Physiological adaptive mechanisms of plants grown in saline soil and implications for sustainable saline agriculture in coastal zone

There is large area of saline abandoned and low-yielding land distributed in coastal zone in the world. Soil salinity which inhibits plant growth and decreases crop yield is a serious and chronic problem for agricultural production. Improving plant salt tolerance is a feasible way to solve this problem. Plant physiological and biochemical responses under salinity stress become a hot issue at present, because it can provide insights into how plants may be modified to become more tolerant. It is generally known that the negative effects of soil salinity on plants are ascribed to ion toxicity, oxidative stress and osmotic stress, and great progress has been made in the study on molecular and physiological mechanisms of plant salinity tolerance in recent years. However, the present knowledge is not easily applied in the agronomy research under field environment. In this review, we simplified the physiological adaptive mechanisms in plants grown in saline soil and put forward a practical procedure for discerning physiological status and responses. In our opinion, this procedure consists of two steps. First, negative effects of salt stress are evaluated by the changes in biomass, crop yield and photosynthesis. Second, the underlying reasons are analyzed from osmotic regulation, antioxidant response and ion homeostasis. Photosynthesis is a good indicator of the harmful effects of saline soil on plants because of its close relation with crop yield and high sensitivity to environmental stress. Particularly, chlorophyll a fluorescence transient has been accepted as a reliable, sensitive and convenient tool in photosynthesis research in recent years, and it can facilitate and enrich photosynthetic research under field environment.     

Current developments in arbuscular mycorrhizal fungi research and its role in salinity stress alleviation: a biotechnological perspective

Arbuscular mycorrhizal fungi (AMF) form widespread symbiotic associations with 80% of known land plants. They play a major role in plant nutrition, growth, water absorption, nutrient cycling and protection from pathogens, and as a result, contribute to ecosystem processes. Salinity stress conditions undoubtedly limit plant productivity and, therefore, the role of AMF as a biological tool for improving plant salt stress tolerance, is gaining economic importance worldwide. However, this approach requires a better understanding of how plants and AMF intimately interact with each other in saline environments and how this interaction leads to physiological changes in plants. This knowledge is important to develop sustainable strategies for successful utilization of AMF to improve plant health under a variety of stress conditions. Recent advances in the field of molecular biology, “omics” technology and advanced microscopy can provide new insight about these mechanisms of interaction between AMF and plants, as well as other microbes. This review mainly discusses the effect of salinity on AMF and plants, and role of AMF in alleviation of salinity stress including insight on methods for AMF identification. The focus remains on latest advancements in mycorrhizal research that can potentially offer an integrative understanding of the role of AMF in salinity tolerance and sustainable crop production.     

Endophytic Bacteria in Plant Salt Stress Tolerance: Current and Future Prospects

Soil salinity is a major limiting factor for crop productivity worldwide and is continuously increasing owing to climate change. A wide range of studies and practices have been performed to induce salt tolerance mechanisms in plants, but their result in crop improvement has been limited due to lack of time and money. In the current scenario, there is increasing attention towards habitat-imposed plant stress tolerance driven by plant-associated microbes, either rhizospheric and/or endophytic. These microbes play a key role in protecting plants against various environmental stresses. Therefore, the use of plant growth-promoting microbes in agriculture is a low-cost and eco-friendly technology to enhance crop productivity in saline areas. In the present review, the authors describe the functionality of endophytic bacteria and their modes of action to enhance salinity tolerance in plants, with special reference to osmotic and ionic stress management. There is concrete evidence that endophytic bacteria serve host functions, such as improving osmolytes, anti-oxidant and phytohormonal signaling and enhancing plant nutrient uptake efficiency. More research on endophytes has enabled us to gain insights into the mechanism of colonization and their interactions with plants. With this information in mind, the authors tried to solve the following questions: (1) how do benign endophytes ameliorate salt stress in plants? (2) What type of physiological changes incur in plants under salt stress conditions? And (3), what type of determinants produced by endophytes will be helpful in plant growth promotion under salt stress?

     

The battle of two ions: Ca2+ signalling against Na+ stress

Soil salinity adversely affects plant growth, crop yield and the composition of ecosystems. Salinity stress impacts plants by combined effects of Na⁺ toxicity and osmotic perturbation. Plants have evolved elaborate mechanisms to counteract the detrimental consequences of salinity. Here we reflect on recent advances in our understanding of plant salt tolerance mechanisms. We discuss the embedding of the salt tolerance‐mediating SOS pathway in plant hormonal and developmental adaptation. Moreover, we review newly accumulating evidence indicating a crucial role of a transpiration‐dependent salinity tolerance pathway, that is centred around the function of the NADPH oxidase RBOHF and its role in endodermal and Casparian strip differentiation. Together, these data suggest a unifying and coordinating role for Ca²⁺ signalling in combating salinity stress at the cellular and organismal level.     

Salinity Adaptation and the Contribution of Parental Environmental Effects in Medicago truncatula

High soil salinity negatively influences plant growth and yield. Some taxa have evolved mechanisms for avoiding or tolerating elevated soil salinity, which can be modulated by the environment experienced by parents or offspring. We tested the contribution of the parental and offspring environments on salinity adaptation and their potential underlying mechanisms. In a two-generation greenhouse experiment, we factorially manipulated salinity concentrations for genotypes of Medicago truncatula that were originally collected from natural populations that differed in soil salinity. To compare population level adaptation to soil salinity and to test the potential mechanisms involved we measured two aspects of plant performance, reproduction and vegetative biomass, and phenological and physiological traits associated with salinity avoidance and tolerance. Saline-origin populations had greater biomass and reproduction under saline conditions than non-saline populations, consistent with local adaptation to saline soils. Additionally, parental environmental exposure to salt increased this difference in performance. In terms of environmental effects on mechanisms of salinity adaptation, parental exposure to salt spurred phenological differences that facilitated salt avoidance, while offspring exposure to salt resulted in traits associated with greater salt tolerance. Non-saline origin populations expressed traits associated with greater growth in the absence of salt while, for saline adapted populations, the ability to maintain greater performance in saline environments was also associated with lower growth potential in the absence of salt. Plastic responses induced by parental and offspring environments in phenology, leaf traits, and gas exchange contribute to salinity adaptation in M. truncatula. The ability of plants to tolerate environmental stress, such as high soil salinity, is likely modulated by a combination of parental effects and within-generation phenotypic plasticity, which are likely to vary in populations from contrasting environments.     

Expression of an Arabidopsis vacuolar sodium/proton antiporter gene in cotton improves photosynthetic performance under salt conditions and increases fiber yield in the field

Drought and salinity are two major limiting factors in crop productivity. One way to reduce crop loss caused by drought and salinity is to increase the solute concentration in the vacuoles of plant cells. The accumulation of sodium ions inside the vacuoles provides a 2-fold advantage: (i) reducing the toxic levels of sodium in cytosol; and (ii) increasing the vacuolar osmotic potential with the concomitant generation of a more negative water potential that favors water uptake by the cell and better tissue water retention under high soil salinity. The success of this approach was demonstrated in several plants, where the overexpression of the Arabidopsis gene AtNHX1 that encodes a vacuolar sodium/proton antiporter resulted in higher plant salt tolerance. Overexpression of AtNHX1 increases sodium uptake in vacuoles, which leads to increased vacuolar solute concentration and therefore higher salt tolerance in transgenic plants. In an effort to engineer cotton for higher drought and salt tolerance, we created transgenic cotton plants expressing AtNHX1. These AtNHX1-expressing cotton plants generated more biomass and produced more fibers when grown in the presence of 200 mM NaCl in greenhouse conditions. The increased fiber yield was probably due to better photosynthetic performance and higher nitrogen assimilation rates observed in the AtNHX1-expressing cotton plants as compared with wild-type cotton plants under saline conditions. Furthermore, the field-grown AtNHX1-expressing cotton plants produced more fibers with better quality, indicating that AtNHX1 can indeed be used for improving salt stress tolerance in cotton.     

Role of Triacontanol in Counteracting the Ill Effects of Salinity in Plants: A Review

Soil salinity is one among the common environmental threats to agriculture. It adversely affects the physio-biochemical processes of plants that eventually lead to the reduction in growth, development and crop productivity. To cope with such adverse conditions, plants develop certain internal mechanisms, but under severe conditions these mechanisms fail to tolerate the salt stress. To overcome this problem, various strategies have been employed that help plants to mitigate salinity effects. Among the various strategies, the application of plant growth regulators (PGRs) has gained significant attention to induce salt tolerance in plants. A number of PGRs have been used so far. Among these, triacontanol (TRIA), a new PGR is gaining a lot of importance to enhance the plant growth, productivity and salinity tolerance in different crops. The utility of TRIA is dependent on its applied concentration. Its lower concentrations generally alleviate the salinity effects. However, the knowledge of its biosynthesis, signalling and its role particularly to mitigate salinity effect remains scanty. In the present article, the focus has been given on the role of exogenous applications of TRIA in the regulation of physio-biochemical characteristics especially plant growth, photosynthesis, nutrient acquisition, oxidative stress, antioxidant systems, compatible solutes, yield attributes and its mode of action in plants under salinity conditions. The salient features of the review may provide new insights on the role of TRIA in countering the ill effect of salinity in different crop plants.

     

Comparative roles of brassinosteroids and polyamines in salt stress tolerance

Salt stress is among the major abiotic stresses that adversely affect the global crop production and its adverse impacts are getting more serious in the regions where saline water is used for irrigation. It induces reactive oxygen species, alters the activity of antioxidant system and adversely affects the process of photosynthesis. Various strategies have been employed to mitigate the deleterious effects of salt stress. Presently, the recommended strategies to overcome the adverse effects of salt stress include the use of tolerant cultivars, ameliorative water management and diverse cultural practices. However, none of these approaches have been found to be fully effective under salt stress conditions. An alternative and technically simpler approach to induce salt stress tolerance is the exogenous application of plant growth regulators (PGRs). This technique has gained significant importance during the past decade. PGRs have been implicated to regulate a wide range of metabolic and physiological activities in plants, ranging from cell division and organogenesis to protection against biotic and abiotic stresses. One of the important factors for enhanced plant productivity by PGRs is their efficiency to overcome the salt-induced stress conditions. Recent findings on the effects of brassinosteroids and polyamines on the salt stress tolerance of crops open new avenues to address the salinity problems. This review enlightens the role of brassinosteroids and polyamines in different plant processes like their role in regulation of photosynthesis, antioxidant systems and other related aspects, thereby improving overall performance of plants.     

Crop production and management under saline conditions

Summary This review evaluates management practices that may minimize yield reduction under saline conditions according to three strategies: (I) control of root-zone salinity; (II) reduced damage to the crop; (III) reduced damage to individual plants. Plant response to salinity is described by an unchanged yield up to a threshold soil salinity (a), then a linear reduction in relative yield (b), to a maximum soil salinity that corresponds to zero yield (Y_o). Strategies I and II do not take into consideration any change in the parameters of the response curve, while strategy III is aimed at modifying them.Control of root zone salinity is obtained by irrigation and leaching. From the review of existing data it is concluded that the effective soil salinity parameter should be taken as the mean electrical conductivity of the saturated paste extract or of the soil solution over time and space. Several irrigation and leaching practices are discussed. It is shown that intermittent leaching is more advantageous than leaching at each irrigation. Specific cultivation and irrigation practices that result in soil salinity reduction adjacent to young seedlings and the use of water of low salinity at specifically sensitive growth stages may be highly beneficial. Recent data do not show that reduced irrigation intervals improve crop response more under saline than under nonsaline irrigation. Alternate use of water of different salt concentrations results in mixing in the soil and the crop responds to the mean water salinity.Reduced damage at the fiel level when soil or irrigation water salinity is too high to maintain full yield of single plants requires a larger crop stand. For row crops reduced inter-row spacing is more effective than reduced intra-row spacing.Reduced damage at the plant level while the salinity tolerance of the plants remains constant shows up in the response curve parameters as larger threshold and slope and constant salinity at zero yield. This is the effect of a reduced atmospheric water demand that results in reduced stress in the plant under given salinity. Management can also change the salt tolerance of the crop. This will show up as higher salinity at zero yield, as well as changes in threshold and slope. Such changes in the response curve were found at different growth stages, under different atmospheric CO₂, under different fertilization, and when sprinkler irrigation was compared with drip irrigation.Contribution from the Agricultural Research Organization, The Volcani Center, Bet Dagan, Israel. No. 1111-E 1984 series.     

Improving salinity tolerance in crop plants: a biotechnological view

Salinity limits the production capabilities of agricultural soils in large areas of the world. Both breeding and screening germplasm for salt tolerance encounter the following limitations: (a) different phenotypic responses of plants at different growth stages, (b) different physiological mechanisms, (c) complicated genotype × environment interactions, and (d) variability of the salt-affected field in its chemical and physical soil composition. Plant molecular and physiological traits provide the bases for efficient germplasm screening procedures through traditional breeding, molecular breeding, and transgenic approaches. However, the quantitative nature of salinity stress tolerance and the problems associated with developing appropriate and replicable testing environments make it difficult to distinguish salt-tolerant lines from sensitive lines. In order to develop more efficient screening procedures for germplasm evaluation and improvement of salt tolerance, implementation of a rapid and reliable screening procedure is essential. Field selection for salinity tolerance is a laborious task; therefore, plant breeders are seeking reliable ways to assess the salt tolerance of plant germplasm. Salt tolerance in several plant species may operate at the cellular level, and glycophytes are believed to have special cellular mechanisms for salt tolerance. Ion exclusion, ion sequestration, osmotic adjustment, macromolecule protection, and membrane transport system adaptation to saline environments are important strategies that may confer salt tolerance to plants. Cell and tissue culture techniques have been used to obtain salt tolerant plants employing two in vitro culture approaches. The first approach is selection of mutant cell lines from cultured cells and plant regeneration from such cells (somaclones). In vitro screening of plant germplasm for salt tolerance is the second approach, and a successful employment of this method in durum wheat is presented here. Doubled haploid lines derived from pollen culture of F₁ hybrids of salt-tolerant parents are promising tools to further improve salt tolerance of plant cultivars. Enhancement of resistance against both hyper-osmotic stress and ion toxicity may also be achieved via molecular breeding of salt-tolerant plants using either molecular markers or genetic engineering.     

Halophytes--an emerging trend in phytoremediation

Halophytic plants are of special interest because these plants are naturally present in environments characterized by an excess of toxic ions, mainly sodium and chloride. Several studies have revealed that these plants may also tolerate other stresses including heavy metals based on the findings that tolerance to salt and to heavy metals may, at least partly, rely on common physiological mechanisms. In addition, it has been shown that salt-tolerant plants may also be able to accumulate metals. Therefore, halophytes have been suggested to be naturally better adapted to cope with environmental stresses, including heavy metals compared to salt-sensitive crop plants commonly chosen for phytoextraction purposes. Thus, potentially halophytes are ideal candidates for phytoextraction orphytostabilization of heavy metal polluted soils and moreover of heavy metal polluted soils affected by salinity. Some halophytes use excretion processes in order to remove the excess of salt ions from their sensitive tissues and in some cases these glandular structures are not always specific to Na+ and Cl- and other toxic elements such as cadmium, zinc, lead, or copper are accumulated and excreted by salt glands or trichomes on the surface of the leaves--a novel phytoremediation process called "phytoexcretion". Finally, the use of halophytes has also been proposed for soil desalination through salt accumulation in the plant tissue or dissolution of soil calcite in the rhizosphere to provide Ca2+ that can be exchanged with Na+ at cation exchange sites.     

Recent developments in understanding salinity tolerance

Salt stress imposes a major environmental threat to agriculture and its adverse impacts are getting more serious problem in regions where saline water is used for irrigation. Therefore, the efforts to increase salt tolerance of crop plants bear remarkable importance to supply sustainable agriculture on marginal lands and could potentially improve crop yield overall. Acclimation of plants to salinized conditions depend upon activation of cascades of molecular networks involved in stress sensing, signal transduction and the expression of specific stress-related genes and metabolites. Adaptational processes are elaborate and more than one gene might be expressed during the acclimation process. Isolation of Salt Overly Sensitive (SOS) genes by sos mutants shed us light on the relationship between ion homeostasis and salinity tolerance. The essential role of antioxidative system to maintain a balance between the overproduction of Reactive Oxygen Species (ROS) and their scavenging to keep them at signaling level for reinstating metabolic homeostasis has already been established. Compatible osmolytes synthesized to maintain equal water potential with the environment under salinity conditions implements another strategy to develop resistance against salinity. With the growing body of information about molecular markers, genomics and post-genomics and thus increasing understanding of signaling pathways and mechanisms that contributes to plant stress responses, significant breakthroughs have been emerged to figure out the mechanism and control of salinity tolerance at molecular level. Many transgenic works were carried out to produce transgenic plants to develop enhanced tolerance to salt stress. However, a few of them seem succeeded to be implemented in salt-affected marginal lands efficiently. This minireview focuses on the recent developments in salinity tolerance research aiming to contribute sustainable food production under salt stress in the face of a globally warming ecosystem.     

Mechanisms of sodium uptake by roots of higher plants

The negative impact of soil salinity on agricultural yields is significant. For agricultural plants, sensitivity to salinity is commonly (but not exclusively) due to the abundance of Na⁺ in the soil as excess Na⁺ is toxic to plants. We consider reducing Na⁺ uptake to be the key, as well as the most efficient approach, to control Na⁺ accumulation in crop plants and hence to improve their salt resistance. Understanding the mechanism of Na⁺ uptake by the roots of higher plants is crucial for manipulating salt resistance. Hence, the aim of this review is to highlight and discuss recent advances in our understanding of the mechanisms of Na⁺ uptake by plant roots at both physiological and molecular levels. We conclude that continued efforts to investigate the mechanisms of root Na⁺ uptake in higher plants are necessary, especially that of low-affinity Na⁺ uptake, as it is the means by which sodium enters into plants growing in saline soils.     

Variation from plant tissue cultures: biotechnological application to improving salinity tolerance

Breeding for salt tolerance in crop plants is envisaged as one way to combat a worldwide problem of increasing soil salinity in agricultural land. Tissue culture techniques may prove valuable as a means of achieving this goal. In this review, reports of the selection and characterization of plant cell cultures tolerant to excess salt are assessed, in the context of variability from tissue culture and the significance of cellular physiological adaptation to salinity. The examples of plant regeneration from salt grown cell cultures are also outlined, with emphasis on correlation to the effect of salt on cell cultures, genetic variability for salt tolerance in vitro, and the value of regenerates in the development of salt tolerant plants.     

蕾期土壤盐度降低后棉花叶片的生理功能恢复

试验于2011—2012年在江苏南京江苏省农业科学院经济作物研究所试验田进行,采用盆栽方法,以鲁棉研37号和苏棉22号为供试材料,设置土壤盐度降低试验(初始土壤含盐量为0.2%,棉花进入二叶期后每7d加入混合盐1次,每次增加0.1%,使土壤含盐量逐渐达到0.5%,蕾期进行盐度降低处理,使土壤含盐量降低到0.2%左右),研究蕾期土壤盐度降低后棉花叶片的生理代谢动态特征。结果表明:土壤盐度降低后,棉花叶片叶绿素(Chl)、类胡萝卜素(Car)含量和Chl/Car升高;净光合速率和气孔导度升高,且分别在土壤盐度降低后第14天和7天接近于低盐对照;土壤盐度降低后棉花叶片超氧化物歧化酶(SOD)和过氧化物酶(POD)活性升高,过氧化氢酶(CAT)活性和丙二醛(MDA)含量降低,MDA含量在土壤盐度降低后第14天接近于低盐对照;土壤盐度降低后棉花叶片中可溶性糖、游离氨基酸和脯氨酸含量降低,且接近于低盐对照。上述结果表明土壤盐度降低后,棉花叶片生理功能逐渐恢复,进而实现棉花生长发育的恢复补偿。棉花叶片生理功能在土壤盐度降低后的恢复能力存在品种间差异,鲁棉研37号较苏棉22号叶片生理功能表现出更强的恢复能力。     

植物耐盐性机理研究进展

在盐胁迫下环境中某些植物会在发生一些变化。从生理学、生物化学、盐胁迫分子学机制的角度对植物对盐胁迫的反应研究进行了回顾,并提供了一些目前知识水平上能增加植物盐耐性的方法。解释了在盐胁迫下植物的离子吸收、相溶性物质、抗氧化酶、植物激素、光合作用等方面的变化规律,其中也有耐盐植物功能调节的研究,这有助于从多学科研究的角度评估盐胁迫的生态重要性。     

A novel in vitro tissue culture approach to study salt stress responses in citrus

In citrus, a major crop throughout the world, growth and yield are seriously affected by salinity. Different approaches, including agronomical, physiological and molecular methods, have been used to address this problem. In this work, an in vitro experimental system has been developed to study the toxic effect of NaCl on three citrus genotypes, avoiding the ion filter that represents the root system. To carry out the experiments, shoots were obtained from nodal segments of Cleopatra mandarin, Carrizo citrange and citrumelo CPB4475 plants growing in a greenhouse. Shoots were cultured in control or NaCl-supplemented media. After testing several salt concentrations, 60 mM NaCl was selected as saline treatment. Shoots accumulated similar levels of chloride when cultured without roots and exhibited similar leaf damage. No increases in malondialdehyde levels were observed in any genotype (as a measure of oxidative stress). Similar patterns of hormonal signalling (in terms of abscisic acid and salicylic acid contents) were exhibited in the three genotypes, despite their different tolerance under field conditions. All data together indicate that, without root system, all genotypes had the same behaviour under salt stress. The in vitro culture system has been proved as a useful tool to study biochemical processes involved in the response of citrus to salt stress.     

Plant Growth-Promoting Rhizobacteria-Mediated Adaptive Responses of Plants Under Salinity Stress

In this recent era, several approaches have been developed to alleviate the adverse effects of salinity stress in different plants. However, some of them are not eco-friendly. In this context, evolving sustainable approaches which enhance the productivity of saline soil without harming the environment are necessary. Many recent studies showed that plant growth-promoting rhizobacteria (PGPR) are known to confer salinity tolerance to plants. Salt-stressed plants inoculated with PGPR enhance the growth and productivity of crops by reducing oxidative damage, maintaining ionic homeostasis, enhancing antioxidant machinery, and regulating gene expressions. The PGPR also regulates the photosynthetic attributes such as net photosynthetic rate, chlorophyll, and carotenoid contents and enhances the salinity tolerance to plants. Moreover, PGPR has a great role in the enhancement of phytohormones and secondary metabolites synthesis in plants under salt stress. This review summarizes the current reports of the application of PGPR in plants under salt stress and discusses the PGPR-mediated mechanisms in plants of salt tolerance. This review also discusses the potential role of PGPR in cross-talk with phytohormones and secondary metabolites to alleviate salt stress and highlights the research gaps where further research is needed.

     

Salinity and its effects on the functional biology of legumes

Salinity has plagued soil fertility and drastically affected growth and survival of glycophytes in irrigated regions of the world since the beginning of recorded history. It is particularly common in arid and semi-arid areas where evapotranspiration exceeds annual precipitation, and where irrigation is therefore necessary to meet crop water needs. Salt buildup in the soils and groundwater has threatened its productivity and sustainability. Plant responses to salt stress include an array of changes at the molecular, biochemical and physiological levels. Salt stress involves a water deficit induced by the salt concentration in the rhizosphere, resulting in disruption of homeostasis and ion distribution in the cell and denaturation of structural and functional proteins. As a consequence, salinity stress often activates cell signaling pathways including those that lead to synthesis of osmotically active metabolites, specific proteins, and certain free radical scavenging enzymes that control ion and water flux and support scavenging of oxygen radicals or chaperones. ROS detoxification forms an important defense against salt stress. Legumes are a key component of sustainable agriculture and can offer many economic and environmental benefits if grown more widely in crop rotations because of their ability to fix nitrogen in the root nodules in a symbiotic interaction with soil rhizobia. Due to their capacity to grow on nitrogen-poor soils, they can be efficiently used for improving saline soil fertility and help to reintroduce agriculture to these lands. However, in legumes, salt stress imposes a significant limitation of productivity related to the adverse effects on the growth of the host plant, the root-nodule bacteria, symbiotic development and finally the nitrogen fixation capacity. This paper reviews responses of legumes to salinity stress with emphasis on physiological and biochemical mechanisms of salt tolerance.     

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.