Abstract
Increased anthropogenic activities over the last decades have led to a gradual increase in cadmium content in the soil, which, due to its high mobility in soil, makes Cd accumulation in plants a serious threat to the health of animals and humans. Plant hormones including melatonin (Mel) and brassinosteroids (BR) are known to provide tolerance against various abiotic stresses. In this work, the role of combined and separate exogenous application of Mel and BR on Cd stress in cherry tomato plants was examined. Cd stress significantly reduced tomato growth by inducing oxidative stress and reduced K+ uptake in roots and shoots. Combined application of Mel and BR reduced detrimental effects of Cd in tomato by (i) reducing Cd accumulation in the shoot; (ii) increasing the activities of different antioxidants (SOD, CAT, APX, GR); (iii) triggering higher expression of genes relating to Cd vacuolar sequestration (Na+/H+ EXCHANGER, SlNHX1; NATURAL RESISTANCE-ASSOCIATED MACROPHAGE PROTEIN 6, SlNRAMP6), and Cd transport and detoxification (HEAVY-METAL-ASSOCIATED 3, SlHMA3; PLANT CADMIUM RESISTANT 2, SlPCR2); and (iv) improving plant K+ homeostasis and contents in root and shoot. The latter trait was associated with the reduced gene expression of K+-permeable outward rectifying channel (SlGORK3), and transcriptional upregulation of high affinity potassium transporter 5 (SIHAK5) under Cd stress. A separate application of Mel and BR showed tissue-specific regulation of tomato growth and Cd tolerance by regulating antioxidant activities, K+ uptake, Cd uptake, and translocation from root to shoot and their endogenous contents. Melatonin per se was more effective in improving Cd tolerance in shoot while beneficial BR effects were more pronounced in roots, and their combined application was effective in both tissues. Taken together, reported results show tissue-specific regulation of Cd tolerance by Mel and BR in cherry tomato plants and demonstrate the efficiency of combined Mel + BR treatment as a practical tool to reduce Cd accumulation and mitigate its negative effects on plant growth.
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Claims 14
Claim · model-written summary
Cd stress significantly reduced tomato growth by inducing oxidative stress.
- Taxon
- tomato
Solanum lycopersicume2 - Treatment
- Cd stresse2
- Outcome
- tomato growthe2
- Direction
- reducede2
- Qualifier
- by inducing oxidative stresse2
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Cd stress in cherry tomato plants”
e2“Cd stress significantly reduced tomato growth by inducing oxidative stress and reduced K+ uptake in roots and shoots”
Claim · model-written summary
Cd stress reduced K+ uptake in roots and shoots of tomato.
- Taxon
- tomato
Solanum lycopersicume2 - Organ / stage
- roots and shootse2
- Treatment
- Cd stresse2
- Outcome
- K+ uptakee2
- Direction
- reducede2
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Cd stress in cherry tomato plants”
e2“Cd stress significantly reduced tomato growth by inducing oxidative stress and reduced K+ uptake in roots and shoots”
Claim · model-written summary
Combined melatonin and brassinosteroid application reduced Cd accumulation in the shoot of tomato.
- Taxon
- tomato
Solanum lycopersicume3 - Organ / stage
- shoote3
- Treatment
- Combined application of Mel and BRe3
- Outcome
- Cd accumulatione3
- Direction
- reducinge3
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Cd stress in cherry tomato plants”
e3“Combined application of Mel and BR reduced detrimental effects of Cd in tomato by (i) reducing Cd accumulation in the shoot”
Claim · model-written summary
Combined melatonin and brassinosteroid application increased the activities of different antioxidants in tomato.
- Taxon
- tomato
Solanum lycopersicume1 - Treatment
- Combined application of Mel and BRe3
- Outcome
- activities of different antioxidantse4
- Direction
- increasinge4
- Qualifier
- SOD, CAT, APX, GRe4
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Cd stress in cherry tomato plants”
e3“Combined application of Mel and BR reduced detrimental effects of Cd in tomato by (i) reducing Cd accumulation in the shoot”
e4“increasing the activities of different antioxidants (SOD, CAT, APX, GR)”
Claim · model-written summary
Combined melatonin and brassinosteroid application raised expression of SlNHX1 in tomato under Cd stress.
- Taxon
- tomato
Solanum lycopersicume1 - Treatment
- Combined application of Mel and BRe3
- Gene
- SlNHX1e5
- Outcome
- expression of genes relating to Cd vacuolar sequestratione5
- Direction
- highere5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Cd stress in cherry tomato plants”
e3“Combined application of Mel and BR reduced detrimental effects of Cd in tomato by (i) reducing Cd accumulation in the shoot”
e5“higher expression of genes relating to Cd vacuolar sequestration (Na+/H+ EXCHANGER, SlNHX1; NATURAL RESISTANCE-ASSOCIATED MACROPHAGE PROTEIN 6, SlNRAMP6), and Cd transport and detoxification (HEAVY-METAL-ASSOCIATED 3, SlHMA3; PLANT CADMIUM RESISTANT 2, SlPCR2)”
Claim · model-written summary
Combined melatonin and brassinosteroid application raised expression of SlNRAMP6 in tomato under Cd stress.
- Taxon
- tomato
Solanum lycopersicume1 - Treatment
- Combined application of Mel and BRe3
- Gene
- SlNRAMP6e5
- Outcome
- expression of genes relating to Cd vacuolar sequestratione5
- Direction
- highere5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Cd stress in cherry tomato plants”
e3“Combined application of Mel and BR reduced detrimental effects of Cd in tomato by (i) reducing Cd accumulation in the shoot”
e5“higher expression of genes relating to Cd vacuolar sequestration (Na+/H+ EXCHANGER, SlNHX1; NATURAL RESISTANCE-ASSOCIATED MACROPHAGE PROTEIN 6, SlNRAMP6), and Cd transport and detoxification (HEAVY-METAL-ASSOCIATED 3, SlHMA3; PLANT CADMIUM RESISTANT 2, SlPCR2)”
Claim · model-written summary
Combined melatonin and brassinosteroid application raised expression of SlHMA3 in tomato.
- Taxon
- tomato
Solanum lycopersicume1 - Treatment
- Combined application of Mel and BRe3
- Gene
- SlHMA3e5
- Outcome
- Cd transport and detoxificatione5
- Direction
- highere5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Cd stress in cherry tomato plants”
e3“Combined application of Mel and BR reduced detrimental effects of Cd in tomato by (i) reducing Cd accumulation in the shoot”
e5“higher expression of genes relating to Cd vacuolar sequestration (Na+/H+ EXCHANGER, SlNHX1; NATURAL RESISTANCE-ASSOCIATED MACROPHAGE PROTEIN 6, SlNRAMP6), and Cd transport and detoxification (HEAVY-METAL-ASSOCIATED 3, SlHMA3; PLANT CADMIUM RESISTANT 2, SlPCR2)”
Claim · model-written summary
Combined melatonin and brassinosteroid application raised expression of SlPCR2 in tomato.
- Taxon
- tomato
Solanum lycopersicume1 - Treatment
- Combined application of Mel and BRe3
- Gene
- SlPCR2e5
- Outcome
- Cd transport and detoxificatione5
- Direction
- highere5
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Cd stress in cherry tomato plants”
e3“Combined application of Mel and BR reduced detrimental effects of Cd in tomato by (i) reducing Cd accumulation in the shoot”
e5“higher expression of genes relating to Cd vacuolar sequestration (Na+/H+ EXCHANGER, SlNHX1; NATURAL RESISTANCE-ASSOCIATED MACROPHAGE PROTEIN 6, SlNRAMP6), and Cd transport and detoxification (HEAVY-METAL-ASSOCIATED 3, SlHMA3; PLANT CADMIUM RESISTANT 2, SlPCR2)”
Claim · model-written summary
Higher K+ homeostasis in tomato under Cd stress was associated with reduced SlGORK3 gene expression.
- Taxon
- tomato
Solanum lycopersicume1 - Treatment
- Cd stresse6
- Gene
- SlGORK3e6
- Outcome
- gene expressione6
- Direction
- reducede6
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Cd stress in cherry tomato plants”
e6“The latter trait was associated with the reduced gene expression of K+-permeable outward rectifying channel (SlGORK3), and transcriptional upregulation of high affinity potassium transporter 5 (SIHAK5) under Cd stress”
Claim · model-written summary
Higher K+ homeostasis in tomato under Cd stress was associated with upregulated SIHAK5 expression.
- Taxon
- tomato
Solanum lycopersicume1 - Treatment
- Cd stresse6
- Gene
- SIHAK5e6
- Outcome
- gene expressione6
- Direction
- upregulatione6
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Cd stress in cherry tomato plants”
e6“The latter trait was associated with the reduced gene expression of K+-permeable outward rectifying channel (SlGORK3), and transcriptional upregulation of high affinity potassium transporter 5 (SIHAK5) under Cd stress”
Claim · model-written summary
Melatonin alone was more effective than BR in improving Cd tolerance in tomato shoot.
- Taxon
- tomato
Solanum lycopersicume1 - Organ / stage
- shoote7
- Treatment
- Melatonin per see7
- Outcome
- Cd tolerancee7
- Direction
- more effectivee7
- Comparator
- BRe7
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Cd stress in cherry tomato plants”
e7“Melatonin per se was more effective in improving Cd tolerance in shoot while beneficial BR effects were more pronounced in roots”
Claim · model-written summary
BR alone was more effective than melatonin in improving Cd tolerance in tomato roots.
- Taxon
- tomato
Solanum lycopersicume1 - Organ / stage
- rootse7
- Treatment
- BRe7
- Outcome
- Cd tolerancee7
- Direction
- more pronouncede7
- Comparator
- Melatonin per see7
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Cd stress in cherry tomato plants”
e7“Melatonin per se was more effective in improving Cd tolerance in shoot while beneficial BR effects were more pronounced in roots”
Claim · model-written summary
Combined melatonin and BR application was effective in both tomato roots and shoots against Cd stress.
- Taxon
- tomato
Solanum lycopersicume1 - Organ / stage
- both tissuese2
- Treatment
- combined applicatione2
- Outcome
- Cd tolerancee7
- Direction
- effectivee2
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Cd stress in cherry tomato plants”
e7“Melatonin per se was more effective in improving Cd tolerance in shoot while beneficial BR effects were more pronounced in roots”
e2“their combined application was effective in both tissues”
Claim · model-written summary
Combined melatonin and brassinosteroid application improved K+ homeostasis and contents in tomato root and shoot.
- Taxon
- tomato
Solanum lycopersicume1 - Organ / stage
- root and shoote2
- Treatment
- Combined application of Mel and BRe3
- Outcome
- K+ homeostasis and contentse2
- Direction
- improvinge2
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Cd stress in cherry tomato plants”
e3“Combined application of Mel and BR reduced detrimental effects of Cd in tomato by (i) reducing Cd accumulation in the shoot”
e2“(iv) improving plant K+ homeostasis and contents in root and shoot”
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