Paper · 2026 · Plant Physiology and Biochemistry

Pre-acclimation to low nitrogen enhances drought tolerance in Lolium perenne through integrated metabolic and transcriptional alterations

Hui Zuo; Xinyue Xiong; Yuqian Chen; et al.

13
claims
36
verbatim quotes
0
citations
2
research areas
✓
Open access · cc-by

Abstract

Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne ). While nitrogen management is widely employed in agronomic practice, its specific role in mediating drought adaptation strategies remains unclear. This study aimed to systematically characterize the response of perennial ryegrass to drought stress following low nitrogen pre-acclimation through integrated physiological and transcriptomic analyses during both stress and recovery phases. Our results demonstrated that drought severely impaired photosynthetic capacity and induced oxidative damage in non-acclimated plants, as evidenced by the significant reductions in net photosynthetic rate, stomatal conductance and PSII efficiency ( Fv / Fm ), alongside elevated malondialdehyde (MDA) levels and increased activities of key antioxidant enzymes (e.g., ascorbate peroxidase, peroxidase and catalase). In contrast, low nitrogen pre-acclimation effectively preserved photosynthetic performance under subsequent drought, mitigating declines in gas exchange parameters and maintaining PSII integrity. These pre-acclimated plants also exhibited reduced oxidative stress under drought and superior recovery capacity after rewatering. This enhanced drought tolerance was associated with fructan accumulation and tempered transcriptional responses. Low nitrogen pre-acclimation mitigated the drought-induced transcriptional upheaval, attenuated the activation of hormonal signaling pathways and MAPK cascades, significantly alleviated the downregulation of genes encoding photosynthetic apparatus, stabilized chlorophyll metabolism and optimized carbon-nitrogen balance. These findings reveal a nitrogen-mediated priming mechanism that enhances drought tolerance through integrated metabolic and transcriptional adjustments, providing new insights into the interaction between nutrient signaling and stress resistance, as well as potential strategies for enhancing plant tolerance under climate change. • Low nitrogen pre-acclimation primes drought tolerance in Lolium perenne. • Pre-acclimation mitigates photosynthetic decline and oxidative damage under drought. • Maximized fructan accumulation and optimized C/N balance underpin tolerance. • A unique transcriptional reprogramming drives the primed, coordinated response. • Reveals a nitrogen mediated cross-stress memory mechanism for enhanced tolerance.

Abstract from the paper’s OpenAlex record; open copy licensed cc-by. 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 13

Intervention effectobservedoriginal resultnegativesignificant (as stated)measured

Claim · model-written summary

Drought significantly reduced net photosynthetic rate, stomatal conductance and PSII efficiency in non-acclimated Lolium perenne.

Taxon
Lolium perenneLolium perennee0
Treatment
droughte2
Outcome
net photosynthetic rate, stomatal conductance and PSII efficiencye1
Direction
reductionse1
Qualifier
non-acclimated plantse2

Not reportedComparator

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

Lolium perenne
Lolium perenne · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”

✓ located verbatim in the source · abstract, characters 0–130 · show in text ↑
e1

“as evidenced by the significant reductions in net photosynthetic rate, stomatal conductance and PSII efficiency ( Fv / Fm )”

✓ located verbatim in the source · abstract, characters 654–777 · show in text ↑
e2

“drought severely impaired photosynthetic capacity and induced oxidative damage in non-acclimated plants”

✓ located verbatim in the source · abstract, characters 549–652 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Drought elevated malondialdehyde (MDA) levels in non-acclimated Lolium perenne.

Taxon
Lolium perenneLolium perennee0
Treatment
droughte2
Outcome
malondialdehyde (MDA) levelse3
Direction
elevatede3

Not reportedComparator

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

Lolium perenne
Lolium perenne · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”

✓ located verbatim in the source · abstract, characters 0–130 · show in text ↑
e2

“drought severely impaired photosynthetic capacity and induced oxidative damage in non-acclimated plants”

✓ located verbatim in the source · abstract, characters 549–652 · show in text ↑
e3

“alongside elevated malondialdehyde (MDA) levels”

✓ located verbatim in the source · abstract, characters 779–826 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Drought increased the activities of key antioxidant enzymes in non-acclimated Lolium perenne.

Taxon
Lolium perenneLolium perennee0
Treatment
droughte2
Outcome
activities of key antioxidant enzymese4
Direction
increasede4
Qualifier
ascorbate peroxidase, peroxidase and catalasee4

Not reportedComparator

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

Lolium perenne
Lolium perenne · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”

✓ located verbatim in the source · abstract, characters 0–130 · show in text ↑
e2

“drought severely impaired photosynthetic capacity and induced oxidative damage in non-acclimated plants”

✓ located verbatim in the source · abstract, characters 549–652 · show in text ↑
e4

“increased activities of key antioxidant enzymes (e.g., ascorbate peroxidase, peroxidase and catalase)”

✓ located verbatim in the source · abstract, characters 831–932 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Low nitrogen pre-acclimation preserved photosynthetic performance under subsequent drought in Lolium perenne.

Taxon
Lolium perenneLolium perennee0
Treatment
low nitrogen pre-acclimatione5
Outcome
photosynthetic performancee5
Direction
preservede5
Comparator
non-acclimated plantse2

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

Lolium perenne
Lolium perenne · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”

✓ located verbatim in the source · abstract, characters 0–130 · show in text ↑
e2

“drought severely impaired photosynthetic capacity and induced oxidative damage in non-acclimated plants”

✓ located verbatim in the source · abstract, characters 549–652 · show in text ↑
e5

“low nitrogen pre-acclimation effectively preserved photosynthetic performance under subsequent drought”

✓ located verbatim in the source · abstract, characters 947–1049 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Low nitrogen pre-acclimation mitigated declines in gas exchange parameters under drought.

Taxon
Lolium perenneLolium perennee0
Treatment
low nitrogen pre-acclimatione5
Outcome
declines in gas exchange parameterse6
Direction
mitigatinge6
Comparator
non-acclimated plantse2

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

Lolium perenne
Lolium perenne · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”

✓ located verbatim in the source · abstract, characters 0–130 · show in text ↑
e2

“drought severely impaired photosynthetic capacity and induced oxidative damage in non-acclimated plants”

✓ located verbatim in the source · abstract, characters 549–652 · show in text ↑
e5

“low nitrogen pre-acclimation effectively preserved photosynthetic performance under subsequent drought”

✓ located verbatim in the source · abstract, characters 947–1049 · show in text ↑
e6

“mitigating declines in gas exchange parameters and maintaining PSII integrity”

✓ located verbatim in the source · abstract, characters 1051–1128 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Pre-acclimated Lolium perenne plants showed reduced oxidative stress under drought.

Taxon
Lolium perenneLolium perennee0
Treatment
low nitrogen pre-acclimatione5
Outcome
oxidative stresse7
Direction
reducede7

Not reportedComparator

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

Lolium perenne
Lolium perenne · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”

✓ located verbatim in the source · abstract, characters 0–130 · show in text ↑
e5

“low nitrogen pre-acclimation effectively preserved photosynthetic performance under subsequent drought”

✓ located verbatim in the source · abstract, characters 947–1049 · show in text ↑
e7

“These pre-acclimated plants also exhibited reduced oxidative stress under drought and superior recovery capacity after rewatering”

✓ located verbatim in the source · abstract, characters 1130–1259 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Pre-acclimated Lolium perenne plants showed superior recovery capacity after rewatering.

Taxon
Lolium perenneLolium perennee0
Treatment
low nitrogen pre-acclimatione5
Outcome
recovery capacity after rewateringe7
Direction
superiore7

Not reportedComparator

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

Lolium perenne
Lolium perenne · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”

✓ located verbatim in the source · abstract, characters 0–130 · show in text ↑
e5

“low nitrogen pre-acclimation effectively preserved photosynthetic performance under subsequent drought”

✓ located verbatim in the source · abstract, characters 947–1049 · show in text ↑
e7

“These pre-acclimated plants also exhibited reduced oxidative stress under drought and superior recovery capacity after rewatering”

✓ located verbatim in the source · abstract, characters 1130–1259 · show in text ↑
Associationobservedoriginal resultpositivemeasured

Claim · model-written summary

Drought tolerance in Lolium perenne was associated with fructan accumulation.

Taxon
Lolium perenneLolium perennee0
Compound
fructane7
Outcome
drought tolerancee7

Not reportedDirection

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

Lolium perenne
Lolium perenne · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”

✓ located verbatim in the source · abstract, characters 0–130 · show in text ↑
e7

“This enhanced drought tolerance was associated with fructan accumulation and tempered transcriptional responses”

✓ located verbatim in the source · abstract, characters 1261–1372 · show in text ↑
Intervention effectobservedoriginal resultpositivesignificant (as stated)measured

Claim · model-written summary

Low nitrogen pre-acclimation alleviated the drought-induced downregulation of genes encoding the photosynthetic apparatus in Lolium perenne.

Taxon
Lolium perenneLolium perennee0
Treatment
Low nitrogen pre-acclimatione6
Gene
genes encoding photosynthetic apparatuse6
Outcome
downregulation of genese6
Direction
alleviatede6

Not reportedComparator

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

Lolium perenne
Lolium perenne · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”

✓ located verbatim in the source · abstract, characters 0–130 · show in text ↑
e6

“Low nitrogen pre-acclimation mitigated the drought-induced transcriptional upheaval, attenuated the activation of hormonal signaling pathways and MAPK cascades, significantly alleviated the downregulation of genes encoding photosynthetic apparatus”

✓ located verbatim in the source · abstract, characters 1374–1621 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Low nitrogen pre-acclimation attenuated the activation of hormonal signaling pathways and MAPK cascades under drought in Lolium perenne.

Taxon
Lolium perenneLolium perennee0
Treatment
Low nitrogen pre-acclimatione6
Outcome
activation of hormonal signaling pathways and MAPK cascadese6
Direction
attenuatede6

Not reportedComparator

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

Lolium perenne
Lolium perenne · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”

✓ located verbatim in the source · abstract, characters 0–130 · show in text ↑
e6

“Low nitrogen pre-acclimation mitigated the drought-induced transcriptional upheaval, attenuated the activation of hormonal signaling pathways and MAPK cascades, significantly alleviated the downregulation of genes encoding photosynthetic apparatus”

✓ located verbatim in the source · abstract, characters 1374–1621 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Low nitrogen pre-acclimation mitigated the drought-induced transcriptional upheaval in Lolium perenne.

Taxon
Lolium perenneLolium perennee0
Treatment
Low nitrogen pre-acclimatione6
Outcome
drought-induced transcriptional upheavale6
Direction
mitigatede6

Not reportedComparator

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

Lolium perenne
Lolium perenne · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”

✓ located verbatim in the source · abstract, characters 0–130 · show in text ↑
e6

“Low nitrogen pre-acclimation mitigated the drought-induced transcriptional upheaval, attenuated the activation of hormonal signaling pathways and MAPK cascades, significantly alleviated the downregulation of genes encoding photosynthetic apparatus”

✓ located verbatim in the source · abstract, characters 1374–1621 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Low nitrogen pre-acclimation stabilized chlorophyll metabolism under drought in Lolium perenne.

Taxon
Lolium perenneLolium perennee0
Treatment
Low nitrogen pre-acclimatione6
Outcome
chlorophyll metabolisme7
Direction
stabilizede7

Not reportedComparator

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

Lolium perenne
Lolium perenne · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”

✓ located verbatim in the source · abstract, characters 0–130 · show in text ↑
e6

“Low nitrogen pre-acclimation mitigated the drought-induced transcriptional upheaval, attenuated the activation of hormonal signaling pathways and MAPK cascades, significantly alleviated the downregulation of genes encoding photosynthetic apparatus”

✓ located verbatim in the source · abstract, characters 1374–1621 · show in text ↑
e7

“stabilized chlorophyll metabolism and optimized carbon-nitrogen balance”

✓ located verbatim in the source · abstract, characters 1623–1694 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Low nitrogen pre-acclimation optimized the carbon-nitrogen balance under drought in Lolium perenne.

Taxon
Lolium perenneLolium perennee0
Treatment
Low nitrogen pre-acclimatione6
Outcome
carbon-nitrogen balancee7
Direction
optimizede7

Not reportedComparator

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

Lolium perenne
Lolium perenne · TaxonRasbak · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”

✓ located verbatim in the source · abstract, characters 0–130 · show in text ↑
e6

“Low nitrogen pre-acclimation mitigated the drought-induced transcriptional upheaval, attenuated the activation of hormonal signaling pathways and MAPK cascades, significantly alleviated the downregulation of genes encoding photosynthetic apparatus”

✓ located verbatim in the source · abstract, characters 1374–1621 · show in text ↑
e7

“stabilized chlorophyll metabolism and optimized carbon-nitrogen balance”

✓ located verbatim in the source · abstract, characters 1623–1694 · show in text ↑

nitrogen · physiology: photosynthesis · Lolium perenne (perennial ryegrass)Positive and negative results

4 papers: 3 positive, 1 negative or no effect

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

  1. positive
  2. no effect
  3. positive
See all 4 papers →

nitrogen · physiology: pigments · Lolium perenne (perennial ryegrass)Positive and negative results

4 papers: 3 positive, 1 negative or no effect

2 papers in the same direction as this paper · 1 in the opposite direction

  1. negative
  2. positive
  3. positive
See all 4 papers →

drought · physiology: oxidative · Lolium perenne (perennial ryegrass)Positive and negative results

5 papers: 1 positive, 4 negative or no effect

3 papers in the same direction as this paper · 1 in the opposite direction

  1. positive
  2. negative
  3. negative

1 paper more in this group

See all 5 papers →

nitrogen · molecular: expression · Lolium perenne (perennial ryegrass)Results agree

2 papers: 2 positive, 0 negative or no effect

1 paper in the same direction as this paper

  1. positive
See all 2 papers →

drought · physiology: photosynthesis · Lolium perenne (perennial ryegrass)Results agree

3 papers: 0 positive, 3 negative or no effect

2 papers in the same direction as this paper

  1. negative
  2. negative
See all 3 papers →

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.