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As halophytes grow vigorously in saline soils, they serve as extraordinary resources for the identification and development of new crop systems. Understanding the mechanisms of tolerance of halophytes to salinity in combination with other co-occurring constraints such as drought, flooding, heavy metals and nutrient deficiencies, would facilitate efforts to use halophytes for saline land revegetation, as well as provide new insights that might be considered in future breeding of plants for salt-affected agricultural lands. Recent results suggest that salinity may improve the response of halophytes to other stresses. Some physiological and biochemical mechanisms of tolerance to salinity are common to many halophytes when plants are subjected to salinity, whereas others are specifically amplified under a combination of stresses. Therefore, the response of halophytes to multiple stresses may not reflect an additive effect of these constraints, but rather, constitute specific response to a new situation where many constraints are operating simultaneously. Comparative studies between halophytes and glycophytes have shown that halophytes are better equipped with the mechanisms of cross-stress tolerance and are constitutively prepared for stress. Moreover, other data has shown that the pre-treatment of halophytes with salinity or other constraints in the early stages of development improves their subsequent response to salinity, which suggests the capacity of these plants to ‘memorise’ a previous stress allows them respond positively to subsequent stress.

Though halophytes are naturally adapted to salinity, their salt-tolerance limits are greatly influenced by their provenance and developmental stage. In the present study, physio-biochemical responses of two Tunisian ecotypes of the oilseed coastal halophyte Cakile maritima (Brassicaceae) to salinity (0-400 mM NaCl) were monitored during germination and vegetative growth stages. Tabarka and Jerba seeds were collected from humid or arid climatic areas, respectively. Plant response to salinity appeared to depend on the ecotype and salinity levels. Increasing salinity inhibited germination process. Jerba seeds were found to be more salt tolerant than the Tabarka ones. At the autotrophic stage of growth and under salt-free conditions, Jerba was less productive than Tabarka (in terms of dry matter accumulation), but plant biomass production and leaf expansion (area and number) of the former ecotype were progressively improved by 100 mM NaCl, as compared to the control. In contrast, at the same salt concentration, these parameters decreased under increasing salinity in Tabarka (salt sensitive). Leaf chlorophyll content was reduced at severe salinity, but this effect was more conspicuous in the sensitive Tabarka plants. Na + contents in the Jerba and Tabarka leaves collected from the 400 mM NaCl-treated plants were 17-and 12-fold higher than in the respective controls. This effect was accompanied by a significant reduction in the leaf K + , Mg 2+ and Ca 2+ contents, especially in the salt-treated Tabarka. A significant accumulation of proline and soluble carbohydrates in leaves was found during the period of intensive leaf growth. These organic compounds likely play a role in leaf osmotic adjustment and in protection of membrane stability at severe salinity.

Sesuvium portulacastrum is a halophytic species well adapted to salinity and drought. In order to evaluate the physiological impact of salt on water deficit-induced stress response, we cultivated seedlings for 12 days, in the presence or absence of 100 mmol l(-1) NaCl, on a nutrient solution containing either 0 mmol l(-1) or 25 mmol l(-1) mannitol. Mannitol-induced water stress reduced growth, increased the root/shoot ratio, and led to a significant decrease in water potential and leaf relative water content, whereas leaf Na(+) and K(+) concentrations remained unchanged. The addition of 100 mmol l(-1) NaCl to 25 mmol l(-1) mannitol-containing medium mitigated the deleterious impact of water stress on growth of S. portulacastrum, improved the relative water content, induced a significant decrease in leaf water potential and, concomitantly, resulted in enhancement of overall plant photosynthetic activity (i.e. CO(2) net assimilation rate, stomatal conductance). Presence of NaCl in the culture medium, together with mannitol, significantly increased the level of Na(+) and proline in the leaves, but it had no effect on leaf soluble sugar content. These findings suggest that the ability of NaCl to improve plant performance under mannitol-induced water stress may be due to its effect on osmotic adjustment through Na(+) and proline accumulation, which is coupled with an improvement in photosynthetic activity. A striking recovery in relative water content and growth of the seedlings was also recorded in the presence of NaCl on release of the water stress induced by mannitol.