The physiological and molecular mechanisms of tolerance to osmotic and ionic components of salinity stress are reviewed at the cellular, organ, and whole-plant level. Plant growth responds to salinity in two phases: a rapid, osmotic phase that inhibits growth of young leaves, and a slower, ionic phase that accelerates senescence of mature leaves. Plant adaptations to salinity are of three distinct types: osmotic stress tolerance, Na + or Cl − exclusion, and the tolerance of tissue to accumulated Na + or Cl − . Our understanding of the role of the HKT gene family in Na + exclusion from leaves is increasing, as is the understanding of the molecular bases for many other transport processes at the cellular level. However, we have a limited molecular understanding of the overall control of Na + accumulation and of osmotic stress tolerance at the whole-plant level. Molecular genetics and functional genomics provide a new opportunity to synthesize molecular and physiological knowledge to improve the salinity tolerance of plants relevant to food production and environmental sustainability.

https://rodas5.us.es/file/c2a700e9-8b4a-1970-d343-4a269c302970/1/estres_salino_bibliografia_scorm.zip/files/estres_salino_bibliografia.pdf

Mechanisms of salinity tolerance

We discuss the physical and enzymatic bases of carbone isotope discrimination during photosynthesis, noting how knowledge of discrimination can be used to provide additional insight into photosynthetic metabolism and the environmental influences on that process

Carbon Isotope Discrimination and Photosynthesis

Plant responses to salt and water stress have much in common. Salinity reduces the ability of plants to take up water, and this quickly causes reductions in growth rate, along with a suite of metabolic changes identical to those caused by water stress. The initial reduction in shoot growth is probably due to hormonal signals generated by the roots. There may be salt-specific effects that later have an impact on growth; if excessive amounts of salt enter the plant, salt will eventually rise to toxic levels in the older transpiring leaves, causing premature senescence, and reduce the photosynthetic leaf area of the plant to a level that cannot sustain growth. These effects take time to develop. Salttolerant plants differ from salt-sensitive ones in having a low rate of Na + and Cl ‐ transport to leaves, and the ability to compartmentalize these ions in vacuoles to prevent their build-up in cytoplasm or cell walls and thus avoid salt toxicity. In order to understand the processes that give rise to tolerance of salt, as distinct from tolerance of osmotic stress, and to identify genes that control the transport of salt across membranes, it is important to avoid treatments that induce cell plasmolysis, and to design experiments that distinguish between tolerance of salt and tolerance of water stress.

https://onlinelibrary.wiley.com/doi/pdf/10.1046/j.0016-8025.2001.00808.x

Comparative physiology of salt and water stress

Salt and drought stress signal transduction consists of ionic and osmotic homeostasis signaling pathways, detoxification (i.e., damage control and repair) response pathways, and pathways for growth regulation. The ionic aspect of salt stress is signaled via the SOS pathway where a calcium-responsive SOS3-SOS2 protein kinase complex controls the expression and activity of ion transporters such as SOS1. Osmotic stress activates several protein kinases including mitogen-activated kinases, which may mediate osmotic homeostasis and/or detoxification responses. A number of phospholipid systems are activated by osmotic stress, generating a diverse array of messenger molecules, some of which may function upstream of the osmotic stress-activated protein kinases. Abscisic acid biosynthesis is regulated by osmotic stress at multiple steps. Both ABA-dependent and -independent osmotic stress signaling first modify constitutively expressed transcription factors, leading to the expression of early response transcriptional activators, which then activate downstream stress tolerance effector genes.

/pdf/salt-and-drought-stress-signal-transduction-in-plants-4bj87ki2ww.pdf

Salt and drought stress signal transduction in plants

▪ Abstract Plant responses to salinity stress are reviewed with emphasis on molecular mechanisms of signal transduction and on the physiological consequences of altered gene expression that affect biochemical reactions downstream of stress sensing. We make extensive use of comparisons with model organisms, halophytic plants, and yeast, which provide a paradigm for many responses to salinity exhibited by stress-sensitive plants. Among biochemical responses, we emphasize osmolyte biosynthesis and function, water flux control, and membrane transport of ions for maintenance and re-establishment of homeostasis. The advances in understanding the effectiveness of stress responses, and distinctions between pathology and adaptive advantage, are increasingly based on transgenic plant and mutant analyses, in particular the analysis of Arabidopsis mutants defective in elements of stress signal transduction pathways. We summarize evidence for plant stress signaling systems, some of which have components analogous to t...

Plant cellular and molecular responses to high salinity.

The hypothesis that ABA produced by roots in drying soil is responsible for stomatal closure was tested with grafted plants constructed from the ABA-deficient tomato mutants, sitiens and flacca and their near-isogenic wild-type parent. Three types of experiments were conducted. In the first type, reciprocal grafts were made between the wild type and sitiens or flacca. Stomatal conductance accorded with the genotype of the shoot, not the root. Stomates closed in all of the grafted plants in response to soil drying, regardless of the root genotype, i.e. regardless of the ability of the roots to produce ABA. In the second type of experiment, wild-type shoots were grafted onto a split-root system consisting of one wild-type root grafted to one mutant (flacca or sitiens) root. Water was withheld from one root system, while the other was watered well so that the shoots did not experience any decline in water potential or loss of turgor. Stomates closed to a similar extent when water was withheld from the mutant roots or the wild-type roots. In the third type of experiment, grafted plants with wild-type shoots and either wild-type or sitiens roots were established in pots that could be placed inside a pressure chamber, and the pressure increased as the soil dried so that the shoots remained fully turgid throughout. Stomates closed as the soil dried, regardless of whether the roots were wild type or sitiens. These experiments demonstrate that stomatal closure in response to soil drying can occur in the absence of leaf water deficit, and does not require ABA production by roots. A chemical signal from roots leading to a change in apoplastic ABA levels in leaves may be responsible for the stomatal closure.

Stomatal control in tomato with ABA‐deficient roots: response of grafted plants to soil drying

To examine the factors that affect tolerance to high internal salt concentrations, two tetraploid wheat genotypes that differ in the degree of salt-induced leaf injury (Wollaroi and Line 455) were grown in 150 mM NaCl for 4 weeks. Shoot biomass of both genotypes was substantially reduced by salinity, but genotypic differences appeared only after 3 weeks, when durum cultivar Wollaroi showed greater leaf injury and a greater reduction in biomass than Line 455. Ion accumulation, water relations, chlorophyll fluorescence and gas exchange were followed on one leaf (leaf 3) throughout its life. Salinity caused a large decrease in stomatal conductance (gs) of both genotypes. This was not due to poor water relations, as leaf turgor of both genotypes was higher in the salt treatment than in controls, so chemical signals were likely to have caused the decrease in gs. Reductions in assimilation rate were initially due to gs and, with time, were due to a combination of stomatal and non-stomatal limitations. The non-stomatal limitations were associated with a build up of Na+ above 250 mM. The efficiency of PSII photochemistry in Line 455 was unaffected throughout. However, in Wollaroi, the potential and actual quantum yield of PSII photochemistry began to decline as the leaf aged and the thermal energy dissipation of excess light energy (NPQ) increased. This coincided with high Na+ and Cl– concentrations in the leaf and with chlorophyll degradation, indicating that these later reductions in CO2 assimilation in Wollaroi were a consequence of a direct toxic ion effect. The earlier reduction in CO2 assimilation and greater leaf injury explain why growth of Wollaroi was less than Line 455. The most sensitive indicator of salinity stress was gs, followed by CO2 assimilation, with fluorescence parameters other than NPQ being no more sensitive than chlorophyll itself.

Factors affecting CO2 assimilation, leaf injury and growth in salt-stressed durum wheat.

In many species, salt sensitivity is associated with the accumulation of sodium (Na(+)) in photosynthetic tissues. Na(+) uptake to leaves involves a series of transport steps and so far very few candidate genes have been implicated in the control of these processes. In this study, Na(+) transport was compared in two varieties of durum wheat (Triticum turgidum) L. subsp. durum known to differ in salt tolerance and Na(+) accumulation; the relatively salt tolerant landrace line 149 and the salt sensitive cultivar Tamaroi. Genetic studies indicated that these genotypes differed at two major loci controlling leaf blade Na(+) accumulation (R. Munns, G.J. Rebetzke, S. Husain, R.A. James, R.A. Hare [2003] Aust J Agric Res 54: 627-635). The physiological traits determined by these genetic differences were investigated using measurements of unidirectional (22)Na(+) transport and net Na(+) accumulation. The major differences in Na(+) transport between the genotypes were (1) the rate of transfer from the root to the shoot (xylem loading), which was much lower in the salt tolerant genotype, and (2) the capacity of the leaf sheath to extract and sequester Na(+) as it entered the leaf. The genotypes did not differ significantly in unidirectional root uptake of Na(+) and there was no evidence for recirculation of Na(+) from shoots to roots. It is likely that xylem loading and leaf sheath sequestration are separate genetic traits that interact to control leaf blade Na(+).

Control of Sodium Transport in Durum Wheat

Cropping on saline land is restricted by the low tolerance of crops to salinity and waterlogging. Prospects for improving salt tolerance in wheat and barley include the use of: (i) intra-specific variation, (ii) variation for salt tolerance in the progenitors of these cereals, (iii) wide-hybridisation with halophytic ‘wild’ relatives (an option for wheat, but not barley), and (iv) transgenic techniques. In this review, key traits contributing to salt tolerance, and sources of variation for these within the Triticeae, are identified and recommendations for use of these traits in screening for salt tolerance are summarised. The potential of the approaches to deliver substantial improvements in salt tolerance is discussed, and the importance of adverse interactions between waterlogging and salinity are emphasised. The potential to develop new crops from the diverse halophytic flora is also considered.

Improving salt tolerance of wheat and barley: future prospects

Durum wheat (AB genomes) is more salt-sensitive than bread wheat (ABD genomes), a feature that restricts its expansion into areas with sodic or saline soils. Salt tolerance in bread wheat is linked with a locus on the D genome that results in low Na+ uptake and enhanced K+/Na+ discrimination. In order to introduce salt tolerance into current durum wheats from sources other than the D genome, a search for genetic variation in salt tolerance was made across a wide range of tetraploids representing 5 Triticum turgidum sub-species (durum, carthlicum, turgidum, turanicum, polonicum). Selections were screened for low Na+ uptake and enhanced K+/Na+ discrimination. This was assessed in seedlings grown in 150 mМ NaCl with supplemental Ca2+, by measuring the Na+ and K+ accumulated in the blade of a given leaf over 10 days. Large and repeatable genetic variation was found. Low Na+ accumulation and high K+/Na+ discrimination of similar magnitude to that of bread wheat was found in the sub-species durum. These selections have the potential for improving salt tolerance in durum wheat breeding programs.

Rana Munns

Papers

Stomatal control in tomato with ABA‐deficient roots: response of grafted plants to soil drying

Factors affecting CO2 assimilation, leaf injury and growth in salt-stressed durum wheat.

Control of Sodium Transport in Durum Wheat

Improving salt tolerance of wheat and barley: future prospects

Genetic variation for improving the salt tolerance of durum wheat