Assessment of in vitro growth of apical stem sections and adventitious organogenesis to evaluate salinity tolerance in cultivated tomato

The possible use of in vitro shoot morphogenesis and shoot apex culture to evaluate salt tolerance in cultivated tomato (Lycopersicon esculentum Mill.) has been analyzed, using two cultivars with similar salt tolerance, Pera and Hellfrucht frühstamm (HF). The effect of salt on shoot regeneration was studied by culturing leaf explants on media supplemented with 0, 43, 86, 129 and 172 mM NaCl. The presence of NaCl in the regeneration media at 86 mM strongly inhibited shoot regeneration in the cultivar HF, but not in Pera. However, the substitution of NaCl by mannitol, maintaining the same water potential in the culture media, decreased the regeneration percentage in Pera but did not affect HF. Shoot apices of both cultivars were also subcultured at 6-week intervals, for 4 subcultures, at the same NaCl concentrations as used in the previous experiment, and the shoot growth, leaf and root number, rooted shoot and shoot necrosis were recorded at the end of each subculture. Root formation was the parameter most affected by salt in both cultivars, Pera being more sensitive than HF. The substitution of NaCl by mannitol significantly increased the percentage of rooted shoots in Pera after four subcultures, and slightly decreased this percentage in HF. Shoot necrosis was only observed in the last subculture at NaCl higher than 86 mM, the percentage of necrotic shoots being higher in Pera than in HF (75% and 45%, respectively). The lack of agreement between the results obtained with the in vitro tests, e.g., adventitious shoot formation and growth of apical stem sections, suggests that this approach may not be a reliable tool to evaluate salt tolerance in cultivated tomato. This revised version was published online in June 2006 with corrections to the Cover Date.     

Grafting raises the salt tolerance of tomato through limiting the transport of sodium and chloride to the shoot

With the aim of determining whether grafting could improve salinity tolerance of tomato (Lycopersicon esculentum Mill.), and what characteristics of the rootstock were required to increase the salt tolerance of the shoot, a commercial tomato hybrid (cv. Jaguar) was grafted onto the roots of several tomato genotypes with different potentials to exclude saline ions. The rootstock effect was assessed by growing plants at different NaCl concentrations (0, 25, 50, and 75 mM NaCl) under greenhouse conditions, and by determining the fruit yield and the leaf physiological changes induced by the rootstock after 60 d and 90 d of salt treatment. The grafting process itself did not affect the fruit yield, as non-grafted plants of cv. Jaguar and those grafted onto their own root showed the same yield over time under non-saline conditions. However, grafting raised fruit yield in Jaguar on most rootstocks, although the positive effect induced by the rootstock was lower at 25 mM NaCl than at 50 and 75 mM NaCl. At these higher levels, the plants grafted onto Radja, Pera and the hybrid VolgogradskijxPera increased their yields by approximately 80%, with respect to the Jaguar plants. The tolerance induced by the rootstock in the shoot was related to ionic rather than osmotic stress caused by salinity, as the differential fruit yield responses among graft combinations were mainly related to the different abilities of rootstocks to regulate the transport of saline ions. This was corroborated by the high negative correlation found between fruit yield and the leaf Na(+) or Cl(-) concentrations in salt-treated plants after 90 d of salt treatment. In conclusion, grafting provides an alternative way to enhance salt tolerance, determined as fruit yield, in the tomato, and evidence is reported that the rootstock is able to reduce ionic stress.     

Effect of Short-Term Salinity on Lipid Metabolism and Ion Accumulation in Tomato Roots

To examine the ion accumulation and membrane lipid metabolism in response to salinity we compared two tomato cvs. Pera and Hellfrucht Fruhstamm (HF), considered to be salt-tolerant and sensitive respectively. Na⁺ and K⁺ accumulation was significantly higher in roots of cv. Pera after 24 h and 72 h of 100 mM NaCl. While in cv. HF, a temporary increase in K⁺ accumulation at 24 h was accompanied by a sustained increase in Na⁺ content. Both cultivars enhanced incorporation of [³²P]orthophosphate into phosphatidylinositol 4,5-bisphosphate at 24 h and 72 h of NaCl. In parallel to the increase of phosphatidylinositol 4,5-bisphosphate a decrease in phosphorylation of phosphatidic acid and phosphatidylcholine were observed in the sensitive cv. HF. Structural and signal lipid changes in response to salinity were more evident in the sensitive cv. HF. Salt tolerant cv. Pera accumulated Na⁺ ions in the roots without considerable modifications in lipid metabolism. This revised version was published online in July 2006 with corrections to the Cover Date.     

Genetic variability of NaCl tolerance in tomato

Cultivation of crops in soils with high salt (NaCl) content can affect plant development. We examined the morphological and physiological mechanisms of salt tolerance in tomato. The responses of 72 accessions of tomato (Solanum lycopersicum) to salinity were compared by measuring shoot and root lengths, and fresh shoot and root weights relative to those of controls (plants grown in normal salt levels). All traits were reduced at the seedling stage when salinity levels were increased. The accession x salinity interaction was significant for all traits. Root length had higher heritability than other traits and was used as a selection criterion to identify salt-tolerant and -non-tolerant accessions. On the basis of root length, accessions LA2661, CLN2498A, CLN1621L, BL1176, 6233, and 17870 were considered to be more tolerant than accessions 17902, LO2875 and LO4360. The degree of salt tolerance was checked by analyzing K+ and Na+ concentrations and K+/Na+ ratio in tissues of plants treated with 10 and 15 dS/m salinity levels. Tolerance of these accessions to salinity was most associated with low accumulation of Na+ and higher K+/Na+ ratios.     

Responses to NaCl stress of cultivated and wild tomato species and their hybrids in callus cultures

Summary If in vitro culture is to be used for evaluating the salt tolerance of tomato hybrids and segregant populations in a breeding programme, it is previously necessary to get quick and reliable traits. In this work, growth and physiological responses to salinity of two interspecific hybrids between the cultivated tomato (Lycopersicon esculentum Mill) and its wild salt-tolerant species L pennellii are compared to those of their parents. The leaf callus of the first subculture was grown on media amended with 0, 35, 70, 105, 140, 175 and 210 mM NaCl for 40 days. Relative fresh weight growth of callus in response to increased salinity in the culture medium was much greater in L pennellii than in the tomato cultivars, and greater in the hybrids than in the wild species. Moreover, the different salt tolerance degree of hybrids was related to that of female parents. At high salt levels, only Cl^– accumulation was higher in L pennellii than in tomato cultivars, whereas in the hybrids both Cl^–, and Na⁺ accumulation were higher than in their parents. Proline increased with salinity in the callus of all genotypes; these increases were much higher in the tomato cultivars than in L pennellii, and the hybrids showed a similar response to that of the wild species. Salt-treated callus of the tomato cultivars showed significant increases in valine, isoleucine and leucine contents compared to control callus tissue. In contrast, these amino acids in callus tissues of the wild species and hybrids showed a tendency to decrease with increasing salinity.     

The effectiveness of grafting to improve salt tolerance in tomato when an ‘excluder’ genotype is used as scion

If the main effect of long-term exposure of tomato plants to salinity is the accumulation of toxic concentrations of Na⁺ and Cl⁻ in the leaves, then the selection of ‘excluder’ rootstocks should increase tolerance to salinity in grafted tomato plants, independently of the genotype used as the scion. The question addressed in this study is whether shoot genotypes with an ‘excluder’ character are able to increase their salt tolerance when grafted onto rootstocks of the same characteristics. Moneymaker (with excluder character) was grafted onto two root genotypes, Radja and Pera, selected for their very different ability to regulate the transport of saline ions to the shoot over time. Grafting onto either Pera or Radja improved fruit yield compared to the self-grafted plants of Moneymaker (M/M) when the plants were grown at 50 mM NaCl, whereas there was no effect of either rootstock or of grafting per se (M/M) on fruit yield in the absence of or at 25 mM NaCl. The relationship between the salt responses to mid- and long-term depended on the stress level; after 27 d of 150 mM NaCl treatment, both graft combinations enhanced similarly their salt tolerances as did in the long-term experiment. Moreover, the tolerance induced by rootstock was related to the low rates of saline ion accumulation in their leaves. However, the positive effect of rootstock was only observed with rootstock Pera when the grafted plants were grown at 50 mM NaCl (the same salt level used in the long-term experiment) for 35 d. According to the physiological changes induced by rootstock in the leaves, the different salt responses seem to be due to the fact that the osmotic effect predominated on the toxic effect under these last conditions. Consequently, in order to select rootstocks care must be taken in the timing of any selection process: the stress level and length of exposure to salinity must be sufficient for the true differences in salt tolerance for toxicity to be shown. Taken together, these results show the effectiveness of grafting to enhance fruit yield in tomato and provide evidence that the positive effect induced by rootstock is related to the re-establishment of ionic homeostasis.     

Soil salinity effect on soluble saccharides,phenol, fatty acid and mineral contents,and respiration rate of garlic cultivars

The effect of salinity on contents of water, soluble saccharides, phenols, minerals and on respiration rate in bulbs of five garlic (Allium sativum L.) cultivars differing in salinity tolerance was determined. Cultivar HG-6 was found to be the most tolerant followed by cvs. G-l and G-42, and cv. Aru the least tolerant to salinity. The cultivars which were tolerant showed lesser reduction in water content of the bulbs by salinity. Initial contents of phenolic compounds and sulphur were comparatively low in salinity tolerant cultivars but they increased under high salinity levels whereas reverse was found in salt sensitive cultivars. The fatty acids profile did not show significant changes under saline conditions. Contents of K and Ca were reduced, content of Na was increased and there were no changes in the contents of N, Mn, Cu, Zn and Fe. The changes in soluble saccharides content and respiration rate were not found to be associated with the salinity tolerance.     

In vitro Growth and Leaf Composition of Grapevine Cultivars as Affected by Sodium Chloride

In vitro proliferated shoot culture of six grape genotypes (Vitis vinifera L.) were screened for tolerance to NaCl (0 to 200 mM). The cv. Perlette was found to tolerate 175 mM NaCl followed by cvs. Pusa Seedless and Beauty Seedless 150 mM NaCl. Na, K, Cl, Ca and Mg content increased upto 100 mM NaCl in most of the genotypes. Total sugar and proline content of stem tissue gradually increased under NaCl stress while leaf chlorophyll a+b content declined. Studies suggest that the in vitro screening procedure can be used for ranking the grape genotypes for salinity tolerance.     

Growth and ion accumulation of two rapid-cycling Brassica species differing in salt tolerance

The response of two rapid-cycling Brassica species differing in tolerance to seawater salinity was studied over a period of 24 days. In response to 8 dS m⁻¹ salinity, the two Brassica species showed clear differences in the changes in relative growth rate (RGR), net assimilation rate (NAR) and leaf area ratio (LAR). The RGR of B. napus was slightly reduced by salinity, wheareas the RGR of B. carinata was largely reduced in the early stages of salinization. LAR of B. napus was affected by salinity in the later stages of growth and significantly correlated with the reduction in RGR. On the other hand, the NAR of B. carinata was decreased by salinity, corresponding to the decrease of the RGR of B. carinata. The NAR of B. napus was not significantly affected by salinity according to analysis of covariance. The shoot concentrations of Na, Mg and Cl increased while the concentrations of K and Ca decreased sharply during the first 5 days of salinization; subsequently, all ion concentrations remained relatively constant. The concentrations of Na, K, Ca, Mg and Cl in the root were similarly affected by salinity. There were no significant differences of ion concentrations between species that could be related to the differences in salt tolerance. Thus, the differences in salt tolerance between species can not be related to differences in specific-ion effects, but may be related to some factor that reduces the NAR of B. carinata during the early stages of growth.     

Identification of proteins associated with ion homeostasis and salt tolerance in barley

Identification and characterization of proteins involved in salt tolerance are imperative for revealing its genetic mechanisms. In this study, ionic and proteomic responses of a Tibetan wild barley XZ16 and a well‐known salt‐tolerant barley cv. CM72 were analyzed using inductively coupled plasma‐optical emission spectrometer, 2DE, and MALDI‐TOF/TOF MS techniques to determine salt‐induced differences in element and protein profiles between the two genotypes. In total, 41 differentially expressed proteins were identified in roots and leaves, and they were associated with ion homeostasis, cell redox homeostasis, metabolic process, and photosynthesis. Under salinity stress, calmodulin, Na/K transporters, and H⁺‐ATPases were involved in establishment of ion homeostasis for barley plants. Moreover, ribulose‐1,5‐bisphosphate carboxylase/oxygenase activase and oxygen‐evolving enhancer proteins were significantly upregulated under salinity stress, indicating the great impact of salinity on photosynthesis. In comparison with CM72, XZ16 had greater relative dry weight and lower Na accumulation in the shoots under salinity stress. A higher expression of HvNHX1 in the roots, and some specific proteins responsible for ion homeostasis and cell redox homeostasis, was also found in XZ16 exposed to salt stress. The current results showed that Tibetan wild barley XZ16 and cultivated barley cultivar CM72 differ in the mechanism of salt tolerance.     

Interactive effects of boron and salinity stress on the growth,membrane permeability and mineral composition of tomato and cucumber plants

A greenhouse study was conducted in order to determine interactive effects of NaCl salinity and B on the growth, sodium (Na), chloride (Cl), boron (B), potassium (K) concentrations and membrane permeability of salt resistant Tomato (Lycopersicon esculentum L. cv. Lale F₁) and salt sensitive cucumber (Cucumis sativus L. cv. Santana F₁) plants. Plants were grown in a factorial combination of NaCl (0 and 30 mM for cucumber and 0 and 40 mM for tomato) and B (0, 5, 10 and 20 mg kg^–1 soil). Boron toxicity symptoms appeared at 5 mg kg^–1 B treatments in both plants. Salinity caused an increase in leaf injury due to B toxicity, but it was more severe in cucumber. Dry weights of the plants decreased with the increasing levels of applied B in nonsaline conditions, but the decrease in dry weights due to B toxicity was more pronounced in saline conditions especially in cucumber. Salinity × B interaction on the concentration of B in both plants was found significant. However, increase in B concentrations of tomato decreased under saline conditions when compared to nonsaline conditions. Contrary to this, B concentration of cucumber increased as a result of increasing levels of applied B and salinity. Salinity increased Na and Cl concentrations of both plants.Potassium concentration of tomato was not affected by salinity and B treatments, but K concentration of cucumber was decreased by salinity. Membrane permeability of the plants was increased by salinity while toxic levels of B had no effect on membrane permeability in nonsaline conditions. Membrane permeability was significantly increased in the presence of salinity by the increasing levels of applied B.     

Growth and ion relations in response to combined salinity and waterlogging in the perennial forage legumes Lotus corniculatus and Lotus tenuis

Lotus tenuis (Wadst. & Kit.) is a perennial legume widely grown for pasture in the flood-prone and salt affected Pampa region of Argentina. The physiology of salt and waterlogging tolerance in L. tenuis (four cultivars) was evaluated, and compared with Lotus corniculatus (three cultivars); the most widely cultivated Lotus species. Overall, L. tenuis cultivars accumulated less Na⁺ and Cl⁻, and more K⁺ in shoots than L. corniculatus cultivars, when exposed to 200 mM NaCl for 28 days in aerated or in stagnant solutions. Root porosity was higher in L. tenuis cultivars due to greater aerenchyma formation. In a NaCl dose–response experiment (0–400 mM NaCl in aerated solution), L. tenuis (cv. Chaja) accumulated half as much Cl⁻ in its shoots than L. corniculatus (cv. San Gabriel) at all external NaCl concentrations, and about 30% less shoot Na⁺ in treatments above 250 mM NaCl. Ion distributions in shoots were determined for plants at 200 mM NaCl. L. tenuis (cv. Chaja) again accumulated about half as much Cl⁻ in old leaves, young leaves and stems, compared with concentrations in L. corniculatus (cv. San Gabriel). There were not, however, significant differences between the two species for Na⁺ concentrations in the various shoot tissues. The higher root porosity, and maintenance of lower shoot Cl⁻ and Na⁺ concentrations in L. tenuis, compared with L. corniculatus, contributes to the greater tolerance to combined salt and waterlogging stress in L. tenuis. Moreover, significant variation for tolerance to combined salinity and waterlogging stress was identified within both L. tenuis and L. corniculatus.     

Physiological and biochemical responses of tomato microshoots to induced salinity stress with associated ethylene accumulation

Physiological and biochemical responses of open-pollinated ‘Roma’ and dwarf F1 hybrid ‘Patio’ tomato (Lycopersicon esculentum Mill.) cultivars to in vitro induced salinity were examined in light of the possible contribution of ethylene to these symptoms. Salinity was induced by incorporating 0 (control), 50, 100, 150, or 200 mM NaCl into shoot culture media. Elevated salinity treatments significantly enhanced ethylene accumulation in the headspace and were accompanied by increased leaf epinasty in both cultivars. Growth, leaf cell sap osmolarity, leaf tissue viability and shoot soluble protein content were generally depressed with elevated salinity treatments, whereas electrolyte leakage, membrane injury, raffinose, and total sugars were concomitantly increased. Macronutrients N, P, K, Ca, Mg, and S decreased with elevated salinity in both cultivars and were accompanied by a significant increase in Na content and a sharp decrease in K/Na ratio. Tissue micronutrients, Fe, B, Zn, Mn, and Cu were generally decreased with elevated salinity especially at 100 mM or more. Incorporating ethylene inhibitors CoCl₂ or NiCl₂ at 5.0 or 10.0 mg/l into media supplemented with 100 mM NaCl significantly reduced ethylene accumulation in the headspace and prevented epinasty, but did not eliminate the negative impacts on growth and other physiological parameters caused by salinity treatment in either cultivar. Our results indicate that the increase in ethylene under salinity stress is not the primary factor contributing to salinity’s deleterious effect on tomato plant growth and physiology.     

江苏野生大豆的耐盐性和离子在体内的分布及选择性运输

以相对发芽率和出苗率为指标比较了3个野生大豆(Glycine soja)种群的耐盐性,测定了NaCl胁迫下2个耐盐性不同的野生大豆种群(江苏野生大豆,JWS,耐盐;N23232,盐敏感)植株根、茎和叶片中Na^+、K^+和Cl^-含量的变化。结果表明,JWS的耐盐性最强,盐胁迫抑制野生大豆幼苗生长,使其干物质积累量减少,根冠比上升,对耐盐性弱的N23232抑制作用大于耐盐性强的JWS,不同器官离子含量测定结果表明,盐胁迫下野生大豆茎部Na^+含量最高,耐盐的JWS根系具有积累Na^+和Cl^-的能力,叶片Na^+、Cl含量较低,而盐敏感种群N23232根系中：Na^+、Cl^-含量低于耐盐种群JWS,叶片中Na^+、Cl^-含量则高于JWS,JWS根系对K^+、Na^+吸收的选择性(selectivity ratio,SK,Na)和N23232没有明显差异;但叶片和茎运输的SK,Na明显高于N23232,使地上部K^+／Na^+较高,因此认为野生大豆根系对Na^+、Cl^-的积累及K^+向地上部运输的选择性高是其耐盐性强的主要原因。     

盐胁迫下柚和橘幼苗体内矿质元素变化的比较研究

采用沙培法,对盐胁迫下坪山柚和福橘幼苗体内矿质元素的变化进行了研究。结果表明,随着NaCl浓度的增加,坪山柚和福橘幼苗根部及地上部Na^＋、Cl-含量增加,且相同浓度下,福橘比坪山柚高。40mmol／L NaCI胁迫下,坪山柚和福橘幼苗地上部的K^＋、Fe含量,根部的Ca^2＋、Mg^2＋、Zn含量显著下降,而根部Fe含量及地上部Zn含量显著增加。随NaCl浓度增大,坪山柚根部K^＋含量,地上部Ca^2＋、Mg^2＋含量变化不明显,而福橘根部、地上部上述离子含量在NaCl浓度≥160mmol／L时均显著下降。因此,根部K^＋含量,地上部Ca^2＋、Mg^2＋含量存在品种问差异,或许可作为耐盐性鉴定指标。NaCl胁迫降低坪山柚和福橘幼苗根部及地上部P、Mn含量,而Cu含量在较高浓度NaCl胁迫下显著增加。NaCl胁迫明显降低坪山柚和福橘幼苗地上部K^＋／Na^＋、Ca^2＋／Na^＋和Mg^2＋／Na^＋值,其中K^＋／Na^＋值的变化可考虑作为柑橘耐盐性鉴定的指标。     

Salinity tolerance of normal-fruited and cherry tomato cultivars

The salinity tolerances (NaCl) of 8 normal-fruited tomato cultivars (Lycopersicon esculentum Mill.) and 4 cherry tomato cultivars (L. esculentum var.cerasiforme) were determined by yield-substrate EC response curves, according to the Mass-Hoffman model, modified by van Genuchten and Hoffman (1984). The same model was used to determine the response curves of leaf dry-weight, stem dry-weight, and plant height against substrate EC and also between yield and leaf concentrations of Cl- and Na ions.According to the salinity-threshold (maximum EC-value without yield reduction) and slope (yield decrease per unit EC increase) parameters, determined from the yield-EC response curves, the cherry tomato cultivars were more salt-tolerant than the normal-fruited ones. However, on the basis of vegetative growth characters-EC response curves, cherry tomato cultivars and normal-fruited ones were similarly affected by NaCl.The ranking of the cultivars by their salinity tolerance, determined from the plots of yield vs. leaf concentrations of Cl- and Na ions, was the same as that evaluated from the yield vs. substrate EC plots.     

Overexpression of SlSOS2 (SlCIPK24) confers salt tolerance to transgenic tomato

The Ca(2+)-dependent SOS pathway has emerged as a key mechanism in the homeostasis of Na(+) and K(+) under saline conditions. We have identified and functionally characterized the gene encoding the calcineurin-interacting protein kinase of the SOS pathway in tomato, SlSOS2. On the basis of protein sequence similarity and complementation studies in yeast and Arabidopsis, it can be concluded that SlSOS2 is the functional tomato homolog of Arabidopsis AtSOS2 and that SlSOS2 operates in a tomato SOS signal transduction pathway. The biotechnological potential of SlSOS2 to provide salt tolerance was evaluated by gene overexpression in tomato (Solanum lycopersicum L. cv. MicroTom). The better salt tolerance of transgenic plants relative to non-transformed tomato was shown by their faster relative growth rate, earlier flowering and higher fruit production when grown with NaCl. The increased salinity tolerance of SlSOS2-overexpressing plants was associated with higher sodium content in stems and leaves and with the induction and up-regulation of the plasma membrane Na(+)/H(+) (SlSOS1) and endosomal-vacuolar K(+), Na(+)/H(+) (LeNHX2 and LeNHX4) antiporters, responsible for Na(+) extrusion out of the root, active loading of Na(+) into the xylem, and Na(+) and K(+) compartmentalization.     

Growth and mineral nutrition of six rapid-cycling Brassica species in response to seawater salinity

The growth of six rapid-cycling lines of Brassica species, B. napus, B. campestris, B. nigra, B. juncea, B. oleracea and B. carinata was inhibited by seawater salinity. Based on the change in dry matter reduction relative to the control at varying concentrations of salts (4, 8 and 12 dS m^-1), the relative salt tolerance of these species was evaluated. B. napus and B. carinata were the most tolerant and most sensitive species, respectively, while the other four species were moderately tolerant. The influence of seawater on the concentrations of 12 elements including macronutrients and micronutrients in the shoots of these Brassica plants was characterized to determine the relationship between nutritional disturbance and relative salt tolerance. It was found that seawater salinity had a significant effect on the concentrations of Ca, Mg, K, Cl, Na and total N in the shoots of these plants but only the change in Ca concentration was significantly related to the relative salt tolerance of these six rapid-cycling Brassica species according to a rank analysis of the data. This finding indicates that Ca may play a regulatory role in the responses of Brassica species to saline conditions.     

Additive effects of Na+ and Cl- ions on barley growth under salinity stress

Soil salinity affects large areas of the world's cultivated land, causing significant reductions in crop yield. Despite the fact that most plants accumulate both sodium (Na(+)) and chloride (Cl(-)) ions in high concentrations in their shoot tissues when grown in saline soils, most research on salt tolerance in annual plants has focused on the toxic effects of Na(+) accumulation. It has previously been suggested that Cl(-) toxicity may also be an important cause of growth reduction in barley plants. Here, the extent to which specific ion toxicities of Na(+) and Cl(-) reduce the growth of barley grown in saline soils is shown under varying salinity treatments using four barley genotypes differing in their salt tolerance in solution and soil-based systems. High Na(+), Cl(-), and NaCl separately reduced the growth of barley, however, the reductions in growth and photosynthesis were greatest under NaCl stress and were mainly additive of the effects of Na(+) and Cl(-) stress. The results demonstrated that Na(+) and Cl(-) exclusion among barley genotypes are independent mechanisms and different genotypes expressed different combinations of the two mechanisms. High concentrations of Na(+) reduced K(+) and Ca(2+) uptake and reduced photosynthesis mainly by reducing stomatal conductance. By comparison, high Cl(-) concentration reduced photosynthetic capacity due to non-stomatal effects: there was chlorophyll degradation, and a reduction in the actual quantum yield of PSII electron transport which was associated with both photochemical quenching and the efficiency of excitation energy capture. The results also showed that there are fundamental differences in salinity responses between soil and solution culture, and that the importance of the different mechanisms of salt damage varies according to the system under which the plants were grown.