Abstract
Salinity stress severely limits cucumber (Cucumis sativus L.) productivity by disrupting growth, photosynthesis, and ion homeostasis. This study investigated the potential of foliar-applied allantoin (1 mM) and citrulline (1 mM) to enhance salinity tolerance in hairy cucumber, a salt-sensitive Iranian landrace. A completely randomized design factorial experiment with three replications tested salinity levels (0, 50, 100 mM NaCl) and foliar treatments (control, allantoin, citrulline, and their combination). The results showed that salinity reduced plant height, dry weight, chlorophyll content (SPAD, Fv/Fm), and relative water content (RWC) while increasing electrolyte leakage (EL), oxidative markers (MDA, H2O₂), and Na⁺ accumulation. Allantoin mitigated these effects by enhancing nitrogen (N) metabolism, improving K⁺/Na+ homeostasis, and upregulating osmolyte (proline, sugars) and antioxidant (phenolics, ascorbic acid) accumulation. Citrulline boosted nitric oxide (NO) production, which reduced Na⁺ toxicity, improved stomatal conductance, and activated enzymatic antioxidants (SOD, APX). Their combined application synergistically improved growth (44.24% in yield), photosynthesis (5.12% in Fv/Fm), and RWC (6.45%) while reducing H2O2 (61.53%) and oxidative stress (43.74% MDA). Notably, salt-stressed plants treated with both compounds exhibited elevated fruit levels of health-promoting metabolites (cucurbitacin, citrulline). Moreover, allantoin enhanced N assimilation and polyamine-mediated membrane stability, while citrulline-derived NO optimized ion transport and ROS scavenging. These findings highlight the dual role of allantoin (N metabolism) and citrulline (NO signaling) in conferring salt tolerance and propose their combined foliar application as a sustainable strategy for improving cucumber resilience in saline agriculture. - Allantoin+citrulline application improved cucumber biomass under 100 mM NaCl stress conditions - Allantoin improved K+/Ca+2 homeostasis, counteracting Na⁺ toxicity effects - Citrulline application enhanced stomatal conductance and CO₂ assimilation under stress - ROS-scavenging enzymes activity significantly enhanced by combined application - Combined treatment increased proline and soluble sugars for better osmotic adjustment
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Claims 16
Claim · model-written summary
Salinity reduced plant height and dry weight of cucumber.
- Taxon
- cucumber
Cucumis sativuse0 - Treatment
- salinitye1
- Outcome
- plant height, dry weighte1
- Direction
- reducede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Salinity stress severely limits cucumber (Cucumis sativus L.) productivity”
e1“The results showed that salinity reduced plant height, dry weight, chlorophyll content (SPAD, Fv/Fm), and relative water content (RWC)”
Claim · model-written summary
Salinity reduced chlorophyll content and relative water content of cucumber.
- Taxon
- cucumberCucumis sativuse0
- Treatment
- salinitye1
- Outcome
- chlorophyll content (SPAD, Fv/Fm), and relative water content (RWC)e1
- Direction
- reducede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Salinity stress severely limits cucumber (Cucumis sativus L.) productivity”
e1“The results showed that salinity reduced plant height, dry weight, chlorophyll content (SPAD, Fv/Fm), and relative water content (RWC)”
Claim · model-written summary
Salinity increased electrolyte leakage, oxidative markers and Na+ accumulation in cucumber.
- Taxon
- cucumberCucumis sativuse0
- Treatment
- salinitye1
- Outcome
- electrolyte leakage (EL), oxidative markers (MDA, H2O₂), and Na⁺ accumulatione2
- Direction
- increasinge2
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Salinity stress severely limits cucumber (Cucumis sativus L.) productivity”
e1“The results showed that salinity reduced plant height, dry weight, chlorophyll content (SPAD, Fv/Fm), and relative water content (RWC)”
e2“while increasing electrolyte leakage (EL), oxidative markers (MDA, H2O₂), and Na⁺ accumulation”
Claim · model-written summary
Allantoin mitigated salinity effects on cucumber by enhancing nitrogen metabolism.
- Taxon
- cucumberCucumis sativuse0
- Treatment
- foliar-applied allantoin (1 mM)e4
- Outcome
- mitigated these effectse3
- Qualifier
- by enhancing nitrogen (N) metabolisme3
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Salinity stress severely limits cucumber (Cucumis sativus L.) productivity”
e3“Allantoin mitigated these effects by enhancing nitrogen (N) metabolism, improving K⁺/Na+ homeostasis, and upregulating osmolyte (proline, sugars) and antioxidant (phenolics, ascorbic acid) accumulation”
e4“foliar-applied allantoin (1 mM) and citrulline (1 mM) to enhance salinity tolerance in hairy cucumber”
Claim · model-written summary
Citrulline reduced Na+ toxicity in cucumber through nitric oxide production.
- Taxon
- cucumberCucumis sativuse0
- Treatment
- Citrullinee5
- Outcome
- reduced Na⁺ toxicitye5
- Qualifier
- nitric oxide (NO) productione5
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Salinity stress severely limits cucumber (Cucumis sativus L.) productivity”
e5“Citrulline boosted nitric oxide (NO) production, which reduced Na⁺ toxicity, improved stomatal conductance, and activated enzymatic antioxidants (SOD, APX)”
Claim · model-written summary
Citrulline improved stomatal conductance in cucumber under salinity.
- Taxon
- cucumberCucumis sativuse0
- Treatment
- Citrullinee5
- Outcome
- improved stomatal conductancee5
- Qualifier
- nitric oxide (NO) productione5
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Salinity stress severely limits cucumber (Cucumis sativus L.) productivity”
e5“Citrulline boosted nitric oxide (NO) production, which reduced Na⁺ toxicity, improved stomatal conductance, and activated enzymatic antioxidants (SOD, APX)”
Claim · model-written summary
Citrulline activated enzymatic antioxidants (SOD, APX) in cucumber.
- Taxon
- cucumberCucumis sativuse0
- Treatment
- Citrullinee5
- Outcome
- activated enzymatic antioxidants (SOD, APX)e5
- Qualifier
- nitric oxide (NO) productione5
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Salinity stress severely limits cucumber (Cucumis sativus L.) productivity”
e5“Citrulline boosted nitric oxide (NO) production, which reduced Na⁺ toxicity, improved stomatal conductance, and activated enzymatic antioxidants (SOD, APX)”
Claim · model-written summary
Allantoin enhanced N assimilation in cucumber, with polyamine-mediated membrane stability.
- Taxon
- cucumberCucumis sativuse0
- Treatment
- allantoine6
- Outcome
- N assimilatione6
- Qualifier
- polyamine-mediated membrane stabilitye6
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Salinity stress severely limits cucumber (Cucumis sativus L.) productivity”
e6“Moreover, allantoin enhanced N assimilation and polyamine-mediated membrane stability, while citrulline-derived NO optimized ion transport and ROS scavenging”
Claim · model-written summary
Citrulline-derived NO optimized ion transport and ROS scavenging in cucumber.
- Taxon
- cucumberCucumis sativuse0
- Treatment
- citrulline-derived NOe6
- Outcome
- ion transport and ROS scavenginge6
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Salinity stress severely limits cucumber (Cucumis sativus L.) productivity”
e6“Moreover, allantoin enhanced N assimilation and polyamine-mediated membrane stability, while citrulline-derived NO optimized ion transport and ROS scavenging”
Claim · model-written summary
Combined foliar application improved cucumber growth (yield) by 44.24% compared with control.
- Taxon
- cucumberCucumis sativuse0
- Treatment
- combined applicatione6
- Outcome
- growthe6
- Direction
- improvede6
- Value
- 44.24%e6
- Qualifier
- in yielde6
- Comparator
- controle7
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Salinity stress severely limits cucumber (Cucumis sativus L.) productivity”
e7“foliar treatments (control, allantoin, citrulline, and their combination)”
e6“Their combined application synergistically improved growth (44.24% in yield), photosynthesis (5.12% in Fv/Fm), and RWC (6.45%)”
Claim · model-written summary
Combined foliar application improved photosynthesis (Fv/Fm) by 5.12% compared with control.
- Taxon
- cucumberCucumis sativuse0
- Treatment
- combined applicatione6
- Outcome
- photosynthesise6
- Direction
- improvede6
- Value
- 5.12% in Fv/Fme6
- Comparator
- controle7
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Salinity stress severely limits cucumber (Cucumis sativus L.) productivity”
e7“foliar treatments (control, allantoin, citrulline, and their combination)”
e6“Their combined application synergistically improved growth (44.24% in yield), photosynthesis (5.12% in Fv/Fm), and RWC (6.45%)”
Claim · model-written summary
Combined foliar application improved RWC by 6.45% compared with control.
- Taxon
- cucumberCucumis sativuse0
- Treatment
- combined applicatione6
- Outcome
- RWCe6
- Direction
- improvede6
- Value
- 6.45%e6
- Comparator
- controle7
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Salinity stress severely limits cucumber (Cucumis sativus L.) productivity”
e7“foliar treatments (control, allantoin, citrulline, and their combination)”
e6“Their combined application synergistically improved growth (44.24% in yield), photosynthesis (5.12% in Fv/Fm), and RWC (6.45%)”
Claim · model-written summary
Combined foliar application reduced H2O2 by 61.53% compared with control.
- Taxon
- cucumberCucumis sativuse0
- Treatment
- combined applicatione6
- Outcome
- H2O2e5
- Direction
- reducinge5
- Value
- 61.53%e5
- Comparator
- controle7
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Salinity stress severely limits cucumber (Cucumis sativus L.) productivity”
e7“foliar treatments (control, allantoin, citrulline, and their combination)”
e5“while reducing H2O2 (61.53%) and oxidative stress (43.74% MDA)”
e6“Their combined application synergistically improved growth (44.24% in yield), photosynthesis (5.12% in Fv/Fm), and RWC (6.45%)”
Claim · model-written summary
Combined foliar application reduced oxidative stress (MDA) by 43.74% compared with control.
- Taxon
- cucumberCucumis sativuse0
- Treatment
- combined applicatione6
- Outcome
- oxidative stresse5
- Direction
- reducinge5
- Value
- 43.74%e5
- Comparator
- controle7
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Salinity stress severely limits cucumber (Cucumis sativus L.) productivity”
e7“foliar treatments (control, allantoin, citrulline, and their combination)”
e5“while reducing H2O2 (61.53%) and oxidative stress (43.74% MDA)”
e6“Their combined application synergistically improved growth (44.24% in yield), photosynthesis (5.12% in Fv/Fm), and RWC (6.45%)”
Claim · model-written summary
Salt-stressed cucumber treated with both compounds had elevated fruit levels of health-promoting metabolites.
- Taxon
- cucumberCucumis sativuse0
- Treatment
- both compoundse5
- Outcome
- fruit levels of health-promoting metabolitese5
- Direction
- elevatede5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Salinity stress severely limits cucumber (Cucumis sativus L.) productivity”
e5“salt-stressed plants treated with both compounds exhibited elevated fruit levels of health-promoting metabolites (cucurbitacin, citrulline)”
Claim · model-written summary
Combined treatment increased proline and soluble sugars in salt-stressed cucumber.
- Taxon
- cucumberCucumis sativuse0
- Treatment
- Combined treatmente5
- Outcome
- proline and soluble sugarse5
- Direction
- increasede5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Salinity stress severely limits cucumber (Cucumis sativus L.) productivity”
e5“Combined treatment increased proline and soluble sugars”
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