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.
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Claims 13
Claim · model-written summary
Drought significantly reduced net photosynthetic rate, stomatal conductance and PSII efficiency in non-acclimated Lolium perenne.
- Taxon
- Lolium perenne
Lolium 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.

Verbatim
e0“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”
e1“as evidenced by the significant reductions in net photosynthetic rate, stomatal conductance and PSII efficiency ( Fv / Fm )”
e2“drought severely impaired photosynthetic capacity and induced oxidative damage in non-acclimated plants”
Claim · model-written summary
Drought elevated malondialdehyde (MDA) levels in non-acclimated Lolium perenne.
- Taxon
- Lolium perenne
Lolium perennee0 - Treatment
- droughte2
- Outcome
- malondialdehyde (MDA) levelse3
- Direction
- elevatede3
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”
e2“drought severely impaired photosynthetic capacity and induced oxidative damage in non-acclimated plants”
e3“alongside elevated malondialdehyde (MDA) levels”
Claim · model-written summary
Drought increased the activities of key antioxidant enzymes in non-acclimated Lolium perenne.
- Taxon
- Lolium perenne
Lolium 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.

Verbatim
e0“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”
e2“drought severely impaired photosynthetic capacity and induced oxidative damage in non-acclimated plants”
e4“increased activities of key antioxidant enzymes (e.g., ascorbate peroxidase, peroxidase and catalase)”
Claim · model-written summary
Low nitrogen pre-acclimation preserved photosynthetic performance under subsequent drought in Lolium perenne.
- Taxon
- Lolium perenne
Lolium 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.

Verbatim
e0“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”
e2“drought severely impaired photosynthetic capacity and induced oxidative damage in non-acclimated plants”
e5“low nitrogen pre-acclimation effectively preserved photosynthetic performance under subsequent drought”
Claim · model-written summary
Low nitrogen pre-acclimation mitigated declines in gas exchange parameters under drought.
- Taxon
- Lolium perenne
Lolium 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.

Verbatim
e0“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”
e2“drought severely impaired photosynthetic capacity and induced oxidative damage in non-acclimated plants”
e5“low nitrogen pre-acclimation effectively preserved photosynthetic performance under subsequent drought”
e6“mitigating declines in gas exchange parameters and maintaining PSII integrity”
Claim · model-written summary
Pre-acclimated Lolium perenne plants showed reduced oxidative stress under drought.
- Taxon
- Lolium perenne
Lolium 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.

Verbatim
e0“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”
e5“low nitrogen pre-acclimation effectively preserved photosynthetic performance under subsequent drought”
e7“These pre-acclimated plants also exhibited reduced oxidative stress under drought and superior recovery capacity after rewatering”
Claim · model-written summary
Pre-acclimated Lolium perenne plants showed superior recovery capacity after rewatering.
- Taxon
- Lolium perenne
Lolium 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.

Verbatim
e0“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”
e5“low nitrogen pre-acclimation effectively preserved photosynthetic performance under subsequent drought”
e7“These pre-acclimated plants also exhibited reduced oxidative stress under drought and superior recovery capacity after rewatering”
Claim · model-written summary
Drought tolerance in Lolium perenne was associated with fructan accumulation.
- Taxon
- Lolium perenne
Lolium perennee0 - Compound
- fructane7
- Outcome
- drought tolerancee7
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”
e7“This enhanced drought tolerance was associated with fructan accumulation and tempered transcriptional responses”
Claim · model-written summary
Low nitrogen pre-acclimation alleviated the drought-induced downregulation of genes encoding the photosynthetic apparatus in Lolium perenne.
- Taxon
- Lolium perenne
Lolium 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.

Verbatim
e0“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”
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”
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 perenne
Lolium 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.

Verbatim
e0“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”
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”
Claim · model-written summary
Low nitrogen pre-acclimation mitigated the drought-induced transcriptional upheaval in Lolium perenne.
- Taxon
- Lolium perenne
Lolium 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.

Verbatim
e0“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”
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”
Claim · model-written summary
Low nitrogen pre-acclimation stabilized chlorophyll metabolism under drought in Lolium perenne.
- Taxon
- Lolium perenne
Lolium 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.

Verbatim
e0“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”
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”
e7“stabilized chlorophyll metabolism and optimized carbon-nitrogen balance”
Claim · model-written summary
Low nitrogen pre-acclimation optimized the carbon-nitrogen balance under drought in Lolium perenne.
- Taxon
- Lolium perenne
Lolium 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.

Verbatim
e0“Drought stress is a major constraint on the productivity of temperate forage grasses such as perennial ryegrass ( Lolium perenne )”
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”
e7“stabilized chlorophyll metabolism and optimized carbon-nitrogen balance”
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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.