Abstract
The objective of this study was to identify the soil–root regulatory mechanisms by which nitrogen (N) application at varying soil depths increases nutrient use efficiency and grain yield in wheat. In a two-year field experiment on the Loess Plateau, 240 kg N ha −1 applied in a 1:2:1 ratio to the 8-, 16-, and 24-cm soil layers redistributed NO 3 − –N, soil organic carbon, available P, and enzyme activities relative to application only at the 8-cm depth. These changes were associated with enhanced root growth in deeper soil, including greater specific root length and area, higher nitrate reductase activity, and increased root N and P concentrations. Root functional analyses indicated a shift toward more acquisitive strategies, supported by the upregulation of genes related to lateral root formation and root hair development. The 8–16 cm soil layer contributed most to yield formation, explaining 43.0% of yield variation. Layered N placement increased subsoil nutrient availability, root functional differentiation, and depth-specific transcriptional activation, promoting nutrient accumulation and increasing grain yield.
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Claims 11
Claim · model-written summary
Layered nitrogen placement redistributed soil nitrate nitrogen, organic carbon, available phosphorus, and enzyme activities relative to shallow placement.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- 240 kg N ha −1 applied in a 1:2:1 ratio to the 8-, 16-, and 24-cm soil layerse0
- Outcome
- NO 3 − –N, soil organic carbon, available P, and enzyme activitiese0
- Direction
- redistributede0
- Comparator
- application only at the 8-cm depthe0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“240 kg N ha −1 applied in a 1:2:1 ratio to the 8-, 16-, and 24-cm soil layers redistributed NO 3 − –N, soil organic carbon, available P, and enzyme activities relative to application only at the 8-cm depth”
t0“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”
Claim · model-written summary
The soil changes associated with layered nitrogen placement were linked to enhanced deeper root growth.
- Taxon
- wheat
Triticum aestivumt0 - Organ / stage
- roote0
- Outcome
- root growth in deeper soile0
- Direction
- enhancede0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“These changes were associated with enhanced root growth in deeper soil, including greater specific root length and area, higher nitrate reductase activity, and increased root N and P concentrations”
t0“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”
Claim · model-written summary
Specific root length and area were greater in deeper soil under the changed soil conditions.
- Taxon
- wheat
Triticum aestivumt0 - Organ / stage
- roote0
- Outcome
- specific root length and areae0
- Direction
- greatere0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“These changes were associated with enhanced root growth in deeper soil, including greater specific root length and area, higher nitrate reductase activity, and increased root N and P concentrations”
t0“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”
Claim · model-written summary
Root nitrate reductase activity was higher under the changed soil conditions.
- Taxon
- wheat
Triticum aestivumt0 - Organ / stage
- roote0
- Gene
- nitrate reductasee0
- Outcome
- nitrate reductase activitye0
- Direction
- highere0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“These changes were associated with enhanced root growth in deeper soil, including greater specific root length and area, higher nitrate reductase activity, and increased root N and P concentrations”
t0“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”
Claim · model-written summary
Root nitrogen and phosphorus concentrations increased under the changed soil conditions.
- Taxon
- wheat
Triticum aestivumt0 - Organ / stage
- roote0
- Outcome
- root N and P concentrationse0
- Direction
- increasede0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“These changes were associated with enhanced root growth in deeper soil, including greater specific root length and area, higher nitrate reductase activity, and increased root N and P concentrations”
t0“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”
Claim · model-written summary
Root functional strategies shifted toward greater acquisition.
- Taxon
- wheat
Triticum aestivumt0 - Outcome
- more acquisitive strategiese0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Root functional analyses indicated a shift toward more acquisitive strategies”
t0“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”
Claim · model-written summary
Genes related to lateral root formation and root hair development were upregulated.
- Taxon
- wheat
Triticum aestivumt0 - Organ / stage
- lateral root formation and root hair developmente0
- Outcome
- upregulation of genes related to lateral root formation and root hair developmente0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“supported by the upregulation of genes related to lateral root formation and root hair development”
t0“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”
Claim · model-written summary
The 8–16 cm soil layer explained 43.0% of yield variation and contributed most to yield formation.
- Taxon
- wheat
Triticum aestivumt0 - Outcome
- yield variatione0
- Direction
- contributed moste0
- Value
- 43.0%e0
- Setting
- 8–16 cm soil layere0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The 8–16 cm soil layer contributed most to yield formation, explaining 43.0% of yield variation”
t0“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”
Claim · model-written summary
Layered nitrogen placement increased subsoil nutrient availability, root functional differentiation, and depth-specific transcriptional activation.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- Layered N placemente0
- Outcome
- subsoil nutrient availability, root functional differentiation, and depth-specific transcriptional activatione0
- Direction
- increasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Layered N placement increased subsoil nutrient availability, root functional differentiation, and depth-specific transcriptional activation, promoting nutrient accumulation and increasing grain yield”
t0“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”
Claim · model-written summary
Layered nitrogen placement promoted nutrient accumulation.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- Layered N placemente0
- Outcome
- nutrient accumulatione0
- Direction
- promotinge0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Layered N placement increased subsoil nutrient availability, root functional differentiation, and depth-specific transcriptional activation, promoting nutrient accumulation and increasing grain yield”
t0“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”
Claim · model-written summary
Layered nitrogen placement increased grain yield.
- Taxon
- wheat
Triticum aestivumt0 - Treatment
- Layered N placemente0
- Outcome
- grain yielde0
- Direction
- increasinge0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Layered N placement increased subsoil nutrient availability, root functional differentiation, and depth-specific transcriptional activation, promoting nutrient accumulation and increasing grain yield”
t0“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”