Rice OsCIPK17-OsCBL2/3 module enhances shoot Na+ exclusion and plant salt tolerance in transgenic Arabidopsis.

Soil salinity is detrimental to plant growth and remains a major threat to crop productivity of the world. Plants employ various physiological and molecular mechanisms to maintain growth under salt stress. Identification of genes and genetic loci underlying plant salt tolerance holds the key to breeding salt tolerant crops. CIPK-CBL pathways regulate adaptive responses of plants (especially ion transport) to abiotic stresses via fine-tuned Ca2+ signal transduction. In this study, we showed that over-expression of OsCIPK17 in Arabidopsis enhanced primary root elongation under salt stress, which is in a Ca2+ dependent manner. Further investigation revealed that, under salt stress, OsCIPK17 transcript level was significantly induced and its protein moved from the cytosol to the tonoplast. Using both Y2H and BiFC, tonoplast-localised OsCBL2 and OsCBL3 were shown to interact with OsCIPK17. Interestingly, over-expressing salt-induced OsCBL2 or OsCBL3 in Arabidopsis led to enhanced primary root elongation under salt stress. In this process, OsCIPK17 was shown recruited to the tonoplast (similar to the effect of salt stress). Furthermore, transgenic Arabidopsis lines individually over-expressing OsCIPK17, OsCBL2 and OsCBL3 all demonstrated larger biomass and less Na + accumulation in the shoot under salt stress. All data combined suggest that OsCIPK17- OsCBL2/3 module is a major component of shoot Na+ exclusion and therefore plant salt tolerance, which is through enhanced Na + compartmentation into the vacuole in the root. OsCIPK17 and OsCBL2/3 are therefore potential genetic targets that can be used for delivering salt tolerant rice cultivars.

Comparative analysis of salinity tolerance mechanisms in two maize genotypes: growth performance, ion regulation, and antioxidant responses.

This study investigates the differential responses of two maize genotypes, SC180 and SC168, to salt stress, aiming to elucidate the mechanisms underlying salinity tolerance and identify traits associated with improved stress resilience. Salinity stress, imposed by 150 mM NaCl, adversely affected various growth parameters in both genotypes. SC180 exhibited a more pronounced reduction in shoot length (13.6%) and root length (13.6%) compared to SC168, which showed minimal reductions (3.0% and 2.3%, respectively). Additionally, dry weight losses in SC180's leaves, stems, and roots were significantly greater than those in SC168. Under salinity stress, both genotypes accumulated Na+ in all organs, with SC168 showing higher Na + concentrations. However, K+ levels decreased more significantly in SC180's leaves than in SC168's. The study also assessed physiological responses, noting that SC180 experienced a substantial reduction in relative water content (RWC) in leaves (22.7%), while SC168's RWC remained relatively stable (5.15%). Proline accumulation, a marker for osmotic adjustment, increased 2.3-fold in SC168 compared onefold in SC180. Oxidative stress indicators, such as electrolyte leakage and hydrogen peroxide levels, were elevated in both genotypes under salt stress, with SC180 showing higher increases (48.5% and 48.7%, respectively) than SC168 (35.25% and 22.0%). Moreover, antioxidant enzymes (APX, CAT, POD, SOD, GR) activities were significantly enhanced in SC168 under salinity stress, whereas SC180 showed no significant changes in these activities. Stress indices, used to quantify and compare salinity tolerance, consistently ranked SC168 as more tolerant (average rank = 1.08) compared to SC180 (average rank = 1.92). Correlation analyses further confirmed that SC168's superior tolerance was associated with better Na + regulation, maintenance of K+ levels, and a robust antioxidant defense system. In conclusion, SC168 demonstrated greater resilience to salinity stress, attributed to its efficient ion regulation, stable water status, enhanced osmotic adjustment, and strong antioxidant response. These findings provide valuable insights for breeding and developing salinity-tolerant maize varieties.

Comparing Essentiality of SOS1-Mediated Na+ Exclusion in Salinity Tolerance between Cultivated and Wild Rice Species.

Soil salinity is a major constraint that affects plant growth and development. Rice is a staple food for more than half of the human population but is extremely sensitive to salinity. Among the several known mechanisms, the ability of the plant to exclude cytosolic Na+ is strongly correlated with salinity stress tolerance in different plant species. This exclusion is mediated by the plasma membrane (PM) Na+/H+ antiporter encoded by Salt Overly Sensitive (SOS1) gene and driven by a PM H+-ATPase generated proton gradient. However, it is not clear to what extent this mechanism is operational in wild and cultivated rice species, given the unique rice root anatomy and the existence of the bypass flow for Na+. As wild rice species provide a rich source of genetic diversity for possible introgression of abiotic stress tolerance, we investigated physiological and molecular basis of salinity stress tolerance in Oryza species by using two contrasting pairs of cultivated (Oryza sativa) and wild rice species (Oryza alta and Oryza punctata). Accordingly, dose- and age-dependent Na+ and H+ fluxes were measured using a non-invasive ion selective vibrating microelectrode (the MIFE technique) to measure potential activity of SOS1-encoded Na+/H+ antiporter genes. Consistent with GUS staining data reported in the literature, rice accessions had (~4-6-fold) greater net Na+ efflux in the root elongation zone (EZ) compared to the mature root zone (MZ). Pharmacological experiments showed that Na+ efflux in root EZ is suppressed by more than 90% by amiloride, indicating the possible involvement of Na+/H+ exchanger activity in root EZ. Within each group (cultivated vs. wild) the magnitude of amiloride-sensitive Na+ efflux was higher in tolerant genotypes; however, the activity of Na+/H+ exchanger was 2-3-fold higher in the cultivated rice compared with their wild counterparts. Gene expression levels of SOS1, SOS2 and SOS3 were upregulated under 24 h salinity treatment in all the tested genotypes, with the highest level of SOS1 transcript detected in salt-tolerant wild rice genotype O. alta (~5-6-fold increased transcript level) followed by another wild rice, O. punctata. There was no significant difference in SOS1 expression observed for cultivated rice (IR1-tolerant and IR29-sensitive) under both 0 and 24 h salinity exposure. Our findings suggest that salt-tolerant cultivated rice relies on the cytosolic Na+ exclusion mechanism to deal with salt stress to a greater extent than wild rice, but its operation seems to be regulated at a post-translational rather than transcriptional level.

The mechanistic basis of sodium exclusion in Puccinellia tenuiflora under conditions of salinity and potassium deprivation.

Soil salinity is a significant threat to global agriculture. Understanding salt exclusion mechanisms in halophyte species may be instrumental in improving salt tolerance in crops. Puccinellia tenuiflora is a typical salt-excluding halophytic grass often found in potassium-deprived saline soils. Our previous work showed that P. tenuiflora possesses stronger selectivity for K+ than for Na+ ; however, the mechanistic basis of this phenomenon remained elusive. Here, P. tenuiflora PutHKT1;5 was cloned and the functions of PutHKT1;5 and PutSOS1 were characterized using heterologous expression systems. Yeast assays showed that PutHKT1;5 possessed Na+ transporting capacity and was highly selective for Na+ over K+ . PutSOS1 was located at the plasma membrane and operated as a Na+ /K+ exchanger, with much stronger Na+ extrusion capacity than its homolog from Arabidopsis. PutHKT2;1 mediated high-affinity K+ and Na+ uptake and its expression levels were upregulated by mild salinity and K+ deprivation. Salinity-induced changes of root PutHKT1;5 and PutHKT1;4 transcript levels matched the expression pattern of root PutSOS1, which was consistent with root Na+ efflux. The transcript levels of root PutHKT2;1 and PutAKT1 were downregulated by salinity. Taken together, these findings demonstrate that the functional activity of PutHKT1;5 and PutSOS1 in P. tenuiflora roots is fine-tuned under saline conditions as well as by operation of other ion transporters/channel (PutHKT1;4, PutHKT2;1, and PutAKT1). This leads to the coordination of radial Na+ and K+ transport processes, their loading to the xylem, or Na+ retrieval and extrusion under conditions of mild salinity and/or K+ deprivation.

Rice Na+-Permeable Transporter OsHAK12 Mediates Shoots Na+ Exclusion in Response to Salt Stress.

Soil salinity has become a major stress factor that reduces crop productivity worldwide. Sodium (Na+) toxicity in a number of crop plants is tightly linked with shoot Na+ overaccumulation, thus Na+ exclusion from shoot is crucial for salt tolerance in crops. In this study, we identified a member of the high-affinity K+ transport family (HAK), OsHAK12, which mediates shoots Na+ exclusion in response to salt stress in rice. The Oshak12 mutants showed sensitivity to salt toxicity and accumulated more Na+ in the xylem sap, leading to excessive Na+ in the shoots and less Na+ in the roots. Unlike typical HAK family transporters that transport K+, OsHAK12 is a Na+-permeable plasma membrane transporter. In addition, OsHAK12 was strongly expressed in the root vascular tissues and induced by salt stress. These findings indicate that OsHAK12 mediates Na+ exclusion from shoot, possibly by retrieving Na+ from xylem vessel thereby reducing Na+ content in the shoots. These findings provide a unique function of a rice HAK family member and provide a potential target gene for improving salt tolerance of rice.

Comparative physiological and transcriptomic analysis reveals salinity tolerance mechanisms in Sorghum bicolor (L.) Moench.

MAIN CONCLUSION: Mota Maradi is a sorghum line that exhibits holistic salinity tolerance mechanisms, making it a viable potential donor in breeding efforts for improved sorghum lines. High soil salinity is one of the global challenges for crop growth and productivity. Understanding the salinity tolerance mechanisms in crops is necessary for genetic breeding of salinity-tolerant crops. In this study, physiological and molecular mechanisms in sorghum were identified through a comparative analysis between a Nigerien salinity-tolerant sorghum landrace, Mota Maradi, and the reference sorghum line, BTx623. Significant differences on physiological performances were observed, particularly on growth and biomass gain, photosynthetic rate, and the accumulation of Na+, K+, proline, and sucrose. Transcriptome profiling of the leaves, leaf sheaths, stems, and roots revealed contrasting differentially expressed genes (DEGs) in Mota Maradi and BTx623 which supports the physiological observations from both lines. Among the DEGs, ion transporters such as HKT, NHX, AKT, HAK5, and KUP3 were likely responsible for the differences in Na+ and K+ accumulation. Meanwhile, DEGs involved in photosynthesis, cellular growth, signaling, and ROS scavenging were also identified between these two genotypes. Functional and pathway analysis of the DEGs has revealed that these processes work in concert and are crucial in elevated salinity tolerance in Mota Maradi. Our findings indicate how different complex processes work synergistically for salinity stress tolerance in sorghum. This study also highlights the unique adaptation of landraces toward their respective ecosystems, and their strong potential as genetic resources for future plant breeding endeavors.

Different Rhizospheric pH Conditions Affect Nutrient Accumulations in Rice under Salinity Stress.

This study was conducted to determine the responses to saline-alkaline (SA) stress with regard to nutrient accumulation in two rice varieties having different tolerances to salt-stress. A salinity-tolerant landrace, Pokkali, and a salinity-sensitive variety, PTT1, were exposed to three levels of SA conditions, pH 7.0 (mild), pH 8.0 (moderate), and pH 9.0 (severe), under 50 mM Na stress. The results indicated that Pokkali had comparably greater SA tolerance than PTT1 owing to its higher biomass production. The maintenance of the lower Na/K ratio in Pokkali shoots was achieved by the higher expression of OsHKT1;5 encoding a Na+ transporter in the shoots, OsNHX1 encoding a tonoplast-localized Na+/H+ antiporter in the roots, and OsHAK16 encoding a K+ transporter in the roots under SA conditions. We propose that the high expression of Fe deficiency-responsive genes, OsIRT1, OsIRO2, OsYSL15, OsNAS1, and OsNAS2, in both rice varieties under all SA conditions should contribute to Fe homeostasis in the shoots. In addition, SA treatment increased the concentrations of Ca, Mn, Zn, and Cu in the roots but decreased their concentrations in the shoots of both varieties. Overall, the results indicated that high rhizospheric pH influenced nutrient uptake and translocation from the roots to the shoots in rice.

Evaluating and Screening of Agro-Physiological Indices for Salinity Stress Tolerance in Wheat at the Seedling Stage.

Soil salinity is a worldwide issue that affects wheat production. A comprehensive understanding of salt-tolerance mechanisms and the selection of reliable screening indices are crucial for breeding salt-tolerant wheat cultivars. In this study, 30 wheat genotypes (obtained from a rapid selection of 96 original varieties) were chosen to investigate the existing screening methods and clarify the salinity tolerance mechanisms in wheat. Ten-day-old seedlings were treated with 150 mM NaCl. Eighteen agronomic and physiological parameters were measured. The results indicated that the effects of salinity on the agronomic and physiological traits were significant. Salinity stress significantly decreased K+ content and K+/Na+ ratio in the whole plant, while the leaf K+/Na+ ratio was the strongest determinant of salinity tolerance and had a significantly positive correlation with salt tolerance. In contrast, salinity stress significantly increased Na+ concentration and relative gene expression (TaHKT1;5, TaSOS1, and TaAKT1-like). The Na+ transporter gene (TaHKT1;5) showed a significantly greater increase in expression than the K+ transporter gene (TaAKT1-like). We concluded that Na+ exclusion rather than K+ retention contributed to an optimal leaf K+/Na+ ratio. Furthermore, the present exploration revealed that, under salt stress, tolerant accessions had higher shoot water content, shoot dry weight and lower stomatal density, leaf sap osmolality, and a significantly negative correlation was observed between salt tolerance and stomatal density. This indicated that changes in stomata density may represent a fundamental mechanism by which a plant may optimize water productivity and maintain growth under saline conditions. Taken together, the leaf K+/Na+ ratio and stomatal density can be used as reliable screening indices for salt tolerance in wheat at the seedling stage.

Functions and structure of roots and their contributions to salinity tolerance in plants.

Soil salinity is an increasing threat to the productivity of glycophytic crops worldwide. The root plays vital roles under various stress conditions, including salinity, as well as has diverse functions in non-stress soil environments. In this review, we focus on the essential functions of roots such as in ion homeostasis mediated by several different membrane transporters and signaling molecules under salinity stress and describe recent advances in the impacts of quantitative trait loci (QTLs) or genetic loci (and their causal genes, if applicable) on salinity tolerance. Furthermore, we introduce important literature for the development of barriers against the apoplastic flow of ions, including Na+, as well as for understanding the functions and components of the barrier structure under salinity stress.

Anatomical changes in stem and root of soybean plants submitted to salt stress.

The soybean is a legume that is widely cultivated in many countries due to the high levels of protein and oil contained in its seed, and is used for human and animal nutrition. However, salinity affects more than 800 million hectares worldwide, limiting global agricultural production. The aim of this research was to evaluate the structural behaviour of the roots and stems under progressive salt stress, detailing the possible anatomical modifications to these organs in soybean plants during this stress. The plants were randomized into five treatments (0, 50, 100, 150 and 200 mm NaCl). All the root regions studied and exposed to 100 mm Na+ exhibited increases in the epidermis and endodermis and formation of lysogenic aerenchyma with increasing salinity, revealing the protective roles of these structures in reducing Na+ influx. In the stem, increases in the cortex and pith in the first internode subject to 100 mm Na+ suggest anatomical responses that aim to minimize oxidative stress. Soybean plants subjected to progressive salt stress (>50 mm Na+ ) avoided cavitation and loss of function linked to vessel elements, reducing the metaxylem in all the root and stem regions analysed. Finally, our results confirm anatomical changes to the roots and stems.

Targeting salt stress coping mechanisms for stress tolerance in Brassica: A research perspective.

Brassica genus comprises numerous cultivated brassica species with various economic importance. Salt stress is an overwhelming problem causing serious losses in Brassica species (e.g. B. napus, B. rapa, B. oleracea, B. juncea) growth and grain yield production by inducing ionic and ROS toxicity. Given that a significant variation exists in salt tolerance level in Brassica genus, Brassica species exhibited numerous salt tolerance mechanisms which were either overlooked or given less importance to improve and understand innate salt stress tolerance mechanism in Brassica species. In this review, we tried to highlight the importance and recent findings relating to some overlooked and potential mechanisms such as role of neurotransmitters, and role of cytosolic Ca2+ and ROS as signaling elements to enhance salt stress tolerance. Studies revealed that salt tolerant brassica species retained more K+ in leaf mesophyll which confers overall salinity tolerance in salt tolerance brassica species. Neurotransmitter such as melatonin, dopamiane and eATP regulates K+ and Ca2+ permeable ion channels and plays a very crucial role in ionic homeostasis under salinity stress in brassica. At the end, the numerous possible salt stress agronomic strategies were also discussed to mitigate the severity of the salt stress in Brassica species.

Contribution of two different Na+ transport systems to acquired salinity tolerance in rice.

To elucidate the mechanisms of salt acclimation, physiological parameters of 70 rice varieties were compared under control and salt stress conditions after the acclimation treatment. The results indicated that some rice varieties had the ability to acclimatize to salt stress, exhibiting improved growth following the acclimation treatment under subsequent salinity stress compared to those without acclimation treatment. Conversely, some varieties exhibited reduced growth both with and without acclimation treatment under subsequent salinity stress. Acclimatized varieties had differential patterns of Na+ accumulation in the leaf blades because some varieties reduced Na+ accumulation under salinity stress, whereas others did not. Under salt stress, the acclimatized varieties with low Na+ accumulation in the leaf blades highly induced the expression of the OsHKT1;5 gene in the roots, which may contribute to Na+ exclusion from the shoots. On the other hand, the acclimatized varieties with high Na+ accumulation in the leaf blades exhibited higher induction of the OsNHX1 gene, whose gene product participates in the compartmentalization of Na+ into vacuoles. Thus, rice develops different mechanisms of salinity acclimation using two Na+ transport systems, and active regulation of Na+ transport at the transcription level may be involved in the salt acclimation process and enhance salinity tolerance.

Overexpression of Rice Expansin7 (Osexpa7) Confers Enhanced Tolerance to Salt Stress in Rice.

Expansins are key regulators of cell-wall extension and are also involved in the abiotic stress response. In this study, we evaluated the function of OsEXPA7 involved in salt stress tolerance. Phenotypic analysis showed that OsEXPA7 overexpression remarkably enhanced tolerance to salt stress. OsEXPA7 was highly expressed in the shoot apical meristem, root, and the leaf sheath. Promoter activity of OsEXPA7:GUS was mainly observed in vascular tissues of roots and leaves. Morphological analysis revealed structural alterations in the root and leaf vasculature of OsEXPA7 overexpressing (OX) lines. OsEXPA7 overexpression resulted in decreased sodium ion (Na+) and accumulated potassium ion (K+) in the leaves and roots. Under salt stress, higher antioxidant activity was also observed in the OsEXPA7-OX lines, as indicated by lower reactive oxygen species (ROS) accumulation and increased antioxidant activity, when compared with the wild-type (WT) plants. In addition, transcriptional analysis using RNA-seq and RT-PCR revealed that genes involved in cation exchange, auxin signaling, cell-wall modification, and transcription were differentially expressed between the OX and WT lines. Notably, salt overly sensitive 1, which is a sodium transporter, was highly upregulated in the OX lines. These results suggest that OsEXPA7 plays an important role in increasing salt stress tolerance by coordinating sodium transport, ROS scavenging, and cell-wall loosening.

Domestication influences morphological and physiological responses to salinity in Brassica oleracea seedlings.

Brassica oleracea cultivars include important vegetable and forage crops grown worldwide, whereas the wild counterpart occurs naturally on European sea cliffs. Domestication and selection processes have led to phenotypic and genetic divergence between domesticated plants and their wild ancestors that inhabit coastal areas and are exposed to saline conditions. Salinity is one of the most limiting factors for crop production. However, little is known about how salinity affects plants in relation to domestication of B. oleracea. The objective of this study was to determine the influence of domestication status (wild, landrace or cultivar) on the response of different B. oleracea crops to salinity, as measured by seed germination, plant growth, water content and mineral concentration parameters at the seedling stage. For this purpose, two independent pot experiments were conducted with six accessions of B. oleracea, including cabbage (group capitata) and kale (group acephala), in a growth chamber under controlled environmental conditions. In both taxonomic groups, differences in domestication status and salt stress significantly affected all major process such as germination, changes in dry matter, water relations and mineral uptake. In the acephala experiment, the domestication × salinity interaction significantly affected water content parameters and shoot Na+ allocation. At early stages of development, wild plants are more succulent than cultivated plants and have a higher capacity to maintain lower Na+ concentrations in their shoots in response to increasing levels of salinity. Different responses of domesticated and cultivated accessions in relation to these traits indicated a high level of natural variation in wild B. oleracea. Exclusion of Na+ from shoots and increasing succulence may enhance salt tolerance in B. oleracea exposed to extreme salinity in the long term. The wild germplasm can potentially be used to improve the salt tolerance of crops by the identification of useful genes and incorporation of these into salinity-sensitive cultivars.

Shoot sodium exclusion in salt stressed barley (Hordeum vulgare L.) is determined by allele specific increased expression of HKT1;5.

High affinity potassium transporters (HKT) are recognized as important genes for crop salt tolerance improvement. In this study, we investigated HvHKT1;5 as a candidate gene for a previously discovered quantitative trait locus that controls shoot Na+ and Na+/K+ ratio in salt-stressed barley lines on a hydroponic system. Two major haplotype groups could be distinguished for this gene in a barley collection of 95 genotypes based on the presence of three intronic insertions; a designated haplotype group A (HGA, same as reference sequence) and haplotype group B (HGB, with insertions). HGB was associated with a much stronger root expression of HKT1;5 compared to HGA, and consequently higher K+ and lower Na+ and Cl- concentrations and a lower Na+/K+ ratio in the shoots three weeks after exposure to 200 mM NaCl. Our experimental results suggest that allelic variation in the promoter region of the HGB gene is linked to the three insertions may be responsible for the observed increase in expression of HvHKT1;5 alleles after one week of salt stress induction. This study shows that in barley - similar to wheat and rice - HKT1;5 is an important contributor to natural variation in shoot Na+ exclusion.

Characterization of Na+ exclusion mechanism in rice under saline-alkaline stress conditions.

This study was designed to elucidate the physiological responses of two rice genotypes to different pH levels under high saline stress. A salt-tolerant cultivar, FL478, and a salt-sensitive cultivar, IR29, were exposed to saline-alkaline solutions supplemented with 50 mM Na at pH 9 (severe), pH 8 (moderate), and pH 7 (mild) for three weeks. The results indicated that FL478 is relatively saline-alkaline tolerant compared to IR29, and this was evident from its higher dry mass production, lower Na+ concentration in the leaf blades, and maintenance of water balance under both mild and moderate saline-alkaline stress conditions. In both cultivars, Na+ concentrations in the leaf blades were considerably higher at pH 8 than at pH 7, indicating that high alkaline stress promoted Na+ accumulation under highly saline conditions. FL478 plants had lower Na+/K+ ratios in leaf blades and leaf sheaths than IR29 plants under saline-alkaline stress at both pH 7 and pH 8. To understand the mechanisms behind the difference in saline-alkaline tolerance between the two rice genotypes, transcript levels of the genes encoding Na+ transport proteins were analyzed. In response to mild and moderate saline-alkaline stress conditions, salt-tolerant FL478 had highly induced expression of the OsHKT1;5 gene in the roots, corresponding to lower Na+ accumulation in the leaf blades. Induction of high expression of the OsSOS1 gene in the roots of FL478 implied that Na may be effectively exported from cytosols to apoplasts in the roots resulting in sequestration of Na+ to outside of the roots and loading Na+ in xylem transpiration stream. On the other hand, the salt-sensitive IR29 had lower expression of the genes related to Na+ transporters, such as the OsHKT1;5 gene and the OsSOS1 gene, in the roots, leading to higher Na+ accumulation in the shoots. Expression of the determinant genes for alkaline tolerance, such as K+ and Fe acquisition and acidification of the rhizosphere was highly induced in FL478, but not in IR29. Thus, molecular analysis suggested that genes encoding Na+ transport proteins are involved in regulating Na+ transport under saline-alkaline stress in both salt-tolerant and salt-sensitive rice cultivars, and this is useful information for improving saline-alkaline tolerance traits of rice in the future.

Sustainable wheat (Triticum aestivum L.) production in saline fields: a review.

A large part of global agricultural fields, including the wheat (Triticum aestivum L.) ones, are subjected to various stresses including salinity. Given the increasing world population, finding methods and strategies that can alleviate salinity stress on crop yield production is of outmost importance. The presented review has consulted more than 400 articles related to the clean and sustainable production of wheat in saline fields affected by biological, environmental, economical, and social parameters including the important issue of climate change (global warming). The negative effects of salt stress on plant growth and the techniques, which have been so far detected to alleviate salinity stress on wheat growth have been analyzed and presented. The naturally tolerant species of wheat can use a range of mechanisms to alleviate salinity stress including sodium exclusion, potassium retention, and osmoregulation. However, the following can be considered as the most important techniques to enhance wheat tolerance under stress: (1) the biotechnological (crop breeding), biological (soil microbes), and biochemical (seed priming) methods, (2) the use of naturally tolerant genotypes, and (3) their combined use. The proper handling of irrigation water is also an important subject, which must be considered when planting wheat in saline fields. In conclusion, the sustainable and cleaner production of wheat under salt stress is determined by a combination of different parameters including the biotechnological techniques, which if handled properly, can enhance wheat production in saline fields.

Genome-Wide Identification of Na+/H+ Antiporter (NHX) Genes in Sugar Beet (Beta vulgaris L.) and Their Regulated Expression under Salt Stress.

Salinity is one of the major environment factors that limits the growth of plants and the productivity of crops worldwide. It has been shown that Na+ transporters play a central role in salt tolerance and development of plants. The objective of this study was to identify Na+/H+ antiporter (NHX) genes and investigate their expression patterns in sugar beet (Beta vulgaris L.) subjected to various concentrations of NaCl. A total of five putative NHX genes were identified and distributed on four chromosomes in sugar beet. Phylogenetic analysis revealed that these BvNHX genes are grouped into three major classes, viz Vac- (BvNHX1, -2 and -3), Endo- (BvNHX4), and PM-class NHX (BvNHX5/BvSOS1), and within each class the exon/intron structures are conserved. The amiloride-binding site is found in TM3 at N-terminus of Vac-class NHX proteins. Protein-protein interaction (PPI) prediction suggested that only BvNHX5 putatively interacts with calcineurin B-like proteins (CBL) and CBL-interacting protein kinases (CIPK), implying it might be the primary NHX involved in CBL-CIPK pathway under saline condition. It was also found that BvNHX5 contains one abscisic acid (ABA)-responsive element (ABRE), suggesting that BvNHX5 might be involved in ABA signal responsiveness. Additionally, the qRT-PCR analysis showed that all the BvNHX genes in both roots and leaves are significantly up-regulated by salt, and the transcription levels under high salinity are significantly higher than those under either low or moderate salinity. Taken together, this work gives a detailed overview of the BvNHX genes and their expression patterns under salt stress. Our findings also provide useful information for elucidating the molecular mechanisms of Na+ homeostasis and further functional identification of the BvNHX genes in sugar beet.

Root-zone-specific sensitivity of K+-and Ca2+-permeable channels to H2O2 determines ion homeostasis in salinized diploid and hexaploid Ipomoea trifida.

Polyploids generally possess superior K+/Na+ homeostasis under saline conditions compared with their diploid progenitors. In this study, we identified the physiological mechanisms involved in the ploidy-related mediation of K+/Na+ homeostasis in the roots of diploid (2x) and hexaploid (6x; autohexaploid) Ipomoea trifida, which is the closest relative of cultivated sweet potato. Results showed that 6x I. trifida retained more K+ and accumulated less Na+ in the root and leaf tissues under salt stress than 2x I. trifida. Compared with its 2x ancestor, 6x I. trifida efficiently prevents K+ efflux from the meristem root zone under salt stress through its plasma membrane (PM) K+-permeable channels, which have low sensitivity to H2O2. Moreover, 6x I. trifida efficiently excludes Na+ from the elongation and mature root zones under salt stress because of the high sensitivity of PM Ca2+-permeable channels to H2O2. Our results suggest the root-zone-specific sensitivity to H2O2 of PM K+- and Ca2+-permeable channels in the co-ordinated control of K+/Na+ homeostasis in salinized 2x and 6x I. trifida. This work provides new insights into the improved maintenance of K+/Na+ homeostasis of polyploids under salt stress.