Paper · 2025 · Journal of Pineal Research

Melatonin Modulates Copper Homeostasis Through NO Dependent Manner in Rice ( Oryza sativa L.)

Jing Huang; Hao Yu Wang; Jia Qi Li; et al.

14
claims
31
verbatim quotes
3
citations
2
research areas
—
Not open access

Abstract

Melatonin (MT) plays a critical role in regulating rice responses to heavy metal stress. However, the mechanism by which MT alleviates copper (Cu) toxicity in rice remains insufficiently understood. In this study, excessive Cu increased endogenous MT level in rice roots, accompanied by enhanced gene expression involved in MT biosynthesis, suggesting its role in Cu stress responses. Exogenous MT reduced Cu accumulation and restored root growth in rice under Cu stress, thereby mitigating Cu-induced phytotoxicity. Mechanistically, MT increased the root pectin content and its Cu retention, thereby reducing Cu influx into the cytoplasm. Additionally, exogenous MT upregulated the expression of Heavy Metal ATPase 4 (OsHMA4) involved in vacuolar sequestration, as well as Cu efflux transporter genes OsHMA6 and OsHMA9. Conversely, it downregulated the plasma membrane Cu uptake-associated genes Copper Transporter (OsCOPT1, OsCOPT2, and OsCOPT3), the vacuolar Cu exporter OsCOPT7, and the xylem loading transporter gene OsHMA5, thereby alleviating Cu toxicity in rice. Furthermore, MT elevated nitric oxide (NO) levels in root tips, and application of the exogenous NO scavenger, 2-(4-carboxyphenyl)-4, 4, 5, 5-tetramethylimidazoline-1-oxyl-3-oxide (cPTIO), attenuated the mitigative effect of MT on excess Cu-stressed rice. In summary, MT alleviates Cu toxicity by reducing Cu binding to root cell walls, thereby limiting Cu uptake and translocation from roots to shoots. This process may depend on NO accumulation and provides new insights into the molecular mechanisms underlying MT-induced Cu tolerance in rice.

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Claims 14

Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Excess copper increased endogenous melatonin in rice roots.

Organ / stage
rootse0
Treatment
excessive Cue0
Outcome
endogenous MT levele0
Direction
increasede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“excessive Cu increased endogenous MT level in rice roots”

✓ located verbatim in the source · abstract, characters 214–270 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Excess copper enhanced expression of genes involved in melatonin biosynthesis in rice roots.

Organ / stage
rootse0
Treatment
excessive Cue0
Outcome
gene expression involved in MT biosynthesise0
Direction
enhancede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“excessive Cu increased endogenous MT level in rice roots, accompanied by enhanced gene expression involved in MT biosynthesis”

✓ located verbatim in the source · abstract, characters 214–339 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Exogenous melatonin reduced copper accumulation in stressed rice.

Treatment
Exogenous MTe0
Outcome
Cu accumulatione0
Direction
reducede0
Qualifier
under Cu stresse0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“Exogenous MT reduced Cu accumulation and restored root growth in rice under Cu stress”

✓ located verbatim in the source · abstract, characters 385–470 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Exogenous melatonin restored rice root growth under copper stress.

Treatment
Exogenous MTe0
Outcome
root growthe0
Direction
restorede0
Qualifier
under Cu stresse0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“Exogenous MT reduced Cu accumulation and restored root growth in rice under Cu stress”

✓ located verbatim in the source · abstract, characters 385–470 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Melatonin increased rice root pectin content.

Treatment
MTe0
Outcome
root pectin contente0
Direction
increasede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“MT increased the root pectin content and its Cu retention”

✓ located verbatim in the source · abstract, characters 534–591 · show in text ↑
e1

“in rice under Cu stress”

✓ located verbatim in the source · abstract, characters 447–470 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Melatonin increased copper retention in root pectin.

Treatment
MTe0
Outcome
Cu retentione0
Direction
increasede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“MT increased the root pectin content and its Cu retention”

✓ located verbatim in the source · abstract, characters 534–591 · show in text ↑
e1

“in rice under Cu stress”

✓ located verbatim in the source · abstract, characters 447–470 · show in text ↑
Mechanismobservedoriginal resultpositivemeasured

Claim · model-written summary

Melatonin reduced copper influx into the cytoplasm.

Treatment
MTe2
Outcome
Cu influx into the cytoplasme0
Direction
reducinge0

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“thereby reducing Cu influx into the cytoplasm”

✓ located verbatim in the source · abstract, characters 593–638 · show in text ↑
e1

“in rice under Cu stress”

✓ located verbatim in the source · abstract, characters 447–470 · show in text ↑
e2

“MT increased the root pectin content”

✓ located verbatim in the source · abstract, characters 534–570 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Exogenous melatonin upregulated OsHMA4 expression.

Treatment
exogenous MTe0
Gene
OsHMA4e0
Outcome
expression of Heavy Metal ATPase 4 (OsHMA4)e0
Direction
upregulatede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“exogenous MT upregulated the expression of Heavy Metal ATPase 4 (OsHMA4)”

✓ located verbatim in the source · abstract, characters 654–726 · show in text ↑
e1

“in rice under Cu stress”

✓ located verbatim in the source · abstract, characters 447–470 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Exogenous melatonin upregulated OsHMA6 and OsHMA9 expression.

Treatment
exogenous MTe2
Gene
OsHMA6 and OsHMA9e0
Outcome
Cu efflux transporter genes OsHMA6 and OsHMA9e0
Direction
upregulatede2

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“as well as Cu efflux transporter genes OsHMA6 and OsHMA9”

✓ located verbatim in the source · abstract, characters 763–819 · show in text ↑
e1

“in rice under Cu stress”

✓ located verbatim in the source · abstract, characters 447–470 · show in text ↑
e2

“exogenous MT upregulated”

✓ located verbatim in the source · abstract, characters 654–678 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Exogenous melatonin downregulated OsCOPT1, OsCOPT2, and OsCOPT3 expression.

Treatment
exogenous MTe2
Gene
OsCOPT1, OsCOPT2, and OsCOPT3e0
Outcome
plasma membrane Cu uptake-associated genese0
Direction
downregulatede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“it downregulated the plasma membrane Cu uptake-associated genes Copper Transporter (OsCOPT1, OsCOPT2, and OsCOPT3)”

✓ located verbatim in the source · abstract, characters 833–947 · show in text ↑
e1

“in rice under Cu stress”

✓ located verbatim in the source · abstract, characters 447–470 · show in text ↑
e2

“exogenous MT upregulated”

✓ located verbatim in the source · abstract, characters 654–678 · show in text ↑
e3

“it downregulated”

✓ located verbatim in the source · abstract, characters 833–849 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Exogenous melatonin downregulated OsCOPT7 expression.

Treatment
exogenous MTe2
Gene
OsCOPT7e0
Outcome
vacuolar Cu exporter OsCOPT7e0
Direction
downregulatede3

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“the vacuolar Cu exporter OsCOPT7”

✓ located verbatim in the source · abstract, characters 949–981 · show in text ↑
e1

“in rice under Cu stress”

✓ located verbatim in the source · abstract, characters 447–470 · show in text ↑
e2

“exogenous MT upregulated”

✓ located verbatim in the source · abstract, characters 654–678 · show in text ↑
e3

“it downregulated”

✓ located verbatim in the source · abstract, characters 833–849 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Exogenous melatonin downregulated OsHMA5 expression.

Treatment
exogenous MTe2
Gene
OsHMA5e0
Outcome
xylem loading transporter gene OsHMA5e0
Direction
downregulatede3

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“the xylem loading transporter gene OsHMA5”

✓ located verbatim in the source · abstract, characters 987–1028 · show in text ↑
e1

“in rice under Cu stress”

✓ located verbatim in the source · abstract, characters 447–470 · show in text ↑
e2

“exogenous MT upregulated”

✓ located verbatim in the source · abstract, characters 654–678 · show in text ↑
e3

“it downregulated”

✓ located verbatim in the source · abstract, characters 833–849 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Melatonin elevated nitric oxide levels in rice root tips.

Organ / stage
root tipse0
Treatment
MTe0
Outcome
nitric oxide (NO) levelse0
Direction
elevatede0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“MT elevated nitric oxide (NO) levels in root tips”

✓ located verbatim in the source · abstract, characters 1084–1133 · show in text ↑
e1

“alleviating Cu toxicity in rice.”

✓ located verbatim in the source · abstract, characters 1038–1070 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

The nitric oxide scavenger attenuated the copper-stress mitigation from melatonin in rice.

Treatment
exogenous NO scavengere0
Outcome
mitigative effect of MTe0
Direction
attenuatede0
Qualifier
excess Cu-stressed ricee0

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Oryza sativa
Oryza sativa · Taxon · picture from its Wikimedia Commons categoryDesmanthus4food · CC BY-SA 3.0 us · Wikimedia Commons ↗

Verbatim

e0

“application of the exogenous NO scavenger, 2-(4-carboxyphenyl)-4, 4, 5, 5-tetramethylimidazoline-1-oxyl-3-oxide (cPTIO), attenuated the mitigative effect of MT on excess Cu-stressed rice”

✓ located verbatim in the source · abstract, characters 1139–1325 · show in text ↑

copper atom · molecular: metabolites · Oryza sativa (rice)Positive and negative results

12 papers: 5 positive, 7 negative or no effect

4 papers in the same direction as this paper · 7 in the opposite direction

  1. negative
  2. negative
    B and Cu deficiencies slightly increased NiR activity.

    Nitrate assimilation in rice seedlings as affected by mineral nutrition · 1977

    Opposite directionAbout lacking this nutrient, so it counts the other way round

  3. negative

8 papers more in this group

See all 12 papers →

nitric oxide · molecular: metabolites · Oryza sativa (rice)Positive and negative results

13 papers: 8 positive, 5 negative or no effect

4 papers in the same direction as this paper · 8 in the opposite direction

  1. positive
  2. positive
  3. positive

9 papers more in this group

See all 13 papers →

melatonin · molecular: metabolites · Oryza sativa (rice)Positive and negative results

16 papers: 14 positive, 2 negative or no effect

13 papers in the same direction as this paper · 2 in the opposite direction

  1. no effect
  2. no effect
  3. positive

12 papers more in this group

See all 16 papers →

melatonin · nutrient: content · Oryza sativa (rice)Results agree

13 papers: 13 positive, 0 negative or no effect

12 papers in the same direction as this paper

  1. positive
  2. positive
  3. positive

9 papers more in this group

See all 13 papers →

melatonin · molecular: expression · Oryza sativa (rice)Results agree

20 papers: 20 positive, 0 negative or no effect

19 papers in the same direction as this paper

  1. positive
  2. positive
  3. positive

16 papers more in this group

See all 20 papers →

melatonin · growth · Oryza sativa (rice)Results agree

28 papers: 28 positive, 0 negative or no effect

27 papers in the same direction as this paper

  1. positive
  2. positive
  3. positive

24 papers more in this group

See all 28 papers →

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.