Paper · 2026 · Plant Physiology and Biochemistry

ZmMKK4, a mitogen-activated protein kinase kinase, influences maize kernel size

Haibo Qu; Binyan Yao; Wenhui Rao; et al.

9
claims
18
verbatim quotes
1
citations
4
research areas
✓
Open access · cc-by-nc

Abstract

Maize kernel size is regulated by multiple genetic pathways, including the evolutionarily conserved mitogen-activated protein kinase (MAPK) cascade, which plays crucial roles in plant growth and development. However, the function of MAPKK kinases in maize kernel development remains poorly understood. Here, we identified mitogen-activated protein kinase kinase 4 (ZmMKK4) as a positive regulator of kernel development in maize. ZmMKK4 is localized primarily in the nucleus and cytoplasm and shows high expression levels across multiple tissues, including kernels. Knockout mutants of zmmkk4 resulted in significantly reduced kernel size and weight, accompanied by severe impairments in kernel development. Furthermore, we observed a decrease in starch content accompanied by an increase in protein content in the mutant kernels. In summary, our study elucidates the role of ZmMKK4 in promoting kernel size and modulating starch and protein accumulation in maize, providing important insights into the molecular mechanisms underlying kernel development. These findings offer a theoretical foundation and a candidate gene for further research into maize kernel development and molecular breeding strategies. • ZmMKK4 functions as a positive regulator of maize kernel size. • The zmmkk4 gene knockout mutant has a decreased starch content and an increased protein content. • A candidate gene was provided that can be used for molecular marker-assisted selection or gene editing.

Abstract from the paper’s OpenAlex record; open copy licensed cc-by-nc. 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

Mechanismobservedoriginal resultpositivemeasured

Claim · model-written summary

ZmMKK4 was identified as a positive regulator of maize kernel development.

Taxon
maizeZea mayse0
Gene
ZmMKK4e0
Outcome
kernel developmente0
Direction
positive regulatore0

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

“Here, we identified mitogen-activated protein kinase kinase 4 (ZmMKK4) as a positive regulator of kernel development in maize.”

✓ located verbatim in the source · abstract, characters 302–428 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

ZmMKK4 localized primarily to the nucleus and cytoplasm.

Taxon
maizeZea mayst0
Gene
ZmMKK4e0
Outcome
localized primarily in the nucleus and cytoplasme0

Not reportedDirection

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

“ZmMKK4 is localized primarily in the nucleus and cytoplasm and shows high expression levels across multiple tissues, including kernels.”

✓ located verbatim in the source · abstract, characters 429–564 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 62–67 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

ZmMKK4 was highly expressed across multiple tissues including kernels.

Taxon
maizeZea mayst0
Organ / stage
kernelse0
Gene
ZmMKK4e0
Outcome
expression levelse0
Direction
highe0

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

“ZmMKK4 is localized primarily in the nucleus and cytoplasm and shows high expression levels across multiple tissues, including kernels.”

✓ located verbatim in the source · abstract, characters 429–564 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 62–67 · show in text ↑
Intervention effectobservedoriginal resultnegativesignificant (as stated)measured

Claim · model-written summary

zmmkk4 knockout reduced or impaired kernel size.

Taxon
maizeZea mayst0
Treatment
Knockout mutants of zmmkk4e0
Gene
zmmkk4e0
Outcome
kernel sizee0
Direction
reducede0

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

“Knockout mutants of zmmkk4 resulted in significantly reduced kernel size and weight, accompanied by severe impairments in kernel development.”

✓ located verbatim in the source · abstract, characters 565–706 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 62–67 · show in text ↑
Intervention effectobservedoriginal resultnegativesignificant (as stated)measured

Claim · model-written summary

zmmkk4 knockout reduced or impaired weight.

Taxon
maizeZea mayst0
Treatment
Knockout mutants of zmmkk4e0
Gene
zmmkk4e0
Outcome
weighte0
Direction
reducede0

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

“Knockout mutants of zmmkk4 resulted in significantly reduced kernel size and weight, accompanied by severe impairments in kernel development.”

✓ located verbatim in the source · abstract, characters 565–706 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 62–67 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

zmmkk4 knockout reduced or impaired kernel development.

Taxon
maizeZea mayst0
Treatment
Knockout mutants of zmmkk4e0
Gene
zmmkk4e0
Outcome
kernel developmente0
Direction
impairmentse0

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

“Knockout mutants of zmmkk4 resulted in significantly reduced kernel size and weight, accompanied by severe impairments in kernel development.”

✓ located verbatim in the source · abstract, characters 565–706 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 62–67 · show in text ↑
Observationobservedoriginal resultnegativemeasured

Claim · model-written summary

Mutant kernels showed a decrease in starch content.

Taxon
maizeZea mayst0
Material
mutant kernelse0
Outcome
starch contente0
Direction
decreasee0

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

“Furthermore, we observed a decrease in starch content accompanied by an increase in protein content in the mutant kernels.”

✓ located verbatim in the source · abstract, characters 707–829 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 62–67 · show in text ↑
Observationobservedoriginal resultmeasured

Claim · model-written summary

Mutant kernels showed a increase in protein content.

Taxon
maizeZea mayst0
Material
mutant kernelse0
Outcome
protein contente0
Direction
increasee0

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

“Furthermore, we observed a decrease in starch content accompanied by an increase in protein content in the mutant kernels.”

✓ located verbatim in the source · abstract, characters 707–829 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 62–67 · show in text ↑
Recommendationauthor interpretationoriginal resultbasis n/a

Claim · model-written summary

The authors present ZmMKK4 as a candidate for marker-assisted selection or gene editing.

Taxon
maizeZea mayst0
Gene
ZmMKK4t1
Outcome
candidate genee0

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 candidate gene was provided that can be used for molecular marker-assisted selection or gene editing.”

✓ located verbatim in the source · abstract, characters 1373–1476 · show in text ↑
t0

“maize”

✓ located verbatim in the source · title, characters 62–67 · show in text ↑
t1

“ZmMKK4”

✓ located verbatim in the source · title, characters 0–6 · show in text ↑