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Australian saltbush (Atriplex spp.) survive in exceptionally saline environments and are often used for pasture in semi-arid areas. To investigate the impact of salinity on saltbush root morphology and root exudates, three Australian native saltbush species (Atriplex nummularia , Atriplex amnicola , and Atriplex vesicaria ) were grown in vitro in optimised sterile, semi-hydroponic systems in media supplemented with different concentrations of salt (NaCl). Histological stains and chromatographic techniques were used to characterise the root apical meristem (RAM) type and root exudate composition of the saltbush seedlings. We report that saltbush species have closed-type RAMs, which release border-like cells (BLCs). Monosaccharide content, including glucose and fructose, in the root mucilage of saltbush was found to be uniquely low, suggesting that saltbush may minimise carbon release in polysaccharides of root exudates. Root mucilage also contained notable levels of salt, plus increasing levels of unidentified compounds at peak salinity. Un-esterified homogalacturonan, xyloglucan, and arabinogalactan proteins between and on the surface of BLCs may aid intercellular adhesion. At the highest salinity levels, root cap morphology was altered but root:shoot ratio remained consistent. While questions remain about the identity of some components in saltbush root mucilage other than the key monosaccharides, this new information about root cap morphology and cell surface polysaccharides provides avenues for future research.

Atriplex crassifolia (A. crassifolia) is a locally occurring member of Chenopodiaceae family that has been used in folk medicine for the treatment of joint pain and inflammation. The present study was focused to determine the analgesic and anti-inflammatory potential of the plant. n-hexane (ACNH) and methanol (ACM) extracts of A. crassifolia were evaluated for in vitro anti-inflammatory potential using protein denaturation inhibition assay. In vivo anti-inflammatory potential was determined by oral administration of 250, 500, and 1000 mg/kg/day of extracts against carrageenan and formalin-induced paw edema models. Inflammatory mediators such as TNF-α, IL-10, IL-1β, NF-kB, IL-4, and IL-6 were estimated in blood samples of animals subjected to formalin model of inflammation. Analgesic activity was determined using acetic acid-induced writhing and tail flick assay model. Phytochemical profiling was done by GC-mass spectrophotometer. The results of in vitro anti-inflammatory activity revealed that both ACNH and ACM displayed eminent inhibition of protein denaturation in concentration-dependent manner. In acute in vivo carrageenan-induced paw edema model, both extracts reduced inflammation at 5th and 6th hour of study (p < 0.05). A. crassifolia extracts exhibited significant inhibition against formalin-induced inflammation with maximum effect at 1000 mg/kg. ACNH and ACM significantly augmented the inflammatory mediators (p < 0.05). Levels of TNF-α, IL-6, IL-1β, and NF-kB were reduced, while those of IL-4 and IL-10 were upregulated. ACNH displayed maximum analgesic effect at 1000 mg/kg, while ACM showed potent activity at 500 and 1000 mg/kg. The extracts restored the CBC, TLC and CRP toward normal. GC-MS analysis revealed the presence of compounds like n-hexadecanoic acid, Phytol, (9E,11E)-octadecadienoic acid, 2-hydroxy-1-(hydroxymethyl) ethyl ester, 1-hexacosene, vitamin E, campesterol, stigmasterol, gamma sitosterol in both extracts. These compounds have been reported to suppress inflammation by inhibiting inflammatory cytokines. The current study concludes that A. crassifolia possesses significant anti-nociceptive and anti-inflammatory potential owing to the presence of phytochemicals., (© 2024. The Author(s), under exclusive licence to Springer Nature Switzerland AG.)

Background: The plant kingdom has long been considered a valuable source for therapeutic agents, however, some plant species still untapped and need to be phytochemically and biologically explored. Although several Atriplex species have been investigated in depth, A. leucoclada, a halophytic plant native to Saudi Arabian desert, remains to be explored for its phytochemical content and biological potentials. Herein, the current study investigated the metabolic content and the anti-inflammatory potential of A. leucoclada., Methods: Powdered aerial parts of the plant were defatted with n-hexane then the defatted powder was extracted with 80% methanol. n-Hexane extract (ATH) was analyzed using GC-MS, while the defatted extract (ATD) was subjected to different chromatographic methods to isolate the major phytoconstituents. The structures of the purified compounds were elucidated using different spectroscopic methods including advanced NMR techniques. Anti-inflammatory activity of both extracts against COX-1 and COX-2 enzymes were examined in vitro. Molecular docking of the identified compounds into the active sites of COX-1 and COX-2 enzymes was conducted using pdb entries 6Y3C and 5IKV, respectively., Results: Phytochemical investigation of ATD extract led to purification and identification of nine compounds. Interestingly, all the compounds, except for 20-hydroxy ecdysone (1), are reported for the first time from A. leucoclada, also luteolin (6) and pallidol (8) are isolated for the first time from genus Atriplex. Inhibitory activity of ATD and ATH extracts against COX-1 and COX-2 enzymes revealed concentration dependent activity of both fractions with IC 50 41.22, 14.40 μg/ml for ATD and 16.74 and 5.96 μg/ml for ATH against COX-1 and COX-2, respectively. Both extracts displayed selectivity indices of 2.86 and 2.80, respectively as compared to 2.56 for Ibuprofen indicating a promising selectivity towards COX-2. Molecular docking study supported in vitro testing results, where purified metabolites showed binding affinity scores ranged from -9 to -6.4 and -8.5 to -6.6 kcal/mol for COX-1 and 2, respectively, in addition the binding energies of GC-MS detected compounds ranged from -8.9 to -5.5 and -8.3 to -5.1 kcal/mol for COX-1 and 2, respectively as compared to Ibuprofen (-6.9 and -7.5 kcal/mol, respectively), indicating high binding affinities of most of the compounds. Analysis of the binding orientations revealed variable binding patterns depending on the nature of the compounds. Our study suggested A. leucoclada as a generous source for anti-inflammatory agents., (© 2023. The Author(s).)

Dehydrins form the group II LEA protein family and are known to play multiple roles in plant stress tolerance and enzyme protection. They harbor a variable number of conserved lysine rich motifs (K-segments) and may also contain three additional conserved motifs (Y-, F- and S-segments). In this work, we report the isolation and characterization of an FSK 2 -type dehydrin from the halophytic species Atriplex halimus, which we designate as AhDHN1. In silico analysis of the protein sequence revealed that AhDHN1 contains large number of hydrophilic residues, and is predicted to be intrinsically disordered. In addition, it has an FSK 2 architecture with one F-segment, one S-segment, and two K-segments. The expression analysis showed that the AhDHN1 transcript is induced by salt and water stress treatments in the leaves of Atriplex seedlings. Moreover, circular dichroism spectrum performed on recombinant AhDHN1 showed that the dehydrin lacks any secondary structure, confirming its intrinsic disorder nature. However, there is a gain of α-helicity in the presence of membrane-like SDS micelles. In vitro assays revealed that AhDHN1 is able to effectively protect enzymatic activity of the lactate dehydrogenase against cold, heat and dehydration stresses. Our findings strongly suggest that AhDHN1 can be involved in the adaptation mechanisms of halophytes to adverse environments., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

Key Message: The salt-tolerance of transgenic soybean cleared for environmental release was improved by stable over-expression of AhBADH gene from Atriplex hortensis, which was demonstrated through molecular analysis and field experiments. An effective strategy for increasing the productivity of major crops under salt stress conditions is the development of transgenics that harbor genes responsible for salinity tolerance. Betaine aldehyde dehydrogenase (BADH) is a key enzyme involved in the biosynthesis of the osmoprotectant, glycine betaine (GB), and osmotic balance in plants, and several plants transformed with BADH gene have shown significant improvements in salt tolerance. However, very few field-tested transgenic cultivars have been reported, as most of the transgenic studies are limited to laboratory or green house experiments. In this study, we demonstrated through field experiments that AhBADH from Atriplex hortensis confers salt tolerance when transformed into soybean (Glycine max L.). AhBADH was successfully introduced into soybean by Agrobacterium mediated transformation. A total of 256 transgenic plants were obtained, out of which 47 lines showed significant enhancement of salt tolerance compared to non-transgenic control plants. Molecular analyses of the transgenic line TL2 and TL7 with the highest salt tolerance exhibited stable inheritance and expression of AhBADH in progenies with a single copy insertion. TL1, TL2 and TL7 exhibited stable enhanced salt tolerance and improved agronomic traits when subjected to 300mM NaCl treatment. Currently, the transgenic line TL2 and TL7 with stable enhanced salt tolerance, which have been cleared for environmental release, are under biosafety assessment. TL 2 and TL7 stably expressing AhBADH could then be applied in commercial breeding experiments to genetically improve salt tolerance in soybean., (© 2023. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

Mining activities have led to serious environmental (soil erosion, degradation of vegetation, and groundwater contamination) and human health (musculoskeletal problems, diarrheal conditions, and chronic diseases) issues at desert mining areas in northwest China. Native plant species grown naturally in desert regions show a unique tolerance to arid and semiarid conditions and are potential candidates for soil phytoremediation. Here, an ex situ experiment involving pot planting of seedlings of three native plant species ( Suaeda glauca , Artemisia desertorum , and Atriplex canescens ) was designed to explore their phytoremediation potential and the underlying physiological mechanism. For Zn and Cu, the three plants were all with a biological accumulation coefficient (BAC) greater than 1. For Cd, Ni, and Pb, Atriplex canescens had the highest bioaccumulation concentrations (521.52, 862.23, and 1734.59 mg/kg), with BAC values (1.06, 1.30, 1.25) greater than 1, which indicates that Atriplex canescens could be a broad-spectrum metal extraction plant. Physiological analysis (antioxidation, extracellular secretions, photosynthesis, and hydraulics) showed that the three desert plants exploited their unique strategy to protect against the stress of complex metals in soils. Moreover, the second growing period was the main heavy metal accumulation and extraction stage concomitant with highest water use efficiency (iWUE). Taken together, the three desert plants exhibited the potent heavy metal extraction ability and physiological and ecological adaptability to a harsh polluted environment in arid desert areas, providing potential resources for the bioremediation of metal-contaminated soils in an arid and semiarid desert environment.

This experiment aimed to investigate the effects of substituting barley grain with discarded dates on intake and digestibility, nitrogen (N) utilization and fermentation characteristics of sheep fed Atriplex hay-based diets. Four Barki sheep (50.9 ± 2.93 kg of body weight (BW)) fitted with rumen cannula were used in a 4 × 4 Latin Square design with 23-day periods. Four isonitrogenous (~140 g crude protein (CP)/kg of dry matter (DM)) experimental diets were formulated with Atriplex hay, and dietary sugar content was increased by replacing barley grain with discarded dates (the proportions of discarded dates in the diet were 0, 93, 187 and 280 g/kg DM). Nutrient intakes were not influenced by diet. Organic matter, CP and neutral detergent fibre digestibility increased (p < 0.05) linearly and DM digestibility tended to increase with increased dates inclusion. Ruminal pH and molar proportion of butyrate increased (p < 0.05) linearly with increasing levels of dates inclusion, whereas ruminal ammonia-N decreased (p < 0.05) linearly. Total volatile fatty acid concentration was unaffected. Microbial protein synthesis (MCP) increased linearly and efficiency of microbial protein synthesis tended to increase as discarded dates increased. Total N intake was not affected by diets whereas, urinary N, urea-N and total N execrations linearly decreased (p < 0.05) with higher amounts of dates in the diets. Blood urea-N tended to decrease with increasing levels of dates. It could be concluded that replacing 100% barley grain with discarded dates as a source of soluble carbohydrates can enhance the utilization of N in Atriplex hay-based diet and consequently improve digestibility and MCP., (© 2021 Wiley-VCH GmbH.)

The chemical characteristics of mine tailings, organic amendments (doses), and plants are the critical factors that must be evaluated and monitored to ensure the sustainability of phytostabilization. The aim of this study was to evaluate the mobility of copper (Cu) in mine tailings (MT) of the Zone Central of Chile to which commercial humic substances were added, examining their effect on the uptake of Atriplex halimus. Two commercial humic substances (HS1 and HS2) extracted from leonardite (highly oxidized lignite), of different pH and total organic carbon, were evaluated by adsorption curve for Cu. In columns, soluble Cu, pH, and electrical conductivity in leachates were evaluated for MT, MT + HS1, and MT + HS2, and HS1 and HS2 in doses of 120 mg kg -1 . In pot assay, seeds were germinated directly in MT and cultivated for 140 days with the addition of HS2 in 120 and 240 mg kg -1 . Mine tailing presents high concentration of Cu (2016 ± 223 mg kg -1 , pH 6.3 ± 0.1). The results of sequential extraction indicate that Cu is associated with the sulfide fraction of low risk of mobility. The amount of Cu sorbed by HS1 was higher than that sorbed by HS2, and both humic substances showing better fit to the Freundlich than Langmuir model. Lixiviation of Cu was significantly lower in MT + HS1 (0.166 ± 0.043 mg kg -1 ) and MT + HS2 (0.157 ± 0.018 mg kg -1 ) than in MT (0.251 ± 0.052 mg kg -1 ). Copper concentration in plants reached 185.8 ± 37.8 mg kg -1 in the roots and 32.6 ± 7.4 mg kg -1 in the aerial parts cultivated in MT without effect of the humic substance addition in Cu uptake nor growth. Copper concentrations in the aerial parts were adjusted to sufficient or normal levels in plant. A good management of mine tailings through phytostabilization could consider an adequate mixture of humic substances (to avoid leaching of metals) and an organic amendment that provides essential nutrients and increases biomass generation.

Whole plants and hypocotyl-derived calli of the halophyte plant species Atriplex atacamensis were exposed to 50 μM arsenate (As(V)) or 50 μM arsenite (As(III)). At the whole plant level, As(III) was more toxic than As(V): it reduced plant growth, stomatal conductance, photosystem II efficiency while As(V) did not. In roots, As accumulated to higher level in response to As(III) than in response to As(V). Within root tissues, both arsenate and arsenite were identified in response to each treatment suggesting that oxidation of As(III) may occur. More than 40% of As was bound to the cell wall in the roots of As(V)-treated plants while this proportion strongly decreased in As(III)-treated ones. In leaves, total As and the proportion of As bound to the cell wall were similar in response to As(V) and As(III). Non-protein thiol increased to higher extent in response to As(V) than in response to As(III) while ethylene synthesis was increased in As(III)-treated plants only. Polyamine profile was modified in a contrasting way in response to As(V) and As(III). At the callus level, As(V) and As(III) 50 μM did not reduce growth despite an important As accumulation within tissues. Calli exposed to 50 μM As did not increase the endogenous non-protein thiol. In contrast to the whole plants, arsenite was not more toxic than arsenate at the cell line level and As(V)-treated calli produced higher amounts of ethylene and malondialdehyde. A very high dose of As(V) (1000 μM) strongly reduced callus growth and lead to non-protein thiols accumulation. It is concluded that As(III) was more toxic than As(V) at the plant level but not at the cellular level and that differential toxicity was not fully explained by speciation of accumulated As. Arsenic resistance in A. atacamensis exhibited a cellular component which however did not reflect the behavior of whole plant when exposed to As(V) or As(III).

Mine tailings have adverse chemical and physical conditions, including high concentrations of metals and salts, low organic matter content, and unbalanced rates of nutrients which limit the development of vegetation. A large scale field experiment was conducted to reclaim a tailing pond by triggering the growth of native species by spontaneous colonization by tilling (TL) the tailing pond surface and using marble waste (CaCO 3 ; MW), pig slurry (PS) and their combination (MW + PS) as soil amendments. Soil physicochemical properties and water and DTPA extractable metal concentrations of bulk and rhizosphere soils were analyzed after five year from the application of the treatments. In addition, plants of Atriplex halimus from each treatment were collected and metals in roots, leaves and stems analyzed. Before amendments application, the studied pond showed a neutral pH, high salinity and a moderate organic carbon content. After five years, the pH value was significantly increased only in MW plot. The results showed significant increases of DTPA-extractable Zn in MW and MW + PS plots, Pb in all treatments except MW plot, Cd only in PS plot, and Cu only in MW + PS plot. A. halimus was the most dominant species, growing spontaneously in all plots, with lower vegetation cover in CT and MW plots, 6% and 2% respectively. Application of MW increased leaf Pb accumulation by 2.5-fold and Cd by 55%, when compared to the CT. The high initial salinity and probable substitution of metals by Ca 2+ on exchangeable surfaces of soil particles may be the reasons for higher uptake of metals in MW plot when compared to the other plots. Although this plant is widely utilized in contaminated sites for phytostabilization purposes, it may absorb and translocate high concentrations of metals to the aboveground tissues in saline contaminated sites., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

The survival of Atriplex nummularia plants in saline environments is possible mainly due to the presence of salt-accumulating epidermal vesicles. Commonly, destructive methods, such as plant material maceration and subsequent reading in osmometers, are employed in studies on water relations and osmotic adjustment and are inconvenient due to their underestimation of the total water potential inside the cells, which can cause overestimation of an osmotic adjustment that is not present. As a result, methods that preserve leaf structure, such as pressure-volume (P-V) curves, which take into consideration only the salts that compose the symplastic solution, are more adequate. Thus, the main objectives of this study were to evaluate the effect of determination methods of osmotic potential (Ψ o ) in Atriplex nummularia through destructive and leaf structure-preserving techniques and to determine the water relations of the species under increasing NaCl concentrations. Plants were subjected to daily irrigations, maintaining soil moisture at 80% of field capacity, with solutions of increasing NaCl concentration (0, 0.05, 0.1, 0.2, 0.25 and 0.3 M) for 84 days. Water potential, osmotic potential and osmotic adjustment were determined. In addition, P-V curves were constructed using pressure chambers. Water and osmotic potentials decreased linearly with increasing NaCl concentration in the irrigation solution. The main discrepancies observed were related to the osmotic adjustments determined through maceration and P-V curves. Based on the present research, it was possible to conclude that in studies with species that have salt-accumulating vesicles in the epidermis, such as the plants in the genus Atriplex, constructing P-V curves is more adequate than destructive methods., (Copyright © 2018 Elsevier Masson SAS. All rights reserved.)

Alfin-like (AL) is a small plant-specific gene family with prominent roles in root growth and abiotic stress response. Here, we aimed to identify novel stress tolerance AL genes from the stress-tolerant species Atriplex hortensis. Totally, we isolated four AhAL genes, all encoding nuclear-localized proteins with cis-element-binding and transrepression activities. Constitutive expression of AhAL1 in Arabidopsis facilitated plants to survive under saline condition, while expressing anyone of the other three AhAL genes led to salt-hypersensitive response, indicating functional divergence of AhAL family. AhAL1 also conferred enhanced drought tolerance, as judged from enhanced survival, improved growth, decreased malonaldehyde (MDA) content and reduced water loss in AhAL1-expressing plants compared to WT. In addition, abscisic acid (ABA)-mediated stomatal closure and inhibition of seed germination and primary root elongation were enhanced in AhAL1-transgenic plants. Further analysis demonstrated that AhAL1 could bind to promoter regions of GRF7, DREB1C and several group-A PP2C genes and repress their expression. Correspondingly, the expression levels of positive stress regulator genes DREB1A, DREB2A and three ABFs were all increased in AhAL1-expressing plants. Based on these results, AhAL1 was identified as a novel candidate gene for improving abiotic stress tolerance of crop plants.

The mushroom cultivation industry produces a huge amount of spent mushroom compost (SMC), a wide world agricultural organic waste which causes serious environmental problems. However, this cheap organic waste could be useful in the remediation of contaminated soils. The aim of this work was to assess the potential of SMC in combination with the native shrub Atriplex halimus, to phytoremediate two mine soils contaminated with Cd, Pb and Cu. Firstly, to minimize metal availability in the soil, the optimal doses of SMC were determined. Secondly, a phytoremediation assay in greenhouse conditions was carried out to test the effects of A. halimus in combination with SMC at different doses. The results showed the ability of SMC to reduce soil acidity, the mobility of the metals and the enhancement of A. halimus growth. SMC promoted metal immobilization in the root of A. halimus and decreased the translocation from the roots to the shoots. The combination of SMC amendment and A. halimus produced phytostabilization of the metals in the mine soils assayed. In conclusion, SMC represents an adequate organic solid waste which in combination with A. halimus can reduce the adverse impact caused by the high mobility of metals in acid mine soils.

Most arsenic in surface soil and water exists primarily in its oxidized form, as arsenate (As(V); AsO 4 3- ), which is an analog of phosphate (PO 4 3- ). Arsenate can be taken up by phosphate transporters. Atriplex atacamensis Phil. is native to northern Chile (Atacama Desert), and this species can cope with high As concentrations and low P availability in its natural environment. To determine the impact of P on As accumulation and tolerance in A. atacamensis, the plants were cultivated in a hydroponic system under four treatments: no As(V) addition with 323µM phosphate (control); 1000µM As(V) addition with 323µM phosphate; no As(V) and no phosphate; 1000µM As(V) addition and no phosphate. Phosphate starvation decreased shoot fresh weight, while As(V) addition reduced stem and root fresh weights. Arsenate addition decreased the P concentrations in both roots and leaves, but to a lesser extent than for P starvation. Phosphorus starvation increased the As concentrations in roots, but decreased it in shoots, which suggests that P deficiency reduced As translocation from roots to shoots. Arsenate addition increased total glutathione, but P deficiency decreased oxidized and reduced glutathione in As(V)-treated plants. Arsenate also induced an increase in S accumulation and nonprotein thiol and ethylene synthesis, and a decrease in K concentrations, effects that were similar for the P-supplied and P-starved plants. In contrast, in As(V)-treated plants, P starvation dramatically decreased total soluble protein content and increased lipid peroxidation, compared to plants supplied with P. Phosphorus nutrition thus appears to be an important component of A. atacamensis response to As toxicity., (Copyright © 2017 Elsevier Inc. All rights reserved.)

An ErbB-3-binding protein gene AcEBP1, also known as proliferation-associated 2G4 gene (PA2G4s) belonging to the M24 superfamily, was obtained from the saltbush Atriplex canescens. Subcellular localization imaging showed the fusion protein AcEBP1-eGFP was located in the nucleus of epidermal cells in Nicotiana benthamiana. The AcEBP1 gene expression levels were up-regulated under salt, osmotic stress, and hormones treatment as revealed by qRT-PCR. Overexpression of AcEBP1 in Arabidopsis demonstrated that AcEBP1 was involved in root cell growth and stress responses (NaCl, osmotic stress, ABA, low temperature, and drought). These phenotypic data were correlated with the expression patterns of stress responsive genes and PR genes. The AcEBP1 transgenic Arabidopsis plants also displayed increased sensitivity under low temperature and evaluated resistance to drought stress. Together, these results demonstrate that AcEBP1 negatively affects cell growth and is a regulator under stress conditions., (Copyright © 2016 Elsevier Ireland Ltd. All rights reserved.)

Although the use of saltbush plants in metal phytoremediation is well known, there is little information about the impact of sugarcane vinasse (SCV) and EDTA on metal uptake. Heavily cadmium-polluted soil (38 mg kg(-1) Cd) was used in pot and incubation experiments to investigate the Cd phytoextraction potential of wavy saltbush (Atriplex undulata) and quail saltbush (Atriplex lentiformis). EDTA at rates of 3, 6, and 10 mM kg(-1) soil and SCV at rates of 7, 15, and 30 mL kg(-1) soil were added to the polluted soil. The application of EDTA significantly (P = 0.002) reduced the growth of saltbush plants; on the other hand, SCV improved the growth. Both EDTA and SCV increased the availability and root-to-shoot transfer of Cd. The plants of A. lentiformis grown on the soil amended with the highest rate of SCV were able to remove 20.4 % of the total soil Cd during a period of 9 months. Based on the obtained results, it may be concluded that A. lentiformis and sugarcane vinasse could be more effective in the phytoextraction of Cd from the polluted soils.

The phytochemical investigation of a Tunisian plant Atriplex portulacoides (Chenopodiaceae) led to the isolation of two new compounds designated as portulasoid (2) and septanoecdysone (3) along with the known 20-hydroxyecdysone (20HE) (1). Their chemical structures were elucidated on the basis of extensive spectroscopic methods including ES-HRMS, 1D and 2D-NMR. The isolated compounds were finally tested for their antioxidant activity by using DPPH, ABTS(+), Fe(3+) and catalase assays and also for their antibacterial and anticholinesterase activities., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

Atriplex gmelini is a halophyte and possesses bladder hairs on the leaf surface. It is also known to accumulate the osmoprotectant glycinebetaine (GB). However, it remains unclear whether GB and its biosynthetic enzyme choline monooxygenase (CMO) accumulate in the bladder hairs. Microscopic observation of young leaves showed many bladder hairs on their surfaces, but their total number decreased along with leaf maturity. Sodium Green fluorescent approach revealed Na(+) accumulation in bladder cells of young leaves when A. gmelini was grown at high salinity (250 mM NaCl). Due to fewer bladder hairs in mature leaves, Na(+) accumulation was mostly found in mesophyll cells of mature leaves under high salinity. GB accumulation was found at significant level in both bladder- and laminae-cells without any addition of NaCl and its content increased at high salinity. CMO was not found in bladder hairs or young leaf laminae. Instead, the CMO protein expression was observed in mature leaves and that showed increased accumulation with increasing concentration of NaCl. Furthermore, in situ hybridization experiments revealed the expression of a transporter gene for GB, AgBetT, in the bladder hairs. Based on these results, the synthesis and translocation of GB in A. gmelini were discussed., (Copyright © 2014 Elsevier GmbH. All rights reserved.)

A study was carried out to identify the mechanisms underlying stress caused by Cd and Pb accumulation in leaves of Atriplex halimus L. collected from habitats representing different kinds of pollution. Mean concentrations of Cd and Pb in aerial parts exceeded the critical levels in polluted plants as compared to reference plants. There were significant reduction in guiacol peroxidase, ascorbate peroxidase and glutathione content in most of polluted plants. The results showed increase in superoxide dismutase enzyme in all polluted plants. The significant increment in catalase enzyme, glutathione S-transeferase and ascorbic acid content were observed in most of polluted plants. Results of the nine differential expressed bands showed down regulation of NADH dehydrogenase and Sedoheptulose-bisphosphatase in polluted plants. In contrast, there were six regulated genes in highly polluted plants, representing transcription factors, membrane transporters and ROS detoxification. The transcription level of phytochelatin synthase showed a significant increase in all polluted plants, while heavy metal ATPase transporter expression significantly increased in some polluted plants. In conclusion, A. halimus may use two different strategies against Cd and Pb stress, in which the molecular and physiological features affords similar levels of Cd and Pb tolerance through binding, sequestration and the reduction of harmful effect of heavy metals.

Phytostabilization of heavy metals in contaminated soils should be subject to two conditions, the first is the choice plant must be able to stabilize heavy metals in soil, the second is the plant material which produced from the phytostabilization process must be safe and useful to avoid overload on environmental system. A field experiment was conducted out to evaluate the phytostabilization potential of two halophytes species (Atriplex lentiformis and Atriplex undulata). Compost at rates of 0, 15 and 30 ton ha(-1) was used to examine its role in plant growth and heavy metals uptake. The high rate of compost (30 ton ha(-1)) decreased zinc (Zn) concentrations in the leaves of A. lentiformis and A. undulata by 15.8 and 13.0%, while lead (Pb) in the leaves decreased by 37.6 and 35.2% respectively. Despite the extremely high total heavy metals concentrations in the studied soil, plants of Atriplex were able to grow and maintain shoots metals content below the toxic level and the produced plant materials had a high nutritive value compared to the conventional forage crops. Phosphorus (P) and chloride (Cl) in the roots of Atriplex plants play important function in heavy metals phytostabilization mechanism by the two halophytes plants.

Mine tailings in semiarid regions are highly susceptible to erosion and are sources of dust pollution and potential avenues of human exposure to toxic metals. One constraint to revegetation of tailings by phytostabilization is the absence of microbial communities critical for biogeochemical cycling of plant nutrients. The objective of this study was to evaluate specific genes as in situ indicators of biological soil response during phytoremediation. The abundance and activity of 16S rRNA, nifH, and amoA were monitored during a nine month phytostabilization study using buffalo grass and quailbush grown in compost-amended, metalliferous tailings. The compost amendment provided a greater than 5-log increase in bacterial abundance, and survival of this compost-inoculum was more stable in planted treatments. Despite increased abundance, the activity of the introduced community was low, and significant increases were not detected until six and nine months in quailbush, and unplanted compost and buffalo grass treatments, respectively. In addition, increased abundances of nitrogen-fixation (nifH) and ammonia-oxidizing (amoA) genes were observed in rhizospheres of buffalo grass and quailbush, respectively. Thus, plant establishment facilitated the short term stabilization of introduced bacterial biomass and supported the growth of two key nitrogen-cycling populations in compost-amended tailings.

PsbQ is an extrinsic subunit of the photosystem II in eukaryotic photosynthetic organisms. Numerous studies have demonstrated that PsbQ can stabilize the inorganic cofactors and enhance the oxygen release in PSII. The decrease of photosynthesis rate under salinity condition is normally attributed to the high concentration of injurious ions, such as Na(+) and Cl(-), which accumulate in the chloroplast and damage thylakoid membrane under salinity stress. In this study, AcPsbQ1 was isolated from a halophyte Atriplex canescens cDNA library. The AcPsbQ1 contains an open reading frame of 699 bp encoding a 233 amino acid protein. In order to investigate its function, AcPsbQ1 was cloned and transformed into Saccharomyces cerevisiae INVSc1. The heterologous expression of AcPsbQ1 in transgenic yeast significantly helped to increase the adapting and recovery ability of yeast cells under the salt and drought. Quantitative real-time PCR assay was performed to reveal the expression pattern of AcPsbQ1 under different abiotic stresses. On exposure to NaCl stress, the transcript level of AcPsbQ1 was significantly enhanced. AcPsbQ1 expression level was also up-regulated under drought stress. These results indicated that AcPsbQ1 might involve in the response to salt stress in A. canescens.

We detected absorption and transportation of ions in the leaves of Atriplex nummularia, Atriplex canescens and Lycium barbarum under NaHCO3 stress (300 mmol x L(-1)) by using atomic absorption spectrophotometry and non-invasive ion flux measurement. The results showed that leaves of the A. nummularia, A. canescens and L. barbarum exhibited a high capacity to induce the Na+ accumulation when compared with that of control. The higher the concentration of NaHCO3 treatment, the more Na+ accumulated in the leaves of the three plants under experimental condition. L. barbarum showed a higher Na+ efflux in the mesophyll cells, whereas A. nummularia and A. canescens showed a relative lower efflux. A lower K+ content and a higher Na+/K+ ratio were detected in leaves of A. nummularia and L. barbarum. However, a higher K+ content and a lower Na+/K+ ratio were seen in leaves of A. canescens. Due to induction of Ca2+ efflux under the NaHCO3 treatment, a lower Ca2+ content and a higher Na+/Ca2+ ratio were observed in L. barbarum. On the contrary, a higher Ca2+ influx was observed in A. nummularia and A. canescens. These results suggested that the three shrubs species had different Na+ segmentation strategies. The accumulation of Na+ inhibited Ca2+ absorption in leaves of L. barbarum, while in the A. nummularia and A. canescens, Ca2+ influx induced [Ca2+]cyt which preserved a less-depolarized PM and then inhibited K efflux. The maintaining of cellular K+/Na+ homeostasis in A. nummularia and A. canescens might be achieved by the induction of [Ca2+]cyt under the NaHCO3 treatment.

This study aims to investigate the behavior of Atriplex nummularia under field conditions, including its growth, periodic cuttings, salt extraction, and soil chemical properties monitored for 16 months. Three treatments were evaluated: soil cultivated with Atriplex pruned at 6 and 12 months after transplanting (MAT); soil cultivated with plants that were harvested only at the end of the experiment (16 MAT); and a control (uncultivated soil) with four replications. Soil samplings were taken at 0, 6, 12, and 16 MAT. The samples were taken at depths of 0-20, 20-40, 40-60, and 60-80 cm. Biometric variables for growth were monitored monthly. The shoot was divided into leaves, thin stems (< or = 3 mm diameter), and thick stems (> 3 mm diameter) to determine its content of Ca, Mg, Na, K, and Cl. We concluded that pruning regime for Atriplex was efficient mainly because it stimulated regrowth of less lignified material (leaves and stems < or = 3 mm). We found that elements extracted by plant tissue can be quantified accurately, making them valuable indicators of the efficiency of the recovery process. The use of the Atriplex is recommended because the the possibility of revegetating areas inhospitable to most species used in conventional farming.

To assess metal mobility in pruning waste and biosolids compost (pH 6.9 and total concentration of metals in milligram per kilogram of Cd 1.9, Cu 132, Fe 8,513, Mn 192, Pb 81, and Zn 313), shrubs species Atriplex halimus and Rosmarinus officinalis were transplanted in this substrate and irrigated with citric acid (4 g L(-1), pH 2.9) and nutrient solution daily for 60 days. Citric acid significantly increased the concentrations of soluble Mn and Fe in the nutrient substrate solution measured by suction probes, while other metals did not vary in concentration (Cu and Zn) or were not observed at detectable levels (Cd and Pb). In plants, citric acid significantly increased the concentrations of Cu (2.7 ± 0.1-3.3 ± 0.1 mg kg(-1)), Fe (49.2 ± 5.2-76.8 ± 6.8 mg kg(-1)), and Mn (7.2 ± 1.1-11.4 ± 0.7 mg kg(-1)) in leaves of R. officinalis, whereas the concentration of only Mn (25.4 ± 0.3-42.2 ± 2.9 mg kg(-1)) was increased in A. halimus. Increasing Fe and Mn solubility by citric acid addition indicates the possibility of using it to improve plant nutrition. The mobility of metals in this substrate was influenced for the concentration of the metal, the degree of humification of organic matter and its high Fe content.

The Pre-Andean area of Chile exhibits saline soils of volcanic origin naturally contaminated with arsenic (As), and we hypothesise that revegetation with resistant species may be a valid alternative for soil management in this area. Thus, the xerophytic and halophytic shrubs Atriplex halimus and Atriplex atacamensis were cultivated in containers for 90 days in Pre-Andean soil, As-soil, (111±19 mg As kg(-1), pH8.4±0.1) or control soil (12.7±1.1 mg As kg(-1), pH7.8±0.1) to evaluate As accumulation and resistance using stress bioindicators (chlorophylls, malondialdehyde (MDA) and total thiols). Sequential extraction of As-soil indicated that 52.3% of As was found in the most available fraction. The As distribution was significantly different between the species: A. halimus translocated the As to leaves, whilst A. atacamensis retained the As in roots. At 30 and 90 days, A. halimus showed similar As concentrations in the leaves (approximately 5.5 mg As kg(-1)), and As increased in stems and roots (up to 4.73 and 16.3 mg As kg(-1), respectively). In A. atacamensis, As concentration was lower (2.6 in leaves; 3.2 in stems and 6.9 in roots in mg As kg(-1)). Both species exhibited a high concentration of B in leaves (362-389 mg kg(-1)). If the plants are used for animal feed, it should be considered that A. halimus accumulates higher concentration of As and B in the leaves than A. atacamensis. Neither plant growth nor stress bioindicators were negatively affected by the high levels of available As, with the exception of MDA in the leaves of A. halimus. The results indicate that these plants resist contamination by arsenic, accumulating mainly the metalloid in the roots and can be recommended to generate plant cover in As-contaminated soils in the Pre-Andean region, under saline conditions controlled, preventing the dispersion of this metalloid via wind and leaching., (Copyright © 2013 Elsevier B.V. All rights reserved.)

Tolerance of plants to elevated concentrations of heavy metals in growth media and in its tissues leads to high degrees of metal bioaccumulation, which may pose a risk for humans and animals alike. Therefore, bio-accumulating plants need thorough evaluation from an environmental health point of view. A glasshouse experiment concerning the xerohalophyte Atriplex halimus was carried out to determine its tolerance and capacity to accumulate copper. We investigated the effect of Cu from 0 to 30 mmol l(-1) on the growth, photosynthetic apparatus and nutrient uptake of A. halimus by measuring gas exchange, chlorophyll fluorescence and photoinhibition. We also determined total Cu, sodium, potassium, magnesium, phosphorous, and nitrogen content in the plant. Our results indicated that A. halimus presented a high resistance to Cu-induced stress, since the plants were able to survive at concentrations higher than 15 mmol l(-1) Cu. However, this capacity was not reflected in its ability to accumulate and tolerate greater amounts of Cu in its tissues, since clear phytotoxicity symptoms were detected at tissue concentrations greater than 38 mg kg(-1) Cu. Thus, Cu increment caused a reduction in A. halimus growth, which was related to a decrease in net photosynthetic rate. This reduction was associated with the adverse effect of Cu on the photochemical apparatus and the reduction in the absorption of essential nutrients. The high resistance of A. halimus was largely related with the capacity of this species to avoid the absorption of great amounts of Cu. For all the above reasons, A. halimus could have the characteristics of a Cu-exclusion plant., (Copyright © 2012 Elsevier Inc. All rights reserved.)

Plant hydraulic characteristics were studied in diploid, tetraploid and hexaploid cytotypes of Atriplex canescens (Chenopodiaceae) to investigate the potential physiological basis underlying the intraspecific habitat differentiation among plants of different ploidy levels. Populations of A. canescens from different habitats of the Chihuahuan Desert (New Mexico, USA) were analyzed using flow cytometry to determine ploidy levels. Traits related to xylem water transport efficiency and safety against drought-induced hydraulic failure were measured in both stems and leaves. At the stem level, cytotypes of higher ploidy showed consistently lower leaf-specific hydraulic conductivity but greater resistance to drought-induced loss of hydraulic conductivity. At the leaf level, comparisons in hydraulics between cytotypes did not show a consistent pattern, but exhibited high plasticity to proximal environmental conditions related to soil water availability. The results suggest that a trade-off between stem hydraulic efficiency and safety across ploidy levels underlies niche differentiation among different cytotypes of A. canescens. Polyploidization may have been facilitated by environmental heterogeneity related to water availability, and variation in water-related physiology found in the present study suggests an important functional basis for the niche differentiation and coexistence of A. canescens cytotypes in desert environments., (No claim to original US government works. New Phytologist © 2012 New Phytologist Trust.)

Suitable plant species are able to accumulate heavy metals and to produce biomass useful for non-food purposes. In this study, three endemic Mediterranean plant species, Atriplex halimus, Portulaca oleracea and Medicago lupulina were grown hydroponically to assess their potential use in phytoremediation and biomass production. The experiment was carried out in a growth chamber using half strength Hoagland's solutions separately spiked with 5 concentrations of Pb and Zn (5, 10, 25, 50, and 100 mg L(-1)), and 3 concentrations of Ni (1, 2 and 5 mg L(-1)). Shoot and root biomass were determined and analyzed for their metals contents. A. halimus and M. lupulina gave high shoot biomass with relatively low metal translocation to the above ground parts. Metals uptake was a function of both metals and plant species. It is worth noting that M. lupulina was the only tested plant able to grow in treatment Pb50 and to accumulate significant amount of metal in roots. Plant metal uptake efficiency ranked as follows: A. halimus > M. lupulina > P. oleracea. Due to its high biomass production and the relatively high roots metal contents, A. halimus and M. lupulina could be successfully used in phytoremediation, and in phytostabilization, in particular.

The n-BuOH-soluble fraction of the MeOH-CH2Cl2 (1:1) extract of the aerial parts of Egyptian Atriplex halimus L. yielded two new flavonol glycosides, designated as atriplexoside A (1) [3'-O-methylquercetin-4'-O-beta-apiofuranoside-3-O-(6"-O-alpha-rhamnopyranosyl-beta-glucospyranoside)] and atriplexoside B (2) [3'-O-methylquercetin-4'-O-(5"-O-beta-xylopyranosyl-beta-apiofuranoside)-3-O-(6"-O-alpha-rhamnopyranosyl-beta-glucopyranoside)], together with six known compounds: two phenolic glucosides (3, 4), one ecdysteroid (5), one megastigmane (6) and two methoxylated flavonoid glycosides (7, 8). The structures of the compounds were elucidated by detailed spectroscopic analysis, including HR-ESI-MS and 2D-NMR spectroscopic data. DPPH radical scavenging, antileishmanial and anti-multidrug resistance activities were investigated using the n-BuOH-soluble fraction as well as the isolated compounds. Compound 8 (5-O-methylquercetin-3-O-(6"-O-alpha-rhamnopyranosyl-beta-glucopyranoside) presented marked DPPH radical scavenging, weak antileishmanial and anti-multidrug resistance activity while the other tested compounds showed weaker activities.

Atriplex halimus L. is a xero-halophyte species widespread in the Mediterranean basin. The tolerance to water stress and high salinity of two Atriplex populations from semi-arid (Djelfa) and arid saline (Laghouat) Algerian regions has been investigated in relation with organic solutes and antioxidant systems. Whereas no noticeable difference was observed between the two populations under water stress resulting from withholding watering or PEG treatment, Laghouat plants display significantly higher fresh and dry weights than Djelfa plants when exposed to high salinity. At 300mM NaCl, Laghouat plants exhibit higher concentrations in Na(+), proline and quaternary ammonium compounds, and a higher catalase activity than Djelfa plants. We then analysed the involvement of recently characterized plastidial thiol reductases, peroxiredoxins (Prxs) and methionine sulphoxide reductases (MSRs), key enzymes scavenging organic peroxides and repairing oxidized proteins, respectively. Upon salt treatment (300mM NaCl), we observed higher amounts of PrxQ and over-oxidized 2-Cys Prx in Laghouat than in Djelfa. An increased abundance of plastidial MSRA and a higher total MSR activity were also noticed in Laghouat plants treated with 300mM NaCl compared to Djelfa ones. We propose that mechanisms based on organic solutes and antioxidant enzymes like catalases, peroxiredoxins and MSRs party underlie the better tolerance of the Laghouat population to high salt., (Copyright © 2012 Elsevier GmbH. All rights reserved.)

The accumulation of significant pools of sulfidic sediments in inland wetlands and creeks is an emerging risk for the management of inland waterways. We used replicated plot trials to appraise the viability of various strategies for neutralizing oxidized, acidified sulfidic sediments in a highly degraded wetland. Of the twenty different treatments trialed only addition of calcium hydroxide or calcium carbonate, burning of wood, and planting of Phragmites australis, Typha domingensis and Atriplex nummularia into beds prepared with CaCO3 or P. australis and T. domingensis into beds of sediment and mulch, decreased total actual acidity (TAA) in the top 5 cm of sediment in the first two weeks following treatment. Only the calcium hydroxide treatments and planting of P. australis, T. domingensis and A. nummularia into beds prepared with CaCO3 decreased TAA for a longer period of time (6 months). None of the treatments, except the planting of P. australis into beds prepared with lime, decreased TAA in the 5-30 cm layer of sediments. Therefore, the only effective treatment appears to be the application of highly alkaline ameliorants which need to be transported to the site. A survey of the wetland was undertaken to estimate the total amount of actual and potential acidity stored in the wetland's sediment and overlying water and showed that up to 1200 tonnes of calcium carbonate would be required to neutralise all of the actual and potential acidity in the 10 ha wetland. However, neutralisation of the remaining water in the wetland (about 12.5 ML) would produce approximately 2750 m3 of metal rich sludge (approximately 100 tonnes dry weight) that would require separate disposal., (Crown Copyright © 2012. Published by Elsevier Ltd. All rights reserved.)

Background: Three annual Atriplex species-A. hortensis var. purpurea, A. hortensis var. rubra and A. rosea-growing on soil with various levels of the heavy metals copper, lead, nickel, and zinc, have been investigated., Results: Metal accumulation by Atriplex plants differed among species, levels of polluted soil and tissues. Metals accumulated by Atriplex were mostly distributed in root tissues, suggesting that an exclusion strategy for metal tolerance widely exists in them. The increased concentration of heavy metals in soil led to increases in heavy metal shoot and root concentrations of Ni, Cu, Pb and Zn in plants as compared to those grown on unpolluted soil. Accumulation was higher in roots than shoots for all the heavy metals. None of the plants were suitable for phytoextraction because no hyperaccumulator was identified. However, plants with a high bioconcentration factor and low translocation factor have the potential for phytostabilization. Similarly, the correlation between metal concentrations and translocations in plants (BCFs and TFs) using a linear regression was also statistically significant., Conclusion: Among the plants studied, var. purpurea was the most efficient in accumulating Pb and Zn in its shoots, whereas var. rubra was most suitable for phytostabilization of sites contaminated with Cu and Ni., (Copyright © 2011 Society of Chemical Industry.)

Even at trace levels, arsenic is of environmental and health concern due to its high toxicity. The xerohalophyte plant species Atriplex atacamensis grows on an arsenic-contaminated mining area in North Chile. Young seedlings that were grown from seeds collected from these plants were grown in a nutrient solution under controlled environmental conditions and were exposed for 14 and 28 days to 0, 100 or 1000 μM arsenate. More than 75% of the plants that were exposed to the highest As dose survived until the end of the treatment. The seedling growth was reduced (100 μM As) or inhibited (1000 μM As) in the stress conditions, but the plants were able to efficiently close their stomata and perform osmotic adjustments to avoid secondary water stress. Arsenic accumulated up to 400 μg g(-1) DW in the shoots and 3500 μg g(-1) DW in the roots. Arsenate drastically impaired the P content and increased glycinebetaine content, although no arsenobetaine was found in the tissues. With the exception of arsenite and arsenate, no As-containing organic compound was detected. Arsenic was not excreted by the trichomes that were present at the leaf surface. Although an increase in the total level of non-protein thiols suggested that arsenite fixation on the sulfhydryl groups could occur in the stressed tissues, the majority of the soluble arsenic remained in its oxidized state As(V). Arsenate induced an increase in the free soluble polyamine concentrations in all of the organs, and it increased the proportion of spermidine and spermine and decreased the proportion of putrescine in the polyamine pool. Therefore, it is likely that these polycationic molecules may assist in arsenate sequestration in the stressed tissues, and A. atacamensis may represent a promising plant species that can be tested in field trials for its phytomanagement of As-contaminated sites in desert areas., (Copyright © 2011 Elsevier B.V. All rights reserved.)

The removal of sodium salts from saline soils by salt tolerant crops, as alternative for costly chemical amendments, has emerged as an efficient low cost technology. Lysimeter experiments were carried out on a highly saline sodic soil (ECe = 65.3 dS m(-1), ESP = 27.4, CEC = 47.9 cmole+ kg(-1), and pH = 7.7) and irrigated with canal water (EC = 2.2 dSm(-1), SAR = 4.8) to investigate reclamation efficiency under four different treatments: control (no crop and no gypsum application) (C), gypsum application equivalent to 100% gypsum requirement (G100), planting sea orach (Atriplex halimus) as phytoremediation crop (Cr), planting sea orach with gypsum application equivalent to 50% gypsum requirement (CrG50). Soil salinity (ECe) and exchangeable sodium percentage (ESP) were significantly reduced compared to the control. Average ESP and ECe (dS m(-1)) in the top layer were 9.1, 5.8 (control), 4.8, 3.7 (Cr), 3.3, 3.9 (CrG50), and 3.8, 3.1 (G100), respectively. Atriplex halimus can be recommended as phytoremediation crop to reclaim highly saline sodic clay loam soils.

Proline transporters (ProTs) originally described as highly selective transporters for proline, have been shown to also transport glycinebetaine (betaine). Here we examined and compared the transport properties of Bet/ProTs from betaine accumulating (sugar beet, Amaranthus, and Atriplex,) and non-accumulating (Arabidopsis) plants. Using a yeast mutant deficient for uptake of proline and betaine, it was shown that all these transporters exhibited higher affinity for betaine than proline. The uptake of betaine and proline was pH-dependent and inhibited by the proton uncoupler carbonylcyanide m-chlorophenylhydrazone (CCCP). We also investigated choline transport by using a choline transport-deficient yeast mutant. Results revealed that these transporters exhibited a higher affinity for choline uptake rather than betaine. Uptake of choline by sugar beet BvBet/ProT1 was independent of the proton gradient and the inhibition by CCCP was reduced compared with that for uptake of betaine, suggesting different proton binding properties between the transport of choline and betaine. Additionally, in situ hybridization experiments revealed the localization of sugar beet BvBet/ProT1 in phloem and xylem parenchyma cells., (Copyright © 2011 Elsevier GmbH. All rights reserved.)

The possibility of remediating contaminated soils though the use of high biomass-generating, native plant species capable of removing heavy metals is receiving increased attention. The cadmium (Cd) accumulation capacities of the native Mediterranean, perennial shrubs Atriplex halimus, Phyllirea angustifolia, Rhamnus alaternus and Rosmarinus officinalis were tested by growing transplanted specimens in a pine bark compost substrate (pH 5.6) contaminated with 100 mg Cd kg(-1). After 70 days, only R. alaternus showed reduced growth. The increase in biomass seen in all the test species enhanced the phytoextraction of Cd. However, the species behaved as metal excluders, except for the halophyte A. halimus, which behaved as an indicator plant. In this species the leaf Cd concentration reached 35 mg Cd kg(-1), with the shoot responsible for some 86% of total Cd accumulation. Atriplex halimus showed the highest bioconcentration factor (BCF) (0.36) and leaf Cd transport index (1.68); consequently, this species showed the highest Cd phytoextraction capacity.

Mercury contaminated stockpiles of biosolids (8.4 mg kg⁻¹ Hg) from Melbourne Water's Western Treatment Plant (MW-WTP) were investigated to evaluate the possibility of their Hg chelate-assisted phytoextraction. The effects of ammonium thiosulphate (NH₄)₂S₂O₃, cysteine (Cys), nitrilotriacetic acid (NTA), and potassium iodide (KI) were studied to mobilize Hg and to increase its uptake in plant shoots. Three plant species were selected for this study, one herbaceous and two grasses: Atriplex codonocarpa, Austrodanthonia caespitosa and Vetiveria zizanioides. KI proved to be the best candidate for Hg phytostabilization in biosolids because it facilitated the concentration of this metal mainly in roots. (NH₄)₂S₂O₃ was shown to be the most effective chelating agent among those tested for Hg phytoextraction as it allowed the highest translocation of Hg into the above-ground tissues of the selected plant species. The phytoextraction conditions using A. caespitosa as the best performing plant species were optimized at an (NH₄)₂S₂O₃ concentration of 27 mmol kg⁻¹ and contact time with biosolids of seven day. Monitoring of the Hg concentration in biosolids and in leachate water during a 9-day treatment revealed that the biosolids Hg concentration decreased significantly after the first day of treatment and then it decreased only slightly with time reaching a value of 5.6 mg kg⁻¹ Hg at the end of the 9-day period. From the corresponding results obtained for the leachate water, it was suggested that a relatively large fraction of Hg (0.7 mg kg⁻¹ Hg) was promptly mobilized and consequently the plants were able to take up the metal and translocate it into shoots., (Copyright © 2011 Elsevier B.V. All rights reserved.)

Seed dimorphism provides plants with alternative strategies for survival in unfavorable environments. Here, we investigated the physiological responses and differential gene expression caused by salinity exposure in Atriplex centralasiatica plants grown from the two different seed morphs. Seedlings derived from yellow seeds (YS) showed a greater salt tolerance than those derived from brown seeds (BS). Salt treatment induced nitric oxide (NO) synthesis in roots, and seedlings derived from YS produced greater amounts of NO than did those from BS. Analyses of NO scavenging during salt stress revealed that NO contributed to the differential salt tolerance in seedlings derived from the two seed morphs by modulating antioxidative enzyme activity, hydrogen peroxide accumulation and the ion equilibrium. We also applied transcriptomics and subsequent microarray analysis to evaluate the differential gene expression during salt treatment. These genes encoded proteins related to osmotic and ionic homeostasis, redox equilibrium and signal transduction. A select group of genes including GH3.3, CAT1/2, TIP1, SIHP1 and EXP1 were further confirmed with RT-PCR analysis. These results revealed that the enhanced salt tolerance of seedlings from YS appeared to be governed by a superior ability to achieve ionic homeostasis and redox equilibrium, a rapid response to salt stress, and ultimately better growth potential. NO serves as a vital regulator in these processes.

The secreting glandular trichomes are recognized as an efficient structure that alleviates salt effects on Atriplex halimus. They are found on buds, young green stems, and leaves. They occupy both the leaf surfaces and give them a whitish color. Their histogenesis and ultrastructure were investigated in the third young leaves. They appear in early stage of plant development and its initiation continuous until just the leaf final development state. Each trichome contains two parts; a stalk which has high electron opacity, embedded in epidermal cells, and bears a second one which is unicellular, called bladder cell and has a low electron density. The bladder cell appears as a huge vacuole and the well-reduced cytoplasm which is pushed close to the wall, contains only a few organelles. Concurrently, the use of silver chloride precipitation technique shows that, in secretion process, salt follows a symplasmatic pathway which is consolidated by the presence of numerous plasmodesmata between the stalk cell(s), and the bladder one and the neighboring mesophyll cells. In addition, according to lanthanum-tracer study, salt can be excreted apoplastically. In fact, the heavy element can be transported via endocytosis vesicles, and by Golgi, endoplasmic reticulum, and lysosome (G.E.R.L.) network toward the storage vacuoles.

Drought and salinity induce water deficit, but may also have distinct effects on plant metabolism. To compare their impact on leaf senescence in relation to ABA and ethylene synthesis, young plants of Atriplex halimus L. were exposed to iso-osmotic concentrations of NaCl (160mM) or PEG (15%) in nutrient solution. Plant growth and development were more affected by PEG than by NaCl. Stressed plants remained able to reduce their osmotic potential, but the nature of accumulated organic osmocompatible solutes varied according to the stressing agent. Glycinebetaine accumulated to a greater extent in salt-treated plants than in water-stressed plants. Sodium chloride induced the accumulation of non-reducing sucrose, while PEG-treated plants mainly accumulated reducing glucose and fructose. Abscisic acid (ABA) accumulated in response to salt, while ethylene was synthesized mainly by PEG-treated plants and was involved in the induction of early senescence processes characterized by synthesis of reactive oxygen species, peroxidation of membrane lipids and a decrease in chlorophyll content. ABA sensitivity of stressed tissues was markedly different in response to salt and in response to non-ionic osmotic stress, and exogenous ABA (50μM) had contrasting effects on most physiological parameters depending on the stressing agent. Exogenous ABA induced a decrease in root and shoot growth and sucrose content, and an increase in reactive oxygen species content in salt-stressed plants. In contrast, exogenous ABA increased growth in PEG-treated plants in relation to an improvement of water use efficiency resulting from a more efficient stomatal control. Exogenous ABA increased ethylene synthesis in salt-treated plants, but had only marginal impact on PEG-treated ones. The xero-halophyte A. halimus thus responds in a contrasting way to salt and water stress, through accumulation of distinct osmocompatible solutes and hormonal compounds such as ethylene and ABA could play distinct roles in stress-induced senescence processes., (Copyright © 2010. Published by Elsevier GmbH.)

Beside a direct toxicity, cadmium impact on plants involves both a secondary-induced water stress and an oxidative stress. Proliferating cell lines of Atriplex halimus were selected for their sensitivity or resistance to polyethylene glycol (PEG 10,000, 20%) and then exposed to 100 microM CdCl2 in the simultaneous presence or absence of PEG 20% or 150 mM NaCl. The PEG resistant cell line exhibited a higher growth in the presence of Cd than the sensitive line, although Cd acccumulation was higher in the former than in the latter. Exogenous PEG induced an increase in Cd concentration in the sensitive but not in the resistant cell line while NaCl induced a decrease in Cd accumulation in both cell lines. In the presence of Cd alone, the water content (WC) was higher and the osmotic potential was lower in PEG-sensitive than in PEG resistant line. The presence of PEG in the Cd-containing medium increased the WC and decreased the osmotic potential in PEG-resistant line comparatively to Cd stress alone, while an inverse trend was observed for the sensitive line. The PEG-resistant cell line displayed a higher ability to cope with oxidative stress in relation to an increase of endogenous antioxidants (glutathione and ascorbic acid), a high constitutive superoxide dismutase (EC 1.15.1.1) activity and an efficient Cd-induced increase in glutathione reductase (GR) (EC 1.6.4.1) and ascorbate peroxidase (APX) (EC 1.11.1.11). Cadmium tolerance in PEG-resistant line is thus not related to any strategy of Cd exclusion or osmotic adjustment but to tolerance mechanisms allowing the tissue to restrict the deleterious impact of accumulated Cd., (Copyright 2009 Elsevier GmbH. All rights reserved.)

Background, Aim, and Scope: The success of phytoextraction depends upon the identification of suitable plant species that hyperaccumulate heavy metals and produce large amounts of biomass using established agricultural techniques. In this study, the Mediterranean saltbush Atriplex halimus L., which is a C4 perennial native shrub of Mediterranean basin with an excellent tolerance to drought and salinity, is investigated with the main aim to assess its phytoremediation potential for Pb and Cd removal from contaminated soils. In particular, the influence of soil salinity in metal accumulation has been studied as there is notable evidence that salinity changes the bioavailability of metals in soil and is a key factor in the translocation of metals from roots to the aerial parts of the plant., Materials and Methods: Three pot experiments were conducted under greenhouse conditions for a 10-week period with A. halimus grown in soil artificially polluted with 20 ppm of Cd and/or 800 ppm of Pb and irrigated with three different salt solutions (0.0%, 0.5%, and 3.0% NaCl). Soil measurements for soil characterization were performed with the expiration of the first week of plant exposure to metals and NaCl, and at the end of the experimental period, chlorophyll content, leaf protein content, leaf specific activity of guaiacol peroxidase (EC 1.11.1.7), shoot water content, biomass, and Cd and Pb content in the plant tissues were determined. Additionally, any symptoms of metal or salt toxicity exhibited by the plants were visually noted during the whole experimental period., Results: The experimental data suggest that increasing salinity increases cadmium uptake by A. halimus L. while in the case of lead there was not a clear effect of the presence of salt on lead accumulation in plant tissues. A. halimus developed no visible signs of metal toxicity; only salt toxicity symptoms were observed in plants irrigated with 3% NaCl solutions. Chlorophyll content, leaf protein content, shoot water content, and biomass were not negatively affected by the metals; instead, there was even an increase in the amount of photosynthetic pigments in plants treated with both metals and salinity. The specific activity of guaiacol peroxidase seems to have a general tendency for increase in plants treated with the metals in comparison with the respective controls but a statistically significant difference exists only in plants treated with the metal mixture and saline conditions., Discussion: The data revealed that lead and cadmium accumulation in plant tissues was kept generally at low levels. Salinity was found to have a positive effect on cadmium uptake by the plant and this may be related to a higher bioavailability of the metal in soil due to decreased Cd sorption on soil particles. On the other hand, salinity did not influence in a clear way the uptake of Pb by the plant probably because of lead's limited mobility in soils and plant tissues. Cd and Pd usually decrease the chlorophyll content and biomass and change water relations in plants; however, A. halimus was found not to be affected indicating that it is a Cd- and Pb-tolerant plant. Guaiacol peroxidase activity as one of the parameters expressing oxidative damage and extent of stress in plants was not generally found to be significantly affected under the presence of metals in most plants suggesting that the extent of stress in plants was minimal, while only for plants treated with the metal mixture and low salinity the enzyme activity was elevated confirming that this enzyme serves as an antioxidative tool against the reactive oxygen species produced by the metals., Conclusions: Atriplex halimus L. is a Pb- and Cd-tolerant plant but metal concentrations achieved in plant tissues were kept generally at low levels; however, metal accumulation in shoots, especially for Cd, considered together with its high biomass production, rapid growth, and deep root system able to cope with poor structure and xeric characteristics of several polluted soils suggest that this plant deserves further investigation., Recommendations and Perspectives: Phytoextraction by halophytes is a promising alternative for the remediation of heavy metal contaminated sites affected by salinity since saline depressions often indicate sites of industrial effluents accumulation, contaminated by heavy metals, including Pb and Cd. Halophytes are also promising candidates for the removal of heavy metals from non-saline soils. Furthermore, the use of such plants can be potentially viewed as an alternative method for soil desalination where salt is removed from the soil instead of being washed downwards by water or other solutions.

Batch studies were performed to determine the interference of calcium (Ca) and magnesium (Mg) on the sorption of Cu(II), Cd(II), Cr(III), Cr(VI), Pb(II), and Zn(II) [from CuSO(4), K(2)Cr(2)O(7), Pb(NO(3))(2), Cr(NO(3))(3), ZnCl(2), and Cd(NO(3))(2)] by saltbush (Atriplex canescens) biomass. The results demonstrated that Ca and Mg at concentrations of at least 20 times higher than the concentration of most of the target metals did not interfere with the metal binding. The data show that the batch binding capacity from a multimetal solution at pH 5.0 was (micromol/g) about 260 for Cr(III) and Pb, and about 117, 54, and 49 for Cu, Zn, and Cd, respectively. The use of 0.1M HCl allowed the recovery of 85-100% of the bound Cu, Cr(III), and Pb, and more than 37% of the bound Cd and Zn. The column binding capacity for Pb was about 49 micromol/g from both the single and multimetal solutions, while it was, respectively about 35 and 23 micromol/g for Cr(III). The binding capacity for Cu and Zn from the single and multimetal column experiments was 35 micromol/g and less than 10 micromol/g, respectively. The stripping data from the single column experiment showed that 0.1M HCl allowed the recovery of all the bound Cu and Zn, 90% and 74% of the bound Pb and Cr(VI), respectively, and less than 25% of the bound Cd and Cr(III), while the stripping from the multimetal experiment showed that 0.1M HCl allowed the recovery of all the bound Cu and about 74%, 54%, 43%, and 40% of the bound Pb, Zn, Cd, and Cr(III), respectively.

A field experiment was carried out to determine the growth characteristics and mineral composition of two local halophytes (Atriplex halimus and Salvadora persica) under saline irrigation at Kind Abdulaziz City for Science and Technology (KACST), Research Station Al-Muzahmyia, Riyadh. The experiment treatments were one soil (sandy), four irrigation waters of different salinities (2000, 8000, 12000 and 16000 mg L(-1) TDS), two halophytes (Salvadora persica and Atriplex halimus) and one irrigation level (irrigation at 50% depletion of moisture at field capacity). Mean fresh biomass yield and fresh plant root weight of A. halimus increased while that of S. persica decreased significantly with increasing irrigation water salinity in all the treatments. Soil salinity increased significantly with increasing water salinity. A positive correlation (r = 0.987) existed between the irrigation water salinity and the soil salinity resulting from saline irrigation. The plant tissue protein contents increased in A. halimus, but decreased in S. persica with increasing irrigation water salinity. The Na ion uptake by plant roots was significantly less than K in A. halimus compared to S. persica which indicated adjustment of plants to high soil salinity and high Na ion concentration for better growth. The order of increasing salt tolerance was A. halimus > S. persica under the existing plant growing conditions. Among the two halophytes, A. halimus showed great potential for establishing gene banks of local species, because it has more forage value due to high protein contents than S. persica for range animals.

Soil salinity and drought compromise water uptake and lead to osmotic adjustment in xero-halophyte plant species. These important environmental constraints may also have specific effects on plant physiology. Stress-induced accumulation of osmocompatible solutes was analysed in two Tunisian populations of the Mediteranean shrub Atriplex halimus L.-plants originating from a salt-affected coastal site (Monastir) or from a non-saline semi-arid area (Sbikha)-were exposed to nutrient solution containing either low (40 mM) or high (160 mM) doses of NaCl or 15% polyethylene glycol. The low NaCl dose stimulated plant growth in both populations. Plants from Monastir were more resistant to high salinity and exhibited a greater ability to produce glycinebetaine in response to salt stress. Conversely, plants from Sbikha were more resistant to water stress and displayed a higher rate of proline accumulation. Proline accumulated as early as 24 h after stress imposition and such accumulation was reversible. By contrast, glycinebetaine concentration culminated after 10 d of stress and did not decrease after the stress relief. The highest salt resistance of Monastir plants was not due to a lower rate of Na(+) absorption; plants from this population exhibited a higher stomatal conductance and a prodigal water-use strategy leading to lower water-use efficiency than plants from Sbikha. Exogenous application of proline (1 mM) improved the level of drought resistance in Monastir plants through a decrease in oxidative stress quantified by the malondialdehyde concentration, while the exogenous application of glycinebetaine improved the salinity resistance of Sbikha plants through a positive effect on photosystem II efficiency.

Background and Aims: Atriplex (Halimione) portulacoides is a halophytic, C(3) shrub. It is virtually confined to coastal salt marshes, where it often dominates the vegetation. The aim of this study was to investigate its growth responses to salinity and the extent to which these could be explained by photosynthetic physiology., Methods: The responses of young plants to salinity in the range 0-700 mol m(-3) NaCl were investigated in a glasshouse experiment. The performance of plants was examined using classical growth analysis, measurements of gas exchange (infrared gas analysis), determination of chlorophyll fluorescence characteristics (modulated fluorimeter) and photosynthetic pigment concentrations; total ash, sodium, potassium and nitrogen concentrations, and relative water content were also determined., Key Results: Plants accumulated Na(+) approximately in proportion to external salinity. Salt stimulated growth up to an external concentration of 200 mol m(-3) NaCl and some growth was maintained at higher salinities. The main determinant of growth response to salinity was unit leaf rate. This was itself reflected in rates of CO(2) assimilation, which were not affected by 200 mol m(-3) but were reduced at higher salinities. Reductions in net photosynthetic rate could be accounted for largely by lower stomatal conductance and intercellular CO(2) concentration. Apart from possible effects of osmotic shock at the beginning of the experiment, salinity did not have any adverse effect on photosystem II (PSII). Neither the quantum efficiency of PSII (Phi(PSII)) nor the chlorophyll fluorescence ratio (F(v)/F(m)) were reduced by salinity, and lower mid-day values recovered by dawn. Mid-day F(v)/F(m) was in fact depressed more at low external sodium concentration, by the end of the experiment., Conclusions: The growth responses of the hygro-halophyte A. portulacoides to salinity appear largely to depend on changes in its rate of photosynthetic gas exchange. Photosynthesis appears to be limited mainly through stomatal conductance and hence intercellular CO(2) concentration, rather than by effects on PSII; moderate salinity might stimulate carboxylation capacity. This is in contrast to more extreme halophytes, for which an ability to maintain leaf area can partially offset declining rates of carbon assimilation at high salinity.

Mine tailings are moderately to severely impacted sites that lack normal plant cover, soil structure and development, and the associated microbial community. In arid and semiarid environments, tailings and their associated contaminants are prone to eolian dispersion and water erosion, thus becoming sources of metal contamination. One approach to minimize or eliminate these processes is to establish a permanent vegetation cover on tailings piles. Here we report a revegetation trial conducted at a moderately impacted mine tailings site in southern Arizona. A salt and drought-tolerant plant, four-wing saltbush [Atriplex canescens (Pursh) Nutt.], was chosen for the trial. A series of 3 by 3 m plots were established in quadruplicate on the test site to evaluate growth of four-wing saltbush transplants alone or with compost addition. Results show that >80% of the transplanted saltbush survived after 1.5 yr in both treatments. Enumeration of heterotrophs and community structure analysis were conducted to monitor bacterial community changes during plant establishment as an indicator of plant and soil health. The bacterial community was evaluated using denaturing gradient gel electrophoresis (DGGE) analysis of 16S rDNA PCR gene products from tailings samples taken beneath transplant canopies. Significant differences in heterotrophic counts and community composition were observed between the two treatments and unplanted controls throughout the trial, but treatment effects were not observed. The results suggest that compost is not necessary for plant establishment at this site and that plants, rather than added compost, is the primary factor enhancing bacterial heterotrophic counts and affecting community composition.

Abandoned mine tailings sites in semiarid regions remain unvegetated for extended periods of time and are subject to eolian dispersion and water erosion. This study examines the potential phytostabilization of a lead-zinc mine tailings site using a native, drought-tolerant halophyte, quailbush [Atriplex lentiformis (Torr.) S. Wats.]. In a greenhouse study germination, growth, and metal uptake was evaluated in two compost-amended mine tailings samples, K4 (pH 3) and K6 (pH 6) at 75, 85, 90, 95, and 100% mine tailings, and two controls, off-site and compost. Microbial community changes were monitored by performing MPN analysis of iron- and sulfur-oxidizing bacteria as well as heterotrophic plate counts. Results demonstrate that germination is not a good indicator for phytostabilization since it was only inhibited in the unamended K4 treatment. Plant growth was significantly reduced in 95 and 100% mine tailings, while growth in 75, 85, and 90% treatments was similar to the off-site control. Quailbush accumulated elevated levels of the nutrient metals Na, K, Mn, and Zn in the shoot tissues; however, metal accumulation was generally below the domestic animal toxicity limit. Initially, autotrophic population estimates were four to six logs higher than heterotrophic counts, indicating extremely stressed conditions. However, post-harvest, heterotrophic bacterial counts increased to normal levels (approximately 10(6) CFU g-1 dry tailings) and dominated the rhizosphere. Therefore, with compost amendment, quailbush has good potential as a native species candidate for phytostabilization of mine tailings in semiarid environments.

Fourier transform infrared (FTIR) studies were performed to confirm the chemical modification of saltbush (Atriplex canescens) biomass and to provide information about the identity and binding characteristics of the chemical groups responsible for the binding of Cd(II), Cr(III), and Cr(VI). In addition, studies were performed to determine the optimum time for the binding of the three ions by saltbush biomass, and to study the efficiency of HCl and sodium citrate as stripping agents. The metal quantification was performed using inductively coupled plasma optical emission spectroscopy (ICP-OES). The results showed that 10 min or less is enough to achieve the maximum metal binding, and that aqueous solutions of 0.1 mM HCl or sodium citrate were enough to strip more than 80% of the bound Cd. It was determined that more than 70% of the bound Cr(III) was stripped using 0.1 mM HCl. Chemical modification of carboxyl and ester groups on the biomass was performed. The FTIR results confirmed that the esterification of carboxyl groups and hydrolysis of ester groups in the native biomass had occurred. The direct effect of these modifications on the binding properties of the biomass provided strong evidence that the carboxyl functionality is the main group responsible for binding Cd and Cr(III). However, the IR data showed that for Cr(VI), a different type of functional group is involved.