转碱蓬SsNHX1基因烟草的耐盐抗旱性研究

烟草是重要的模式植物和经济作物,盐害和干旱两种环境因子对其生长发育、产量和品质都危害很大。为了提高烟草的耐盐抗旱性,本研究利用农杆菌介导的遗传转化法在烟草中过量表达了碱蓬液泡膜Na~+/H~+逆向转运基因SsNHX1,对转基因烟草的耐盐及抗旱性进行表型鉴定和各项生化指标的检测,以期得到耐盐抗旱表性良好的SsNHX1转基因烟草。表型分析发现,SsNHX1基因过表达株系L1和L5的抗盐能力比野生型显著提高,表现为盐胁迫条件下仍能保持旺盛的生长且根系的伸长未受抑制。SsNHX1过表达株系在叶片和根系中积累了更多的Na~+和K~+,同时Na~+含量增长速率较快,而K~+含量降低速率较缓,并可维持较高的叶片相对含水量和叶绿素含量,及较低的丙二醛含量和相对电导率。干旱胁迫发现,过表达株系受干旱胁迫程度更小,并在复水后迅速恢复正常生长。同时,过表达株系的丙二醛含量和相对电导率显著低于野生型,且维持了较高的叶片相对含水量及叶绿素含量。这些结果说明SsNHX1基因在烟草中过量表达后,降低了盐胁迫和干旱胁迫对烟草根系及细胞膜的损伤,并通过调节离子含量、降低细胞的渗透势,维持了叶片较高的相对含水量和叶绿素含量,最终提高了烟草的抗盐和抗旱性。     

Saline and alkaline stress genotypic tolerance in sweet sorghum is linked to sodium distribution

Salt and alkali stress limit crop growth and reduce agricultural productivity worldwide, which have led to increased interest in enhancing salt tolerance in crop plants. Sweet sorghum (Sorghum bicolor (Linn.) Moench) is a monocotyledonous crop species that shows greater tolerance to salt–alkali stress than most other crops, although the underlying mechanisms behind this tolerance remain unclear. Therefore, we investigated the effects of salt and alkali stresses on two sweet sorghum varieties M-81E, which is stress tolerant, and 314B, which is stress sensitive. Namely, we surveyed plant growth parameters, measured Na⁺ and K⁺ distributions at the level of the whole plant as well as in three specific tissues, and then determined the activities of H⁺-ATPase, H⁺-PPase and Na⁺/H⁺ exchange in root vacuole membranes under stress conditions. Following treatment of the seedlings for 3 days with salt or alkali solutions, the plant growth was inhibited and Na⁺ levels in the whole plant, leaves, sheath, and roots were increased in both genotypes. Under alkali stress, K⁺ levels in the whole plant, leaves, sheath, and roots were decreased in both genotypes. M-81E roots accumulated significantly higher levels of Na⁺ than leaves, whereas the opposite was true for 314B. Under salt stress, both the hydrolytic and proton-transporting activities of V-H⁺-ATPase were enhanced and Na⁺/H⁺ exchange activity was dramatically upregulated, whereas V-H⁺-PPase activity was decreased. M-81E showed a greater capacity to compartmentalize Na⁺ within root cell vacuoles and maintain higher levels of K⁺ uptake compared with 314B, resulting in higher K⁺/Na⁺ transport selectivity in this genotype. These results also demonstrated that H⁺-ATPase activity and ionic homeostasis (Na⁺/K⁺) were likely important contributors to the tolerance of saline-alkali stress and crucially important for understanding alkaline stress in both crops and wild plants.     

Intra‐specific variation for salt tolerance in a potential oil‐seed crop,brown mustard (Brassica juncea (l.) Czern. and Coss.)

Thirty eight accessions of brown mustard (Brassica juncea (L.) Czern. and Coss.) were screened after two weeks growth in solution culture containing 120 mol m‐³ NaCl. Considerable variation for salt tolerance was observed in this set of germplasm, since some accessions showed relatively vigorous growth in saline medium.

In order to determine the consistency of degree of salt tolerance at different growth stages of crop life cycle two salt tolerant accessions, P‐15 and KS‐51 and two salt sensitive 85362 and 85605 were tested at the adult stage in 0(control), 100 and 200 mol m^‐3 NaCl. Both the tolerant accessions produced significantly greater fresh and dry biomass and had considerably higher seed yield than those of the salt sensitive accessions. Analysis of different ions in the leaves showed that salt tolerant accessions contained greater amounts of Na⁺, K⁺ and Ca²⁺ than the salt sensitive accessions, although they did not differ significantly for leaf Cl^‐. Only one salt tolerant accession P‐15 had greater leaf K/Na ratio and K⁺ versus Na⁺ selectivity compared with the tolerant KS‐51 and the two salt sensitive accessions.

From this study it was established that there is a considerable variation for salt tolerance in B.juncea which can be exploited by selection and breeding for improvement of its salt tolerance. Since the degree of salt tolerance in B.juncea does not change at different growth stages of the crop life cycle, selection for salt tolerance at the initial growth stages could provide individuals that would be tolerant at all other growth stages. Accumulation of Na⁺, K⁺ and Ca²⁺ in the leaves are important components of salt tolerance in B.juncea.     

Salt Tolerance of Seedling and Adult Bread Wheat Plants Based on Ion Contents and Agronomic Traits

Abstract

A greenhouse experiment was conducted on two salt‐tolerant, two moderately tolerant, and two sensitive Iranian and exotic bread wheat cultivars and their F₁ generations to investigate the effect of salt stress on ion contents of young leaves, biomass yield, and salt stress tolerance index. The materials were evaluated in gravel culture under high salinity (EC=22.5 dSm^?1) and nonstress (EC=2.0 dSm^?1) conditions. Results of stress intensity showed that K⁺/Na⁺ ratio, biomass yield, and Na⁺ concentration were most affected by salt stress. There was no genetic relationship between Mg²⁺ and Ca²⁺ contents with salt tolerance. However, strong relationships were observed among K⁺/Na⁺ ratio, biomass yield, and stress tolerance index. Factor analysis revealed four factors, which explained 99.79% of the total variation among characters. Three‐dimensional plots based on the first three factor scores confirmed that the most salt‐tolerant cultivar was Roshan (an old Iranian cultivar), and Roshan×Alvand and Kharchia×Roshan and their reciprocal crosses were the best salt‐tolerant crosses.     

河套灌区向日葵耐盐指标评价

在盐分胁迫下筛选作物不同耐盐指标的适用性,可为耐盐育种和分子标记辅助选择育种提供科学依据。为确定河套灌区向日葵的耐盐指标,本研究以当地主栽品种‘LD 5009’为研究对象,2年共选择14个典型地块作为定位观测点,分析向日葵产量、生物量、株高等12个指标对盐分胁迫的响应,筛选随土壤饱和浸提液电导率(ECe)增加而减小的指标,采用非线性最小二乘数值逼近法建立其随不同土层土壤ECe的S型耐盐方程。结果表明:向日葵产量、生物量、株高、叶面积指数、花盘直径、叶片和茎秆K+含量随土壤盐分的增加而下降。其中,0~20 cm土层ECe与生物量的耐盐函数决定系数最大;盐分胁迫对向日葵叶绿素的合成影响不大;盐分胁迫下,Na+含量逐渐增加,而脯氨酸和SOD含量先增加后减小。因此,盐分胁迫下生物量可作为河套灌区向日葵耐盐性分析的关键指标。     

Potassium–sodium interactions in soil and plant under saline‐sodic conditions

About 7% of the total land around the globe is salt‐affected causing a great loss to agriculture. Salt stress refers to the excessive amount of soluble salts in the root zone which induce osmotic stress and ion toxicity in the growing plant. Among toxic ions, sodium (Na⁺) has the most adverse effects on plant growth by its detrimental influence on plant metabolism in inhibiting enzyme activities. An optimal potassium (K⁺) : Na⁺ ratio is vital to activate enzymatic reactions in the cytoplasm necessary for maintenance of plant growth and yield development. Although most soils have adequate amounts of K⁺, in many soils available K⁺ has become insufficient because of large amounts of K⁺ removal by high‐yielding crops. This problem is exacerbated under sodic or saline‐sodic soil conditions as a consequence of K⁺‐Na⁺ antagonism. Here K⁺ uptake by plants is severely affected by the presence of Na⁺ in the nutrient medium. Due to its similar physicochemical properties, Na⁺ competes with K⁺ in plant uptake specifically through high‐affinity potassium transporters (HKTs) and nonselective cation channels (NSCCs). Membrane depolarization caused by Na⁺ makes it difficult for K⁺ to be taken up by K⁺ inward‐rectifying channels (KIRs) and increases K⁺ leakage from the cell by activating potassium outward‐rectifying channels (KORs). Minimizing Na⁺ uptake and preventing K⁺ losses from the cell may help to maintain a K⁺ : Na⁺ ratio optimum for plant metabolism in the cytoplasm under salt‐stress conditions. It would seem a reasonable assumption therefore that an increase in the concentration of K⁺ in salt‐affected soils may support enhanced K⁺ uptake and reduce Na⁺ influx via HKTs and NCCSs. Although very useful information is available regarding K⁺‐Na⁺ homeostasis indicating their antagonistic effect in plants, current knowledge in applied research is still inadequate to recommend application of potassium fertilizers to alleviate Na⁺ stress in plants under sodic and saline‐sodic conditions. Nevertheless some encouraging results regarding alleviation of Na⁺ stress by potassium fertilization provide the motivation for conducting further studies to improve our understanding and perspectives for potassium fertilization in sodic and saline‐sodic environments.     

Growth and photosynthesis of salt‐stressed sunflower (Helianthus annuus) plants as affected by foliar‐applied different potassium salts

In order to assess the effectiveness of foliar‐applied potassium (K⁺, 1.25%) using different salts (KCl, KOH, K₂CO₃, KNO₃, KH₂PO₄, and K₂SO₄) in ameliorating the inhibitory effect of salt stress on sunflower plants, a greenhouse experiment was conducted. Sodium chloride (150 mM) was applied through the rooting medium to 18 d–old plants and after 1 week of salt treatment; different K⁺‐containing salts were applied twice in 1‐week interval as a foliar spray. Salt stress adversely affected the growth, yield components, gas exchange, and water relations, and also caused nutrient imbalance in sunflower plants. However, foliar‐applied different sources of potassium improved shoot and root fresh and shoot dry weights, achene yield, 100‐achene weight, photosynthetic rate, transpiration rate, stomatal conductance, water‐use efficiency, relative water content, and leaf and root K⁺ concentrations of sunflower plants grown under saline conditions. Under nonsaline conditions, improvement in shoot fresh weight, achene yield, 100‐achene weight, photosynthetic and transpiration rates, and root Na⁺ concentration was observed due to foliar‐applied different K sources. Of the different salts, K₂SO₄, KH₂PO₄, KNO₃, and K₂CO₃ were more effective than KCl and KOH in improving growth and some key physiological processes of sunflower plants.     

作物相对耐盐性的研究──Ⅱ．不同栽培作物的耐盐性差异

本文通过盆栽生物试验,对小麦、大豆、棉花、玉米等栽培作物的苗期耐盐性进行了研究.结果表明:棉花较为耐盐,玉米、小麦次之,大豆耐盐性最差.不同作物各组织中钠的浓度和累积量随盐度增加而剧增.小麦、大豆、棉花根系吸收钠后,不同程度地向地上部分转移;玉米根系吸收钠后,多累积在根系中.不同作物各组织中钾的浓度随盐度增加变化不大.但累积量剧减;钙的浓度和累积量随盐度增加都有不同程度的减少.作物根系吸收钾、钙后,向地上部分运输,因而地上部分组织中钾、钙累积量多于根系中钾、钙累积量.作物体内K/Na比随盐度增加而降低.本文还对不同栽培作物耐盐性差异的机理进行了探讨.     

Salinity‐induced changes in growth,superoxide dismutase activity,and ion content of two olive cultivars

Olive trees (Olea europaea L.) are considered moderately tolerant to salinity, with clear differences found among cultivars. One‐year‐old self‐rooted olive plants of the Croatian cv. Oblica and Italian cv. Leccino were grown for 90 d in nutrient solutions containing 0, 66, or 166 mM NaCl, respectively. The shoot length and the number of nodes and leaves for both cultivars were not affected by salinity up to 66 mM NaCl. However, at 166 mM NaCl, growth of Leccino was reduced earlier and to a higher extent than growth of Oblica. After 10 d of exposure to 66 and 166 mM NaCl, increased activity of superoxide dismutase (SOD) was observed in Leccino, whereas there was almost no response in Oblica. Reduced SOD activity in Leccino at 166 mM NaCl was observed after prolonged stress (90 d), whereas in Oblica SOD was increased at 66 mM compared to control or 166 mM NaCl. Electrolyte and K⁺ leakage were increased and relative water content decreased as NaCl concentration increased with similar intensity of response measured in both cultivars. Oblica exhibited an ability to keep a higher K⁺ : Na⁺ ratio at all salinity levels compared to Leccino, but since no difference was found in leaf K⁺ concentration, this was mainly achieved by less Na⁺ ions reaching the younger leaves. The antioxidative system represents a component of the complex olive salt‐tolerance mechanism, and it seems that the role of SOD in protection from oxidative stress depends on sodium accumulation in leaves.     

Sodium removal from a calcareous saline–sodic soil through leaching and plant uptake during phytoremediation

Saline–sodic and sodic soils are characterized by the occurrence of sodium (Na⁺) to levels that can adversely affect several soil properties and growth of most crops. As a potential substitute of cost‐intensive chemical amelioration, phytoremediation of such soils has emerged as an efficient and low‐cost strategy. This plant‐assisted amelioration involves cultivation of certain plant species that can withstand ambient soil salinity and sodicity levels. It relies on enhanced dissolution of native calcite within the root zone to provide adequate Ca²⁺ for the Na⁺ Ca²⁺ exchange at the cation exchange sites. There is a lack of information for the Na⁺ balance in terms of removal from saline–sodic soils through plant uptake and leaching during the phytoremediation process. We carried out a lysimeter experiment on a calcareous saline–sodic soil [pH of saturated soil paste (pH_s) = 7.2, electrical conductivity of the saturated paste extract (EC_e) = 4.9 dS m⁻¹, sodium adsorption ratio (SAR) = 15.9, CaCO₃ = 50 g kg⁻¹]. There were three treatments: (1) control (without application of a chemical amendment or crop cultivation), (2) soil application of gypsum according to the gypsum requirement of the soil and (3) planting of alfalfa (Medicago sativa L.) as a phytoremediation crop. The efficiency of treatments for soluble salt and Na⁺ removal from the soil was in the order: gypsum ≈ alfalfa > control. In the phytoremediation treatment, the amount of Na⁺ removed from the soil through leaching was found to be the principal cause of reduction in salinity and sodicity. Copyright © 2003 John Wiley & Sons, Ltd.     

Selective restriction of root to shoot ion transport by cotyledon node zone in Kochia sieversiana may contribute to its tolerance to salt and alkali stresses

This study is aimed to examine if cotyledon node zone may play a role in salt and/or alkali tolerance. Seedlings of halophyte plant Kochia sieversiana and glycophyte plant Lycopersicon esculintum Mill were treated with salt and alkali respectively, xylem sap was collected from above or below cotyledon node zone, and components and contents of inorganic ions in the sap were examined. When compared with that collected from below cotyledon node zone, xylem sap collected from above cotyledon node zone in K. sieversiana contains less Na⁺ under both salt and alkali stresses, and less chloride (Cl^–) under salt treatment. Both Na⁺ and Cl^– remain nearly the same in xylem sap collected from below and above cotyledon node zone in L. esculintum Mill. Cotyledon node zone in K. sieversiana selectively restricted ion transport under both salt and alkali stresses, which may represent a novel mechanism of salt and alkali resistance in halophyte plants.     

作物对盐分的吸收及其盐害的预测预报

通过温室和田间试验,研究小麦和甜菜在盐胁迫下的生长及其对盐分的吸收。结果表明,小麦耐热略低于甜菜,Ｎａ＾＋抑制小麦和甜菜对Ｋ＾＋和Ｃａ＾２＋的吸收,小麦和甜菜的相对干物质重与土壤含盐量的关系符合Ｍａａｓ－Ｈｏｆｆｍａｎ模型,小麦和甜菜叶Ｎａ＾＋含量与土壤含盐量呈显著正相关,且与干物质重的关系也符合Ｍａａｓ－Ｈｏｆｆｍａｎ模型,文中提出了利用作物叶的Ｎａ＾＋含量与相对干物质重之间的Ｍａａｓ－Ｈｏｆｆ     

Salt‐resistant and salt‐sensitive wheat genotypes show similar biochemical reaction at protein level in the first phase of salt stress

Salinity has a two‐phase effect on plant growth, an osmotic effect due to salts in the outside solution and ion toxicity in a second phase due to salt build‐up in transpiring leaves. To elucidate salt‐resistance mechanisms in the first phase of salt stress, we studied the biochemical reaction of salt‐resistant and salt‐sensitive wheat (Triticum aestivum L.) genotypes at protein level after 10 d exposure to 125 mM–NaCl salinity (first phase of salt stress) and the variation of salt resistance among the genotypes after 30 d exposure to 125 mM–NaCl salinity (second phase of salt stress) in solution culture experiments in a growth chamber. The three genotypes differed significantly in absolute and relative shoot and root dry weights after 30 d exposure to NaCl salinity. SARC‐1 produced the maximum and 7‐Cerros the minimum shoot dry weights under salinity relative to control. A highly significant negative correlation (r² = –0.99) was observed between salt resistance (% shoot dry weight under salinity relative to control) and shoot Na⁺ concentration of the wheat genotypes studied. However, the salt‐resistant and salt‐sensitive genotypes showed a similar biochemical reaction at the level of proteins after 10 d exposure to 125 mM NaCl. In both genotypes, the expression of more than 50% proteins was changed, but the difference between the genotypes in various categories of protein change (up‐regulated, down‐regulated, disappeared, and new‐appeared) was only 1%–8%. It is concluded that the initial biochemical reaction to salinity at protein level in wheat is an unspecific response and not a specific adaptation to salinity.     

Seawater‐irrigation effects on growth,ion concentration,and photosynthesis of transgenic poplar overexpressing the Na+/H+ antiporter AtNHX1

Tonoplast Na⁺/H⁺ antiporters increase the salt resistance of various plant species, but very little is known about the role of these antiporters in the salt resistance of trees. Understanding the physiological responses of plants to salinity stress is of paramount importance in examining the salt resistance of transgenic plants. In this study, the wild‐type poplar (WT; Populus × euramericana var. Neva) and its transgenic varieties (TR) that overexpress the AtNHX1 gene were exposed to various seawater concentrations (0%, 10%, 20%, and 30%) for 30 d to determine the effects of seawater on seedling growth, ion content, and photosynthetic productivity. Results show that TR plants grew much better than WT under saline conditions. Differences between WT and TR in most parameters were significant after 30 d exposure to 20% and 30% seawater concentrations. The dry weight of TR was higher than that of WT for each seawater treatment. Transgenic variety was able to maintain higher photosynthetic ability than WT upon exposure to salinity and maintained higher K⁺ concentrations and K⁺ : Na⁺ ratio but had less Cl^– compared with WT. This suggests that AtNHX1 has a critical role in the regulation of K⁺ homeostasis, which in turn affects plant K⁺ nutrition and salt resistance.     

盐胁迫下氮素形态对油菜和水稻幼苗离子运输和分布的影响

【目的】土壤盐碱化是制约农作物产量的主要因素之一,盐胁迫影响养分运输和分布,造成植物营养失衡,导致作物发育迟缓,植株矮小,严重威胁着我国的粮食生产。在必需营养元素中,氮素是需求量最大的元素,NO-3和NH+4是植物吸收氮素的两种离子形态。植物对盐胁迫的响应受到不同形态氮素的调控,研究不同形态氮素营养下植物的耐盐机制对提高植物耐盐性及产量具有重要的意义。【方法】本文以喜硝植物油菜(Brassica napus L.)和喜铵植物水稻(Oryza sativa L.)为试验材料,采用室内营养液培养方法,研究了NO-3和NH+4对Na Cl胁迫下油菜及水稻苗期生长状况、对Na+运输和积累的影响,以对照与盐胁迫植株生物量之差与Na+积累量之差的比值,评估Na+对植株的伤害程度。【结果】1)在非盐胁迫条件下,硝态氮营养显著促进油菜和水稻根系的生长;盐胁迫条件下,油菜和水稻生物量均显著受到抑制,Na Cl对供应铵态氮营养植株的抑制更为显著。2)盐胁迫条件下,两种供氮形态下,油菜和水稻植株Na+含量均显著增加,硝态氮营养油菜叶柄Na+显著高于铵态氮营养,叶柄Na+含量/叶片Na+含量大于铵营养油菜,硝态氮营养水稻根系Na+含量显著低于铵营养,地上部则相反。3)铵营养油菜和水稻Na+伤害度显著高于硝营养植株。4)盐胁迫条件下,硝态氮营养油菜地上部和水稻根系K+含量均显著高于铵态氮营养。5)盐胁迫条件下,硝营养油菜和水稻木质部Na+浓度,韧皮部Na+和K+浓度及水稻木质部K+浓度均高于铵营养植株。【结论】与铵营养相比,硝营养油菜和水稻具有更好的耐盐性。硝态氮处理油菜叶柄Na+显著高于铵态氮处理,能够截留Na+向叶片运输。同时,供应硝态氮营养更有利于油菜和水稻吸收K+,有助于维持植物体内离子平衡。盐胁迫下,硝营养油菜和水稻木质部Na+浓度,韧皮部Na+和K+浓度及水稻木质部K+浓度均高于铵营养植株,表明硝态氮营养油菜和水稻木质部-韧皮部对离子有较好的调控能力,是其耐盐性高于铵营养的原因之一。     

Responses of Batis maritima plants challenged with up to two‐fold seawater NaCl salinity

Batis maritima is a promising halophyte for sand‐dune stabilization and saline‐soil reclamation. This species has also applications in herbal medicine and as an oilseed crop. Here, we address the plant response to salinity reaching up to two‐fold seawater concentration (0–1000 mM NaCl), with a particular emphasis on growth, water status, mineral nutrition, proline content, and photosystem II integrity. Plant biomass production was maximal at 200 mM NaCl, and the plants survived even when challenged with 1000 mM NaCl. Plant water status was not impaired by the high accumulation of sodium in shoots, suggesting that Na⁺ compartmentalization efficiently took place in vacuoles. Concentrations of Mg²⁺ and K⁺ in shoots were markedly lower in salt‐treated plants, while that of Ca²⁺ was less affected. Soluble‐sugar and chlorophyll concentrations were hardly affected by salinity, whereas proline concentration increased significantly in shoots of salt‐treated plants. Maximum quantum efficiency (F_v/F_m), quantum yield of PSII (Φ_PSII), and electron‐transport rate (ETR) were maximal at 200–300 mM NaCl. Both nonphotochemical quenching (NPQ) and photochemical quenching (qP) were salt‐independent. Interestingly, transferring the plants previously challenged with supraoptimal salinities (400–1000 mM NaCl) to the optimal salinity (200 mM NaCl) substantially restored their growth activity. Altogether, our results indicate that B. maritima is an obligate halophyte, requiring high salt concentrations for optimal growth, and surviving long‐term extreme salinity. Such a performance could be ascribed to the plant capability to use sodium for osmotic adjustment, selective absorption of K⁺ over Na⁺ in concomitance with the stability of PSII functioning, and the absence of photosynthetic pigment degradation.     

Evaluation of Salt Tolerance and Its Relationship with Carbon Isotope Discrimination and Physiological Parameters of Barley Genotypes

Soil management through the cultivation of salt-tolerant plants is a practical approach to combat soil salinization. In this study, salt tolerance of 35 barley (Hordeum vulgare L.) genotypes was tested at four salinity levels (0, 100, 200, and 300 mM NaCl in Hoagland nutrient solution) at two growth stages (germination and vegetative). The relationship between salinity tolerance and carbon isotope discrimination (CID) was also accessed. Results of the study carried out under laboratory conditions showed that a negative linear relationship was observed between salt concentration and germination as well as other growth parameters. Some genotypes showed good salt tolerance at germination but failed to survive at seedling stage. However, five genotypes, namely, Jau-83, Pk-30109, Pk-30118, 57/2D, and Akermanns Bavaria showed better tolerance to salinity (200 mM) both at germination and at vegetative growth stage. The salt tolerance of these barley genotypes was significantly correlated with minimum decrease in K⁺:Na⁺ ratio in plant tissue with increase in the root zone salinity. However, the case was reversed in sensitive genotypes. CID was decreased linearly with increase in root zone salinity. However, salt-tolerant genotypes maintained their turgor by osmotic adjustment and by minimum increase in diffusive resistance and showed minimum reduction in CID (Δ) with gradual increase in rooting medium salt concentration. Results suggested that the tolerant genotypes make osmotic adjustments by selective uptake of K⁺ and by maintaining a higher K⁺:Na⁺ ratio in leaves. Moreover, CID technique can also be good criteria for screening of salt-tolerant germplasm.     

THE ROLE OF CALCIUM IN PLANTS' SALT TOLERANCE

High concentrations of sodium (Na) are toxic to most plant species, making soil salinity a major abiotic stress in plant productivity world wide. It has been shown that, calcium (Ca²⁺) is an important determinant for plant salt tolerance and confers protective effects on plants under growing in sodic soils. Calcium plays an essential role in processes that preserve the structural and functional integrity of plant cell membranes, stabilizes cell wall structures, regulates ion transport and selectivity, and controls ion-exchange behavior as well as cell wall enzyme activities. The nature of these responses will vary depending on the plant genotype. One of the essential functions of Ca²⁺ is acting as a second messenger in stress signaling. Genetic evidence suggests that perception of salt stress leads to a cytosolic calcium-signal that activates the calcium sensor protein SOS₃. SOS₃ binds to and activates a ser/thr protein kinase SOS₂. The activated SOS₂ kinase regulates the activities of SOS₁, a plasma membrane Na⁺/H⁺ antiporter, and NHX₁, a tonoplast Na⁺/H⁺ antiporter. This results in either Na⁺ efflux out of cytosol or its compartmentation in vacuole.     

Proton release by N2‐fixing plant roots: A possible contribution to phytoremediation of calcareous sodic soils

With a world‐wide occurrence on about 560 million hectares, sodic soils are characterized by the occurrence of excess sodium (Na⁺) to levels that can adversely affect crop growth and yield. Amelioration of such soils needs a source of calcium (Ca²⁺) to replace excess Na⁺ from the cation exchange sites. In addition, adequate levels of Ca²⁺ in ameliorated soils play a vital role in improving the structural and functional integrity of plant cell walls and membranes. As a low‐cost and environmentally feasible strategy, phytoremediation of sodic soils — a plant‐based amelioration — has gained increasing interest among scientists and farmers in recent years. Enhanced CO₂ partial pressure (P_CO2) in the root zone is considered as the principal mechanism contributing to phytoremediation of sodic soils. Aqueous CO₂ produces protons (H⁺) and bicarbonate (HCO₃^‐). In a subsequent reaction, H⁺ reacts with native soil calcite (CaCO₃) to provide Ca²⁺ for Na⁺ Ca²⁺ exchange at the cation exchange sites. Another source of H⁺ may occur in such soils if cropped with N₂‐fixing plant species because plants capable of fixing N₂ release H⁺ in the root zone. In a lysimeter experiment on a calcareous sodic soil (pH_s = 7.4, electrical conductivity of soil saturated paste extract (EC_e) = 3.1 dS m^‐1, sodium adsorption ratio (SAR) = 28.4, exchangeable sodium percentage (ESP) = 27.6, CaCO₃ = 50 g kg^‐1), we investigated the phytoremediation ability of alfalfa (Medicago sativa L.). There were two cropped treatments: Alfalfa relying on N₂ fixation and alfalfa receiving NH₄NO₃ as mineral N source, respectively. Other treatments were non‐cropped, including a control (without an amendment or crop), and soil application of gypsum or sulfuric acid. After two months of cropping, all lysimeters were leached by maintaining a water content at 130% waterholding capacity of the soil after every 24±1 h. The treatment efficiency for Na⁺ removal in drainage water was in the order: sulfuric acid > gypsum = N₂‐fixing alfalfa > NH4NO3‐fed alfalfa > control. Both the alfalfa treatments produced statistically similar root and shoot biomass. We attribute better Na+ removal by the N₂‐fixing alfalfa treatment to an additional source of H⁺ in the rhizosphere, which helped to dissolve additional CaCO₃ and soil sodicity amelioration.