Paper · 2026 · Scientia Horticulturae

Effects of summer nitrogen application rate on citrus shoot-root-soil interactions

Wenfeng Li; Qianzi Wang; Yongzhong Liu; et al.

18
claims
36
verbatim quotes
1
citations
5
research areas
✓
Open access · cc-by-nc-nd

Abstract

• Identified a critical pure nitrogen threshold of 3–4 g plant −1 for regulating summer shoot growth and maintaining physiological health in potted 'Newhall' navel orange seedlings. • Elucidated the complex regulatory mechanism of summer shoot growth, demonstrating joint mediation by root-zone nitrogen availability, root system architectural plasticity, and the intricate ABA-Pro-Chl metabolic balance. • Developed a robust multiple regression predictive model integrating soil available nitrogen (NO 3 − -N and NH 4 + -N) and root morphological parameters to effectively forecast summer shoot growth dynamics (R 2 = 0.53–0.72). Nitrogen (N) is the primary nutrient that initiates the summer-flush (SF) in citrus trees. However, the integrated "soil-root-shoot" feedback mechanism regulating SF intensity remains poorly understood in fertigation-based systems. To quantify dose-dependent effects on SF morphogenesis, nutrient allocation, root-tissue, and soil N concentrations, we conducted a greenhouse pot experiment with two-year-old 'Newhall' navel orange grafted onto Poncirus trifoliata rootstocks. The plants were subjected to six summer N rates (pure N 1, 2, 3, 4, 5, and 7 g plant −1 ). The results showed that: (1) Reducing N supply progressively delayed SF emergence and advanced maturation, thereby shortening the flush period; (2) Compared to high N treatment (5–7 g plant −1 ), moderate N (3–4 g plant −1 ) application treatment significantly suppressed SF growth, reducing shoot number, length, basal diameter, leaf number, emergence rate and biomass by 42.86 %, 8.99 %, 9.76 %, 16.74 %, 29.37 %, and 47.59 %, respectively. Conversely, this optimal N range maximized the net increments in plant height (1.96 cm) and main stem diameter (1.52 cm); (3) Within this range, the summer shoots maintained medium-to-high levels of nutrient content (N, phosphorus, potassium) and physiologically active substances (abscisic acid, chlorophyll, proline), while root growth was moderated to a medium-low level. Meanwhile, the nutrient status of the root zone soil and roots remained in the medium-to-high range; (4) Root zone soil ammonium nitrogen and nitrate nitrogen concentrations and root length density were strongly correlated with SF architectural traits. Multiple regression models incorporating these variables explained >50 % of the variance in SF phenotype. This regulation was attributed to an N-driven "soil-root-shoot" feedback mechanism, where the root zone inorganic N pool, root morphology, and the balance of key shoot physiologically active substances collectively defined summer-flush growth patterns. Therefore, it is advisable to apply 3–4 g N plant −1 as the ideal summer rate for potted citrus seedlings in controlled environments. This effectively maintains a balance between vegetative growth and nutrient homeostasis through the identified "soil-root-shoot" feedback loop. However, these conclusions need to be verified in orchard-grown, fruit-bearing trees to substantiate their relevance in commercial cultivation systems before broader implementation.

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 18

Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Reducing nitrogen supply progressively delayed summer flush emergence in citrus.

Taxon
citrusCitrust0
Treatment
Reducing N supplye1
Outcome
SF emergencee1
Direction
delayede1

Not reportedComparator

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

Citrus
Citrus · Taxon · picture of the genus (one of its species)Ellen Levy Finch ( Elf ) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“citrus”

✓ located verbatim in the source · title, characters 47–53 · show in text ↑
e1

“Reducing N supply progressively delayed SF emergence and advanced maturation, thereby shortening the flush period”

✓ located verbatim in the source · abstract, characters 1227–1340 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Reducing nitrogen supply advanced maturation of the summer flush, shortening the flush period.

Taxon
citrusCitrust0
Treatment
Reducing N supplye1
Outcome
maturatione1
Direction
advancede1

Not reportedComparator

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

Citrus
Citrus · Taxon · picture of the genus (one of its species)Ellen Levy Finch ( Elf ) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“citrus”

✓ located verbatim in the source · title, characters 47–53 · show in text ↑
e1

“Reducing N supply progressively delayed SF emergence and advanced maturation, thereby shortening the flush period”

✓ located verbatim in the source · abstract, characters 1227–1340 · show in text ↑
Intervention effectobservedoriginal resultsignificant (as stated)measured

Claim · model-written summary

Moderate N (3–4 g per plant) significantly suppressed summer flush growth relative to high N (5–7 g per plant), reducing shoot number by 42.86 %.

Taxon
citrusCitrust0
Treatment
moderate N (3–4 g plant −1 ) application treatmente1
Outcome
shoot numbere1
Direction
reducinge1
Value
42.86 %e1
Comparator
high N treatment (5–7 g plant −1 )e1

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

Citrus
Citrus · Taxon · picture of the genus (one of its species)Ellen Levy Finch ( Elf ) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“citrus”

✓ located verbatim in the source · title, characters 47–53 · show in text ↑
e1

“Compared to high N treatment (5–7 g plant −1 ), moderate N (3–4 g plant −1 ) application treatment significantly suppressed SF growth, reducing shoot number, length, basal diameter, leaf number, emergence rate and biomass by 42.86 %, 8.99 %, 9.76 %, 16.74 %, 29.37 %, and 47.59 %, respectively”

✓ located verbatim in the source · abstract, characters 1346–1639 · show in text ↑
Intervention effectobservedoriginal resultsignificant (as stated)measured

Claim · model-written summary

Moderate N (3–4 g per plant) significantly suppressed summer flush growth relative to high N (5–7 g per plant), reducing length by 8.99 %.

Taxon
citrusCitrust0
Treatment
moderate N (3–4 g plant −1 ) application treatmente1
Outcome
lengthe1
Direction
reducinge1
Value
8.99 %e1
Comparator
high N treatment (5–7 g plant −1 )e1

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

Citrus
Citrus · Taxon · picture of the genus (one of its species)Ellen Levy Finch ( Elf ) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“citrus”

✓ located verbatim in the source · title, characters 47–53 · show in text ↑
e1

“Compared to high N treatment (5–7 g plant −1 ), moderate N (3–4 g plant −1 ) application treatment significantly suppressed SF growth, reducing shoot number, length, basal diameter, leaf number, emergence rate and biomass by 42.86 %, 8.99 %, 9.76 %, 16.74 %, 29.37 %, and 47.59 %, respectively”

✓ located verbatim in the source · abstract, characters 1346–1639 · show in text ↑
Intervention effectobservedoriginal resultsignificant (as stated)measured

Claim · model-written summary

Moderate N (3–4 g per plant) significantly suppressed summer flush growth relative to high N (5–7 g per plant), reducing basal diameter by 9.76 %.

Taxon
citrusCitrust0
Treatment
moderate N (3–4 g plant −1 ) application treatmente1
Outcome
basal diametere1
Direction
reducinge1
Value
9.76 %e1
Comparator
high N treatment (5–7 g plant −1 )e1

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

Citrus
Citrus · Taxon · picture of the genus (one of its species)Ellen Levy Finch ( Elf ) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“citrus”

✓ located verbatim in the source · title, characters 47–53 · show in text ↑
e1

“Compared to high N treatment (5–7 g plant −1 ), moderate N (3–4 g plant −1 ) application treatment significantly suppressed SF growth, reducing shoot number, length, basal diameter, leaf number, emergence rate and biomass by 42.86 %, 8.99 %, 9.76 %, 16.74 %, 29.37 %, and 47.59 %, respectively”

✓ located verbatim in the source · abstract, characters 1346–1639 · show in text ↑
Intervention effectobservedoriginal resultsignificant (as stated)measured

Claim · model-written summary

Moderate N (3–4 g per plant) significantly suppressed summer flush growth relative to high N (5–7 g per plant), reducing leaf number by 16.74 %.

Taxon
citrusCitrust0
Treatment
moderate N (3–4 g plant −1 ) application treatmente1
Outcome
leaf numbere1
Direction
reducinge1
Value
16.74 %e1
Comparator
high N treatment (5–7 g plant −1 )e1

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

Citrus
Citrus · Taxon · picture of the genus (one of its species)Ellen Levy Finch ( Elf ) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“citrus”

✓ located verbatim in the source · title, characters 47–53 · show in text ↑
e1

“Compared to high N treatment (5–7 g plant −1 ), moderate N (3–4 g plant −1 ) application treatment significantly suppressed SF growth, reducing shoot number, length, basal diameter, leaf number, emergence rate and biomass by 42.86 %, 8.99 %, 9.76 %, 16.74 %, 29.37 %, and 47.59 %, respectively”

✓ located verbatim in the source · abstract, characters 1346–1639 · show in text ↑
Intervention effectobservedoriginal resultsignificant (as stated)measured

Claim · model-written summary

Moderate N (3–4 g per plant) significantly suppressed summer flush growth relative to high N (5–7 g per plant), reducing emergence rate by 29.37 %.

Taxon
citrusCitrust0
Treatment
moderate N (3–4 g plant −1 ) application treatmente1
Outcome
emergence ratee1
Direction
reducinge1
Value
29.37 %e1
Comparator
high N treatment (5–7 g plant −1 )e1

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

Citrus
Citrus · Taxon · picture of the genus (one of its species)Ellen Levy Finch ( Elf ) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“citrus”

✓ located verbatim in the source · title, characters 47–53 · show in text ↑
e1

“Compared to high N treatment (5–7 g plant −1 ), moderate N (3–4 g plant −1 ) application treatment significantly suppressed SF growth, reducing shoot number, length, basal diameter, leaf number, emergence rate and biomass by 42.86 %, 8.99 %, 9.76 %, 16.74 %, 29.37 %, and 47.59 %, respectively”

✓ located verbatim in the source · abstract, characters 1346–1639 · show in text ↑
Intervention effectobservedoriginal resultsignificant (as stated)measured

Claim · model-written summary

Moderate N (3–4 g per plant) significantly suppressed summer flush growth relative to high N (5–7 g per plant), reducing biomass by 47.59 %.

Taxon
citrusCitrust0
Treatment
moderate N (3–4 g plant −1 ) application treatmente1
Outcome
biomasse1
Direction
reducinge1
Value
47.59 %e1
Comparator
high N treatment (5–7 g plant −1 )e1

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

Citrus
Citrus · Taxon · picture of the genus (one of its species)Ellen Levy Finch ( Elf ) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“citrus”

✓ located verbatim in the source · title, characters 47–53 · show in text ↑
e1

“Compared to high N treatment (5–7 g plant −1 ), moderate N (3–4 g plant −1 ) application treatment significantly suppressed SF growth, reducing shoot number, length, basal diameter, leaf number, emergence rate and biomass by 42.86 %, 8.99 %, 9.76 %, 16.74 %, 29.37 %, and 47.59 %, respectively”

✓ located verbatim in the source · abstract, characters 1346–1639 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

The optimal N range maximized the net increment in plant height (1.96 cm).

Taxon
citrusCitrust0
Treatment
optimal N rangee1
Outcome
plant heighte1
Direction
maximizede1
Value
1.96 cme1

Not reportedComparator

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

Citrus
Citrus · Taxon · picture of the genus (one of its species)Ellen Levy Finch ( Elf ) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“citrus”

✓ located verbatim in the source · title, characters 47–53 · show in text ↑
e1

“this optimal N range maximized the net increments in plant height (1.96 cm) and main stem diameter (1.52 cm)”

✓ located verbatim in the source · abstract, characters 1653–1761 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

The optimal N range maximized the net increment in main stem diameter (1.52 cm).

Taxon
citrusCitrust0
Treatment
optimal N rangee1
Outcome
main stem diametere1
Direction
maximizede1
Value
1.52 cme1

Not reportedComparator

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

Citrus
Citrus · Taxon · picture of the genus (one of its species)Ellen Levy Finch ( Elf ) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“citrus”

✓ located verbatim in the source · title, characters 47–53 · show in text ↑
e1

“this optimal N range maximized the net increments in plant height (1.96 cm) and main stem diameter (1.52 cm)”

✓ located verbatim in the source · abstract, characters 1653–1761 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Within the optimal N range, summer shoots maintained medium-to-high levels of nutrient content (N, phosphorus, potassium).

Taxon
citrusCitrust0
Outcome
nutrient contente1
Qualifier
medium-to-high levelse1

Not reportedDirection

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

Citrus
Citrus · Taxon · picture of the genus (one of its species)Ellen Levy Finch ( Elf ) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“citrus”

✓ located verbatim in the source · title, characters 47–53 · show in text ↑
e1

“the summer shoots maintained medium-to-high levels of nutrient content (N, phosphorus, potassium) and physiologically active substances (abscisic acid, chlorophyll, proline)”

✓ located verbatim in the source · abstract, characters 1786–1959 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Within the optimal N range, summer shoots maintained medium-to-high levels of physiologically active substances (abscisic acid, chlorophyll, proline).

Taxon
citrusCitrust0
Outcome
physiologically active substancese1
Qualifier
medium-to-high levelse1

Not reportedDirection

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

Citrus
Citrus · Taxon · picture of the genus (one of its species)Ellen Levy Finch ( Elf ) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“citrus”

✓ located verbatim in the source · title, characters 47–53 · show in text ↑
e1

“the summer shoots maintained medium-to-high levels of nutrient content (N, phosphorus, potassium) and physiologically active substances (abscisic acid, chlorophyll, proline)”

✓ located verbatim in the source · abstract, characters 1786–1959 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Within the optimal N range, root growth was moderated to a medium-low level.

Taxon
citrusCitrust0
Outcome
root growthe1
Qualifier
medium-low levele1

Not reportedDirection

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

Citrus
Citrus · Taxon · picture of the genus (one of its species)Ellen Levy Finch ( Elf ) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“citrus”

✓ located verbatim in the source · title, characters 47–53 · show in text ↑
e1

“root growth was moderated to a medium-low level”

✓ located verbatim in the source · abstract, characters 1967–2014 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

The nutrient status of the root zone soil and roots remained in the medium-to-high range.

Taxon
citrusCitrust0
Organ / stage
root zone soil and rootse1
Outcome
nutrient statuse1
Qualifier
medium-to-high rangee1

Not reportedDirection

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

Citrus
Citrus · Taxon · picture of the genus (one of its species)Ellen Levy Finch ( Elf ) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“citrus”

✓ located verbatim in the source · title, characters 47–53 · show in text ↑
e1

“the nutrient status of the root zone soil and roots remained in the medium-to-high range”

✓ located verbatim in the source · abstract, characters 2027–2115 · show in text ↑
Associationobservedoriginal resultmeasured

Claim · model-written summary

Root zone soil ammonium and nitrate nitrogen concentrations and root length density were strongly correlated with summer flush architectural traits.

Taxon
citrusCitrust0
Organ / stage
Root zone soile1
Outcome
SF architectural traitse1
Qualifier
strongly correlatede1

Not reportedDirection

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

Citrus
Citrus · Taxon · picture of the genus (one of its species)Ellen Levy Finch ( Elf ) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“citrus”

✓ located verbatim in the source · title, characters 47–53 · show in text ↑
e1

“Root zone soil ammonium nitrogen and nitrate nitrogen concentrations and root length density were strongly correlated with SF architectural traits”

✓ located verbatim in the source · abstract, characters 2121–2267 · show in text ↑
Method / resourceobservedoriginal resultpositivemeasured

Claim · model-written summary

Multiple regression models incorporating soil nitrogen and root variables explained more than 50% of the variance in summer flush phenotype.

Taxon
citrusCitrust0
Outcome
variance in SF phenotypee1
Value
>50 %e1

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

Citrus
Citrus · Taxon · picture of the genus (one of its species)Ellen Levy Finch ( Elf ) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“citrus”

✓ located verbatim in the source · title, characters 47–53 · show in text ↑
e1

“Multiple regression models incorporating these variables explained >50 % of the variance in SF phenotype”

✓ located verbatim in the source · abstract, characters 2269–2373 · show in text ↑
Mechanismauthor interpretationoriginal resultmeasured

Claim · model-written summary

The regulation of summer flush growth is attributed to an N-driven soil-root-shoot feedback in which the root zone inorganic N pool, root morphology and the balance of shoot active substances jointly define growth patterns.

Taxon
citrusCitrust0
Outcome
summer-flush growth patternse1
Qualifier
root zone inorganic N pool, root morphology, and the balance of key shoot physiologically active substancese1

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

Citrus
Citrus · Taxon · picture of the genus (one of its species)Ellen Levy Finch ( Elf ) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“citrus”

✓ located verbatim in the source · title, characters 47–53 · show in text ↑
e1

“where the root zone inorganic N pool, root morphology, and the balance of key shoot physiologically active substances collectively defined summer-flush growth patterns”

✓ located verbatim in the source · abstract, characters 2459–2626 · show in text ↑
Recommendationauthor interpretationoriginal resultbasis n/a

Claim · model-written summary

Applying 3–4 g N per plant is advisable as the ideal summer rate for potted citrus seedlings in controlled environments.

Taxon
citrus seedlingsCitruse1
Treatment
3–4 g N plant −1e1
Outcome
ideal summer ratee1

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

Citrus
Citrus · Taxon · picture of the genus (one of its species)Ellen Levy Finch ( Elf ) · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

t0

“citrus”

✓ located verbatim in the source · title, characters 47–53 · show in text ↑
e1

“it is advisable to apply 3–4 g N plant −1 as the ideal summer rate for potted citrus seedlings in controlled environments”

✓ located verbatim in the source · abstract, characters 2639–2760 · show in text ↑

nitrogen · growth · CitrusPositive and negative results

17 papers: 15 positive, 2 negative or no effect

14 papers in the same direction as this paper · 2 in the opposite direction

  1. positive
  2. negative
  3. negative

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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.