High-affinity K<Superscript>+</Superscript> transporter <Emphasis Type="Italic">PhaHAK5</Emphasis> is expressed only in salt-sensitive reed plants and shows Na<Superscript>+</Superscript> permeability under NaCl stress

To understand the mechanism of ion homeostasis in salt tolerant and sensitive plants, we isolated cDNAs for K⁺ transporter PhaHAK1-u and PhaHAK5-u from reed plants. PhaHAK1-u belongs to group I and PhaHAK5-u belongs to group IV by phylogenetic analysis, respectively. PhaHAK5-u is predicted to be a plasma membrane transporter, and shows high-affinity K⁺ transporter. Expression of PhaHAK5 was found in salt-sensitive reed plants, but not in any parts of salt-tolerant reed plants maintained under both control and K⁺ starvation conditions. Under the NaCl stress, the K⁺ uptake ability of the yeast strain expressing PhaHAK5-u was remarkably lower than that of the yeast strain expressing PhaHAK1-u, and PhaHAK5-u showed Na⁺ permeability. These results suggest that PhaHAK5 is one of the routes by which Na⁺ enters cells.     

NaCl胁迫对不同耐盐性玉米自交系萌动种子和幼苗离子稳态的影响

盐胁迫影响植物组织的离子分布,不同品种间存在差异。以玉米耐盐自交系81162和8723及盐敏感自交系P138为材料,研究了不同浓度(0、60、140、220 mmol/L)Na Cl胁迫下萌动期种子和幼苗的不同部位中Na+、K+、Ca2+含量以及K+/Na+和Ca2+/Na+比值的变化,旨在探讨不同自交系耐盐性差异的原因。结果表明,在萌动种子中,3个玉米自交系中的Na+积累量表现为种皮胚胚乳,K+累积表现为胚种皮胚乳;幼苗中,Na+积累表现为根茎叶。随着Na Cl浓度的增加,3个玉米自交系萌动种子和幼苗中的Na+含量逐渐升高,但是萌动种子中耐盐自交系81162和8723的Na+增加幅度小于盐敏感自交系P138,Na+含量小于盐敏感自交系P138;幼苗中耐盐自交系81162和8723的Na+增加幅度大于盐敏感自交系P138,幼苗根中Na+含量大于盐敏感自交系P138;茎叶中的Na+含量小于盐敏感自交系P138。随着Na Cl浓度的增加,萌动种子和幼苗中的K+和Ca2+含量逐渐降低。K+离子在耐盐自交系81162和8723萌动种子和幼苗中的降低幅度小于盐敏感自交系P138;Ca2+离子在耐盐自交系81162和8723幼苗中的降低幅度小于盐敏感自交系P138;而在萌动种子中3个自交系Ca2+的流失差异不大。耐盐自交系81162和8723萌动种子和幼苗中K+含量都大于盐敏感自交系P138。耐盐自交系81162和8723的萌动种子和幼苗根中Ca2+含量都大于盐敏感自交系P138;幼苗叶片中则小于盐敏感自交系P138。萌动种子和幼苗中K+/Na+和Ca2+/Na+均随着Na Cl浓度的升高而降低,K+/Na+比值表现为耐盐自交系81162和8723大于盐敏感自交系P138。耐盐自交系81162和8723通过调节离子平衡维持萌动种子和幼苗中较高的K+/Na+比值从而提高耐盐性。     

Salt Tolerance in Aquatic Macrophytes: Ionic Relation and Interaction

Effects of seawater salinity (SWS) and pure NaCl on the intracellular contents of Na⁺, K⁺, Mg²⁺, Ca²⁺, chlorophylls (Chl) and carotenoids (Car) were studied in three submerged aquatic macrophytes, Hydrilla verticillata, Najas indica and Najas gramenia, which differed in their tolerance to salinity. NaCl resulted in significant increase in Chl/Car ratio in the salt-sensitive H. verticillata and moderately salt-tolerant N. indica, but not in the salt-tolerant N. gramenia. SWS treatment did not result in any significant change in the ratio. The intracellular content of Na⁺ increased significantly in all the test plants upon exposure to both NaCl and SWS. The content of K⁺ decreased significantly in these plants upon salinity treatment, except in N. gramenia. The contents of Ca²⁺ and Mg²⁺ decreased significantly upon NaCl treatment and remained unchanged or increased upon SWS treatment. No relationship between salt tolerance and K⁺/Na⁺ ratio was observed. The maintenance of a minimal level of K⁺ was observed to be the most probable requirement of salt tolerance in aquatic macrophytes.     

短期NaCl胁迫对不同小麦品种幼苗K+吸收和Na+、K+积累的影响

为研究盐胁迫下小麦幼苗生长及Na⁺、K⁺的吸收和积累规律,以中国春、洲元9369和长武134等3种耐盐性不同小麦品种为材料,采用非损伤微测技术检测盐胁迫2 d后的根系K⁺离子流变化,并对植株体内的Na⁺、K⁺含量进行测定。结果表明:短期(2d)盐胁迫对小麦生长有抑制作用,且对根系的抑制大于地上部,耐盐品种下降幅度小于盐敏感品种。盐胁迫下,小麦根际的 K⁺大量外流,盐敏感品种中国春K⁺流速显著高于耐盐品种长武134,最高可达15倍。小麦幼苗地上部分和根系均表现为Na⁺积累增加,K⁺积累减少,Na⁺/K⁺比随盐浓度增加而上升。中国春限Na⁺能力显著低于长武134,Na⁺/K⁺则显著高于长武134。综上所述,盐胁迫下造成小麦组织器官中Na⁺/K⁺比上升的主要原因是根系K⁺大量外流和Na⁺的过量积累,耐盐性不同的小麦品种间差异显著,并认为根系对K⁺的保有能力可能是作物耐盐性评价的一个重要指标。     

Responses of salt-tolerant and intolerant wheat genotypes to sodium chloride: Photosynthesis,antioxidants activities,and yield

Physiological responses of two wheat (Triticum aestivum L.) genotypes (salt-tolerant DK961 and salt-sensitive JN17) to increased salt concentrations (50, 100, 150 mM NaCl: NaCl⁵⁰, NaCl¹⁰⁰, NaCl¹⁵⁰) were studied. Photosynthetic capacity, irradiance response curves, contents of soluble sugars, proteins, and chlorophyll (Chl), K⁺/Na⁺ ratio, and activities of antioxidant enzymes (superoxide dismutase, peroxidase, and catalase) in flag leaves were measured on 7 d after anthesis. In control (NaCl⁰) plants, non-significant (p>0.05) differences were found in gas exchange and saturation irradiance (SI) between salt-tolerant (ST) and salt-sensitive (SS) wheat genotypes. However, we found higher soluble sugar and protein contents, K⁺/Na⁺ ratio, and antioxidant enzyme activities, but lower Chl content and yield in ST wheat. Salinity stresses remarkably increased soluble sugar and protein contents and the antioxidant activities, but decreased K⁺/Na⁺ ratio, Chl contents, SI, photosynthetic capacities, and yield, the extent being considerably larger in JN17 than DK961. Although the soluble sugar and protein contents and the antioxidant activities of JN17 elevated more evidently under salt stresses, those variables never reached the high levels of DK961. The antioxidant enzyme activities of SS wheat increased in NaCl⁵⁰ and NaCl¹⁰⁰, but decreased rapidly when the NaCl concentration reached 150 mM. Thus the ST wheat could maintain higher grain yield than the SS one by remaining higher osmoregulation and antioxidative abilities, which led to higher photosynthetic capacity. Hence the ST wheat could harmonize the relationship between CO₂ assimilation (source) and the grain yield (sink) under the experimental conditions.     

Salt-tolerance in plants. II. In vitro translation of m-RNAs from salt-tolerant and salt-sensitive plants on wheat germ ribosomes. Responses to ions and compatible organic solutes

Abstract Messenger RNA from salt-sensitive and salt-tolerant plants Triticum aestivum. Beta vulgaris, Pisum sativum, Chenopodium album and Atriplex nummularia was translated in vitro in a wheatgerm translation system. The optimal monovalent and divalent ion concentrations for translation were independent of the salt tolerance of the plants from which the m-RNAs were derived. Translation was optimal in 100 120 mol m⁻³ potassium acetate and 1.5–2.0 mol m⁻³ Mg²⁺. Substitution of Na⁺ for K⁺, or of Cl⁻ for acetate, was inhibitory. The pattern of polypeptides synthesized from cytoplasmic m-RNAs of salt-sensitive and salt-tolerant plants remained constant in all the conditions examined. The effects of adding the ‘compatible' organic solutes glycine-betaine and mannitol were examined in the wheat-germ system primed with RNA from the leaves of Triticum aestivum or Beta vulgaris. The rate of translation, the optimum ionic concentrations and the distribution of polypeptide products were maintained in organic solute concentrations of up to 500 mol m⁻³. Proline above 300 mol m⁻³ and surcose above 100 mol m⁻³ did inhibit translation. The results indicate that translation in plants is unlikely in cytoplasmic K⁺ concentrations exceeding 180 mol m⁻³, but would proceed in the presence of up to 500 mol m⁻³ mannitol or glyinebetaine, or of up to 300 mol m⁻³ proline.     

Alleviation of detrimental effects of NaCl by silicon nutrition in salt-sensitive and salt-tolerant genotypes of sugarcane (Saccharum officinarum L.)

Salt stress is a major environmental factor which adversely affects the crop yield and quality. However, adequate regulation of mineral nutrients may ameliorate the deleterious effects of salts and help to sustain crop productivity under salt stress. Salt-sensitive (SPF 213) and salt-tolerant (HSF 240) sugarcane genotypes were grown in gravel at 0 and 100 mM NaCl by supplying 0, 1.4 mM, 2.1 mM and 2.8 mM of Si as calcium silicate. Results revealed that plants treated with NaCl alone showed a significant (P?≤?0.05) reduction in dry matter production, K⁺ concentration, cane yield and juice quality in both genotypes but the magnitude of reduction was relatively more in salt-sensitive genotype than salt-tolerant. Addition of Si significantly (P?≤?0.05) reduced the uptake and translocation of Na⁺ but increased K⁺ concentrations particularly in shoots of both sugarcane genotypes. Cane yield and yield attributes were significantly (P?≤?0.05) higher where Si was added. Juice quality characteristics were significantly (P?≤?0.05) improved in salt-sensitive and salt-tolerant sugarcane genotypes with the application of Si. The results suggested that added Si interacted with Na⁺, reduced its uptake and transport to shoots and consequently improved cane yield and juice quality in salt-sensitive and salt-tolerant sugarcane genotypes under salt stress.     

The Effect of Salinity on Growth, Cation Content, Na+-Uptake and Translocation in Salt-Sensitive and Salt-Tolerant Plantago Species

Growth of salt-sensitive Plantago media L. and salt-tolerant P. coronopus L. and P. maritima L. was followed under saline conditions. Growth was reduced according to the ecological features of these species: P. media was sensitive to 25 mM NaCl, while P, coronopus and P. maritima could grow in 150 mM and 300 mM NaCl, respectively. The three Plantago species accumulated Na⁺ in the shoot and maintained a relatively low Na⁺ level in the root. K⁺. Mg²⁺ and Ca²⁺ levels of both shoots and roots decreased with increasing salinity. The results suggest that the difference between salt-resistant and salt-sensitive species is located in the ion secretory system which is involved in the ion translocation from the root to the shoot rather than in the primary uptake process through the plasmalemma of the cortical cells.     

Effects of salinity on the contents of polyamines and some other compounds in sunflower plants differing in salt tolerance

The effects of salt stress (50, 100, and 150 mM NaCl) on the levels of free, bound, and total polyamines were studied in the leaf tissues of salt-tolerant (Coban) and salt-sensitive (Sanbro) cultivars of sunflower (Helianthus annuus L.) plants grown for 15 or 25 days under salinity. The amounts of free, acid-soluble bound, and total spermine increased in leaf tissues of sunflower plants subjected to salt stress while the levels of other polyamines decreased or no significant changes occurred. The increase in some PA titers suggests their potential role in overcoming the adverse effect of salinity stress. The salt sensitivity of the sunflower plants was associated with the excessive accumulation of total polyamines in the leaf tissues of salt-sensitive cultivar (Sanbro) under saline condition. The content of other compounds such as proline, protein, and ions (Na⁺, K⁺, Cl^-, K⁺/Na⁺) in leaf tissue changed depending on salt concentration and the cultivars used.From Fiziologiya Rastenii, Vol. 52, No. 1, 2005, pp. 36–42.Original English Text Copyright © 2005 by Mutlu, Bozcuk.This article was presented by the authors in English.     

Physiological Characteristics and Alternative Respiratory Pathway under Salt Stress in Two Wheat Cultivars Differing in Salt Tolerance

Changes in respiratory pathway, dry weight, contents of proline, ATP, Na⁺and K⁺were investigated under five salinity treatments in the leaves of plants of spring wheat (Triticum aestivumL.). Two cultivars differing in salt resistance, namely, 89122 (salt-tolerant) and Longchun13 (salt-sensitive), were used. The decrease in dry weight and K⁺content was observed with the increasing NaCl concentrations, but more in cv. 89122 plants than in Longchun13 plants. Contents of proline and Na⁺in both cvs increased greatly, but the former increased more in 89122 while the latter more in Longchun13 plants. In all salinity treatments tested, a salt-induced increase in the activity of the alternative pathway was found, although cytochrome pathway (CP) still remained the main electron transport pathway. ATP production changed in parallel with CP operation. Cv. 89122 plants could produce more ATP than cv. Longchun13 plants exposed to each salinity treatment and their ATP generation could even be stimulated in contrast to its rapidly decline in Longchun13 plants with increased salinity stress. The possible relationship between respiration metabolism and above mentioned physiological changes is discussed.     

Influence of NaCl-salinity on growth,photosynthesis, water relations and solute accumulation in <Emphasis Type="Italic">Phragmites australis</Emphasis>

The aim of this study was to investigate the effects of NaCl-salinity on the physiological attributes in common reed, Phragmites australis (Cav.) Trin. ex Steudel. Plants grew optimally under salinity treatment with standard nutrient solution without added salt and at NaCl concentrations up to 100 mM. Applied for 21 days, NaCl-salinity (300 and 500 mM) caused a significant reduction in growth allocation of all different tissues of P. australis. Shoot growth of reed plants displayed a highly significant correlation with plant–water relations and photosynthetic parameters. The net photosynthetic rate and stomatal conductance of reed plants treated with NaCl-salinity at varying osmotic potential (ψ_π) of nutrient solutions were positively correlated, and the former variable also had a strong positive relationship with transpiration rate. Leaf water potential and ψ_π followed similar trends and declined significantly as ψ_π of watering solutions was lowered. The increase in total inorganic nutrients resulting from increased Na⁺ and Cl⁻ in all tissues and K⁺, Ca²⁺ and Mg²⁺ concentrations were maintained even at the most extreme salt concentration. Common reed exhibited high K⁺/Na⁺ and Ca²⁺/Na⁺ selectivity ratios over a wide range of salinities under NaCl-salinity. These findings suggest that reed plants were able to adapt well to high salinities by lowering their leaf ψ_π and the adjustment of osmotically active solutes in the leaves.     

Salt tolerance in rice in vitro: Implication of accumulation of Na+, K+ and proline

The implication of accumulation of both inorganic (Na⁺, K⁺) and organic (proline) solutes were evaluated in unadapted and NaCl-adapted callus of a salt-sensitive (Basmati 370) and a salt-tolerant (SR-26B) cultivar of rice (Oryza sativa L.) after a NaCl shock. Accumulation of Na⁺,K⁺ and/or proline in callus was co relatable and the relative presence of these components in tissues after shock treatment was found to be important factors to support differential regrowth capacities of the shock treated calluses. Presence or retention of K⁺ in rice callus was a key factor for salt tolerance as it was observed to be positively correlated with growth in both the varieties. The results indicated that K⁺ was the first candidate to counteract the negative water potential of outside milieu, while proline was probably the last metabolic device that rice calluses opted for when exposed to salt stress.     

Salinity resistance in Zea mays: fluxes of potassium, sodium and chloride, cytoplasmic concentrations and microsomal membrane lipids

Abstract Cytoplasmic concentrations, fluxes of K⁺, Na⁺ and Cl and microsomal membrane lipids were investigated in a salt-sensitive and salt-resistant variety of Zea mays. The salt resistance of Protador relative to LG_H (salt-sensitive) appears to be related to higher K⁺ fluxes and cytoplasmic concentrations, and lower Na⁺ and Cl fluxes and cytoplasmic concentrations, when grown in NaCl. There were no apparent differences in the simple chemical composition of root microsomal membrane lipids between the two varicties, neither were these affected by salt.     

Effect of salt stress on photosynthesis and physiological parameters of three contrasting barley genotypes

In order to understand the physiological traits important in conferring salt tolerance in three barley genotypes, this study was performed under field conditions with three water salinity levels (2, 10, and 18 dS m–1). High salinity decreased net photosynthetic rate, transpiration rate, and stomatal conductance, K⁺ concentration, K⁺:Na⁺ ratio, and grain yield, but increased electrolyte leakage and Na⁺ content. Under 10 and 18 dS m–1 salinity, Khatam (salt-tolerant) had the maximum stomatal conductance, K⁺, K⁺:Na⁺ ratio, and the grain yield, and a minimum Na⁺ content and electrolyte leakage, whereas Morocco (salt-sensitive) had the lowest net photosynthetic rate, stomatal conductance, K⁺ content, K⁺:Na⁺ ratio, and grain yield, and the highest Na⁺ content and electrolyte leakage. This study showed that tolerant genotypes of barley may avoid Na⁺ accumulation in aboveground parts, facilitating a higher photosynthetic rate and higher grain yield.     

Expression of the AKT1-type K+ channel gene from Puccinellia tenuiflora, PutAKT1, enhances salt tolerance in Arabidopsis

Potassium channels are important for many physiological functions in plants, one of which is to regulate plant adaptation to stress conditions. In this study, a K⁺ channel PutAKT1 cDNA was isolated from the salt-tolerant plant Puccinellia tenuiflora. A phylogenetic analysis showed that PutAKT1 belongs to the AKT1-subfamily in the Shaker K⁺ channel family. PutAKT1 was localized in the plasma membrane and it was preferentially expressed in the roots. The expression of PutAKT1 was induced by K⁺-starvation stress in the roots and was not down-regulated by the presence of excess Na⁺. Arabidopsis plants over-expressing PutAKT1 showed enhanced salt tolerance compared to wild-type plants as shown by their shoot phenotype and dry weight. Expression of PutAKT1 increased the K⁺ content of Arabidopsis under normal, K⁺-starvation, and NaCl-stress conditions. Arabidopsis expressing PutAKT1 also showed a decrease in Na⁺ accumulation both in the shoot and in the root. These results suggest that PutAKT1 is involved in mediating K⁺ uptake (i) both in low- and in high-affinity K⁺ uptake range, and (ii) unlike its homologs in rice, even under salt-stress condition.     

Four Pathogenic Candida Species Differ in Salt Tolerance

The virulence of Candida species depends on many environmental conditions, including extracellular pH and concentration of alkali metal cations. Tests of the tolerance/sensitivity of four pathogenic Candida species (C. albicans, C. dubliniensis, C. glabrata, and C. parapsilosis) to alkali metal cations under various growth conditions revealed significant differences among these species. Though all of them can be classified as rather osmotolerant yeast species, they exhibit different levels of tolerance to different salts. C. parapsilosis and C. albicans are the most salt-tolerant in general; C. dubliniensis is the least tolerant on rich YPD media and C. glabrata on acidic (pH 3.5) minimal YNB medium. C. dubliniensis is relatively salt-sensitive in spite of its ability to maintain as high intracellular K⁺/Na⁺ ratio as its highly salt-tolerant relative C. albicans. On the other hand, C. parapsilosis can grow in the presence of very high external NaCl concentrations in spite of its high intracellular Na⁺ concentrations (and thus lower K⁺/Na⁺ ratio) and thus resembles salt-tolerant (halophilic) Debaryomyces hansenii.     

Selection of callus cultures of sugarcane (Saccharum sp.) tolerant to NaCl and their response to salt stress

Stable callus cultures tolerant to NaCl (68 mM) were developed from salt-sensitive sugarcane cultivar CP65-357 by in vitro selection process. The accumulation of both inorganic (Na⁺, Cl⁻ and K⁺) and organic (proline and soluble sugars) solutes was determined in selected and non-selected calli after a NaCl shock in order to evaluate their implication in in vitro salt tolerance of the selected lines. Both salt-tolerant and non-selected calli showed similar relative fresh weight growth in the absence of NaCl. No growth reduction was observed in salt-tolerant calli while a significant reduction about 32% was observed in nonselected ones when both were cultivated on 68 mM NaCl. Accumulation of Na⁺ was similar in both salt-tolerant and non-selected calli in the presence of NaCl. Accumulation of Cl⁻ was lower in NaCl-tolerant than in non-selected calli while proline and soluble sugars were more accumulated in salt-tolerant than in non-selected calli when both were exposed to salt. K⁺ level decreased more severely in non-selected calli than in NaCl-tolerant ones after NaCl shock. The results indicated that K⁺ and Cl⁻ may play a key role in in vitro salt-tolerance in sugarcance cell lines obtained by in vitro selection and that organic solutes could contribute mainly to counteract the negative water potential of the outside medium.     

Salt-tolerant reed plants contain lower Na<Superscript>+</Superscript> and higher K<Superscript>+</Superscript> than salt-sensitive reed plants

Reed plants (Phragmites australis Trinius) grow not only in fresh and brackish water areas but also in arid and high salinity regions. Reed plants obtained from a riverside (Utsunomiya) were damaged by 257 mM NaCl, whereas desert plants (Nanpi) were not. When the plants were grown under salt stress, the shoots of the Utsunomiya plants contained high levels of sodium and low levels of potassium, whereas the upper part of the Nanpi plants contained low levels of sodium and high levels of potassium. One month salt stress did not affect potassium contents in either Utsunomiya or Nanpi plants, but it did dramatically increase sodium contents only in the Utsunomiya plants. The ratio of K⁺ to Na⁺ was maintained at a high level in the upper parts of the Nanpi plants, whereas the ratio markedly decreased in the Utsunomiya plants in the presence of NaCl. Accumulation of Na⁺ in the roots and Na⁺ efflux from the roots were greater in the Nanpi plants than in the Utsunomiya plants. These results suggest that the salt tolerance mechanisms of Nanpi reed plants include an improved ability to take up K⁺ to prevent an influx of Na⁺ and an improved ability to exclude Na⁺ from the roots.