Salinity effects on polyphenol content and antioxidant activities in leaves of the halophyte Cakile maritima.

Cakile maritima is a local oilseed halophyte exhibiting potential for secondary metabolite production. In the present study, plant growth, leaf polyphenol content and antioxidant activity were comparatively analyzed in two C. maritima Tunisian accessions (Jerba and Tabarka, respectively sampled from arid and humid bioclimatic stages) under salt constraint. Three-week-old plants were subjected to 0, 100, and 400 mM NaCl for 28 days under glasshouse conditions. A significant variability in salt response was found between both accessions: while Tabarka growth (shoot biomass, leaf expansion) was significantly restricted at 100 and 400 mM NaCl, compared to the control, Jerba growth increased at 100mM before declining at 400 mM NaCl. The better behaviour of Jerba salt-challenged plants, compared to those of Tabarka, may be related to their higher polyphenol content (1.56- and 1.3-fold the control, at 100 and 400 mM NaCl respectively) and antioxidant activity (smaller IC(50) values for both 1,1-diphenyl-2-picrylhydrazyl and superoxide scavenging), associated with lower leaf MDA accumulation (ca. -66% of the control at 100mM NaCl). Taken together, our findings suggest that halophytes may be interesting for production of antioxidant compounds, and that the accession-dependent capacity to induce antioxidative mechanisms in response to salt, may result in a corresponding variability for growth sustainability.     

ABA, GA3, and nitrate may control seed germination of Crithmum maritimum (Apiaceae) under saline conditions

Impaired germination is common among halophyte seeds exposed to salt stress, partly resulting from the salt-induced reduction of the growth regulator contents in seeds. Thus, the understanding of hormonal regulation during the germination process is a main key: (i) to overcome the mechanisms by which NaCl-salinity inhibit germination; and (ii) to improve the germination of these species when challenged with NaCl. In the present investigation, the effects of ABA, GA₃, NO⁻₃, and NH⁺₄ on the germination of the oilseed halophyte Crithmum maritimum (Apiaceae) were assessed under NaCl-salinity (up to 200 mM NaCl). Seeds were collected from Tabarka rocky coasts (N-W of Tunisia). The exogenous application of GA₃, nitrate (either as NaNO₃ or KNO₃), and NH₄Cl enhanced germination under NaCl salinity. The beneficial impact of KNO₃ on germination upon seed exposure to NaCl salinity was rather due to NO⁻₃ than to K⁺, since KCl failed to significantly stimulate germination. Under optimal conditions for germination (0 mM NaCl), ABA inhibited germination over time in a dose dependent manner, but KNO₃ completely restored the germination parameters. Under NaCl salinity, the application of fluridone (FLU) an inhibitor of ABA biosynthesis, stimulated substantially seed germination. Taken together, our results point out that NO⁻₃ and GA₃ mitigate the NaCl-induced reduction of seed germination, and that NO⁻₃ counteracts the inhibitory effect of ABA on germination of C. maritimum. To cite this article: A. Atia et al., C. R. Biologies 332 (2009).     

Interactive effects of salinity and phosphorus availability on growth, water relations, nutritional status and photosynthetic activity of barley (Hordeum vulgare L.)

The interactive effects of salinity and phosphorus availability on growth, water relations, nutritional status and photosynthetic activity were investigated in barley (Hordeum vulgare L. cv. Manel). Seedlings were grown hydroponically under low or sufficient phosphorus (P) supply (5 or 180 μmol KH(2) PO(4) plant(-1) week(-1) , respectively), with or without 100 mm NaCl. Phosphorus deficiency or salinity significantly decreased whole plant growth, leaf water content, leaf osmotic potential and gas exchange parameters, with a more marked impact of P stress. The effect of both stresses was not additive since the response of plants to combined salinity and P deficiency was similar to that of plants grown under P deficiency alone. In addition, salt-treated plants exposed to P deficiency showed higher salt tolerance compared to plants grown with sufficient P supply. This was related to plant ability to significantly increase root:shoot DW ratio, root length, K(+)/Na(+) ratio, leaf proline and soluble sugar concentrations and total non-enzymatic antioxidant capacity, together with restricting Na(+) accumulation in the upper leaves. As a whole, our results indicate that under concomitant exposure to both salt and P deficiency, the impact of the latter constraint is pre-dominant.     

Comparative salt tolerance analysis between Arabidopsis thaliana and Thellungiella halophila, with special emphasis on K(+)/Na(+) selectivity and proline accumulation

The eco-physiology of salt tolerance, with an emphasis on K⁺ nutrition and proline accumulation, was investigated in the halophyte Thellungiella halophila and in both wild type and eskimo-1 mutant of the glycophyte Arabidopsis thaliana, which differ in their proline accumulation capacity. Plants cultivated in inert sand were challenged for 3 weeks with up to 500 mM NaCl. Low salinity significantly decreased A. thaliana growth, whereas growth restriction was significant only at salt concentrations equal to or exceeding 300 mM NaCl in T. halophila. Na⁺ content generally increased with the amount of salt added in the culture medium in both species, but T. halophila showed an ability to control Na⁺ accumulation in shoots. The analysis of the relationship between water and Na⁺ contents suggested an apoplastic sodium accumulation in both species; this trait was more pronounced in A. thaliana than in T. halophila. The better NaCl tolerance in the latter was associated with a better K⁺ supply, resulting in higher K⁺/Na⁺ ratios. It was also noteworthy that, despite highly accumulating proline, the A. thaliana eskimo-1 mutant was the most salt-sensitive species. Taken together, our findings indicate that salt tolerance may be partly linked to the plants’ ability to control Na⁺ influx and to ensure appropriate K⁺ nutrition, but is not linked to proline accumulation.     

Application of municipal solid waste compost reduces the negative effects of saline water in Hordeum maritimum L

The efficiency of composted municipal solid wastes (MSW) to reduce the adverse effects of salinity was investigated in Hordeum maritimum under greenhouse conditions. Plants were cultivated in pots filled with soil added with 0 and 40tha(-1) of MSW compost, and irrigated twice a week with tap water at two salinities (0 and 4gl(-1) NaCl). Harvests were achieved at 70 (shoots) and 130 (shoots and roots) days after sowing. At each cutting, dry weight (DW), NPK nutrition, chlorophyll, leaf protein content, Rubisco (ribulose-bisphosphate carboxylase/oxygenase) capacity, and contents of potential toxic elements were determined. Results showed that compost supply increased significantly the biomass production of non salt-treated plants (+80%). This was associated with higher N and P uptake in both shoots (+61% and +80%, respectively) and roots (+48% and +25%, respectively), while lesser impact was observed for K+. In addition, chlorophyll and protein contents as well as Rubisco capacity were significantly improved by the organic amendment. MSW compost mitigated the deleterious effect of salt stress on the plant growth, partly due to improved chlorophyll and protein contents and Rubisco capacity (-15%, -27% and -14%, respectively, in combined treatment, against -45%, -84% and -25%, respectively, in salt-stressed plants without compost addition), which presumably favoured photosynthesis and alleviated salt affect on biomass production by 21%. In addition, plants grown on amended soil showed a general improvement in their heavy metals contents Cu2+, Pb2+, Cd2+, and Zn2+ (in combined treatment: 190%, 53%, 168% and 174% in shoots and 183%, 42%, 42% and 114% in roots, respectively) but remained lower than phytotoxic values. Taken together, these findings suggest that municipal waste compost may be safely applied to salt-affected soils without adverse effects on plant physiology.     

Relationship between the photosynthetic activity and the performance of Cakile maritima after long-term salt treatment

Cakile maritima is a halophyte with potential for ecological, economical and medicinal uses. We address here the impact of salinity on its growth, photosynthesis and seed quality. Whole plant growth rate and shoot development were stimulated at moderate salinity (100–200 m M NaCl) and inhibited at higher salt concentrations. Although diminished in the presence of salt, potassium and calcium uptake per unit of root biomass was maintained at relatively high value, while nutrient-use efficiency (NUE) was improved in salt-treated plants. Chl and carotenoid concentrations decreased at extreme salinities, but anthocyanin concentration continuously grew with salinity. Net photosynthetic rate (A), stomatal conductance, maximum quantum efficiency of PSII and quantum yield were stimulated in the 100–200 m M NaCl range. Higher salinity adversely affected gas exchange and changed PSII functional characteristics, resulting in a reduction of A per leaf area unit. This phenomenon was associated with increased non-photochemical quenching. Harvest index, silique number and seeds per fruit valve were maximal at 100 m M NaCl. Despite the decreasing salt accumulation gradient from the vegetative to the reproductive organs, high salinities were detrimental for the seed viability and increased the proportion of empty siliques. Overall, the salt-induced changes in the plant photosynthetic activity resulted into analogous responses at the vegetative and reproductive stages. The enhancement of NUE, the absence of pigment degradation, the reduction of water loss and the concomitant PSII protection from photodamage through thermal dissipation of excess excitation significantly accounted for Cakile survival capacity at high salinity.     

Comparative Ni tolerance and accumulation potentials between Mesembryanthemum crystallinum (halophyte) and Brassica juncea: Metal accumulation,nutrient status and photosynthetic activity

Saline soils often constitute sites of accumulation of industrial and urban wastes contaminated by heavy metals. Halophytes, i.e. native salt-tolerant species, could be more suitable for heavy metal phytoextraction from saline areas than glycophytes, most frequently used so far. In the framework of this approach, we assess here the Ni phytoextraction potential in the halophyte Mesembryanthemum crystallinum compared with the model species Brassica juncea. Plants were hydroponically maintained for 21 days at 0, 25, 50, and 100 μM NiCl₂. Nickel addition significantly restricted the growth activity of both species, and to a higher extent in M. crystallinum, which did not, however, show Ni-related toxicity symptoms on leaves. Interestingly, photosynthesis activity, chlorophyll content and photosystem II integrity assessed by chlorophyll fluorescence were less impacted in Ni-treated M. crystallinum as compared to B. juncea. The plant mineral nutrition was differently affected by NiCl₂ exposure depending on the element, the species investigated and even the organ. In both species, roots were the preferential sites of Ni²⁺ accumulation, but the fraction translocated to shoots was higher in B. juncea than in M. crystallinum. The relatively good tolerance of M. crystallinum to Ni suggests that this halophyte species could be used in the phytoextraction of moderately polluted saline soils.     

Physiological and biochemical traits involved in the genotypic variability to salt tolerance of Tunisian Cakile maritima

Cakile maritima (family: Brassicaceae) was collected from three provenances belonging to different bioclimatic stages (humid, semi arid and arid) in Tunisia to study their eco‐physiological and biochemical responses to salinity. Seedlings were cultivated on inert sand for 20 days under NaCl treatments (0, 100, 200, 400 mm NaCl). Plant response to salinity was provenance‐ and salt‐dependent. At 100 mm NaCl, growth parameters (leaf biomass, area, number per plant and relative growth rate) were improved in plants from Jerba (originating from arid bioclimatic stage) compared with the control, while growth was reduced in those from Tabarka (from humid area). High salt levels (400 mm NaCl) decreased the plant growth in the three provenances, but plants in Tabarka were the most salt sensitive. The relative salt tolerance of plants from Jerba and Bekalta provenances was associated with low levels of malondialdehyde as well as of electrolyte leakage and endoproteolytic activity. Salt reduced leaf hydration, the decrease in water content being dose‐dependent and more pronounced in Tabarka. Increase in salinity led to significant increase in leaf succulence and decrease in leaf water potential, especially in Jerba plants. The plants from the latter displayed the highest leaf levels of Na⁺ and Cl^?, proline, soluble carbohydrates, soluble proteins, and polyphenols. Overall, the higher salt tolerance of plants from Jerba provenance, and to a lower extent of those from Bekalta, may be partly related to their better capacity for osmotic adjustment and to limit oxidative damage when salt‐challenged.     

Comparison of two chickpea varieties regarding their responses to direct and induced Fe deficiency

The effects of Fe deficiency (whether direct or bicarbonate-induced) on plant morphology, growth parameters, photosynthesis-related pigment contents, gas exchange, and water relations were addressed in two contrasting chickpea varieties (INRAT88 and Chetoui, respectively tolerant and sensitive to Fe deficiency). A marked decrease in the whole plant Fe content was observed in the Fe deprived plants of both varieties, especially the bicarbonate-treated ones, which showed a slower growth development and water deficit stress symptoms (increased leaf tissue osmolality associated with decreased shoot height, increased leaf mass to area ratio, and decreased water content). Both Fe shortage and bicarbonate addition resulted in both varieties in the decline of the photosynthetic pigment contents, contributing to lower photosynthetic efficiency (φ_c) and lower net photosynthesis (A). Fe deficiency reduced the water use efficiency and physiological availability of water too. However, INRAT88 was more tolerant to Fe deficiency than Chetoui, by maintaining a higher growth rate associated with lower respiration rate (R_D), higher chlorophyll a and b concentrations, higher A, lower transpiration rate (E) and a higher water use efficiency (A/E). The present data suggest that the efficient utilisation of Fe for the synthesis of chlorophyll together with the effective control of electron-transport chains at chloroplasts (high A) and mitochondria (low R_D) may account for the higher tolerance of INRAT88 to direct Fe deficiency. Further investigations with respect to oxidative stress and ROS generation, or about photorespiration would be helpful for a better understanding of their interaction with Fe deficiency in this grain legume.     

Risk of municipal solid waste compost and sewage sludge use on photosynthetic performance in common crop (<Emphasis Type="Italic">Triticum durum</Emphasis>)

The effect of composted municipal solid waste (MSW) and sewage sludge (SS) on photosynthetic activity of wheat (Triticum durum L.) was investigated. Chlorophyll fluorescence and gas exchange parameters were assessed following application of up to 300 t ha^?1 of MSW compost or SS. 100 t ha^?1 MSW compost was optimal for the plant growth, which showed 78% stimulation as compared to the control. This was associated with higher maximum quantum efficiency (F _v /F _m) of photosystem II (PSII) and the actual quantum efficiency of PSII open centers at light adapted state (ΔF/\(F_{\rm m}^{\prime}\)). Maximal values of net photosynthetic rate and stomatal conductance were recorded at 100 t ha^?1 MSW compost (+40 and +116%, respectively). Ribulose bisphosphate carboxylase/oxygenase (RubisCO) activity was also significantly stimulated at 100 t ha^?1, while less significant impact was found in SS treatment. A marked accumulation of Ni, Pb, Cu, and Zn in concomitance with membrane lipid peroxidation were observed at 200–300 t ha^?1 MSW compost and SS, resulting in lower photosynthetic activity and altered PSII functional integrity. Altogether, these results suggest that the MSW compost at 100 t ha^?1 would be suitable for wheat cultivation, within the critical limits of heavy metal accumulation. However, long-term field experiments seem necessary to more accurately evaluate the safety of MSW application.