Halophytes, plants that survive to reproduce in environments where the salt concentration is around 200 mM NaCl or more, constitute about 1% of the worlds flora. Some halophytes show optimal growth in saline conditions; others grow optimally in the absence of salt. However, the tolerance of all halophytes to salinity relies on controlled uptake and compartmentalization of Na+, K+ and Cl- and the synthesis of organic compatible solutes, even where salt glands are operative. Although there is evidence that different species may utilize different transporters in their accumulation of Na+, in general little is known of the proteins and regulatory networks involved. Consequently, it is not yet possible to assign molecular mechanisms to apparent differences in rates of Na+ and Cl- uptake, in root-to-shoot transport (xylem loading and retrieval), or in net selectivity for K+ over Na+. At the cellular level, H+-ATPases in the plasma membrane and tonoplast, as well as the tonoplast H+-PPiase, provide the transmembrane proton motive force used by various secondary transporters. The widespread occurrence, taxonomically, of halophytes and the general paucity of information on the molecular regulation of tolerance mechanisms persuade us that research should be concentrated on a number of model species that are representative of the various mechanisms that might be involved in tolerance.

Salinity tolerance in halophytes

Salinity is an ever-present threat to crop yields, especially in countries where irrigation is an essential aid to agriculture. Although the tolerance of saline conditions by plants is variable, crop species are generally intolerant of one-third of the concentration of salts found in seawater. Attempts to improve the salt tolerance of crops through conventional breeding programmes have met with very limited success, due to the complexity of the trait: salt tolerance is complex genetically and physiologically. Tolerance often shows the characteristics of a multigenic trait, with quantitative trait loci (QTLs) associated with tolerance identified in barley, citrus, rice, and tomato and with ion transport under saline conditions in barley, citrus and rice. Physiologically salt tolerance is also complex, with halophytes and less tolerant plants showing a wide range of adaptations. Attempts to enhance tolerance have involved conventional breeding programmes, the use of in vitro selection, pooling physiological traits, interspecific hybridization, using halophytes as alternative crops, the use of marker-aided selection, and the use of transgenic plants. It is surprising that, in spite of the complexity of salt tolerance, there are commonly claims in the literature that the transfer of a single or a few genes can increase the tolerance of plants to saline conditions. Evaluation of such claims reveals that, of the 68 papers produced between 1993 and early 2003, only 19 report quantitative estimates of plant growth. Of these, four papers contain quantitative data on the response of transformants and wild-type of six species without and with salinity applied in an appropriate manner. About half of all the papers report data on experiments conducted under conditions where there is little or no transpiration: such experiments may provide insights into components of tolerance, but are not grounds for claims of enhanced tolerance at the whole plant level. Whether enhanced tolerance, where properly established, is due to the chance alteration of a factor that is limiting in a complex chain or an effect on signalling remains to be elucidated. After ten years of research using transgenic plants to alter salt tolerance, the value of this approach has yet to be established in the field.

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Improving crop salt tolerance

http://nhjy.hzau.edu.cn/kech/ssyy/qysd/njsl/21.pdf

Potential biochemical indicators of salinity tolerance in plants

This review describes physiological mechanisms and selectable indicators of gene action, with the aim of promoting new screening methods to identify genetic variation for increasing the salt tolerance of cereal crops. Physiological mechanisms that underlie traits for salt tolerance could be used to identify new genetic sources of salt tolerance. Important mechanisms of tolerance involve Na+ exclusion from the transpiration stream, sequestration of Na+ and Cl- in the vacuoles of root and leaf cells, and other processes that promote fast growth despite the osmotic stress of the salt outside the roots. Screening methods for these traits are discussed in relation to their use in breeding, particularly with respect to wheat. Precise phenotyping is the key to finding and introducing new genes for salt tolerance into crop plants.

Approaches to increasing the salt tolerance of wheat and other cereals

This paper discusses whole-plant responses to salinity in order to answer the question of what process limits growth of non-halophytes in saline soils. Leaf growth is more sensitive to salinity than root growth, so we focus on the process or processes that might limit leaf expansion. Effects of short-term exposure (days) are considered separately from long-term exposure (weeks to years). The answer in the short term is probably the water status of the root and we suggest that a message from the root is regulating leaf expansion. The answer to what limits growth in the long term may be the maximum salt concentration tolerated by the fully expanded leaves of the shoot; if the rate of leaf death approaches the rate of new leaf expansion, the photosynthetic area will eventually become too low to support continued growth.

Whole-plant responses to salinity

The terrestrial halophyte, Salicornia bigelovii Torr., was evaluated as an oilseed crop for direct seawater irrigation during 6 years of field trials in an extreme coastal desert environment. Yields of seed and biomass equated or exceeded freshwater oilseed crops such as soybean and sunflower. The seed contained 26 to 33 percent oil, 31 percent protein, and was low in fiber and ash (5 to 7 percent). The oil and meal were extracted by normal milling equipment, and the oil was high in linoleic acid (73 to 75 percent) and could replace soybean oil in chicken diets. The meal had antigrowth factors, attributed to saponins, but could replace soybean meal in chicken diets amended with the saponin antagonist, cholesterol. Salicornia bigelovii appears to be a potentially valuable new oilseed crop for subtropical coastal deserts.

https://science.sciencemag.org/content/sci/251/4997/1065.full.pdf

Salicornia bigelovii Torr.: An Oilseed Halophyte for Seawater Irrigation

A clear correlation between the presence of stomata on both surfaces and factors such as habitat, growth form, and physiology has yet to emerge in the literature. However, certain loose trends with these factors are evident, and these are reviewed along with evidence for hypostomaty as the primitive form. It is proposed that the effect of developing stomata on the upper surface as well as the lower is to increase maximum leaf conductance to CO2. Plants with a high photosynthetic capacity, living in full-sun environments, and experiencing rapidly fluctuating or continuously available soil water (as opposed to seasonal or long-term soil water depletion), are identified as deriving an adaptive advantage from a high maximum leaf conductance. The correlation between groups of plants fitting the above conditions and those noted to be amphistomatic is remarkable. Plants not fitting the conditions are found to be largely hypostomatic.

The adaptive significance of amphistomatic leaves

Irrigating crops with seawater

Salicornia bigelovii Torr. plants were grown in 5, 200, or 600 mol m-3 NaCl, representing suboptimal, optimal, and supraoptimal salinities, respectively. Shoot fresh and dry mass of plants grown at optimal salinity were more than 2 x higher than those grown at the other two salinities. In spite of the comparable growth reductions at sub- and supraoptimal salinities, the physiological responses, and presumably the causes of the growth reductions, were not the same at those two salinities. Water and osmotic potentials of the shoots decreased significantly with increasing salinity, but turgor potentials did not differ significantly among treatments. Differences in photosynthetic rates were not consistent with the differences in growth. Rates were significantly higher in plants grown at 5 mol m-3 NaCl when expressed relative to photosynthetic area, but when photosynthesis was expressed relative to the amount of chlorophyll, no significant differences were found among salinities. Stomatal conductance decreased...

Growth and Physiology of Salicornia bigelovii Torr. at Suboptimal Salinity

. Growth rates and levels of minerals, Na+, K+, Mg++, Ca++, and water were measured in dicotyledonous halophytes grown along a salinity gradient from fresh water to 720 mol m−3 NaCl in a controlled environment greenhouse. Ten test species from the families Chenopodiaceae, Aizoaceae, and Batidaceae exhibited growth stimulation by 180 mol m−3 NaCl and were classified as euhalophytes. Ten others from the families Chenopodiaceae, Aizoaceae, Asteraceae, Brassicaceae, Polygonaceae, Boraginaceae, Malvaceae, and Plumbaginaceae showed their best growth on fresh water and were classified miohalophytes. Salt, and particularly sodium, accumulated in all halophytes but to a significantly greater extent among euhalophytes than miohalophytes. The water content of most species increased when grown on 180 mol m−3 NaCl compared to fresh water; but at higher salinities some of the species underwent dehydration. Dehydration of the succulent S. europaca was not coupled to a proportional decrease in growth. Water content and cation accumulation in euhalophytes appeared to be coordinated to produce a constant osmotic potential gradient within the shoot tissues relative to the external salinity. In contrast, miohalophytes did not appear to regulate osmotic potential as closely as euhalophytes.

James W. O'Leary

Papers

Salicornia bigelovii Torr.: An Oilseed Halophyte for Seawater Irrigation

The adaptive significance of amphistomatic leaves

Irrigating crops with seawater

Growth and Physiology of Salicornia bigelovii Torr. at Suboptimal Salinity

Relationship between salt accumulation and water content of dicotyledonous halophytes