Paper · 2026 · PLoS ONE

Copper causes reduced nitrogen fixation but does not accumulate in the nodules of the legume Lotus japonicus

Kathryn Lamoureux; Sheila M. Macfie

10
claims
20
verbatim quotes
0
citations
2
research areas
✓
Open access · cc-by

Abstract

Copper can be a soil contaminant at concentrations that are toxic to plants, particularly because of copper-induced oxidative stress. The legume-rhizobia partnership that allows for biological nitrogen fixation is sensitive to oxidative stress, and this study investigates if copper acts directly on the machinery of nitrogen fixation, or indirectly via toxicity to the entire plant. When Lotus japonicus inoculated with its rhizobial partner Mesorhizobium loti was exposed to 300 or 450 µM of copper, biomass was reduced by 30-40% in the shoots, 40-55% in the roots, and 40-60% in the nodules relative to control. While concentrations of copper in shoots and roots increased in proportion to the amount of copper in the growth medium, concentrations of copper in nodules did not vary in response to copper treatment. Malondialdehyde, a marker of oxidative damage, in the nodule similarly did not vary with copper treatment. However, nitrogen fixation and ascorbate peroxidase activity decreased by 40-45% and 40-60%, respectively, which can be indicators of early nodule senescence. This would suggest that copper-induced reduction in nodule activity is not directly due to oxidative stress in the nodules; it is due to stress on the host plant that limits its ability to support its symbionts.

Abstract from the paper’s OpenAlex record; open copy licensed cc-by. Read at the source ↗

The text the model read, with the source record’s inline markup (italics, sub- and superscripts) rendered and its character entities decoded; quote offsets refer to the raw text.

Claims 10

Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Copper reduced Lotus japonicus shoot biomass by 30-40% relative to control in copper-exposed nodulated plants

Taxon
Lotus japonicusLotus japonicuse1
Organ / stage
shootse2
Treatment
300 or 450 µM of coppere1
Interactor
Mesorhizobium lotiMesorhizobium lotie1symbiont
Outcome
biomasse2
Direction
reducede2
Value
30-40%e2
Comparator
controle2

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

Lotus japonicus
Lotus japonicus · Taxonharum.koh · CC BY-SA 4.0 · iNaturalist ↗
Mesorhizobium loti
Mesorhizobium loti · Interactor · picture from its Wikimedia Commons categoryNinjatacoshell · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e1

“When Lotus japonicus inoculated with its rhizobial partner Mesorhizobium loti was exposed to 300 or 450 µM of copper”

✓ located verbatim in the source · abstract, characters 384–500 · show in text ↑
e2

“biomass was reduced by 30-40% in the shoots, 40-55% in the roots, and 40-60% in the nodules relative to control”

✓ located verbatim in the source · abstract, characters 502–613 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Copper reduced Lotus japonicus root biomass by 40-55% relative to control in copper-exposed nodulated plants

Taxon
Lotus japonicusLotus japonicuse1
Organ / stage
rootse2
Treatment
300 or 450 µM of coppere1
Interactor
Mesorhizobium lotiMesorhizobium lotie1symbiont
Outcome
biomasse2
Direction
reducede2
Value
40-55%e2
Comparator
controle2

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

Lotus japonicus
Lotus japonicus · Taxonharum.koh · CC BY-SA 4.0 · iNaturalist ↗
Mesorhizobium loti
Mesorhizobium loti · Interactor · picture from its Wikimedia Commons categoryNinjatacoshell · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e1

“When Lotus japonicus inoculated with its rhizobial partner Mesorhizobium loti was exposed to 300 or 450 µM of copper”

✓ located verbatim in the source · abstract, characters 384–500 · show in text ↑
e2

“biomass was reduced by 30-40% in the shoots, 40-55% in the roots, and 40-60% in the nodules relative to control”

✓ located verbatim in the source · abstract, characters 502–613 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Copper reduced Lotus japonicus nodule biomass by 40-60% relative to control

Taxon
Lotus japonicusLotus japonicuse1
Organ / stage
nodulese2
Treatment
300 or 450 µM of coppere1
Interactor
Mesorhizobium lotiMesorhizobium lotie1symbiont
Outcome
biomasse2
Direction
reducede2
Value
40-60%e2
Comparator
controle2

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

Lotus japonicus
Lotus japonicus · Taxonharum.koh · CC BY-SA 4.0 · iNaturalist ↗
Mesorhizobium loti
Mesorhizobium loti · Interactor · picture from its Wikimedia Commons categoryNinjatacoshell · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e1

“When Lotus japonicus inoculated with its rhizobial partner Mesorhizobium loti was exposed to 300 or 450 µM of copper”

✓ located verbatim in the source · abstract, characters 384–500 · show in text ↑
e2

“biomass was reduced by 30-40% in the shoots, 40-55% in the roots, and 40-60% in the nodules relative to control”

✓ located verbatim in the source · abstract, characters 502–613 · show in text ↑
Associationobservedoriginal resultmeasured

Claim · model-written summary

Copper concentrations in Lotus japonicus shoots and roots increased in proportion to the copper supply

Taxon
Lotus japonicusLotus japonicuse1
Treatment
the amount of copper in the growth mediume3
Outcome
concentrations of copper in shoots and rootse3
Direction
increased in proportion toe3

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

Lotus japonicus
Lotus japonicus · Taxonharum.koh · CC BY-SA 4.0 · iNaturalist ↗

Verbatim

e1

“When Lotus japonicus inoculated with its rhizobial partner Mesorhizobium loti was exposed to 300 or 450 µM of copper”

✓ located verbatim in the source · abstract, characters 384–500 · show in text ↑
e3

“While concentrations of copper in shoots and roots increased in proportion to the amount of copper in the growth medium”

✓ located verbatim in the source · abstract, characters 615–734 · show in text ↑
Associationobservedoriginal resultno effectmeasured

Claim · model-written summary

Copper concentrations in Lotus japonicus nodules did not vary with copper treatment

Taxon
Lotus japonicusLotus japonicuse1
Organ / stage
nodulese4
Treatment
copper treatmente4
Outcome
concentrations of copper in nodulese4
Direction
did not varye4

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

Lotus japonicus
Lotus japonicus · Taxonharum.koh · CC BY-SA 4.0 · iNaturalist ↗

Verbatim

e1

“When Lotus japonicus inoculated with its rhizobial partner Mesorhizobium loti was exposed to 300 or 450 µM of copper”

✓ located verbatim in the source · abstract, characters 384–500 · show in text ↑
e4

“concentrations of copper in nodules did not vary in response to copper treatment”

✓ located verbatim in the source · abstract, characters 736–816 · show in text ↑
Associationobservedoriginal resultno effectmeasured

Claim · model-written summary

Malondialdehyde in Lotus japonicus nodules did not vary with copper treatment

Taxon
Lotus japonicusLotus japonicuse1
Organ / stage
nodulee5
Treatment
copper treatmente5
Outcome
Malondialdehydee5
Direction
did not varye5

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

Lotus japonicus
Lotus japonicus · Taxonharum.koh · CC BY-SA 4.0 · iNaturalist ↗

Verbatim

e1

“When Lotus japonicus inoculated with its rhizobial partner Mesorhizobium loti was exposed to 300 or 450 µM of copper”

✓ located verbatim in the source · abstract, characters 384–500 · show in text ↑
e5

“Malondialdehyde, a marker of oxidative damage, in the nodule similarly did not vary with copper treatment”

✓ located verbatim in the source · abstract, characters 818–923 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Copper exposure decreased nitrogen fixation in Lotus japonicus by 40-45%

Taxon
Lotus japonicusLotus japonicuse1
Treatment
300 or 450 µM of coppere1
Interactor
Mesorhizobium lotiMesorhizobium lotie1symbiont
Outcome
nitrogen fixatione6
Direction
decreasede6
Value
40-45%e6
Qualifier
which can be indicators of early nodule senescencee6

Not reportedComparator

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

Lotus japonicus
Lotus japonicus · Taxonharum.koh · CC BY-SA 4.0 · iNaturalist ↗
Mesorhizobium loti
Mesorhizobium loti · Interactor · picture from its Wikimedia Commons categoryNinjatacoshell · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e1

“When Lotus japonicus inoculated with its rhizobial partner Mesorhizobium loti was exposed to 300 or 450 µM of copper”

✓ located verbatim in the source · abstract, characters 384–500 · show in text ↑
e6

“nitrogen fixation and ascorbate peroxidase activity decreased by 40-45% and 40-60%, respectively, which can be indicators of early nodule senescence”

✓ located verbatim in the source · abstract, characters 934–1082 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Copper exposure decreased ascorbate peroxidase activity in Lotus japonicus nodules by 40-60%

Taxon
Lotus japonicusLotus japonicuse1
Treatment
300 or 450 µM of coppere1
Interactor
Mesorhizobium lotiMesorhizobium lotie1symbiont
Outcome
ascorbate peroxidase activitye6
Direction
decreasede6
Value
40-60%e6

Not reportedComparator

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

Lotus japonicus
Lotus japonicus · Taxonharum.koh · CC BY-SA 4.0 · iNaturalist ↗
Mesorhizobium loti
Mesorhizobium loti · Interactor · picture from its Wikimedia Commons categoryNinjatacoshell · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e1

“When Lotus japonicus inoculated with its rhizobial partner Mesorhizobium loti was exposed to 300 or 450 µM of copper”

✓ located verbatim in the source · abstract, characters 384–500 · show in text ↑
e6

“nitrogen fixation and ascorbate peroxidase activity decreased by 40-45% and 40-60%, respectively, which can be indicators of early nodule senescence”

✓ located verbatim in the source · abstract, characters 934–1082 · show in text ↑
Mechanismauthor interpretationoriginal resultmeasured

Claim · model-written summary

Copper-induced reduction in nodule activity is not directly due to oxidative stress in the nodules

Taxon
Lotus japonicusLotus japonicuse1
Outcome
copper-induced reduction in nodule activitye7
Qualifier
not directly due to oxidative stress in the nodulese7

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

Lotus japonicus
Lotus japonicus · Taxonharum.koh · CC BY-SA 4.0 · iNaturalist ↗

Verbatim

e1

“When Lotus japonicus inoculated with its rhizobial partner Mesorhizobium loti was exposed to 300 or 450 µM of copper”

✓ located verbatim in the source · abstract, characters 384–500 · show in text ↑
e7

“This would suggest that copper-induced reduction in nodule activity is not directly due to oxidative stress in the nodules; it is due to stress on the host plant that limits its ability to support its symbionts”

✓ located verbatim in the source · abstract, characters 1084–1294 · show in text ↑
Mechanismauthor interpretationoriginal resultmeasured

Claim · model-written summary

Copper-induced reduction in nodule activity is due to stress on the host plant that limits its support of symbionts

Taxon
Lotus japonicusLotus japonicuse1
Outcome
copper-induced reduction in nodule activitye7
Qualifier
stress on the host plant that limits its ability to support its symbiontse7

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

Lotus japonicus
Lotus japonicus · Taxonharum.koh · CC BY-SA 4.0 · iNaturalist ↗

Verbatim

e1

“When Lotus japonicus inoculated with its rhizobial partner Mesorhizobium loti was exposed to 300 or 450 µM of copper”

✓ located verbatim in the source · abstract, characters 384–500 · show in text ↑
e7

“This would suggest that copper-induced reduction in nodule activity is not directly due to oxidative stress in the nodules; it is due to stress on the host plant that limits its ability to support its symbionts”

✓ located verbatim in the source · abstract, characters 1084–1294 · show in text ↑