Paper · 2026 · PLANT PHYSIOLOGY

Citric acid alters Arabidopsis root morphology and development through ROS-dependent and ROS-independent mechanisms

Tao Zhang; Jesus T Peng; Jie Chu; et al.

11
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22
verbatim quotes
0
citations
3
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Open access · cc-by-nc-nd

Abstract

Citric acid is an integral component of primary metabolism and cellular energetics and plays extensive roles in other cellular processes, such as signaling, chelating, and exudation. Here, we characterized the unique effects that citric acid has on Arabidopsis (Arabidopsis thaliana) root structure and development. In particular, we investigated how citric acid modifies 2 root types in opposing ways: by inhibiting primary root growth while promoting anchor root growth. To understand the mechanisms driving these different growth patterns within the same organism, we analyzed nutrient and transcriptomic responses to citric acid treatment in anchor roots and primary roots. High-spatial resolution elemental analysis revealed that root meristems and the root-hypocotyl junction are regions of strong nutrient enrichment, but that citric acid treatment has little effect on nutrient levels in these regions. Transcriptional analysis revealed major differences between primary roots and anchor roots in response to citric acid. In particular, citric acid acted as a reactive oxygen species (ROS) scavenger through increased Class III peroxidase transcription, effectively reducing H2O2 levels both in vitro and in vivo. Altering the ROS balance at the root-hypocotyl junction was sufficient to induce anchor root formation. Citric acid treatment also differentially upregulated lignin biosynthesis, lignin assembly, and ETHYLENE RESPONSE FACTOR 115 expression in primary roots and anchor roots. ETHYLENE RESPONSE FACTOR 115 regulates the quiescent center and root columella, and we found that citric acid treatment induces developmental defects in this tissue. Overall, this study reveals that a vital organic acid produced and secreted at relatively high concentrations has both widespread and specific effects on plant development and root architecture.

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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 11

Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Citric acid inhibited primary root growth.

Taxon
ArabidopsisArabidopsist0
Treatment
citric acide0
Outcome
primary root growthe0
Direction
inhibitinge0

Not reportedComparator

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

Arabidopsis
Arabidopsis · Taxon · picture of the genus (one of its species)Original uploader was Brona at en.wikipedia . User:Roepers at nl.wikipedia · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“citric acid modifies 2 root types in opposing ways: by inhibiting primary root growth while promoting anchor root growth”

✓ located verbatim in the source · abstract, characters 351–471 · show in text ↑
t0

“Arabidopsis”

✓ located verbatim in the source · title, characters 19–30 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Citric acid promoted anchor root growth.

Taxon
ArabidopsisArabidopsist0
Treatment
citric acide0
Outcome
anchor root growthe0
Direction
promotinge0

Not reportedComparator

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

Arabidopsis
Arabidopsis · Taxon · picture of the genus (one of its species)Original uploader was Brona at en.wikipedia . User:Roepers at nl.wikipedia · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“citric acid modifies 2 root types in opposing ways: by inhibiting primary root growth while promoting anchor root growth”

✓ located verbatim in the source · abstract, characters 351–471 · show in text ↑
t0

“Arabidopsis”

✓ located verbatim in the source · title, characters 19–30 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Root meristems and root hypocotyl junction had strong nutrient enrichment.

Taxon
ArabidopsisArabidopsist0
Organ / stage
root meristems and the root-hypocotyl junctione0
Outcome
nutrient enrichmente0
Direction
stronge0

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

Arabidopsis
Arabidopsis · Taxon · picture of the genus (one of its species)Original uploader was Brona at en.wikipedia . User:Roepers at nl.wikipedia · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“root meristems and the root-hypocotyl junction are regions of strong nutrient enrichment”

✓ located verbatim in the source · abstract, characters 735–823 · show in text ↑
t0

“Arabidopsis”

✓ located verbatim in the source · title, characters 19–30 · show in text ↑
Intervention effectobservedoriginal resultno effectmeasured

Claim · model-written summary

Citric acid had little effect on nutrient levels in these root regions.

Taxon
ArabidopsisArabidopsist0
Treatment
citric acid treatmente0
Outcome
nutrient levelse0
Direction
little effecte0

Not reportedComparator

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

Arabidopsis
Arabidopsis · Taxon · picture of the genus (one of its species)Original uploader was Brona at en.wikipedia . User:Roepers at nl.wikipedia · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“citric acid treatment has little effect on nutrient levels in these regions”

✓ located verbatim in the source · abstract, characters 834–909 · show in text ↑
t0

“Arabidopsis”

✓ located verbatim in the source · title, characters 19–30 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Primary and anchor roots had markedly different transcriptional responses to citric acid.

Taxon
ArabidopsisArabidopsist0
Treatment
citric acide0
Outcome
differences between primary roots and anchor rootse0

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

Arabidopsis
Arabidopsis · Taxon · picture of the genus (one of its species)Original uploader was Brona at en.wikipedia . User:Roepers at nl.wikipedia · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“major differences between primary roots and anchor roots in response to citric acid”

✓ located verbatim in the source · abstract, characters 945–1028 · show in text ↑
t0

“Arabidopsis”

✓ located verbatim in the source · title, characters 19–30 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Citric acid increased Class III peroxidase transcription.

Taxon
ArabidopsisArabidopsist0
Treatment
citric acide0
Gene
Class III peroxidasee0
Outcome
Class III peroxidase transcriptione0
Direction
increasede0

Not reportedComparator

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

Arabidopsis
Arabidopsis · Taxon · picture of the genus (one of its species)Original uploader was Brona at en.wikipedia . User:Roepers at nl.wikipedia · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“citric acid acted as a reactive oxygen species (ROS) scavenger through increased Class III peroxidase transcription”

✓ located verbatim in the source · abstract, characters 1045–1160 · show in text ↑
t0

“Arabidopsis”

✓ located verbatim in the source · title, characters 19–30 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Citric acid reduced H2O2 levels in vitro and in vivo.

Taxon
ArabidopsisArabidopsist0
Outcome
H2O2 levelse0
Direction
reducinge0
Qualifier
both in vitro and in vivoe0

Not reportedTreatmentComparator

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

Arabidopsis
Arabidopsis · Taxon · picture of the genus (one of its species)Original uploader was Brona at en.wikipedia . User:Roepers at nl.wikipedia · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“reducing H2O2 levels both in vitro and in vivo”

✓ located verbatim in the source · abstract, characters 1174–1220 · show in text ↑
t0

“Arabidopsis”

✓ located verbatim in the source · title, characters 19–30 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Changing ROS balance at the root hypocotyl junction induced anchor root formation.

Taxon
ArabidopsisArabidopsist0
Organ / stage
root-hypocotyl junctione0
Treatment
Altering the ROS balancee0
Outcome
anchor root formatione0

Not reportedComparatorDirection

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

Arabidopsis
Arabidopsis · Taxon · picture of the genus (one of its species)Original uploader was Brona at en.wikipedia . User:Roepers at nl.wikipedia · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Altering the ROS balance at the root-hypocotyl junction was sufficient to induce anchor root formation”

✓ located verbatim in the source · abstract, characters 1222–1324 · show in text ↑
t0

“Arabidopsis”

✓ located verbatim in the source · title, characters 19–30 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Citric acid differentially upregulated lignin biosynthesis and assembly.

Taxon
ArabidopsisArabidopsist0
Organ / stage
primary roots and anchor rootse0
Treatment
Citric acid treatmente0
Outcome
lignin biosynthesis, lignin assemblye0
Direction
upregulatede0

Not reportedComparator

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

Arabidopsis
Arabidopsis · Taxon · picture of the genus (one of its species)Original uploader was Brona at en.wikipedia . User:Roepers at nl.wikipedia · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Citric acid treatment also differentially upregulated lignin biosynthesis, lignin assembly, and ETHYLENE RESPONSE FACTOR 115 expression in primary roots and anchor roots”

✓ located verbatim in the source · abstract, characters 1326–1495 · show in text ↑
t0

“Arabidopsis”

✓ located verbatim in the source · title, characters 19–30 · show in text ↑
Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Citric acid differentially upregulated ETHYLENE RESPONSE FACTOR 115 expression.

Taxon
ArabidopsisArabidopsist0
Treatment
Citric acid treatmente0
Gene
ETHYLENE RESPONSE FACTOR 115e0
Outcome
ETHYLENE RESPONSE FACTOR 115 expressione0
Direction
upregulatede0

Not reportedComparator

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

Arabidopsis
Arabidopsis · Taxon · picture of the genus (one of its species)Original uploader was Brona at en.wikipedia . User:Roepers at nl.wikipedia · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“Citric acid treatment also differentially upregulated lignin biosynthesis, lignin assembly, and ETHYLENE RESPONSE FACTOR 115 expression in primary roots and anchor roots”

✓ located verbatim in the source · abstract, characters 1326–1495 · show in text ↑
t0

“Arabidopsis”

✓ located verbatim in the source · title, characters 19–30 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Citric acid caused developmental defects in the quiescent center and columella.

Taxon
ArabidopsisArabidopsist0
Organ / stage
quiescent center and root columellae0
Treatment
citric acid treatmente0
Outcome
developmental defectse0

Not reportedComparatorDirection

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

Arabidopsis
Arabidopsis · Taxon · picture of the genus (one of its species)Original uploader was Brona at en.wikipedia . User:Roepers at nl.wikipedia · CC BY-SA 3.0 · Wikimedia Commons ↗

Verbatim

e0

“ETHYLENE RESPONSE FACTOR 115 regulates the quiescent center and root columella, and we found that citric acid treatment induces developmental defects in this tissue”

✓ located verbatim in the source · abstract, characters 1497–1661 · show in text ↑
t0

“Arabidopsis”

✓ located verbatim in the source · title, characters 19–30 · show in text ↑

citric acid · molecular: expression · ArabidopsisPositive and negative results

2 papers: 1 positive, 1 negative or no effect

1 in the opposite direction

  1. negative
See all 2 papers →

citric acid · growth · ArabidopsisPositive and negative results

2 papers: 1 positive, 1 negative or no effect

1 in the opposite direction

  1. negative
See all 2 papers →

reactive oxygen species · growth · ArabidopsisPositive and negative results

9 papers: 7 positive, 2 negative or no effect

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

  1. negative
  2. negative
  3. positive

5 papers more in this group

See all 9 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.