Salt tolerance in Aster tripolium L. II. Ionic regulation

Abstract Measurements of tissue ion contents (Na, K and Cl) were carried out at frequent intervals on plants of Aster tripolium L. grown at a range of salinities for 36 d. Aster tripolium behaved as a typical halophyte showing high levels of inorganic ion accumulation even at low salinities. As salinity increased Na replaced K to a large extent in the shoot but root K was unaffected up to 500 mol m^?3 external NaCl. Shoot (Na + K) concentration on a tissue water basis was maintained constant in all treatments throughout the experiment, whereas shoot (Na + K) on a dry weight basis showed marked fluctuations in some treatments. An increase in (Na + K) per gram dry weight was, however, accompanied by a parallel increase in fresh weight: dry weight (FW : DW) ratio. Transport of (Na + K) to the shoot per unit root weight changed during the experiment in the manner expected, given the observed changes in shoot relative growth rate and FW : DW to result in a constant shoot (Na + K) concentration on a water basis. Chloride was the major balancing anion in the shoot at high salinity, but never accounted for more than 38% of the (Na + K) found in the root tissue. At all salinities (Na + K) salts accounted for the majority of the measured shoot sap osmotic potential. The interactions between salinity, growth, ion transport and osmotic adjustment are discussed.     

Differential growth of some grapevine varieties in Syria in response to salt <Emphasis Type="Italic">in vitro</Emphasis>

Summary Growth sensitivity of four local grapevine (Vitis vinifera) varieties, Ashlamesh, Helwani, Kassofee, and Khoudeiry, were evaluated for salt. They were cultured on DSD1 medium until rooting stage, then they were transferred to a liquid DSD1 medium containing 0, 10, 20, 30, 40, 80, 120, or 150 mM NaCl for 30 d. The shoot length and leaf number of Ashlamesh, Helwani, and Kassofee were significantly increased at 10 and/or 30 mM NaCl, whereas, 150 mM NaCl decreased shoot length of all varieties except Kassofee. The presence of NaCl at 80 mM or higher concentrations decreased the chlorophyll content and root number of all varieties, while 30 mM NaCl increased root number of Kassofec.     

Salt tolerance in Aster tripolium L. III. Na and K fluxes in intact seedlings

Abstract Uptake and transport of Na and K was studied using the radioactive tracers ²²Na and ⁴²K in intact Aster tripolium L. seedlings grown at two salinities CS 10 and CS 100, (containing 10mol m^?1 and 100 mol m^?3 Na, respectively, together with other major ions in the proportions found in sea water). At both salinities a much greater proportion of the Na than K taken up by the plant was subsequently transported to the shoot. Most ⁴²K fluxes were reduced by about 40% in CS 100 plants relative to CS 10 except root accumulation which increased. Experiments involving changing the salinity from CS 10 to CS 100 showed that ⁴²K fluxes remained constant for at least 40 h, indicating that competition with Na for uptake sites was not the cause of the reduced flux in CS 100 plants. ²²Na fluxes responded immediately to a change in salinity with all fluxes increasing six-fold when the salinity was raised. When the salinity was lowered, however, root accumulation returned to the level in CS 10 control plants whereas transport to the shoot was inhibited by the previous high salinity treatment, being reduced to only 35% of the rate in CS 10 plants. The time courses of osmotic adjustment and Na accumulation following an increase in salinity were found to be very similar, with sufficient Na being accumulated to account for the observed increase in sap osmotic pressure.     

Ionic regulation of the unicellular green algadunaliella tertiolecta

Summary Different techniques were investigated in order to determine the Na, K and Cl concentrations ofDunaliella tertiolecta cells adapted to a large range of salinity (20 to 1640 mM NaCl). The K cell concentrations were 6 to 13 times higher than the K concentration of the external medium (11 mM). The The Na and Cl cell concentrations, on the other hand, were lower than in the external medium at all salinities tested. Considerable differences in the absolute values of Na and Cl were, however, found according to the technique employed. These results are interpreted in terms of compartmentalization of the cells (at least two compartments). It is postulated that the larger compartment regulates its ion concentrations, maintaining low Na and Cl and high K concentrations, whereas the second compartment equilibrates with the external medium. The cation permeability of the membrane limiting the regulating compartment is altered by the antibiotics nystatin and monensin. Incubation of cells in K-free medium leads to a decrease of K and to an increase of the cell Na, this effect being reversed by addition of KCl to the medium. A good correlation is found between gain of K and loss of Na, suggesting a stoichiometric exchange of these two ions. The magnitude of this apparent Na/K exchange increases as the salinity increases. The external K concentration necessary to mediate half-saturation of the Na/K exchange is a function of the NaCl concentration of the adaptation medium. This Na/K exchange is partially light-dependant and inhibited by cold, cyanide and DCCD. It is suggested that this mechanism helps in the regulation of the ionic composition ofDunaliella cells.     

Plant growth and ion relations in lucerne (Medicago sativa L.) in response to the combined effects of NaCl and P

The combined effect of NaCl and P on the growth of lucerne was studied in two hydroponic greenhouse experiments. NaCl concentrations were identical in each experiment (0, 50 and 100 mM NaCl) while external P concentrations were low (viz. 0.002, 0.02 and 0.2 mM measured as 0.006, 0.026 and 0.2 mM, respectively) in one experiment and higher (0.5 and 5.0 mM) in the second. Plant biomass was reduced more by the low P levels than by high concentrations of NaCl. A significant NaCl^*P effect was found where external P concentrations were low (0.006–0.2 mM) but there was no difference in plant production between the two P concentrations of 0.5 and 5.0 mM. Shoot and root concentrations of Na and Cl increased significantly with increasing NaCl concentration in both experiments and there were some differences in the concentrations of these ions at different external P levels. At low P, NaCl had no significant effect on shoot concentrations of P; however, root P concentrations tended to decrease with increasing NaCl level. Increasing external P from 0.006 to 0.2 mM led to significant increases in P concentrations in both roots and shoots. At higher P, concentrations of P in both the shoots and the roots did not differ with external NaCl or P conditions. Our results illustrate the complex relationship that exists between NaCl and P at low P levels. We conclude that high or non-limiting concentrations of P (0.2 – 5.0 mM) do not affect lucerne's response to NaCl.     

Response of seeds ofLupinus termis andVicia faba to the interactive effect of salinity and ascorbic acid or pyridoxine

The interactive effect of salinity and presoaking in ascorbic acid or phyridoxine on germination, seedling growth, and some relevant metabolic changes ofLupinus termis andVicia faba seeds were studied. Germination studies indicated that broad bean tolerated NaCl salinity up to 240mM NaCl and lupin to 200mM NaCl. The lengths of roots and shoots and their water content, as well as dry matter yield, remained more or less unchanged up to the level of 80mM NaCl. Salinity induced marked progressive increases of carbohydrates and proline in broad bean and soluble protein in lupin seedlings, irrespective of the salinity level used. The other organic solutes (soluble protein in broad bean and carbohydrates in lupin seedlings) remained more or less unchanged at low and moderate levels of NaCl. However, under the higher salinity levels, in lupin the losses in carbohydrates were accompanied by increases in soluble protein, whereas in broad bean an opposite effect was obtained. The level of 40mM NaCl had a pronounced stimulatory effect on the all the variables studied. Presoaking seeds in either ascorbic acid or pyridoxine counteracted the adverse effects of salinity on germination and seedling growth as well as on some metabolic mechanisms of lupin and broad bean plants. The importance of these processes to the salinity tolerance of broad bean and lupin have been discussed.     

Response of tomato cultivars to salinity

The responses of five tomato cultivars (L. esculentum Mill) of different degrees of salt tolerance were examined over a range of 0 to 140 mM NaCl applied for 3 and 10 weeks. Judged by both Na and Cl accumulations and maintenance of K, Ca and Mg contents with increasing salinity, the most tolerant cultivars (Pera and GC-72) showed different responses. The greater salt tolerance of cv Pera was associated with a higher Cl and Na accumulation and a lower K content in the shoot than those found in the other cultivars, typical of a halophytic response to salinity. However, the greater salt tolerance of cv GC-72 was associated with a retention of Na and Cl in the root, restriction of their translocation to the shoot and maintenance of potassium selectivity under saline conditions. The salt tolerance mechanisms that operated in the remaining cultivars were similar to that of cv GC-72, as at first they excluded Na and Cl from the shoots, accumulating them in the roots; with longer treatment, the ability to regulate Na and Cl concentrations in the plant was lost only in the most salt sensitive cultivar (Volgogradskij), resulting in a massive influx of both ions into the shoot.The salt sensitivity of some tomato cultivars to salinity could be due to both the toxic effect of Na and Cl ions and nutritional imbalance induced by salinity, as plant growth was inversely correlated with Na and Cl contents and directly correlated with K and Ca contents. This study displays that there is not a single salt tolerance mechanism, since different physiological responses among tomato cultivars have been found.     

Effect of CaSO4 and NaCl on mineral content ofLeucaena leucocephala

The effects of varying CaSO₄ and NaCl levels on the nutrient content ofLeucaena leucocephala were established by examining the concentrations of Na, Ca, Cl, K and Mg in leucaena roots, stems and leaves. Leucaena was grown in nutrient solution at four levels of CaSO₄ (0.5, 1.0, 2.5 and 5.0 mM) and NaCl (1, 25, 50 and 100 mM), in randomized blocks with five replications. Leucaena excluded sodium from stems and leaves when NaCl concentration was 50 mM or less. Sodium uptake decreased as CaSO₄ concentration increased. Calcium uptake was affected by NaCl concentration when substrate CaSO₄ concentration was 0.5 mM. At this level, 100 mM NaCl caused a marked decrease in leaf calcium and a marked increase in leaf Cl. In all other treatments, Cl uptake was not affected by CaSO₄ concentration. Potassium uptake was strongly depressed as NaCl concentration increased at low Ca concentration, but this effect was offset at high Ca. Magnesium uptake decreased as CaSO₄ levels increased.     

Response of three wheat genotypes grown under saline medium to low/high potassium levels

A water culture experiment was conducted, to study the response of three wheat genotypes (Sarsabz, Kiran-95 and Pasban-90) to low and high potassium levels, (0.01 and 10 mM KC1) grown under two salinity concentrations (50 mM and 150 mM, NaCI). The results showed that the presence of sufficient potassium in the growth medium was found to bring good effects on plant growth. The data showed that shoot length of Kiran-95 growing under two salinities and associated with low and high potassium was quite satisfactory followed by Sarsabz and Pasban-90. Ionic content in plant shoots also varied with the increase in salinity levels of the medium. Potassium content in plant shoot was strongly regulated by Na⁺ ions, showing gradually decrease in K with the increase in Na accumulation in shoot. Under high salinities Kiran-95 had maximum K content in both low/high K supply, followed by Sarsabz and Pasban-90. This ability of Kiran-95 to maintain optimum K level may be the reason of its better survival.     

The effects of salinity on growth,nodulation and nitrogen fixation ofCasuarina equisetifolia

The growth, nodulation and nitrogen fixation ofCasuarina equisetifolia were compared at six levels (0–500mM NaCl) of salinity in sand culture. Dry weight of nodules, shoots and roots and N content of shoots increased at intermediate levels of salinity (50–100 mM) but decreased at 500 mM NaCl. Nodulation occurred at all NaCl levels, but at 500mM NaCl level, the nodule dry weight declined by 50% from the control. Increasing NaCl concentration of up to 200mM had little effect on the N₂-fixation rate, but at 500mM NaCl level the rate decreased to 40% of the control value.     

Combined effect of salinity and hypoxia in wheat (Triticum aestivum L.) and wheat-Thinopyrum amphiploids

The effects of sodium chloride salinity and hypoxia were studied in eight wheat lines and three wheat-Thinopyrum amphiploids in vermiculite-gravel culture. The lines were treated with either 100 or 150 mol m^–3 NaCl with and without hypoxia. Saline hypoxic conditions significantly reduced the vegetative growth, water use, grain and straw yields for all wheat varieties except the amphiploids, whereas NaCl or hypoxia alone had less pronounced effects. In addition, saline hypoxic stress reduced K⁺ concentration and increased significantly the Na⁺ and Cl^– concentrations in cell sap expressed from leaves. There was more Na⁺ and Cl^– accumulation in wheats than the amphiploids in hypoxic conditions at 150 mol m^–3 NaCl. Of the wheats, Pato was the most sensitive at all stress levels while aTriticum aestivum cv. Chinese Spring ×Thinopyrum elongatum amphiploid was the most tolerant of the three amphiploids.     

Counteraction of Salinity Stress on Wheat Plants by Grain Soaking in Ascorbic Acid, Thiamin or Sodium Salicylate

The interactive effects of salinity stress (40, 80, 120 and 160 mM NaCl) and ascorbic acid (0.6 mM), thiamin (0.3 mM) or sodium salicylate (0.6 mM) were studied in wheat (Triticum aestivum L.). The contents of cellulose, lignin of either shoots or roots, pectin of root and soluble sugars of shoots were lowered with the rise of NaCl concentration. On the other hand, the contents of hemicellulose and soluble sugars of roots, starch and soluble proteins of shoots, proline of either shoots or roots, and amino acids of roots were raised. Also, increasing NaCl concentration in the culture media increased Na⁺ and Ca²⁺ accumulation and gradually lowered K⁺ and Mg²⁺ concentration in different organs of wheat plant. Grain soaking in ascorbic acid, thiamin or sodium salicylate could counteract the adverse effects of NaCl salinity on the seedlings of wheat plant by suppression of salt stress induced accumulation of proline.     

Agronomical and physiological characterization of salinity tolerance in a commercial tomato hybrid

The salt tolerance of the commercial F1 tomato hybrid (Lycopersicon esculentum Mill) Radja (GC-793) has been agronomically and physiologically evaluated under greenhouse conditions, using a control (nutrient solution), a moderate (70 mM NaCl added to the nutrient solution) and a high salt level (140 mM NaCl), applied for 130 days. The results show that Radja is a Na⁺-excluder genotype, tolerant to moderate salinity. Fruit yield was reduced by 16% and 60% and the shoot biomass by 30% and more than 75% under moderate and high salinities, respectively. At 90 days of salt treatment (DST), the mature leaves feeding the 4th truss at fruiting accumulated little Na⁺ (178 mmol kg^-1 DW). At this time, the sucrose concentration in these leaves even increased with moderate salinity and the amino acid proline was not accumulated under salt conditions as compared to control. At 130 DST, Na⁺ was accumulated mainly by the roots in proportion to the salt level applied, while in leaves appreciable amounts were found only at high salinity (452 mmol kg^-1 DW). In the leaves, Cl^- was always accumulated in proportion to the salt level and in a very much greater amounts than Na⁺ (until 1640 mmol kg^-1 DW). The sucrose content was reduced in all plants by salinity, and was distributed preferentially toward the distal stem and peduncle of a truss at fruiting under moderate salinity, and toward the basal stem and root at high salinity. Moreover, proline was accumulated in different organs of the plant only at high salinity, coinciding with Na⁺ accumulation in leaves. Attempts are made to find a clear relationship between physiological behaviour triggered by stress and the agronomical behaviour, in order to assess the validity of physiological traits used for salt-tolerance selection and breeding in tomato.     

Salinity-calcium interactions on growth and ionic concentration of Citrus plants

Two-year-old Navel orange scions (Citrus sinensis (L.) Osbeck) budded to either Cleopatra mandarin (C. reticulata) and Troyer citrange (C. sinensis × P. trifoliata) rootstocks were used in this experiment. Cleopatra manda in rootstock was considered more tolerant to salinity than Troyer citrange, and this property was attributed to a greater capacity to exclude chloride ions.Plants were grown under glasshouse conditions and supplied with nutrient solution containing either no or 45 mM NaCl. Calcium concentration was increased from 3 to 30 mM. Sodium, potassium, calcium and chloride concentrations in plant organs were analyzed after 90 days of treatment.Supplemental Ca was found to mitigate the adverse effects of salinity on plant growth, defoliation or leaf injury.Chemical analysis indicated that in plants grafted on Troyer citrange Ca restricted uptake and subsequent translocation of Na to the leaves and increased K concentration in both roots and leaves. However, in Cleopatra mandarin-grafted plants increasing Ca levels seemed to reduce transport of Na from roots to leaves, and Na accumulation in roots was associated with reduced concentration of K in this rootstock.Organ chloride analysis showed that Cl accumulation in leaves of plants grafted on both rootstocks was reduced when external Ca concentration increased, whereas Cl concentration in roots remained constant or increased. The data of distribution of Cl in plants showed that a high external Ca level increased Cl accumulation in the basal stem and roots, and reduced the transport of Cl from roots to leaves.     

Salt Tolerance in the Triticeae: The Contribution of the D Genome to Cation Selectivity in Hexaploid Wheat

Inorganic cation concentrations were measured in shoots of hexaploidbread wheat (Triticum aestivum L.) and its presumed ancestorsgrown at 100 mol m^–3 external NaCl. Aegilops squarrosaand T. aestivum had high K/Na ratios while T. dicoccoides andAe. speltoides had low K/Na ratios. T. monococcum although havinga high K/Na ratio, had the highest total salt load of the fivespecies tested. The effect of the D genome (from Ae. squarrosa)was further investigated in seedlings of synthetic hexaploidwheats, and was again found to improve cation selectivity. Differentresponses were obtained from root and shoot tissue in this experiment.One synthetic hexaploid and its constituent parents were grownto maturity at 100 mol m^-3 NaCl and the yields recorded. Despitecomplications due to increased tillering in the stressed hexaploid,it was possible to show that the addition of the D genome enhancedyield characteristics in the hexaploid wheat. An experimentwith synthetic hexaploids derived from the tetraploid wheatvariety "Langdon" and several Ae. squarrosa accessions revealeddifferences in vegetative growth rates between the differentsynthetic hexaploids in the presence or absence of 150 or 200mol m^–3 external NaCl. The possibility of transferringsalt tolerance genes from Ae. squarrosa to hexaploid wheat usingsynthetic hexaploids as bridging species is discussed. Key words: Salt stress, wheat, D genome, Aegiops squarrosa, synthetic hexaploids     

Influence of aqueous leaf extract of common lambsquarters and NaCl salinity on the germination, growth, and nutrient content of wheat

Aqueous leaf extract of common lambsquarters (Chenopodium album L.) was evaluated alone or in combination with NaCl salinity for its influence on germination, seedling growth and contents of Na, K, and Ca by shoots and roots of wheat. The leaf extract and NaCl alone or in combination did not have any significant effect on germination, but the shoot and root lengths of seedlings and their dry weight decreased significantly by the treatments. Root growth was affected more than the shoot. The combination of leaf extract and NaCl drastically reduced the growth more than the separate effects of these stress treatments. The incorporation of leaf extract in the growth media decreased the content of Na by shoot, whereas the contents of K and Ca increased. NaCl treatments in combination with leaf extract increased the content of Na. Similar increases were observed for K and Ca in shoot, while these nutrients were increased in roots compared to control.     

Interaction of NaCl and Cd stress on compartmentation pattern of cations,antioxidant enzymes and proteins in leaves of two wheat genotypes differing in salt tolerance

Physiological mechanisms of salinity–Cd interactions were investigated in inter- and intracellular leaf compartments of salt-tolerant wheat × Lophopyrum elongatum (Host) A. Löve (syn. Agropyron elongatum) amphiploid and its salt-sensitive wheat parent (Triticum aestivum L. cv Chinese Spring). In comparison with the intracellular fluid, only very low Na⁺ concentrations (up to about 4 mM) were found in the intercellular leaf compartment of wheat after a 75 mM supply of NaCl. NaCl salinity led to a higher Cd concentration in leaves of the salt-sensitive genotype. Cd in the intercellular leaf compartment was not detectable. Higher K⁺ concentrations in the intercellular leaf compartment of the salt-sensitive genotype suggest a higher plasma membrane permeability caused by NaCl + Cd stress. Ascorbate peroxidase (APX) activity was increased in leaves of the salt-sensitive genotype under the combined NaCl and Cd stress. The highest non-specific peroxidase activities were detected under the combined stresses. It is suggested that NaCl and Cd stress in combination enhance the production of oxygen radicals and H₂O₂, especially in leaves of the salt-sensitive genotype. As a consequence, disturbed membrane function may cause elevated Cd concentrations in the intracellular leaf compartment under salinity. Cd did not change protein concentration and pattern in leaves. The protein content in inter-and intracellular leaf compartments of both genotypes was increased under salinity. A different protein pattern was obtained in inter- and intracellular leaf compartments. Thus, several physiological interactions between NaCl stress and Cd were found in the two wheat genotypes.     

Differential Solute Regulation in Leaf Blades of Various Ages in Salt-Sensitive Wheat and a Salt-Tolerant Wheat x Lophopyrum elongatum (Host) A. Love Amphiploid

Leaf blades of different ages from a salt-tolerant wheat x Lophopyrum elongatum (Host) A. Love (syn. Agropyron elongatum Host) amphiploid and its salt-sensitive wheat parent (Triticum aestivum L.cv Chinese Spring) were compared for their ionic relations, organic solute accumulation, and sap osmotic potential ([pi]sap). The plants were grown for 18 d in nonsaline (1.25 mM Na+) and salinized (200 mM NaCl) nutrient solutions. The response of leaf blades to NaCl salinity depended greatly on their age or position on the main stem. Na and proline levels were highest in the oldest leaf blade and progressively lower in younger ones. Glycine betaine and asparagine levels were highest in the youngest blade. The [pi]sap was similar for corresponding leaf blades of both genotypes, but contributions of various solutes to the difference in [pi]sap between blades from control and 200 mM NaCl treatments differed greatly. The NaCl-induced decline in [pi]sap of the youngest leaf blade of Chinese Spring was predominately due to the accumulation of Na and to a lesser extent asparagine; in the amphiploid, it was due to a combination of glycine betaine, K, Na, and asparagine. Proline contributed little in the youngest blade of either genotype. In the older blades Na was the major solute contributing to the decline in [pi]sap. Thus, the maintenance of low Na and high K levels and the accumulation of glycine betaine in the young leaf tissues contributed to the NaCl tolerance of the amphiploid. No such role was evident for proline.     

Interactive effects of boron and salinity stress on the growth,membrane permeability and mineral composition of tomato and cucumber plants

A greenhouse study was conducted in order to determine interactive effects of NaCl salinity and B on the growth, sodium (Na), chloride (Cl), boron (B), potassium (K) concentrations and membrane permeability of salt resistant Tomato (Lycopersicon esculentum L. cv. Lale F₁) and salt sensitive cucumber (Cucumis sativus L. cv. Santana F₁) plants. Plants were grown in a factorial combination of NaCl (0 and 30 mM for cucumber and 0 and 40 mM for tomato) and B (0, 5, 10 and 20 mg kg^–1 soil). Boron toxicity symptoms appeared at 5 mg kg^–1 B treatments in both plants. Salinity caused an increase in leaf injury due to B toxicity, but it was more severe in cucumber. Dry weights of the plants decreased with the increasing levels of applied B in nonsaline conditions, but the decrease in dry weights due to B toxicity was more pronounced in saline conditions especially in cucumber. Salinity × B interaction on the concentration of B in both plants was found significant. However, increase in B concentrations of tomato decreased under saline conditions when compared to nonsaline conditions. Contrary to this, B concentration of cucumber increased as a result of increasing levels of applied B and salinity. Salinity increased Na and Cl concentrations of both plants.Potassium concentration of tomato was not affected by salinity and B treatments, but K concentration of cucumber was decreased by salinity. Membrane permeability of the plants was increased by salinity while toxic levels of B had no effect on membrane permeability in nonsaline conditions. Membrane permeability was significantly increased in the presence of salinity by the increasing levels of applied B.     

The effect of increasing NaCl levels on the potassium utilization efficiency of tomatoes grown under low-K stress

Tomato strains were grown under low-K stress (71 μM K) over a wide range of external Na levels (from 0.014 to 27.8 mM Na) to measure strain response in Na substitution capacity in relation to Na concentration. Relative differences among strains for Na substitution capacity were similar at all Na levels except for the minus Na control treatment. Successive doubling of external Na concentration over the range of Na levels tested resulted in a positive linear response in plant dry weight, under low-K stress, with a similar slope for all five strains. The five strains also were grown at a toxic Na level (87 mM Na) under low K and adequate K conditions. Plant dry weight was not reduced at the toxic Na level relative to the minus Na control when the strains were grown under low-K stress; however, plant dry weight was reduced an average of fifty-five percent at the toxic Na level relative to the control when the strains were grown under adequate K conditions. There was no relationship between Na substitution capacity of strains grown under low-K stress and tolerance to toxic Na levels under adequate K conditions.