the source · abstract, characters 675–1117 · show in text ↑
t0

“wheat”

✓ located verbatim in the source · title, characters 67–72 · show in text ↑
Associationobservedoriginal resultpositivesimulated

Claim · model-written summary

The SEM linked microbial community composition and diversity to enhanced yield.

Interactor
microbial community compositionse0e1t0symbiont
Outcome
yielde0e1t0
Direction
enhancede0e1t0

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

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Structural equation model (SEM) revealed that soil N and root enzyme activity mediated microbial community compositions and diversity, which collectively enhanced both yield and NUE.”

✓ located verbatim in the source · abstract, characters 1471–1653 · show in text ↑
e1

“Compared with N0 (0 kg ha −1 ), moderate N application (180–240 kg ha −1 ) optimized soil nutrient availability, as evidenced by increasing available nitrogen (AN) and ammonium-N (NH 4 + -N) by 17.42 %–27.33 % and 12.24 %–6 2.49 %, respectively, and enhanced root morphology, with root vessel diameter (RVD), root length density (RLD), and root weight density (RWD) rising by 5.12 %–21.57 %, 14.17 %–65.77 %, and 3.46 %–62.10 %, respectively.”

✓ located verbatim in the source · abstract, characters 675–1117 · show in text ↑
t0

“wheat”

✓ located verbatim in the source · title, characters 67–72 · show in text ↑
Associationobservedoriginal resultpositivesimulated

Claim · model-written summary

The SEM linked microbial community composition and diversity to enhanced NUE.

Interactor
microbial community compositionse0e1t0symbiont
Outcome
NUEe0e1t0
Direction
enhancede0e1t0

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

Triticum aestivum
Triticum aestivum · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Structural equation model (SEM) revealed that soil N and root enzyme activity mediated microbial community compositions and diversity, which collectively enhanced both yield and NUE.”

✓ located verbatim in the source · abstract, characters 1471–1653 · show in text ↑
e1

“Compared with N0 (0 kg ha −1 ), moderate N application (180–240 kg ha −1 ) optimized soil nutrient availability, as evidenced by increasing available nitrogen (AN) and ammonium-N (NH 4 + -N) by 17.42 %–27.33 % and 12.24 %–6 2.49 %, respectively, and enhanced root morphology, with root vessel diameter (RVD), root length density (RLD), and root weight density (RWD) rising by 5.12 %–21.57 %, 14.17 %–65.77 %, and 3.46 %–62.10 %, respectively.”

✓ located verbatim in the source · abstract, characters 675–1117 · show in text ↑
t0

“wheat”

✓ located verbatim in the source · title, characters 67–72 · show in text ↑

nitrogen · soil: biology · Triticum aestivum (wheat)Positive and negative results

29 papers: 23 positive, 6 negative or no effect

22 papers in the same direction as this paper · 5 in the opposite direction · 1 reporting no difference from an alternative

  1. negative
  2. no effect
  3. negative

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nitrogen · soil: chemistry · Triticum aestivum (wheat)Positive and negative results

49 papers: 38 positive, 11 negative or no effect

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

  1. no effect
  2. no effect
  3. negative

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nitrogen · growth · Triticum aestivum (wheat)Positive and negative results

551 papers: 426 positive, 125 negative or no effect

425 papers in the same direction as this paper · 116 in the opposite direction · 9 reporting no difference from an alternative

  1. positive
  2. negative
  3. no effect

547 papers more in this group

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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.