Thriving under Salinity: Growth, Ecophysiology and Proteomic Insights into the Tolerance Mechanisms of Obligate Halophyte Suaeda fruticosa.

Studies on obligate halophytes combining eco-physiological techniques and proteomic analysis are crucial for understanding salinity tolerance mechanisms but are limited. We thus examined growth, water relations, ion homeostasis, photosynthesis, oxidative stress mitigation and proteomic responses of an obligate halophyte Suaeda fruticosa to increasing salinity under semi-hydroponic culture. Most biomass parameters increased under moderate (300 mmol L-1 of NaCl) salinity, while high (900 mmol L-1 of NaCl) salinity caused some reduction in biomass parameters. Under moderate salinity, plants showed effective osmotic adjustment with concomitant accumulation of Na+ in both roots and leaves. Accumulation of Na+ did not accompany nutrient deficiency, damage to photosynthetic machinery and oxidative damage in plants treated with 300 mmol L-1 of NaCl. Under high salinity, plants showed further decline in sap osmotic potential with higher Na+ accumulation that did not coincide with a decline in relative water content, Fv/Fm, and oxidative damage markers (H2O2 and MDA). There were 22, 54 and 7 proteins in optimal salinity and 29, 46 and 8 proteins in high salinity treatment that were up-regulated, down-regulated or exhibited no change, respectively, as compared to control plants. These data indicate that biomass reduction in S. fruticosa at high salinity might result primarily from increased energetic cost rather than ionic toxicity.

Effects of Chloride and Sulfate Salts on Seed Germination and Seedling Growth of Ballota hirsuta Benth. and Myrtus communis L.

Soil salinity is a well-known abiotic factor affecting the germination and seedling growth of various plant species. Therefore, we evaluated the effects of different chloride salts (NaCl, KCl and MgCl2) and sulfate salts (Na2SO4, K2SO4 and MgSO4) on the seed germination and early seedling growth of two important ethnomedicinal shrubs of North Africa and the Mediterranean basin (Ballota hirsuta and Myrtus communis). Seeds of these species were subjected to five salinity levels (0-100 mM) and incubated at 20 °C under a light regime (12 h photoperiod). Both species demonstrated their highest germination percentage under control conditions (i.e., without salinity). However, as salinity levels increased, the germination percentages for both species decreased, regardless of the type of salt used. Cations appeared to be more determinative than the anions in regulating the seed germination of both species. M. communis seeds displayed greater sensitivity to sodium (Na+) salts, especially when accompanied with chloride (Cl-) anions. At the higher salt concentrations (75 and 100 mM), Na+ salts had a more pronounced inhibitory effect on M. communis seedling growth compared to potassium (K+) and magnesium (Mg2+) salts. Conversely, Mg2+ salts were more detrimental to seedling growth in B. hirsuta. Based on our results, it can be concluded that both of these species are able to tolerate a moderate level of salinity. Overall, B. hirsuta may be a promising choice for rehabilitating the soils dominated by chloride salts, while M. communis could be utilized for restoring sulfate-dominated soils.

Salt-Induced Modulation of Ion Transport and PSII Photoprotection Determine the Salinity Tolerance of Amphidiploid Brassicas.

Brassica species show varying levels of resistance to salt stress. To understand the genetics underlying these differential stress tolerance patterns in Brassicas, we exposed two widely cultivated amphidiploid Brassica species having different genomes, Brassica juncea (AABB, n = 18) and Brassica napus (AACC, n = 19), to elevated levels of NaCl concentration (300 mM, half the salinity of seawater). B. juncea produced more biomass, an increased chlorophyll content, and fewer accumulated sodium (Na+) and chloride (Cl-) ions in its photosynthesizing tissues. Chlorophyll fluorescence assays revealed that the reaction centers of PSII of B. juncea were more photoprotected and hence more active than those of B. napus under NaCl stress, which, in turn, resulted in a better PSII quantum efficiency, better utilization of photochemical energy with significantly reduced energy loss, and higher electron transport rates, even under stressful conditions. The expression of key genes responsible for salt tolerance (NHX1 and AVP1, which are nuclear-encoded) and photosynthesis (psbA, psaA, petB, and rbcL, which are chloroplast-encoded) were monitored for their genetic differences underlying stress tolerance. Under NaCl stress, the expression of NHX1, D1, and Rubisco increased several folds in B. juncea plants compared to B. napus, highlighting differences in genetics between these two Brassicas. The higher photosynthetic potential under stress suggests that B. juncea is a promising candidate for genetic modifications and its cultivation on marginal lands.

Effects of Salinity Stress on Chloroplast Structure and Function.

Salinity is a growing problem affecting soils and agriculture in many parts of the world. The presence of salt in plant cells disrupts many basic metabolic processes, contributing to severe negative effects on plant development and growth. This review focuses on the effects of salinity on chloroplasts, including the structures and function of these organelles. Chloroplasts house various important biochemical reactions, including photosynthesis, most of which are considered essential for plant survival. Salinity can affect these reactions in a number of ways, for example, by changing the chloroplast size, number, lamellar organization, lipid and starch accumulation, and interfering with cross-membrane transportation. Research has shown that maintenance of the normal chloroplast physiology is necessary for the survival of the entire plant. Many plant species have evolved different mechanisms to withstand the harmful effects of salt-induced toxicity on their chloroplasts and its machinery. The differences depend on the plant species and growth stage and can be quite different between salt-sensitive (glycophyte) and salt-tolerant (halophyte) plants. Salt stress tolerance is a complex trait, and many aspects of salt tolerance in plants are not entirely clear yet. In this review, we discuss the different mechanisms of salt stress tolerance in plants with a special focus on chloroplast structure and its functions, including the underlying differences between glycophytes and halophytes.

Spatial and seasonal metal variation, bioaccumulation and biomonitoring potential of halophytes from littoral zones of the Karachi Coast.

Coastal wetlands primarily serve as natural sinks of trace metals and their importance for phytoremediation is well known at global level. There are some reports on trace metal availability in soil sediments of the Karachi coast but studies on accumulation and translocation to harvestable (Shoot) parts among halophytes of the littoral zones have not been conducted. Hence, phytoremediation potential of six naturally existing halophytes (Aeluropus lagopoides, Arthrocnemum macrostachyum, Atriplex stocksii, Avicennia marina, Cressa cretica and Suaeda fruticosa) was assessed for cleaning metal (Mn, Zn, Pb and Cr) polluted soils of the Karachi coast. Seasonal (winters, pre and post-monsoon summers) and spatial (three littoral zones: viz., site - I: Sandspit, site - II: Do-Dariya/Clifton and site - III: Korangi creek) variations in soil and plant metals of the Karachi coast were studied. Soil Zn, Pb and Cr were generally higher in winters, Mn and organic matter in summers (7-11%) while pH values ranged between 7.15 and 7.5 in all seasons at site - III. All tested species had potential for cleaning Pb through their harvestable part (shoots) with A. stocksii as prominent candidate (16 mg kg-1) at site - I. Cressa cretica emerged as exclusive candidate for Zn phytoremediation (96 mg kg-1) at site - I, while S. fruticosa, A. macrostachyum and A. lagopoides showed bioaccumulation in pre-monsoon summers at site II. Aeluropus lagopoides with higher Mn in post monsoon summers (62 at site - III and 53 mg kg-1 at site - II) and Cr (7.1 mg kg-1 at site - II and 14 mg kg-1 at site - III) appeared exclusive bioindicator with potential of for cleaning all metals (Mn, Zn, Pb and Cr) at different sites. Metal bioaccumulation at study sites appeared species specific and varied seasonally among tested halophytes.

Comparative assessment of heavy metal accumulation and bio-indication in coastal dune halophytes.

This study aimed at assessing heavy metals (Fe, Mn, Zn, Cu, Cr and Pb) in four perennial halophytes (viz. Heliotropium bacciferum, Halopyrum mucronatum, Ipomoea pes-caprae and Salsola imbricata) growing at two sites on the Karachi coast. Site - II, closer to the Industrial area had higher bioavailability as well as translocation factor (TF) for most of the heavy metals and Na+ where soil sediments had lower pH (approximately 7.5), higher salinity (EC) and organic matter (OM). Site - I which was far from Industrial area had comparatively higher bio-concentration factor (BCF) and lower TF for metal ions and soil pH of 8.1-9. Metal accumulation in plants was both site and species specific. Extractable concentration of shoot Pb in all tested halophytes was above normal of the threshold values (i.e., >0.3 mg kg-1) while Mn (<50 mg kg-1) and Cu (<40 mg kg-1) were within permissible limits. Salsola imbricata had highest Na+ at both sites (site - I = 73; site - II = 98 mg kg-1) with and 10 mg kg-1 extractable shoot Pb at site - I. Ipomea pes-caprae also accumulated shoot Pb higher than normal (site - I = 3.3; site - II = 0.8 mg kg-1) with lowest Na+ content. Heliotropium bacciferum had higher extractable Pb (site - I = 10.5; II = 2.75) with >20 mg kg-1 Na+ in shoot while maintaining > 1 TF for Pb, Cu, Mn and Zn at site - I and all tested metals at site - II. Halopyrum mucronatum had highest shoot Fe (644 mg kg-1), Zn (63 mg kg-1) and Cr (9.2 mg kg-1) at site - II and above threshold values of Pb at both sites (site - I = 8.2; site - II = 2.5 mg kg-1) which makes this species an ideal bio-indicator candidate while other species could be potentially used for Pb phytoremediation.

Excreting and non-excreting grasses exhibit different salt resistance strategies.

The combination of traits that makes a plant successful under saline conditions varies with the type of plant and its interaction with the environmental conditions. Knowledge about the contribution of these traits towards salt resistance in grasses has great potential for improving the salt resistance of conventional crops. We attempted to identify differential adaptive response patterns of salt-excreting versus non-excreting grasses. More specifically, we studied the growth, osmotic, ionic and nutrient (carbon/nitrogen) relations of two salt-excreting (Aeluropus lagopoides and Sporobolus tremulus) and two non-excreting (Paspalum paspalodes and Paspalidium geminatum) perennial C4 grasses under non-saline and saline (0, 200 and 400 mM NaCl) conditions. Growth and relative growth rate decreased under saline conditions in the order P. geminatum > S. tremulus = A. lagopoides > P. paspalodes. The root-to-shoot biomass allocation was unaffected in salt-excreting grasses, increased in P. paspalodes but decreased in P. geminatum. Salt-excreting grasses had a higher shoot/root Na(+) ratio than non-excreting grasses. K(+), Ca(2+) and Mg(2+) homoeostasis remained undisturbed among test grasses possibly through improved ion selectivity with rising substrate salinity. Salt-excreting grasses increased leaf succulence, decreased ψs and xylem pressure potential, and accumulated proline and glycinebetaine with increasing salinity. Higher salt resistance of P. paspalodes could be attributed to lower Na(+) uptake, higher nitrogen-use efficiency and higher water-use efficiency among the test species. However, P. geminatum was unable to cope with salt-induced physiological drought. More information is required to adequately document the differential strategies of salt resistance in salt-excreting and non-excreting grasses.

The influence of genes regulating transmembrane transport of Na+ on the salt resistance of Aeluropus lagopoides.

Plantlets of Aeluropus lagopoides (Linn.) Trin. Ex Thw. were grown at different NaCl concentrations (26, 167, 373 and 747mM) for 3, 7 and 15 days; their growth, osmotic adjustment, gas exchange, ion compartmentalisation and expression of various genes related to Na+ flux was studied. Plantlets showed optimal growth in non-saline (control; 26mM NaCl) solutions, whereas CO2/H2O gas exchange, leaf water concentration and water use efficiency decreased under all salinity treatments, accompanied by increased leaf senescence, root ash, sodium content and leaf osmolality. A decrease in malondialdehyde (MDA) content with time was correlated with Na+ accumulation in the leaf apoplast and a concomitant increase in Na+ secretion rate. A. lagopoides accumulated a higher concentration of Na+ in root than in leaf vacuoles, corresponding with higher expression of V-NHX and lower expression of PM-NHX in root than leaf tissue. It appears that V-ATPase plays a vital role during Na+ transport by producing an electromotive force, driving ion transport. Leaf calcium increased with increasing salinity, with more rapid accumulation at high salinity than at low salinity, indicating a possible involvement of Ca2+ in maintaining K+:Na+ ratio. Our results suggest that A. lagopoides successfully compartmentalised Na+ at salinities up to 373mM NaCl by upregulating the gene expression of membrane linked transport proteins (V-NHX and PM-NHX). At higher salinity (747mM NaCl), a reduction in the expression of V-NHX and PM-NHX in leaves without any change in the rate of salt secretion, is a possible cause of the toxicity of NaCl.