Abstract
Heavy metal contamination, particularly from cadmium (Cd) and lead (Pb), poses significant risks to soil and water resources and leads to severe damage in plants. This study investigated the physiological and molecular mechanisms of the responses of tomato ( Solanum lycopersicum L.) seedlings to Cd and Pb stress by applying 50 mg/L Cd, 100 mg/L Pb, and a combination of 50 mg/L Cd + 100 mg/L Pb. The goal was to understand how these heavy metals impact the growth, antioxidant systems, and secondary metabolic pathways in tomato seedlings. The results showed that compared with the control, Cd + Pb stress significantly increased the content of soluble sugar by 37.40% and 33.46% on days 5 and 15, respectively, and the content of proline by 77.91% to 93.91% during the entire period in tomato seedlings. It also elevated electrical leakage by 110.52% on day 15, maintained the levels of malondialdehyde close to the control, enhanced the activities of superoxide dismutase by 33.32% on day 10 and 11.22% on day 15, peroxidase by 42.15% on day 10, and catalase by 90.78% on day 10. Additionally, it reduced the contents of hydrogen peroxide by 15.47% to 29.64% and the rate of formation of superoxide anions by 26.34% to 53.47% during the entire period of treatment. The transcriptomic analysis revealed a significant differential expression of the genes involved in pathways, such as phenylalanine, glutathione, arginine and proline, and nitrogen metabolism. These genes included PALs , HDCs , GGCT , ODC1 , LAPs , SMS , and SAMDC . Notably, transcription factors, such as ERF109 , ARF9 , GRF3 , GRF4 , GRF7 , and GRF9 , were also significantly regulated. The study concluded that Cd and Pb stress enhanced the osmoregulatory and antioxidant defense systems in tomato seedlings, which may contribute to their tolerance to heavy metal stress. Future research could explore the application of these findings to develop strategies to improve the resistance of plants to contamination with heavy metals.
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Claims 14
Claim · model-written summary
content of soluble sugar changed under combined Cd and Pb stress by 37.40%.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- Cd + Pb stresse2
- Outcome
- content of soluble sugare0
- Direction
- increasede0
- Value
- 37.40%e0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Cd + Pb stress significantly increased the content of soluble sugar by 37.40% and 33.46% on days 5 and 15, respectively”
t0“tomato seedlings”
e2“Cd + Pb stress”
Claim · model-written summary
content of soluble sugar changed under combined Cd and Pb stress by 33.46%.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- Cd + Pb stresse2
- Outcome
- content of soluble sugare0
- Direction
- increasede0
- Value
- 33.46%e0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Cd + Pb stress significantly increased the content of soluble sugar by 37.40% and 33.46% on days 5 and 15, respectively”
t0“tomato seedlings”
e2“Cd + Pb stress”
Claim · model-written summary
content of proline changed under combined Cd and Pb stress by 77.91% to 93.91%.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- Cd + Pb stresse2
- Outcome
- content of prolinee0
- Direction
- increaseinferrede0
- Value
- 77.91% to 93.91%e0
- Qualifier
- during the entire periode0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“content of proline by 77.91% to 93.91% during the entire period in tomato seedlings”
t0“tomato seedlings”
e2“Cd + Pb stress”
Claim · model-written summary
electrical leakage changed under combined Cd and Pb stress by 110.52%.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- Cd + Pb stresse2
- Outcome
- electrical leakagee0
- Direction
- elevatede0
- Value
- 110.52%e0
- Qualifier
- day 15e0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“elevated electrical leakage by 110.52% on day 15”
t0“tomato seedlings”
e2“Cd + Pb stress”
Claim · model-written summary
levels of malondialdehyde changed under combined Cd and Pb stress.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- Cd + Pb stresse2
- Outcome
- levels of malondialdehydee0
- Direction
- close to the controle0
- Comparator
- controle0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“maintained the levels of malondialdehyde close to the control”
t0“tomato seedlings”
e2“Cd + Pb stress”
Claim · model-written summary
activities of superoxide dismutase changed under combined Cd and Pb stress by 33.32%.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- Cd + Pb stresse2
- Outcome
- activities of superoxide dismutasee0
- Direction
- enhancede0
- Value
- 33.32%e0
- Qualifier
- day 10e0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“enhanced the activities of superoxide dismutase by 33.32% on day 10 and 11.22% on day 15”
t0“tomato seedlings”
e2“Cd + Pb stress”
Claim · model-written summary
activities of superoxide dismutase changed under combined Cd and Pb stress by 11.22%.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- Cd + Pb stresse2
- Outcome
- activities of superoxide dismutasee0
- Direction
- enhancede0
- Value
- 11.22%e0
- Qualifier
- day 15e0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“enhanced the activities of superoxide dismutase by 33.32% on day 10 and 11.22% on day 15”
t0“tomato seedlings”
e2“Cd + Pb stress”
Claim · model-written summary
peroxidase changed under combined Cd and Pb stress by 42.15%.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- Cd + Pb stresse2
- Outcome
- peroxidasee0
- Direction
- increaseinferrede0
- Value
- 42.15%e0
- Qualifier
- day 10e0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“peroxidase by 42.15% on day 10”
t0“tomato seedlings”
e2“Cd + Pb stress”
Claim · model-written summary
catalase changed under combined Cd and Pb stress by 90.78%.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- Cd + Pb stresse2
- Outcome
- catalasee0
- Direction
- increaseinferrede0
- Value
- 90.78%e0
- Qualifier
- day 10e0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“catalase by 90.78% on day 10”
t0“tomato seedlings”
e2“Cd + Pb stress”
Claim · model-written summary
contents of hydrogen peroxide changed under combined Cd and Pb stress by 15.47% to 29.64%.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- Cd + Pb stresse2
- Outcome
- contents of hydrogen peroxidee0
- Direction
- reducede0
- Value
- 15.47% to 29.64%e0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“reduced the contents of hydrogen peroxide by 15.47% to 29.64%”
t0“tomato seedlings”
e2“Cd + Pb stress”
Claim · model-written summary
rate of formation of superoxide anions changed under combined Cd and Pb stress by 26.34% to 53.47%.
- Taxon
- tomato
Solanum lycopersicumt0 - Treatment
- Cd + Pb stresse2
- Outcome
- rate of formation of superoxide anionse0
- Direction
- decreaseinferrede0
- Value
- 26.34% to 53.47%e0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“rate of formation of superoxide anions by 26.34% to 53.47%”
t0“tomato seedlings”
e2“Cd + Pb stress”
Claim · model-written summary
differential expression of the genes.
- Taxon
- tomato
Solanum lycopersicumt0 - Outcome
- differential expression of the genese0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“significant differential expression of the genes involved in pathways, such as phenylalanine, glutathione, arginine and proline, and nitrogen metabolism”
t0“tomato seedlings”
Claim · model-written summary
These genes included PALs , HDCs , GGCT , ODC1 , LAPs , SMS , and SAMDC.
- Taxon
- tomato
Solanum lycopersicumt0 - Gene
- PALse0
- Outcome
- These genes included PALs , HDCs , GGCT , ODC1 , LAPs , SMS , and SAMDCe0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“These genes included PALs , HDCs , GGCT , ODC1 , LAPs , SMS , and SAMDC”
t0“tomato seedlings”
Claim · model-written summary
transcription factors.
- Taxon
- tomato
Solanum lycopersicumt0 - Gene
- ERF109e0
- Outcome
- transcription factorse0
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“transcription factors, such as ERF109 , ARF9 , GRF3 , GRF4 , GRF7 , and GRF9 , were also significantly regulated”
t0“tomato seedlings”
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