Soil salinization is a severe abiotic stress that negatively affects plant growth and development, leading to physiological abnormalities and ultimately threatening global food security. The condition arises from excessive salt accumulation in the soil, primarily due to anthropogenic activities such as irrigation, improper land uses, and overfertilization. The presence of Na⁺, Cl⁻, and other related ions in the soil above normal levels can disrupt plant cellular functions and lead to alterations in essential metabolic processes such as seed germination and photosynthesis, causing severe damage to plant tissues and even plant death in the worst circumstances. To counteract the effects of salt stress, plants have developed various mechanisms, including modulating ion homeostasis, ion compartmentalization and export, and the biosynthesis of osmoprotectants. Recent advances in genomic and proteomic technologies have enabled the identification of genes and proteins involved in plant salt-tolerance mechanisms. This review provides a short overview of the impact of salinity stress on plants and the underlying mechanisms of salt-stress tolerance, particularly the functions of salt-stress-responsive genes associated with these mechanisms. This review aims at summarizing recent advances in our understanding of salt-stress tolerance mechanisms, providing the key background knowledge for improving crops’ salt tolerance, which could contribute to the yield and quality enhancement in major crops grown under saline conditions or in arid and semiarid regions of the world.

Climate change has increased the overall impact of abiotic stress conditions such as drought, salinity, and extreme temperatures on plants. Abiotic stress adversely affects the growth, development, crop yield, and productivity of plants. When plants are subjected to various environmental stress conditions, the balance between the production of reactive oxygen species and its detoxification through antioxidant mechanisms is disturbed. The extent of disturbance depends on the severity, intensity, and duration of abiotic stress. The equilibrium between the production and elimination of reactive oxygen species is maintained due to both enzymatic and non-enzymatic antioxidative defense mechanisms. Non-enzymatic antioxidants include both lipid-soluble (α-tocopherol and β-carotene) and water-soluble (glutathione, ascorbate, etc.) antioxidants. Ascorbate peroxidase (APX), superoxide dismutase (SOD), catalase (CAT), and glutathione reductase (GR) are major enzymatic antioxidants that are essential for ROS homeostasis. In this review, we intend to discuss various antioxidative defense approaches used to improve abiotic stress tolerance in plants and the mechanism of action of the genes or enzymes involved.

The Arabidopsis gene AtNHX1 encodes a vacuolar membrane-bound sodium/proton (Na⁺/H⁺) antiporter that transports Na⁺ into the vacuole and exports H⁺ into the cytoplasm. The Arabidopsis gene SOS1 encodes a plasma membrane-bound Na⁺/H⁺ antiporter that exports Na⁺ to the extracellular space and imports H⁺ into the plant cell. Plants rely on these enzymes either to keep Na⁺ out of the cell or to sequester Na⁺ into vacuoles to avoid the toxic level of Na⁺ in the cytoplasm. Overexpression of AtNHX1 or SOS1 could improve salt tolerance in transgenic plants, but the improved salt tolerance is limited. NaCl at concentration >200 mM would kill AtNHX1-overexpressing or SOS1-overexpressing plants. Here it is shown that co-overexpressing AtNHX1 and SOS1 could further improve salt tolerance in transgenic Arabidopsis plants, making transgenic Arabidopsis able to tolerate up to 250 mM NaCl treatment. Furthermore, co-overexpression of AtNHX1 and SOS1 could significantly reduce yield loss caused by the combined stresses of heat and salt, confirming the hypothesis that stacked overexpression of two genes could substantially improve tolerance against multiple stresses. This research serves as a proof of concept for improving salt tolerance in other plants including crops.

Abiotic stresses are major limiting factors that pose severe threats to agricultural production. Conventional breeding has significantly improved crop productivity in the last century, but traditional breeding has reached its maximum capacity due to the multigenic nature of abiotic stresses. Alternatively, biotechnological approaches could provide new opportunities for producing crops that can adapt to the fast-changing environment and still produce high yields under severe environmental stress conditions. Many stress-related genes have been identified and manipulated to generate stress-tolerant plants in the past decades, which could lead to further increase in food production in most countries of the world. This review focuses on the recent progress in using transgenic technology and gene editing technology to improve abiotic stress tolerance in plants, and highlights the potential of using genetic engineering to secure food and fiber supply in a world with an increasing population yet decreasing land and water availability for food production and fast-changing climate that will be largely hostile to agriculture.

Protein phosphatase 2A (PP2A) is an enzyme consisting of three subunits: a scaffolding A subunit, a regulatory B subunit and a catalytic C subunit. PP2As were shown to play diverse roles in eukaryotes. In this study, the function of the Arabidopsis PP2A‐C5 gene that encodes the catalytic subunit 5 of PP2A was studied using both loss‐of‐function and gain‐of‐function analyses. Loss‐of‐function mutant pp2a‐c5‐1 displayed more impaired growth during root and shoot development, whereas overexpression of PP2A‐C5 conferred better root and shoot growth under different salt treatments, indicating that PP2A‐C5 plays an important role in plant growth under salt conditions. Double knockout mutants of pp2a‐c5‐1 and salt overly sensitive (sos) mutants sos1‐1, sos2‐2 or sos3‐1 showed additive sensitivity to NaCl, indicating that PP2A‐C5 functions in a pathway different from the SOS signalling pathway. Using yeast two‐hybrid analysis, four vacuolar membrane chloride channel (CLC) proteins, AtCLCa, AtCLCb, AtCLCc and AtCLCg, were found to interact with PP2A‐C5. Moreover, overexpression of AtCLCc leads to increased salt tolerance and Cl⁻ accumulation in transgenic Arabidopsis plants. These data indicate that PP2A‐C5‐mediated better growth under salt conditions might involve up‐regulation of CLC activities on vacuolar membranes and that PP2A‐C5 could be used for improving salt tolerance in crops.