Arguments for the use of physiological criteria for improving the salt tolerance in crops

Efforts to develop new crop varieties with improved salt tolerance have been intensified over the past 15–20 years. Despite the existence of genetic variation for salt tolerance within species, and many methods available for expanding the source of genetic variation, there is only a limited number of varieties that have been developed with improved tolerance. These new varieties have all been based upon selection for agronomic characters such as yield or survival in saline conditions. That is, based upon characters that integrate the various physiological mechanisms responsible for tolerance. Yet over the same time period, knowledge of physiological salt responses has increased substantially.Selection and breeding to increase salt tolerance might be more successful if selection is based directly on the physiological mechanisms or characters conferring tolerance. Basic questions associated with using physiological selection criteria are discussed in the paper. These are centred around the need for genetic variation, the importance of the targeted mechanism, the ease of detection of the physiological mechanism (including the analytical requirements) and the breeding strategy. Many mechanisms, including ion exclusion, ion accumulation, compatible solute production and osmotic adjustment have been associated with genetic variation in salt tolerance. Yet their successful use in improving salt tolerance, via physiological selection criteria, is largely non-existent. Consideration is given to the role of physiological criteria in the short and long term in improving salt tolerance. In several glycophytic species, particularly legumes, physiological selection based on ion exclusion from the shoots shows promise. Recent results for white clover indicate the potential for using a broad physiological selection criterion of restricted Cl accumulation in the shoots, with scope for future refinement based upon the specific physiological characters that combined result in ion exclusion.     

Some mechanisms of salt tolerance in crop plants

Summary In the first part of this review the main features of salt tolerance in higher plants are discussed. The hypothesis of intracellular compartmentation of solutes is used as a basis for models of tolerance mechanisms operating in roots and in leaves. Consideration is given to the implications of the various mechanisms for the yield potential of salt-tolerant crop plants.Some work on the more salt-tolerant members of the Triticeae is then described. The perennial speciesElytrigia juncea andLeymus sabulosus survive prolonged exposure to 250 mol m^–3 NaCl, whereas the annual Triticum species are severely affected at only 100 mol m^–3 NaCl. In the perennial species the tissue ion levels are controlled within narrow limits. In contrast, the more susceptible wheats accumulate far more sodium and chloride than is needed for osmotic adjustment, and the effects of salt stress increase with time of exposure.Two different types of salt tolerance are exhibited in plants capable of growing at high salinities. In succulent Chenopodiaceae, for example, osmotic adjustment is achieved mainly by accumulation of high levels of sodium and chloride in the shoots, accompanied by synthesis of substantial amounts of the compatible solute glycinebetaine. This combination of mechanisms allows high growth rates, in terms of both fresh and dry weight. At the opposite end of the spectrum of salt tolerance responses are the halophytic grasses, which strictly limit the influx of salts into the shoots, but suffer from very much reduced growth rates under saline conditions. Another variation is shown in those species that possess salt glands. The development and exploitation of crop plants for use on saline soils is discussed in relation to the implications of these various mechanisms.     

In vitro selection for salt tolerance in crop plants: Theoretical and practical considerations

Summary In recent years attempts have been made to supplement traditional breeding for the production of salt-tolerant plants with variability existing in cell culture. The potential causes suggested as an explanation for the limited success of the in vitro approach include: a) lack, or loss during selection, of regeneration capability; b) the development of epigenetically adapted cells; c) lack of correlation between the mechanisms of tolerance operating in cultured cells and mechanisms that operate in cells in the intact plant; and d) multigenicity of salt tolerance. The recent successful production of healthy, fertile, and genetically stable salt-tolerant regenerants from cells obtained from highly morphogenic explants which are selected early in culture (using one-step or short-term strategies) for salt tolerance, together with the demonstration that salt-sensitive plants can become tolerant by mutations in one or few genes, suggest that some of the potential limitations can be overcome and that some of them may not exist at all.     

A method for screening rice plants for salt tolerance

A number of varieties of rice, a halophyte, Sesuvium portulacastrum and a glycophyte, Phaseolus vulgaris were grown in culture solution containing a range of concentrations of NaCl. Growth of the plants and internal sodium concentrations of the roots were measured after 14 days. The electrical potential difference (PD) between the external solution and the vacuole of the outer cells of the root was also measured. This enabled the driving force on sodium at the cell membranes to be calculated using the Nernst equation. It was found that Sesuvium and those varieties of rice that had previously shown salt tolerance generated relatively negative PDs and large driving forces tending to exclude sodium from the root. This suggested that a simple measurement of PD for plants grown in a given concentration of NaCl over a given period of time would provide a fairly rapid screening method for salt tolerance in rice and possibly other species also. T J Flowers Section editor     

Studies on the salt tolerance of some flower crops grown under glass

Summary The salt sensitivity of carnations, gerberas, anthuriums, chrysanthemums and hippeastrums was studied in an investigation. The crops were grown in basins filled with soil and irrigated with the aid of low level sprinklers. Two different cultivars of each crop were included in the experiments. The crops were irrigated with water containing different levels of salts. The EC of the irrigation water (EC_w) ranged between 0.2 and 3.9 mS.cm^–1 at 25°C.The salt applications had a deleterious effect on the development of all the flower crops used in the investigation. Carnations and chrysanthemums proved to be the least sensitive. Gerberas and hippeastrums showed a medium sensitivity and anthuriums proved to be the most salt sensitive. The latter crop also showed a specific sensitivity to sodium chloride.The salinity threshold values were low for most crops,i.e. EC_w<0.6. Slightly higher values were found for carnations only. The salinity decrease values of EC_w showed very wide variations and ranged from 6 to 34%. A method is proposed for the conversion of EC_w values into EC values for the saturation extract (EC_e) with the aid of the soil analytical results.The results of tissue analyses showed that the sodium and chloride contents in particular were affected by the salt applications. The effects of the salt applications on the uptake of the major nutrient elements varied from crop to crop.     

Response to short-term inundation with isoosmotic solutions of seawater and sorbitol in a C4 nonhalophyte: evidence for a salt tolerance mechanism

Summary To examine the importance of Na⁺ and Cl^- to osmotic adjustment in a salt-tolerant ecotype of the C₄ nonhalophyte Andropogon glomeratus, plants were watered with sorbitol, a neutral osmoticum, and synthetic seawater, for five days. Gas exchange measurements were made during the course of the watering treatment and during a recovery period following treatment. Leaf osmotic adjustment occurred only in plants watered with seawater, and was associated with an increase in Na⁺ and Cl^- concentrations. Estimates of the molar concentrations indicated these ions could account for 95% of the leaf osmotic adjustment. Net photosynthetic CO₂ uptake was less effected during the watering treatment, and photosynthetic recovery was greater following the treatment in plants watered with seawater. Photosynthetic inhibition was related primarily to metabolic factors, including a decrease in carboxylation efficiency. A model is presented for a mechanism promoting tolerance to transient seawater inundation in A. glomeratus.     

Some physiological and morphological characteristics of citrus plants for drought resistance

Tolerance and avoidance mechanisms to drought stress were studied in 6-month-old plants of Newhall orange (Citrus sinensis (L.) Osbeck) and Ellendale tangor (orange × mandarin hybrid) (Citrus sinensis (L) Osbeck × Citrus reticulata Blanco) during a drought/rewatering cycle under controlled conditions. Drought stress did not promote osmotic adjustment, while elastic adjustment (tissue elasticity increase) was noted in stressed orange and tangor plants. Both citrus plants showed a parallel decrease in leaf conductance (g₁) and leaf water potential (Ψ₁) under water stress. Tangor plants had a more efficient water conservative strategy than orange, based on the characteristics of canopy architecture (lower canopy area and a more closed canopy with leaves nearly vertically oriented) together with a significant decrease in cuticular transpiration rates (TRc) under stress.     

Altered physiology in trehalose-producing transgenic tobacco plants: Enhanced tolerance to drought and salinity stresses

Transgenic tobaccoNicotiana tabacum L. var. SR1) plants that over-express theEscherichia coli trehalose-6-phosphate synthase (TPS) gene(otsA) synthesized small amounts of trehalose (<400 μg g^-1 leaf) while non-transformants produced no detectable trehalose. Some transgenic plants expressing a high level ofotsA exhibited stunted growth and morphologically altered leaves. We tested F₂2 homozygous plants devoid of phenotypic changes to determine their physiological responses to dehydration and salinity stresses. All transgenic plants maintained better leaf turgidity under a limited water supply or after treatment with polyethylene glycol (PEG). Furthermore, fresh weight was maintained at higher levels after either treatment. The initial leaf water potential was higher in transgenic plants than non-transformants, but, in both plant types, was decreased to a comparable degree following dehydration. When grown with 250 mM NaCl, transgenic plants exhibited a significant delay in leaf withering and chlorosis, as well as more efficient seed germination. Our results suggest that either trehalose or trehalose-6-phosphate can act as an osmoprotective molecule without maintaining water potential, in contrast to other osmolytes. Furthermore, both appear to protect young embryos under unfavorable water status to ensure subsequent germination.     

植物耐盐的分子生物学基础

植物分子水平的耐盐研究是近年的研究热点,通过综述与耐盐有关的几种重要分子的性质和作用,总结了几种与植物耐盐有关的基因以及它们在盐分胁迫下的表达和调控。     

Macroevolutionary patterns of salt tolerance in angiosperms

Background Halophytes are rare, with only 0·25 % of angiosperm species able to complete their life cycle in saline conditions. This could be interpreted as evidence that salt tolerance is difficult to evolve. However, consideration of the phylogenetic distribution of halophytes paints a different picture: salt tolerance has evolved independently in many different lineages, and halophytes are widely distributed across angiosperm families. In this Viewpoint, I will consider what phylogenetic analysis of halophytes can tell us about the macroevolution of salt tolerance.Hypothesis Phylogenetic analyses of salt tolerance have shown contrasting patterns in different families. In some families, such as chenopods, salt tolerance evolved early in the lineage and has been retained in many lineages. But in other families, including grasses, there have been a surprisingly large number of independent origins of salt tolerance, most of which are relatively recent and result in only one or a few salt-tolerant species. This pattern of many recent origins implies either a high transition rate (salt tolerance is gained and lost often) or a high extinction rate (salt-tolerant lineages do not tend to persist over macroevolutionary timescales). While salt tolerance can evolve in a wide range of genetic backgrounds, some lineages are more likely to produce halophytes than others. This may be due to enabling traits that act as stepping stones to developing salt tolerance. The ability to tolerate environmental salt may increase tolerance of other stresses or vice versa.Conclusions Phylogenetic analyses suggest that enabling traits and cross-tolerances may make some lineages more likely to adapt to increasing salinization, a finding that may prove useful in assessing the probable impact of rapid environmental change on vegetation communities, and in selecting taxa to develop for use in landscape rehabilitation and agriculture.