Modulation of ion transporter genes of salt-stressed sorghum (Sorghum bicolor L. Moench) by foliar application of digitoxin.

A meeting on the molecular basis of ionic homeostasis and salt tolerance in plants took place in Madrid, Spain, October 22–24, 2001. This meeting was organized by Eduardo Blumwald (Davis, CA) and Alonso Rodriguez‐Navarro (Madrid, Spain) at the Centre for International Meetings on Biology (‘Instituto Juan March de Estudios e Investigaciones’). ![][1] Ionic homeostasis is a fundamental cellular phenomenon. All living cells maintain an intracellular ionic composition compatible with their constituent molecules, and this requires the regulation of multiple membrane transporters and signal transduction pathways. Other biophysical parameters such as turgor and electrical potential are also part of this essential regulation. How ionic homeostasis is achieved, however, is not completely understood. Although most transporters have already been identified, their physiological function is only starting to be demonstrated and the receptors and most components of the regulatory pathways that effect ionic homeostasis remain unknown. In the case of plants, this problem is related to mineral nutrition and salinity tolerance, both of which have great relevance for agriculture. In fact, as demonstrated by this meeting, salinity stress has been one of the keys to opening the black box of ionic homeostasis in general. Another has been the novel molecular genetics of the plant Arabidopsis thaliana . Of course, other approaches have also contributed to our present understanding of ion homeostasis in plants and were represented at the meeting. For further details, see Blumwald (2000), Hasegawa et al . (2000), Bohnert et al . (2001), Serrano and Rodriguez‐Navarro (2001) and Zhu (2001). ### Some physiology of salt tolerance Salt stress is an important threat to the future of agriculture in many productive areas of the planet. In countries such as Australia and Pakistan, salinity is already a national concern, as it was in the past in ancient Mesopotamia. Areas of California and the Mediterranean region are also threatened. … [1]: /embed/graphic-1.gif

Comparative analyses of plant responses to drought and salt stress in related taxa: A useful approach to study stress tolerance mechanisms.

The improved resistance to high salinity induced by trehalose is associated with ionic regulation and osmotic adjustment in Catharanthus roseus

Drought and salinity are the main abiotic stresses limiting crop yield and quality worldwide. Improving food production in drought- and salt-prone areas is the key to meet the increasing food demands in near future. It has been widely reported that silicon (Si), a second most abundant element in soil, could reduce drought and salt stress in plants. Here, we reviewed the emerging role of Si in enhancing drought and salt tolerance in plants and highlighted the mechanisms through which Si could alleviate both drought and salt stress in plants. Silicon application increased plant growth, biomass, photosynthetic pigments, straw and grain yield, and quality under either drought or salt stress. Under both salt and drought stress, the key mechanisms evoked are nutrient elements homeostasis, modification of gas exchange attributes, osmotic adjustment, regulating the synthesis of compatible solutes, stimulation of antioxidant enzymes, and gene expression in plants. In addition, Si application decreased Na(+) uptake and translocation while increased K(+) uptake and translocation under salt stress. However, these mechanisms vary with plant species, genotype, growth conditions, duration of stress imposed, and so on. This review article highlights the potential for improving plant resistance to drought and salt stress by Si application and provides a theoretical basis for application of Si in saline soils and arid and semiarid regions worldwide. This review article also highlights the future research needs about the role of Si under drought stress and in saline soils.

Background Salinity is one of the most lethal abiotic stresses which affect multiple aspects of plant physiology. Natural variations in plant germplasm are a great resource that could be exploited for improvement in salt tolerance. Eruca sativa (E. sativa) exhibits tolerance to abiotic stresses. However, thorough evaluation of its salt stress tolerance and screening for traits that could be reliably applied for salt tolerance needs to be studied. The current study was designed to characterize 25 E. sativa accessions, originating from diverse geographical regions of Pakistan, for the salt stress tolerance. Methods Salt stress (150 mM NaCl) was applied for 2 weeks to the plants at four leaf stage in hydroponics. Data of the following morpho-physiological traits were collected from control and treated plants of all the accessions: root length (RL), shoot length (SL), plant height (PH), leaf number (LN), leaf area (LA), fresh weight (FW), dry weight (DW), chlorophyl content (SPAD), electrolyte leakage (EL), relative water content (RWC), gas exchange parameters and mineral ion content. Salt tolerance was determined based on membership function value (MFV) of the tested traits. Results Compared with control, the salt-stressed group had significantly reduced mean SL, RL, PH, LN, LA, FW, DW and SPAD. NaCl treatment triggered a slight increase in EL in few accessions. Mean RWC of control and treated groups were not significantly different although few accessions exhibited variation in this trait. Salt stress caused a significant reduction in photosynthesis rate (PR), transpiration rate (TR) and stomatal conductance (SC) but intercellular CO2 (Ci) was not significantly different between control and treated groups. Compared with control, the salt-stressed plants accumulated significantly higher Na+, K+ and Ca2+ while significantly lower Mg2+. K+/Na+ ratio was significantly decreased in salt-stressed plants compared with control. Importantly, significant inter-accession variations were found for all the tested traits. The principal component analysis identified SL, RL, PH, LN, LA, FW, DW and PR as the most significant traits for resolving inter-accession variability. Based on MFV of the tested traits, accessions were categorized into five standard groups. Among 25 accessions, one accession was ranked as highly tolerant, four as tolerant while 15 accessions were ranked as moderately tolerant. Of the remaining five accessions, four were ranked as sensitive while one accession as highly sensitive. Conclusion E. sativa accessions were found to exhibit significant genetic diversity in all the tested traits. A few most significant traits for dissecting the genetic variability were identified that could be used for future large-scale germplasm screening in E. sativa. Salt tolerant accessions could be a good resource for future breeding programs aiming to improve salt stress tolerance.

Amaranthus cruentus L. is a forage species, with grains that exhibit excellent nutritional characteristics, being the 'BRS Alegria' the first cultivar recommended for cultivation in Brazil. This study aimed at evaluating the effect of salt stress on the growth and concentrations of organic and inorganic solutes in Amaranthus cruentus L. ('BRS Alegria' cultivar). Height, stem diameter, number of leaves, leaf, stem and root dry mass, leaf area, relative water content and membrane integrity percentage, as well as soluble carbohydrate, free amino acid, soluble protein, free proline, Na+, Cl- and K+ contents, were evaluated in different plant organs. Salinity significantly reduced the biomass yield and leaf area from the treatment with 25 mM of NaCl, indicating that the 'BRS Alegria' cultivar is sensitive to salt stress. Soluble carbohydrates in the leaves decreased by 59 %, while the other organic solutes showed no substantial increases. These results, coupled with the reduction in the relative water content and membrane integrity, suggest a low ability of this cultivar to adjust osmotically under salt stress. The K+/Na+ ratio abruptly decreased in 25 mM of NaCl, suggesting an ionic imbalance, which may partially explain the salt-induced growth reduction.

Salt stress is one of the devastating factors that hampers growth and productivity of soybean. Use of Pseudomonas pseudoalcaligenes to improve salt tolerance in soybean has not been thoroughly explored yet. Therefore, we observed the response of hydroponically grown soybean plants, inoculated with halotolerant P. pseudoalcaligenes (SRM-16) and Bacillus subtilis (SRM-3) under salt stress. In vitro testing of 44 bacterial isolates revealed that four isolates showed high salt tolerance. Among them, B. subtilis and P. pseudoalcaligenes showed ACC deaminase activity, siderophore and indole acetic acid (IAA) production and were selected for the current study. We determined that 106 cells/mL of B. subtilis and P. pseudoalcaligenes was sufficient to induce tolerance in soybean against salinity stress (100 mM NaCl) in hydroponics by enhancing plant biomass, relative water content and osmolytes. Upon exposure of salinity stress, P. pseudoalcaligenes inoculated soybean plants showed tolerance by the increased activities of defense related system such as ion transport, antioxidant enzymes, proline and MDA content in shoots and roots. The Na+ concentration in the soybean plants was increased in the salt stress; while, bacterial priming significantly reduced the Na+ concentration in the salt stressed soybean plants. However, the antagonistic results were observed for K+ concentration. Additionally, soybean primed with P. pseudoalcaligenes and exposed to 100 mM NaCl showed a new protein band of 28 kDa suggesting that P. pseudoalcaligenes effectively reduced salt stress. Our results showed that salinity tolerance was more pronounced in P. pseudoalcaligenes as compared to B. subtilis. However, a detailed study at molecular level to interpret the mechanism by which P. pseudoalcaligenes alleviates salt stress in soybean plants need to be explored.

The cyclic nucleotide cyclic guanosine monophosphate (cGMP) is a powerful cell signaling molecule involved in biotic and abiotic stress perception and signal transduction. In the model plant Arabidopsis thaliana, salt and osmotic stress rapidly induce increase in cGMP which plays role by modulating the activity of monovalent cation transporters, possibly by direct binding to these proteins and by altering the expression of many abiotic stress responsive genes. In a recent study, a membrane permeable analogue of cGMP (8-bromo-cGMP) was found to have a promotive effect on soluble sugar, flavonoids and lignin content, and membrane integrity in Solanum lycopersicum seedlings under salt stress. However, it remains to be elucidated how salt stress affects the endogenous cGMP level in S. lycopersicum and if Br-cGMP-induced improvement in salt tolerance in S. lycopersicum involves altered cation fluxes. The current study was conducted to answer these questions. A rapid increase (within 30 s) in endogenous cGMP level was determined in S. lycopersicum roots after treatment with 100 mM NaCl. Addition of membrane permeable Br-cGMP in growth medium remarkably ameliorated the inhibitory effects of NaCl on seedlings’ growth parameters, chlorophyll content and net photosynthesis rate. In salt stressed plants, Br-cGMP significantly decreased Na+ content by reducing its influx and increasing efflux while it improved plants K+ content by reducing its efflux and enhancing influx. Furthermore, supplementation with Br-cGMP improved plant’s proline content and total antioxidant capacity, resulting in markedly decreased electrolyte leakage under salt stress. Br-cGMP increased the expression of Na+/H+ antiporter genes in roots and shoots of S. lycopersicum growing under salt stress, potentially enhancing plant’s ability to sequester Na+ into the vacuole. The findings of this study provide insights into the mechanism of cGMP-induced salt stress tolerance in S. lycopersicum.

Seed priming with Beauveria bassiana improves growth and salt stress response in rice

Streptomyces rimosus rhizobacteria and Glomus mosseae mycorrhizal fungus inoculation alleviate salinity stress in grapevine through morphophysiological changes and nutritional balance

Ion transport is the fundamental factor determining salinity tolerance in plants. The Review starts from differences in ion transport between salt tolerant halophytes and salt-sensitive plants with an emphasis on transport of potassium and sodium via plasma membranes. The comparison provides introductory information for increasing salinity tolerance. Effects of salt stress on ion transport properties of membranes show huge opportunities for manipulating ion fluxes. Further steps require knowledge about mechanisms of ion transport and individual genes of ion transport proteins. Initially, the Review describes methods to measure ion fluxes, the independent set of techniques ensures robust and reliable basement for quantitative approach. The Review briefly summarizes current data concerning Na+ and K+ concentrations in cells, refers to primary thermodynamics of ion transport and gives special attention to individual ion channels and transporters. Simplified scheme of a plant cell with known transport systems at the plasma membrane and tonoplast helps to imagine the complexity of ion transport and allows choosing specific transporters for modulating ion transport. The complexity is enhanced by the influence of cell size and cell wall on ion transport. Special attention is given to ion transporters and to potassium and sodium transport by HKT, HAK, NHX, and SOS1 proteins. Comparison between non-selective cation channels and ion transporters reveals potential importance of ion transporters and the balance between the two pathways of ion transport. Further on the Review describes in detail several successful attempts to overexpress or knockout ion transporters for changing salinity tolerance. Future perspectives are questioned with more attention given to promising candidate ion channels and transporters for altered expression. Potential direction of increasing salinity tolerance by modifying ion channels and transporters using single point mutations is discussed and questioned. An alternative approach from synthetic biology is to create new regulation networks using novel transport proteins with desired properties for transforming agricultural crops. The approach had not been widely used earlier; it leads also to theoretical and pure scientific aspects of protein chemistry, structure-function relations of membrane proteins, systems biology and physiology of stress and ion homeostasis. Summarizing, several potential ways are aimed at required increase in salinity tolerance of plants of interest.

Bacillus velezensis HR6-1 enhances salt tolerance in tomato by increasing endogenous cytokinin content and improving ROS scavenging.

High salinity and nitrogen (N) deficiency in soil are two key factors limiting crop productivity, and they usually occur simultaneously. Here we firstly found that H(+) -PPase is involved in salt-stimulated NO3 (-) uptake in the euhalophyte Salicornia europaea. Then, two genes (named SeVP1 and SeVP2) encoding H(+) -PPase from S. europaea were characterized. The expression of SeVP1 and SeVP2 was induced by salt stress and N starvation. Both SeVP1 or SeVP2 transgenic Arabidopsis and wheat plants outperformed the wild types (WTs) when high salt and low N occur simultaneously. The transgenic Arabidopsis plants maintained higher K(+) /Na(+) ratio in leaves and exhibited increased NO3 (-) uptake, inorganic pyrophosphate-dependent vacuolar nitrate efflux and assimilation capacity under this double stresses. Furthermore, they had more soluble sugars in shoots and roots and less starch accumulation in shoots than WT. These performances can be explained by the up-regulated expression of ion, nitrate and sugar transporter genes in transgenic plants. Taken together, our results suggest that up-regulation of H(+) -PPase favours the transport of photosynthates to root, which could promote root growth and integrate N and carbon metabolism in plant. This work provides potential strategies for improving crop yields challenged by increasing soil salinization and shrinking farmland.

Acclimation to salt stress in plants is regulated by complex signaling pathways involving endogenous phytohormones. The signaling role of salicylic acid (SA) in regulating crosstalk between endogenous plant growth regulators’ levels was investigated in barley (Hordeum vulgare L. ‘Ince’; 2n = 14) leaves and roots under salt stress. Salinity (150 and 300 mM NaCl) markedly reduced leaf relative water content (RWC), growth parameters, and leaf water potential (LWP), but increased proline levels in both vegetative organs. Exogenous SA treatment did not significantly affect salt-induced negative effects on RWC, LWP, and growth parameters but increased the leaf proline content of plants under 150 mM salt stress by 23.1%, suggesting that SA enhances the accumulation of proline, which acts as a compatible solute that helps preserve the leaf’s water status under salt stress. Changes in endogenous phytohormone levels were also investigated to identify agents that may be involved in responses to increased salinity and exogenous SA. Salt stress strongly affected endogenous cytokinin (CK) levels in both vegetative organs, increasing the concentrations of CK free bases, ribosides, and nucleotides. Indole-3-acetic acid (IAA, auxin) levels were largely unaffected by salinity alone, especially in barley leaves, but SA strongly increased IAA levels in leaves at high salt concentration and suppressed salinity-induced reductions in IAA levels in roots. Salt stress also significantly increased abscisic acid (ABA) and ethylene levels; the magnitude of this increase was reduced by treatment with exogenous SA. Both salinity and SA treatment reduced jasmonic acid (JA) levels at 300 mM NaCl but had little effect at 150 mM NaCl, especially in leaves. These results indicate that under high salinity, SA has antagonistic effects on levels of ABA, JA, ethylene, and most CKs, as well as basic morphological and physiological parameters, but has a synergistic effect on IAA, which was well exhibited by principal component analysis (PCA).

Ectopic expression of BrALDH7B2 gene encoding an antiquitin from Brassica rapa confers tolerance to abiotic stresses and improves photosynthetic performance under salt stress in tobacco