An inland and a coastal population of the Mediterranean xero-halophyte species Atriplex halimus L. differ in their ability to accumulate proline and glycinebetaine in response to salinity and water stress

Soil salinity and drought compromise water uptake and lead toosmotic adjustment in xero-halophyte plant species. These importantenvironmental constraints may also have specific effects onplant physiology. Stress-induced accumulation of osmocompatiblesolutes was analysed in two Tunisian populations of the Mediteraneanshrub Atriplex halimus L.—plants originating from a salt-affectedcoastal site (Monastir) or from a non-saline semi-arid area(Sbikha)—were exposed to nutrient solution containingeither low (40 mM) or high (160 mM) doses of NaCl or 15% polyethyleneglycol. The low NaCl dose stimulated plant growth in both populations.Plants from Monastir were more resistant to high salinity andexhibited a greater ability to produce glycinebetaine in responseto salt stress. Conversely, plants from Sbikha were more resistantto water stress and displayed a higher rate of proline accumulation.Proline accumulated as early as 24 h after stress impositionand such accumulation was reversible. By contrast, glycinebetaineconcentration culminated after 10 d of stress and did not decreaseafter the stress relief. The highest salt resistance of Monastirplants was not due to a lower rate of Na⁺ absorption; plantsfrom this population exhibited a higher stomatal conductanceand a prodigal water-use strategy leading to lower water-useefficiency than plants from Sbikha. Exogenous application ofproline (1 mM) improved the level of drought resistance in Monastirplants through a decrease in oxidative stress quantified bythe malondialdehyde concentration, while the exogenous applicationof glycinebetaine improved the salinity resistance of Sbikhaplants through a positive effect on photosystem II efficiency. Key words: Atriplex halimus, glycinebetaine, halophyte, NaCl, osmotic adjustment, proline, salinity, water stress     

Possible mechanisms of increasing salt tolerance in lentil plants after pre-exposure to low salt concentration

The main objective of the present study is to test the effect of short preexposure of lentil (Lens culinaris Medik.) plants to low salt concentration on shoot growth, oxidative stress and activity of antioxidant enzymes under high salt stress. To fulfill this objective, lentil plants were pretreated with 10 mM NaCl for 3 days and then they were exposed to high salt concentration of 300 mM for 7 days. After that, shoot growth was evaluated in terms of shoot length, fresh and dry weight. Biochemical changes in terms of oxidative stress and activity of antioxidant enzymes were also assessed in lentil plants. The shoot growth of lentil plants preexposed to low salt concentration was significantly enhanced under high salt stress, whereas it was severely retarded in lentil plants that were directly exposed to high salt stress. Moreover, lipid peroxidation and the accumulation of hydrogen peroxide were highly reduced in the shoot of lentil plants preexposed to low salt concentration. The activity of antioxidant enzymes (catalase and superoxide dismutase) was significantly higher in the shoot of lentil plants preexposed to low salt concentration than those directly exposed to high salt concentration. Overall, the results revealed an enhanced salt tolerance in lentil plants after short exposure to low salt concentration.     

Introduction of a Na+/H+ antiporter gene from Atriplex gmelini confers salt tolerance to rice

We engineered a salt-sensitive rice cultivar (Oryza sativa cv. Kinuhikari) to express a vacuolar-type Na+/H+ antiporter gene from a halophytic plant, Atriplex gmelini (AgNHX1). The activity of the vacuolar-type Na+/H+ antiporter in the transgenic rice plants was eight-fold higher than that in wild-type rice plants. Salt tolerance assays followed by non-stress treatments showed that the transgenic plants overexpressing AgNHX1 could survive under conditions of 300 mM NaCl for 3 days while the wild-type rice plants could not. These results indicate that overexpression of the Na+/H+ antiporter gene in rice plants significantly improves their salt tolerance.     

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.     

Photosynthetic activity and leaf antioxidative responses of Atriplex portulacoides subjected to extreme salinity

Responses of Atriplex portulacoides upon 40-day-long exposure to salinity (0?C1,000?mM NaCl) were investigated. Mother plants originated from a sabkha located in a semi-arid region of Tunisia. The plant relative growth rate and leaf expansion increased significantly at 200?mM NaCl but decreased at higher salinities. Interestingly, the plants survived salinity as high as 1,000?mM NaCl without displaying salt-induced toxicity symptoms. Despite significant increase in leaf Na⁺ and Cl^? concentrations upon salt treatment, no significant effect on leaf relative water content was registered. Chlorophyll contents and the gas exchange parameters showed a significant stimulation at the optimal salinity (200?mM NaCl) followed by a decline at higher salinities. Extreme salinity hardly impacted the maximal efficiency of photosystem II photochemistry (F _v/F _m), but a marked decrease in the relative quantum yield of photosystem II (??_PSII) was observed, along with a significant increase in non-photochemical quenching (NPQ). Leaf malondialdehyde and carotenoid contents were generally unaffected following salt exposure, whereas those of anthocyanins, polyphenols, and proline increased significantly, being maximal at 1,000?mM NaCl. Leaf superoxide dismutase (EC 1.15.1.1), ascorbate peroxidase (EC 1.11.1.11), and glutathione reductase (EC 1.6.4.2) activities were significantly stimulated by salinity, whereas catalase (EC 1.11.1.6) activity was maximal in the 0?C400?mM NaCl range. As a whole, protecting the photosynthetic machinery from salt-induced photodamage together with the sustained antioxidant activity may account for the performance of A. portulacoides under high salinity.     

Plant growth promoting rhizobacteria alleviate salinity induced negative effects on growth, oil content and physiological status in Mentha arvensis

This study was aimed to investigate the effect of inoculation on three salt-tolerant, plant-growth-promoting rhizobacteria (PGPR) STR2 (Bacillus pumilus), STR8 (Halomonas desiderata) and STR36 (Exiguobacterium oxidotolerans), for their growth-promoting potential and efficacy in augmenting salt tolerance in Mentha arvensis, an essential oil-bearing crop and natural source of l-menthol, under varying levels of NaCl stress (0, 100, 300 and 500 mM) imposed through irrigating water. Increase in the levels of salt concentration led to a decrease in the growth, fresh weight, leaf–stem ratio, oil content and yield. However, the negative effects of salinity were observed to be convalesced in the PGPR inoculated plants. At salinity levels of 100 and 300 mM NaCl, H. desiderata inoculated plants recorded the highest herb yield, whereas at 500 mM NaCl, the plants inoculated with E. oxidotolerans yielded maximum herb. The oil content in non-inoculated, salt-stressed plants was observed to be 0.46, 0.42 and 0.35 % at 100, 300 and 500 mM NaCl, respectively, whereas the plants inoculated with H. desiderata recorded the oil content of 0.71 and 0.60 and 0.48 % at similar levels of NaCl stress, respectively. The halotolerant PGPR minimized the deleterious effects of salt toxicity producing at par or higher yields at lower and medium salinity levels (100, 300 mM NaCl) than the un-inoculated non-salt-stressed plants through improved foliar nutrient uptake and enhanced antioxidant machinery. Based on the results of the experiments reported herein, the use of salt-tolerant, plant-growth-promoting bacteria may provide an effective means of facilitating M. arvensis growth in salt-stressed environments.     

Role of Salicylic Acid in Promoting Salt Stress Tolerance and Enhanced Artemisinin Production in Artemisia annua L.

In the present investigation, the role of salicylic acid (SA) in inducing salinity tolerance was studied in Artemisia annua L., which is a major source of the antimalarial drug artemisinin. SA, when applied at 1.00 mM, provided considerable protection against salt stress imposed by adding 50, 100, or 200 mM NaCl to soil. Salt stress negatively affected plant growth as assessed by length and dry weight of shoots and roots. Salinity also reduced the values of photosynthetic attributes and total chlorophyll content and inhibited the activities of nitrate reductase and carbonic anhydrase. Furthermore, salt stress significantly increased electrolyte leakage and proline content. Salt stress also induced oxidative stress as indicated by the elevated levels of lipid peroxidation compared to the control. A foliar spray of SA at 1.00 mM promoted the growth of plants, independent of salinity level. The activity of antioxidant enzymes, namely, catalase, peroxidase, and superoxide dismutase, was upregulated by salt stress and was further enhanced by SA treatment. Artemisinin content increased at 50 and 100 mM NaCl but decreased at 200 mM NaCl. The application of SA further enhanced artemisinin content when applied with 50 and 100 mM NaCl by 18.3 and 52.4%, respectively. These results indicate that moderate saline conditions can be exploited to obtain higher artemisinin content in A. annua plants, whereas the application of SA can be used to protect plant growth and induce its antioxidant defense system under salt stress.     

Effect of salt stress on growth parameters, enzymatic antioxidant system, and lipid peroxidation in wild chicory (Cichorium intybus L.)

Efficient utilization of saline land for food cultivation can increase agricultural productivity and rural income. To obtain information on the salt tolerance/susceptibility of wild chicory (Cichorium intybus L.), the influence of salinity (0–260 mM NaCl) on chicory seed germination and that of two salinity levels of irrigation water (100 and 200 mM NaCl) on plant growth, antioxidative enzyme activity, and accumulation of proline and malondialdehyde (MDA) were investigated. The trials were performed outdoors, in pots placed under a protective glass covering, for two consecutive years. Seeds showed a high capacity to germinate in saline conditions. The use of 100 mM NaCl solution resulted in 81 % germination, whereas seed germinability decreased below 40 % using salt concentrations above 200 mM NaCl. Wild chicory showed tolerance to medium salinity (100 mM NaCl), whereas a drastic reduction in biomass was observed when 200 mM NaCl solution was used for irrigation. MDA, present in higher amounts in leaves than in roots, decreased in both tissues under increasing salinity. Proline content increased remarkably with the level of salt stress, more so in roots than in leaves. In salt stress conditions, the activity of antioxidant enzymes (APX, CAT, POD, SOD) was enhanced. The electrophoretic patterns of the studied enzymes showed that the salinity of irrigation water affected only the intensity of bands, but did not activate new isoforms. Our results suggest that wild chicory is able to grow in soil with moderate salinity by activating antioxidative responses both in roots and leaves.     

Over-expression of Osmotin Induces Proline Accumulation and Confers Tolerance to Osmotic Stress in Transgenic Tobacco

Osmotin has been implicated in conferring tolerance to drought and salt stress in plants. We have over-expressed the osmotin gene under the control of constitutive CaMV 35S promoter in transgenic tobacco, and studied involvement of the protein in imparting tolerance to salinity and drought stress. The transgenic plants exhibited retarded leaf senescence and improved germination on a medium containing 200mM NaCl. Further, the transgenics maintained higher leaf relative water content (RWC), leaf photosynthesis and free proline content than the wild type plants during water stress and after recovery from stress. When subjected to salt stress (200mM NaCl), the transgenic plants accumulated significantly more proline than the wild type plants. These results suggest the involvement of the osmotin-induced increase in proline in imparting tolerance to salinity and drought stress in transgenic plants over-expressing the osmotin gene.     

Sea fennel (<Emphasis Type="Italic">Crithmum maritimum</Emphasis> L.) under salinity conditions: a comparison of leaf and root antioxidant responses

The present study was carried out to compare the effect of NaCl on growth, cell membrane damage, and antioxidant defences in the halophyte Crithmum maritimum L. (sea fennel). Physiological and biochemical changes were investigated under control (0 mM NaCl) and saline conditions (100 and 300 mM NaCl). Biomass and growth of roots were more sensitive to NaCl than leaves. Roots were distinguished from leaves by increased electrolyte leakage and high malondialdehyde (MDA) concentration. Superoxide dismutase (SOD), catalase (CAT) and ascorbate peroxidase (APX) activities, ascorbic acid (AA) and glutathione (GSH) concentrations were lower in the roots than in the leaves of control plants. The different activity patterns of antioxidant enzymes in response to 100 and 300 mM NaCl indicated that leaves and roots reacted differently to salt stress. Leaf CAT, APX and glutathione reductase (GR) activities were lowest at 300 mM NaCl, but they were unaffected by 100 mM NaCl. Only SOD activity was reduced in the latter treatment. Root SOD activity was significantly decreased in response to 300 mM NaCl and root APX activity was significantly higher in plants treated with 100 and 300 mM compared to the controls. The other activities in roots were insensitive to salt. The concentration of AA decreased in leaves at 100 and 300 mM NaCl, and in roots at 300 mM NaCl, when compared to control plants. The concentrations of GSH in NaCl-treated leaves and roots were not significantly different from the controls. In both organs, AA and GSH were predominating in the total pool in ascorbic acid and glutathione, under control or saline conditions.     

Growth responses of Stylosanthes humilis (Fabaceae) populations to saline stress

The growth responses to salinity of seedlings of six populations of Stylosanthes humilis from three ecogeographic regions of Northeast Brazil, characterized by wet or semi-arid climate, were analyzed following 28 days in solution cultures at concentrations of 0, 40, 80 and 120 mM NaCl. Root and shoot mass, shoot length and characters of foliar damage (number of leaves with chlorosis and necrosis) of the populations were more affected by increasing NaCl than numbers of leaves and branches and root length. Although S. humilis has been considered a salt sensitive legume, significant differences in salt tolerance between populations were found. The estimated concentrations which reduced shoot dry mass by 50 and 25% varied between populations from 84 to 108 and from 49 to 83 mM NaCl, respectively. Salt tolerance in S. humilis during the initial growth stage was lower than the reported one for germination. With one exception, populations from semi-arid climate with saline soils showed higher salt tolerance than those from non saline soils. The results suggest that salt tolerance in these populations is mainly associated with the occurrence of salinity in the soil of their provenance.     

Physiological responses to salt stress in young umbu plants

Soil salinity affects plant growth and development due to harmful ion effects and water stress caused by reduced osmotic potential in the soil solution. In order to evaluate the effects of salt stress in young umbu plants, research was performed in green house conditions at the Laboratory of Plant Physiology at Federal Rural University of Pernambuco, Brazil. Growth, stomatal behaviour, water relations, and both inorganic and organic solutes were studied aiming for a better understanding of the responses of umbu plants to increasing salinity. Plants were grown in washed sand with Hoagland and Arnon nutrient solution with 0, 25, 50, 75, and 100 mM NaCl. Growth, leaf water potential, transpiration, and diffusive resistance were evaluated. Na⁺, K⁺, Cl⁻, soluble carbohydrates, and free amino acid contents were measured in several plant organs. Most variables were affected with salinity above 50 mM NaCl showing decreases in: number of leaves, plant height, stems diameter, and dry masses, and increases in root-to-shoot ratio. Reductions in ψ_pd were observed in plants grown under 75 and 100 mM NaCl. All salt levels above zero increased Na⁺ and Cl⁻ contents in leaves. However, K⁺ content was not affected. Na⁺ and Cl⁻ in stems and roots reached saturation in treatments above 50 mM NaCl. Organic solute accumulation in response to salt stress was not observed in umbu plants. These results suggest that umbu plants tolerate salt levels up to 50 mM NaCl without showing significant physio-morphological alterations.     

A DESD-box helicase functions in salinity stress tolerance by improving photosynthesis and antioxidant machinery in rice (Oryza sativa L. cv. PB1)

The exact mechanism of helicase-mediated salinity tolerance is not yet understood. We have isolated a DESD-box containing cDNA from Pisum sativum (Pea) and named it as PDH45. It is a unique member of DEAD-box helicase family; containing DESD instead of DEAD/H. PDH45 overexpression driven by constitutive cauliflower mosaic virus-35S promoter in rice transgenic [Oryza sativa L. cv. Pusa Basmati 1 (PB1)] plants confers salinity tolerance by improving the photosynthesis and antioxidant machinery. The Na⁺ ion concentration and oxidative stress parameters in leaves of the NaCl (0, 100 or 200 mM) treated PDH45 overexpressing T₁ transgenic lines were lower as compared to wild type (WT) rice plants under similar conditions. The 200 mM NaCl significantly reduced the leaf area, plant dry mass, net photosynthetic rate (P_N), stomatal conductance (gs), intercellular CO₂ (Ci), chlorophyll (Chl) content in WT plants as compared to the transgenics. The T₁ transgenics exhibited higher glutathione (GSH) and ascorbate (AsA) contents under salinity stress. The activities of antioxidant enzymes viz. superoxide dismutase (SOD), ascorbate peroxidase (APX), guaiacol peroxidase (GPX) and glutathione reductase (GR) were significantly higher in transgenics; suggesting the existence of an efficient antioxidant defence system to cope with salinity induced-oxidative damage. Yeast two-hybrid assay indicated that the PDH45 protein interacts with Cu/Zn SOD, adenosine-5′-phosphosulfate-kinase, cysteine proteinase and eIF(4G), thus confirming the involvement of ROS scavenging machinery in the transgenic plants to provide salt tolerance. Furthermore, the T₂ transgenics were also able to grow, flower, and set viable seeds under continuous salinity stress of 200 mM NaCl. This study provides insights into the mechanism of PDH45 mediated salinity stress tolerance by controlling the generation of stress induced reactive oxygen species (ROS) and also by protecting the photosynthetic machinery through a strengthened antioxidant system.     

Elucidation of physiological and biochemical mechanisms of an endemic halophyte Centaurea tuzgoluensis under salt stress

In this study, physiological and biochemical responses of Centaurea tuzgoluensis, a Turkish endemic halophyte, to salinity were studied. Therefore, the changes in shoot growth, leaf relative water content (RWC), ion concentrations, lipid peroxidation, hydroxyl (OH^.) radical scavenging activity, proline (Pro) content, and antioxidant system [superoxide dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX) and glutathione reductase (GR)] were investigated. The 60 days (d) old C. tuzgoluensis seedlings were subjected to 0, 150 and 300 mM NaCl for 7 d and 14 d. The relative shoot growth was generally did not change in the 150 mM NaCl, but reduced with 300 mM NaCl stress at 7 d and 14 d. RWC was higher in 150 mM NaCl-treated leaves than that of 300 mM NaCl. Salinity decreased K⁺/Na⁺ ratio, but increased Na⁺, Cl^?, Ca⁺² and Na⁺/Cl^? ratio in the leaves. On the other hand, it did not change or increase the K⁺ content at 150 and 300 mM NaCl, respectively. MDA content in the 150 and 300 mM NaCl-treated leaves remained close to control at 7 d. This was related to enhanced activities of SOD, CAT, APX and GR enzymes, and their isoenzymes especially Fe-SOD in the leaves. On the other hand, the higher sensitivity to 300 mM NaCl at 14 d was associated with inadequate increase in antioxidant enzymes and the decreased OH radical scavenging activity. All these results suggest that C. tuzgoluensis has different antioxidant metabolisms between short- (7 d) and long-term (14 d) salt treatments and salinity tolerance of C. tuzgoluensis might be closely related to increased capacity of antioxidative system to scavenge reactive oxygen species (ROS) and accumulation of osmoprotectant proline under salinity conditions.     

Salt tolerance is unrelated to carbohydrate metabolism in cowpea cultivars

Under salinity stress, plants commonly accumulate carbohydrates for osmotic adjustment to balance the excess accumulated ions and to protect biomolecules. We selected two cowpea cultivars with contrasting response to salinity, Pitiúba (salt-tolerant) and TVu (salt-sensitive), to investigate whether the salt tolerance could be associated with changes in carbohydrate accumulation and metabolism in leaves and roots during a long-term experiment. Two salt treatments (0 and 75 mM NaCl) were applied to 10-day-old plants grown in nutrient solution for 24 days. Despite some changes in carbohydrate accumulation and carbohydrate metabolism enzymes induced by salt stress, no consistent alterations in carbohydrates could be found in leaves or roots in this study. Therefore, we suggest that tolerance to salt stress is largely unrelated to carbohydrate accumulation in cowpea.     

Involvement of thioredoxin y2 in the preservation of leaf methionine sulfoxide reductase capacity and growth under high light

Methionine (Met) in proteins can be oxidized to two diastereoisomers of methionine sulfoxide, Met‐S‐O and Met‐R‐O, which are reduced back to Met by two types of methionine sulfoxide reductases (MSRs), A and B, respectively. MSRs are generally supplied with reducing power by thioredoxins. Plants are characterized by a large number of thioredoxin isoforms, but those providing electrons to MSRs in vivo are not known. Three MSR isoforms, MSRA4, MSRB1 and MSRB2, are present in Arabidopsis thaliana chloroplasts. Under conditions of high light and long photoperiod, plants knockdown for each plastidial MSR type or for both display reduced growth. In contrast, overexpression of plastidial MSRBs is not associated with beneficial effects in terms of growth under high light. To identify the physiological reductants for plastidial MSRs, we analyzed a series of mutants deficient for thioredoxins f, m, x or y. We show that mutant lines lacking both thioredoxins y1 and y2 or only thioredoxin y2 specifically display a significantly reduced leaf MSR capacity (–25%) and growth characteristics under high light, related to those of plants lacking plastidial MSRs. We propose that thioredoxin y2 plays a physiological function in protein repair mechanisms as an electron donor to plastidial MSRs in photosynthetic organs.     

Anatomical changes induced by increasing NaCl salinity in three fodder shrubs, <Emphasis Type="Italic">Nitraria retusa</Emphasis>, <Emphasis Type="Italic">Atriplex halimus</Emphasis> and <Emphasis Type="Italic">Medicago arborea</Emphasis>

Nitraria retusa and Atriplex halimus (xero-halophytes) plants were grown in the range 0–800 mM NaCl while Medicago arborea (glycophyte) in 0–300 mM NaCl. Plants were harvested after 120 days of salt-treatment. The present study was designed to study the effect of salinity on root, stem and leaf anatomy, water relationship, and plant growth in greenhouse conditions. Salinity induced anatomical changes in the roots, stems and leaves. The cuticle and epidermis of N. retusa and A. halimus stems were unaffected by salinity. However, root anatomical parameters (root cross section area, cortex thickness and stele to root area ratio), and stem anatomical parameters (stem cross section area and cortex area) were promoted at 100–200 mM NaCl. Indicating that low to moderate salinity had a stimulating effect on root and stem growth of these xero-halophytic species. At higher salinities, root and stem structures were altered significantly, and their percentages of reduction were higher in A. halimus than in N. retusa whereas, in M. arborea, they were strongly altered as salinity rose. NaCl (100–300 mM) reduced leaf water content by 21.2–56.2% and specific leaf area by 51–88.1%, while increased leaf anatomical parameters in M. arborea (e.g. increased thickness of upper and lower epidermis, palisade and spongy mesophyll, entire lamina, and increased palisade to spongy mesophyll ratio). Similar results were evidenced in A. halimus leaves with salinity exceeding 100 mM NaCl. Leaves of N. retusa were thinner in salt-stressed plants while epidermis thickness and water content was unaffected by salinity. The size of xylem vessel was unchanged under salinity in the leaf’s main vein of the three species while we have increased number in M. arborea leaf main vein in the range of 200–300 mM NaCl. A longer distance between leaf vascular bundle, a reduced size and increased number of xylem vessel especially in stem than in root vascular system was evidenced in M. arborea treated plants and only at (400–800 mM) in the xero-halophytic species. The effects of NaCl toxicity on leaf, stem and root ultrastructure are discussed in relation to the degree of salt resistance of these three species. Our results suggest that both N. retusa and A. halimus show high tolerance to salinity while M. arborea was considered as a salt tolerant species.     

Salt-spray stimulated growth in strand-line species

The response to salt spray and soil salinity of two sand dune strandline species ( Cakile maritima Scop. and Salsola kali L.) and two salt marsh strand-line species ( Atriplex hastata L. and A. littoralis L.) was compared in sand-compost cultures. The growth of the salt-marsh species remained unaffected, while the growth of the sand dune species Cakile maritima was strongly reduced by NaCl (150 and 300 m M ) absorbed via the root system. All four species were resistant to airborne salinity, and under conditions of low soil fertility, salt spray increased the dry matter production, especially of the sand dune species. Mineral analysis revealed foliar uptake of Na, K, Cl, Ca and Mg. Na and Cl ions absorbed from seawater droplets induced succulence. Both salt spray and soil salt increased the methylated quaternary ammonium compound content in the shoot tissue. Under non-saline conditions a considerable amount of these osmotic solutes was still present, while turgor pressure potential in these plants was rather low. The relation between salt, compatible osmotic solutes, turgor pressure potential and growth is discussed. Next to the major constituents of seawater, Na and Cl, especially magnesium and to a lesser extent, calcium, accumulated in the shoot tissue. Based on the positive growth response of the sand dune species to airborne salt, they should be termed 'aerohalophytes', whereas 'soil halophytes' should be used when referring to the Atriplex species, which are more specifically adapted to the increased salinity of salt marsh soils.     

Glutamine synthetase and glutamate dehydrogenase contribute differentially to proline accumulation in leaves of wheat (Triticum aestivum) seedlings exposed to different salinity

To investigate the roles of ammonium-assimilating enzymes in proline synthesis under salinity stress, the activities of glutamine synthetase (GS; EC 6.3.1.2) and NADH-dependent glutamate dehydrogenase (NADH-GDH; EC 1.4.1.2) were determined in leaves of wheat (Triticum aestivum) seedlings exposed to salt stress at 150 and 300 mM NaCl for 5d. At the lower salinity, only GS activity increased markedly. At 300 mM NaCl, however, NADH-GDH activity increased while GS activity decreased. A significant accumulation of proline was found only at high-salinity exposure while glutamate, a proline precursor, increased dramatically under both low and high salinity. These data suggests that GS-catalysis might be the main glutamate synthesis pathway under low salinity. At 300 mM NaCl, glutamate seems to be preferentially produced through the process catalyzed by NADH-GDH. The increase of ammonium in salinity-stressed wheat seedlings might have resulted from increased photorespiration, which is responsible for the higher NADH-GDH activity. The activity of Delta(1)-pyrroline-5-carboxylate reductase (P5CR; EC 1.5.1.2) was significantly enhanced at 300 mM NaCl but remained unchanged at 150 mM. Delta(1)-Pyrroline-5-carboxylate synthetase (P5CS) activity did not show a specific response, indicating that P5CR might be the limiting step in proline synthesis from glutamate at high salinity.     

The role of salinity in seed maturation of the euhalophyte Suaeda salsa

Controlled conditions were used to investigate how salinity maintains the salt tolerance of seeds and seedlings of the euhalophyte Suaeda salsa. Seeds were harvested from S. salsa plants that had been treated with 1 or 500 mM NaCl for 113 days in a glasshouse. The results showed that high salinity (500 mM NaCl) increased chlorophyll concentration and oxygen production in embryos of maturing seeds. At 500 mM NaCl, the phosphatidylglycerol and sulfoquinovosyldiacylglycerol levels and the digalactosyldiacylglycerol/monogalactosyldiacylglycerol ratio were higher in young seedlings derived from seeds whose source plants were cultured in 500 mM rather than in 1 mM NaCl. When seeds were incubated with 600 mM NaCl, the conductivity and malondialdehyde concentration in the embryos was greater if the source plants had been cultured in 1 mM rather than in 500 mM NaCl. The opposite pattern was evident for seedling survival and shoot weight. In conclusion, salinity during seed maturation may increase the salt tolerance of seeds and seedlings by increasing the oxygen production in the embryos of the maturing seeds and by changing the lipid composition of membranes in the seedlings.