Abstract
• Salt stress significantly decreased maize seed germination and seedling growth. • Stomatal size and K + /Na + ratios were significantly decreased under salt stress. • Under salt stress, MDA and H 2 O 2 increased, whereas antioxidant enzymes decreased. • Proline and glycine-betaine increased germination, stomatal size, and K + /Na + ratios. • Proline and glycine-betaine reduced salt-induced H 2 O 2 and MDA by boosting antioxidants. Osmolytes proline (Pro) and glycine betaine (GB) have been reported to impart tolerance against salinity stress in many plants. However, there is no report available on the combined application of Pro and GB to mitigate salinity-induced growth inhibition in maize. Consequently, the goal of the current study is to assess Pro's and GB's potential as priming and exogenous agents in maize under salt stress. Therefore, the present study was conducted using a petri dish and hydroponic pot experiment to evaluate the morpho-physiological and biochemical characteristics of maize plants subjected to salt stress, with the addition of Pro and GB. The seeds of maize were germinated under 20 mM of each Pro or GB, with or without 120 mM salinity stress. The findings demonstrated that salt stress lessened the final germination percentage (FGP) (52.9 %), photosynthetic pigments (40.0 %), relative water content (RWC) (20.2 %), stomatal size (59.3 %), and leaf and root K + /Na + ratios of maize seedlings compared to control. In addition, compared to the control, higher levels of malondialdehyde (MDA) (24.2 %) and hydrogen peroxide (H 2 O 2 ) (25.7 %) were observed, whereas lower levels of catalase (CAT) (43.8 %) and peroxidase (POX) (30.4 %) were noted. The priming with Pro, GB, Pro+GB significantly increased FGP, germination index (GI), shoot-root biomass, seed vigor index (SVI), and reduced mean germination time (MGT) under salt stress. Foliar application of Pro, GB, or Pro+GB resulted in a significant increment in chlorophyll contents, RWC, K + absorption, and stomatal size under salt stress. Furthermore, supplementing with Pro, GB, or Pro+GB reduced the accumulation of H 2 O 2 (24.0, 23.3, and 31.1 %, respectively) and MDA (22.8, 17.2, and 32.1 %, respectively) caused by salt and augmented the levels of CAT (33.3, 22.8, and 45.2 %, respectively) and POX (36.4, 23.5, and 47.2 %, respectively) in the leaves. Taken together, the current study's findings indicate that combining Pro and GB is one of the most effective techniques for improving the salinity-tolerant seed germination and seedling traits of maize plants. Consequently, this study recommends that Pro and GB can be used as seed priming and exogenous agents to help maize grow faster in salt-stress situations.
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Claims 20
Claim · model-written summary
Salt stress reduced maize final germination percentage (FGP) by 52.9 % compared with control.
- Treatment
- salt stresse0
- Outcome
- final germination percentage (FGP)e0
- Direction
- lessenede0
- Value
- 52.9 %e0
- Comparator
- controle0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The findings demonstrated that salt stress lessened the final germination percentage (FGP) (52.9 %), photosynthetic pigments (40.0 %), relative water content (RWC) (20.2 %), stomatal size (59.3 %), and leaf and root K + /Na + ratios of maize seedlings compared to control.”
Claim · model-written summary
Salt stress reduced maize photosynthetic pigments by 40.0 % compared with control.
- Treatment
- salt stresse0
- Outcome
- photosynthetic pigmentse0
- Direction
- lessenede0
- Value
- 40.0 %e0
- Comparator
- controle0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The findings demonstrated that salt stress lessened the final germination percentage (FGP) (52.9 %), photosynthetic pigments (40.0 %), relative water content (RWC) (20.2 %), stomatal size (59.3 %), and leaf and root K + /Na + ratios of maize seedlings compared to control.”
Claim · model-written summary
Salt stress reduced maize relative water content (RWC) by 20.2 % compared with control.
- Treatment
- salt stresse0
- Outcome
- relative water content (RWC)e0
- Direction
- lessenede0
- Value
- 20.2 %e0
- Comparator
- controle0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The findings demonstrated that salt stress lessened the final germination percentage (FGP) (52.9 %), photosynthetic pigments (40.0 %), relative water content (RWC) (20.2 %), stomatal size (59.3 %), and leaf and root K + /Na + ratios of maize seedlings compared to control.”
Claim · model-written summary
Salt stress reduced maize stomatal size by 59.3 % compared with control.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The findings demonstrated that salt stress lessened the final germination percentage (FGP) (52.9 %), photosynthetic pigments (40.0 %), relative water content (RWC) (20.2 %), stomatal size (59.3 %), and leaf and root K + /Na + ratios of maize seedlings compared to control.”
Claim · model-written summary
Salt stress reduced leaf and root K + /Na + ratios in maize seedlings.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“The findings demonstrated that salt stress lessened the final germination percentage (FGP) (52.9 %), photosynthetic pigments (40.0 %), relative water content (RWC) (20.2 %), stomatal size (59.3 %), and leaf and root K + /Na + ratios of maize seedlings compared to control.”
Claim · model-written summary
Under salt stress maize had higher malondialdehyde (MDA) by 24.2 % than control.
Not reportedTaxonTreatment
Verbatim
e0“In addition, compared to the control, higher levels of malondialdehyde (MDA) (24.2 %) and hydrogen peroxide (H 2 O 2 ) (25.7 %) were observed, whereas lower levels of catalase (CAT) (43.8 %) and peroxidase (POX) (30.4 %) were noted.”
t0“maize”
Claim · model-written summary
Under salt stress maize had higher hydrogen peroxide (H 2 O 2 ) by 25.7 % than control.
Not reportedTaxonTreatment
Verbatim
e0“In addition, compared to the control, higher levels of malondialdehyde (MDA) (24.2 %) and hydrogen peroxide (H 2 O 2 ) (25.7 %) were observed, whereas lower levels of catalase (CAT) (43.8 %) and peroxidase (POX) (30.4 %) were noted.”
t0“maize”
Claim · model-written summary
Under salt stress maize had lower catalase (CAT) by 43.8 % than control.
Not reportedTaxonTreatment
Verbatim
e0“In addition, compared to the control, higher levels of malondialdehyde (MDA) (24.2 %) and hydrogen peroxide (H 2 O 2 ) (25.7 %) were observed, whereas lower levels of catalase (CAT) (43.8 %) and peroxidase (POX) (30.4 %) were noted.”
t0“maize”
Claim · model-written summary
Under salt stress maize had lower peroxidase (POX) by 30.4 % than control.
Not reportedTaxonTreatment
Verbatim
e0“In addition, compared to the control, higher levels of malondialdehyde (MDA) (24.2 %) and hydrogen peroxide (H 2 O 2 ) (25.7 %) were observed, whereas lower levels of catalase (CAT) (43.8 %) and peroxidase (POX) (30.4 %) were noted.”
t0“maize”
Claim · model-written summary
Pro priming increased germination and biomass indices and reduced germination time under salt stress.
- Treatment
- Proe0
- Outcome
- FGP, germination index (GI), shoot-root biomass, seed vigor index (SVI)e0
- Direction
- increasede0
- Qualifier
- under salt stresse0
Not reportedTaxonComparator
Verbatim
e0“The priming with Pro, GB, Pro+GB significantly increased FGP, germination index (GI), shoot-root biomass, seed vigor index (SVI), and reduced mean germination time (MGT) under salt stress.”
t0“maize”
Claim · model-written summary
GB priming increased germination and biomass indices and reduced germination time under salt stress.
- Treatment
- GBe0
- Outcome
- FGP, germination index (GI), shoot-root biomass, seed vigor index (SVI)e0
- Direction
- increasede0
- Qualifier
- under salt stresse0
Not reportedTaxonComparator
Verbatim
e0“The priming with Pro, GB, Pro+GB significantly increased FGP, germination index (GI), shoot-root biomass, seed vigor index (SVI), and reduced mean germination time (MGT) under salt stress.”
t0“maize”
Claim · model-written summary
Pro+GB priming increased germination and biomass indices and reduced germination time under salt stress.
- Treatment
- Pro+GBe0
- Outcome
- FGP, germination index (GI), shoot-root biomass, seed vigor index (SVI)e0
- Direction
- increasede0
- Qualifier
- under salt stresse0
Not reportedTaxonComparator
Verbatim
e0“The priming with Pro, GB, Pro+GB significantly increased FGP, germination index (GI), shoot-root biomass, seed vigor index (SVI), and reduced mean germination time (MGT) under salt stress.”
t0“maize”
Claim · model-written summary
Pro foliar application increased chlorophyll, water content, potassium absorption, and stomatal size under salt stress.
- Treatment
- Proe0
- Outcome
- chlorophyll contents, RWC, K + absorption, and stomatal sizee0
- Direction
- incremente0
- Qualifier
- under salt stresse0
Not reportedTaxonComparator
Verbatim
e0“Foliar application of Pro, GB, or Pro+GB resulted in a significant increment in chlorophyll contents, RWC, K + absorption, and stomatal size under salt stress.”
t0“maize”
Claim · model-written summary
GB foliar application increased chlorophyll, water content, potassium absorption, and stomatal size under salt stress.
- Treatment
- GBe0
- Outcome
- chlorophyll contents, RWC, K + absorption, and stomatal sizee0
- Direction
- incremente0
- Qualifier
- under salt stresse0
Not reportedTaxonComparator
Verbatim
e0“Foliar application of Pro, GB, or Pro+GB resulted in a significant increment in chlorophyll contents, RWC, K + absorption, and stomatal size under salt stress.”
t0“maize”
Claim · model-written summary
Pro+GB foliar application increased chlorophyll, water content, potassium absorption, and stomatal size under salt stress.
- Treatment
- Pro+GBe0
- Outcome
- chlorophyll contents, RWC, K + absorption, and stomatal sizee0
- Direction
- incremente0
- Qualifier
- under salt stresse0
Not reportedTaxonComparator
Verbatim
e0“Foliar application of Pro, GB, or Pro+GB resulted in a significant increment in chlorophyll contents, RWC, K + absorption, and stomatal size under salt stress.”
t0“maize”
Claim · model-written summary
Pro reduced salt-induced H 2 O 2 accumulation in maize leaves by 24.0%.
Not reportedTaxonComparator
Verbatim
e0“Furthermore, supplementing with Pro, GB, or Pro+GB reduced the accumulation of H 2 O 2 (24.0, 23.3, and 31.1 %, respectively) and MDA (22.8, 17.2, and 32.1 %, respectively) caused by salt and augmented the levels of CAT (33.3, 22.8, and 45.2 %, respectively) and POX (36.4, 23.5, and 47.2 %, respectively) in the leaves.”
t0“maize”
Claim · model-written summary
GB reduced salt-induced H 2 O 2 accumulation in maize leaves by 23.3%.
Not reportedTaxonComparator
Verbatim
e0“Furthermore, supplementing with Pro, GB, or Pro+GB reduced the accumulation of H 2 O 2 (24.0, 23.3, and 31.1 %, respectively) and MDA (22.8, 17.2, and 32.1 %, respectively) caused by salt and augmented the levels of CAT (33.3, 22.8, and 45.2 %, respectively) and POX (36.4, 23.5, and 47.2 %, respectively) in the leaves.”
t0“maize”
Claim · model-written summary
Pro+GB reduced salt-induced H 2 O 2 accumulation in maize leaves by 31.1 %%.
- Organ / stage
- leavese0
- Treatment
- Pro+GBe0
- Outcome
- accumulation of H 2 O 2e0
- Direction
- reducede0
- Value
- 31.1 %e0
Not reportedTaxonComparator
Verbatim
e0“Furthermore, supplementing with Pro, GB, or Pro+GB reduced the accumulation of H 2 O 2 (24.0, 23.3, and 31.1 %, respectively) and MDA (22.8, 17.2, and 32.1 %, respectively) caused by salt and augmented the levels of CAT (33.3, 22.8, and 45.2 %, respectively) and POX (36.4, 23.5, and 47.2 %, respectively) in the leaves.”
t0“maize”
Claim · model-written summary
Pro reduced salt-induced MDA accumulation in maize leaves by 22.8%.
Not reportedTaxonComparator
Verbatim
e0“Furthermore, supplementing with Pro, GB, or Pro+GB reduced the accumulation of H 2 O 2 (24.0, 23.3, and 31.1 %, respectively) and MDA (22.8, 17.2, and 32.1 %, respectively) caused by salt and augmented the levels of CAT (33.3, 22.8, and 45.2 %, respectively) and POX (36.4, 23.5, and 47.2 %, respectively) in the leaves.”
t0“maize”
Claim · model-written summary
GB reduced salt-induced MDA accumulation in maize leaves by 17.2%.
Not reportedTaxonComparator
Verbatim
e0“Furthermore, supplementing with Pro, GB, or Pro+GB reduced the accumulation of H 2 O 2 (24.0, 23.3, and 31.1 %, respectively) and MDA (22.8, 17.2, and 32.1 %, respectively) caused by salt and augmented the levels of CAT (33.3, 22.8, and 45.2 %, respectively) and POX (36.4, 23.5, and 47.2 %, respectively) in the leaves.”
t0“maize”
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