The HKT protein family plays a vital role in plant responses to salt stress by mediating sodium (Na+) and potassium (K+) transport and maintaining Na+-K+ balance. Ipomoea pes-caprae (IPC), a pantropical creeping plant distributed along coastal regions in tropical and subtropical zones, exhibits exceptional salt tolerance. Understanding its salt tolerance mechanisms provides valuable insights for developing salt-tolerant crops and identifying candidate genes for genetic engineering. In this study, we identified two HKT genes, IpcHKT1;1 and IpcHKT1;2, in IPC. Phylogenetic analysis with HKT genes from other Ipomoea species revealed that all analyzed species contain two HKT genes located adjacently on the same chromosome. Comparative analysis of conserved motifs and intron-exon structures indicated that, despite their close evolutionary relationship, the HKT genes in IPC may exhibit functional divergence. Promoter analysis showed that their regulatory regions are enriched with cis-elements associated with responses to biotic and abiotic stresses, hormonal signaling, and growth, highlighting functional diversity within the HKT family. Subcellular localization experiments demonstrated that IpcHKT1;1 and IpcHKT1;2 are ion transporters localized to the plasma membrane. Heterologous expression in yeast confirmed their role in Na+/K+ symporter. Furthermore, RT-qPCR analysis revealed distinct expression patterns under salt stress: IpcHKT1;2 was significantly upregulated in roots, while IpcHKT1;1 expression was transitionally downregulated at 400 mM NaCl treatment. Prolonged high expression of IpcHKT1;2 in roots suggests its critical role in sustained salt stress tolerance. These findings provide new insights into the molecular mechanisms of salt tolerance in IPC. The identification of IpcHKT1;1 and IpcHKT1;2 as key players in salt stress responses offers promising genetic resources for enhancing crop resilience to soil salinity, addressing challenges associated with global salinization.

Globally, salt stress is one of the most significant abiotic stresses limiting crop production in dry-land regions. Nowadays, growing crops in dry-land regions under saline irrigation is the main focus. Soil amendment with organic materials has shown the potential to mitigate the adverse effects of salinity on plants. This study aimed to examine the ameliorative impact of soil amendment (manure + sandy, compost + sandy, clay + sandy and sandy soil) on the growth, yield, physiological, and biochemical attributes of Hedysarum scoparium Fisch. et Mey (HS) and Avena sativa L. (OT) under fresh and saline water irrigation in dry-land regions. The results showed that salt stress negatively affected both plant species' growth, physiological traits, yield, and chloride ions. In response to saline irrigation, plants of both species increased catalase (CAT) and ascorbate peroxidase (APX) activities as part of a self-defense mechanism to minimize damage. Salt stress also significantly raised levels of hydrogen peroxide (H2O2), malondialdehyde (MDA), and chloride ions (Cl). However, soil amendment treatments like manure + sandy and compost + sandy soil countered the negative effects of saline irrigation, significantly improving plant growth and yield compared with sandy soil. Thus, organic soil amendment is a promising strategy for sustainable crop production under saline irrigation in dry-land regions. This study provides valuable insights into enhancing agricultural production by fostering resilient halophytes and salt-tolerant plant species in challenging environments.

Salinization is a significant global issue causes irreversible damage to plants by reducing osmotic potential, inhibiting seed germination, and impeding water uptake. Seed germination, a crucial step towards the seedling stage is regulated by several hormones and genes, with the balance between abscisic acid and gibberellin being the key mechanism that either promotes or inhibits this process. Additionally, mucilage, a gelatinous substance, is known to provide protection against drought, herbivory, soil adhesion, and seed sinking. However, limited information is available on the structure and thickness of seed mucilage in halophytes under different salinity conditions. In this study, the mucilage structure of the extreme halophyte Schrenkiella parvula was compared with the glycophyte Arabidopsis thaliana in response to salinity. We found differences in the expression levels of genes such as ABI5, RGL2, DOG1, ENO2, and DHAR2, which are involved in seed germination and antioxidant activity, as well as in the mucilage structure of seeds of S. parvula and A. thaliana seeds at different salt concentrations. The responses of seed germination of S. parvula to salinity indicate that it is more salt-tolerant than A. thaliana. Additionally, it was found that S. parvula mucilage decreased under salt conditions but not under mannitol conditions, whereas in A. thaliana mucilage did not change under both conditions, which is one of the adaptation strategies of S. parvula to salt conditions. We believe that these fundamental analyzes will provide a foundation for future molecular and biochemical studies comparing the responses of crops and halophytes to salinity stress.

Date palm (Phoenix dactylifera L.) is an important crop in arid regions and it is well adapted to desert ecosystems. To understand its remarkable ability to grow and yield in water-limited environments, we conducted experiments in which water was withheld for up to 4 weeks. In response to drought, root, rather than leaf, osmotic strength increased, with organic solutes such as sugars and amino acids contributing more to the osmolyte increase than minerals. Consistently, carbon and amino acid metabolism was acclimated toward biosynthesis at both the transcriptional and translational levels. In leaves, a remodeling of membrane systems was observed, suggesting changes in thylakoid lipid composition which, together with the restructuring of the photosynthetic apparatus, indicated an acclimation preventing oxidative damage. Thus, xerophilic date palm avoids oxidative damage under drought by combined prevention and rapid detoxification of oxygen radicals. Although minerals were expected to serve as cheap key osmotics, date palm also relies on organic osmolytes for osmotic adjustment in the roots during early drought acclimation. The diversion of these resources away from growth is consistent with the date palm strategy of generally slow growth in harsh environments and clearly indicates a trade-off between growth and stress-related physiological responses. Osmotic strength of date palm roots increases with soil desiccation, for which the accumulation of organic osmolytes, such as sugars and amino acids, is essential to complement energetically cheap mineral osmotics.

Soil salinization negatively affects plant growth and threatens food security. Halotolerant plant growth-promoting bacteria (PGPB) can alleviate salt stress in plants via diverse mechanisms. In the present study, we isolated salt-tolerant bacteria with phosphate-solubilizing abilities from the rhizosphere of Salix linearistipularis, a halophyte distributed in saline-alkali soils. Strain A103 showed high phosphate solubilization activity and was identified as Enterobacter asburiae based on genome analysis. In addition, it can produce indole-3-acetic acid (IAA), siderophores, and 1-aminocyclopropane-1-carboxylate (ACC) deaminase. Genome mining has also revealed the presence of several functional genes involved in the promotion of plant growth. Inoculation with A103 markedly improved alfalfa growth in the presence of 100 mM NaHCO3. Under alkali stress, the shoot and root dry weights after bacterial inoculation improved by 42.9 % and 21.9 %, respectively. Meanwhile, there was a 35.9-37.1 % increase in the shoot and root lengths after treatment with A103 compared to the NaHCO3-treated group. Soluble sugar content, peroxidase and catalase activities increased in A103-inoculated alfalfa under alkaline stress. A significant decrease in the malondialdehyde content was observed after treatment with strain A103. Metabolomic analysis indicated that strain A103 positively regulated alkali tolerance in alfalfa through the accumulation of metabolites, such as homocarnosine, panthenol, and sorbitol, which could reduce oxidative damage and act as osmolytes. These results suggest that halophytes are valuable resources for bioprospecting halotolerant beneficial bacteria and that the application of halotolerant growth-promoting bacteria is a natural and efficient strategy for developing sustainable agriculture.

There is still a need to investigate the relationships between glycophytes and halophytes and the many biotic and abiotic factors in their natural environments. Therefore, we study the effects of the type of environment on the ecophysiological responses and condition of the glycophyte Elder Sambucus nigra L., the macrophyte Common Reed Phragmites australis (Cav.) Trin. ex Steud., the facultative halophyte Weeping Alkaligrass Puccinellia distans (Jacq.) Parl, and the obligate halophyte Common Glasswort Salicornia europaea L. in a saline-disturbed anthropogenic region of central Poland. We analyzed the effects of salinity, acidity, and soil organic matter on shoot length, lipoperoxidation, and proline in roots and green parts, and evaluated plant responses to environmental disturbance, which allowed for the comparison of adaptation strategies. The studies were carried out in (1) sodium production (near sodium factories), (2) anthropogenic environments (waste dumps, agroecosystems, calcium deposits, post-production tanks), (3) wetland environments (near river channels and riparian areas), and (4) control (natural, unpolluted environments). Green parts of plants are better suited to indicate environmental stress than roots. Their higher structural MDA membrane damage is related to the transport of toxic ions to the shoots by a rapid transpiration stream in the xylem. We found high salinity to be the main factor inducing growth and found it to be correlated with the high pH effect on proline increase in glycophytes (Elder, Reed) and Weeping Alkaligrass, in contrast to Common Glasswort. We suggest that proline accumulation allows osmotic adjustment in the green parts of reeds and alkaligrasses, but may have another function (in Elder). Common Glasswort accumulates large amounts of Na+, which is energetically more effective than proline accumulation for osmotic adjustment. Organic matter affects plant growth and proline levels, but soil salinity and pH alter nutrient availability. Plant distribution along the salinity gradient indicates that Elder is the most salt-sensitive species compared to Reed, Alkaligrass, and Glasswort. Salinity and the lack of control of thick reeds, which compete with other plant groups, affect the distribution of halophytes in saline environments.

In semi-arid Mediterranean regions, particularly in some wetland soils, salinity is thought to be an indicator of low-quality soils. In this study, a characterization is presented of the soils surrounding El Hito saline pond (Castilla La Mancha, Central Spain), an ecological halophyte niche within a natural semi-arid steppe land. The main aim is to classify the salt-affected soils and their morphology, genesis, and physico-chemical properties. Four soil profiles were opened with a backhoe machine for sampling and subsequent description on the basis of their pedogenetic morphology. Systematic surface sampling was also performed. Standard methods were followed to measure the soil properties of 27 samples. Overall electrical conductivity (EC) and pH levels of the wetland were mapped (using ArcGIS 3.1.3). Soil salinity at elevated levels was detected, inhibiting plant uptake of water and nutrients. Distinct sub-areas of extreme elevated surface salinity providing specialized plant habitats and poor soil structure were observed, as well as a mainly whitened-yellowish-greenish soil colour due to salt accumulation and poor drainage. The soils also showed alkaline pH values. In most samples, the pH was over 8.5, and EC was higher than 4 (dS m-1), and in several samples higher than 20 (dS m-1). A low sodium (Na) content was detected in the saturation extract where magnesium (Mg+) was the dominant soluble cation, followed by both calcium (Ca+) and sodium (Na+), and then potassium (K+), present in lower proportions. Sulphate (SO42-) and then chloride (Cl-) anions were dominant, although carbon trioxide, (CO3-) and carbonate (CO32-) anions were also present. The percentages of organic carbon (C) were very low, while total nitrogen (TN) and available phosphorus (P) were higher in the upper horizons, suggesting a degree of eutrophication. The present work will increase the existing knowledge about the role of El Hito saline pond, that play a vital ecological role in the broader biosphere, providing new suggestions to readers on how this knowledge can be used to improve these types of ecosystems. In particular, the agricultural pesticides and fertilizers continuously damage the soil fertility as evidenced by the high content of soluble phosphorus found in some points of the Hito saline pond.

Soil salinity hampers the survival and productivity of crops. To minimize salt-associated damages in plant, better salt management practices in agriculture have become a prerequisite. Seed priming with different halo-agents is a technique, which improves the primed plant's endurance to tackle sodium. Salt tolerance is achieved in tolerant plants through fundamental physiological mechanisms- ion-exclusion and tissue tolerance, and salt-tolerant plants may (Na+ accumulators) or may not (Na+ excluders) allow sodium movement to leaves. While Na+ excluders depend on ion exclusion in roots, Na+ accumulators are proficient Na+ managers that can compartmentalize Na+ in leaves and use them beneficially as inexpensive osmoticum. Salt-sensitive plants are Na+ accumulators, but their inherent tissue tolerance ability and ion-exclusion process are insufficient for tolerance. Seed priming with different halo-agents aids in 'rewiring' of the salt tolerance mechanisms of plants. The resetting of the salt tolerance mechanism is not universal for every halo-agent and might vary with halo-agents. Here, we review the physiological mechanisms that different halo-agents target to confer enhanced salt tolerance in primed plants. Calcium and potassium-specific halo-agents trigger Na+ exclusion in roots, thus ensuring a low amount of Na+ in leaves. In contrast, Na+-specific priming agents favour processes for Na+ inclusion in leaves, improve plant tissue tolerance or vacuolar sequestration, and provide the greatest benefit to salt-sensitive and sodium accumulating plants. Overall, this review will help to understand the underlying mechanism behind plant's inherent nature towards salt management and its amelioration with different halo-agents, which helps to optimize crop stress performance. Understanding plants' inherent response towards the ion- Na+ and selection of priming agents, both are complementary for optimization of crop performance under stress.

Plant growth-promoting rhizosphere bacteria (PGPR) are increasingly considered as highly efficient bioagents. They confer for instance better salt tolerance to host plants while improving soil enzyme activities and microbiome diversity, which are indicators of soil biological health and fertility. Here, the aim was to investigate the efficiency of native bacteria from saline rhizosphere of Hordeum marinum, in alleviating the adverse effects of salinity in this facultative halophyte with fodder potential. Following identification and characterization for salt tolerance capacity and in vitro plant growth promoting potential, the bacteria strain Bacillus pumilus, was selected for its ameliorative effects on H. marinum when cultivated under NaCl salinity (200 mM) in a greenhouse experiment. Salinity significantly restricted growth, induced oxidative stress and disturbed the mineral nutrition of H. marinum, whereas inoculation with B. pumilus significantly restored growth, as leaf elongation rate and plant dry weight. Similarly, the highest accumulation of K+, N and P were observed in inoculated plants challenged with salinity. This was concomitant with reduced MDA and H2O2 levels, thereby preventing oxidative damage in H. marinum plants cultivated under salt stress. Furthermore, B. pumilus strain also overcame salt impact via indirect mechanisms by activating soil enzymes (dehydrogenase, beta-glucosidase and acid phosphatase) and microbial biomass C production. Overall, our data further highlight the environmental significance of microorganisms inhabiting extreme environments like sabkhas as promising agents for biological approaches aiming at saline soil rehabilitation and improving plant productivity when challenged with salinity, due to their beneficial effect on both plant and the rhizosphere.

Aeluropus lagopoides, a dominant palatable species in various sabkha and coastal regions of Saudi Arabia, can withstand harsh saline environments through phenotypic plasticity. When subjected to grazing, how A. lagopoides adapt phenotypically is currently unknown. There is a breakage in the chain of study on the spatial and temporal expansion strategy of A. lagopoides plants when subjected to different grazing stresses in different saline soil habitats. A grazing experiment was conducted to investigate the phenotypic plasticity and resource allocation pattern response of A. lagopoides in different saline soils. Individual A. lagopoides rhizomes from five saline regions were grown and exposed to varied grazing treatments in the form of clipping, viz; light, moderate, and heavy grazing, as compared to a grazing exclusion control. Our results showed that heavy grazing/clipping significantly decreased the shoot system and above-ground biomass in high-saline region plants in the early season. Plant length, root length, root and shoot biomass, the number of stolons, average stolon length, leaf area, and SLA of A. lagopiodes responded significantly to grazing intensities. A. lagopoides from the Qareenah, Qaseem, and Jizan regions were more tolerant to light grazing than A. lagopoides from the Salwa and Jouf regions. Light grazing showed significantly good re-growth, especially during the late season. Light grazing decreased the synthesis of chlorophyll content. Also, A. lagopiodes reduced the risk caused by reactive oxygen species via the increased accumulation of proline content. Overall, plants adapted to different morphological and physiological strategies to tolerate different levels of grazing intensities by adapting their morphological attributes. Though heavy grazing damages the plant, light and moderate grazing can be allowed to maintain the productivity and economic benefits of sabka habitats where soil conditions are moderately saline.