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. 2021 Mar 19;22(6):3154. doi: 10.3390/ijms22063154

Table 2.

The use of PGPR in seed biopriming technique for improving salinity stress tolerance in plants.

PGPR Treatments GP PhoPs AEs MDA Ion Content Pro Ethylene Sources
S. maltophilia SBP-9.
Bacterized wheat seeds with S. maltophilia SBP-9 for 1 h.
Control [102]
0 mM NaCl + SBP-9 ↑ 15% SL, ↑ 10% RL, ↑ 12% SFW,
↑ 17% SDW, ↑ 33% RFW,
↑ 9% RDW
↑ 8% Tchl ↑ 33% SOD
↑ 20% CAT
↑ 33% POD
↓ 27% In shoot: ↓ 4% Na+, ↑ 12% K+, Na+/K+ ratio ~0.54 ↓ 10% N/A
150 mM NaCl + Unprimed ↓ 11% SL, ↓ 5% RL, ↓ 8% SFW,
↓ 24% SDW, ↓ 11% RFW,
↓ 23% RDW
↓ 15% Tchl ↑ 58% SOD
↑ 7% CAT
↑ 67% POD
↑ 17% In shoot: ↑ 48% Na+, ↓ 17% K+, Na+/K+ ratio ~1.12 ↑ 74%
150 mM NaCl + SBP-9 ↑ 4% SL, ↑ 15% RL, ↑ 4% SFW,
↑ 7% SDW, ↑ 22% RFW, ↓ 5% RDW
↑ 5% Tchl ↑ 133% SOD
↑ 93% CAT
↑ 133% POD
↓ 13% In shoot: ↑ 20% Na+, ↑ 3% K+ Na+/K+ ratio ~0.73 ↑ 23%
200 mM NaCl + Unprimed ↓ 37% SL, ↓ 20% RL, ↓ 32% SFW,
↓ 38% SDW, ↓ 33% RFW,
↓ 64% RDW
↓ 59% Tchl ↑ 217% SOD
↑ 100% CAT
↑ 192% POD
↑ 93% In shoot: ↑ 107% Na+, ↓ 25% K+, Na+/K+ ratio ~1.73 ↑ 165%
200 mM NaCl + SBP-9 ↓ 11% SL, ↓ 5% RL, ↓ 16% SFW,
↓ 17% SDW, ↓ 6% RFW,
↓ 41% RDW
↓ 39% Tchl ↑ 350% SOD
↑ 180% CAT
↑ 283% POD
↑ 50% In shoot: ↑ 54% Na+, ↓ 3% K+ Na+/K+ ratio ~1 ↑ 110%
Enterobacter P23.
Seeds of Oryza sativa cv. Ratna treated with bacterial suspension.
Control [49]
150 mM NaCl + Unprimed ↓ 51% GP, ↓ 97% SVI, ↓ 45% SFW,
↓ 58% SDW, ↓ 33% SL,
↓ 39% RFW, ↓ 63% RDW, ↓ 44% RL
↓ 54% Chl a
↓ 80% Chl b
↓ 56% Tchl
↑ 120% SOD
↑ 112% CAT
↑ 174% POD
↑ 700% PPO
↑ 300% N/A ↑ 175% N/A
150 mM NaCl + P23 ↓ 22% GP, ↓ 58% SVI, ↓ 16% SFW,
↓ 11% SL, ↓ 23% SDW,
↓ 15% RFW, ↓ 30% RDW, ↓ 11% RL
↓ 13% Chl a
↓ 10% Chl b
↓ 8% Tchl
↑ 32% SOD
↑ 46% CAT
↑ 70% POD
↑ 300% PPO
↑ 195% ↑ 75%


B. pumilus FAB10.
Wheat seeds cv. 343 treated with FAB10.
Control [51]
75 mM NaCl + Unprimed ↓ 17% SL, ↓ 35% RL, ↓ 49% SDW,
↓ 53% RDW, ↓ 35% SpDW,
↓ 21% GY, ↓ 17% GPr
N/A ↑ 20% SOD
↑ 40% CAT
↑ 50% GR
↑ 189% N/A ↑ 105%
125 mM NaCl + Unprimed ↓ 24% SL, ↓ 49% RL, ↓ 42% SDW,
↓ 67% RDW, ↓ 48% SpDW,
↓ 31% GY, ↓ 22% GPr
↑ 23% SOD
↑ 80% CAT
↑ 75% GR
↑ 189% ↑ 146%
250 mM NaCl + Unprimed ↓ 35% SL, ↓ 52% RL, ↓ 40% SDW,
↓ 76% RDW, ↓ 61% SpDW,
↓ 41% GY, ↓ 29% GPr
↑ 25% SOD
↑ 80% CAT
↑ 75% GR
↑ 260% ↑ 171%
75 mM NaCl + FAB10 ↓ 13% SL, ↓ 11% RL, ↓ 13% SDW,
↓ 20% RDW, ↓ 4% SpDW, ↓ 3% GY, ↓ 11% GPr
↑ 5% SOD
↓ 20% CAT
↓ 25% GR
↑ 103% ↑ 77%
125 mM NaCl + FAB10 ↓ 17% SL, ↓ 22% RL, ↓ 18% SDW,
↓ 43% RDW, ↓ 9% SpDW,
↓ 13% GY, ↓ 16% GPr
↑ 10% SOD
↑ 20% CAT
↑ 0% GR
↑ 180% ↑ 123%
250 mM NaCl + FAB10 ↓ 25% SL, ↓ 24% RL, ↓ 18% SDW,
↓ 51% RDW, ↓ 61% SpDW,
↓ 25% GY, ↓ 27% GPr
↑ 12% SOD
↑ 20% CAT
↑ 0% GR
↑ 237% ↑ 139%

Consortium (R. leguminosarum +
A. chroococcum)
and/or Mel.
Vicia faba seeds were treated with the consortium as peat-based inoculant and/or Mel
solution
Saline soil Control [124]
25 µM Mel + Unprimed ↑ 6% SL, ↑ 37% NL, ↑ 18% SFW,
↑ 24% SDW, ↑ 23% Y
↑ 6% Chl a
↑ 11% Chl b
↑ 9% Car
N/A N/A N/A ↑ 17% N/A
50 µM Mel + Unprimed ↑ 24% SL, ↑ 56% NL, ↑ 41% SFW,
↑ 36% SDW, ↑ 41% Y
↑ 30% Chl a
↑ 26% Chl b
↑ 18% Car
↑ 30%
100 µM Mel + Unprimed ↑ 41% SL, ↑ 93% NL, ↑ 72% SFW,
↑ 78% SDW, ↑ 58% Y
↑ 44% Chl a
↑ 80% Chl b
↑ 35% Car
↑ 39%
0 µM Mel + Primed ↑ 18% SL, ↑ 43% NL, ↑ 25% SFW,
↑ 36% SDW, ↑ 48% Y
↑ 31% Chl a
↑ 35% Chl b
↑ 28% Car
↑ 44%
25 µM Mel + Primed ↑ 30% SL, ↑ 79% NL, ↑ 49% SFW,
↑ 56% SDW, ↑ 71% Y
↑ 42% Chl a
↑ 59% Chl b
↑ 43% Car
↑ 57%
50 µM Mel + Primed ↑ 70% SL, ↑ 97% NL, ↑ 68% SFW,
↑ 68% SDW, ↑ 82% Y
↑ 56% Chl a
↑ 107% Chl b
↑ 66% Car
↑ 89%
100 µM Mel + Primed ↑ 74% SL, ↑ 118% NL, ↑ 98% SFW, ↑ 104% SDW, ↑ 96% Y ↑ 71% Chl a
↑ 118% Chl b
↑ 71% Car
↑ 110%
Non-saline soil Control
25 µM Mel + Unprimed ↑ 6% SL, ↑ 14% NL, ↑ 10% SFW,
↑ 21% SDW, ↑ 18% Y
↑ 9% Chl a
↑ 10% Chl b
↑ 4% Car
↑ 5%
50 µM Mel + Unprimed ↑ 12% SL, ↑ 35% NL, ↑ 23% SFW,
↑ 31% SDW, ↑ 29% Y
↑ 11% Chl a
↑ 24% Chl b
↑ 21% Car
↑ 11%
100 µM Mel + Unprimed ↑ 16% SL, ↑ 49% NL, ↑ 38% SFW,
↑ 69% SDW, ↑ 32% Y
↑ 16% Chl a
↑ 34% Chl b
↑ 24% Car
↑ 45%
0 µM Mel + Primed ↑ 20% SL, ↑ 28% NL, ↑ 14% SFW,
↑ 27% SDW, ↑ 17% Y
↑ 17% Chl a
↑ 24% Chl b
↑ 21% Car
↑ 36%
25 µM Mel + Primed ↑ 29% SL, ↑ 49% NL, ↑ 24% SFW,
↑ 46% SDW, ↑ 25% Y
↑ 23% Chl a
↑ 30% Chl b
↑ 30% Car
↑ 94%
50 µM Mel + Primed ↑ 45% SL, ↑ 55% NL, ↑ 33% SFW,
↑ 57% SDW, ↑ 38% Y
↑ 26% Chl a
↑ 42% Chl b
↑ 33% Car
↑ 110%
100 µM Mel + Primed ↑ 49% SL, ↑ 76% NL, ↑ 52% SFW,
↑ 87% SDW, ↑ 45% Y
↑ 30% Chl a
↑ 29% Chl b
↑ 40% Car
↑ 139%
A. aneurinilyticus ACC02,
Paenibacillus ACC06 and Consortium (ACC02+ ACC06).
French bean seeds inoculated with ACC02, ACC06 and consortium (ACC02 + ACC06)
Control [88]
0 mM NaCl + ACC02 ↑ 10% SL, ↑ 50% RL, ↑ 158% SFW,
↑ 10% SDW, ↑ 50% RFW,
↑ 21% RDW
↑ 36% Tchl N/A N/A N/A N/A ↑ 9%
0 mM NaCl + ACC06 ↑ 30% SL, ↑ 30% RL, ↑ 216% SFW,
↑ 10% SDW, ↑ 60% RFW,
↑ 14% RDW
↑ 29% Tchl ↓ 9%
0 mM NaCl + Consortium ↑ 50% SL, ↑ 70% RL, ↑ 233% SFW,
↑ 80% SDW, ↑ 90% RFW,
↑ 85% RDW
↑ 57% Tchl ↑ 27%
25 mM NaCl + Unprimed
25 mM NaCl + ACC02 ↑ 33% SL, ↑ 79% RL, ↑ 120% SFW,
↑ 300% SDW, ↑ 46% RFW,
↑ 182% RDW
↑ 28% Tchl N/A N/A N/A N/A ↓ 38%
25 mM NaCl + ACC06 ↑ 47% SL, ↑ 58% RL, ↑ 120% SFW,
↑ 350% SDW, ↑ 36% RFW,
↑ 142% RDW
↑ 35% Tchl ↓ 42%
25 mM NaCl + Consortium ↑ 60% SL, ↑ 110% RL, ↑ 255% SFW, ↑ 425% SDW, ↑ 81% RFW,
↑ 220% RDW
↑ 57% Tchl ↓ 61%

P. fluorescence,
B. pumilus, E. aurantiacum and consortium (P. fluorescence + B. pumilus + E.aurantiacum)
Wheat seeds soaked in bacterial inoculant containing single PGPR strains or the consortium of three bacterial cultures for 2 h. Saline soil ECe 13.41
Unprimed seeds [122]
Seeds primed
with P. fluorescence
Galaxy-13:
↑ 5% SL, ↑ 7% RL, ↑ 3% SFW,
↑ 2% SDW, ↑ 33% 100 GW,
↓ 13% RFW, ↓ 29% RDW
Aas-11:
↑ 11% SL, ↑ 24% RL, ↑ 48% SFW,
↑ 144% RFW, ↑ 57% SDW,
↑ 75% RDW, ↑ 23% 100 GW
N/A Galaxy-13:
↓ 30% SOD
↓ 0% POD
↑ 27% CAT
Aas-11:
↓ 57% SOD
↓ 14% POD
↑ 25% CAT
N/A Galaxy-13:
In root:
↑ 50% Na+, ↑ 40% K+,
Na+/K+ ratio ~0.19
In shoot: ↑ 28% Na+, ↑ 23% K+, Na+/K+ ratio ~3.9
Aas-11:
In root: ↓ 13% Na+, ↑ 99% K+, Na+/K+ ratio ~0.16.
In shoot: ↑ 92% Na+, ↑ 16% K+, Na+/K+ ratio ~3.4
Galaxy-13: ↓ 20%
Aas-11:
↑ 33%
N/A
Seeds primed with B. pumilus Galaxy-13:
↓ 7% SL, ↓ 18% SFW, ↓ 26% SDW,
↓ 8% RFW, ↓ 57% RDW,
↑ 67% RL, 31% 100 GW
Aas-11:
↑ 13% SL, ↑ 21% RL, ↑ 61% SFW,
↑ 678% RFW, ↑ 66% SDW,
↑ 838% RDW, ↑ 53% 100 GW
Galaxy-13:
↓ 35% SOD
↓ 5% POD
↑ 4% CAT
Aas-11:
↓ 65% SOD
↓ 38% POD
↑ 35% CAT
Galaxy-13:
In root: ↑ 0% Na+, ↑ 34% K+, Na+/K+ ratio ~0.13
In shoot: ↓ 8% Na+, ↓ 19% K+, Na+/K+ ratio ~4.3
Aas-11:
In root: ↓ 13% Na+, ↑ 195% K+, Na+/K+ ratio ~0.11.
In shoot: ↑ 59% Na+, ↓ 12% K+, Na+/K+ ratio ~3.7
Galaxy-13:
↑ 287%
Aas-11:
↑ 150%
Seeds primed with E.aurantiacum Galaxy-13:
↑ 6% SL, ↑ 47% RL, ↑ 3% SFW,
↑ 49% 100 GW, ↓ 17% RFW,
↓ 2% SDW, ↓ 28% RDW
Aas-11:
↑ 10% SL, ↑ 7% RL, ↑ 52% SFW,
↑ 511% RFW, ↑ 71% SDW,
↑ 713% RDW, ↑ 47% 100 GW
Galaxy-13:
↑ 2% SOD
↑ 48% CAT
↓ 43% POD
Aas-11:
↓ 65% SOD
↓ 57% POD
↓ 5% CAT
Galaxy-13:
In root: ↑ 33% Na+, ↑ 34% K+, Na+/K+ ratio ~0.18
In shoot: ↑ 27% Na+, ↑ 0% K+, Na+/K+ ratio ~4.77.
Aas-11:
In root: ↓ 13% Na+, ↑ 286% K+, Na+/K+ ratio ~0.08
In shoot: ↑ 40% Na+, ↑ 22% K+, Na+/K+ ratio ~2.36
Galaxy-13: ↑ 227%
Aas-11:
↑ 110%
Seeds primed with a consortium Galaxy-13:
↓ 1% SL, ↑ 73% RL, ↑ 6% SFW,
↑ 30% RFW, ↑ 7% SDW,
↑ 43% RDW, ↑ 53% 100 GW
Aas-11:
↑ 13% SL, ↑ 3% RL, ↑ 65% SFW,
↑ 556% RFW, ↑ 77% SDW,
↑ 725% RDW, ↑ 48% 100 GW
Galaxy-13:
↑ 37% SOD
↓ 32% POD
↓ 6% CAT
Aas-11:
↓ 57% SOD
↑ 24% POD
↑ 28% CAT
Galaxy-13:
In root: ↑ 0% Na+, ↑ 114% K+, Na+/K+ ratio ~0.08
In shoot: ↑ 17% Na+, ↑ 15% K+, Na+/K+ ratio ~3.8
Aas-11:
In root: ↑ 0% Na+, ↑ 173% K+, Na+/K+ ratio ~0.13
In shoot: ↑ 68% Na+, ↑ 30% K+, Na+/K+ ratio ~2.67
Galaxy-13:
↑ 327%
Aas-11:
↑ 17%

Sphingobacterium BHU-AV3.
Bacterized tomato seeds with BHU-AV3 for 24h
Control [47]
200 mM NaCl + Unprimed ↓ 52% SL, ↓ 49% RL,
↓ 54% TPDW
↓ 44% Tchl In shoot:
↑ 90% SOD
↑ 260% POD
↑ 100% PPO
In root:
↑ 83% SOD
↑ 100% POD
↑ 53% PPO
N/A In shoot: ↑ 258% Na+, ↓ 63% K+, Na+/K+ ratio ~3.6
In root: ↑ 190% Na+, ↓ 53% K+, Na+/K+ ratio ~3.5
In shoot:
↑ 153%
In root:
↑ 56%
N/A
0 mM NaCl + BHU-AV3 ↓ 11.3% SL, ↑ 16% RL,
↑ 11% TPDW
↑ 5% Tchl In shoot:
↓ 20% SOD
↓ 0% POD
↓ 25% PPO
In root:
↑ 16% SOD
↑ 6% POD
↓ 12% PPO
In shoot: ↑ 9% Na+, ↑ 9% K+, Na+/K+ ratio ~0.3
In root: ↓ 5% Na+, ↑ 8% K+, Na+/K+ ratio ~0.5
In shoot:
↓ 7%
In root:
↓ 5%
200 mM NaCl + BHU-AV3 ↓ 30% SL, ↓ 22% RL,
↓ 29% TPDW
↓ 14% Tchl In shoot:
↑ 10% SOD
↑ 1000% POD
↑ 50% PPO
In root:
↑ 117% SOD
↑ 200% POD
↑ 71% PPO
In shoot: ↑ 130% Na+, ↓ 20% K+, Na+/K+ ratio ~1
In root: ↑ 115% Na+, ↓ 24% K+, Na+/K+ ratio ~1.6
In shoot:
↑ 84%
In root:
↑ 111%
K. sacchari MSK1.
Mung bean seeds primed with MSK1
Control [36]
50 mM NaCl + Unprimed ↓ 8% SL, ↓ 8% RL, ↓ 5% SDW,
↓ 5% RDW, ↓ 15% SY, ↓ 3% GP
↓ 15% Tchl
↓ 4% Car
↑ 5% GR,
↑ 33% CAT
↑ 13% SOD
↑ 23% APX
↑ 32% In shoor: ↑ 100% Na+, ↑ 44% K+, Na+/K+ ratio ~0.46, ↓ 4% N,
↓ 19% P
↑ 63% N/A
100 mM NaCl + Unprimed ↓ 16% SL, ↓ 20% RL, ↓ 10% SDW,
↓ 15% RDW, ↓ 21% SY, ↓ 6% GP
↓ 35% Tchl
↓ 9% Car
↑ 15% GR
↑ 58% CAT
↑ 39% SOD
↑ 45% APX
↑ 47% In shoot: ↑ 200% Na+, ↑ 100% K+, Na+/K+ ratio ~0.5, ↓ 4% N,
↓ 28% P
↑ 88%
200 mM NaCl + Unprimed ↓ 24% SL, ↓ 28% RL, ↓ 21% SDW,
↓ 35% RDW, ↓ 26% SY, ↓ 8% GP
↓ 42% Tchl
↓ 19% Car
↑ 35% GR
↑ 108% CAT
↑ 52% SOD
↑ 73% APX
↑ 84% In shoot: ↑ 450% Na+, ↑ 222% K+, Na+/K+ ratio ~0.57, ↓ 12% N,
↓ 44% P
↑ 213%
400 mM NaCl + Unprimed ↓ 41% SL, ↓ 52% RL, ↓ 34% SDW,
↓ 55% RDW, ↓ 34% SY, ↓ 26% GP
↓ 62% Tchl
↓ 33% Car
↑ 64% GR
↑ 208% CAT
↑ 96% SOD
↑ 102% APX
↑ 153% In shoot: ↑ 800% Na+, ↑ 378% K+, Na+/K+ ratio ~0.63, ↓ 21% N,
↓ 59% P
↑ 350%
0 mM NaCl + MSK1 ↑ 5% SL, ↑ 12% RL, ↑ 7% SDW,
↑ 15% RDW, ↑ 9% SY, ↑ 7% GP
↑ 29% Tchl
↑ 7% Car
↓ 9% GR
↓ 33% CAT
↓ 22% SOD
↓ 9% APX
↓ 37% In shoot: ↓ 67% Na+, ↓ 22% K+, Na+/K+ ratio ~0.14, ↑ 9% N,
↑ 15% P
↓ 25%
50 mM NaCl + MSK1 ↓ 3% SL, ↑ 4% RL, ↓ 2% SDW,
↑ 3% RDW, ↑ 10% SY, ↑ 2% GP
↓ 3% Tchl
↓ 0% Car
↑ 2% GR
↑ 8% CAT
↑ 9% SOD
↑ 9% APX
↑ 11% In shoot: ↑ 33% Na+, ↑ 22% K+, Na+/K+ ratio ~0.36, ↓ 1% N,
↓ 41% P
↑ 30%
100 mM NaCl + MSK1 ↓ 8% SL, ↓ 12% RL, ↓ 7% SDW,
↓ 10% RDW, ↓ 19% SY, ↓ 5% GP
↓ 31% Tchl
↓ 8% Car
↑ 11% GR
↑ 50% CAT
↑ 22% SOD
↑ 32% APX
↑ 37% In shoot: ↑ 183% Na+, ↑ 89% K+, Na+/K+ ratio ~0.5, ↓ 4% N,
↓ 22% P
↑ 75%
200 mM NaCl + MSK1 ↓ 22% SL, ↓ 16% RL, ↓ 17% SDW,
↓ 25% RDW, ↓ 24% SY, ↓ 7% GP
↓ 35% Tchl
↓ 13% Car
↑ 33% GR
↑ 100% CAT
↑ 48% SOD
↑ 64% APX
↑ 79% In shoot: ↑ 433% Na+, ↑ 211% K+, Na+/K+ ratio ~0.57, ↓ 9% N,
↓ 37% P
↑ 200%
400 mM NaCl + MSK1 ↓ 35% SL, ↓ 24% RL, ↓ 32% SDW,
↓ 48% RDW, ↓ 32% SY, ↓ 25% GP
↓ 54% Tchl
↓ 27% Car
↑ 60% GR
↑ 192% CAT
↑ 91% SOD
↑ 91% APX
↑ 137% In shoot: ↑ 783% Na+, ↑ 367% K+, Na+/K+ ratio ~0.63, ↓ 19% N,
↓ 57% P
↑ 325%

Note: All calculations in the Table 2 represent the comparisons between the treated plants and the control plants (non-stress conditions and un-priming) except that the control plants in the study of Nawaz et al. (2020) [122] were cultivated in the saline soil ECe ~13.41. The up arrowhead (↑) indicates an increase in a tested parameter as compared to the control. The down arrowhead (↓) displays a reduction in a tested parameter relative to the control. Abbreviation in the Table 2: A. calcoaceticus, Acinetobacter calcoaceticus; A. aneurinilyticus, Aneurinibacillus aneurinilyticus; A. chroococcum, Azotobacter chroococcum; AEs, Antioxidant enzymes; APX, Ascorbate peroxidase; B. pumilus, Bacillus pumilus; Car, Carotenoids; CAT, Catalase; Chl, Chlorophyll; E. aurantiacum, Exiguobacterium aurantiacum; GP, Germination percentage; GPr, Grain protein; GR, Glutathione reductase; GW, Grain weight; GY, Grain yield; K. sacchari, Kosakonia sacchari; MDA, Malondialdehyde; Mel, Melatonin; N/A, Not available; NL, Number of leaves per plant; P. fluorescence, Pseudomonas fluorescence; PhoPs, Photosynthetic pigments; POD, Peroxidase; P. putida, Pseudomonas putida; PPO, Polyphenol oxidase; RDW, Root dry weight; RFW, Root fresh weight; R. leguminosarum, Rhizobium leguminosarum; RL, Root length; SDW, Shoot dry weight; SFW, Shoot fresh weight; SL, Shoot length; S. maltophilia, Stenotrophomonas maltophilia; SOD, Superoxide dismutase; SpDW, Spike dry weight; SVI, Seedling vigor index; SY, Seed yield; Tchl, Total chlorophyll; TPDW, Total plant dry weight; Y, Yield.