Root-specific upregulation of the Na+/K+ transport genes mitigates salt stress in blueberry

doi:10.1016/j.jia.2025.12.004

Journal of Integrative Agriculture

2026, Vol. 25

Issue (8): 3240-3253 DOI: 10.1016/j.jia.2025.12.004

Horticulture

Advanced Online Publication | Current Issue | Archive | Adv Search

Root-specific upregulation of the Na⁺/K⁺ transport genes mitigates salt stress in blueberry

Huifang Song^1*, Bingshuai Du^1*, Xinyan Zhao¹, Kaiyue Feng¹, Lingyun Zhang^{1, 2#}, Yibo Cao^{1, 2#}

¹State Key Laboratory of Efficient Production of Forest Resources/Key Laboratory of Forest Silviculture and Conservation of the Ministry of Education/College of Forestry, Beijing Forestry University, Beijing 100083, China

²Research & Development Center of Blueberry, Beijing Forestry University, Beijing 100083, China

Highlights
● Ion transport pathways were enriched under salt stress, and root-upregulated Na⁺/K⁺ transport genes promoted blueberry salt tolerance.
● Leaf Na⁺ and K⁺ contents strongly correlated with VcHAK5, VcNHX1, and VcNHX2 expression.
● Knockdown of VcHAK5, VcNHX1, and VcNHX2 increased salt sensitivity in blueberry calli.
● Reciprocal grafting showed that root Na⁺/K⁺ homeostasis is critical for salt tolerance.

Abstract
References

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摘要

土壤盐渍化日益成为限制高丛蓝莓（Vaccinium corymbosum）生产的重要因素，但其在盐胁迫下的生理与分子应答机制仍缺乏系统认识。为此，本研究以耐盐品种‘Duke’与盐敏感品种‘Sweetheart’的根和叶为材料，对差异表达基因进行了基因功能富集（GO）分析和加权基因共表达网络分析（WGCNA）。GO分析结果表明，盐胁迫下根部的离子转运及细胞离子稳态相关通路显著富集。WGCNA 结果显示，Na⁺与K⁺含量与核心基因 VcHAK5、VcNHX1、VcNHX2 的表达呈高度相关，这些基因在耐盐品种根部显著上调。功能上，VcHAK5 编码质膜上的K⁺选择性转运蛋白，VcNHX1 和 VcNHX2 编码液泡膜上的Na⁺/K⁺-H⁺ 逆向转运蛋白。在蓝莓愈伤组织中分别沉默这些基因，转基因愈伤表现为盐敏感。此外，耐盐差异品种间的互换枕木嫁接试验表明，降低根部Na⁺含量及Na⁺/K⁺比值是实现耐盐性的关键。本研究首次系统揭示了蓝莓对盐胁迫的响应机制，鉴定了耐盐相关的关键靶基因，并强调了耐盐根系在整体耐盐性形成中的重要作用。

Abstract

Soil salinization is a growing challenge for highbush blueberry (Vaccinium corymbosum) production, but knowledge about its physiological and molecular responses to salt stress remains limited. To address this, we performed Gene Ontology (GO) enrichment analysis and weighted gene co-expression network analysis (WGCNA) on differentially expressed genes from the roots and leaves of the salt-tolerant cultivar ‘Duke’ and the salt-sensitive cultivar ‘Sweetheart’. GO analysis revealed significant enrichment of ion transport and cellular ion homeostasis in the roots under salt stress. WGCNA identified a strong correlation between Na⁺ and K⁺ contents and the expression of hub genes VcHAK5, VcNHX1, and VcNHX2 under salt stress. These genes were significantly upregulated in the roots of the salt-tolerant cultivar. VcHAK5 encodes a K⁺-selective ion transporter in the plasma membrane, and VcNHX1 and VcNHX2 encode Na⁺/H⁺ and K⁺/H⁺ antiporters in the tonoplast. Knockdown mutants of these genes in blueberry calli showed hypersensitivity to salt stress. Furthermore, reciprocal grafting between salt-sensitive and salt-tolerant blueberry cultivars demonstrated that lower root Na⁺ content and Na⁺/K⁺ ratios are crucial for salt tolerance. This study provides the first comprehensive insights into blueberry responses to salt stress, identifies target genes and highlights the critical role of salt-resistant roots.

Keywords: blueberry soil salinization salt tolerance ion transport grafting

Received: 27 March 2025 Accepted: 25 September 2025 Online: 08 December 2025

Fund:

This work was supported by the Key Research and Development Plan of Shandong Province, China (2024LZGCQY025),
the National Natural Science Foundation of China (32101555), and the 5·5 Engineering Research & Innovation Team Project of Beijing Forestry University (BLRC2023B08).

About author: Huifang Song, E-mail: songhuifang1010@bjfu.edu.cn; #Correspondence Yibo Cao, E-mail: caoyibo@bjfu.edu.cn; Lingyun Zhang, Tel: +86-10-62336044, E-mail: lyzhang@bjfu.edu.cn *These authors contributed equally to this study.

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Huifang Song, Bingshuai Du, Xinyan Zhao, Kaiyue Feng, Lingyun Zhang, Yibo Cao. 2026. Root-specific upregulation of the Na⁺/K⁺ transport genes mitigates salt stress in blueberry. Journal of Integrative Agriculture, 25(8): 3240-3253.

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