HvNax3—a locus controlling shoot sodium exclusion derived from wild barley (Hordeum vulgare ssp. spontaneum)

Previous work identified the wild barley (Hordeum vulgare ssp. spontaneum) accession CPI-71284-48 as being capable of limiting sodium (Na⁺) accumulation in the shoots under saline hydroponic growth conditions. Quantitative trait locus (QTL) analysis using a cross between CPI-71284-48 and a selection of the cultivated barley (H. vulgare ssp. vulgare) cultivar Barque (Barque-73, a moderate Na⁺ excluder) attributed the control of the Na⁺ exclusion trait from CPI-71284-48 to a single locus on the short arm of chromosome 7H, which was named HvNax3. The locus reduced shoot Na⁺ accumulation by 10–25% in plants grown in 150 mM NaCl. Markers generated using colinearity with rice and Brachypodium, together with the analysis of introgression lines and F₂ and F₃ families, enabled HvNax3 to be mapped to a 1.3-cM interval. Genes from the corresponding rice and Brachypodium intervals encode 16 different classes of proteins and include several plausible candidates for HvNax3. The potential of HvNax3 to provide a useful trait contributing to salinity tolerance in cultivated barley is discussed.     

A novel protein kinase involved in Na+ exclusion revealed from positional cloning

Salinity is a major abiotic stress which affects crop plants around the world, resulting in substantial loss of yield and millions of dollars of lost revenue. High levels of Na⁺ in shoot tissue have many adverse effects and, crucially, yield in cereals is commonly inversely proportional to the extent of shoot Na⁺ accumulation. We therefore need to identify genes, resistant plant cultivars and cellular processes that are involved in salinity tolerance, with the goal of introducing these factors into commercially available crops. Through the use of an Arabidopsis thaliana mapping population, we have identified a highly significant quantitative trait locus (QTL) linked to Na⁺ exclusion. Fine mapping of this QTL identified a protein kinase (AtCIPK16), related to AtSOS2, that was significantly up‐regulated under salt stress. Greater Na⁺ exclusion was associated with significantly higher root expression of AtCIPK16, which is due to differences in the gene's promoter. Constitutive overexpression of the gene in Arabidopsis leads to plants with significant reduction in shoot Na⁺ and greater salinity tolerance. amiRNA knock‐downs of AtCIPK16 in Arabidopsis show a negative correlation between the expression levels of the gene and the amount of shoot Na⁺. Transgenic barley lines overexpressing AtCIPK16 show increased salinity tolerance.     

Control of xylem Na+ loading and transport to the shoot in rice and barley as a determinant of differential salinity stress tolerance

Control of xylem Na⁺ loading has often been named as the essential component of salinity tolerance mechanism. However, it is less clear to what extent the difference in this trait may determine differential salinity tolerance between species. In this study, barley (Hordeum vulgare L. cv. CM72) and rice (Oryza sativa L. cv. Dongjin) plants were grown under two levels of salinity. Na⁺ and K⁺ concentrations in the xylem sap, and shoot and root tissues were measured at different time points after stress onset. Salt‐exposed rice plants prevented xylem Na⁺ loading for several days, but failed to control this process in the longer term, ultimately resulting in a massive Na⁺ shoot loading. Barley plants quickly increased xylem Na⁺ concentration and its delivery to the shoot (most likely for the purpose of osmotic adjustment) but were able to reduce this process later on, keeping most of accumulated Na⁺ in the root, thus maintaining non‐toxic shoot Na⁺ level. Rice plants increased shoot K⁺ concentration, while barley plants maintained higher root K⁺ concentration. Control of xylem Na⁺ loading is remarkably different between rice and barley; this difference may differentiate the extent of the salinity tolerance between species. This trait should be investigated in more detail to be used in the breeding programs aimed to improve salinity tolerance in crops.     

Identification of quantitative trait loci for ion homeostasis and salt tolerance in barley (Hordeum vulgare L.)

Ion homeostasis is considered to be one of the most important mechanisms underlying salt stress tolerance. We used the Steptoe × Morex barley doubled haploid population to screen for genetic variation in response to salinity stress at an early development stage in a hydroponics system, focusing on ion homeostasis. Salinity induced a strong adverse effect on growth of the parents and their derived population, with Steptoe as the more tolerant parent. Steptoe maintained higher concentrations of K⁺, Na⁺ and Cl^? in the roots and a similar shoot/root ion ratio (<1) under stress conditions compared to control conditions. In contrast, Morex had higher concentrations of these ions in the shoots under stress and a doubled shoot/root ion ratio relative to control conditions, indicating that salt exclusion might contribute to the higher tolerance of Steptoe. Correlation and path analysis demonstrated that shoot Cl^? contents most strongly affected salt tolerance and suggest that both Na⁺ and Cl^? contents are important for salinity stress tolerance in barley. We identified 11 chromosomal regions involved in the control of the variation observed for salt tolerance and various salt stress response traits, including Na⁺, Cl^? and K⁺ contents in shoots. Two specific regions on chromosomes 2H and 3H were found controlling ion contents and salt tolerance, pointing to genes involved in ion homeostasis that contribute to salt tolerance.     

A major terminal drought tolerance QTL of pearl millet is also associated with reduced salt uptake and enhanced growth under salt stress

The performance of a major quantitative trait locus (QTL) of terminal drought tolerance (DT) of pearl millet was assessed under salt stress. The test-cross hybrids of the QTL donor parent (drought tolerant, PRLT 2/89-33), QTL recipient parent (drought sensitive, H 77/833-2), and a set of six near isogenic lines introgressed with a terminal DT-QTL (QTL-NILs) were evaluated for germination and seedling emergence at 7 days after sowing (DAS) in Petri plates at four salinity levels, and at vegetative (24 DAS) and maturity stages at three salinity and alkalinity levels. Na⁺ and K⁺ accumulation, their compartmentation in different plant parts, and their effects on growth and yield parameters were evaluated. The DT-QTL donor parent and QTL-NILs accumulated less Na⁺ in shoot parts at seedling, vegetative and maturity stages, and also partitioned the accumulated Na⁺ more into nodes and internodes and less into leaves than the drought-sensitive recurrent parent. The pattern of reduced salt accumulation in the drought-tolerant parent and QTL-NILs was consistently associated with better growth and productivity in saline and alkaline treatments. It is concluded that the DT-QTL contributed by PRLT 2/89-33 exerted favourable effects on growth and productivity traits under salt stress by limiting Na⁺ accumulation in leaves.     

How much sodium accumulation is necessary for salt tolerance in subspecies of the halophyte Atriplex canescens?

Two sympatric subspecies of the xerohalophyte Atriplex canescens Pursh. (Nutt.) were compared for 84 d in outdoor salinity trials in their native coastal desert environment in Sonora, Mexico. Subspecies linearis grows naturally on sea water in the high intertidal zone of estuaries while subspecies canescens grows on dunes. In lysimeter pot experiments, ssp. linearis exhibited 50% growth reduction when the mean root zone salinity reached 1160 mol m⁻³ NaCl compared to just 760 mol m⁻³ for ssp. canescens. When irrigated with sea water in a flood plot, ssp. linearis had 50% higher growth rates than ssp. canescens. The specialization of ssp. linearis for a saline environment was associated with greater net transport of Na⁺ from root to shoot, greater Na⁺ accumulation in the leaves and a higher Na:K ratio in the leaves compared to ssp. canescens. On the other hand, the two subspecies achieved approximately the same degree of osmotic adjustment in the leaves, equal to two to three times the external salinity, and had similar water use efficiencies. Even at relatively low salinities, both subspecies accumulated larger quantities of Na⁺ for osmotic adjustment than K⁺. The results suggest that breeding for Na⁺ accumulation rather than exclusion might be the more effective strategy for improving salt tolerance of conventional crop plants.     

Ecosystem changes in Galápagos highlands by the invasive tree Cinchona pubescens

Background and aims

Salinity is an increasing problem for agricultural production worldwide. Understanding how Na⁺ enters plants is important if reducing Na⁺ influx, a key component of the regulation of Na⁺ accumulation in plants and improving salt tolerance of crop plants, is to be achieved. Our previous work indicated that two distinct low-affinity Na⁺ uptake pathways exist in the halophyte Suaeda maritima. Here, we report the external NaCl concentration at which uptake switches from pathway 1 to pathway 2 and the kinetics of the interaction between external K⁺ concentration and Na⁺ uptake and accumulation in S. maritima in order to determine the roles of K⁺ transporters or channels in low-affinity Na⁺ uptake.

Methods

Na⁺ influx, Na⁺ and K⁺ accumulations in S. maritima exposed to various concentrations of NaCl (0–200 mM) were analyzed in the absence and presence of the inhibitors TEA and Ba⁺ (5 mM TEA or 3 mM Ba²⁺) or KCl (0, 10 or 50 mM).

Results

Our earlier proposal was confirmed and extended that there are two distinct low-affinity Na⁺ uptake pathways in S. maritima: pathway 1 might be mediated by a HKT-type transporter under low salinity conditions and pathway 2 by an AKT1-type channel or a KUP/HAK/KT type transporter under high salinity conditions. The external NaCl concentration at which two distinct low-affinity Na⁺ uptake switches from pathway 1 to pathway 2, the ‘turning point’, is between 90 and 95 mM. Over a short period (12 h) of Na⁺ and K⁺ treatments, a low concentration of K⁺ (10 mM) facilitated Na⁺ uptake by S. maritima under high salinity (100–200 mM NaCl), whether or not the plants had been subjected to a longer (3 d) period of K⁺ starvation. The kinetics suggests that low concentration of K⁺ (10 mM) might activate AKT1-type channels or KUP/HAK/KT-type transporters under high salinity (100–200 mM NaCl).

Conclusions

The turning-point of external NaCl concentrations for the two low-affinity Na⁺ uptake pathways in Suaeda maritima is between 90 and 95 mM. A low concentration of K⁺ (10 mM) might activate AKT1 or KUP/HAK/KT and facilitate Na⁺ uptake under high salinity (100–200 mM NaCl). The kinetics of K⁺ on Na⁺ uptake and accumulation in S maritima are also consistent with there being two low-affinity Na⁺ uptake pathways.     

Identification of quantitative trait loci contributing to yield and seed quality parameters under terminal drought in barley advanced backcross lines

Drought is a major limiting factor for barley production, especially in the primary areas of its cultivation. Wild barley represents a major source of favourable alleles for increasing the genetic variation for multiple traits including resistance to both biotic and abiotic stresses. We used advanced backcross quantitative trait locus (AB-QTL) analysis of a BC3-doubled haploid population developed between the cultivated parent Brenda (Hordeum vulgare ssp. vulgare) and the wild accession HS584 (H. vulgare ssp. spontaneum) to study the contribution of wild barley in improving various agronomic and seed quality traits under post-anthesis drought. The experiment was carried out at two different locations (IPK, Gatersleben and Nordsaat, Böhnshausen) and terminal drought was imposed by withholding water or spraying with potassium iodide at 10 days after flowering under greenhouse or field conditions, respectively. QTL analysis indicated that wild barley contributed favourably to most of the traits studied under both control and drought conditions. A total of seven hot-spot QTL regions with co-localizing QTL for various traits harboured more than 80 % of the stable QTL detected in the present study. For yield and thousand-grain weight and their respective drought tolerance indices, most of the QTL were derived from Brenda. On the other hand, for traits like seed length and seed nitrogen content, all the QTL were contributed by HS584, the parent having higher trait value. A significantly reduced carbon/nitrogen (C/N) ratio in the selected contrasting inferior lines compared to superior ones suggests that C/N ratio could be a potential parameter for screening not just seed quality parameters but also grain weight performance under terminal drought.     

Oxygen-Dependent Exclusion of Sodium Ions from Shoots by Roots of Zea mays (cv Pioneer 3906) in Relation to Salinity Damage

Using radio-tracers, we measured Na⁺ and K⁺ accumulation in roots and transport to shoots in Zea mays (cv Pioneer 3906) as a function of NaCl concentration and O₂ partial pressure in the nutrient solution. Under fully aerobic conditions, roots partially excluded Na⁺ from the shoots over a wide range of NaCl concentration (0.2-200 millimolar). With root anoxia, the exclusion mechanism broke down so that much greater amounts of Na⁺ reached the shoots, with simultaneous inhibition of K⁺ transport. The ratio Na⁺/K⁺ entering the shoot consequently increased 90 to 200 times. Increases in Na⁺ transport were first detected when the O₂ partial pressure was reduced from ambient (21% v/v) to 15%, whereas K⁺ transport was not inhibited until O₂ concentrations were <5%. Since soil O₂ deficiency can often accompany high salinity in irrigation agriculture, failure of the Na⁺ exclusion mechanism may be a contributory factor in salinity damage of salt-sensitive glycophytes.     

Accumulation of potassium and sodium by barley roots in a k-na replacement series

Excised roots of barley (Hordeum vulgare, var. Campana) were incubated for 24 hr in solutions containing constant total concentrations of KCl and NaCl but in which the mole fractions of K and Na were varied in replacement series. In solutions containing 1, 10, or 50 mm concentrations of K⁺ plus Na⁺, total cation accumulation was dependent upon the total salt concentration but was relatively independent of the mole fractions of K⁺ and Na⁺. These results imply that accumulation of K⁺ and Na⁺ was limited by a common factor. In solutions containing 0.01 mm K⁺ plus Na⁺ there was a strong preference for K⁺ over Na⁺ and the sum of K⁺ and Na⁺ accumulation increased with increasing K⁺ concentration.     

Evaluation of salinity tolerance in seedlings of sugar beet (Beta vulgaris L.) cultivars using proline, soluble sugars and cation accumulation criteria

Salinity tolerance of sugar beet (Beta vulgaris L.) cultivars in terms of growth, proline and soluble sugars concentrations, and Na⁺/K⁺ and Na⁺/Ca²⁺ ratios were analyzed in this study. Three-week-old seedlings of three sugar beet cultivars, ‘Gantang7’, ‘SD13829’, and ‘ST21916’, differing in salinity tolerance, were treated with 0, 50, 100, and 200 mM NaCl. Plant shoots and roots were harvested at 7 days after treatment and subjected to analysis. Low concentration of NaCl (50 mM) enhanced fresh and dry weights of shoot and root in ‘Gantang7’, whereas high one (200 mM) reduced growth in all cultivars and the less reduction was observed in ‘ST21916’. Shoot proline was strongly induced by salinity stress in both ‘Gantang7’ and ‘SD13829’, while it remained unchanged in ‘ST21916’. The addition of 50 mM NaCl significantly increased shoot soluble sugars concentrations in ‘Gantang7’ while it had no significant effects in the other two cultivars. ‘Gantang7’ also showed a higher level of root soluble sugars concentration as compared to the other two cultivars. At 50 mM NaCl, the lower shoot Na⁺ concentration, and the higher shoot K⁺ and root Ca²⁺ concentration in ‘Gantang7’ resulted in the lower shoot Na⁺/K⁺ and root Na⁺/Ca²⁺ ratio. However, ‘SD13829’ maintained a lower Na⁺/K⁺ ratio in both shoot and root when subjected to 200 mM NaCl treatment. According to comprehensive evaluation on salinity tolerance, it is clear that ‘Gantang7’ is more tolerant to salinity than the other two cultivars. Therefore, it is suggested that ‘Gantang7’ should be more suitable for cultivating in the arid and semi-arid irrigated regions.     

New perspective on the mechanism of alleviating salt stress by spermidine in barley seedlings

Salt stress is considered to be a major limiting factor for plant growth and crop productivity. Salt injuries in plants are mostly due to excess Na⁺ entry. A possible survival strategy of plants under saline environments is the effective compartmentation of excess Na⁺ by sequestering Na⁺ in roots and inhibiting transport of Na⁺ from roots to shoots. Our previous study showed that exogenous application of polyamines (PAs) could attenuate salt injuries in barley plants. In order to further understand such protective roles of PAs against salt stress, the effects of spermidine (Spd) on sodium and potassium distribution in barley (Hordeum vulgare L.) seedlings under saline conditions were investigated. The results showed that exogenous application of Spd induced reductions in Na⁺ levels in roots and shoots with comparison of NaCl-treated plants, while no significant changes in K⁺ levels were observed. Correspondingly, the plants treated with Spd exogenously maintained high values of [K⁺]/[Na⁺] as compared with salt-stressed plants. Moreover, it was shown by X-ray microanalysis that K⁺ and Na⁺ accumulated mainly in the exodermal intercellular space and cortical cells of roots under salinity stress, and low accumulation was observed in endodermal cells and stelar parenchyma, indicating Casparian bands possibly act as ion transport barriers. Most importantly, Spd treatment further strengthened this barrier effects, leading to inhibition of Na⁺ transport into shoots. These results suggest that, by reinforcing barrier effects of Casparian bands, exogenous Spd inhibits Na⁺ transport from roots to shoots under conditions of high salinity which are beneficial for attenuating salt injuries in barley seedlings.     

Sodium and potassium compartmentation and transport in the roots of intact lettuce plants

In this report, we consider the accumulation in roots, and transport to the shoot, of Na⁺ and K⁺ in intact lettuce plants (Lactuca sativa cv Black-seeded Simpson). Plants were grown in modified Hoagland medium supplemented with 10 moles NaCl per cubic meter. At this salinity, significant levels of Na⁺ were accumulated in roots and shoots, but there was no reduction in plant growth. Transport characteristics for both Na⁺ and K⁺ were qualitatively similar to those previously reported, for Spergularia marina, indicating that the results obtained with these experimental protocols are not limited to one unconventional experimental plant. The most pronounced difference in transport of the two ions was evident when transport was followed in a chase period after a 10 minute uptake pulse. For Na⁺, there was an initially rapid, but small, loss of label to the medium, and very little movement to the shoot. For K⁺, little label was lost from the plants, but translocation to the shoot proceeded for at least 60 minutes. The transport systems were further distinguished by treating the roots during labeling with 20 micrograms per milliliter cycloheximide. For K⁺, both uptake and translocation were reduced by about 50%. For Na⁺, root accumulation was stimulated more than five-fold, while transport to the shoot was reduced about 20%. Cycloheximide also modified the Na⁺ transport characteristics such that continued translocation occurred during the chase period of pulse-chase studies.     

Evidence that arbuscular mycorrhizal and phosphate-solubilizing fungi alleviate NaCl stress in the halophyte Kosteletzkya virginica: nutrient uptake and ion distribution within root tissues

The effects of an arbuscular mycorrhizal (AM) fungus, Glomus mosseae, and a phosphate-solubilizing microorganism (PSM), Mortierella sp., and their interactions, on nutrient (N, P and K) uptake and the ionic composition of different root tissues of the halophyte Kosteletzkya virginica (L.), cultured with or without NaCl, were evaluated. Plant biomass, AM colonization and PSM populations were also assessed. Salt stress adversely affected plant nutrient acquisition, especially root P and K, resulting in an important reduction in shoot dry biomass. Inoculation of the AM fungus or/and PSM strongly promoted AM colonization, PSM populations, plant dry biomass, root/shoot dry weight ratio and nutrient uptake by K. virginica, regardless of salinity level. Ion accumulation in root tissues was inhibited by salt stress. However, dual inoculation of the AM fungus and PSM significantly enhanced ion (e.g., Na⁺, Cl^?, K⁺, Ca²⁺, Mg²⁺) accumulation in different root tissues, and maintained lower Na⁺/K⁺ and Ca²⁺/Mg²⁺ ratios and a higher Na⁺/Ca²⁺ ratio, compared to non-inoculated plants under 100 mM NaCl conditions. Correlation coefficient analysis demonstrated that plant (shoot or root) dry biomass correlated positively with plant nutrient uptake and ion (e.g., Na⁺, K⁺, Mg²⁺ and Cl^?) concentrations of different root tissues, and correlated negatively with Na⁺/K⁺ ratios in the epidermis and cortex. Simultaneously, root/shoot dry weight ratio correlated positively with Na⁺/Ca²⁺ ratios in most root tissues. These findings suggest that combined AM fungus and PSM inoculation alleviates the deleterious effects of salt on plant growth by enabling greater nutrient (e.g., P, N and K) absorption, higher accumulation of Na⁺, K⁺, Mg²⁺ and Cl^? in different root tissues, and maintenance of lower root Na⁺/K⁺ and higher Na⁺/Ca²⁺ ratios when salinity is within acceptable limits.     

Differences in efficient metabolite management and nutrient metabolic regulation between wild and cultivated barley grown at high salinity

Physiological and biochemical responses of Hordeum maritimum and H. vulgare to salt stress were studied over a 60‐h period. Growth at increasing salinity levels (0, 100, 200 and 300 mM NaCl) was assessed in hydroponic culture. H. maritimum was shown to be a true halophyte via its typical behaviour at high salinity. Shoot growth of cultivated barley was gradually reduced with increasing salinity, whereas that of wild barley was enhanced at 100 and 200 mm NaCl then slightly reduced at 300 mM NaCl. The higher salt tolerance of H. maritimum as compared to H. vulgare was due to its higher capacity to maintain cell turgor under severe salinity. Furthermore, H. maritimum exhibited fine regulation of Na⁺ transport from roots to shoots and, unlike H. vulgare, it accumulated less Na⁺ in shoots than in roots. In addition, H. maritimum can accumulate more Na⁺ than K⁺ in both roots and shoots without the appearance of toxicity symptoms, indicating that Na⁺ was well compartmentalized within cells and substituted K⁺ in osmotic adjustment. The higher degree of salt tolerance of H. maritimum is further demonstrated by its economic strategy: at moderate salt treatment (100 mm NaCl), it used inorganic solutes (such as Na⁺) for osmotic adjustment and kept organic solutes and a large part of the K⁺ for metabolic activities. Indeed, K⁺ use efficiency in H. maritimum was about twofold that in H. vulgare; the former started to use organic solutes as osmotica only at high salinity (200 and 300 mm NaCl). These results suggest that the differences in salt tolerance between H. maritimum and H. vulgare are partly due to (i) differences in control of Na⁺ transport from roots to shoots, and (ii) H. maritimum uses Na⁺ as an osmoticum instead of K⁺ and organic solutes. These factors are differently reflected in growth.     

Comparison of salt tolerance and osmotic adjustment of low-sodium and high-sodium subspecies of the C4 halophyte, Atriplex canescens

The relationship between Na⁺ accumulation and salt tolerance was tested by comparing subspecies of the halophyte, Atriplex canescens (fourwing saltbush), that differed markedly in Na⁺ content and Na:K ratios. Above ground tissues of one low-sodium and two high-sodium subspecies were compared with respect to cation accumulation, osmotic adjustment and growth along a salinity gradient in greenhouse trials. Plants of each subspecies were grown for 80 d on 2.2, 180, 540 and 720 mol m^?3 NaCl. At harvest, A. canescens ssp. canescens had significantly lower Na⁺ levels, higher K⁺ levels and lower Na:K ratios in leaf and stem tissues than A. canescens ssp. macropoda and linearis over the salinity range (P < 0.05 or 0.01). Na:K ratios in leaves of the latter two, high-sodium, subspecies were approximately 2 on the lowest salinity treatment and ranged from 5 to 10 on the more saline solutions. By contrast, Na:K ratios in leaves of the low-sodium subspecies canescens, were only 0.4 on the lowest salinity and ranged narrowly from 1.7 to 2.3 at higher salinities. However, despite different patterns of Na⁺ and K⁺ accumulation, all three subspecies exhibited equally high salt tolerance and had similar osmotic pressures in their leaves or stems over the salinity range. Contrary to expectations, high salt tolerance was not necessarily dependent on high levels of Na⁺ accumulation in this species.     

Functional analyses of a putative plasma membrane Na+/H+ antiporter gene isolated from salt tolerant Helianthus tuberosus

Jerusalem artichokes (Helianthus tuberosus L.) can tolerate relatively higher salinity, drought and heat stress. In this paper, we report the cloning of a Salt Overly Sensitive 1 (SOS1) gene encoding a plasma membrane Na⁺/H⁺ antiporter from a highly salt-tolerant genotype of H. tuberosus, NY1, named HtSOS1 and characterization of its function in yeast and rice. The amino acid sequence of HtSOS1 showed 83.4 % identity with the previously isolated SOS1 gene from the Chrysanthemum crassum. The mRNA level in the leaves of H. tuberosus was significantly up-regulated by presence of high concentrations of NaCl. Localization analysis using rice protoplast expression showed that the protein encoded by HtSOS1 was located in the plasma membrane. HtSOS1 partially suppressed the salt sensitive phenotypes of a salt sensitive yeast strain. In comparison with wild type (Oryza sativa L., ssp. Japonica. cv. Nipponbare), the transgenic rice expressed with HtSOS1 could exclude more Na⁺ and accumulate more K⁺. Expression of HtSOS1 decreased Na⁺ content much larger in the shoot than in the roots, resulting in more water content in the transgenic rice than WT. These data suggested that HtSOS1 may be useful in transgenic approaches to improving the salinity tolerance of glycophyte.     

Effect of salt stress on photosynthesis and physiological parameters of three contrasting barley genotypes

In order to understand the physiological traits important in conferring salt tolerance in three barley genotypes, this study was performed under field conditions with three water salinity levels (2, 10, and 18 dS m–1). High salinity decreased net photosynthetic rate, transpiration rate, and stomatal conductance, K⁺ concentration, K⁺:Na⁺ ratio, and grain yield, but increased electrolyte leakage and Na⁺ content. Under 10 and 18 dS m–1 salinity, Khatam (salt-tolerant) had the maximum stomatal conductance, K⁺, K⁺:Na⁺ ratio, and the grain yield, and a minimum Na⁺ content and electrolyte leakage, whereas Morocco (salt-sensitive) had the lowest net photosynthetic rate, stomatal conductance, K⁺ content, K⁺:Na⁺ ratio, and grain yield, and the highest Na⁺ content and electrolyte leakage. This study showed that tolerant genotypes of barley may avoid Na⁺ accumulation in aboveground parts, facilitating a higher photosynthetic rate and higher grain yield.     

Responses of Atriplex spongiosa and Suaeda monoica to Salinity

The growth and tissue water, K⁺, Na⁺, Cl⁻, proline and glycinebetaine contents of the shoots and roots of two Chenopodiaceae, Atriplex spongiosa and Suaeda monoica have been measured over a range of external NaCl salinities. Both species showed some fresh weight response to low salinity mainly due to increased succulence. S. monoica showed both a greater increase in succulence (at low salinities) and tolerance of high salinities than A. spongiosa. Both species had high affinities for Na⁺ and maintained constant but low shoot K⁺ contents with increasing salinity. These trends were more marked with S. monoica in which Na⁺ stimulated the accumulation of K⁺ in roots. An association between high leaf Na⁺ accumulation, high osmotic pressure, succulence, and a positive growth response at low salinities was noted. Proline accumulation was observed in shoot tissues with suboptimal water contents. High glycinebetaine contents were found in the shoots of both species. These correlated closely with the sap osmotic pressure and it is suggested that glycinebetaine is the major cytoplasmic osmoticum (with K⁺ salts) in these species at high salinities. Na⁺ salts may be preferentially utilized as vacuolar osmotica.     

Sodium exclusion QTL associated with improved seedling growth in bread wheat under salinity stress

Worldwide, dryland salinity is a major limitation to crop production. Breeding for salinity tolerance could be an effective way of improving yield and yield stability on saline-sodic soils of dryland agriculture. However, this requires a good understanding of inheritance of this quantitative trait. In the present study, a doubled-haploid bread wheat population (Berkut/Krichauff) was grown in supported hydroponics to identify quantitative trait loci (QTL) associated with salinity tolerance traits commonly reported in the literature (leaf symptoms, tiller number, seedling biomass, chlorophyll content, and shoot Na⁺ and K⁺ concentrations), understand the relationships amongst these traits, and determine their genetic value for marker-assisted selection. There was considerable segregation within the population for all traits measured. With a genetic map of 527 SSR-, DArT- and gene-based markers, a total of 40 QTL were detected for all seven traits. For the first time in a cereal species, a QTL interval for Na⁺ exclusion (wPt-3114-wmc170) was associated with an increase (10%) in seedling biomass. Of the five QTL identified for Na⁺ exclusion, two were co-located with seedling biomass (2A and 6A). The 2A QTL appears to coincide with the previously reported Na⁺ exclusion locus in durum wheat that hosts one active HKT1;4 (Nax1) and one inactive HKT1;4 gene. Using these sequences as template for primer design enabled mapping of at least three HKT1;4 genes onto chromosome 2AL in bread wheat, suggesting that bread wheat carries more HKT1;4 gene family members than durum wheat. However, the combined effects of all Na⁺ exclusion loci only accounted for 18% of the variation in seedling biomass under salinity stress indicating that there were other mechanisms of salinity tolerance operative at the seedling stage in this population. Na⁺ and K⁺ accumulation appear under separate genetic control. The molecular markers wmc170 (2A) and cfd080 (6A) are expected to facilitate breeding for salinity tolerance in bread wheat, the latter being associated with seedling vigour.