Abstract
Cadmium (Cd) and the freeze-thaw cycle (FTC) are serious concerns worldwide, and plants at high altitudes are particularly vulnerable to these stresses. This study aims to evaluate the role of Betaine (B) in increasing Secale cereale L. tolerance to FTC and Cd stress. In this study, Secale cereale L . was chosen as a model to investigate the effect of B application on seedlings exposed to FTC and Cd stress. Photosynthetic and non-photosynthetic parameters were measured. An integrated biomarker response (IBR) model and molecular docking were also used for a comprehensive assessment. The results showed that Cd and FTC significantly elevated malondialdehyde (MDA), hydrogen peroxide (H 2 O 2 ), ascorbate peroxidase (APX), glutamate decarboxylase (GAD), nonprotein thiol (NPT), phytochelatins (PCs), and reduced glutathione (GSH) activities. Combined stress resulted in significant increases of 9.42%, 10.7%, and 4.6% in GAD, H2O2, and NPT, respectively, and increases of 2.5% and 16.1% in PCs and MDA activities, respectively. FTC and Cd adversely affected photosynthetic parameters, causing decreases of 19.4%, 58.2%, and 56.3% in water use efficiency (WUE), transpiration rate (Tr), and photosynthetic rate (Pn), respectively, while intercellular CO₂ concentration (Ci) increased by 61.7%. With exogenous B application, antioxidant enzyme activities were elevated, while MDA, H 2 O 2 , and relative conductivity (RC) were reduced. These results suggest that B is a capable agent for mitigating abiotic stress in cold-exposed and Cd-contaminated environments. The outcomes provide essential insights for future research on ecology, environmental pollution, and management strategies to enhance plant resilience. • Freeze–thaw cycles and cadmium pose combined ecological stress to rye seedlings. • Dual stress elevates oxidative damage and disrupts photosynthetic performance. • Betaine mitigates Cd–FTC toxicity by strengthening antioxidant defenses.
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Claims 13
Claim · model-written summary
Cd and FTC significantly elevated ascorbate peroxidase (APX) activity in rye seedlings.
- Taxon
- Secale cereale
Secale cerealet0 - Treatment
- Cd and FTCe1
- Outcome
- ascorbate peroxidase (APX)e1
- Direction
- significantly elevatede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Interactive effects of cadmium stress and freeze–thaw cycles on Secale cereale L.: Protective role of exogenous betaine”
e1“The results showed that Cd and FTC significantly elevated malondialdehyde (MDA), hydrogen peroxide (H 2 O 2 ), ascorbate peroxidase (APX), glutamate decarboxylase (GAD), nonprotein thiol (NPT), phytochelatins (PCs), and reduced glutathione (GSH) activities”
Claim · model-written summary
Cd and FTC significantly elevated reduced glutathione (GSH) activity in rye seedlings.
- Taxon
- Secale cereale
Secale cerealet0 - Treatment
- Cd and FTCe1
- Outcome
- reduced glutathione (GSH)e1
- Direction
- significantly elevatede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Interactive effects of cadmium stress and freeze–thaw cycles on Secale cereale L.: Protective role of exogenous betaine”
e1“The results showed that Cd and FTC significantly elevated malondialdehyde (MDA), hydrogen peroxide (H 2 O 2 ), ascorbate peroxidase (APX), glutamate decarboxylase (GAD), nonprotein thiol (NPT), phytochelatins (PCs), and reduced glutathione (GSH) activities”
Claim · model-written summary
Combined stress increased glutamate decarboxylase (GAD) by 9.42%.
- Taxon
- Secale cereale
Secale cerealet0 - Treatment
- Combined stresse1
- Outcome
- GADe1
- Direction
- increasese1
- Value
- 9.42%e1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Interactive effects of cadmium stress and freeze–thaw cycles on Secale cereale L.: Protective role of exogenous betaine”
e1“Combined stress resulted in significant increases of 9.42%, 10.7%, and 4.6% in GAD, H2O2, and NPT, respectively”
Claim · model-written summary
Combined stress increased hydrogen peroxide (H2O2) by 10.7%.
- Taxon
- Secale cereale
Secale cerealet0 - Treatment
- Combined stresse1
- Outcome
- H2O2e1
- Direction
- increasese1
- Value
- 10.7%e1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Interactive effects of cadmium stress and freeze–thaw cycles on Secale cereale L.: Protective role of exogenous betaine”
e1“Combined stress resulted in significant increases of 9.42%, 10.7%, and 4.6% in GAD, H2O2, and NPT, respectively”
Claim · model-written summary
Combined stress increased nonprotein thiol (NPT) by 4.6%.
- Taxon
- Secale cereale
Secale cerealet0 - Treatment
- Combined stresse1
- Outcome
- NPTe1
- Direction
- increasese1
- Value
- 4.6%e1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Interactive effects of cadmium stress and freeze–thaw cycles on Secale cereale L.: Protective role of exogenous betaine”
e1“Combined stress resulted in significant increases of 9.42%, 10.7%, and 4.6% in GAD, H2O2, and NPT, respectively”
Claim · model-written summary
Combined stress increased phytochelatins (PCs) by 2.5%.
- Taxon
- Secale cereale
Secale cerealet0 - Treatment
- Combined stresse1
- Outcome
- PCse2
- Direction
- increasese2
- Value
- 2.5%e2
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Interactive effects of cadmium stress and freeze–thaw cycles on Secale cereale L.: Protective role of exogenous betaine”
e1“Combined stress resulted in significant increases of 9.42%, 10.7%, and 4.6% in GAD, H2O2, and NPT, respectively”
e2“increases of 2.5% and 16.1% in PCs and MDA activities, respectively”
Claim · model-written summary
Combined stress increased malondialdehyde (MDA) by 16.1%.
- Taxon
- Secale cereale
Secale cerealet0 - Treatment
- Combined stresse1
- Outcome
- MDAe2
- Direction
- increasese2
- Value
- 16.1%e2
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Interactive effects of cadmium stress and freeze–thaw cycles on Secale cereale L.: Protective role of exogenous betaine”
e1“Combined stress resulted in significant increases of 9.42%, 10.7%, and 4.6% in GAD, H2O2, and NPT, respectively”
e2“increases of 2.5% and 16.1% in PCs and MDA activities, respectively”
Claim · model-written summary
FTC and Cd decreased water use efficiency (WUE) by 19.4%.
- Taxon
- Secale cereale
Secale cerealet0 - Treatment
- FTC and Cde1
- Outcome
- water use efficiency (WUE)e1
- Direction
- decreasese1
- Value
- 19.4%e1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Interactive effects of cadmium stress and freeze–thaw cycles on Secale cereale L.: Protective role of exogenous betaine”
e1“FTC and Cd adversely affected photosynthetic parameters, causing decreases of 19.4%, 58.2%, and 56.3% in water use efficiency (WUE), transpiration rate (Tr), and photosynthetic rate (Pn), respectively”
Claim · model-written summary
FTC and Cd decreased transpiration rate (Tr) by 58.2%.
- Taxon
- Secale cereale
Secale cerealet0 - Treatment
- FTC and Cde1
- Outcome
- transpiration rate (Tr)e1
- Direction
- decreasese1
- Value
- 58.2%e1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Interactive effects of cadmium stress and freeze–thaw cycles on Secale cereale L.: Protective role of exogenous betaine”
e1“FTC and Cd adversely affected photosynthetic parameters, causing decreases of 19.4%, 58.2%, and 56.3% in water use efficiency (WUE), transpiration rate (Tr), and photosynthetic rate (Pn), respectively”
Claim · model-written summary
FTC and Cd decreased photosynthetic rate (Pn) by 56.3%.
- Taxon
- Secale cereale
Secale cerealet0 - Treatment
- FTC and Cde1
- Outcome
- photosynthetic rate (Pn)e1
- Direction
- decreasese1
- Value
- 56.3%e1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Interactive effects of cadmium stress and freeze–thaw cycles on Secale cereale L.: Protective role of exogenous betaine”
e1“FTC and Cd adversely affected photosynthetic parameters, causing decreases of 19.4%, 58.2%, and 56.3% in water use efficiency (WUE), transpiration rate (Tr), and photosynthetic rate (Pn), respectively”
Claim · model-written summary
FTC and Cd increased intercellular CO2 concentration (Ci) by 61.7%.
- Taxon
- Secale cereale
Secale cerealet0 - Treatment
- FTC and Cde1
- Outcome
- intercellular CO₂ concentration (Ci)e2
- Direction
- increasede2
- Value
- 61.7%e2
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Interactive effects of cadmium stress and freeze–thaw cycles on Secale cereale L.: Protective role of exogenous betaine”
e1“FTC and Cd adversely affected photosynthetic parameters, causing decreases of 19.4%, 58.2%, and 56.3% in water use efficiency (WUE), transpiration rate (Tr), and photosynthetic rate (Pn), respectively”
e2“intercellular CO₂ concentration (Ci) increased by 61.7%”
Claim · model-written summary
Exogenous betaine application elevated antioxidant enzyme activities in rye seedlings.
- Taxon
- Secale cereale
Secale cerealet0 - Treatment
- exogenous B applicatione1
- Outcome
- antioxidant enzyme activitiese1
- Direction
- elevatede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Interactive effects of cadmium stress and freeze–thaw cycles on Secale cereale L.: Protective role of exogenous betaine”
e1“With exogenous B application, antioxidant enzyme activities were elevated, while MDA, H 2 O 2 , and relative conductivity (RC) were reduced”
Claim · model-written summary
Exogenous betaine application reduced MDA, H2O2 and relative conductivity in rye seedlings.
- Taxon
- Secale cereale
Secale cerealet0 - Treatment
- exogenous B applicatione1
- Outcome
- MDA, H 2 O 2 , and relative conductivity (RC)e1
- Direction
- reducede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“Interactive effects of cadmium stress and freeze–thaw cycles on Secale cereale L.: Protective role of exogenous betaine”
e1“With exogenous B application, antioxidant enzyme activities were elevated, while MDA, H 2 O 2 , and relative conductivity (RC) were reduced”
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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.