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Diversity, distribution and roles of osmoprotective compounds accumulated in halophytes under abiotic stress.

BACKGROUND AND AIMS Osmolytes are low-molecular-weight organic solutes, a broad group that encompasses a variety of compounds such as amino acids, tertiary sulphonium and quaternary ammonium compounds, sugars and polyhydric alcohols. Osmolytes are accumulated in the cytoplasm of halophytic species in order to balance the osmotic potential of the Na(+) and Cl(-) accumulated in the vacuole. The advantages of the accumulation of osmolytes are that they keep the main physiological functions of the cell active, the induction of their biosynthesis is controlled by environmental cues, and they can be synthesized at all developmental stages. In addition to their role in osmoregulation, osmolytes have crucial functions in protecting subcellular structures and in scavenging reactive oxygen species. SCOPE This review discusses the diversity of osmolytes among halophytes and their distribution within taxonomic groups, the intrinsic and extrinsic factors that influence their accumulation, and their role in osmoregulation and osmoprotection. Increasing the osmolyte content in plants is an interesting strategy to improve the growth and yield of crops upon exposure to salinity. Examples of transgenic plants as well as exogenous applications of some osmolytes are also discussed. Finally, the potential use of osmolytes in protein stabilization and solvation in biotechnology, including the pharmaceutical industry and medicine, are considered.

Combined effects of long-term salinity and soil drying on growth, water relations, nutrient status and proline accumulation of Sesuvium portulacastrum

The interaction between soil drying and salinity was studied in the perennial halophyte, Sesuvium portulacastrum. Rooted cuttings were individually cultivated for three months in silty-sandy soil under two irrigation modes: 100 and 25% of field capacity (FC). The amount of the evapotranspirated water was replaced by a nutrient solution containing either 0 or 100 mM NaCl. Whole-plant growth, leaf water content, leaf water potential (Psi(w)), and Na+, K+, and proline concentrations in the tissues were measured. When individually applied, both drought and salinity significantly restricted whole-plant growth, with a more marked effect of the former stress. However, the effects of the two stresses were not additive on whole-plant biomass or on leaf expansion. Root growth was more sensitive to salt than to soil drying, the latter being even magnified by the adverse impact of salinity. Leaf water content was significantly reduced following exposure to water-deficit stress, but was less affected in salt-treated plants. When simultaneously submitted to water-deficit stress and salinity, plants displayed higher values of water and potassium use efficiencies, leaf proline and Na+ concentrations, associated with lower leaf water potential (-1.87 MPa), suggesting the ability of S. portulacastrum to use Na+ and proline for osmotic adjustment.

Effects of water deficit stress on growth, water relations and osmolyte accumulation in Medicago truncatula and M. laciniata populations

The effects of water stress were investigated in two Tunisian Medicago truncatula populations collected from arid (Mt-173) and sub-humid (Mt-664) climates and two Tunisian M. laciniata populations originating from arid (Ml-173) and semi-arid (Ml-345) regions. After a pre-treatment phase (24 days after sowing, DAS) of watering at 100% of field capacity (FC), the plants were either irrigated at 100% FC or at only 33% FC. After 12 days of treatment (36 DAS), one lot of dehydrated plants was rewatered at 100% FC. A final harvest was carried out after 24 days of treatment (48 DAS). Measured parameters were total dry weight (TDW), root shoot ratio (RSR), leaf relative water content (RWC), osmotic potential (OP), photosynthetic parameters (CO(2) net assimilation A, stomatal conductance g(s) and transpiration E), malondialdehyde (MDA) concentration and leaf contents in inorganic (Na(+) and K(+)) and organic solutes (proline and soluble sugars). Under water deficit conditions, compared to M. laciniata, M. truncatula populations showed a higher reduction in TDW, A, g(s) and RWC associated with a higher increase in MDA concentration. Thus, the relative tolerance of M. laciniata populations to water shortage would be related to their lower intrinsic growth rate and stomatal control of gas exchange. TDW, A, g(s), E and RWC were more decreased by water deficit in Ml-345 than in Ml-173. Drought tolerance of Ml-173 was found to be associated with a more pronounced decrease of OP and a lower reduction in RWC due to the accumulation of solutes such as proline, soluble sugars and K(+). In addition, Ml-173 showed the highest water use efficiency values (WUE) and the lowest MDA concentrations under water deficit stress.

Differential response to water deficit stress in alfalfa (Medicago sativa) cultivars: Growth, water relations, osmolyte accumulation and lipid peroxidation

The present study was fixed as objective to compare the response to water deficit (33% of field capacity, FC) stress of eight cultivars of Medicago sativa, originating from the Mediterranean basin. Comparison was performed on some key parameters such as growth, relative water content, leaf water potential, MDA tissue content, electrolyte leakage and proline and soluble sugar tissue concentrations. In all cultivars, water deficit stress reduced the whole plant growth, increased the root/shoot DW ratio and led to a significant decrease in leaf relative water content and leaf water potential. In water-stressed plants and for the majority of cultivars, proline and soluble sugar concentrations increased significantly. The Tamantit cultivar originating from Algeria showed the better tolerance to water deficit stress. Some criteria are concomitant with this tolerance: (1) a high biomass production under stress conditions when compared to other cultivars (2) the root preferential development, (3) the better ability to accumulate proline and soluble sugars, and (4) the aptitude to protect its photosynthetic apparatus against the oxidative stress generated by the water deficit stress. Ecotipo Siciliano, originating from Italy was the most sensitive cultivar. The increase of proline and soluble sugars concentrations upon water deficit stress particularly in the most tolerant cultivar suggested their involvement in the osmotic adjustment.

Medicago sativa - Sinorhizobium meliloti Symbiosis Promotes the Bioaccumulation of Zinc in Nodulated Roots

In this study we investigated effects of Zn supply on germination, growth, inorganic solutes (Zn, Ca, Fe, and Mg) partitioning and nodulation of Medicago sativa This plant was cultivated with and without Zn (2 mM). Treatments were plants without (control) and with Zn tolerant strain (S532), Zn intolerant strain (S112) and 2 mM urea nitrogen fertilisation. Results showed that M. sativa germinates at rates of 50% at 2 mM Zn. For plants given nitrogen fertilisation, Zn increased plant biomass production. When grown with symbionts, Zn supply had no effect on nodulation. Moreover, plants with S112 showed a decrease of shoot and roots biomasses. However, in symbiosis with S532, an increase of roots biomass was observed. Plants in symbiosis with S. meliloti accumulated more Zn in their roots than nitrogen fertilised plants. Zn supply results in an increase of Ca concentration in roots of fertilised nitrogen plants. However, under Zn supply, Fe concentration decreased in roots and increased in nodules of plants with S112. Zn supply showed contrasting effects on Mg concentrations for plants with nitrogen fertilisation (increase) and plants with S112 (decrease). The capacity of M. sativa to accumulate Zn in their nodulated roots encouraged its use in phytostabilisation processes.

Effect of nitrogen deficiency, salinity and drought on proline metabolism in Sesuvium portulacastrum

Drought and high salinity are responsible for large decreases in crop productivity all over theworld [1]. These losses of crop yield result from limitations of plant development through excessive ion accumulation, water deficit and mineral deficiencies [2]. Under these prevalent stresses, tolerant plants adopt various strategies with a wide range of biochemical to physiological and morphological adaptations [3]. Morphological ones include modifications in growth and allocation of assimilates towards roots for an efficient exploitation of soil nutrients [4]. The physiological strategy is represented by a higher selectivity for K+ over Na+ [5], an increase in K+-use efficiency [6], and the synthesis of organic osmolytes, with low molecular weight, for osmo-protection [7]. These osmolytes are sugars, polyols, amino acids, tertiary and quarternary ammonium, and tertiary sulphonium compounds [8].

Differential responses to phosphorus availability in the halophytes Aeluropus Littoralis, Catapodium rigidum, and Hordeum maritimum

ABSTRACT The objective of this experiment was to explore the variability of response to phosphorus (P) availability in three fodder Poaceae species native of saline biotopes, Aeluropus littoralis, Catapodium rigidum, and Hordeum maritimum. The behavior of these wild species has been compared to that of a cultivated one, Hordeum vulgare. Plants were grown in plastic pots filled with inert sand. The experiment was carried out in a greenhouse with six P levels: 0, 5, 15, 60, 180, and 360 µM P concentration supplied as KH2PO4. Results showed that the growth of the four species is strongly affected by the low phosphorus availability in the medium. Furthermore, to reach 80% of their maximum yield expressed as shoot dry weight (DW), a soil phosphorus concentration of approximately 40 µM P concentration is sufficient for Hordeum maritimum, whereas approximately 66, 120, and 132 µM P concentration are needed for C. rigidum, A. littoralis, and Hordeum vulgare plants, respectively. Thus, our results showed that spontaneous species are more tolerant to P deficiency than cultivated barley. The relative performance of the three wild Poaceae species is likely related to their capacity to maintain higher P acquisition efficiency (PAE) under low P concentration conditions. These native Poaceae species may therefore represent genetic resources for the improvement of barley.

Effects of salt treatment on growth, lipid membrane peroxidation, polyphenol content, and antioxidant activities in leaves of Sesuvium portulacastrum L.

ABSTRACT Changes in growth, leaf contents of proline, oxidative stress-related parameters, and phenolic compounds, and antioxidant activities were investigated in the halophyte species Sesuvium portulacastrum L. under saline conditions. Rooted cuttings were individually cultivated in sandy soil. After five weeks of pre-treatment, seedlings were submitted during one month to different salt concentrations ranging from 0 to 800 mM NaCl. The plant growth was significantly improved by salt at 200–600 mM concentration. This trend was associated with (i) the stimulation of photosynthetic activity, (ii) the protection of membrane integrity (leaf MDA content 50% lower than the control), and (iii) higher total antioxidant activity, especially at 400 mM NaCl. At this salt concentration plants accumulated high contents of proline, polyphenols, antocyanins, and carotenoids. These compounds could be implied in the protection of the photosynthetic system and in the improvement of growth. Exposure to 800 mM NaCl impaired significantly photosynthesis, proline, polyphenol, antocyanin, and carotenoid accumulation. Yet, the strong antiradical activity (DPPH) observed at this extreme salinity might partly explain the plant survival. S. portulacastrum could be used in the rehabilitation and the stabilisation of saline or saline arid land. Additionally, under saline conditions, S. portulacastrum accumulate a large amount of proline and exhibits important antioxidant potentialities.

Insights into the physiological responses of the facultative halophyte Aeluropus littoralis to the combined effects of salinity and phosphorus availability.

In this work, we investigate the physiological responses to P deficiency (5μM KH2PO4=D), salt stress (400mM NaCl=C+S), and their combination (D+S) on the facultative halophyte Aeluropus littoralis to understand how plants adapt to these combined stresses. When individually applied, both P deficiency and salinity significantly restricted whole plant growth, with a more marked effect of the latter stress. However, the effects of the two stresses were not additive in plant biomass production since the response of plants to combined salinity and P deficiency was similar to that of plants grown under salt stress alone. In addition the observed features under salinity alone are kept when plants are simultaneously subjected to the combined effects of salinity and P deficiency such as biomass partitioning; the synthesis of proline and the K(+)/Na(+) selectivity ratio. Thus, increasing P availability under saline conditions has no significant effect on salt tolerance in this species. Plants cultivated under the combined effects of salinity and P deficiency exhibited the lowest leaf water potential. This trend was associated with a high accumulation of Na(+), Cl(-) and proline in shoots of salt treated plants suggesting the involvement of these solutes in osmotic adjustment. Proline could be involved in other physiological processes such as free radical scavenging. Furthermore, salinity has no significant effect on phosphorus acquisition when combined with a low P supply and it significantly decreased this parameter when combined with a sufficient P supply. This fact was probably due to salts effect on P transporters. In addition, shoot soluble sugars accumulation under both P deficiency treatments with and without salt likely play an important role in the adaptation of A. littoralis plants to P shortage applied alone or combined with salinity. Moreover, there was a strong correlation between shoot and root intracellular acid phosphatase activity and phosphorus use efficiency which strengthens the assumption that intracellular acid phosphatase enzymes are involved in P remobilization in this species. Finally, our results showed that P availability has no significant effect on salt excretion in A. littorlais which suggests that independently of the P status in the plant, excretion remains priority over other functions requiring energy such as growth. This result could also indicate that salt excretion is not energy-dependent in this species.

Differential performance of two forage species, Medicago truncatula and Sulla carnosa, under water-deficit stress and recovery

Abstract. The response patterns during water deficit stress and subsequent recovery of two forage species, Medicago truncatula and Sulla carnosa, were studied. After germination and pre-treatment, seedlings were individually cultivated for two months under two irrigation modes: 100% and 33% of field capacity. Measured parameters were plant growth, water relations, leaf osmotic potential, lipid peroxidation, and leaf inorganic (Na+ and K+) and organic (proline and soluble sugars) solute contents, as well as delta-1-pyrroline-5-carboxylate synthase (P5CS) and proline dehydrogenase (PDH) activities. Our results showed that under control conditions, and in contrast to roots, no significant differences were observed in shoot biomass production between the two species. However, when subjected to water-deficit stress, M. truncatula appeared to be more tolerant than S. carnosa (reduction by 50 and 70%, respectively). In the two studied species, water-deficit stress led to an increase in root/shoot ratio and leaf proline and soluble sugar contents, and a decrease in leaf osmotic potential. Enzymatic assay revealed that in the two species, P5CS activity was stimulated whereas that of PDH was inhibited under stress conditions. Despite greater accumulation of proline, sugar, and potassium in leaves of S. carnosa, M. truncatula was more tolerant to water deficit. This was essentially due to its capacity to control tissue hydration and water-use efficiency, in addition to its greater ability to protect membrane integrity. Following stress relief, M. truncatula and S. carnosa showed partial re-establishment of growth capacity.