Paper · 2025 · Journal of Integrative Agriculture

Phosphorus application enhances floret development and grain formation, stabilizing wheat yield under drought stress through improved photosynthetic performance and nutrient accumulation

Yanyan Zhang; Yahui Li; Jiahao Liu; et al.

20
claims
40
verbatim quotes
2
citations
3
research areas
✓
Open access · cc-by-nc-nd

Abstract

1. Phosphorus application alleviates drought stress by increasing grain number and stabilizing yield. 2. Phosphorus application regulates floret development in inferior spikelets, enhancing the formation of fertile florets. 3. Phosphorus increases grain number by modulating photosynthetic performance and nutrient accumulation. Phosphorus application mitigates drought stress and promotes wheat spike and floret differentiation and development; however, the mechanisms by which phosphorus regulates floret-to-grain transition under drought stress to maintain stable yield remain unclear. This study aimed to elucidate the nutritional and physiological mechanisms through which phosphorus application enhances floret development and grain setting, thereby reducing the adverse effects of drought stress in field-grown wheat. Three water regimes (W0, severe drought; W1, moderate drought; W2, well-watered) and two phosphorus levels (P0, no phosphorus application; P1, standard phosphorus application) were applied using two wheat varieties—large-spike Zhoumai 16 (V1) and multi-spike Yumai 49-198 (V2)—over two growing seasons (2020–2022) under field conditions. The experiment systematically assessed the impact of phosphorus on fertile floret dynamics, leaf photosynthetic characteristics, and nutrient uptake and translocation across different water treatments. Results indicated that drought stress inhibited leaf photosynthetic traits in both varieties, impairing aboveground dry matter production and nutrient absorption, ultimately restricting floret development and grain set and significantly reducing yield. Phosphorus application increased chlorophyll (Chl) content, net photosynthetic rate (P n ), maximum photochemical efficiency ( F v / F m ), and actual photochemical efficiency (ΦPSII), thereby promoting dry matter, nitrogen, and phosphorus accumulation in all organs and increasing the ratio of phosphorus allocation to spikes relative to non-spike tissues. Compared with P0, phosphorus application increased spike number, grains per spike, and thousand-grain weight (TGW) in both varieties, with grain number per spike rising by 9.63–18.68% (V1) and 10.88–18.81% (V2), and yield gains ranging from 14.48–33.84% (V1) and 13.23–29.29% (V2). Differences among water treatments and varieties revealed that phosphorus exerted a stronger regulatory effect under moderate drought (W1) than under severe drought (W0) or well-watered (W2) conditions, and its influence was more pronounced in the large-spike variety V1 than in the multi-spike variety V2. While phosphorus application enhanced photosynthetic performance and nutrient accumulation similarly to irrigation, it exhibited contrasting patterns in phosphorus partitioning and TGW regulation. This study provides theoretical insight and technical guidance for improving floret development, increasing grain number and yield, and counteracting drought stress in wheat production through phosphorus-mediated regulation of carbon, nitrogen, and phosphorus nutrition.

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 20

Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Drought stress reduced leaf photosynthetic traits.

Treatment
drought stresse0
Outcome
leaf photosynthetic traitse0
Direction
inhibitede0

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

“drought stress inhibited leaf photosynthetic traits in both varieties, impairing aboveground dry matter production and nutrient absorption, ultimately restricting floret development and grain set and significantly reducing yield”

✓ located verbatim in the source · abstract, characters 1388–1616 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Drought stress reduced aboveground dry matter production.

Treatment
drought stresse0
Outcome
aboveground dry matter productione0
Direction
impairinge0

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

“drought stress inhibited leaf photosynthetic traits in both varieties, impairing aboveground dry matter production and nutrient absorption, ultimately restricting floret development and grain set and significantly reducing yield”

✓ located verbatim in the source · abstract, characters 1388–1616 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Drought stress reduced nutrient absorption.

Treatment
drought stresse0
Outcome
nutrient absorptione0
Direction
impairinge0

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

“drought stress inhibited leaf photosynthetic traits in both varieties, impairing aboveground dry matter production and nutrient absorption, ultimately restricting floret development and grain set and significantly reducing yield”

✓ located verbatim in the source · abstract, characters 1388–1616 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Drought stress reduced floret development and grain set.

Treatment
drought stresse0
Outcome
floret development and grain sete0
Direction
restrictinge0

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

“drought stress inhibited leaf photosynthetic traits in both varieties, impairing aboveground dry matter production and nutrient absorption, ultimately restricting floret development and grain set and significantly reducing yield”

✓ located verbatim in the source · abstract, characters 1388–1616 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultnegativesignificant (as stated)measured

Claim · model-written summary

Drought stress reduced yield.

Treatment
drought stresse0
Outcome
yielde0
Direction
reducinge0

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

“drought stress inhibited leaf photosynthetic traits in both varieties, impairing aboveground dry matter production and nutrient absorption, ultimately restricting floret development and grain set and significantly reducing yield”

✓ located verbatim in the source · abstract, characters 1388–1616 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Phosphorus application increased chlorophyll (Chl) content.

Treatment
Phosphorus applicatione0
Outcome
chlorophyll (Chl) contente0
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

“Phosphorus application increased chlorophyll (Chl) content, net photosynthetic rate (P n ), maximum photochemical efficiency ( F v / F m ), and actual photochemical efficiency (ΦPSII)”

✓ located verbatim in the source · abstract, characters 1618–1801 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Phosphorus application increased net photosynthetic rate (P n ).

Treatment
Phosphorus applicatione0
Outcome
net photosynthetic rate (P n )e0
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

“Phosphorus application increased chlorophyll (Chl) content, net photosynthetic rate (P n ), maximum photochemical efficiency ( F v / F m ), and actual photochemical efficiency (ΦPSII)”

✓ located verbatim in the source · abstract, characters 1618–1801 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Phosphorus application increased maximum photochemical efficiency ( F v / F m ).

Treatment
Phosphorus applicatione0
Outcome
maximum photochemical efficiency ( F v / F m )e0
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

“Phosphorus application increased chlorophyll (Chl) content, net photosynthetic rate (P n ), maximum photochemical efficiency ( F v / F m ), and actual photochemical efficiency (ΦPSII)”

✓ located verbatim in the source · abstract, characters 1618–1801 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Phosphorus application increased actual photochemical efficiency (ΦPSII).

Treatment
Phosphorus applicatione0
Outcome
actual photochemical efficiency (ΦPSII)e0
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

“Phosphorus application increased chlorophyll (Chl) content, net photosynthetic rate (P n ), maximum photochemical efficiency ( F v / F m ), and actual photochemical efficiency (ΦPSII)”

✓ located verbatim in the source · abstract, characters 1618–1801 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Phosphorus promoted dry matter, nitrogen and phosphorus accumulation in all organs.

Outcome
dry matter, nitrogen, and phosphorus accumulatione0
Direction
promotinge0

Not reportedTreatmentComparator

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

“promoting dry matter, nitrogen, and phosphorus accumulation in all organs and increasing the ratio of phosphorus allocation to spikes relative to non-spike tissues”

✓ located verbatim in the source · abstract, characters 1811–1974 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Phosphorus increased the ratio of phosphorus allocated to spikes.

Outcome
ratio of phosphorus allocation to spikese0
Direction
increasinge0
Comparator
non-spike tissuese0

Not reportedTreatment

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

“promoting dry matter, nitrogen, and phosphorus accumulation in all organs and increasing the ratio of phosphorus allocation to spikes relative to non-spike tissues”

✓ located verbatim in the source · abstract, characters 1811–1974 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Phosphorus increased spike number relative to P0.

Treatment
phosphorus applicatione0
Outcome
spike numbere0
Direction
increasede0
Comparator
P0e0

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 P0, phosphorus application increased spike number, grains per spike, and thousand-grain weight (TGW) in both varieties, with grain number per spike rising by 9.63–18.68% (V1) and 10.88–18.81% (V2), and yield gains ranging from 14.48–33.84% (V1) and 13.23–29.29% (V2)”

✓ located verbatim in the source · abstract, characters 1976–2256 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Phosphorus increased thousand-grain weight relative to P0.

Treatment
phosphorus applicatione0
Outcome
thousand-grain weight (TGW)e0
Direction
increasede0
Comparator
P0e0

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 P0, phosphorus application increased spike number, grains per spike, and thousand-grain weight (TGW) in both varieties, with grain number per spike rising by 9.63–18.68% (V1) and 10.88–18.81% (V2), and yield gains ranging from 14.48–33.84% (V1) and 13.23–29.29% (V2)”

✓ located verbatim in the source · abstract, characters 1976–2256 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Phosphorus increased grains per spike by 9.63–18.68% in V1.

Material
V1e0
Treatment
phosphorus applicatione0
Outcome
grain number per spikee0
Direction
risinge0
Value
9.63–18.68%e0
Comparator
P0e0

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 P0, phosphorus application increased spike number, grains per spike, and thousand-grain weight (TGW) in both varieties, with grain number per spike rising by 9.63–18.68% (V1) and 10.88–18.81% (V2), and yield gains ranging from 14.48–33.84% (V1) and 13.23–29.29% (V2)”

✓ located verbatim in the source · abstract, characters 1976–2256 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Phosphorus increased grains per spike by 10.88–18.81% in V2.

Material
V2e0
Treatment
phosphorus applicatione0
Outcome
grain number per spikee0
Direction
risinge0
Value
10.88–18.81%e0
Comparator
P0e0

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 P0, phosphorus application increased spike number, grains per spike, and thousand-grain weight (TGW) in both varieties, with grain number per spike rising by 9.63–18.68% (V1) and 10.88–18.81% (V2), and yield gains ranging from 14.48–33.84% (V1) and 13.23–29.29% (V2)”

✓ located verbatim in the source · abstract, characters 1976–2256 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Phosphorus increased yield by 14.48–33.84% in V1.

Material
V1e0
Treatment
phosphorus applicatione0
Outcome
yield gainse0
Direction
gainse0
Value
14.48–33.84%e0
Comparator
P0e0

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 P0, phosphorus application increased spike number, grains per spike, and thousand-grain weight (TGW) in both varieties, with grain number per spike rising by 9.63–18.68% (V1) and 10.88–18.81% (V2), and yield gains ranging from 14.48–33.84% (V1) and 13.23–29.29% (V2)”

✓ located verbatim in the source · abstract, characters 1976–2256 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Phosphorus increased yield by 13.23–29.29% in V2.

Material
V2e0
Treatment
phosphorus applicatione0
Outcome
yield gainse0
Direction
gainse0
Value
13.23–29.29%e0
Comparator
P0e0

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 P0, phosphorus application increased spike number, grains per spike, and thousand-grain weight (TGW) in both varieties, with grain number per spike rising by 9.63–18.68% (V1) and 10.88–18.81% (V2), and yield gains ranging from 14.48–33.84% (V1) and 13.23–29.29% (V2)”

✓ located verbatim in the source · abstract, characters 1976–2256 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Phosphorus had a stronger regulatory effect under moderate drought than severe drought or well-watered conditions.

Treatment
phosphoruse0
Outcome
regulatory effecte0
Direction
strongere0
Qualifier
moderate drought (W1)e0
Comparator
severe drought (W0) or well-watered (W2) conditionse0

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

“phosphorus exerted a stronger regulatory effect under moderate drought (W1) than under severe drought (W0) or well-watered (W2) conditions”

✓ located verbatim in the source · abstract, characters 2321–2459 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Phosphorus influence was greater in V1 than V2.

Material
large-spike variety V1e0
Outcome
influencee0
Direction
more pronouncede0
Comparator
multi-spike variety V2e0

Not reportedTreatment

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

“its influence was more pronounced in the large-spike variety V1 than in the multi-spike variety V2”

✓ located verbatim in the source · abstract, characters 2465–2563 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑
Intervention effectobservedoriginal resultmixedmeasured

Claim · model-written summary

Phosphorus application and irrigation showed contrasting phosphorus partitioning and thousand-grain weight regulation.

Outcome
phosphorus partitioning and TGW regulatione0
Direction
contrastinge0

Not reportedTreatmentComparator

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

“it exhibited contrasting patterns in phosphorus partitioning and TGW regulation”

✓ located verbatim in the source · abstract, characters 2681–2760 · show in text ↑
t0

“wheat yield”

✓ located verbatim in the source · title, characters 84–95 · show in text ↑

phosphorus · physiology: photosynthesis · Triticum aestivum (wheat)Results differ

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

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phosphorus · physiology: pigments · Triticum aestivum (wheat)Positive and negative results

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drought · nutrient: content · Triticum aestivum (wheat)Positive and negative results

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drought · physiology: photosynthesis · Triticum aestivum (wheat)Positive and negative results

246 papers: 41 positive, 205 negative or no effect

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phosphorus · yield · Triticum aestivum (wheat)Positive and negative results

572 papers: 524 positive, 48 negative or no effect

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drought · yield · Triticum aestivum (wheat)Positive and negative results

632 papers: 95 positive, 537 negative or no effect

536 papers in the same direction as this paper · 95 in the opposite direction

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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.