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Halophytes are plants able to tolerate high salt concentrations but no clear definition was retained for them. In literature, there are more studies that showed saltenhanced tolerance to other abiotic stresses compared to investigations that found enhanced salt tolerance by other abiotic stresses in halophytes. The phenomenon by which a plant resistance to a stress induces resistance to another is referred to as cross-tolerance. In this work, we reviewed cross-tolerance in halophytes at the physiological, biochemical, and molecular levels. A special attention was accorded to the cross-tolerance between salinity and organic pollutants that could allow halophytes a higher potential of xenobiotic phytoremediation in comparison with glycophytes.
The aim of this investigation was to identify the growth limiting factors in Arabidopsis thaliana subjected to a mild salt stress. Two natural accessions (Col and N1438) were compared. In spite of their morphological and developmental similarity, they have been previously shown to differ in the response of their superoxide dismutase genes to salt stress (Physiol Plant 132:293–305, 2008). Thirty-day-old seedlings were grown for 15 days using a split-root configuration in which the root system was divided into two equal parts: the first was immersed in a complete nutrient solution with 50 mM NaCl added, while the second part was immersed in either complete or incomplete K-, Ca-, or N-free medium. Using this approach, we demonstrated that K+ and Ca2+ uptake was impaired in the roots subjected to NaCl. There was no indication of the salt-induced inhibition of N uptake. If K+ or Ca2+ were available from salt-free medium, plants were able to grow at normal rate and accumulate large amounts of Na+ in the shoots. These results indicate that the sensitivity of Arabidopsis growth to mild salinity was probably due to an inhibition of K+ or Ca2+ root transport by salt rather than due to salt accumulation in shoots. Furthermore, the salt sensitivity of ion transport in roots seemed to depend on the genotype, since K+ was limiting for Col growth, in contrast to N1438, the growth of which was limited by Ca2+.
Seeds of four lettuce (Lactuca sativa L.) varieties (Romaine, Augusta, Vista and Verte) differing in their salt sensitivity were sown at 0 (Control), 50, 100 and 150 mM NaCl. The final germination percentage decreased with the increasing salinity and was annulated at the highest salt concentration in Vista and Verte, the most sensitive varieties. However, in the less sensitive ones, Romaine and Augusta, it was slightly modified at 50 and 100 mM NaCl and then decreased by 50% compared with the control, at 150 mM. The effects of NaCl 100 mM on seedling growth, phytase activities, phytate and inorganic phosphorus contents were studied in Romaine and Vista showing different behaviours towards salinity. Radicle and hypocotyl length and fresh and dry weights were reduced by salt treatment in both varieties. In addition, radicle phytase activity exhibited an increase in Romaine (less sensitive) and a decrease in Vista (more sensitive). In hypocotyl, this activity showed no difference with the control in the two varieties. However, in cotyledons, and during early hours after germination, salinity decreased phytase activity in both varieties whereas in the later hours (72–96 h) this activity reached the value of the control in Romaine. The enhancement of phytase activity was concomitant with an increase in orthophosphate content and a decrease in phytate reserve. These results suggest that salt presence in the medium delays Pi remobilization from phytate stock, but stimulates assimilation of phosphorus more than its accumulation in the organs of the two lettuce varieties.
Mentha pulegium L. is a medicinal and aromatic plant belonging to the Labiatae family present in the humid to the arid bioclimatic regions of Tunisia. We studied the effect of different salt concentrations on plant growth, mineral composition and antioxidant responses. Physiological and biochemical parameters were assessed in the plant organs after 2 weeks of salt treatment with 25, 50, 75 and 100 mM NaCl. Results showed that, growth was reduced even by 25 mM, and salt effect was more pronounced in shoots (leaves and stems) than in roots. This growth decrease was accompanied by a restriction in tissue hydration and K⁺ uptake, as well as an increase in Na⁺ levels in all organs. Considering the response of antioxidant enzymes to salt, leaves and roots reacted differently to saline conditions. Leaf and root guaiacol peroxidase activity showed an increase by different concentration of NaCl, but superoxide dismutase activity in the same organs showed a slight modification in NaCl-treated leaves and roots. Moreover, polyphenol contents and antioxidant activity were analysed in M. pulegium leaves and roots under salt constraint. The analysis showed an increase of total polyphenol content (2.41–8.17 mg gallic acid equivalent g⁻¹ dry weight) in leaves. However, methanol extract of leaves at 100 mM NaCl displayed the highest DPPH scavenging ability with the lowest IC₅₀ value (0.27 µg ml⁻¹) in comparison with control which exhibited IC₅₀ equal to 0.79 µg ml⁻¹ .
In the present work, we investigated the effect of salt stress on the distribution of safflower (Carthamus tinctorius L.) antioxidant system in relation to leaf age. The study was carried out under growth chamber conditions using seedlings of three cultivars which were subjected to 0 and 50 mM NaCl for 3 weeks. Leaf growth, water content, lipid peroxidation, and phenolic compound (total polyphenols, total flavonoids, and proanthocyanidins) concentration were measured at two leaf stages (young and old leaves). Leaf growth was affected by salinity only in Kairouan cultivar that also showed a significant decrease in old leaf water content. By contrast, Gabes and Tazarka cultivars maintained their old leaf water content constant and showed a reduction in that of young leaves. This could be attributed to a higher aptitude of the latter two cultivars to use absorbed sodium and chloride for osmotic adjustment in old leaves, keeping potassium for specific functions. Salt-induced lipid peroxidation was observed only in old leaves, whereas the accumulation of the major phenolic compounds under saline conditions was higher in young leaves, except in Gabes cultivar where no significant difference was found between the two leaf stages. A significant variability was also found between the three cultivars. The better behavior of salt-challenged leaves of Gabes and Tazarka cultivars compared to that of Kairouan cultivar may be related to their higher water content and the accumulation of polyphenols, in particular flavonoids that were shown to be efficiently involved in the restriction of salt-induced oxidative damages.
In the present work, magnesium deficiency effects were studied in Sulla carnosa plants grown in nutrient solution containing 1.50, 0.05, 0.01, and 0.00 mM Mg²⁺. After 5 weeks of treatment, fully expanded leaves were harvested to study their morphological and ultrastructural changes, as well as their carbohydrate, pigment, and Mg²⁺ concentrations. In control plants, leaves were well developed with remarkable green color. Down to 0.05 mM Mg²⁺, no chlorosis symptom was recorded, but below this concentration, mature leaves showed an appearance of interveinal chlorosis that was much more pronounced at 0.00 mM Mg²⁺ with the development of necrotic spots. Optima of chlorophyll a, chlorophyll b, and carotenoid concentrations were observed at 0.05 and 1.50 mM Mg²⁺; leaf magnesium concentration was severely reduced at 0.05 mM Mg²⁺. A significant decrease in pigment concentrations was noticed at 0.01 mM Mg²⁺, but the lowest values were recorded at 0.00 mM Mg²⁺. Enzymatic assays showed an increase in the accumulation of soluble sugars and starch with decreasing Mg²⁺ concentration. These results were in accordance with those of ultrastructural studies that revealed a marked alteration of chloroplasts in leaves of deficient plants. These chloroplasts were round and bigger as a result of a massive accumulation of oversized starch grains with disrupted thylakoids. As a whole, 1.50, 0.05, and 0.01 mM Mg²⁺ were found optimal, suboptimal, and deficient concentrations, respectively, the latter showing no significant difference with absolute Mg²⁺ absence (0.00 mM Mg²⁺).
Seedlings of two Tunisian Carthamus tinctorius L. provenances (Kairouan and Tazarka) differing in salt sensitivity were hydroponically grown at 0 and 50 mM NaCl over 21 days. Leaves of Kairouan (saltsensitive) showed a 48% restriction in their growth at 50 mM NaCl although they accumulated less sodium than those of Tazarka (less salt-sensitive) that maintained an unchanged growth. Salt treatment induced oxidative stress in C. tinctorius and the effect was more pronounced in the leaves of the more salt sensitive provenance, Kairouan. Both provenances exhibited a stimulation of antioxidant enzyme activities with higher catalase (CAT) and superoxide dismutase (SOD) activities in Tazarka and higher peroxidase (POD) activity in Kairouan. But, it seems that antioxidant activities were more correlated with polyphenol content. Actually, leaves of Tazarka experienced higher polyphenol and antioxidant activity than Kairouan at 50 mM NaCl. Hence, moderate salinity (3 g NaCl L⁻¹) enhanced bioactive molecule yield in the less salt sensitive provenance, Tazarka. In addition, C. tinctorius was found rich in ascorbic acid, but the moderate salt stress enhanced its production only in the sensitive provenance.
In the present work, separate and combined effects of excessive potassium and magnesium deficiency on safflower (Carthamus tinctorius) were studied. Four treatments were considered: C (control treatment: complete medium containing 1.5 mM Mg), ?KCl (excessive potassium treatment: complete medium added with 60 mMKCl), -Mg (Mg-deficient treatment: containing 0.1 mM Mg), and DS (double stress treatment: Mg-deficient medium (0.1 mM Mg) added with 60 mM KCl. Excessive potassium effect on plant growth was more pronounced than that of Mg deficiency. The two stresses impaired differently plant organs; KCl application affected more roots than shoots, whereas Mg deficiency reduced only leaf biomass. Gas exchange and pigment concentrations and patterns were severely impaired by KCl and mainly by interactive effects of the two stresses. This led to obvious lipid peroxidation and electrolyte leakage. Mg deficiency did not induce lipid peroxidation and electrolyte leakage, but as applied with excessive potassium, it doubled the effect of the latter. Mineral analyses showed that major cation nutrition was disturbed by KCl and combined stresses and at a lower level by magnesium deficiency. Plants did not show an enhanced selectivity of Mg and Ca over K but they improved their use efficiencies.
The aim of the present work was to check whether carbohydrate metabolism and partitioning contribute to the higher salt tolerance of the facultative halophyte Hordeum marinum compared to the glycophyte Hordeum vulgare. Seedlings with the same size from the two species were hydroponically grown at 0 (control), 150, and 300 mM NaCl for 3 weeks. H. marinum maintained higher relative growth rate, which was concomitant with a higher aptitude to maintain better shoot tissue hydration and membrane integrity under saline conditions compared to H. vulgare. Gas exchanges were reduced in the two species under saline conditions, but an increase in their water use efficiency was recorded. H. marinum exhibited an increase in leaf soluble sugar concentrations under saline conditions together with an enhancement in the transglucosidase DPE2 (EC 2.4.1.25) activity at 300 mM NaCl. However, H. vulgare showed a high increase in starch phosphorylase (EC 2.4.1.1) activity under saline conditions together with a decrease in leaf glucose and starch concentrations at 300 mM NaCl. In roots, both species accumulated glucose and fructose at 150 mM NaCl, but H. marinum exhibited a marked decrease in soluble sugar concentrations and an increase in starch concentration at 300 mM NaCl. Our data constitute an initiation to the involvement of carbohydrate metabolism and partitioning in salt responses of barley species and further work is necessary to elucidate how their flexibility confers higher tolerance to H. marinum compared to H. vulgare.
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