Paper · 2026 · Plant Signaling & Behavior

ZmWRKY104–ZmCCaMK interaction enhances brassinosteroid-promoted salt tolerance in maize ( Zea mays L.) via antioxidant defense

Li-li Zhao

9
claims
16
verbatim quotes
1
citations
3
research areas
✓
Open access · cc-by

Abstract

Brassinosteroid (BR)-mediated salt tolerance is a crucial mechanism for maize (Zea mays L.) adaptation to saline-alkaline environments. This study aimed to elucidate the molecular mechanism underlying BR-induced salt tolerance in maize, focusing on the regulatory roles of ZmWRKY104 and ZmCCaMK. Key results showed that ZmWRKY104 directly interacts with ZmCCaMK in the nucleus in a non-phosphorylation-dependent manner, forming a novel regulatory module. BR treatment upregulates ZmWRKY104 expression, and overexpression of ZmWRKY104 significantly enhances the activities of antioxidant enzymes (APX and SOD). Co-expression of ZmWRKY104 and ZmCCaMK synergistically promotes the antioxidant defense system in maize. Transgenic maize overexpressing ZmWRKY104 exhibits obvious salt tolerance advantages under 100 mM NaCl stress compared to wild-type plants, including reduced leaf yellowing, increased plant height and root length, as well as decreased electrolyte leakage (EL) and malondialdehyde (MDA) content. Collectively, this study identifies a novel non-phosphorylation-dependent WRKY-CCaMK regulatory module in the BR signaling pathway, which enhances BR-induced maize salt tolerance by synergistically activating antioxidant defense. The findings highlight ZmWRKY104 as a candidate gene and provide a potential molecular mechanism for salt-tolerant maize breeding in saline-alkaline regions of northern China.

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 9

Observationobservedoriginal resultmeasured

Claim · model-written summary

ZmWRKY104 interacts directly with ZmCCaMK in the nucleus without depending on phosphorylation.

Taxon
maizeZea mayst0
Gene
ZmWRKY104e0
Outcome
interacts with ZmCCaMK in the nucleuse0
Qualifier
non-phosphorylation-dependent mannere0

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“ZmWRKY104 directly interacts with ZmCCaMK in the nucleus in a non-phosphorylation-dependent manner”

✓ located verbatim in the source · abstract, characters 320–418 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 82–87 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Brassinosteroid treatment upregulates ZmWRKY104 expression.

Taxon
maizeZea mayst0
Treatment
BR treatmente0
Gene
ZmWRKY104e0
Outcome
ZmWRKY104 expressione0
Direction
upregulatese0

Not reportedComparator

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“BR treatment upregulates ZmWRKY104 expression”

✓ located verbatim in the source · abstract, characters 455–500 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 82–87 · show in text ↑
Intervention effectobservedoriginal resultpositivesignificant (as stated)measured

Claim · model-written summary

ZmWRKY104 overexpression significantly increases APX and SOD antioxidant enzyme activities.

Taxon
maizeZea mayst0
Treatment
overexpression of ZmWRKY104e0
Gene
ZmWRKY104e0
Outcome
activities of antioxidant enzymes (APX and SOD)e0
Direction
enhancese0

Not reportedComparator

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“overexpression of ZmWRKY104 significantly enhances the activities of antioxidant enzymes (APX and SOD)”

✓ located verbatim in the source · abstract, characters 506–608 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 82–87 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Co-expression of ZmWRKY104 and ZmCCaMK synergistically promotes antioxidant defense in maize.

Taxon
maizeZea mayse0
Treatment
Co-expression of ZmWRKY104 and ZmCCaMKe0
Gene
ZmWRKY104 and ZmCCaMKe0
Outcome
antioxidant defense systeme0
Direction
promotese0

Not reportedComparator

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“Co-expression of ZmWRKY104 and ZmCCaMK synergistically promotes the antioxidant defense system in maize.”

✓ located verbatim in the source · abstract, characters 610–714 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Under 100 mM NaCl, ZmWRKY104-overexpressing maize had less leaf yellowing than wild type.

Taxon
maizeZea mayse0
Treatment
overexpressing ZmWRKY104e0
Outcome
leaf yellowinge0
Direction
reducede0
Qualifier
under 100 mM NaCl stresse0
Comparator
wild-type plantse0

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“Transgenic maize overexpressing ZmWRKY104 exhibits obvious salt tolerance advantages under 100 mM NaCl stress compared to wild-type plants, including reduced leaf yellowing”

✓ located verbatim in the source · abstract, characters 715–887 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Under salt stress, ZmWRKY104-overexpressing maize had greater plant height and root length than wild type.

Taxon
maizeZea mayst0
Treatment
overexpressing ZmWRKY104e1
Outcome
plant height and root lengthe0
Direction
increasede0
Qualifier
under 100 mM NaCl stresse0
Comparator
wild-type plantse0

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“under 100 mM NaCl stress compared to wild-type plants, including reduced leaf yellowing, increased plant height and root length”

✓ located verbatim in the source · abstract, characters 800–927 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 82–87 · show in text ↑
e1

“Transgenic maize overexpressing ZmWRKY104 exhibits obvious salt tolerance advantages under 100 mM NaCl stress compared to wild-type plants, including reduced leaf yellowing”

✓ located verbatim in the source · abstract, characters 715–887 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Under salt stress, ZmWRKY104-overexpressing maize had lower electrolyte leakage and malondialdehyde content than wild type.

Taxon
maizeZea mayst0
Treatment
overexpressing ZmWRKY104e1
Outcome
electrolyte leakage (EL) and malondialdehyde (MDA) contente0
Direction
decreasede0
Comparator
wild-type plantse0

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“under 100 mM NaCl stress compared to wild-type plants, including reduced leaf yellowing, increased plant height and root length, as well as decreased electrolyte leakage (EL) and malondialdehyde (MDA) content”

✓ located verbatim in the source · abstract, characters 800–1008 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 82–87 · show in text ↑
e1

“Transgenic maize overexpressing ZmWRKY104 exhibits obvious salt tolerance advantages under 100 mM NaCl stress compared to wild-type plants, including reduced leaf yellowing”

✓ located verbatim in the source · abstract, characters 715–887 · show in text ↑
Mechanismauthor interpretationoriginal resultpositivemeasured

Claim · model-written summary

Authors identify a non-phosphorylation-dependent WRKY-CCaMK module that enhances brassinosteroid-induced salt tolerance through antioxidant defense.

Taxon
maizeZea mayse0
Gene
WRKY-CCaMKe0
Outcome
maize salt tolerancee0
Qualifier
by synergistically activating antioxidant defensee0

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“a novel non-phosphorylation-dependent WRKY-CCaMK regulatory module in the BR signaling pathway, which enhances BR-induced maize salt tolerance by synergistically activating antioxidant defense”

✓ located verbatim in the source · abstract, characters 1046–1238 · show in text ↑
Recommendationauthor interpretationoriginal resultbasis n/a

Claim · model-written summary

Authors propose ZmWRKY104 as a candidate for salt-tolerant maize breeding in northern China.

Taxon
maizeZea mayse0
Gene
ZmWRKY104e0
Outcome
salt-tolerant maize breedinge0
Setting
northern Chinae0

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

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

e0

“ZmWRKY104 as a candidate gene and provide a potential molecular mechanism for salt-tolerant maize breeding in saline-alkaline regions of northern China”

✓ located verbatim in the source · abstract, characters 1263–1414 · show in text ↑

brassinolide · molecular: expression · Zea mays (maize)Results agree

4 papers: 4 positive, 0 negative or no effect

3 papers in the same direction as this paper

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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.