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.
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Claims 9
Claim · model-written summary
ZmWRKY104 interacts directly with ZmCCaMK in the nucleus without depending on phosphorylation.
- 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.

Verbatim
e0“ZmWRKY104 directly interacts with ZmCCaMK in the nucleus in a non-phosphorylation-dependent manner”
t0“maize”
Claim · model-written summary
Brassinosteroid treatment upregulates ZmWRKY104 expression.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“BR treatment upregulates ZmWRKY104 expression”
t0“maize”
Claim · model-written summary
ZmWRKY104 overexpression significantly increases APX and SOD antioxidant enzyme activities.
- 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.

Verbatim
e0“overexpression of ZmWRKY104 significantly enhances the activities of antioxidant enzymes (APX and SOD)”
t0“maize”
Claim · model-written summary
Co-expression of ZmWRKY104 and ZmCCaMK synergistically promotes antioxidant defense in maize.
- 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.

Verbatim
e0“Co-expression of ZmWRKY104 and ZmCCaMK synergistically promotes the antioxidant defense system in maize.”
Claim · model-written summary
Under 100 mM NaCl, ZmWRKY104-overexpressing maize had less leaf yellowing than wild type.
- 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.

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”
Claim · model-written summary
Under salt stress, ZmWRKY104-overexpressing maize had greater plant height and root length than wild type.
- 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.

Verbatim
e0“under 100 mM NaCl stress compared to wild-type plants, including reduced leaf yellowing, increased plant height and root length”
t0“maize”
e1“Transgenic maize overexpressing ZmWRKY104 exhibits obvious salt tolerance advantages under 100 mM NaCl stress compared to wild-type plants, including reduced leaf yellowing”
Claim · model-written summary
Under salt stress, ZmWRKY104-overexpressing maize had lower electrolyte leakage and malondialdehyde content than wild type.
- 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.

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”
t0“maize”
e1“Transgenic maize overexpressing ZmWRKY104 exhibits obvious salt tolerance advantages under 100 mM NaCl stress compared to wild-type plants, including reduced leaf yellowing”
Claim · model-written summary
Authors identify a non-phosphorylation-dependent WRKY-CCaMK module that enhances brassinosteroid-induced salt tolerance through antioxidant defense.
- 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.

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”
Claim · model-written summary
Authors propose ZmWRKY104 as a candidate for salt-tolerant maize breeding in northern China.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

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”
The same questions in other papers
brassinolide · molecular: expression · Zea mays (maize)Results agree
3 papers in the same direction as this paper
- positive
- positive
- positive
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.