The significant rise in soil salinity has had detrimental effects on global agricultural production, negatively impacting overall plant health and leading to a decline in productivity. As a protective response, plants have developed diverse regulatory mechanisms to counteract these adverse conditions. The mechanisms help mitigate damage caused by both osmotic and ionic stress resulting from high salinity. Given the severe threat this poses to global food security and the well-being of the world's population, scientists have dedicated decades of research to understanding how to manage salt stress. Numerous mechanisms have been identified and studied to enhance plant salt tolerance and alleviate the damage caused by salt stress. This review examines recent advancements in molecular regulatory mechanisms underlying plant salt, including salt uptake and transport, salt sensing and signalling, hormonal regulation, epigenetic modifications, genetic adaptation, and posttranslational modifications. Although current knowledge has advanced our understanding, critical gaps and controversies remain, such as the stability of epigenetic memory, the trade-off between stress tolerance and growth, hormonal crosstalk, and novel genes with uncharacterised roles in salt tolerance. To resolve these questions, further research employing techniques like GWAS, transcriptomics, transgenic and genome-editing technologies, as well as studies on energy allocation and hormonal regulation, is essential. A deeper exploration of these complex, synergistic mechanisms will pave the way for enhancing plant resilience and ensuring adaptation to increasingly challenging environmental conditions.
Among various abiotic stresses, secondary soil salinization poses a significant threat to plant productivity and survival. Cultivated chrysanthemums (Chrysanthemum morifolium), widely grown as ornamental crops, are highly susceptible to salt stress, and their complex polyploid genome complicates the identification of stress resistance genes. In contrast, C. indicum, a native diploid species with robust stress tolerance, serves as a valuable genetic resource for uncovering stress-responsive genes and improving the resilience of ornamental chrysanthemum cultivars. In this study, we cloned, overexpressed (OE-CiHY5), and silenced (RNAi-CiHY5) the CiHY5 gene in C. indicum. OE-CiHY5 plants exhibited larger leaves, sturdier stalks, and higher chlorophyll content compared to wild-type plants, while RNAi-CiHY5 plants displayed weaker growth. Under salt stress, OE-CiHY5 plants demonstrated significantly improved growth, enhanced osmotic adjustment, and effective ROS scavenging. In contrast, RNAi-CiHY5 plants were more sensitive to salinity, showing higher electrolyte leakage and impaired osmotic regulation. Transcriptomic analyses revealed that CiHY5 regulates key hormonal pathways such as zeatin (one of cytokinins), abscisic acid and jasmonic acid, as well as metabolic pathways, including photosynthesis, carbohydrate metabolism, which collectively contribute to the enhanced stress resilience of OE-CiHY5 plants. Promoter-binding assays further confirmed that CiHY5 directly interacts with the CiABF3 promoter, highlighting its critical role in ABA signaling. Evolutionary analyses showed that HY5 is conserved across plant lineages, from early algae to advanced angiosperms, with consistent responsiveness to salt and other abiotic stresses in multiple Chrysanthemum species. These findings establish CiHY5 as a key regulator of salt tolerance in C. indicum, orchestrating a complex network of hormonal and metabolic pathways to mitigate salinity-induced damage. Given the conserved nature of HY5 and its responsiveness to various stresses, HY5 gene provides valuable insights into the molecular mechanisms underlying stress adaptation and represents a promising genetic target for enhancing salt stress resilience in chrysanthemums.
Background Cotton is a vital economic crop and reserve material and a pioneer crop planted on saline-alkaline soil. Improving the tolerance of cotton to saline alkaline environments is particularly important. Results Salt-tolerant and salt-sensitive cotton plants at the three-leaf stage were subjected to 200 mM NaCl stress treatment, thereafter, microstructural observations beside physiological and biochemical analyses were performed on cotton leaves at 0 h (CK), 48 h (NaCl) and re-watering (RW) for 48 h. Salt stress altered microstructural observations and physiological and biochemical in ST and SS (p < 0.05). After re-watering, ST recovered fully, while SS sustained permanent oxidative and structural damage, indicating distinct salt tolerance. Transcriptome analysis was performed on cotton leaves under salt stress and re-watering conditions. KEGG analysis revealed that the response of cotton to salt stress and its adaptation to re-watering may be related to major protein families such as photosynthesis (ko 00195), photosynthesis-antenna protein (ko 00196), plant hormone signal transduction (ko 04075), starch and sucrose metabolism (ko 00500), and porphyrin and chlorophyll metabolism (ko 00860). A gray coexpression module associated with cotton restoration under salt stress was enriched according to WGCNA. Conclusions Salt stress did not only affect the physiological and biochemical levels of cotton but also induced structural changes in cells and tissues. Re-watering was relatively effective in stabilizing the physiological and biochemical parameters, as well as the leaf microstructure, of cotton plants under salt stress. WGCNA revealed enriched gray coexpression modules related to the recovery of cotton plants under salt stress, and screening of the pivotal genes in the gray module revealed five critical hubs, namely, GH_A01G1528, GH_A08G2688, GH_D08G2683, GH_D01G1620 and GH_A10G0617. Overall, our findings can provide new insights into enhancing cotton salt tolerance and exploring salt tolerance genes in cotton,including screening cotton genetic resources using those potential responsive genes. This study provides a theoretical basis for further exploration of the molecular mechanism of cotton salt tolerance and genetic resources for breeding salt-tolerant cotton.
Soybean, a globally significant and versatile crop, serves as a vital source of both oil and protein. However, environmental factors such as soil salinization pose substantial challenges to its cultivation, adversely affecting both yield and quality. Enhancing the salt tolerance of soybeans can mitigate yield losses and promote the development of the soybean industry. Members of the plant-specific transcription factor family NAC play crucial roles in plant adaptation to abiotic stress conditions. In this study, we screened the soybean GmNAC family genes potentially involved in the salt stress response and identified 18 GmNAC genes that may function during the early stages of salt stress. Among these, the GmNAC035 gene exhibited a rapid increase in expression within one hour of salt treatment, with its expression being induced by abscisic acid (ABA) and methyl jasmonate (MeJA), suggesting its significant role in the soybean salt stress response. We further elucidated the role of GmNAC035 in soybean salt tolerance. GmNAC035, a nuclear-localized transcriptional activator, enhances salt tolerance when overexpressed in Arabidopsis, reducing oxidative damage and boosting the expression of stress-responsive genes. It achieves this by regulating key stress response pathways, including the SOS pathway, calcium signaling, and ABA signaling. These findings highlight the potential of GmNAC035 as a genetic engineering target to improve crop salt tolerance.
Soil salinization is a severe environmental issue limiting the growth and yield of crops worldwide. Subsurface drip irrigation with micro-nano bubble hydrogen water (SDH) is an innovative way to realize the role of hydrogen gas (H2) in improving plant resistance to salt stress in practical agricultural productions. Nonetheless, limited information is available on how SDH affects the plant salt tolerance performance. Especially, the underlying physiological respond, hormone-regulated and soil microbial-mediated mechanisms have not been reported so far. In this study, the effects of SDH on lettuce (Lactuca sativa L.) growth, photosynthesis, root development, antioxidant system, phytohormone, and soil microbial community were investigated under normal and salt stress conditions. The results showed that, with salt stress, SDH significantly enhanced the lettuce fresh weight, photosynthesis activity, and root growth. The leaf antioxidant enzyme activities increased and reactive oxygen species (ROS) content decreased to reduce the oxidative damage. The decreased malondialdehyde (MDA) content indicated a low membrane lipid peroxidation responsible for cellular damage. SDH also helped to maintain osmotic homeostasis, which was reflected by the increased soluble protein (SP) content. Reduced Na+/ K+ ratio and ROS did not trigger excessive production of stress response hormones abscisic acid (ABA) and jasmonic acid (JA), which alleviated the mediated growth inhibition effects. SDH enriched the abundance of the plant growth-promoting rhizobacteria (PGPR) in the soil, such as Arthrobacter and Pseudomonas. That might be the reason for explaining the increase in hormone indoleacetic acid (IAA) in lettuce and 1-aminocyclopropane-1carboxylate (ACC) deaminase activity in the soil, which was beneficial for inhibiting ethylene production and promoting plant growth. Under the normal condition, variations of physiological and growth indicators as affected by SDH were similar to those under the salt stress condition, except for root development. High concentration of dissolved hydrogen gas in water might expel the oxygen. The induced soil anoxic environment limited oxygen diffusion, in turn inhibited root respiration and growth. The effect of hydrogen concentration on the plant tolerance against salt stress under different salt content could be further studied.
Chrysanthemum, a valuable ornamental flower, has limited salinity tolerance, which restricts its cultivation in salt-stressed conditions. In this study, we investigated the salt tolerance of a population derived from the salttolerant germplasm Chrysanthemum yantaiense. The parents and 91 offspring were subjected to 300 mM NaCl concentrations for 30 days. Based on the observed changes in growth and the degree of damage caused by salt stress, 15 high-resistant, 52 moderate-resistant, and 16 low-resistant strains were identified. Two offspring (i.e., YS-58 and YS-123) with contrasting salt tolerance were subjected to 15 days of salt stress, with phenotypic, physiological, and biochemical responses assessed at 5, 10, and 15 days. YS-58 demonstrated greater resilience, maintaining higher shoot fresh weight by day 10, and exhibiting significantly less growth impairment in both aboveground and belowground by day 15 compared to YS-123. Under salt stress, YS-58 accumulated lower Na* levels in leaves, while sustaining higher K* content in roots and stems. Additionally, YS-58 showed elevated proline levels, reduced soluble sugar content, and decreased malondialdehyde (MDA) accumulation, along with enhanced superoxide dismutase (SOD) activity relative to YS-123. Understanding these mechanisms will provide insights into how chrysanthemums survive under saline conditions, potentially enabling large-scale cultivation in saline soils.
Chloride (Cl-) ions cause major damage to crops in saline soils. Understanding the key factors that influence Cl- uptake and translocation will aid the breeding of more salt-tolerant crops. Here, using genome-wide association study and transcriptomic analysis, we identified a NITRATE TRANSPORTER 1 (NRT1)/PEPTIDE TRANSPORTER family (NPF) protein, GmNPF7.5, as the dominant gene locus influencing Cl- homeostasis in soybean (Glycine max). A natural SNP variation resulted in two haplotypes (GmNPF7.5HapA and GmNPF7.5HapB), which was associated with Cl- content. GmNPF7.5HapA mediated Cl- or nitrate (NO3-) uptake in a pH-dependent manner and exhibited higher permeability for Cl- over NO3-. The suppression of GmNPF7.5HapA expression decreased Cl- accumulation and salt damage in plants, whereas its overexpression showed the opposite effects. The elite haplotype GmNPF7.5HapB diminished Cl- transport activity independently from NO3- permeability, thus enhancing soybean salt tolerance. Furthermore, the protein kinase GmPI4K gamma 4 could phosphorylate GmNPF7.5, which repressed Cl- uptake without affecting NO3- permeability. Our findings define a regulatory mechanism for Cl- control under NaCl stress, providing a strategy for the improvement of salt tolerance in soybean plants.
Rising soil salinity poses significant challenges to Mediterranean viticulture. While some rootstocks effectively reduce salt accumulation in grafted scions, the mechanisms and performance of novel rootstocks remain largely unexplored. This study compared two novel M-series rootstocks (M2, M4) with established commercial rootstocks (1103 Paulsen, R110) to evaluate their physiological responses and salt tolerance under irrigation with varying salinity levels (0, 25, 50, and 75 mM NaCl) over 5 months. Growth parameters, photosynthetic efficiency, chlorophyll content (SPAD), ion homeostasis, and visual symptoms were monitored. Results revealed genotype-specific strategies: 1103 Paulsen exhibited robust photosynthetic efficiency and ion exclusion, maintaining growth and chlorophyll stability; M2 demonstrated superior biomass retention and moderate ion compartmentalization but showed reduced photosynthetic performance at higher salinity levels; R110 displayed effective ion management at moderate salinity but experienced significant growth reduction under severe stress; and M4 was the most sensitive, with severe reductions in growth and ion homeostasis. Organ-specific responses highlighted roots acting as primary ion reservoirs, particularly for sodium and calcium; leaves exhibited high potassium and chloride concentrations, critical for photosynthesis but prone to ionic imbalance under stress; and stems and wood played a buffering role, compartmentalizing excess sodium and minimizing damage to photosynthetic tissues. The reported findings provide valuable insights for rootstock selection and breeding programs, particularly for regions facing increasing soil and water salinization challenges.
Introduction Salt stress has emerged as a predominant abiotic factor that jeopardizes global crop growth and yield. The plant hormone salicylic acid (SA) has notable potential in mitigating salt toxicity, yet its mechanism in enhancing the salinity tolerance of tobacco plants is not well explored. Methods This study aimed to assess the potential benefits of exogenous SA application (1.0 mM) on tobacco seedlings subjected to saline soil conditions. Results The foliar spray of SA partially mitigated these salt-induced effects, as evidenced by a reduction of malondialdehyde content, and improvements of leaf K+/Na+ ratios, pigment biosynthesis, and electron transport efficiency under NaCl stress. Additionally, SA increased the contents of total phenolic compound and soluble protein by 16.2% and 28.7% to alleviate NaCl-induced oxidative damage. Under salt stressed conditions, the activities of antioxidant enzymes, including superoxide dismutase, ascorbate peroxidase, catalase, and peroxidase increased by 4.2%similar to 14.4% in SA sprayed tobacco seedlings. Exogenous SA also increased ascorbate and glutathione levels and reduced their reduced forms by increasing the activities of glutathione reductase, ascorbate peroxidase, monodehydroascorbate reductase and dehydroascorbate reductase. qRT-PCR analysis revealed that the key genes regulating SA biosynthesis, carbon assimilation, the antioxidant system and the ascorbate-glutathione cycle were activated by SA under conditions of salt stress. Discussion Our study elucidates the physiological and molecular mechanisms of exogenous SA in enhancing plant salt tolerance and provides a practical basis for crop improvement in saline environments.
Soil salinity is a major global challenge affecting agricultural productivity and food security. This study explores innovative strategies to improve salt tolerance in soybean (Glycine max), a crucial crop in the global food supply. This study investigates the synergistic effects of S-nitroso glutathione (GSNO) and silicon on enhancing salt tolerance in soybean (Glycine max). Two soybean cultivars, Seonpung (salt-tolerant) and Cheongja (salt-sensitive), were analyzed for various physiological, biochemical, and genetic traits under salt stress. The results showed that the combined GSNO and Si treatment significantly improved several key traits, including plant height, relative water content, root development, nodule numbers, chlorophyll content, and stomatal aperture, under both control and salt stress conditions. Additionally, this treatment optimized ion homeostasis by enhancing the Na/K ratio and Ca content, while reducing damage markers such as electrolyte leakage, malondialdehyde, and hydrogen peroxide. The stress-responsive compounds, including proline, ascorbate peroxidase, and water-soluble proteins, were elevated under stress conditions, indicating improved tolerance. Gene expression analysis revealed significant upregulation of genes such as GmNHX1, GmSOS2, and GmAKT1, associated with salt stress response, while GmNIP2.1, GmNIP2.2, and GmLBR were downregulated in both varieties. Notably, the salt-sensitive variety Cheongja exhibited higher electrolyte leakage and oxidative damage compared to the salt-tolerant Seonpung. These findings suggest that the combination of GSNO and silicon enhances salt tolerance in soybean by improving physiological resilience, ion homeostasis, and stress-responsive gene expression.