Paper · 2026 · The Crop Journal

Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots

Yang Zhou; Pengjia Wu; Muhammad Fraz Ali; et al.

11
claims
22
verbatim quotes
0
citations
4
research areas
✓
Open access · cc-by-nc-nd

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.

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

The text the model read, with the source record’s inline markup (italics, sub- and superscripts) rendered and its character entities decoded; quote offsets refer to the raw text.

Claims 11

Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Layered nitrogen placement redistributed soil nitrate nitrogen, organic carbon, available phosphorus, and enzyme activities relative to shallow placement.

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.

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

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”

✓ located verbatim in the source · abstract, characters 250–455 · show in text ↑
t0

“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”

✓ located verbatim in the source · title, characters 0–145 · show in text ↑
Associationobservedoriginal resultpositivemeasured

Claim · model-written summary

The soil changes associated with layered nitrogen placement were linked to enhanced deeper root growth.

Organ / stage
roote0
Outcome
root growth in deeper soile0
Direction
enhancede0

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 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”

✓ located verbatim in the source · abstract, characters 457–654 · show in text ↑
t0

“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”

✓ located verbatim in the source · title, characters 0–145 · show in text ↑
Associationobservedoriginal resultpositivemeasured

Claim · model-written summary

Specific root length and area were greater in deeper soil under the changed soil conditions.

Organ / stage
roote0
Outcome
specific root length and areae0
Direction
greatere0

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 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”

✓ located verbatim in the source · abstract, characters 457–654 · show in text ↑
t0

“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”

✓ located verbatim in the source · title, characters 0–145 · show in text ↑
Associationobservedoriginal resultpositivemeasured

Claim · model-written summary

Root nitrate reductase activity was higher under the changed soil conditions.

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.

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

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”

✓ located verbatim in the source · abstract, characters 457–654 · show in text ↑
t0

“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”

✓ located verbatim in the source · title, characters 0–145 · show in text ↑
Associationobservedoriginal resultpositivemeasured

Claim · model-written summary

Root nitrogen and phosphorus concentrations increased under the changed soil conditions.

Organ / stage
roote0
Outcome
root N and P concentrationse0
Direction
increasede0

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 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”

✓ located verbatim in the source · abstract, characters 457–654 · show in text ↑
t0

“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”

✓ located verbatim in the source · title, characters 0–145 · show in text ↑
Observationobservedoriginal resultpositivemeasured

Claim · model-written summary

Root functional strategies shifted toward greater acquisition.

Outcome
more acquisitive strategiese0

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 functional analyses indicated a shift toward more acquisitive strategies”

✓ located verbatim in the source · abstract, characters 656–733 · show in text ↑
t0

“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”

✓ located verbatim in the source · title, characters 0–145 · show in text ↑
Observationobservedoriginal resultpositivemeasured

Claim · model-written summary

Genes related to lateral root formation and root hair development were upregulated.

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.

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

Verbatim

e0

“supported by the upregulation of genes related to lateral root formation and root hair development”

✓ located verbatim in the source · abstract, characters 735–833 · show in text ↑
t0

“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”

✓ located verbatim in the source · title, characters 0–145 · show in text ↑
Associationobservedoriginal resultpositivemeasured

Claim · model-written summary

The 8–16 cm soil layer explained 43.0% of yield variation and contributed most to yield formation.

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.

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

Verbatim

e0

“The 8–16 cm soil layer contributed most to yield formation, explaining 43.0% of yield variation”

✓ located verbatim in the source · abstract, characters 835–930 · show in text ↑
t0

“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”

✓ located verbatim in the source · title, characters 0–145 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Layered nitrogen placement increased subsoil nutrient availability, root functional differentiation, and depth-specific transcriptional activation.

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.

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

Verbatim

e0

“Layered N placement increased subsoil nutrient availability, root functional differentiation, and depth-specific transcriptional activation, promoting nutrient accumulation and increasing grain yield”

✓ located verbatim in the source · abstract, characters 932–1131 · show in text ↑
t0

“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”

✓ located verbatim in the source · title, characters 0–145 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Layered nitrogen placement promoted nutrient accumulation.

Treatment
Layered N placemente0
Outcome
nutrient accumulatione0
Direction
promotinge0

Not reportedComparator

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

“Layered N placement increased subsoil nutrient availability, root functional differentiation, and depth-specific transcriptional activation, promoting nutrient accumulation and increasing grain yield”

✓ located verbatim in the source · abstract, characters 932–1131 · show in text ↑
t0

“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”

✓ located verbatim in the source · title, characters 0–145 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Layered nitrogen placement increased grain yield.

Treatment
Layered N placemente0
Outcome
grain yielde0
Direction
increasinge0

Not reportedComparator

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

“Layered N placement increased subsoil nutrient availability, root functional differentiation, and depth-specific transcriptional activation, promoting nutrient accumulation and increasing grain yield”

✓ located verbatim in the source · abstract, characters 932–1131 · show in text ↑
t0

“Layered nitrogen placement increases wheat yield by transcriptionally regulating root functional strategies and reshaping soil enzymatic hotspots”

✓ located verbatim in the source · title, characters 0–145 · show in text ↑