Stoichiometric variation of halophytes in response to changes in soil salinity

• Variation in soil salt may change the stoichiometry of a halophyte by altering plant ecophysiology, and exert different influences on various plant organs, which has potentially important consequences for the nutrition of consumers as well as nutrient cycling in a saline ecosystem.
• Using a greenhouse pot experiment, we investigated the effect of salinity variability on the growth and stoichiometry of different organs of Suaeda glauca and Salicornia europaea – two dominant species of important ecological and economic value in the saline ecosystem.
• Our results showed that appropriate salt stimulated the growth of both species during the vigorous growth period, while high salt suppressed growth. Na significantly increased with increased salt in the culture, whereas concentrations of other measured elements and K:Na ratio for both species significantly decreased at low salt treatments, and became more gradual under higher salt conditions. Furthermore, with the change of salt in culture, variations in leaf (degenerated leaf for S. europaea, considered as young stem) stoichiometry, except N:P ratio, were large and less in stems (old stems for S. europaea) than in roots, reflecting physiological and biochemical reactions in the leaf in response to salt stress, supported by sharp changes in trends.
• These results suggest that appropriate saline conditions can enhance biological C fixation of halophytes; however, increasing salt could affect consumer health and decrease cycling of other nutrients in saline ecosystems.

     

Quantification and fatty acid profiles of sulfolipids in two halophytes and a glycophyte grown under different salt concentrations

This study was aimed at understanding the role of sulfolipids in salt tolerance mechanisms of the halophytes Aster tripolium L., Compositae, and Sesuvium portulacastrum L., Aizoaceae, and of the glycophyte Arabidopsis thaliana (L.) Heynh., Brassicaceae. In Aster and Sesuvium the sulfolipid contents increased significantly under salt stress conditions (517 mM or 864 mM). In Arabidopsis, changes in sulfolipid contents were not observed (NaCl up to 100 mM). The fatty acid profile of sulfoquinovosyldiacylglycerol (SQDG) in Aster was modified with increasing NaCl concentrations. LC-MS analyses of sulfolipids from Aster and Sesuvium revealed the presence of 18:3/18:3 and 16:0/18:3 molecules. Obviously, the function of sulfolipids during salt stress differs between halophytic species and between halophytes and glycophytes where sulfolipid accumulation was not observed.     

Role of organic acids in the formation of the ionic composition in developing glycophyte leaves

The ionic composition in the leaves of some glycophyte plants (Phaseolus vulgaris L., Lycopersicon esculentum L., and Amaranthus cruentus L.) was studied during leaf development. Plants were grown in a stationary hydroponic culture; a growth medium contained equimolar concentrations of inorganic ions (NO ₃ ^? , Cl^?, SO ₄ ^2? , H₂PO ₄ ^? , K⁺, Ca²⁺, Mg²⁺, and Na⁺) equal to 5 mg-equiv./l for each ion. In the juvenile leaf, the main ions were K⁺ and water-soluble anions of organic acids represented mainly by di-and tricarboxylic acids in kidney bean and tomato and oxalic acid in amaranth. An increase in the total amount of organic anions, coinciding with the accumulation of bivalent cations, was registered in leaves of glycophytes during their development. Mature and senescing leaves of tomato and kidney bean accumulated mainly di-and tricarboxylic acid salts with the prevalence of Ca²⁺ ions. In amaranth leaves, the formation of water-insoluble (acid-soluble) oxalate pool containing Ca²⁺ ions (mature leaves) or Ca²⁺ and Mg²⁺ ions (senescing leaves) was registered. The priority role of the metabolism of organic acids in the formation of the ionic composition of glycophyte leaves during their development is discussed. It is supposed that the species-specific ionic composition of glycophyte leaves at different developmental stages is due mainly to the pattern of carbon metabolism causing the accumulation either of di-and tricarboxylic acids or oxalic acid.     

Growth,ionic response,and gene expression of shoots and roots of perennial ryegrass under salinity stress

Growth, ionic responses, and expression of candidate genes to salinity stress were examined in two perennial ryegrass accessions differing in salinity tolerance. The salinity tolerant (PI265349) and sensitive accessions (PI231595) were subjected to 75-mM NaCl for 14 days in a growth chamber. Across two accessions, salinity stress increased shoot dry weight and concentrations of malondialdehyde (MDA) and Na⁺ in the shoots and roots, but decreased shoot Ca²⁺ and root K⁺ concentrations. Salinity stress also increased root expressions of SOS1, PIP1, and TIP1. Plant height and chlorophyll content were unaffected by salinity stress in the tolerant accession but significantly decreased in the sensitive accession. Shoot MDA content did not change in the tolerant accession but increased in the sensitive accession. A more dramatic increase in Na⁺ was found in the roots of the sensitive accession. Relative to the control, salinity stress reduced expression of SOS1, NHX1, PIP1, and TIP1 in the shoots but increased expression of these genes in the roots of the tolerant accession. Expression levels of SOS1 increased in the roots and expression of NHX1 increased in the shoots but decreased in the roots of the sensitive accession under salinity stress. A decline in PIP1 expression in the shoots and dramatic increases in TIP expression in both shoots and roots were found in the sensitive accession under salinity stress. The results suggested maintenance of plant growth and leaf chlorophyll content, lesser Na⁺ accumulation in the roots, and lower lipid peroxidation in the shoots which could be associated with salinity tolerance. The decreased expressions of SOS1, NHX1, and TIP1 in the shoots, and increased expressions of NHX1 and PIP1 in the roots might also be related to salinity tolerance in perennial ryegrass.     

Morpho-physiological response to vertically heterogeneous soil salinity of two glycophyte woody taxa,Salix matsudana x S. alba and Eucalyptus camaldulensis Dehnh

Plant and Soil - Growth and physiology of Salix matsudana x S. alba and E. camaldulensis were evaluated in vertical saline gradients to test whether growth is determined by the mean salinity of...     

A comparative study of the hormonal response to high temperatures and stress reiteration in three Labiatae species

Mediterranean plants are usually exposed to a combination of stresses, which may occur simultaneously or at different times throughout their life. Here, the hormonal response to high temperatures was compared in plants of three Labiatae species, including rosemary (Rosmarinus officinalis L.), sage (Salvia officinalis L.) and lemon balm (Melissa officinalis L.). Plants exposed to increasing temperatures for 5 days were subsequently exposed to heat stress and compared to plants experiencing heat stress for the first time (controls). Despite the three species showing a similar tolerance to a single heat stress event, stress recurrence had a different impact on each species. Lemon balm was the most sensitive species to stress reiteration, showing decreased relative water content upon heat stress repetition, together with enhanced levels of α-tocopherol and salicylic acid (SA). Some acclimation responses were observed in rosemary and sage, including improved water contents and reduced jasmonic acid levels in rosemary, and reduced abscisic acid (ABA) and malondialdehyde (MDA) levels in sage plants previously exposed to high temperatures. Furthermore, the response of plants to a combination of heat stress and water deficit was evaluated in plants previously exposed to heat stress and compared to controls. Rosemary and sage were much more resistant than lemon balm, which died when stresses were combined. Despite stress pre-exposure not having any effect on plant performance in terms of F_v/F_m, MDA and relative water content in rosemary and sage, it resulted in higher α-tocopherol levels in both species. The hormonal response differed between species: while the hormonal content did not change in sage, rosemary showed increased ABA and decreased SA levels as a result of repeated stress exposure. Overall, different stress imprints particular to each species and stress scenario were found in α-tocopherol and hormone levels, which led to similar protective effects in rosemary and sage.     

Changes in PS II heterogeneity in response to osmotic and ionic stress in wheat leaves (Triticum aestivum)

High salt stress involves ionic stress as well as osmotic stress. In this work we have tried to differentiate between the ionic and osmotic effects of salt stress on the basis of their ability to cause changes in PS II heterogeneity. PS II heterogeneity is found to vary with environmental conditions. Osmotic stress caused no change in the Q(B) reducing side heterogeneity and a reversible change in antenna heterogeneity. The number of Q(B) non-reducing centers increased under ionic stress but were unaffected by osmotic stress. On the other hand ionic stress led to a partially irreversible change in Q(B) reducing side heterogeneity and a reversible change in antenna heterogeneity. In response to both ionic and osmotic effect, there is conversion of active PS IIα centres to inactive PSIIβ and γ centres.     

Solute adjustments in leaves of two species of wheat at two different stages of growth in response to salinity

Changes in leaf solute concentrations in response to salinity were measured at two growth stages in two species of wheat, Triticum turgidum L. cv. Aldura (Durum group) and Triticum aestivum L., cv. Probred that differed in their salt tolerances. Both species at 55 days of age were Na⁺-excluders, but the concentration of Na⁺ was 10 times higher in T. turgidum than T. aestivum at low to moderate levels of stress. The ratio then decreased until it was 2:1 at – 1.2 MPa. In T. turgidum, K⁺ concentrations decreased with increasing Na⁺ concentrations so that the sum of the two cations remained constant at all stress levels, but in T. aestivum K⁺ decreased more rapidly than Na⁺ increased. In both species growing in media at 0 to –0.6 MPa, the amounts of Mg²⁺ and Ca²⁺ in 55-day-old plants that could be extracted with hot water were below 0.1 mmol (g dry weight)^?1. Then, as osmotic potentials of media decreased further, hot water-extractable Ca²⁺ increased greatly until, at – 1.2 MPa, Ca²⁺ concentrations were almost equal to the sum of Na⁺ and K⁺. In the range of 0 to –1.0 MPa, the ratio of Cl^? to total cationic charge remained constant at 1:6 in T. aestivum and 1:2 in T. turgidum. However, at – 1.2 MPa, the ratio in both species had changed to 2:3. Sucrose and betaine concentrations were 4 and 48 μmol (g dry weight)^?1, respectively, in non-stressed plants of both species. At – 1.2 MPa, sucrose had increased 30-fold but betaine had increased only 2.5-fold. Proline increased exponentially relative to foliar Na⁺ in T. turgidum. In T. aestivum only plants grown at –1.2 MPa contained sufficient Na⁺ to stimulate the accumulation of proline. Although the quantities of the solutes in leaves of non-stressed 96-day-old plants differed from those in non-stressed younger plants, the patterns of change of organic solutes as the older plants were subjected to increasing saline stresses were the same as in younger plants with the exception of sucrose. Sucrose concentrations were much higher in leaves of non-stressed older plants and this sugar first increased and then decreased with decreasing osmotic potentials of media.     

A comparative study of the response of the roots and shoots of sunflower and maize to mechanical stimulation

Despite numerous studies of the effects of mechanical stimulationon plant shoots, the response of roots to mechanical stimulationhas largely been neglected. In this study the effects of shootflexure on the morphology and mechanics of two contrasting speciesof herbaceous angiosperm, growing in a glasshouse were compared:maize (Zea mays), a monocot; and sunflower (Helianthus annuusL.) a dicot. Mechanical stimulation affected the root more than the shootcomponents. Root systems of mechanicallystressed sunflowershad a greater angle of spread and increased root number. Aswell as large morphological and weight effects, with increasesover the control of 33% in the length of rigid root and 38%in the dry weight of lateral roots, in sunflowers, there werealso mechanical effects. In both species roots of flexed plantswere more rigid, stronger and composed of stiffer material andtheir root systems also provided greater anchorage strength.In contrast, there was only a small reduction in shoot weightand shoot height in flexed plants and no effects on mechanicalproperties. There were differences in behaviour between species; maize rootmorphology responded less than that of sunflowers to mechanicalstimulation. The basal diameter of roots increased by only 8%compared with 16% in sunflowers, though the roots of both speciesshowed similar increases in material stiffness. This differenceis related to the lack of secondary thickening in the monocotscompared with the dicot sunflowers. Key words: Thigmomorphogenesis, Helianthus annuus L., Zea mays, anchorage, lodging     

Proteome-level changes in the roots of Pisum sativum in response to salinity

We initiated a proteomics-based approach to identify root proteins affected by salinity in pea (Pisum sativum cv. Cutlass). Salinity stress was imposed either on 2-wk old pea plants by watering with salt water over 6 wk or by germinating and growing pea seeds for 7 days in Petri dishes. Concentrations of NaCl above 75 mM had significant negative effects on growth and development of peas in both systems. Salinity-induced root proteome-level changes in pea were investigated by 2-D electrophoresis of proteins from control, 75 and 150 mM NaCl-treated plants and seedlings. The majority of the protein spots visualised showed reproducible abundance in root protein extracts from whole plants and seedlings. Of these proteins, 35 spots that exhibited significant changes in abundance due to NaCl treatment were selected for identification using ESI-Q-TOF MS/MS. The identities of these proteins, which include pathogenesis-related (PR) 10 proteins, antioxidant enzymes such as superoxide dismutase (SOD) as well as nucleoside diphosphate kinase (NDPK) are presented, and the roles of some of them in mediating responses of pea to salinity are discussed. This is the first report of salinity-induced changes in the root proteome of pea that suggests a potential role for PR10 proteins in salinity stress responses. Our findings also suggest the possible existence of a novel signal transduction pathway involving SOD, H₂O₂, NDPK and PR10 proteins with a potentially crucial role in abiotic stress responses.     

A comparative study of two RNase isozyme activities inNicotiana glutinosa leaves

Effects of some chemicals on the activities of two ribonuclease isozymes, M and F, in the leaves ofNicotiana glutinosa were investigated. CHX and AMD suppressed the increase of M isozyme activity induced with mock-inoculation. On the other hand, CHX suppressed slightly the increase of F izozyme activity induced with TMV-inoculation, but AMD and 5-FU stimulated it.     

Disentangling the contributions of osmotic and ionic effects of salinity on stomatal,mesophyll, biochemical and light limitations to photosynthesis

There are conflicting opinions on the relative importance of photosynthetic limitations under salinity. Quantitative limitation analysis of photosynthesis provides insight into the contributions of different photosynthetic limitations, but it has only been applied under saturating light conditions. Using experimental data and modelling approaches, we examined the influence of light intensity on photosynthetic limitations and quantified the osmotic and ionic effects of salinity on stomatal (L_S), mesophyll (L_M), biochemical (L_B) and light (L_L) limitations in cucumber (Cucumis sativus L.) under different light intensities. Non‐linear dependencies of L_S, L_M and L_L to light intensity were found. Osmotic effects on L_S and L_M increased with the salt concentration in the nutrient solution (S_s) and the magnitude of L_M depended on light intensity. L_S increased with the Na⁺ concentration in the leaf water (S_l) and its magnitude depended on S_s. Biochemical capacity declined linearly with S_l but, surprisingly, the relationship between L_B and S_l was influenced by S_s. Our results suggest that (1) improvement of stomatal regulation under ionic stress would be the most effective way to alleviate salinity stress in cucumber and (2) osmotic stress may alleviate the ionic effects on L_B but aggravate the ionic effects on L_S.     

Cellular redox regulation,signaling, and stress response in plants

Cellular and organellar redox states, which are characterized by the balance between oxidant and antioxidant pool sizes, play signaling roles in the regulation of gene expression and protein function in a wide variety of plant physiological processes including stress acclimation. Reactive oxygen species (ROS) and ascorbic acid (AsA) are the most abundant oxidants and antioxidants, respectively, in plant cells; therefore, the metabolism of these redox compounds must be strictly and spatiotemporally controlled. In this review, we provided an overview of our previous studies as well as recent advances in (1) the molecular mechanisms and regulation of AsA biosynthesis, (2) the molecular and genetic properties of ascorbate peroxidases, and (3) stress acclimation via ROS-derived oxidative/redox signaling pathways, and discussed future perspectives in this field.     

Genetically modified cyanobacteriumNostoc muscorum overproducing proline in response to salinity and osmotic stresses

In the parentNostoc muscorum an active proline oxidase enzyme is required to assimilate exogenous proline as a fixed nitrogen source. Cyanobacterial mutants, resistant to growth inhibitory action of proline analogue L-azetidine-2-carboxylate (Ac-R), were deficient in proline oxidase activity, and were over-accumulators of proline. Proline over-accumulation, resulting either from mutational acquisition of the Ac-R phenotype, or from salinity-induced uptake of exogenous proline, confirmed enhanced salinity/osmotic tolerance in the mutant strain. The nitrogenase activity and photosynthetic O₂ evolution of the parent were sensitive to both salinity as well as osmotic stresses than of Ac-R mutant strain. In addition, the mutation to Ac-resistant phenotype showed no alteration in salinity inducible potassium transport system in the cyanobacterium.     

Antioxidative response to cadmium in roots and leaves of tomato plants

Treatment of tomato seedlings (Lycopersicon esculentum Mill. cv. 63/5 F1) with increasing CdCl₂ concentrations in the culture medium resulted in Cd accumulation more important in roots than in leaves. Biomass production was severely inhibited, even at low Cd concentration. Cd reduced chlorophyll content in leaves and enhanced lipid peroxidation. An increase in antioxidative enzyme (superoxide dismutase, ascorbate peroxidase, guaiacol peroxidase, glutathione reductase) activities was more pronounced in leaves than in roots, while catalase activity increased only in roots. In addition, changes in isoenzyme composition were observed using the non-denaturing polyacrylamid gel electrophoresis.     

Response of photosynthetic performance,water relations and osmotic adjustment to salinity acclimation in two wheat cultivars

Experiment was conducted to identify the impacts of the salinity acclimation process on the photosynthetic efficiency, osmotic adjustment, membrane integrity, and yield components in two wheat cultivars differing in their salinity tolerance. The design of the experiment was factorial randomized block, where genotype is factor 1 and acclimation treatments represent factor 2. Genotypes were grown from emergence to 30 days after sowing (DAS) by irrigating with tap water [electrical conductivity (EC) of 0.776 dS m^?1]. Thereafter, both the genotypes were divided into two groups and exposed to either irrigation with sublethal level of salinity EC of 2.09 or 3.76 dS m^?1 for 21 days. At booting stage (65 DAS), both groups were subjected to lethal level of salinity stress EC of 12 dS m^?1 for 21 days, followed by irrigation with tap water till maturity. Non-acclimated plants were irrigated with tap water from emergence to 65 days, then directly irrigated with lethal level of salinity for 21 days, followed by irrigation with tap water till maturity. The control plants were continuously irrigated with tap water from emergence until maturity. The non-acclimated plants had decreased electron transport rates at the donor and acceptor side of PSII and PSI in Giza 168, and decreased electron transport rates at PSII acceptor side in Sakha 8 compared to control plants. In both genotypes, the non-acclimated plants had decreased chlorophyll a, b, carotenoid, proline and total soluble sugar concentration, relative water content, membrane stability index, yield and yield components compared with acclimated plants. While, osmotic potential and lipid peroxidation showed an opposite trend. Overall, acclimation treatment (EC of 2.09 dS m^?1) during vegetative stage alleviated the inhibitory effects of lethal level of salinity stress at booting stage through enhanced photosynthetic efficiency and osmotic adjustment, resulting in increased membrane integrity, biomass production and grain yield than in non-acclimated plants.