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Salt glands in exo-recretohalophytes: Development, physiological functions, and prospects for improving crop salt tolerance

Limin Wang1†, Junyan Xie1†, Yiping Zou1†, Chunliang Yao1†, Hai Fan1†, Chenqi Shen1, Wenyan Zhou1, Jingran Qin1, Xinke Zhang1, Baoshan Wang1, Jian Zhang1* and Guoliang Han1,2*   

  1. 1. Shandong Provincial Key Laboratory of Stress Plant Biology and Genetic Improvement, College of Life Sciences, Shandong Normal University, Jinan 250014, China;
    2. Dongying Institute, Shandong Normal University, Dongying 257000, China;
    †These authors contributed equally to this work.;
    *Correspondences: Guoliang Han (gl_han@sdnu.edu.cn, Dr. Han is fully responsible for the distribution of all materials associated with this article); Jian Zhang (zhangjian@sdnu.edu.cn)
  • Received:2025-10-26 Accepted:2025-12-23 Online:2026-01-19
  • Supported by:
    This work was supported by the National Natural Science Research Foundation of China (project no. U24A20613 and 32000209), the Natural Science Research Foundation of Shandong Province (project no. ZR2023YQ021), the Taishan Scholar project of Shandong province (project no. tsqn202408335 and tsqnz20240810), and the Key R&D Program of Shandong Province, China (project no.2024SFGC0404).

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Abstract: In exo-recretohalophytes, specialized structures known as salt glands secrete excess salt ions from plant tissues, thereby maintaining intracellular ion homeostasis and sustaining normal metabolism under salt stress. Based on their cellular composition, salt glands can be unicellular, bicellular, or multicellular, and they originate from undifferentiated precursor cells known as multipotent epidermal stem cells. A complex regulatory network drives the division and differentiation of these cells into functional salt-secreting structures. Three hypotheses have been proposed to explain the physiological mechanisms underlying salt secretion by salt glands, each supported by experimental evidence: The osmotic mechanism, the reverse pinocytosis mechanism, and the animal-like fluid transport mechanism. This review summarizes the structural characteristics, developmental processes, salt secretion mechanisms, and potential applications of salt glands in exo-recretohalophytes, providing a foundation for future studies on salt gland biology and their utilization in developing salt-tolerant crops.

Key words: exo-recretohalophyte, regulatory network, salt gland, salt secretion, salt stress

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