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Supplemental Information
Chemical Priming Against Multiple Abiotic Stresses: Mission Possible?Andreas Savvides1,2, Shawkat Ali3, Mark Tester3, Vasileios Fotopoulos1
1Department of Agricultural Sciences, Biotechnology and Food Science, Cyprus University of Technology, 3603 Lemesos, Cyprus.2Agrisearch Innovations Ltd, 2108 Nicosia, Cyprus. 3Center for Desert Agriculture, Division of Biological and Environmental Sciences and Engineering, King Abdullah University of Science and Technology, Thuwal 23955-6900, Saudi Arabia.Correspondence: [email protected] (V. Fotopoulos)
Table 1. The Effects Of Plant Pre-Treatment With The Chemical Agents Sodium Nitroprusside (SNP; NO Donor), Hydrogen Peroxide (H2O2), Sodium Hydrosulfide
(NaHS; H2S Donor), Melatonin, And Polyamines On The Molecular, Biochemical, Physiological, And Growth Traits Of Plants Subjected To Different Abiotic Stresses
Compared With Those Of Plants Without Pre-Treatment.
Priming
agent
Abiotic
stress
Plant
species
TissueMethod of
applicatio
n
Agent
treatment
duration/
interval
duration/Str
ess duration
Agent
concen-
tration
Molecular effects Biochemical effects
Physiological
and growth
effects
Summary/highlights Refs
Sodium Salinity Rice Treated: Nutrient 2d/0d/8d Tested: -↑Expression of stress-related -↑SOD, POX, CAT, APX, GR activities -↑surviving - Alleviation of the stress impacts is limited within S1
nitroprusside
(SNP)
Roots
Sampled:
Leaves
solution 0-1000μM
Optimum:
1μΜ
Toxic:
100 &
1000μM
genes (SPS, P5CS, HSP26) green parts (%),
PSII quantum
yield
a certain agent concentration range.
Citrus Treated:
Roots
Sampled:
Leaves
Incubation 2d/0d/16d Tested:
100μM
-↓DNA strand cleavage, protein
oxidation (carbonylation)
-↑SOD, CAT, APX, GR activities,
[ASC/DHA]a,
[GSH/GSSG], ∙OH scavenging activity
--- - Long-lasting systemic prime effect on the
antioxidant-related tolerance mechanism.
S2
Wheat Treated:
Seed
Sampled:
Germinating
seeds
Presoaking 20h/0d/5d Tested:
0.1mM
--- -↑ATP synthesis, [soluble sugars], [K+],
α-amylase activity, SOD & CAT activities,
-↓[seed starch], [Na+], [MDA], H2O2 &
O2•− release rate in mitochondria
-↑seed
respiration rate,
germination
rate, coleoptile
& radicle DW
-↓seed DW
- Alleviation of the impacts on seed germination,
seedling establishment & cell membrane integrity
- Enhanced antioxidant capacity & maintained
[Na+]/ [K+] balance.
S3
Wheat Treated:
Roots portion
Sampled:
Leaves
Incubation 1d/0d/4d Tested:
1mM
--- - ↑[AsA], [GSH], [GSH/GSSG], MDHAR,
DHAR, GR, GST, GPX, CAT, Gly I, Gly II
activities
- ↓[H2O2], [MDA]
-↓ Leaf
chlorosis/yellowi
ng
- Exogenous NO rendered the plants more
tolerant to salt-induced oxidative damage by
enhancing their antioxidant defense and MG
detoxification systems
S4
Citrus Treated:
Roots
Sampled:
Roots & leaves
Incubation 48h/0d/8d Tested:
100μM
- ↓ H2O2 and NO sources-related
transcripts after chemical agent
treatment in leaves & at the end
of the salt stress in roots, protein
carbonylation & nitration in leaves,
protein nitrosylation in roots
-↑ H2O2 and NO sources-related
transcripts at the end of the salt
stress in leaves, protein
nitrosylation in leaves
- ↑ local & systemic [H2O2] & [NO], [Chl],
- ↓ EL
- ↑
Photosynthetic
capacity
- No leaf
necrosis or
rolling
- H2O2 and NO generation are interlinked in
citrus, forming part of an active interplay
mechanism at the whole-plant level. This
mechanism involves spatio-temporal organization
and tissue-specific control at the transcriptional
level.
- ROS/RNS-mediated protein post-translational
modifications are a key molecular strategy for
signaling transduction and salinity acclimation.
S5
Strawberry Treated: Incubation Tested: Tested: - ↑Expression of genes associated - =[NO], [H2O2] -↑RWC, Ψw, - Alleviation of the stress impacts on leaf turgor, S6
Roots
Sampled:
Leaves
2d/0 or 7d/8d
Optimum:
2d/0d/8d
100μM with enzymatic antioxidants (cAPX,
CAT, GR, MnSOD, MDHAR, DHAR)
and ASC, GSH and RNS
biosynthesis
-↑[Chl], [Crt], [ASC/DHA],
[GSH/GSSG]
-↓EL, [MDA]
photosynthetic
electron
transport rate
-↓leaf wilting,
necrosis
photosynthesis and cell membrane integrity
- Redox regulation & antioxidant signaling are
key-targets of NO priming against salt stress
Drought Rice Treated:
Seed or leaves
Sampled:
leaves
Tested:
Presoaking
or foliar
spraying
Optimum:
F. spraying
2/---/21 seed
0/---/21 foliar
Tested:
0-150 μM
Optimum:
100 μM
--- - ↑[Free proline]
- ↓EL, [H2O2], [MDA]
- ↑
Photosynthetic
rate, RWC, Ψw,
Ψπ, Ψp,
seedling FW,
DW and height
- ↓ gs
- Maintenance of tissue water potential, enhanced
antioxidant system, improved stability of cellular
membranes, enhanced photosynthetic capacity
S7
Trifoliate
orange
Treated:
Shoot & roots
Sampled:
leaves
Spraying &
soil irrigation
14d/3d/7d Tested:
100 μM
--- -↑[NO], [Chl], SOD, POD, CAT activities
-↓ EL, [H2O2], [O2•−]
-↑
photosynthetic
rate
-↓ stomatal
aperture, gs
- Modulation of stomatal response, activation of
the antioxidant enzymes
S8
Heat Rice Treated:
Roots
Sampled:
Leaves
Nutrient
solution
Tested:
2d/0d/5h
Tested:
0-1000μM
Optimum:
1μΜ
-↑Expression of stress-related
genes (SPS, P5CS, HSP26)
-↑SOD, POX, CAT, APX, GR activities -↑surviving
green parts (%),
PSII quantum
yield
- Alleviation of the stress impacts is limited within
a certain agent concentration range.
S1
Maize Treated:
---
Sampled:
Roots &
coleoptiles
--- 6h/0d/18h Tested:
0-400μM
Optimum:
150μM
Toxic:
≥200μM
--- -↑ L-DES activity, [H2S]
-↓EL, [MDA]
-↑Seedlings
survival
percentage,
tissue vitality
- Improved heat tolerance in a concentration-
dependent manner
- Crosstalk between NO and H2S in the
acquisition of heat stress tolerance
S9
Chilling Cucumber Treated:
Shoot
Sampled:
Leaves
Spraying 3d/0d/2d Tested:
0-2mM
Optimum:
1mM
--- - =[soluble protein]
-↑[Chl], [soluble sugars], SOD, POD, CAT,
GR activities
-↓[MDA]
-↓ Chilling
damage index
(chlorosis,
crinkled leaf
- Improved chilling tolerance by the activation of
antioxidant enzymes and osmotic adjustment
substances.
S10
Toxic:
2mM
edge, wilting,
dead leaves)
Wheat Treated:
Seed
Sampled:
Seeds & leaves
Presoaking 20h/0d/--- Tested:
0.1mM
- ↑ SOD encoding gene (Mn-SOD)
expression in seeds & seedlings
-↑α-amylase activity, SOD activity in
seeds & seedlings, [Chl], [soluble
sugars], [sucrose]
-↓[H2O2], [MDA], [starch]
-↑ Germination
rate, seed, seed
respiration rate,
seedling
establishment
and growth
- Improved cold tolerance by promoting β-
amylase activity hydrolysis of starch into
soluble sugars seed respiration & growth
& by enhancing the antioxidant system
S11
Tomato Treated:
Seed
Sampled:
Germinating
seeds &
seedlings
Presoaking 1d/0d/10d Tested:
200μΜ
--- -↑α-amylase, β-amylase activities,
[soluble sugars], seed [NO]
-↑ Germination
capacity, index,
seedling root
and shoot
length
- Improved cold tolerance at least partly by
promoting α- & β-amylase activity
S12
Excess
light
Tall Fescue Treated:
Root
Sampled:
Leaves
Nutrient
solution
---/---/7d Tested:
0-500μM
Optimum:
100μM
Toxic:
≥200μM
--- -↑SOD, CAT, APX, GR activities, NO
release
-↓EL, [MDA], [H2O2], [O2•−], LOX & NOS
activity
--- - Enhanced antioxidant enzymes activity &
reduced [ROS] & LOX activity prevented lipid
peroxidation and membrane damage
S13
Heavy
metals
(Cd)
Alfalfa Treated:
Roots
Sampled:
Roots
Nutrient
solution
6h/0h/12h Tested:
100μM
-↑Transcript levels of Cu/Zn-SOD
and POD
-↑SOD & POD activities, [NO] at the
onset of stress
-↓[TBARS], LOX activity, [Cd], [NO] at the
end of stress
-↑ Plant FW - Improved tolerance may be related with:
suppressed Cd uptake, enhanced antioxidant
system.
S14
Hydrogen
peroxide
Salinity Rice Treated:
Roots
Nutrient
solution
Tested:
2d/0d/8d
Tested:
0-1000μM
-↑Expression of stress-related
genes (SPS, P5CS, HSP26)
-↑SOD, POX, CAT, APX, GR activities -↑surviving
green parts (%),
- Alleviation of the stress impacts is limited within
a certain agent concentration range.
S1
(H2O2) Sampled:
Leaves
Optimum:
10μΜ
PSII quantum
yield
Citrus Treated:
Roots
Sampled:
Leaves
Incubation Tested:
8h/0d/16d
Tested:
10mM
-↓DNA strand cleavage, protein
oxidation (carbonylation)
-↑SOD, CAT, APX, GR activities,
[ASC/DHA],
[GSH/GSSG], ∙OH scavenging activity
--- - Long-lasting systemic prime effect on the
antioxidant-related defense mechanism.
S2
Wheat Treated:
Seed
Sampled:
Leaves/ shoot
Presoaking 8h/0h/15d after
germination
Tested:
0-120μM
Optimum:
80μM
-↑Expression of heat-stable (stress)
proteins (32kDa, 52kDa)
- = shoot [Na+]
-↑shoot [K+], [K+]:[Na+], [Ca2+],
[NO3-], [PO43-]
-↓ shoot [H2O2], membrane
permeability, ion leakage
-↑ Germination
rate, shoot FW &
DW, leaf area,
net
photosynthesis,
gs, Ψw, Ψp
- H2O2-enhanced tolerance is linked to the
activation of antioxidants, maintaining turgor and
meeting plant nutritional requirement.
S15
Citrus Treated:
Roots
Sampled:
Roots & leaves
Incubation 8h/0d/8d Tested:
10mM
- ↓ H2O2 and NO sources-related
transcripts after chemical agent
treatment in leaves & at the end
of the salt stress in roots, protein
carbonylation & nitration in leaves,
protein nitrosylation in roots
-↑ H2O2 and NO sources-related
transcripts at the end of the salt
stress in leaves, protein
nitrosylation in leaves
- ↑ local & systemic [H2O2] & [NO], [Chl],
- ↓ EL
- ↑
Photosynthetic
capacity
- No leaf
necrosis or
rolling
- H2O2 and NO generation are interlinked in
citrus, forming part of an active interplay
mechanism at the whole-plant level. This
mechanism involves spatio-temporal organization
and tissue-specific control at the transcriptional
level.
- ROS/RNS-mediated protein post-translational
modifications are a key molecular strategy for
signaling transduction and salinity acclimation.
S5
Strawberry Treated:
Roots
Sampled:
Leaves
Incubation Tested:
8h/0 or 7d/8d
Optimum:
8h/0d/8d
Tested:
10mM
-↑Expression of genes associated
with enzymatic antioxidants (cAPX,
CAT, GR, MnSOD, MDHAR, DHAR)
and ASC, GSH and RNS
biosynthesis
- =[NO], [H2O2]
-↑[Chl], [Crt], [ASC/DHA],
[GSH/GSSG]
-↓EL, [MDA]
-↑RWC, Ψw,
photosynthetic
electron
transport rate
-↓leaf wilting,
necrosis
- Redox regulation & antioxidant signaling are
key-targets of H2O2 priming against salt stress
S6
Ginseng Treated:
Roots
Sampled:
Shoot & roots
Nutrient
solution
2d/0d/7d Tested:
0.05-250μM
Optimum:
100μM
-↑Expression of defense-related
genes (PGIP, ribonuclease-2 (PR-
10), chitinase (PR-2), calmodulin,
sesquiterpene synthase,
-↑APX, CAT, POX activities, [Chl], [Crt],
[Proline]
-↓[H2O2], [O2•−], [MDA]
-↑Plant height,
plant DW, RWC
- Alleviation of the stress impacts is related to
enhanced antioxidant system and osmotic
adjustment.
S16
Toxic:
>100μM
spermidine synthase)
Drought Wheat Treated:
Seed
Sampled:
Leaves/
Seedlings
Presoaking 6h/---/--- (exp.
ended when 3rd
leaf was fully
expanded)
Tested:
0-140mM
Optimum:
80mM
Toxic:
>80mM
--- -↑[Proline], CAT, APX activities
-↓[H2O2], [MDA]
-↑ Germination
rate, net
photosynthesis,
leaf area,
seedling DW,
WUE
- Alleviation of the stress impacts is related to
enhanced antioxidant system and osmotic
adjustment.
S17
Soybean Treated:
Shoot
Sampled:
Leaves
Spraying Sprayed
once/0d/8d
Tested:
1mM
-↑mRNA levels of GmMIPS2 & GolS
(encode key-enzymes for
oligosaccharides biosynthesis)
-↑[myo-inositol] & [galactinol] -↑ RWC, gs, net
photosynthesis,
-↓ Leaf wilting
- Enhanced tolerance due to the water content
maintenance caused by enhanced
oligosaccharides biosynthesis rather than by
rapid stomatal closure
S18
Heat Rice Treated:
Roots
Sampled:
Leaves
Nutrient
solution
2d/0d/5h Tested:
0-1000μM
Optimum:
10μΜ
-↑Expression of stress-related
genes (SPS, P5CS, HSP26)
-↑SOD, POX, CAT, APX, GR activities -↑surviving
green parts (%),
PSII quantum
yield
- Alleviation of the stress impacts is limited within
a certain agent concentration range
S1
Cucumber Treated:
Shoot
Sampled:
Leaves
Spraying Sprayed
once/12h/3d
Tested:
1.5mM
--- -↑GSH-Px, DHAR, MDHAR, GR, SOD, APX
-↓[H2O2], [MDA]
-↓abnormal
chloroplasts
- Protection of chloroplasts’ ultrastructure by the
enhanced antioxidant activity & decreased lipid
peroxidation
S19
Chilling Tomato Treated:
Roots
Sampled:
Leaves & roots
Incubation 1h/4d/16h Tested:
1mM
Optimum:
--- - = leaf APX, CAT activities, leaf [H2O2],
-↑leaf [anthocyanins], root [proline], root
APX, CAT activities
-↓root [H2O2], leaf & root [MDA]
-↑ RWC - Alleviation of the chilling stress maybe due to
enhanced antioxidant system and cellular
osmotic adjustment
S20
Excess
light
Tobacco Treated:
Shoot
Sampled:
Leaves
Spraying Sprayed
once/1d/1d
Tested:
0-500mM
Optimum:
5mM
--- -↑ CAT, APX & POX activities, GPX protein
levels, [GSH/GSSG]
-↓[MDA]
--- - Alleviation of the stress impacts is limited within
a certain agent concentration range
S21
Toxic:
500mM
Heavy
metals
(Cd)
Rice Treated:
---
Sampled:
Shoot & roots
--- 1d/0d/7d Tested:
100μΜ
--- - = Root [Cd], root APX, leaf GST, leaf
CAT activities, leaf [GSH], leaf & root
[GSSG], leaf & root [AsA], leaf [H2O2]
-↑ leaf & root SOD, root CAT, leaf & root
GPX, leaf APX, root GST activities, root
[GSH]
-↓ Shoot [Cd], Shoot [Cd] / Root [Cd],
root [H2O2], leaf & root [MDA]
-↑Shoot & root
length & DW
- The enhanced tolerance maybe due to a
stimulated antioxidant system and Cd
sequestration.
- The synthesis of GSH and GST were also
enhanced, which seems to regulate Cd
distribution in rice seedlings
S22
(Cr(VI)) Canola Treated:
Roots
Sampled:
Shoot & roots
Nutrient
solution
24h/0d/7d Tested:
200μM
-↑expression level of BnMP1 mRNA - = Root [Cr], SOD & CAT activities
-↑ Shoot [Cr], [Chl], POD & APX activities,
[Protein-bound & non-protein thiols]
-↓[MDA]
-↑Seedlings FW
& DW
- Thiol-containing chelators may play an
important role in the sequestration of Cr
S23
Sodium
hydrosulfide
(NaHS)
Salinity Strawberry Treated:
Roots
Sampled:
Leaves
Incubation Tested:
2d/0d/7d or
2d/3d/7d
Optimum:
2d/0d/7d
Tested:
100 μΜ
- Coordinated regulation of
multiple transcriptional
pathways
key antioxidant, ascorbate and
glutathione biosynthesis, RNS
biosynthesis, dehydration-
responsive element binding factor,
(DREB), and salt overly sensitive
(SOS) pathway genes.
- ↑[H2S], [ASC/DHA], [GSH/GSSG]
- ↓[MDA], [ROS], [RNS]
- ↑ RWC,
photosynthetic
capacity, gs, Ψw
- ↓ Wilting, foliar
injury
- Oxidative and nitrosative cellular damage
control through the increased performance of
antioxidative mechanisms and the coordinated
regulation of the SOS pathway
- Primed state is not long-lasting
S24
Drought Wheat Treated:
Roots
Sampled:
Leaves
Incubation 12h/0h/24h Tested:
0.5-2mM
Optimum:
1mM
Toxic:
≥1.5mM
--- - = MDHAR, GalLDH activity
- ↑[AsA], [GSH], [total ascorbate], [total
glutathione], APX, GR, DHAR, γ-ECS
activities
- ↓[MDA], EL
--- - H2S enhances the antioxidant ability
and protects wheat seedlings against oxidative
stress
induced by water stress
S25
Strawberry Treated:
Roots
Sampled:
Incubation Tested:
2d/0d/7d or
2d/3d/7d
Tested:
100 μΜ
Coordinated regulation of multiple
transcriptional pathwayskey
antioxidant, ascorbate and
-↑[H2S], [ASC/DHA], [GSH/GSSG]
-↓[MDA], [ROS], [RNS]
- ↑ RWC,
photosynthetic
capacity, gs, Ψw
- Oxidative and nitrosative cellular damage
control through the increased performance of
antioxidative mechanisms and the coordinated
S24
Leaves Optimum:
2d/0d/7d
glutathione biosynthesis, RNS
biosynthesis, dehydration-
responsive element binding factor,
(DREB), and salt overly sensitive
(SOS) pathway genes.
- ↓ Wilting, foliar
injury
regulation of the SOS pathway
- Primed states is not long-lasting
Heat Maize Treated:
---
Sampled:
Roots &
coleoptiles
--- 6h/0d/18h Tested:
0-1500μM
Optimum:
500μM
Toxic:
≥1000μM
--- -↑ L-DES activity, [H2S]
-↓EL, [MDA]
-↑Seedlings
survival
percentage,
tissue vitality
- H2S improves heat tolerance in a concentration-
dependent manner
S9
Strawberry Treated:
Roots
Sampled
Leaves
Incubation 48h/0h/8h Tested:
100μΜ
- Induction of gene expression
levels of enzymatic antioxidants
(cAPX, CAT, MnSOD, GR), heat
shock proteins (HSP70, HSP80,
HSP90), aquaporins (PIP)
- ↑[ASC], [GSH], [ASC/DHA], [GSH/GSSG]
- ↓ [H2S], [H2O2], [NO], [DHA], [GSSG],
[MDA]
- ↑
Photosynthetic
capacity
- ↓ Leaf wilting,
curling
- H2S root pretreatment activates a coordinated
network of heat shock defense-related pathways
at a transcriptional level and systemically
protects strawberry plants from heat
shock-induced damage.
S26
Chilling Grapevine Treated:
Leaves
Sampled
Leaves
Spraying 1h/0h/--- Tested:
0.1mM
-↑expression levels of VvICE1 &
VvCBF3
- ↑SOD activity
- ↓[O2•−], [MDA]
--- - H2S is involved in grapevine chilling stress
response by modulating VvICE1 and VvCBF3
transcription.
S27
Excess
light
Dendrobium
officinale
Treated:
Leaves
Sampled:
Leaves
Spraying Sprayed
once/0h/4h
Tested:
0- 600μM
Optimum:
200mM
Toxic:
≥400mM
--- - ↑ CAT, SOD, POD activities
- ↓ MDA content
- ↑
Photosynthetic
capacity
- ↓
Photoinhibition
- Enhanced tolerance against high light stress by
possibly increasing activities of antioxidant
enzymes and enhancing photosynthetic electron
transfer.
S28
Heavy
metals
(Al
toxicity)
Wheat Treated:
Seed
Sampled:
Seed
Incubation 12h/0h/48h Tested:
0-1.5mM
Optimum:
0.6mM
--- - ↑ amylase, esterase activities, APX,
CAT, POD, SOD activities, [H2S]
-↓[H2O2], [O2•−], [MDA], LOX activity,
[Al]
- ↑ Seed
germination %,
radicle &
coleoptile
length, radicle
-The H2S donor-induced lowering of H2O2 and
MDA levels could be attributed to the increased
activities of ROS scavenging enzymes and the
decreased activity of LOX.
S29
number
(Cd) Alfalfa Treated:
Roots
Sampled:
Roots
Nutrient
solution
6h/0h/12h Tested:
0-100μM
Optimum:
100μM
-↑Transcript levels of Cu/Zn-SOD
and POD
-↑SOD & POD activities, [NO] at the
onset of stress
-↓[TBARS], LOX activity, [Cd], [NO] at the
end of stress
-↑ Plant FW - Improved tolerance may be related with:
suppressed Cd uptake, enhanced antioxidant
system.
- NO might be a component of the H2S-induced
cytoprotective role
S14
(Cd) Populus
euphratica
Treated:
Cells
Sampled:
Cells
Medium 6h/0h/72h Tested:
0-200μM
Optimum:
50-100μM
--- -↑ APX, CAT, GR activities, vacuolar [Cd] /
cytoplasmic [Cd]
-↓ [H2O2], [MDA], Cd influx through
plasma membrane
- ↓ Programmed
cell death
- H2S attenuated Cd toxicity
by 1) up-regulating activity of antioxidant
enzymes, 2) reducing Cd entry into the
cytoplasm, and 3) increasing Cd sequestration
into the vacuole.
S30
Melatonin Salinity Malus Treated:
Roots
Sampled:
Leaves & roots
Nutrient
solution
10d/0d/15d Tested:
0.1μM
- ↑ transcripts of MDNHX1,
MdAKT1 in the leaves
- ↑ [Chl], dehydrogenase activity, APX,
CAT, POD activities, root & stem [Na+],
root & stem & leaves [K+], leaves
[K+]/[Na+]
- ↓ EL, [H2O2], leaves [Na+]
- ↑ Plant height,
leaf number,
plant FW & DW,
net
photosynthesis,
- Melatonin is involved in the activation of
antioxidants & the modulation of ion
homeostasis as shown by the up-regulation of the
ion transporters in leaves
S31
Soybean Treated:
Seeds
Sampled:
Seeds & leaves
Coating ---/7d/22d Tested:
0-200 μM
Optimum:
50-100μM
-↑Many stress-responsive genes
e..g. ferredoxin gene (PetF),
ascorbate metabolism-related
genes (VTC4, APX4).
- ↑ [Chl]
- ↓ EL, [H2O2]
- ↑ Biomass,
shoot height,
leaf area
- Melatonin up-regulates many salt-inhibited
genes and may increase salt tolerance partly via
antioxidant capacity enhancement
S32
Bermudagras
s
Treated:
Roots
Sampled:
Leaves
Irrigation 7d/0d/25d Tested:
0- 100 μΜ
Optimum:
20, 100 μM
- ↑ Many stress-responsive genes:
e.g. CBF/DREB genes and target
genes, heat shock TFs, zing finger
TFs, WRKY, MYB, bHLH genes and
hormone-related genes
- ↑ [Melatonin], [Chl] SOD, CAT, POD
activities, GSH redox state, [metabolites]
e.g. amino acids, sugars, sugar alcohols
- ↓ [H2O2], [O2•−], EL, [MDA]
- ↑Survival rate,
plant height,
plant weight
- Melatonin is involved in the activation of
antioxidants, modulation of metabolic
homeostasis & extensive transcriptional
reprogramming.
S33
Drought Bermudagras
s
Treated:
Root
Sampled:
Irrigation 7d/0d/21d/4d
recovery
Tested:
0- 100 μΜ
Optimum:
Same as salt stress Same as salt stress Same as salt
stress
Same as salt stressS33
Leaves 20, 100 μM
Malus Treated:
Roots
Sampled:
Leaves
Nutrient
solution
10d/0d/6d Tested:
100 μΜ
-↓ Expression of melatonin
synthesis genes, ABA biosynthesis
genes, ABA catabolism genes
- ↑CAT, POD, APX enzymes activity, [Chl],
[Crt]
- ↓EL, [H2O2], [ABA]
- ↑ RWC, net
photosynthesis,
gs, stomatal
size and
aperture
- Melatonin enhances ABA degradation and
suppresses its synthesis and thus less H2O2
accumulates in guard cells, directly scavenges
H2O2, up-regulates the antioxidant system
S34
Heat Phacelia
tanacetifolia
Treated:
Seed
Sampled:
Germinating
seeds
Presoaking 2d/2h/7d Tested:
0-90μΜ
Optimum:
0.3-6μΜ
--- --- - ↑ germination
percentage &
rate,
- Melatonin enhances heat stress toleranceS35
Chilling Wheat Treated:
Shoot
Sampled:
Leaves
Spraying 12h/0d/3d Tested:
0,1mM
- ↑Rubisco expression - ↑[Chl], [Crt], [Proline], [soluble protein],
[Phenolic compounds], carbohydrate
accumulation, SOD, GPX, APX, GR
activities, [AsA], [GSH]
- ↓ [DHA], DHA/AsA, GSSG/GSH, [H2O2],
[O2•−], [MDA]
- ↑ RWC, leaf
area
- Melatonin directly scavenges H2O2, enhances
the antioxidant system and photosynthesis and
increase the cellular accumulation of
osmoprotectants.
S36
Arabidopsis Treated:
Roots
Sampled:
Seedlings
Culture
medium
Tested:
1d/0d/3d or
1d/0d/5d
Tested:
0 - 400 μM
Optimum:
10 - 40 μM
Toxic:
>100μM
- ↑ expression of:
C-repeat-binding factors (CBFs),
Drought Response Element
Binding factors (DREBs), a cold-
responsive gene, COR15a, a
transcription factor involved in
freezing and drought-stress
tolerance CAMTA1 and
transcription activators of reactive
oxygen species (ROS)-related
antioxidant genes, ZAT10 and
ZAT12
--- - ↑ FW, primary
root length, and
shoot height
- Melatonin induces
cold signaling genes up-regulation resulting in
cold-protecting compounds biosynthesis
S37
Freezing Bermudagras Treated: Irrigation 7/0/21cold/ Same as salt stress Same as salt stress Same as salt Same as salt stressS33
s Roots
Sampled:
Leaves
8freezing stress
Heavy
metal
(Cu
toxicity)
Red cabbage Treated:
Seed
Sampled:
Germinating
seeds &
seedlings
Hydropriming ---/3d/5d seed
exp.
---/3d/8d
seedling exp.
Tested:
1-100 μM
Optimum:
1-10 μM
Deleterious
:
100 μM
- ↓ DNA endoreduplication blocking - ↓ Lipid peroxidation ([TBARS]) - ↑ Seed
germination
percentage and
rate, seedling
FW
- Melatonin may enhance tolerance to Cu partly
by enhancing the antioxidant system
S38
Polyamines
(Spermidine)
Salinity Rice
(tolerant-
sensitive)
Treated:
Shoot
Sampled:
Grains
Spraying 7d/0d/--- Tested:
1mM
--- - ↓[Na+]/[K+] - ↑ Grain yield
(62% sensitive,
16% tolerant)
- Spermidine largely enhanced yield.
- Spermidine enhanced ion homeostasis.
S39
(Spermidine) Rice Treated:
Seed
Sampled:
Leaves
Presoaking 12h/17d/10d Tested:
1mM
--- - ↑ [Phenolic compounds], [Chl],
[anthocyanins], antioxidant capacity
- ↓[H2O2], [Na+]/[K+]
- ↑ shoot FW,
DW, length
- Long-lasting priming phenomenon
- Spermidine enhances salt tolerance by
increasing antioxidant capacity and ion
homeostasis
S40
(Spermidine) Salinity
&
alkalinity
Tomato Treated:
Seed
Sampled:
Roots
Presoaking 10h/-/6d Tested:
0.25mM
--- - = [free Putrescine]
- ↑ [Free spermidine], [free spermine]
- ↑ Root DW - Exogenous Spd can regulate the metabolic
status of polyamines
S41
(Spermine) Drought Citrus Treated:
Roots
Sampled:
Leaves
Incubation 5d/12h/12h Tested:
1mM
- ↑ Expression levels of ADC &
SPMS (polyamines biosynthesis)
- ↑ [Polyamines], SOD, POD activities
- ↓ EL, [H2O2], [O2•−]
- ↓ Water loss,
stomatal
aperture
- Spm pretreatment causes enhanced
endogenous polyamines, effective ROS
scavenging, and stimulated stomatal closure
S42
(Spermine) Soybean Treated:
Shoot
Sampled:
Pods
Spraying 3d/14d/--- Tested:
0.1mM
Optimum:
--- - = [GSH]
- ↑SOD, CAT activities, [polyphenols],
[Protein]
- ↓ [ABA], [MDA]
- ↑ Pods & seeds
FW,
- Long-lasting tolerance enhancement
- Increased yield
S43
(Spermidine) White clover Treated: Presoaking 3h/0d/7d Tested: - = Transcript level of α-amylase - ↑ α-amylase, β-amylase activities, seed
[reducing sugars], [fructose], [glucose],
- ↑ germination
%, vigor, index,
-The discrepancy between gene expression and
enzyme activity indicates enzyme activity
S44
Seed
Sampled:
Seed &
seedlings
30μM - ↑ Transcript level of β-amylase,
relative expression ratio of Cu/Zn
SOD, CAT, POD, APX
SOD, CAT, POD, APX activities, [AsA],
[AsA]/[DHA], [GSH], [GSH]/[GSSG]
- ↓ seed [starch], [MDA], EL, [H2O2],
[O2•−], [DHA]
rate root
viability, length,
seedling FW,
DW
changes were
not only caused by mRNA levels, but also
regulated at the post-transcriptional level and
influenced by
cellular metabolism
- the function of Spd is
double 1) a stress-protecting compound and 2)
signaling molecule
(Putrescine) Flooding Onion Treated:
Roots
Sampled:
Leaves
Soil
drenching
24h/0d/10d Tested:
0-3mM
Optimum:
2mM
--- - ↑ [Chl], scavenging activity, metal
chelating capacity
- ↓ [H2O2], [O2•−],
- ↑ RWC, plant
height,
photosynthetic
capacity
- Putrescine protective
action against flooding may be related to its iron
binding capacity & enhancement of antioxidant
system
S45
(Spermine, T-
spermine,
spermidine,
putrescine)
Heat Arabidopsis Treated:
Roots
Sampled:
Seedlings
Incubation ---/0h/2h Tested:
0-1mM
Optimum:
0.1mM
Toxic:
≥0.5mM
--- - ↑ [Chl] - ↑ vigorously
growing leaves
& roots
- ↓ cotyledons
chlorosis
- Spermine alleviated heat stress (HS) effects
best
- Spermine increases the HS response at
transcriptional
and translational levels (from research on
transgenic & mutant plants)
S46
(Spermidine) Rice Treated:
Shoot
Sampled:
Leaves
Spraying 24h/0h/48h/
recovery 48h
Tested:
0-2mM
Optimum:
1mM
--- - ↑ [Chl], [AsA], [GSH], AsA & GSH redox
status, SOD, CAT, GPX, GST, APX, DHAR,
GR, GLY I, GLY II activities, [free
spermidine], [soluble conjugated
spermidine]
- ↓ [MDA], [H2O2], [proline], LOX activity
- ↑ leaf FW - Enhancement of the antioxidative and
glyoxalase systems by spermidine rendered rice
seedlings more tolerant to heat stress
S47
(Spermine) Heat &
Drought
Trifoliate
orange
Treated:
Roots
Sampled:
Leaves
Incubation 30h/0h/3h Tested:
1mM
- ↑ expression of heat shock
proteins, abscisic acid (ABA)-
responsive element binding factor,
9-cic-epoxycarotenoid
dioxygenase 3
- ↑ SOD, POD, CAT activities
- ↓ EL, [H2O2], [O2•−],
- ↓ leaf wilting,
water loss
- Polyamines may confer tolerance to multiple
simultaneously applied stresses
S48
(Putrescine,
spermidine)
Chilling Cucumber Treated:
Shoot
Sampled:
Spraying Sprayed
once/12h/3d/
recovery 1d
Tested:
0.5mM
--- - ↑ endogenous [polyamines], [soluble
proteins], SOD, POD, CAT, APX activities
- ↓ EL, [MDA], [H2O2]
--- - Enhanced cold tolerance may be partly related
to enhanced antioxidant capacity and
osmoprotectants
S49
Leaves
(Putrescine,
spermidine,
spermine, 1,3-
diaminopropan
e,
diethylenetriam
ine)
Freezing Wheat Treated:
Shoot
Sampled:
Leaves
Spraying Sprayed
once/24h /24h
Tested:
100mg/l
--- - ↑ [Chl] , [Proline]
- ↓ EL
- ↑ shoot FW,
survival rate
- Polyamines are capable of preventing frost
injuries
- Diethylenetriamine was the most effective
S50
(Spermine)
Heavy
metals
(Cd)
Wheat Treated:
Leaf segments
Sampled:
Leaf segments
Incubation 6h/0h/14h Tested:
1mM
--- - = [GSH], APX activity
- ↑ [spermidine], [spermine], GR activity
- ↓ [TBARS], [H2O2]
--- - Spermine enhances Cd stress toleranceS51
(Spermine) (Cu) Wheat Treated:
Leaf segments
Sampled:
Leaf segments
Incubation 6h/0h/14h Tested:
1mM
--- - = [GSH], APX activity
- ↑[Putrescine], [spermidine], [spermine],
GR activity
- ↓ [TBARS] , [H2O2]
--- - Spermine enhances Cu stress tolerance partly
by enhancing antioxidant capacity
S51
Abbreviations: ABA: abscisic acid, APX: ascorbate peroxidase, AsA: ascorbic acid, ASC: reduced ascorbate, ASC/DHA: ascorbate redox state, CAT: catalase, Chl: chlorophyll, Crt: Carotenoids, DAO: diamine oxidase, DHA: dehydroascorbate, DHAR: dehydroascorbate reductase, DW: dry weight, EL: electrolyte leakage, FW: fresh weight, GalLDH: L-Galactono-1,4-lactone dehydrogenase, gs: stomatal conductance, GPX: glutathione peroxidase, GR: glutathione reductase, GSH: reduced glutathione, GSH/GSSG: glutathione redox state, GSH-Px: glutathione peroxidase, GSSG: oxidized glutathione, Gly I: glyoxalase I, Gly II: glyoxalase II, GST: glutathione S-transferase, L-DES: L-cystine desulfhydrase, LOX: lipoxygenase, MDA: malondialdehyde, MDHAR: monodehydrascorbate reductase, MG: methylglyoxal, NOS: nitric oxide synthase, ODC, ornithine decarboxylase, PAO: polyamine oxidase, POD: peroxidase, POX: guaiacol peroxidase, RLW: leaf relative water content, RNS: reactive nitrogen species, ROS: reactive oxygen species, SAMDC: S-adenosylmethionine decarboxylase, SOD: superoxide dismutase, TBARS: thiobarbituric acid reactive substances, WUE: water use efficiency, γ-ECS: gamma-glutamylcysteine synthetase, Ψp: pressure potential, Ψw: leaf water potential, Ψπ: osmotic potentiala[ ]: concentration, ↑: increase, ↓: decrease, =: not altered,
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