World population is expected to reach 9.2 × 109 people by 2050. Feeding them will require a boost in crop productivity using innovative approaches. Current agricultural production is very dependent on large amounts of inputs and water availability is a major limiting factor. In addition, the loss of genetic diversity and the threat of climate change make a change of paradigm in plant breeding and agricultural practices necessary. Average yields in all major crops are only a small fraction of record yields, and drought and soil salinity are the main factors responsible for yield reduction. Therefore there is the need to enhance crop productivity by improving crop adaptation. Here we review the present situation and propose the development of crops tolerant to drought and salt stress for addressing the challenge of dramatically increasing food production in the near future. The success in the development of crops adapted to drought and salt depends on the efficient and combined use of genetic engineering and traditional breeding tools. Moreover, we propose the domestication of new halophilic crops to create a ‘saline agriculture’ which will not compete in terms of resources with conventional agriculture.
We have studied the responses to changing environmental conditions of five halophytes in a Mediterranean salt marsh, during a 2-year period. Salt tolerance in succulent dicotyledonous halophytes is mostly dependent on compartmentalisation of toxic ions in vacuoles and biosynthesis of osmolytes for osmotic adjustment – mechanisms that appear to be constitutive in the most tolerant taxa – while monocots avoid excessive ion transport to the plant aerial parts. Contrary to what has been described for salt treatments under artificial conditions, the selected halophytes are not affected by oxidative stress in their natural habitat, and do not need to activate antioxidant defence mechanisms.
Nerium oleander is an ornamental species of high aesthetic value, grown in arid and semi-arid regions because of its drought tolerance, which is also considered as relatively resistant to salt; yet the biochemical and molecular mechanisms underlying oleander’s stress tolerance remain largely unknown. To investigate these mechanisms, one-year-old oleander seedlings were exposed to 15 and 30 days of treatment with increasing salt concentrations, up to 800 mM NaCl, and to complete withholding of irrigation; growth parameters and biochemical markers characteristic of conserved stress-response pathways were then determined in stressed and control plants. Strong water deficit and salt stress both caused inhibition of growth, degradation of photosynthetic pigments, a slight (but statistically significant) increase in the leaf levels of specific osmolytes, and induction of oxidative stress—as indicated by the accumulation of malondialdehyde (MDA), a reliable oxidative stress marker—accompanied by increases in the levels of total phenolic compounds and antioxidant flavonoids and in the specific activities of ascorbate peroxidase (APX) and glutathione reductase (GR). High salinity, in addition, induced accumulation of Na+ and Cl- in roots and leaves and the activation of superoxide dismutase (SOD) and catalase (CAT) activities. Apart from anatomical adaptations that protect oleander from leaf dehydration at moderate levels of stress, our results indicate that tolerance of this species to salinity and water deficit is based on the constitutive accumulation in leaves of high concentrations of soluble carbohydrates and, to a lesser extent, of glycine betaine, and in the activation of the aforementioned antioxidant systems. Moreover, regarding specifically salt stress, mechanisms efficiently blocking transport of toxic ions from the roots to the aerial parts of the plant appear to contribute to a large extent to tolerance in Nerium oleander.
We have performed an extensive study on the responses to salt stress in four related Limonium halophytes with different geographic distribution patterns, during seed germination and early vegetative growth. The aims of the work were twofold: to establish the basis for the different chorology of these species, and to identify relevant mechanisms of salt tolerance dependent on the control of ion transport and osmolyte accumulation. Seeds were germinated in vitro, in the presence of increasing NaCl concentrations, and subjected to “recovery of germination” tests; germination percentages and velocity were determined to establish the relative tolerance and competitiveness of the four Limonium taxa. Salt treatments were also applied to young plants, by 1-month irrigation with NaCl up to 800 mM; then, growth parameters, levels of monovalent and divalent ions (in roots and leaves), and leaf contents of photosynthetic pigments and common osmolytes were determined in control and stressed plants of the four species. Seed germination is the most salt-sensitive developmental phase in Limonium. The different germination behavior of the investigated species appears to be responsible for their geographical range size: L. narbonense and L. virgatum, widespread throughout the Mediterranean, are the most tolerant and the most competitive at higher soil salinities; the endemic L. santapolense and L. girardianum are the most sensitive and more competitive only at lower salinities. During early vegetative growth, all taxa showed a strong tolerance to salt stress, although slightly higher in L. virgatum and L. santapolense. Salt tolerance is based on the efficient transport of Na+ and Cl− to the leaves and on the accumulation of fructose and proline for osmotic adjustment. Despite some species-specific quantitative differences, the accumulation patterns of the different ions were similar in all species, not explaining differences in tolerance, except for the apparent activation of K+ transport to the leaves at high external salinity, observed only in the most tolerant L. narbonense and L. virgatum. This specific response may be therefore relevant for salt tolerance in Limonium. The ecological implications of these results, which can contribute to a more efficient management of salt marshes conservation/regeneration programs, are also discussed.
A general response of plants to high soil salinity relies on the cellular accumulation of osmolytes, which help the plant to maintain osmotic balance under salt stress condition and/or act as ‘osmoprotectants’ with chaperon or reactive oxygen species (ROS) scavenging activities. Yet the ecological relevance of this response for the salt tolerance mechanisms of halophytes in their natural habitats remains largely unknown. In this review, we describe and discuss published data supporting the participation of compatible solutes in those mechanisms, with especial focus on soluble carbohydrates. Evidence for a functional role of carbohydrates in salt tolerance include: (i) relatively high levels of specific sugars and polyols have been detected in many halophytic taxa; (ii) an increase in salt tolerance has often been observed in parallel with increased intracellular levels of particular soluble carbohydrates, in transgenic plants overexpressing the corresponding biosynthetic enzymes; (iii) there are several examples of genes involved in carbohydrate metabolism which are induced under salt stress conditions; (iv) specific sugars or polyols have been shown to accumulate in different halophytes upon controlled salt treatments; and (v) although very few field studies on environmentally induced carbohydrate changes in halophytes exist, in general they also support the involvement of this type of osmolytes in salt stress tolerance mechanisms. We also highlight the complexities of unequivocally attributing carbohydrates a biological role in salt tolerance mechanisms of a given tolerant species. It is proposed that research on halophytes in their natural ecosystems should be intensified, correlating seasonal changes in carbohydrate contents with the degree of environmental stress affecting the plants. This could be an important complement to experiments made under more controlled (but artificial) conditions, such as laboratory set-ups.
The effects of salt and water stress on growth and several stress markers were investigated in cherry tomato plants. Some growth parameters (stem length and number of leaves) and chlorophyll contents were determined every third day during plant growth, and leaf material was collected after 25 and 33 days of treatment. Both stresses inhibited plant growth; chlorophyll levels, however, decreased only in response to high NaCl concentrations. Proline contents largely increased in leaves of stressed plants, reaching levels high enough to play a major role in cellular osmotic adjustment. Despite reports indicating that tomato does not synthesize glycine betaine, the stress-induced accumulation of this osmolyte was detected in cherry tomato, albeit at lower concentration than that of proline. Therefore, it appears that the plants are able to synthesise glycine betaine as a secondary osmolyte under strong stress conditions. Total sugars levels, on the contrary, decreased in stress-treated plants. Both stress treatments caused secondary oxidative stress in the plants, as indicated by a significant increase in malondialdehyde (MDA) contents. Water stress led to an increase in total phenolics and flavonoid contents and a reduction of carotenoid levels in the leaves; flavonoids also increased under high salinity conditions.
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