Abstract
The DIR gene family, which encodes Dirigent proteins, plays a crucial role in plant development and stress responses. However, the functions and mechanisms of this family in maize remained underexplored. This study identified ZmDIR11, a member of the maize Dirigent protein family, and explored its role in drought tolerance. ZmDIR11 shared conserved regions with homologous proteins in wheat, rice, and Arabidopsis. RT-qPCR analysis revealed that ZmDIR11 expression is upregulated in leaves under drought and PEG stress, while subcellular localization confirmed its presence in the endoplasmic reticulum. Virus-induced gene silencing (VIGS) and EMS mutagenesis demonstrated that ZmDIR11 silencing or mutation significantly reduces drought tolerance in maize seedlings, indicating its positive regulatory role. Silencing or mutation of ZmDIR11 led to decreased growth parameters (plant height, root length, fresh weight, dry weight, and chlorophyll content) under drought stress, alongside a reduced antioxidant capacity, as evidenced by increased levels of MDA and ROS and decreased activities of SOD, CAT, and POD. Furthermore, ZmDIR11 mutation impaired the synthesis of ABA and zeatin, downregulating key genes in the ABA (ZmABA1, ZmNCED3, ZmSnRK2.6) and zeatin (ZmIPT4, ZmCKO5, ZmCKO4b) pathways. Drought-responsive genes (ZmRD20, ZmRD22, ZmDREB2A) and lignan biosynthesis genes (Zm4CL, ZmC3H, ZmCAD1) are also downregulated in ZmDIR11 mutants. In conclusion, ZmDIR11 enhances drought tolerance in maize by regulating antioxidant capacity, ABA and zeatin synthesis, and lignan metabolism. This study provides new insights into the role of DIR genes in drought tolerance and offers a potential genetic resource for breeding drought-resistant maize varieties.
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Claims 12
Claim · model-written summary
ZmDIR11 expression increased in maize leaves under drought and PEG stress.
- Organ / stage
- leavese0
- Treatment
- drought and PEG stresse0
- Gene
- ZmDIR11e0
- Outcome
- ZmDIR11 expressione0
- Direction
- upregulatede0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“ZmDIR11 expression is upregulated in leaves under drought and PEG stress”
t0“Maize”
Claim · model-written summary
ZmDIR11 was localized to the endoplasmic reticulum.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“subcellular localization confirmed its presence in the endoplasmic reticulum”
t0“ZmDIR11”
t1“Maize”
Claim · model-written summary
Silencing or mutation of ZmDIR11 significantly reduced maize seedling drought tolerance.
- Organ / stage
- seedlingse0
- Treatment
- ZmDIR11 silencing or mutatione0
- Gene
- ZmDIR11e0
- Outcome
- drought tolerancee0
- Direction
- reducese0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“ZmDIR11 silencing or mutation significantly reduces drought tolerance in maize seedlings”
Claim · model-written summary
ZmDIR11 silencing or mutation decreased maize height, root length, and fresh weight under drought.
- Treatment
- Silencing or mutation of ZmDIR11e0
- Gene
- ZmDIR11e0
- Outcome
- plant height, root length, fresh weighte0
- Direction
- decreasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Silencing or mutation of ZmDIR11 led to decreased growth parameters (plant height, root length, fresh weight, dry weight, and chlorophyll content) under drought stress”
t0“Maize”
Claim · model-written summary
ZmDIR11 silencing or mutation decreased maize dry weight and chlorophyll content under drought.
- Treatment
- Silencing or mutation of ZmDIR11e0
- Outcome
- dry weight, and chlorophyll contente0
- Direction
- decreasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Silencing or mutation of ZmDIR11 led to decreased growth parameters (plant height, root length, fresh weight, dry weight, and chlorophyll content) under drought stress”
t0“Maize”
Claim · model-written summary
ZmDIR11 silencing or mutation increased MDA and ROS levels under drought.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Silencing or mutation of ZmDIR11 led to decreased growth parameters (plant height, root length, fresh weight, dry weight, and chlorophyll content) under drought stress, alongside a reduced antioxidant capacity, as evidenced by increased levels of MDA and ROS and decreased activities of SOD, CAT, and POD”
t0“Maize”
Claim · model-written summary
ZmDIR11 silencing or mutation decreased SOD, CAT, and POD activities under drought.
- Treatment
- Silencing or mutation of ZmDIR11e0
- Outcome
- activities of SOD, CAT, and PODe0
- Direction
- decreasede0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Silencing or mutation of ZmDIR11 led to decreased growth parameters (plant height, root length, fresh weight, dry weight, and chlorophyll content) under drought stress, alongside a reduced antioxidant capacity, as evidenced by increased levels of MDA and ROS and decreased activities of SOD, CAT, and POD”
t0“Maize”
Claim · model-written summary
ZmDIR11 mutation impaired ABA and zeatin synthesis.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“ZmDIR11 mutation impaired the synthesis of ABA and zeatin”
t0“Maize”
Claim · model-written summary
ZmDIR11 mutation downregulated ABA pathway genes.
- Treatment
- ZmDIR11 mutatione0
- Gene
- ZmABA1, ZmNCED3, ZmSnRK2.6e0
- Outcome
- key genes in the ABAe0
- Direction
- downregulatinge0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“ZmDIR11 mutation impaired the synthesis of ABA and zeatin, downregulating key genes in the ABA (ZmABA1, ZmNCED3, ZmSnRK2.6) and zeatin (ZmIPT4, ZmCKO5, ZmCKO4b) pathways”
t0“Maize”
Claim · model-written summary
ZmDIR11 mutation downregulated zeatin pathway genes.
- Treatment
- ZmDIR11 mutatione0
- Gene
- ZmIPT4, ZmCKO5, ZmCKO4be0
- Outcome
- zeatin (ZmIPT4, ZmCKO5, ZmCKO4b) pathwayse0
- Direction
- downregulatinge0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“ZmDIR11 mutation impaired the synthesis of ABA and zeatin, downregulating key genes in the ABA (ZmABA1, ZmNCED3, ZmSnRK2.6) and zeatin (ZmIPT4, ZmCKO5, ZmCKO4b) pathways”
t0“Maize”
Claim · model-written summary
Drought-responsive genes were downregulated in ZmDIR11 mutants.
- Material
- ZmDIR11 mutantse0
- Gene
- ZmRD20, ZmRD22, ZmDREB2Ae0
- Outcome
- Drought-responsive genese0
- Direction
- downregulatede0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Drought-responsive genes (ZmRD20, ZmRD22, ZmDREB2A) and lignan biosynthesis genes (Zm4CL, ZmC3H, ZmCAD1) are also downregulated in ZmDIR11 mutants”
t0“Maize”
Claim · model-written summary
Lignan biosynthesis genes were downregulated in ZmDIR11 mutants.
- Material
- ZmDIR11 mutantse0
- Gene
- Zm4CL, ZmC3H, ZmCAD1e0
- Outcome
- lignan biosynthesis genese0
- Direction
- downregulatede0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Drought-responsive genes (ZmRD20, ZmRD22, ZmDREB2A) and lignan biosynthesis genes (Zm4CL, ZmC3H, ZmCAD1) are also downregulated in ZmDIR11 mutants”
t0“Maize”
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