Abstract
Soil contamination by heavy metals, particularly lead (Pb), which is considered the second most toxic metal, poses serious risks to plants and humans due to its accumulation from various anthropogenic activities. Strigolactones (SLs) are a novel class of terpenoid lactones that play a vital role in regulating plant growth and development, particularly under stress conditions. This study aimed to investigate the impact of exogenous SL applications on plant growth and various physiological, biochemical, and molecular parameters in lettuce subjected to Pb stress. Pb stress harmed plant growth, whereas SL treatments improved growth parameters under both control and Pb stress conditions. While Pb stress increased the electrical conductivity (EC), malondialdehyde (MDA) and hydrogen peroxide (H2O2) content, SL applications caused a decrease in these parameters. Pb stress negatively affected chlorophyll content, whereas SL applications reduced negative effect. Pb caused an increase in superoxide dismutase (SOD), catalase (CAT), peroxidase (POD) ascorbate peroxidase (APX), glutathione reductase (GR), glutathione s-transferase (GST) activities and Glucose-6-Phosphate Dehydrogenase (G6PD), 6-Phosphogluconate Dehydrogenase (6GPD). SL treatments significantly increased the activity of antioxidant enzymes in both control and Pb stress conditions. However, Pb stressed plants had lower nitrate reductase activity (NRA) than the control plants while SL treatments increased NRA compared to the non-treatments. Pb stress significantly reduced the uptake of essential nutrients in lettuce seedlings, whereas exogenous SL applications improved nutrient accumulation, particularly under Pb-stressed conditions. Additionally, mRNA expression profiles of nine stress-related genes in different tissues of lettuce were determined. Only Pb stress significantly decreased the expression of genes, particularly LsCCD8 and LsD14, in both tissues. The combined Pb and SL treatment significantly increased the expression of LsMAX2 in both tissues. These results suggest that exogenous SL applications can be an effective strategy to mitigate Pb-induced stress in lettuce by enhancing plant tolerance at physiological, biochemical, and molecular levels.
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Claims 17
Claim · model-written summary
Pb stress harmed lettuce plant growth.
- Taxon
- lettuce
Lactuca sativat0 - Treatment
- Pb stresse1
- Outcome
- plant growthe1
- Direction
- harmede1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tolerance to lead stress on lettuce (Lactuca sativa L.) plants”
e1“Pb stress harmed plant growth, whereas SL treatments improved growth parameters under both control and Pb stress conditions”
Claim · model-written summary
SL treatments improved lettuce growth parameters under both control and Pb stress conditions.
- Taxon
- lettuce
Lactuca sativat0 - Treatment
- SL treatmentse1
- Outcome
- growth parameterse1
- Direction
- improvede1
- Qualifier
- under both control and Pb stress conditionse1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tolerance to lead stress on lettuce (Lactuca sativa L.) plants”
e1“Pb stress harmed plant growth, whereas SL treatments improved growth parameters under both control and Pb stress conditions”
Claim · model-written summary
Pb stress increased electrical conductivity, malondialdehyde and hydrogen peroxide content in lettuce.
- Taxon
- lettuce
Lactuca sativat0 - Treatment
- Pb stresse2
- Outcome
- electrical conductivity (EC), malondialdehyde (MDA) and hydrogen peroxide (H2O2) contente2
- Direction
- increasede2
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tolerance to lead stress on lettuce (Lactuca sativa L.) plants”
e2“While Pb stress increased the electrical conductivity (EC), malondialdehyde (MDA) and hydrogen peroxide (H2O2) content, SL applications caused a decrease in these parameters”
Claim · model-written summary
SL applications decreased electrical conductivity, malondialdehyde and hydrogen peroxide content in lettuce.
- Taxon
- lettuce
Lactuca sativat0 - Treatment
- SL applicationse2
- Outcome
- electrical conductivity (EC), malondialdehyde (MDA) and hydrogen peroxide (H2O2) contente2
- Direction
- decreasee2
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tolerance to lead stress on lettuce (Lactuca sativa L.) plants”
e2“While Pb stress increased the electrical conductivity (EC), malondialdehyde (MDA) and hydrogen peroxide (H2O2) content, SL applications caused a decrease in these parameters”
Claim · model-written summary
Pb stress negatively affected chlorophyll content in lettuce.
- Taxon
- lettuce
Lactuca sativat0 - Treatment
- Pb stresse3
- Outcome
- chlorophyll contente3
- Direction
- negatively affectede3
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tolerance to lead stress on lettuce (Lactuca sativa L.) plants”
e3“Pb stress negatively affected chlorophyll content, whereas SL applications reduced negative effect”
Claim · model-written summary
SL applications reduced the negative effect of Pb stress on lettuce chlorophyll content.
- Taxon
- lettuce
Lactuca sativat0 - Treatment
- SL applicationse3
- Outcome
- chlorophyll contente3
- Direction
- reducede3
- Qualifier
- negative effecte3
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tolerance to lead stress on lettuce (Lactuca sativa L.) plants”
e3“Pb stress negatively affected chlorophyll content, whereas SL applications reduced negative effect”
Claim · model-written summary
Pb increased activities of superoxide dismutase, catalase and peroxidase in lettuce.
- Taxon
- lettuce
Lactuca sativat0 - Treatment
- Pbe5
- Gene
- superoxide dismutase (SOD), catalase (CAT), peroxidase (POD)e5
- Outcome
- activitiese5
- Direction
- increasee5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tolerance to lead stress on lettuce (Lactuca sativa L.) plants”
e5“Pb caused an increase in superoxide dismutase (SOD), catalase (CAT), peroxidase (POD) ascorbate peroxidase (APX), glutathione reductase (GR), glutathione s-transferase (GST) activities and Glucose-6-Phosphate Dehydrogenase (G6PD), 6-Phosphogluconate Dehydrogenase (6GPD)”
Claim · model-written summary
Pb increased activities of ascorbate peroxidase, glutathione reductase and glutathione s-transferase in lettuce.
- Taxon
- lettuce
Lactuca sativat0 - Treatment
- Pbe5
- Gene
- ascorbate peroxidase (APX), glutathione reductase (GR), glutathione s-transferase (GST)e5
- Outcome
- activitiese5
- Direction
- increasee5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tolerance to lead stress on lettuce (Lactuca sativa L.) plants”
e5“Pb caused an increase in superoxide dismutase (SOD), catalase (CAT), peroxidase (POD) ascorbate peroxidase (APX), glutathione reductase (GR), glutathione s-transferase (GST) activities and Glucose-6-Phosphate Dehydrogenase (G6PD), 6-Phosphogluconate Dehydrogenase (6GPD)”
Claim · model-written summary
Pb increased activities of glucose-6-phosphate dehydrogenase and 6-phosphogluconate dehydrogenase in lettuce.
- Taxon
- lettuce
Lactuca sativat0 - Treatment
- Pbe5
- Gene
- Glucose-6-Phosphate Dehydrogenase (G6PD), 6-Phosphogluconate Dehydrogenase (6GPD)e5
- Outcome
- activitiese5
- Direction
- increasee5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tolerance to lead stress on lettuce (Lactuca sativa L.) plants”
e5“Pb caused an increase in superoxide dismutase (SOD), catalase (CAT), peroxidase (POD) ascorbate peroxidase (APX), glutathione reductase (GR), glutathione s-transferase (GST) activities and Glucose-6-Phosphate Dehydrogenase (G6PD), 6-Phosphogluconate Dehydrogenase (6GPD)”
Claim · model-written summary
SL treatments significantly increased antioxidant enzyme activity in both control and Pb-stressed lettuce.
- Taxon
- lettuce
Lactuca sativat0 - Treatment
- SL treatmentse6
- Outcome
- activity of antioxidant enzymese6
- Direction
- increasede6
- Qualifier
- in both control and Pb stress conditionse6
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tolerance to lead stress on lettuce (Lactuca sativa L.) plants”
e6“SL treatments significantly increased the activity of antioxidant enzymes in both control and Pb stress conditions”
Claim · model-written summary
Pb-stressed lettuce plants had lower nitrate reductase activity than control plants.
- Taxon
- lettuce
Lactuca sativat0 - Treatment
- Pb stressed plantse7
- Outcome
- nitrate reductase activity (NRA)e7
- Direction
- lowere7
- Comparator
- the control plantse7
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tolerance to lead stress on lettuce (Lactuca sativa L.) plants”
e7“Pb stressed plants had lower nitrate reductase activity (NRA) than the control plants while SL treatments increased NRA compared to the non-treatments”
Claim · model-written summary
SL treatments increased nitrate reductase activity compared with non-treated lettuce.
- Taxon
- lettuce
Lactuca sativat0 - Treatment
- SL treatmentse7
- Outcome
- NRAe7
- Direction
- increasede7
- Comparator
- the non-treatmentse7
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tolerance to lead stress on lettuce (Lactuca sativa L.) plants”
e7“Pb stressed plants had lower nitrate reductase activity (NRA) than the control plants while SL treatments increased NRA compared to the non-treatments”
Claim · model-written summary
Pb stress significantly reduced nutrient uptake in lettuce seedlings.
- Taxon
- lettuce
Lactuca sativat0 - Organ / stage
- seedlingse0
- Treatment
- Pb stresse0
- Outcome
- uptake of essential nutrientse0
- Direction
- reducede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tolerance to lead stress on lettuce (Lactuca sativa L.) plants”
e0“Pb stress significantly reduced the uptake of essential nutrients in lettuce seedlings”
Claim · model-written summary
Exogenous SL applications improved nutrient accumulation, particularly under Pb stress.
- Taxon
- lettuce
Lactuca sativat0 - Treatment
- exogenous SL applicationse1
- Outcome
- nutrient accumulatione1
- Direction
- improvede1
- Qualifier
- particularly under Pb-stressed conditionse1
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tolerance to lead stress on lettuce (Lactuca sativa L.) plants”
e1“whereas exogenous SL applications improved nutrient accumulation, particularly under Pb-stressed conditions”
Claim · model-written summary
Pb stress significantly decreased expression of LsCCD8 and LsD14 in both lettuce tissues.
- Taxon
- lettuce
Lactuca sativat0 - Organ / stage
- both tissuese2
- Treatment
- Pb stresse2
- Gene
- LsCCD8 and LsD14e2
- Outcome
- expression of genese2
- Direction
- decreasede2
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tolerance to lead stress on lettuce (Lactuca sativa L.) plants”
e2“Only Pb stress significantly decreased the expression of genes, particularly LsCCD8 and LsD14, in both tissues”
Claim · model-written summary
Combined Pb and SL treatment significantly increased LsMAX2 expression in both lettuce tissues.
- Taxon
- lettuce
Lactuca sativat0 - Organ / stage
- both tissuese3
- Treatment
- combined Pb and SL treatmente3
- Gene
- LsMAX2e3
- Outcome
- expressione3
- Direction
- increasede3
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
t0“tolerance to lead stress on lettuce (Lactuca sativa L.) plants”
e3“The combined Pb and SL treatment significantly increased the expression of LsMAX2 in both tissues”
Claim · model-written summary
Exogenous SL applications are an effective strategy to mitigate Pb-induced stress in lettuce.
- Taxon
- lettuce
Lactuca sativae4 - Outcome
- mitigate Pb-induced stresse4
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e4“exogenous SL applications can be an effective strategy to mitigate Pb-induced stress in lettuce”
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