HgS2, a novel salt-responsive gene from the Halophyte Halogeton glomeratus, confers salt tolerance in transgenic Arabidopsis.

Halophyte Halogeton glomeratus mostly grows in saline desert areas in arid and semi-arid regions and is able to adapt to adverse conditions such as salinity and drought. Earlier transcriptomic studies revealed activation of the HgS2 gene in the leaf of H. glomeratus seedlings when exposed to saline conditions. To identify the properties of HgS2 in H. glomeratus, we used yeast transformation and overexpression in Arabidopsis. Yeast cells genetically transformed with HgS2 exhibited K+ uptake and Na+ efflux compared with control (empty vector). Stable overexpression of HgS2 in Arabidopsis improved its resistance to salt stress and led to a notable rise in seed germination in salinity conditions compared to the wild type (WT). Transgenic Arabidopsis regulated ion homeostasis in plant cells by increasing Na+ absorption and decreasing K+ efflux in leaves, while reducing Na+ absorption and K+ efflux in roots. In addition, overexpression of HgS2 altered transcription levels of stress response genes and regulated different metabolic pathways in roots and leaves of Arabidopsis. These results offer new insights into the role of HgS2 in plants' salt tolerance.

Klenkia sesuvii sp. nov., isolated from leaves of halophyte Sesuvium portulacastrum.

During a study on the diversity of culturable actinobacteria from coastal halophytes in Thailand, strain LSe6-5T was isolated from leaves of sea purslane (Sesuvium portulacastrum L.), and a polyphasic approach was employed to determine its taxonomic position. The 16S rRNA gene sequences analysis indicated that the strain was most closely related to Klenkia brasiliensis Tu 6233T (99.2 %), Klenkia marina YIM M13156T (99.1 %), and Klenkia terrae PB261T (98.7 %). The genome of strain LSe6-5T was estimated to be 4.33 Mbp in size, with DNA G+C contents of 74.3%. A phylogenomic tree based on whole-genome sequences revealed that strain LSe6-5T formed a clade with Klenkia marina DSM 45722T, indicating their close relationship. However, the average nucleotide identity (ANI)-blast, ANI-MUMmer, and dDDH values between strain LSe6-5T with K. marina DSM 45722T (87.1, 88.9, and 33.0 %) were below the thresholds of 95-96 % ANI and 70 % dDDH for identifying a novel species. Furthermore, strain LSe6-5T showed morphological and chemotaxonomic characteristics of the genus Klenkia. Cells were motile, rod-shaped, and Gram-stain-positive. Optimal growth of strain LSe6-5T occurred at 28 °C, pH 7.0, and 0-3 % NaCl. The whole-cell hydrolysates contained meso-diaminopimelic acid as the diagnostic diamino acid, with galactose, glucose, mannose, and ribose as whole-cell sugars. The predominant menaquinones were MK-9(H4) and MK-9(H0). The polar lipid profile was composed of diphosphatidylglycerol, hydroxyphosphatidylethanolamine, phosphatidylinositol, glycophosphatidylinositol, an unidentified phospholipid, and an unidentified lipid. Major cellular fatty acids were iso-C15 : 0, iso-C16 : 0, and iso-C17 : 0. From the distinct phylogenetic position and combination of genotypic and phenotypic characteristics, it is supported that strain LSe6-5T represents a novel species of the genus Klenkia, for which the name Klenkia sesuvii sp. nov. is proposed. The type strain is strain LSe6-5T (=TBRC 16417T= NBRC 115929T).

Effect of salt-alkali stress on seed germination of the halophyte Halostachys caspica.

The increasing global phenomenon of soil salinization has prompted heightened interest in the physiological ecology of plant salt and alkali tolerance. Halostachys caspica belonging to Amaranthaceae, an exceptionally salt-tolerant halophyte, is widely distributed in the arid and saline-alkali regions of Xinjiang, in Northwest China. Soil salinization and alkalinization frequently co-occur in nature, but very few studies focus on the interactive effects of various salt and alkali stress on plants. In this study, the impacts on the H. caspica seed germination, germination recovery and seedling growth were investigated under the salt and alkali stress. The results showed that the seed germination percentage was not significantly reduced at low salinity at pH 5.30-9.60, but decreased with elevated salt concentration and pH. Immediately after, salt was removed, ungerminated seeds under high salt concentration treatment exhibited a higher recovery germination percentage, indicating seed germination of H. caspica was inhibited under the condition of high salt-alkali stress. Stepwise regression analysis indicated that, at the same salt concentrations, alkaline salts exerted a more severe inhibition on seed germination, compared to neutral salts. The detrimental effects of salinity or high pH alone were less serious than their combination. Salt concentration, pH value, and their interactions had inhibitory effects on seed germination, with salinity being the decisive factor, while pH played a secondary role in salt-alkali mixed stress.

The WRKY gene family in the halophyte Limonium bicolor: identification, expression analysis, and regulation of salt stress tolerance.

KEY MESSAGE: 63 L. bicolor WRKY genes were identified and their informatics was analyzed. The results suggested that the LbWRKY genes involved in the development and salt secretion of salt glands in L. bicolor. Salt stress, as a universal abiotic stress, severely inhibits the growth and development of plants. WRKY transcription factors play a vital role in plant growth and development, as well as in response to various stresses. Nevertheless, little is known of systematic genome-wide analysis of the WRKY genes in Limonium bicolor, a model recretohalophyte. In this study, 63 L. bicolor WRKY genes were identified (LbWRKY1-63), which were unevenly distributed across seven chromosomes and one scaffold. Based on the structural and phylogenetic characteristics, 63 LbWRKYs are divided into three main groups. Cis-elements in the LbWRKY promoters were related to growth and development, phytohormone responses, and stress responses. Colinearity analysis showed strong colinearity between LbWRKYs and GmWRKYs from soybean (Glycine max). Therefore, LbWRKY genes maybe have similar functions to GmWRKY genes. Expression analysis showed that 28 LbWRKY genes are highly expressed in roots, 9 in stems, 26 in leaves, and 12 in flowers and most LbWRKY genes responded to NaCl, ABA, and PEG6000. Silencing LbWRKY10 reduced salt gland density and salt secretion ability of leaves, and the salt tolerance of the species. Consistent with this, genes associated with salt gland development were markedly down-regulated in the LbWRKY10-silenced lines. Our findings suggested that the LbWRKY genes involved in the development and salt secretion of salt glands in L. bicolor. Our research provides new insights into the functions of the WRKY family in halophytes.

Gordonia prachuapensis sp. nov. and Gordonia sesuvii sp. nov., two novel actinobacteria isolated from mangrove sediments and leaves of halophyte Sesuvium portulacastrum in Thailand.

A polyphasic approach was used to characterize two novel actinobacterial strains, designated PKS22-38T and LSe1-13T, which were isolated from mangrove soils and leaves of halophyte Sesuvium portulacastrum (L.), respectively. Phylogenetic analyses based on 16S rRNA gene sequences showed that they belonged to the genus Gordonia and were most closely related to three validly published species with similarities ranging from 98.6 to 98.1 %. The genomic DNA G+C contents of strains PKS22-38T and LSe1-13T were 67.3 and 67.2 mol%, respectively. The average nucleotide identity (ANI) and digital DNA-DNA hybridization (dDDH) values between the two strains were 93.3 and 54.9 %, respectively, revealing that they are independent species. Meanwhile, the ANI and dDDH values between the two novel strains and closely related type strains were below 80.5 and 24.0 %, respectively. Strains PKS22-38T and LSe1-13T contained C16 : 0, C18 : 1 ω9c and C18 : 0 10-methyl (TBSA) as the major fatty acids and diphosphatidylglycerol, phosphatidylethanolamine and phosphatidylinositol as the main phospholipids. The predominant menaquinone was MK-9(H2). Based on phenotypic, chemotaxonomic, phylogenetic and genomic data, strains PKS22-38T and LSe1-13T are considered to represent two novel species within the genus Gordonia, for which the names Gordonia prachuapensis sp. nov. and Gordonia sesuvii sp. nov. are proposed, with strain PKS22-38T (=TBRC 17540T=NBRC 116256T) and strain LSe1-13T (=TBRC 17706T=NBRC 116396T) as the type strains, respectively.

Molecular cloning and characterization of a salt overly sensitive3 (SOS3) gene from the halophyte Pongamia.

A high concentration of sodium (Na+) is the primary stressor for plants in high salinity environments. The Salt Overly Sensitive (SOS) pathway is one of the best-studied signal transduction pathways, which confers plants the ability to export too much Na+ out of the cells or translocate the cytoplasmic Na+ into the vacuole. In this study, the Salt Overly Sensitive3 (MpSOS3) gene from Pongamia (Millettia pinnata Syn. Pongamia pinnata), a semi-mangrove, was isolated and characterized. The MpSOS3 protein has canonical EF-hand motifs conserved in other calcium-binding proteins and an N-myristoylation signature sequence. The MpSOS3 gene was significantly induced by salt stress, especially in Pongamia roots. Expression of the wild-type MpSOS3 but not the mutated nonmyristoylated MpSOS3-G2A could rescue the salt-hypersensitive phenotype of the Arabidopsis sos3-1 mutant, which suggested the N-myristoylation signature sequence of MpSOS3 was required for MpSOS3 function in plant salt tolerance. Heterologous expression of MpSOS3 in Arabidopsis accumulated less H2O2, superoxide anion radical (O2-), and malondialdehyde (MDA) than wild-type plants, which enhanced the salt tolerance of transgenic Arabidopsis plants. Under salt stress, MpSOS3 transgenic plants accumulated a lower content of Na+ and a higher content of K+ than wild-type plants, which maintained a better K+/Na+ ratio in transgenic plants. Moreover, no development and growth discrepancies were observed in the MpSOS3 heterologous overexpression plants compared to wild-type plants. Our results demonstrated that the MpSOS3 pathway confers a conservative salt-tolerant role and provided a foundation for further study of the SOS pathway in Pongamia.

How the Ethylene Biosynthesis Pathway of Semi-Halophytes Is Modified with Prolonged Salinity Stress Occurrence?

The mechanism of ethylene (ET)-regulated salinity stress response remains largely unexplained, especially for semi-halophytes and halophytes. Here, we present the results of the multifaceted analysis of the model semi-halophyte Mesembryanthemum crystallinum L. (common ice plant) ET biosynthesis pathway key components' response to prolonged (14 days) salinity stress. Transcriptomic analysis revealed that the expression of 3280 ice plant genes was altered during 14-day long salinity (0.4 M NaCl) stress. A thorough analysis of differentially expressed genes (DEGs) showed that the expression of genes involved in ET biosynthesis and perception (ET receptors), the abscisic acid (ABA) catabolic process, and photosynthetic apparatus was significantly modified with prolonged stressor presence. To some point this result was supported with the expression analysis of the transcript amount (qPCR) of key ET biosynthesis pathway genes, namely ACS6 (1-aminocyclopropane-1-carboxylate synthase) and ACO1 (1-aminocyclopropane-1-carboxylate oxidase) orthologs. However, the pronounced circadian rhythm observed in the expression of both genes in unaffected (control) plants was distorted and an evident downregulation of both orthologs' was induced with prolonged salinity stress. The UPLC-MS analysis of the ET biosynthesis pathway rate-limiting semi-product, namely of 1-aminocyclopropane-1-carboxylic acid (ACC) content, confirmed the results assessed with molecular tools. The circadian rhythm of the ACC production of NaCl-treated semi-halophytes remained largely unaffected by the prolonged salinity stress episode. We speculate that the obtained results represent an image of the steady state established over the past 14 days, while during the first hours of the salinity stress response, the view could be completely different.

Halophytic Bacterial Endophyte Microbiome from Coastal Desert-Adapted Wild Poaceae Alleviates Salinity Stress in the Common Wheat Triticum aestivum L.

Abiotic stresses threaten the strategic crops of Poaceae (Gramineae) worldwide. Habitat-adapted microbiome of wild plants has the potential to alleviate abiotic stresses in alternate hosts. Persian Gulf's coastal deserts are colonized by halophyte plants hosting habitat-adapted halophytic microbiota. Here, endophytic bacteria from wild Poaceae in coastal deserts of the north Persian Gulf at Hormozgan province, Iran, were isolated and screened for mitigating salinity stress in wheat. Accordingly, seven dominant species of wild Poaceae in the region, i.e., Aeloropus lagopoides, Aeloropus litiralis, Chrysopogon aucheri, Cymbopogon olivieri, Desmostachya sp., Halopayrum mucronatum, and Sporbuls arabicus, were explored. In total, 367 endophytic bacteria were isolated, 90 of which tolerated 2.5-M NaCl. Of these, 38 strains were selected based on their bioactivity and applied for in vitro wheat-interaction assays under 250-mM NaCl stress. Five superior strains promoted seed germination and growth indices in rain-fed winter wheat cv. Sardari, i.e., Bacillus subtilis B14, B19, & B27, Bacillus sp. B21, and Bacillus licheniformis Ba38. In planta assays in saline soil (2.7 dS m-1) using the superior strains indicated that Bacillus sp. B21 and Bacillus licheniformis Ba38 increased germination and root and shoot lengths and their dry and fresh weights in wheat seedlings. Moreover, phenolics and flavonoids contents of wheat seedlings were influenced by endophyte application. Thus, the coastal desert-adapted microbiome of wild Poaceae could alleviate abiotic stress and promote growth in cultivated species of Poaceae, such as wheat.

From swamp to field: how genes from mangroves and its associates can enhance crop salinity tolerance.

Salinity stress is a critical challenge in crop production and requires innovative strategies to enhance the salt tolerance of plants. Insights from mangrove species, which are renowned for their adaptability to high-salinity environments, provides valuable genetic targets and resources for improving crops. A significant hurdle in salinity stress is the excessive uptake of sodium ions (Na+) by plant roots, causing disruptions in cellular balance, nutrient deficiencies, and hampered growth. Specific ion transporters and channels play crucial roles in maintaining a low Na+/K+ ratio in root cells which is pivotal for salt tolerance. The family of high-affinity potassium transporters, recently characterized in Avicennia officinalis, contributes to K+ homeostasis in transgenic Arabidopsis plants even under high-salt conditions. The salt overly sensitive pathway and genes related to vacuolar-type H+-ATPases hold promise for expelling cytosolic Na+ and sequestering Na+ in transgenic plants, respectively. Aquaporins contribute to mangroves' adaptation to saline environments by regulating water uptake, transpiration, and osmotic balance. Antioxidant enzymes mitigate oxidative damage, whereas genes regulating osmolytes, such as glycine betaine and proline, provide osmoprotection. Mangroves exhibit increased expression of stress-responsive transcription factors such as MYB, NAC, and CBFs under high salinity. Moreover, genes involved in various metabolic pathways, including jasmonate synthesis, triterpenoid production, and protein stability under salt stress, have been identified. This review highlights the potential of mangrove genes to enhance salt tolerance of crops. Further research is imperative to fully comprehend and apply these genes to crop breeding to improve salinity resilience.

Floristic changes and environmental drivers of soil fungi and archaea in different salt-tolerant plant communities in the intertidal habitat of coastal wetlands.

Microorganisms are crucial elements of terrestrial ecosystems, which play significant roles in improving soil physicochemical properties, providing plant growth nutrients, degrading toxic and harmful chemicals, and biogeochemical cycling. Variations in the types and quantities of root exudates among different plants greatly alter soil physicochemical properties and result in variations in the diversity, structure, and function of soil microorganisms. Not much is understood about the differences of soil fungi and archaea communities for different plant communities in coastal wetlands, and their response mechanisms to environmental changes. In this study, fungal and archaea communities in soils of Suaeda salsa, Phragmites australis, and Spartina alterniflora in the intertidal habitat of coastal wetlands were selected for research. Soil fungi and archaea were analyzed for diversity, community structure, and function using high throughput ITS and 16S rRNA gene sequencing. The study revealed significant differences in fungi and archaea's diversity and community structure in the rhizosphere soil of three plant communities. At the same time, there is no significant difference in the functional groups. SOM, TP, AP, MC, EC and SOM, TN, TP, AP, MC, EC are the primary environmental determinants affecting changes in soil fungal and archaeal communities, respectively. Variations in the diversity, community structure, and ecological functions of fungi and archaea can be used as indicators characterizing the impact of external disturbances on the soil environment, providing a theoretical foundation for the effective utilization of soil microbial resources, thereby achieving the goal of environmental protection and health promotion.

Enhanced photochemical efficiency of PSII in Prosopis juliflora suggests contribution to invasion advantage over native C3 xero-halophytes under salt stress.

Chlorophyll a fluorescence parameters related to PSII photochemistry, photoprotection and photoinhibition were investigated in four C3 plant species growing in their natural habitat: Prosopis juliflora ; Abutilon indicum ; Salvadora persica ; and Phragmites karka . This study compared the light reaction responses of P. juliflora , an invasive species, with three native co-existing species, which adapt to varying water deficit and high salt stress. Chlorophyll a fluorescence quenching analyses revealed that P. juliflora had the highest photochemical quantum efficiency and yield, regulated by higher fraction of open reaction centres and reduced photoprotective energy dissipation without compromising the integrity of photosynthetic apparatus due to photoinhibition. Moreover, the elevated values of parameters obtained through polyphasic chlorophyll a fluorescence induction kinetics, which characterise the photochemistry of PSII and electron transport, highlighted the superior performance index of energy conservation in the transition from excitation to the reduction of intersystem electron carriers for P. juliflora compared to other species. Enhanced pigment contents and their stoichiometry in P. juliflora apparently contributed to upregulating fluxes and yields of energy absorbance, trapping and transport. This enhanced photochemistry, along with reduced non-photochemical processes, could explain the proclivity for invasion advantage in P. juliflora across diverse stress conditions.

Assessing the potential of Australian indigenous edible halophytes as salt substitutes: From wild to plate.

Increased salt (sodium chloride (NaCl)) consumption contributes to high blood pressure, increasing the risk of cardiovascular disease. Reducing the intake of NaCl could result in significant public health benefits. Australian grown halophytes are consumed traditionally by indigenous communities as food and medicine. The importance of halophytes has been recently "rediscovered" due to their salty taste and crunchy texture. This study aimed to assess the potential of Australian indigenous edible halophytes (AIEH) as salt substitutes. A benchtop test was carried out to establish a sensory lexicon of four important AIEH (samphire, seapurslane, seablite, and saltbush) and to select the most promising halophyte based on sensory attributes and nutritional composition. Samphire and saltbush, the most common and commercially important halophytes, were used as comparisons. Semolina was used to prepare the halophyte-based test food for the benchtop sensory study. Results of the formal sensory study showed that the growing location of samphire and saltbush can significantly affect their sensory attributes. Samphire had the most favorable sensory attributes and nutritional quality, with dry herb and bran aroma and flavor, whereas the saltbush test food preparations had herbaceous, minty dry wood, and green fruit aroma and flavor. The "optimal" concentration of added freeze-dried samphire/saltbush powder was determined based on the saltiness perception of the NaCl-semolina formulation (0.3% table salt equivalent to 1% samphire freeze-dried powder and 1.4%-2.0% saltbush freeze-dried powder, respectively). This study provided novel and crucial information on the potential use of AIEH as natural salt substitutes. PRACTICAL APPLICATION: There is an increasing demand for natural salt substitutes. Halophytes are salt tolerant plants that sustain in arid or semiarid areas and have the potential to be used as natural salt substitutes. To the best of our knowledge, this is the first study reporting the sensory profiles of four important Australian indigenous edible halophytes (samphire, seapurslane, seablite, and saltbush). This study also demonstrated how different growing locations can affect the sensory attributes of halophytes and subsequently their potential food applications. Our findings provide critical information and data to further study halophytes in the context of novel food applications.

In vivo Effects of Salicornia herbacea and Calystegia soldanella Extracts for Memory Improvement.

The global elderly population, aged 65 and over, reached approximately 10% in 2020, and this proportion is expected to continue rising. Therefore, the prevalence of neurodegenerative diseases such as Parkinson's disease (PD), which are characterized by declining memory capabilities, is anticipated to increase. In a previous study, we successfully restored the diminished memory capabilities in a fruit fly model of PD by administering an omija extract. To identify functional ingredients that can enhance memory akin to the effects of the omija extract, we conducted screenings by administering halophyte extracts to the PD model. Halophytes are plants that thrive in high-salt environments, and given Korea's geographic proximity to the sea on three sides, it serves as an optimal hub for the utilization of these plants. Upon examining the effects of the oral administration of 12 halophyte extracts, Salicornia herbacea and Calystegia soldanella emerged as potential candidates for ameliorating memory loss in PD model flies. Moreover, our findings suggested that C. soldanella, but not S. herbacea, can mitigate oxidative stress in DJ-1ß mutants.

Effects of short-term nitrogen and phosphorus addition on soil bacterial community of different halophytes.

Soil microbial community composition and diversity are often affected by nutrient enrichment, which may influence soil microbes to affect nutrient cycling and plant community structure. However, the response of soil bacteria to nitrogen (N) and phosphorus (P) addition and whether it is influenced by plants remains unclear. By 16S rRNA sequencing, we investigated the response of the rhizosphere and bulk soil bacterial communities of different halophytes (salt-rejecting, salt-absorbing, and salt-secreting plant) in the Yellow River Delta to short-term N and P addition. The response of rhizosphere bacterial diversity to N and P addition was opposite in Phragmites communis and Suaeda salsa. N addition increased the rhizosphere soil bacterial α-diversity of S. salsa and Aeluropus sinensis, while P addition decreased the rhizosphere bacterial α-diversity bacteria of S. salsa. The N and P addition had a weak effect on the rhizosphere bacterial community composition and a significant effect on the bulk soil bacterial community composition of halophytes. The S. salsa and P. communis bulk soil bacterial community were mainly influenced by P addition, while it was influenced by N addition in A. sinensis. N and P addition reduced the difference in bacterial community composition between the two types of soil. N and P addition increased the eutrophic taxa (Proteobacteria and Bacteroidetes) and decreased the oligotrophic taxa (Acidobacteria). Redundancy analysis showed that soil organic matter, salt, and total N content had significant effects on the bacterial community composition. The results clarify that the response of soil bacterial communities to N and P additions is inconsistent across the three halophyte soils, and the effect of plant species on the bacterial community was stronger than short-term N and P addition. IMPORTANCE: The bulk soil bacterial community was more affected by nutrient addition. Nitrogen (N) and phosphorus (P) have different effects on bacterial community. Soil organic matter is a key factor influencing the response of bacterial community to nutrient addition. N and P influence on bacterial community changes with plants.

Salt-tolerant plant growth-promoting Pseudomonas atacamensis KSS-6 in combination with organic manure enhances rice yield, improves nutrient content and soil properties under salinity stress.

In the current study salt tolerant-plant growth-promoting rhizobacteria (ST-PGPR) Pseudomonas atacamensis KSS-6, selected on the basis of prominent plant growth-promoting (PGP) and stress tolerance properties was tested as bioinoculant to improve yield of rice grown in saline soil. The ST-PGPR KSS-6 was capable of maintaining the PGP traits up to 200 mM NaCl, however, higher salt stress conditions affected these activities. The study was designed to determine the effect of developed talc-based bioformulation using KSS-6 along with organic manure (OM) on growth and yield of paddy under saline conditions. Bioformulation broadcasting was also done to examine the effect on soil properties. It was found that the combinatorial treatment showed positive impact on growth and yield of rice under saline conditions. Co-application of KSS-6 with OM showed maximum increment in growth, chlorophyll content, plant fresh weight, and dry weight as compared to untreated control plants. Furthermore, the combinatorial treatment improved the nutrient content (P, K, Zn, Fe, Mg, and Mn) by more than 35% and enhanced the biochemical parameters such as proline, flavonoids, carbohydrates, protein, dietary fiber, and antioxidant content of rice grains by more than 32%. Soil parameters including pH and electrical conductivity (EC), moisture content, total organic carbon, OM, sodium, and chloride ions were also improved upon treatment. There was significant lowering of EC from 7.43 to 4.3 dS/m when combination of OM and bacteria were applied. These findings suggest that the application of KSS-6 in the form of bioinoculant could be a promising strategy to mitigate negative impacts of salt stress and enhance the yield and nutritional properties of rice grown in degraded and saline soil.

Combining different species in restoration is not always the right decision: Monocultures can provide higher ecological functions than intercropping in a desert ecosystem.

Vegetation restoration in deserts is challenging due to these ecosystems' inherent fragility and harsh environmental conditions. One approach for active restoration involves planting native species, which can accelerate the recovery of ecosystem functions. To ensure the effectiveness of this process, carefully selecting species for planting is crucial. Generally, it is expected that a more diverse mix of species in the plantation will lead to the recovery of a greater number of ecosystem functions, especially when the selected species have complementary niche traits that facilitate maximum cooperation and minimize competition among them. In this study, we evaluated the planting of two native species from the hyper-desert of Taklamakan, China, which exhibit marked morpho-physiological differences: a phreatophytic legume (Alhagi sparsifolia) and a halophytic non-legume (Karelinia caspia). These species were grown in both monoculture and intercrop communities. Monoculture of the legume resulted in the highest biomass accumulation. Intercropping improved several ecosystem functions in the 50 cm-upper soil, particularly those related to phosphorus (P), carbon (C), and sulfur (S) concentrations, as well as soil enzyme activities. However, it also increased soil sodium (Na+) concentration and pH. Halophyte monocultures enhanced ecological functions associated with nitrogen concentrations in the upper soil and with P, S, C, and cation concentrations (K+, Ca2+, Mg2+, Cu2+, Fe2+, Zn2+, Co2+, Ni2+), along with enzyme activities in the deep soil. It also maximized Na+ accumulation in plant biomass. In summary, we recommend legume monoculture when the primary goal is to optimize biomass accumulation. Conversely, halophyte monoculture is advisable when the objective is to extract sodium from the soil or enhance ecosystem functions in the deep soil. Intercropping the two species is recommended to maximize the ecosystem functions of the upper soil, provided there is no salinization risk. When planning restoration efforts in desert regions, it is essential to understand the impact of each species on ecosystem function and how complementary species behave when intercropped. However, these interactions are likely species- and system-specific, highlighting the need for more work to optimize solutions for different arid ecosystems.

Overexpression of a plasmalemma Na+/H+ antiporter from the halophyte Nitraria sibirica enhances the salt tolerance of transgenic poplar.

The plasmalemma Na+/H+ antiporter Salt Overly Sensitive 1 (SOS1) is responsible for the efflux of Na+ from the cytoplasm, an important determinant of salt resistance in plants. In this study, an ortholog of SOS1, referred to as NsSOS1, was cloned from Nitraria sibirica, a typical halophyte that grows in deserts and saline-alkaline land, and its expression and function in regulating the salt tolerance of forest trees were evaluated. The expression level of NsSOS1 was higher in leaves than in roots and stems of N. sibirica, and its expression was upregulated under salt stress. Histochemical staining showed that ß-glucuronidase (GUS) driven by the NsSOS1 promoter was strongly induced by abiotic stresses and phytohormones including salt, drought, low temperature, gibberellin, and methyl jasmonate, suggesting that NsSOS1 is involved in the regulation of multiple signaling pathways. Transgenic 84 K poplar (Populus alba × P. glandulosa) overexpressing NsSOS1 showed improvements in survival rate, root biomass, plant height, relative water levels, chlorophyll and proline levels, and antioxidant enzyme activities versus non-transgenic poplar (NT) under salt stress. Transgenic poplars accumulated less Na+ and more K+ in roots, stems, and leaves, which had a lower Na+/K+ ratio compared to NT under salt stress. These results indicate that NsSOS1-mediated Na+ efflux confers salt tolerance to transgenic poplars, which show more efficient photosynthesis, better scavenging of reactive oxygen species, and improved osmotic adjustment under salt stress. Transcriptome analysis of transgenic poplars confirmed that NsSOS1 not only mediates Na+ efflux but is also involved in the regulation of multiple metabolic pathways. The results provide insight into the regulatory mechanisms of NsSOS1 and suggest that it could be used to improve the salt tolerance of forest trees.

Stress salinity in plants: New strategies to cope with in the foreseeable scenario.

The excess of salts in soils causes stress in most plants, except for some halophytes that can tolerate higher levels of salinity. The excess of Na+ generates an ionic imbalance, reducing the K+ content and altering cellular metabolism, thus impacting in plant growth and development. Additionally, salinity in soil induces water stress due to osmotic effects and increments the production of reactive oxygen species (ROS) that affect the cellular structure, damaging membranes and proteins, and altering the electrochemical potential of H+, which directly affects nutrient absorption by membrane transporters. However, plants possess mechanisms to overcome the toxicity of the sodium ions, such as internalization into the vacuole or exclusion from the cell, synthesis of enzymes or protective compounds against ROS, and the synthesis of metabolites that help to regulate the osmotic potential of plants. Physiologic and molecular mechanisms of salinity tolerance in plants will be addressed in this review. Furthermore, a revision of strategies taken by researchers to confer salt stress tolerance on agriculturally important species are discussed. These strategies include conventional breeding and genetic engineering as transgenesis and genome editing by CRISPR/Cas9.

The NAC gene family in the halophyte Limonium bicolor: Identification, expression analysis, and regulation of abiotic stress tolerance.

NAC transcription factors regulate plant growth, development, and stress responses. However, the number, types, and biological functions of Limonium bicolor LbNAC genes have remained elusive. L. bicolor secretes excessive salt ions through salt glands on its stems and leaves to reduce salt-induced damage. Here, we identified 63 NAC members (LbNAC1-63) in L. bicolor, which were unevenly distributed across eight chromosomes. Cis-elements in the LbNAC promoters were related to growth and development, stress responses, and phytohormone responses. We observed strong colinearity between LbNACs and GmNACs from soybean (Glycine max). Thus, LbNAC genes may share similar functions with GmNAC genes. Expression analysis indicated that 16 LbNAC genes are highly expressed in roots, stems, leaves, and flowers, whereas 17 LbNAC genes were highly expressed throughout salt gland development, suggesting that they may regulate this developmental stage. Silencing LbNAC54 in L. bicolor decreased salt gland density, salt secretion from leaves, and overall salt tolerance. In agreement, genes related to salt gland development were significantly downregulated in LbNAC54-silenced lines. Our findings shed light on LbNAC genes and help elucidate salt gland development and salt secretion in L. bicolor. Our data also provide insight into NAC functions in halophytes.

Sulfur-Oxidizing Bacteria Alleviate Salt and Cadmium Stress in Halophyte Tripolium pannonicum (Jacq.) Dobrocz.

The aim of this study was to investigate how introducing halophilic sulfur-oxidizing bacteria (SOB) Halothiobacillus halophilus to the growth substrate affects the physiological and biochemical responses of the halophyte Tripolium pannonicum (also known as sea aster or seashore aster) under salt and cadmium stress conditions. This study assessed the plant's response to these stressors and bacterial inoculation by analyzing various factors including the accumulation of elements such as sodium (Na), chloride (Cl), cadmium (Cd) and sulfur (S); growth parameters; levels of photosynthetic pigments, proline and phenolic compounds; the formation of malondialdehyde (MDA); and the plant's potential to scavenge 2,2-Diphenyl-1-picrylhydrazyl (DPPH). The results revealed that bacterial inoculation was effective in mitigating the deleterious effect of cadmium stress on some growth criteria. For instance, stem length was 2-hold higher, the growth tolerance index was 3-fold higher and there was a 20% increase in the content of photosynthetic pigments compared to non-inoculated plants. Furthermore, the SOB contributed to enhancing cadmium tolerance in Tripolium pannonicum by increasing the availability of sulfur in the plant's leaves, which led to the maintenance of an appropriate, about 2-fold-higher level of phenolic compounds (phenylpropanoids and flavonols), as well as chloride ions. The level of MDA decreased after bacterial application in all experimental variants except when both salt and cadmium stress were present. These findings provide novel insights into how halophytes respond to abiotic stress following inoculation of the growth medium with sulfur-oxidizing bacteria. The data suggest that inoculating the substrate with SOB has a beneficial effect on T. pannonicum's tolerance to cadmium stress.