Abstract
The exclusion of sodium ions (Na+) from the shoot tissue, termed shoot Na+ exclusion, underlies a core mechanism of crop salt tolerance. Recent studies have shown that the HAK (High-Affinity K+ Transporter) family Na+ transporters play a key role in shoot Na+ exclusion of various crops, however, it is unknown whether and how this type of transporter is post-transcriptionally regulated. Here, we show that two closely related SnRK2 kinases, designated as ZmSnRK2.9 and ZmSnRK2.10, promote shoot Na+ exclusion and salt tolerance by activating the Na+ transporter ZmHAK4 in maize. Under salt conditions, the kinase activity of ZmSnRK2.9 and ZmSnRK2.10 is activated, then they interact with and phosphorylate ZmHAK4 at Ser5, increasing the Na+ transport activity of ZmHAK4, which in turn promotes salt tolerance by improving the exclusion of Na+ from the shoot tissue. Furthermore, we show that a 20-bp deletion that occurred naturally in the ZmSnRK2.10 promoter decreases its transcript level, resulting in an increased shoot Na+ content under salt conditions. Our findings support a breeding program that can utilize the favorable alleles of ZmHAK4 and ZmSnRK2.10 to enhance both the transcriptional and post-transcriptional activation of ZmHAK4, thus advancing the development of salt-tolerant maize. The authors demonstrated that maize SnRK2 kinases (ZmSnRK2.9/2.10) phosphorylate ZmHAK4, reducing shoot sodium and enhancing salt tolerance, and identified a favorable allele of ZmSnRK2.10 that provides a target for breeding salt-tolerant maize.
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Claims 7
Claim · model-written summary
ZmSnRK2.9 and ZmSnRK2.10 promote shoot sodium exclusion and salt tolerance through ZmHAK4.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“two closely related SnRK2 kinases, designated as ZmSnRK2.9 and ZmSnRK2.10, promote shoot Na+ exclusion and salt tolerance by activating the Na+ transporter ZmHAK4 in maize.”
Claim · model-written summary
Salt conditions activate both kinase activities.
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“Under salt conditions, the kinase activity of ZmSnRK2.9 and ZmSnRK2.10 is activated”
t0“maize”
Claim · model-written summary
The kinases interact with and phosphorylate ZmHAK4 at Ser5.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“they interact with and phosphorylate ZmHAK4 at Ser5”
t0“maize”
Claim · model-written summary
Phosphorylation increases ZmHAK4 sodium transport activity.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“increasing the Na+ transport activity of ZmHAK4”
t0“maize”
Claim · model-written summary
A natural 20-bp promoter deletion lowers ZmSnRK2.10 transcript level.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“a 20-bp deletion that occurred naturally in the ZmSnRK2.10 promoter decreases its transcript level”
t0“maize”
Claim · model-written summary
The promoter deletion increases shoot sodium content under salt conditions.
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“a 20-bp deletion that occurred naturally in the ZmSnRK2.10 promoter decreases its transcript level, resulting in an increased shoot Na+ content under salt conditions.”
t0“maize”
Claim · model-written summary
The authors recommend favorable ZmHAK4 and ZmSnRK2.10 alleles for salt-tolerant maize breeding.
- Outcome
- utilize the favorable alleles of ZmHAK4 and ZmSnRK2.10e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e0“utilize the favorable alleles of ZmHAK4 and ZmSnRK2.10 to enhance both the transcriptional and post-transcriptional activation of ZmHAK4”
t0“maize”
The same questions in other papers
salinity · physiology: enzyme activity · Zea mays (maize)Positive and negative results
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