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Volume 14
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10.3390/plants14233658
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Article

Cloning and Functional Verification of Salt Tolerance Gene HbNHX2 in Hordeum brevisubulatum

by
Mingzhi Zhang
†,
Mei Yang
†,
Wenjie Zhao
,
Hang Yin
*,
Xinyi Zhang
,
Bing Li
,
Muzhapaer Tuluhong
,
Baiji Wang
,
Shanshui Zheng
and
Guowen Cui
*
College of Animal Science and Technology, Northeast Agricultural University, Harbin 150030, China
*
Authors to whom correspondence should be addressed.
†
These authors contributed equally to this work.
Plants 2025, 14(23), 3658; https://doi.org/10.3390/plants14233658 (registering DOI)
Submission received: 29 October 2025 / Revised: 27 November 2025 / Accepted: 28 November 2025 / Published: 30 November 2025
(This article belongs to the Section Plant Genetics, Genomics and Biotechnology)
Versions Notes

Abstract

A high salt environment seriously affects the physiological metabolism and yield of plants. Hordeum brevisubulatum (Trin.) Link has high biomass and important ecological, feeding and economic values, but its growth conditions have serious saline-alkali effects. The NHX gene family plays a vital role in regulating intracellular Na+/K+ balance, pH homeostasis, and vesicle and protein transport in plants. In this study, the HbNHX2 gene was cloned from Hordeum brevisubulatum and functionally characterized through phenotypic, physiological, and molecular analyses in transgenic tobacco. Expression profiling revealed that HbNHX2 was most abundant in spikes and least abundant in root tips, and the expression level was significantly induced under salt stress. Overexpression of HbNHX2 led to decreased malondialdehyde (MDA) and superoxide anion (O2−) levels, while it enhanced the activities of superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT). Additionally, the levels of glutathione (GSH), soluble proteins, proline, and chlorophyll were also increased. Several stress-responsive genes, including CBL1, ERF2, BI-1, Cu/Zn-SOD, Mn-SOD, POD, GR1, KC1, TPK1, TIP, P5CS, BAS1, STN7 and LTP1, were significantly upregulated, while SERK3B was downregulated. These findings suggest that HbNHX2 enhances plant salt tolerance by maintaining osmotic balance, scavenging reactive oxygen species (ROS), and regulating stress-responsive gene expression. This study provides new insights into the molecular mechanism of salt tolerance in Hordeum brevisubulatum and lays a foundation for breeding salt-tolerant forage crops.
Keywords: Hordeum brevisubulatum; membrane-bound Na+/H+ antiporters; HbNHX2 gene; physiological mechanism; salt stress Hordeum brevisubulatum; membrane-bound Na+/H+ antiporters; HbNHX2 gene; physiological mechanism; salt stress

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MDPI and ACS Style

Zhang, M.; Yang, M.; Zhao, W.; Yin, H.; Zhang, X.; Li, B.; Tuluhong, M.; Wang, B.; Zheng, S.; Cui, G. Cloning and Functional Verification of Salt Tolerance Gene HbNHX2 in Hordeum brevisubulatum. Plants 2025, 14, 3658. https://doi.org/10.3390/plants14233658

AMA Style

Zhang M, Yang M, Zhao W, Yin H, Zhang X, Li B, Tuluhong M, Wang B, Zheng S, Cui G. Cloning and Functional Verification of Salt Tolerance Gene HbNHX2 in Hordeum brevisubulatum. Plants. 2025; 14(23):3658. https://doi.org/10.3390/plants14233658

Chicago/Turabian Style

Zhang, Mingzhi, Mei Yang, Wenjie Zhao, Hang Yin, Xinyi Zhang, Bing Li, Muzhapaer Tuluhong, Baiji Wang, Shanshui Zheng, and Guowen Cui. 2025. "Cloning and Functional Verification of Salt Tolerance Gene HbNHX2 in Hordeum brevisubulatum" Plants 14, no. 23: 3658. https://doi.org/10.3390/plants14233658

APA Style

Zhang, M., Yang, M., Zhao, W., Yin, H., Zhang, X., Li, B., Tuluhong, M., Wang, B., Zheng, S., & Cui, G. (2025). Cloning and Functional Verification of Salt Tolerance Gene HbNHX2 in Hordeum brevisubulatum. Plants, 14(23), 3658. https://doi.org/10.3390/plants14233658

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