Mechanism of tolerance in rice varieties differing in sodicity tolerance

Summary The performance of two rice varieties selected for sodicity tolerance and sensitivity was studied in pot culture. Tolerant variety CSR-1 showed much better survival, plant growth and yield as compared to the sensitive one (Basmati-370). Sensitive variety was characterized by higher reduction in growth yield, higher Na and lower K concentrations under sodicity in all the plant organs. Highest Na concentrations are in sensitive variety and its distribution is in the order of Senescent yellow leaves > Earheads > Stem > Younger green leaves. Analysis of different leaves revealed leaf to leaf gradients in terms of Na and K and Na/K ratio were lowest in the youngest leaf and highest in the oldest surviving leaf and these values were much higher in the sensitive variety. This showed that Na ions are mainly localised in the older leaves, which are shed and provide a secondary process of tolerance. Thus the tolerant variety is best marked by maintenance of K levels and much lower increases in Na in all the plant organs, especially earheads, inspite of their growing in highly sodic soils.     

獐茅盐腺形态结构及其泌盐性

显微观察结果显示,獐茅盐腺为典型的双细胞结构,主要分布在叶脉附近,这样有利于它快速收集来自根部的盐离子。同时,X-ray微区分析结果表明,獐茅盐腺可以有效地将Na^＋从表皮细胞和叶肉细胞转运到基细胞,由帽细胞分泌到体外,从而降低植物体内盐分水平以适应盐渍环境。用不同盐类处理材料,獐茅对Na^＋、K^＋和Ca^2＋的吸收和分泌均表现为具有不同的选择性,分泌量的顺序为Na^＋〉K^＋〉Ca^2＋,并且仅仅24h内,盐分的分泌量就已经超过了叶片内的含量,结果植株体内的总离子含量水平几乎不变,说明獐茅具有较强的泌盐能力。     

A model for the evolution of pathogen-induced leaf shedding

A size-structured plant population model is developed to study the evolution of pathogen-induced leaf shedding under various environmental conditions. The evolutionary stable strategy (ESS) of the leaf shedding rate is determined for two scenarios: i) a constant leaf shedding strategy and ii) an infection load driven leaf shedding strategy.
The model predicts that ESS leaf shedding rates increase with nutrient availability. No effect of plant density on the ESS leaf shedding rate is found even though disease severity increases with plant density.
When auto-infection, that is increased infection due to spores produced on the plant itself, plays a key role in further disease increase on the plant, shedding leaves removes disease that would otherwise contribute to disease increase on the plant itself. Consequently leaf shedding responses to infections may evolve. When external infection, that is infection due to immigrant spores, is the key determinant, shedding a leaf does not reduce the force of infection on the leaf shedding plant. In this case leaf shedding will not evolve.
Under a low external disease pressure adopting an infection driven leaf shedding strategy is more efficient than adopting a constant leaf shedding strategy, since a plant adopting an infection driven leaf shedding strategy does not shed any leaves in the absence of infection, even when leaf shedding rates are high. A plant adopting a constant leaf shedding rate sheds the same amount of leaves regardless of the presence of infection.
Based on the results we develop two hypotheses that can be tested if the appropriate plant material is available.     

Penetration, Retention and Transport of Foliar Applied Single Salts of Na, K, Rb and Cs

The penetration, retention and transport of foliar applied chloride salts of Na-22, K-43, Rb-86, and Cs-134 were studied under controlled environmental conditions. Penetration although occurring at different rates was found to be non-selective and appears to be a passive free diffusion process. Retention in the treated area is similar for Na, K and Rb and greater for Cs. Transport of these elements away from that area and in the plant was greatest and fastest for Na and Rb and slower for Cs and K. Distribution in the plants following leaf treatment was similar for K, Rb and Cs but no significant upward transport was noted for Na. Losses from roots were greatest for Na. No significant K could be detected in the culture medium 4 days after treatment of the leaves.     

Na+/H+ exchange activity in the plasma membrane of Arabidopsis

In plants, Na+/H+ exchangers in the plasma membrane are critical for growth in high levels of salt, removing toxic Na+ from the cytoplasm by transport out of the cell. The molecular identity of a plasma membrane Na+/H+ exchanger in Arabidopsis (SOS1) has recently been determined. In this study, immunological analysis provided evidence that SOS1 localizes to the plasma membrane of leaves and roots. To characterize the transport activity of this protein, purified plasma membrane vesicles were isolated from leaves of Arabidopsis. Na+/H+ exchange activity, monitored as the ability of Na to dissipate an established pH gradient, was absent in plants grown without salt. However, exchange activity was induced when plants were grown in 250 mm NaCl and increased with prolonged salt exposure up to 8 d. H+-coupled exchange was specific for Na, because chloride salts of other monovalent cations did not dissipate the pH gradient. Na+/H+ exchange activity was dependent on Na (substrate) concentration, and kinetic analysis indicated that the affinity (apparent Km) of the transporter for Na+ is 22.8 mm. Data from two experimental approaches supports electroneutral exchange (one Na+ exchanged for one proton): (a) no change in membrane potential was measured during the exchange reaction, and (b) Na+/H+ exchange was unaffected by the presence or absence of a membrane potential. Results from this research provide a framework for future studies into the regulation of the plant plasma membrane Na+/H+ exchanger and its relative contribution to the maintenance of cellular Na+ homeostasis during plant growth in salt.     

Trade-off between leaf chlorophyll and betacyanins in Suaeda salsa in the Liaohe estuary wetland in northeast China

Aims Pigment composition is an important functional trait that can be affected by environmental factors. The objective of this study was to investigate the effect of soil salinity on pigment composition in Suaeda salsa by comparing chlorophyll and betacyanin content in the Liaohe estuary wetland, a typical coastal wetland in northeast China.Methods We investigated the plant biomass, percentage of red leaves and pigment content (chlorophyll a, chlorophyll b and betacyanins) in S. salsa in intertidal and supratidal zones of the upper, middle and lower reaches of the Liaohe estuary wetlands. The Na + content of both the soil and plant was also measured. Full analysis of variance and multivariate analysis were used to compare differences in pigment content and Na + content between the supratidal and intertidal zones.Important findings Pigment composition was significantly affected by soil salinity. With increasing soil salinity, the percentage of red leaves was higher in the intertidal zone than in the supratidal zone. In all three reaches, plants had lower chlorophyll a content and higher betacyanin content in the intertidal zone than in the supratidal zone. Compared to chlorophyll a, chlorophyll b was less sensitive to soil salinity. There were no differences in chlorophyll b content between the intertidal and supratidal zones in the upper and lower reaches. Furthermore, pigment composition was associated with both the plant tissue and soil Na + content. Compared to the supratidal zone, the intertidal zone had a higher Na + content in plants. There was a negative relationship between plant chlorophyll content and soil Na + content, but a positive relationship between betacyanin content and soil Na + content. Overall, the results indicated that there might be a trade-off between leaf chlorophyll and betacyanin content in S. salsa to maintain its growth and survival in high salinity environments.     

The arabidopsis Na+/H+ exchanger AtNHX1 catalyzes low affinity Na+ and K+ transport in reconstituted liposomes.

In saline environments, plants accumulate Na(+) in vacuoles through the activity of tonoplast Na(+)/H(+) antiporters. The first gene for a putative plant vacuolar Na(+)/H(+) antiporter, AtNHX1, was isolated from Arabidopsis and shown to increase plant tolerance to NaCl. However, AtNHX1 mRNA was up-regulated by Na(+) or K(+) salts in plants and substituted for the homologous protein of yeast to restore tolerance to several toxic cations. To study the ion selectivity of the AtNHX1 protein, we have purified a histidine-tagged version of the protein from yeast microsomes by Ni(2+) affinity chromatography, reconstituted the protein into lipid vesicles, and measured cation-dependent H(+) exchange with the fluorescent pH indicator pyranine. The protein catalyzed Na(+) and K(+) transport with similar affinity in the presence of a pH gradient. Li(+) and Cs(+) ions were also transported with lower affinity. Ion exchange by AtNHX1 was inhibited 70% by the amiloride analog ethylisopropyl-amiloride. Our data indicate a role for intracellular antiporters in organelle pH control and osmoregulation.     

Why does Viola hondoensis (Violaceae) shed its winter leaves in spring?

? Premise of the study: Viola hondoensis is a perennial herb that inhabits the understory of temperate, deciduous forests. It is an evergreen plant with a leaf life span that is shorter than a year. Its summer leaves are produced in spring and shed in autumn; winter leaves are produced in autumn and shed in spring. Here we asked why the plant sheds its winter leaves in spring, though climate conditions improve from spring to summer. We proposed four hypotheses for the cause of shedding: (1) changes in seasonal environment such as day length or air temperature, (2) shading by canopy deciduous trees, (3) self-shading by taller summer leaves, and (4) competition for nutrients between summer and winter leaves. ? Methods: To test these hypotheses, we manipulated the environment of winter leaves: (1) plants were transplanted to the open site where there was no shading by canopy trees. (2) Petioles of summer leaves were anchored to the soil surface to avoid shading of winter leaves. (3) Sink organs were removed to eliminate nutrient competition. ? Key results: Longevity of winter leaves was extended when shading by summer leaves was eliminated and when sink organs were removed, but not when plants were transplanted to the open site. ? Conclusion: We conclude that the relative difference in light availability between summer and winter leaves is a critical factor for regulation of leaf shedding, consistent with the theory of maximization of the whole-plant photosynthesis.     

Arrested Leaf Abscission in the Non-Abscising Variety of Pubescent Birch: Developmental,Morphological and Hormonal Aspects

A novel type of abscission development, arrested abscission,is described for a rare variety of pubescent birch (Betula pubescensEhrh. f. hibernifolia Ulvinen), the leaves of which dehydrateand remain attached to the plant. Anatomical examination ofthe abscission zone revealed that its development is temporallyand spatially similar to that of normal leaves of Betula pubescens.Floodingand exogenous ethylene were effective in inducing shedding ofthe leaves at the beginning of the growing season, butthis effectwas lost in senescing leaves. Leaf hydration and abscissionwere retained in the presence of external abscisic acid (ABA)inthe non-abscising variety, which also had a lower level of endogenousABA. The observed responses together with the alteredhormonallevel in this variety suggest that apart from ethylene, ABAis involved in autumnal abscission indirectly by regulatingtheleaf water status. Key words: ABA deficiency, abscisic acid, abscission zone, ethylene, senescenc     

Nutritional characteristics in leaves of native plants grown in acid sulfate,peat, sandy podzolic,and saline soils distributed in Peninsular Thailand

Acid sulfate soils, peat soils, sandy podzolic, and saline soils are widely distributed in Peninsular Thailand. Native plants adapted to such problem soils have grown well, and showed no symptom of mineral deficiency or toxicity. Dominant plants growing in low pH soils (acid sulfate and peat) were Melastoma marabathricum and Melaleuca cajuputi. Since M. marabathricum accumulated a huge amount of aluminum (Al) in leaves, especially in new growing leaves, it can be designated an Al accumulator plant. While M. cajuputi did not accumulate Al in shoot, it can be designated an Al excluder plant. Both plant species adapted well to low pH soils, though a different strategy was used for Al. On the other hand, in acid sulfate and peat soils, M. cajuputi, Panicum repens, Cyperus haspan, and Ischaemum aristatum accumulated large amounts of Na in the leaves (or shoots), even in soil with low exchangeable Na concentration. Thus, when growing in the presence of high Al and Na concentration in soils, plant species have developed two opposite strategies: (1) Al or Na accumulation in the leaf and (2) Al or Na exclusion from the leaf. Al concentration in leaves had a negative relationship with the other mineral nutrients except for N and Mn, and Na concentration in leaves also had a negative relationship with P, Zn, Mn, Cu, and Al. Consequently, Al and Na accumulator plants are characterized by their exclusion of other minerals from their leaves.     

Effects of salinity on growth and cation accumulation of Sporobolus virginicus (Poaceae)

Optimal growth of euhalophytes requires moderate concentrations of salt and, in dicotyledons, is associated with succulence and accumulation of Na(+) in plant tissues. However, reports of salt-stimulated growth in monocotyledons are rare. Relative growth rate (RGR), biomass accumulation, and water content were studied in Sporobolus virginicus (Poaceae), a C(4) chloridoid grass, grown hydroponically with different concentrations of NaCl. Cation concentrations were determined by atomic absorption spectrophotometry. Optimal growth occurred at 100-150 mmol/L NaCl and was not dependent on nitrogen levels or accompanied by accumulation of Na(+) in leaves. Biomass accumulation and RGR in plants grown at 450 mmol/L NaCl were greater than in plants grown at 5 mmol/L. The Na?:?K ratios were lower in leaves than in roots, indicating discrimination in Na(+) and K(+) transport. Secretion of Na(+) increased from 166.5 to 336.7 mmol · g(-1) dry biomass · d(-1) as the NaCl concentration of the nutrient solution increased from 125 mmol/L to 450 mmol/L. Water concentrations of leaves and shoots were significantly greater in plants grown at optimal levels of salinity than in plants grown at lower or higher salinities. These results demonstrate salt-stimulated growth in a monocotyledon.     

Effects of sodium and potassium chlorides on the growth and accumulation of mineral ions by Cyperus involucratus Rottb.

The response of the large emergent aquatic plant Cyperus involucratus Rottb. to NaCl and KCl salinities was studied under glasshouse conditions. Growth was not affected greatly by 50 mM NaCl or 100 mM KCl, but plants treated with 100 mM NaCl or 200 mM KCl were stunted and showed symptoms of salt toxicity. NaCl salinity was more detrimental than KCl, and Cl was more damaging to plant growth than K, but less damaging than Na. Salinity had no effect on plant succulence. NaCl and KCl treatments caused a decrease in concentrations of Ca and Mg in plant tissue, and NaCl-stressed plants accumulated nitrate-N. Na was largely excluded from leaves, whereas K was not. Concentrations up to 4% Cl, 8% K and 2.3% Na (dry wt. basis) were recorded in above-ground plant organs. C. involucratus is potentially a useful plant for inclusion in aquatic plant filters to treat wastewaters, and harvestable parts of the plant (culms and leaves) had over 60% of the total plant K, Na and Cl. The results are discussed with respect to the use of C. involucratus in systems for wastewater reclamation.     

盐胁迫下盐地碱蓬体内无机离子含量分布特点的研究

用不同浓度NaCl溶液处理盐地碱蓬（Suaeda salsa)植株后,测定并比较老叶、幼叶及根部的无机离子含量和对K的选择性,叶片及根部的Na^ 、Cl^-含量随盐度的增加而升高,且累积趋势相似,盐胁迫下根部Na^ 、Cl^-及总离子含量（K^ 、Na^ 、Ca^2 ,NO3^-,Cl^-)明显低于叶片,说明盐地碱蓬地盐胁迫下,以叶片优先积累大量离子（如Na^ ,Cl^-) 为其适应特征。NaCl处理下,叶片的K^ ,Ca^2 含量低于对照,但随盐度的增加保持相对稳定,而根部K^ 含量,K/Na比、对K的选择性则高于叶片,这对盐胁迫下地上部的K^ 亏缺有一定补偿作用。低盐度处理（100mmol/LNaCl)促进NO3^-的吸收,另外随盐度的增加,叶片渗透势下降,渗透调节能力增强,幼叶渗透势低于老叶,但渗透调节能力相同。     

Determination of critical K concentrations in sap from individual leaves or from whole plants using K-interruption experiments

Summary A new method is described for estimating critical K concentrations from K interruption experiments using only 2 treatments. Frequent measurements are made of the growth and K concentration of plants subjected to either continued or interrupted K supply and the data used to define the relation between relative yield and K concentration for the K-deficient plants. Critical concentrations are estimated from the results using a mathematical model of plant growth to interpolate over the critical concentration region of the curve. The method has the advantage that the critical concentrations are determined at the exact time that growth is affected. The method was tested using data from previously published experiments with lettuce in which the concentrations of K were measured in sap from both the total shoot and from individual leaf petioles. The model accurately predicted the form of the relationship between relative yield and K concentration for the total shoot and for young expanding leaves, but consistently deviated from the data for recently matured ones. Average estimates of critical concentration ranged fromca. 18 to 34 mmoll⁻¹ in the young leaves and from 48 to 67 mmoll⁻¹ in the mature ones when Na salts were present or absent respectively. The values for total shoot sap were similar to those for mature leaves. The critical concentrations for young expanding leaves were virtually identical to the minimum believed to be needed for the maintenance of important biochemical processes in individual cells, and suggests that a single critical K concentration for plant sap might apply to a wide range of crops provided an actively growing part of the plant is sampled.     

NaCl处理对盐地碱蓬开花及Na＋、K＋含量的影响

为探讨盐地碱蓬（Suaeda salsa）开花与盐的关系, 研究了1 （对照）、200和400 mmol·L-1 NaCl处理对盐地碱蓬不同分枝和单株开花数目、叶片和茎及花器官中的Na＋、K＋含量及Na＋/K＋比的影响。结果表明： 与对照相比, 200 mmol·L-1 NaCl处理下盐地碱蓬分枝及单株开花数目增加最显著, 单株开花数目增加了69.90%, 花器官中的Na＋含量、Na＋/K＋比分别增加了1.41倍和1.77倍, 而一级叶片的Na＋含量、Na＋/K＋比分别增加了3.96倍和4.96倍, 茎中Na＋含量、Na＋/K＋比分别增加了7.00倍和12.39倍。400 mmol·L-1 NaCl处理下, 单株开花数目增加了19.00%, 花器官中的Na＋含量、Na＋/K＋比分别增加了2.09倍和3.21倍, 而一级叶片的Na＋含量、Na＋/K＋比分别增加了4.28倍和6.50倍, 茎中Na＋含量、Na＋/K＋比分别增加了7.65和15.40倍。这些结果表明, NaCl处理下真盐生植物盐地碱蓬可能通过把Na＋区域化到茎叶而动员其中的K＋到花器官中维持花器官中合适的Na＋/K＋比而促进开花。     

Salinity-induced changes in essential oil,pigments and salts accumulation in sweet basil (Ocimum basilicum) in relation to alterations of morphological development

The objective of the project was to study salinity-induced effects on essential oil, pigments and salts accumulation in sweet basil (Ocimum basilicum, the cultivar Perrie) in relation to the alteration of plant morphological development and yield production. Hydroponically grown plants were exposed to one of six NaCl concentrations (1, 25, 50, 75, 100 and 130 mM NaCl). Inhibitory effects of salinity on biomass production of the shoot and the root, and area of individual leaves were apparent already under cultivation with 25 mM NaCl. Elevation of salinity from 1 to 100 mM NaCl induced 63% and 61% reductions in fresh and dry herb biomass production, respectively. The stress-induced reduction of foliage biomass sourced mainly from inhibition of leaf area development rather than reduction of internode and leaf number. Cl and Na concentrations in the leaves, stems and roots increased with elevation of NaCl concentration in the cultivation solution. While the extent of Cl accumulation was leaves>stems>roots, Na was largely excluded from the leaves and was preferentially accumulated in roots and the stems, potentially accounting for the moderate sensitivity of the leaf tissue to salinity. Salt stress increased the contents of essential oil and carotenoids in the leaves that may further account for the moderate sensitivity of sweet basil to salinity and suggest a potential for agro-industrial production. A twofold increase in both carotenoid concentration and the percent of essential oil in the fresh tissue was observed by elevation of the salinity from 1 to 130 mM NaCl. Overall, the stress induced increase of the percent of essential oil in the tissue in the salinity range 1–75 mM NaCl was about 50%, and thereby compensated for the similar reduction of biomass production in this salinity range, so that oil production on per plant basis was not reduced by salinity.     

星星草耐盐碱生理机制再探讨

用不同浓度Na2CO3处理星星草幼苗,7d后测定叶片相对电导率、渗透势、光合速率（Pn）、气孔导度（Gs）、胞间CO2浓度（Ci）、蒸腾强度（Tr）、水分利用效率（WUE）等生理指标,以研究星星草抗盐碱的生理机制。结果表明,低浓度Na2CO3胁迫下,星星草幼苗可通过渗透调节来减轻细胞所受的伤害,因而膜透性受影响较小,相对电导率上升较少,而Gs基本保持不变,Pn、Tr还有所促进,WUE有所增加,这些结果说明,星星草幼苗对低浓度碱性盐具有一定抗性;但犀星草幼苗的这种抗性有一定限度,随着Na2CO3胁迫浓度的增加,Gs、Tr、Pn依次受到抑制,最终膜透性急剧增加,说明星星草幼苗已受损严重,这些变化具有较强的顺序性。     

OSMOTIC ADJUSTMENT OF PLANTS TO SALINE MEDIA. II. DYNAMIC PHASE

Bernstein , Leon . (U. S. Salinity Lab., Riverside, Calif.) Osmotic adjustment of plants to saline media. II. Dynamic phase. Amer. Jour. Bot. 50(4): 360–370. Illus. 1963.—The time-course of osmotic adjustment in bean and pepper plants to increased salinity of the medium was determined by periodic sampling of plants following salt additions to the medium. Bean plants adjusted to increases of 1 atm OP within a day, the adjustment in roots occurring primarily at night following salt addition at 6 pm , whereas leaves and stems made most of their adjustment in the daytime. Pepper plants did not adjust completely to 1.5 atm NaCl additions in 48 hr, but OP increased by about the same amount in both species (0.5—1.0 atm per day). Diurnal fluctuations in OP of leaves and stems of both species and in roots of pepper were matched by parallel fluctuations in K concentrations. Added NaCl caused increased concentrations of K in leaves and stems which were more or less replaced by more slowly absorbed ions, Ca and Mg in bean leaves and Na in bean stems. Other salts produced comparable immediate effects on K level, but K was replaced more rapidly if the cation added was readily accumulated by the bean (Ca). In roots, Na uptake predominated if Na salts were added but K uptake was important on the CaCl₂ treatment. The K effects suggest a passive distribution of K between the cell and the medium.     

Growth Adaptability and Salt Tolerance Osmoregulation of Aneurolepidium chinense Grown on Saline Grassland

The growth of Aneurolepidium chinense Kitag. grown on different saline soil of a Northeast grassland of China was studied and the K+,Na+proline and citric acid accumulated in plants or their organs were determined . The results showed that A. chinense had a strong adaptability to saline habitat and could grow on soil with a salt content of 0. 088%--1.63%,and pH ranging from 8.3 to 9.8. Sodium,proline and citric acid were accumulated by plants for osmotic adjustment when the salinity and Na+content of the soil increased. A. chinense absorbed Na+ rapidly from the soil and excluded K+ when the soil Na+ concentration was less than 20 μmol/g. Sodium was adsorbed slowly and K+ remained constant in plants when Na+ concentrations of soil were 20–80 μmol/g,and proline and citric acid contents were markedly increased. Sodium and K+ decreased slightly in plant tissue and organic solutes increased when Na+ concentrations exceeded 80 μmol/g in soil. Most K+ was distributed in young leaves which were metabolically active. The major sites of Na+ accumulation were in the roots and older mature leaves;whereas proline and citric acid accumulation occurred primarily in young leaves,mature leaves and stems. Tillering buds had strong ability to absord and accumulate K+,Na+ and Proline.     

Effects of salt secretion on psychrometric determinations of water potential of cotton leaves

Thermocouple psychrometers gave lower estimates of water potential of cotton leaves than did a pressure chamber. This difference was considerable for turgid leaves, but progressively decreased for leaves with lower water potentials and fell to zero at water potentials below about −10 bars. The conductivity of washings from cotton leaves removed from the psychrometric equilibration chambers was related to the magnitude of this discrepancy in water potential, indicating that the discrepancy is due to salts on the leaf surface which make the psychrometric estimates too low. This error, which may be as great as 400 to 500%, cannot be eliminated by washing the leaves because salts may be secreted during the equilibration period. Therefore, a thermocouple psychrometer is not suitable for measuring the water potential of cotton leaves when it is above about −10 bars.