Abstract
Polyamines (PAs) play a vital role in response to abiotic stress of higher plants. However, limited studies examined the interplay between PAs and signal molecules. Therefore, in this study, we tried to elucidate the crosstalk between PAs, abscisic acid (ABA), nitric oxide (NO) and hydrogen peroxide (H2O2) under chilling stress conditions were determined using tomato seedlings (Lycopersicon esculentum Mill.) cv. Moneymaker). The study showed that during chilling stress (4 oC, 0 h, 12 h and 24 h), application of Spd and Spm could elevate NO and H2O2 levels, enhance nitrite reductase (NR), nitric oxide synthase (NOS)-like, and polyamine oxidase (PAO) activities, upregulate LeNR relative expression, but did not influence LeNOS1 expression. While putrescine (Put) treatment had no this obvious impact in tomato leaves. Moreover, H2O2 might act as an upstream signal to stimulate NO production under chilling stress. In recovery period (25/15 oC, 10 h), above-mentioned parameters induced by PAs application were back to the control levels. In turn, seedlings pretreated with sodium nitroprusside (SNP, an NO donor) improved Put and Spd levels through increasing arginine decarboxylase (LeADC, LeADC1), ornithine decarboxylase (LeODC) and Spd synthase (LeSPDS) expressions in tomato leaves throughout treatment period. Put content appeared increases first, and then Spd content began to rise under chilling stress. Exogenously applied NO did not improved S-adenosylmethionine decarboxylase (LeSAMDC) and Spm synthase (LeSPMS) expressions, which led to constant Spm level under chilling stress and recover period. On the other hand, exogenous Put significantly increased ABA content and LeNCED1 transcript level, ABA treatment could alleviate the electrolyte leakage (EL) induced by D-arginine (D-Arg, an inhibitor of Put). Taken together, it is concluded that Spd and Spm enhanced NO production in tomato seedlings in H2O2-dependent way under chilling stress, which through NR and NOS pathways. ABA is involved in Put induced chilling stress tolerance. Furthermore, NO could improve the Put and the Spd contents under chilling stress.
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Claims 20
Claim · model-written summary
Spd application elevated NO and H2O2 levels during chilling stress.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Treatment
- Spde0
- Outcome
- NO and H2O2 levelse0
- Direction
- elevatee0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“application of Spd and Spm could elevate NO and H2O2 levels”
Claim · model-written summary
Spm application elevated NO and H2O2 levels during chilling stress.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Treatment
- Spme0
- Outcome
- NO and H2O2 levelse0
- Direction
- elevatee0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“application of Spd and Spm could elevate NO and H2O2 levels”
Claim · model-written summary
Spd application enhanced NR, NOS-like and PAO activities during chilling stress.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Treatment
- Spde0
- Outcome
- nitrite reductase (NR), nitric oxide synthase (NOS)-like, and polyamine oxidase (PAO) activitiese0
- Direction
- enhancee0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“application of Spd and Spm could elevate NO and H2O2 levels, enhance nitrite reductase (NR), nitric oxide synthase (NOS)-like, and polyamine oxidase (PAO) activities”
Claim · model-written summary
Spm application enhanced NR, NOS-like and PAO activities during chilling stress.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Treatment
- Spme0
- Outcome
- nitrite reductase (NR), nitric oxide synthase (NOS)-like, and polyamine oxidase (PAO) activitiese0
- Direction
- enhancee0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“application of Spd and Spm could elevate NO and H2O2 levels, enhance nitrite reductase (NR), nitric oxide synthase (NOS)-like, and polyamine oxidase (PAO) activities”
Claim · model-written summary
Spd application upregulated LeNR relative expression but did not influence LeNOS1 expression.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Treatment
- Spde1
- Gene
- LeNRe0
- Outcome
- LeNR relative expressione0
- Direction
- upregulatee0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“upregulate LeNR relative expression, but did not influence LeNOS1 expression”
e1“application of Spd and Spm”
Claim · model-written summary
Spm application upregulated LeNR relative expression.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Treatment
- Spme1
- Gene
- LeNRe0
- Outcome
- LeNR relative expressione0
- Direction
- upregulatee0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“upregulate LeNR relative expression”
e1“application of Spd and Spm”
Claim · model-written summary
Spd and Spm did not influence LeNOS1 expression.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Gene
- LeNOS1e0
- Outcome
- LeNOS1 expressione0
- Direction
- did not influencee0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“did not influence LeNOS1 expression”
Claim · model-written summary
Put treatment lacked the obvious effects seen for Spd and Spm in tomato leaves.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Organ / stage
- tomato leavese0
- Treatment
- putrescine (Put) treatmente0
- Outcome
- no this obvious impacte0
- Direction
- noe0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“putrescine (Put) treatment had no this obvious impact in tomato leaves”
Claim · model-written summary
H2O2 may stimulate NO production upstream under chilling stress.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Compound
- H2O2e0
- Outcome
- NO productione0
- Qualifier
- under chilling stresse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“H2O2 might act as an upstream signal to stimulate NO production under chilling stress”
Claim · model-written summary
Parameters induced by polyamine application returned to control levels during recovery.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Outcome
- above-mentioned parameterse0
- Direction
- back to the control levelse0
- Qualifier
- recovery period (25/15 oC, 10 h)e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“In recovery period (25/15 oC, 10 h), above-mentioned parameters induced by PAs application were back to the control levels”
Claim · model-written summary
SNP pretreatment improved Put and Spd levels in tomato leaves.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Organ / stage
- tomato leavese0
- Treatment
- sodium nitroprusside (SNP, an NO donor)e0
- Outcome
- Put and Spd levelse0
- Direction
- improvede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“seedlings pretreated with sodium nitroprusside (SNP, an NO donor) improved Put and Spd levels through increasing arginine decarboxylase (LeADC, LeADC1), ornithine decarboxylase (LeODC) and Spd synthase (LeSPDS) expressions in tomato leaves throughout treatment period”
Claim · model-written summary
SNP pretreatment increased LeADC, LeADC1, LeODC and LeSPDS expression.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Treatment
- sodium nitroprusside (SNP, an NO donor)e0
- Outcome
- arginine decarboxylase (LeADC, LeADC1), ornithine decarboxylase (LeODC) and Spd synthase (LeSPDS) expressionse0
- Direction
- increasinge0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“seedlings pretreated with sodium nitroprusside (SNP, an NO donor) improved Put and Spd levels through increasing arginine decarboxylase (LeADC, LeADC1), ornithine decarboxylase (LeODC) and Spd synthase (LeSPDS) expressions”
Claim · model-written summary
Put content rose before Spd content under chilling stress.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Outcome
- Put contente0
- Direction
- increasese0
- Qualifier
- firste0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“Put content appeared increases first, and then Spd content began to rise under chilling stress”
Claim · model-written summary
Spd content rose after Put content under chilling stress.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Outcome
- Spd contente0
- Direction
- risee0
- Qualifier
- thene0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“Put content appeared increases first, and then Spd content began to rise under chilling stress”
Claim · model-written summary
Exogenous NO did not improve LeSAMDC or LeSPMS expression.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Treatment
- NOe0
- Outcome
- S-adenosylmethionine decarboxylase (LeSAMDC) and Spm synthase (LeSPMS) expressionse0
- Direction
- did not improvede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“Exogenously applied NO did not improved S-adenosylmethionine decarboxylase (LeSAMDC) and Spm synthase (LeSPMS) expressions”
Claim · model-written summary
Spm level remained constant during chilling stress and recovery.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Outcome
- Spm levele0
- Direction
- constante0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“constant Spm level under chilling stress and recover period”
Claim · model-written summary
Exogenous Put significantly increased ABA content.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Treatment
- Pute0
- Outcome
- ABA contente0
- Direction
- increasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“exogenous Put significantly increased ABA content and LeNCED1 transcript level”
Claim · model-written summary
Exogenous Put significantly increased LeNCED1 transcript level.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Treatment
- Pute0
- Gene
- LeNCED1e0
- Outcome
- LeNCED1 transcript levele0
- Direction
- increasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“exogenous Put significantly increased ABA content and LeNCED1 transcript level”
Claim · model-written summary
ABA treatment alleviated D-Arg-induced electrolyte leakage.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Treatment
- ABA treatmente0
- Outcome
- electrolyte leakage (EL)e0
- Direction
- alleviatee0
- Qualifier
- D-arginine (D-Arg, an inhibitor of Put)e0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“ABA treatment could alleviate the electrolyte leakage (EL) induced by D-arginine (D-Arg, an inhibitor of Put)”
Claim · model-written summary
Spd and Spm enhanced NO production through an H2O2-dependent pathway under chilling stress.
- Taxon
- tomato seedlings (Lycopersicon esculentum Mill.)
Solanum lycopersicumt0 - Treatment
- Spd and Spme0
- Compound
- H2O2e0
- Outcome
- NO productione0
- Qualifier
- NR and NOS pathwayse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tomato seedlings (Lycopersicon esculentum Mill.)”
e0“Spd and Spm enhanced NO production in tomato seedlings in H2O2-dependent way under chilling stress, which through NR and NOS pathways”
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8 of 11 groups shown: named compounds and organisms first, then groups whose results differ (checked claim by claim), then those with positive and negative results, then the smaller groups.
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.