Genomic analysis reveals phylogeny of Zygophyllales and mechanism for water retention of a succulent xerophyte.

Revealing the genetic basis for stress-resistant traits in extremophile plants will yield important information for crop improvement. Zygophyllum xanthoxylum, an extant species of the ancient Mediterranean, is a succulent xerophyte that can maintain a favorable water status under desert habitats; however, the genetic basis of this adaptive trait is poorly understood. Furthermore, the phylogenetic position of Zygophyllales, to which Z. xanthoxylum belongs, remains controversial. In this study, we sequenced and assembled the chromosome-level genome of Z. xanthoxylum. Phylogenetic analysis showed that Zygophyllales and Myrtales form a separated taxon as a sister to the clade comprising fabids and malvids, clarifying the phylogenetic position of Zygophyllales at whole-genome scale. Analysis of genomic and transcriptomic data revealed multiple critical mechanisms underlying the efficient osmotic adjustment using Na+ and K+ as "cheap" osmolytes that Z. xanthoxylum has evolved through the expansion and synchronized expression of genes encoding key transporters/channels and their regulators involved in Na+/K+ uptake, transport, and compartmentation. It is worth noting that ZxCNGC1;1 (cyclic nucleotide-gated channels) and ZxCNGC1;2 constituted a previously undiscovered energy-saving pathway for Na+ uptake. Meanwhile, the core genes involved in biosynthesis of cuticular wax also featured an expansion and upregulated expression, contributing to the water retention capacity of Z. xanthoxylum under desert environments. Overall, these findings boost the understanding of evolutionary relationships of eudicots, illustrate the unique water retention mechanism in the succulent xerophyte that is distinct from glycophyte, and thus provide valuable genetic resources for the improvement of stress tolerance in crops and insights into the remediation of sodic lands.

Massive increases in C31 alkane on Zygophyllum xanthoxylum leaves contribute to its excellent abiotic stress tolerance.

BACKGROUND AND AIMS: Desert plants possess excellent water-conservation capacities to survive in extreme environments. Cuticular wax plays a pivotal role in reducing water loss through plant aerial surfaces. However, the role of cuticular wax in water retention by desert plants is poorly understood. METHODS: We investigated leaf epidermal morphology and wax composition of five desert shrubs from north-west China and characterized the wax morphology and composition for the typical xerophyte Zygophyllum xanthoxylum under salt, drought and heat treatments. Moreover, we examined leaf water loss and chlorophyll leaching of Z. xanthoxylum and analysed their relationships with wax composition under the above treatments. KEY RESULTS: The leaf epidermis of Z. xanthoxylum was densely covered by cuticular wax, whereas the other four desert shrubs had trichomes or cuticular folds in addition to cuticular wax. The total amount of cuticular wax on leaves of Z. xanthoxylum and Ammopiptanthus mongolicus was significantly higher than that of the other three shrubs. Strikingly, C31 alkane, the most abundant component, composed >71 % of total alkanes in Z. xanthoxylum, which was higher than for the other four shrubs studied here. Salt, drought and heat treatments resulted in significant increases in the amount of cuticular wax. Of these treatments, the combined drought plus 45 °C treatment led to the largest increase (107 %) in the total amount of cuticular wax, attributable primarily to an increase of 122 % in C31 alkane. Moreover, the proportion of C31 alkane within total alkanes remained >75 % in all the above treatments. Notably, the water loss and chlorophyll leaching were reduced, which was negatively correlated with C31 alkane content. CONCLUSION: Zygophyllum xanthoxylum could serve as a model desert plant for study of the function of cuticular wax in water retention because of its relatively uncomplicated leaf surface and because it accumulates C31 alkane massively to reduce cuticular permeability and resist abiotic stressors.

Physiological and transcriptomic analyses provide insight into thermotolerance in desert plant Zygophyllum xanthoxylum.

BACKGROUND: Heat stress has adverse effects on the growth and reproduction of plants. Zygophyllum xanthoxylum, a typical xerophyte, is a dominant species in the desert where summer temperatures are around 40 °C. However, the mechanism underlying the thermotolerance of Z. xanthoxylum remained unclear. RESULTS: Here, we characterized the acclimation of Z. xanthoxylum to heat using a combination of physiological measurements and transcriptional profiles under treatments at 40 °C and 45 °C, respectively. Strikingly, moderate high temperature (40 °C) led to an increase in photosynthetic capacity and superior plant performance, whereas severe high temperature (45 °C) was accompanied by reduced photosynthetic capacity and inhibited growth. Transcriptome profiling indicated that the differentially expressed genes (DEGs) were related to transcription factor activity, protein folding and photosynthesis under heat conditions. Furthermore, numerous genes encoding heat transcription shock factors (HSFs) and heat shock proteins (HSPs) were significantly up-regulated under heat treatments, which were correlated with thermotolerance of Z. xanthoxylum. Interestingly, the up-regulation of PSI and PSII genes and the down-regulation of chlorophyll catabolism genes likely contribute to improving plant performance of Z. xanthoxylum under moderate high temperature. CONCLUSIONS: We identified key genes associated with of thermotolerance and growth in Z. xanthoxylum, which provide significant insights into the regulatory mechanisms of thermotolerance and growth regulation in Z. xanthoxylum under high temperature conditions.

Effect of phosphorus efficiency on elemental stoichiometry of two shrubs.

Phosphorus (P) is an important nutrient that can restrict plant growth. However, the influence of P deficiency on elemental homeostasis and application of the growth rate hypothesis in higher plants remain to be assessed. Two shrubs, Zygophyllum xanthoxylum and Nitraria tangutorum, were used as experiment material and subjected to five P addition treatments: 0, 17.5, 35.0, 52.5 and 70.0 mg P·kg-1 soil. The biomass and relative growth rate of Z. xanthoxylum did not change with altered P supply. There was no significant difference in P concentration among the treatments for Z. xanthoxylum, but N. tangutorum showed an upward trend. The P stoichiometric homeostasis of Z. xanthoxylum was higher than that of N. tangutorum. For Z. xanthoxylum, available P in the rhizosphere improved significantly under extreme P deficiency conditions, and P concentrations in all treatments were lower than in N. tangutorum, showing that Z. xanthoxylum had stronger P absorption and P utilization capacity. No relationships between growth rate and C:N:P ratios were found in Z. xanthoxylum. The strong P efficiency, and high and stable dry matter accumulation, are likely contributors in maintaining stoichiometric homeostasis. In addition, the relatively high biomass accumulation and high P utilization efficiency for Z. xanthoxylum does not support the growth rate hypothesis for this species.

Transcriptomic and metabolomic analysis reveals the role of CoA in the salt tolerance of Zygophyllum spp.

BACKGROUND: Zygophyllum is an important medicinal plant, with notable properties such as resistance to salt, alkali, and drought, as well as tolerance of poor soils and shifting sand. However, the response mechanism of Zygophyllum spp. to abiotic stess were rarely studied. RESULTS: Here, we aimed to explore the salt-tolerance genes of Zygophyllum plants by transcriptomic and metabolic approaches. We chose Z. brachypterum, Z. obliquum and Z. fabago to screen for salt tolerant and sensitive species. Cytological observation showed that both the stem and leaf of Z. brachypterum were significantly thicker than those of Z. fabago. Then, we treated these three species with different concentrations of NaCl, and found that Z. brachypterum exhibited the highest salt tolerance (ST), while Z. fabago was the most sensitive to salt (SS). With the increase of salt concentration, the CAT, SOD and POD activity, as well as proline and chlorophyll content in SS decreased significantly more than in ST. After salt treatment, the proportion of open stomata in ST decreased significantly more than in SS, although there was no significant difference in stomatal number between the two species. Transcriptomic analysis identified a total of 11 overlapping differentially expressed genes (DEGs) in the leaves and roots of the ST and SS species after salt stress. Two branched-chain-amino-acid aminotransferase (BCAT) genes among the 11 DEGs, which were significantly enriched in pantothenate and CoA biosynthesis, as well as the valine, leucine and isoleucine biosynthesis pathways, were confirmed to be significantly induced by salt stress through qRT-PCR. Furthermore, overlapping differentially abundant metabolites showed that the pantothenate and CoA biosynthesis pathways were significantly enriched after salt stress, which was consistent with the KEGG pathways enriched according to transcriptomics. CONCLUSIONS: In our study, transcriptomic and metabolomic analysis revealed that BCAT genes may affect the pantothenate and CoA biosynthesis pathway to regulate the salt tolerance of Zygophyllum species, which may constitute a newly identified signaling pathway through which plants respond to salt stress.

Responses of leaf C:N:P stoichiometry to water supply in the desert shrub Zygophyllum xanthoxylum.

Based on the elemental composition of major biochemical molecules associated with different biological functions, the 'growth rate hypothesis' proposed that organisms with a higher growth rate would be coupled to lower C:N, especially lower C:P and N:P ratios. However, the applicability of the growth rate hypothesis for plants is unclear, especially for shrubs growing under different water supply. We performed an experiment with eight soil moisture levels (soil water content: 4%, 6%, 8%, 13%, 18%, 23%, 26% and 28%) to evaluate the effects of water availability on leaf C:N:P stoichiometry in the shrub Zygophyllum xanthoxylum. We found that leaves grew slowly and favored accumulation of P over C and N under both high and low water supply. Thus, leaf C:P and N:P ratios were unimodally related to soil water content, in parallel with individual leaf area and mass. As a result, there were significant positive correlations between leaf C:P and N:P with leaf growth (u). Our result that slower-growing leaves had lower C:P and N:P ratios does not support the growth rate hypothesis, which predicted a negative association of N:P ratio with growth rate, but it is consistent with recent theoretical derivations of growth-stoichiometry relations in plants, where N:P ratio is predicted to increase with increasing growth for very low growth rates, suggesting leaf growth limitation by C and N rather than P for drought and water saturation.

ZxAKT1 is essential for K+ uptake and K+ /Na+ homeostasis in the succulent xerophyte Zygophyllum xanthoxylum.

The inward-rectifying K+ channel AKT1 constitutes an important pathway for K+ acquisition in plant roots. In glycophytes, excessive accumulation of Na+ is accompanied by K+ deficiency under salt stress. However, in the succulent xerophyte Zygophyllum xanthoxylum, which exhibits excellent adaptability to adverse environments, K+ concentration remains at a relatively constant level despite increased levels of Na+ under salinity and drought conditions. In this study, the contribution of ZxAKT1 to maintaining K+ and Na+ homeostasis in Z. xanthoxylum was investigated. Expression of ZxAKT1 rescued the K+ -uptake-defective phenotype of yeast strain CY162, suppressed the salt-sensitive phenotype of yeast strain G19, and complemented the low-K+ -sensitive phenotype of Arabidopsis akt1 mutant, indicating that ZxAKT1 functions as an inward-rectifying K+ channel. ZxAKT1 was predominantly expressed in roots, and was induced under high concentrations of either KCl or NaCl. By using RNA interference technique, we found that ZxAKT1-silenced plants exhibited stunted growth compared to wild-type Z. xanthoxylum. Further experiments showed that ZxAKT1-silenced plants exhibited a significant decline in net uptake of K+ and Na+ , resulting in decreased concentrations of K+ and Na+ , as compared to wild-type Z. xanthoxylum grown under 50 mm NaCl. Compared with wild-type, the expression levels of genes encoding several transporters/channels related to K+ /Na+ homeostasis, including ZxSKOR, ZxNHX, ZxSOS1 and ZxHKT1;1, were reduced in various tissues of a ZxAKT1-silenced line. These findings suggest that ZxAKT1 not only plays a crucial role in K+ uptake but also functions in modulating Na+ uptake and transport systems in Z. xanthoxylum, thereby affecting its normal growth.

Co-expression of tonoplast Cation/H(+) antiporter and H(+)-pyrophosphatase from xerophyte Zygophyllum xanthoxylum improves alfalfa plant growth under salinity, drought and field conditions.

Salinity and drought are major environmental factors limiting the growth and productivity of alfalfa worldwide as this economically important legume forage is sensitive to these kinds of abiotic stress. In this study, transgenic alfalfa lines expressing both tonoplast NXH and H(+)-PPase genes, ZxNHX and ZxVP1-1 from the xerophyte Zygophyllum xanthoxylum L., were produced via Agrobacterium tumefaciens-mediated transformation. Compared with wild-type (WT) plants, transgenic alfalfa plants co-expressing ZxNHX and ZxVP1-1 grew better with greater plant height and dry mass under normal or stress conditions (NaCl or water-deficit) in the greenhouse. The growth performance of transgenic alfalfa plants was associated with more Na(+), K(+) and Ca(2+) accumulation in leaves and roots, as a result of co-expression of ZxNHX and ZxVP1-1. Cation accumulation contributed to maintaining intracellular ions homoeostasis and osmoregulation of plants and thus conferred higher leaf relative water content and greater photosynthesis capacity in transgenic plants compared to WT when subjected to NaCl or water-deficit stress. Furthermore, the transgenic alfalfa co-expressing ZxNHX and ZxVP1-1 also grew faster than WT plants under field conditions, and most importantly, exhibited enhanced photosynthesis capacity by maintaining higher net photosynthetic rate, stomatal conductance, and water-use efficiency than WT plants. Our results indicate that co-expression of tonoplast NHX and H(+)-PPase genes from a xerophyte significantly improved the growth of alfalfa, and enhanced its tolerance to high salinity and drought. This study laid a solid basis for reclaiming and restoring saline and arid marginal lands as well as improving forage yield in northern China.

Profiling of rutin-mediated alleviation of cadmium-induced oxidative stress in Zygophyllum fabago.

Zygophyllum fabago grows in arid, saline soil, or disturbed sites, such as former industrial or mining areas. This species is able to grow in coarse mineral substrates contaminated with heavy metals. To investigate the effects of the flavonoid rutin (Rtn) on certain heavy metal stress responses such as antioxidant defense systems and water status, seedlings were subjected to 100 and 200 µM CdCl2 treatment without or with 0.25 and 1 mM Rtn for 7 and 14 d (days). Cd stress decreased growth (RGR), water content (RWC), leaf osmotic potential (Ψ(Π)), and chlorophyll fluorescence, all of which could be partly alleviated by addition of Rtn. Activities of superoxide dismutase, peroxidase (POX), ascorbate peroxidase, and glutathione reductase increased within the first 7 d after exposure to Cd. However, failure of antioxidant defense in the scavenging of reactive oxygen species (ROS) was evidenced by an abnormal rise in superoxide anion radical ( O2(•-)) and hydrogen peroxide contents and a decline in hydroxyl radical (OH(•)) scavenging activity, resulting in enhancement of lipid peroxidation (TBARS) as a marker of Cd-induced oxidative stress. However, exogenously applied Rtn considerably improved the stress tolerance of plants via a reduction in Cd accumulation, modulation of POX activity, increase of proline (Pro) content, decrease in TBARS and ROS content and consequent lowering of oxidative damage of membrane. Overall, 0.25 and 1 mM Rtn could protect Z. fabago from the harmful effects of 100 µM Cd-induced oxidative stress throughout the experiment.

Pb-induced responses in Zygophyllum fabago plants are organ-dependent and modulated by salicylic acid.

Zygophyllum fabago is a promising species for restoring heavy metal (HM) polluted soils, although the mechanisms involved in HM tolerance in this non-model plant remain largely unknown. This paper analyses the extent to which redox-active compounds and enzymatic antioxidants in roots, stems and leaves are responsible for Pb tolerance in a metallicolous ecotype of Z. fabago and the possible influence of salicylic acid (SA) pretreatment (24 h, 0.5 mM SA) in the response to Pb stress. SA pretreatment reduced both the accumulation of Pb in roots and even more so the concentration of Pb in aerial parts of the plants, although a similar drop in the content of chlorophylls and in the maximum quantum yield of photosystem II was observed in both Pb- and SA-Pb-treated plants. Pb increased the endogenous free SA levels in all organs and this response was enhanced in root tissues upon SA pretreatment. Generally, Pb induced a reduction in catalase, ascorbate peroxidase and glutathione reductase specific activities, whereas dehydroascorbate reductase was increased in all organs of control plants. SA pretreatment enhanced the Pb-induced H2O2 accumulation in roots by up-regulating Fe-superoxide dismutase isoenzymes. Under Pb stress, the GSH redox ratio remained highly reduced in all organs while the ascorbic acid redox ratio dropped in leaf tissues where a rise in lipid peroxidation products and electrolyte leakage was observed. Finally, an organ-dependent accumulation of proline and ß-carboline alkaloids was found, suggesting these nitrogen-redox-active compounds could play a role in the adaptation strategies of this species to Pb stress.

Syrian bean-caper (Zygophyllum fabago L.) improves organic matter and other properties of mine wastes deposits.

The omni-presence of Zygophyllum fabago L. (Syrian bean-caper) natural colonies in post mining areas prompted us to investigate its contributions to reclamation of mine wastes deposits in southeast Spain. Select plant-related (edaphic) characteristics and bio- and water soluble-Cd, Cu, Pb and Zn in rhizosphere of Z. fabago were compared to deposits one year since application of pig slurry and marble waste. Total N in rhizosphere increased up to a factor of 20X (339 vs 17 mg N kg(-1)) in El Gorguel and 27X (85 vs 3.1 mg N kg(-1)) in El Lirio sites. Organic matter accumulation in rhizosphere from litter and roots of Z. fabago increased organic C from 6.6 to 19.5 g kg(-1) in El Gorguel and from 2.1 to 5.7 g kg(-1) in El Lirio in one year. Dissolution of inorganic C takes place due to organic acids from root exudates of Z. fabago. Reduction in bio-available Cd, Cu, Pb, and Zn in rhizosphere of Z. fabago at El Lirio is attributed to increase in pH from 5.3 to 7.7 through marble waste addition, although increased cation exchange capacity may also have played a role. Addition of marble waste to encourage colonization by Z. fabago in acidic mine wastes deposits was recommended.

Differential cadmium and zinc distribution in relation to their physiological impact in the leaves of the accumulating Zygophyllum fabago L.

Cadmium and zinc share many similar physiochemical properties, but their compartmentation, complexation and impact on other mineral element distribution in plant tissues may drastically differ. In this study, we address the impact of 10 µm Cd or 50 µm Zn treatments on ion distribution in leaves of a metallicolous population of the non-hyperaccumulating species Zygophyllum fabago at tissue and cell level, and the consequences on the plant response through a combined physiological, proteomic and metabolite approach. Micro-proton-induced X-ray emission and laser ablation inductively coupled mass spectrometry analyses indicated hot spots of Cd concentrations in the vicinity of vascular bundles in response to Cd treatment, essentially bound to S-containing compounds as revealed by extended X-ray absorption fine structure and non-protein thiol compounds analyses. A preferential accumulation of Zn occurred in vascular bundle and spongy mesophyll in response to Zn treatment, and was mainly bound to O/N-ligands. Leaf proteomics and physiological status evidenced a protection of photosynthetically active tissues and the maintenance of cell turgor through specific distribution and complexation of toxic ions, reallocation of some essential elements, synthesis of proteins involved in photosynthetic apparatus or C-metabolism, and metabolite synthesis with some specificities regarding the considered heavy metal treatment.

LC-ESI-TOF-MS identification of bioactive secondary metabolites involved in the antioxidant, anti-inflammatory and anticancer activities of the edible halophyte Zygophyllum album Desf.

In this work, liquid chromatography coupled to electrospray time-of-flight mass spectrometry (LC-ESI-TOF-MS) has been applied to screen bioactive metabolites in shoot extract of the medicinal halophyte Zygophyllum album. Among 10 compounds identified (saponins, flavonoids and sterols) five were reported for the first time in Z. album. Furthermore, novel biological activities of hexane, dichloromethane and methanolic extracts were assessed. Results showed that methanolic extract exhibit the highest antioxidant activity using in vitro ORAC test and anti-inflammatory activity, inhibiting by 84.8% NO release in RAW264.7 macrophages. However, dichloromethane extract proved the utmost antioxidant activity in cell (WS1) based-assay (IC50=57 µg/ml) and interesting anticancer capacity against human lung carcinoma (A-549) and colon adenocarcinoma (DLD-1) cells (IC50=37 and 48 µg/ml, respectively). These findings can be attributed to the presence of triterpenes, flavonoids and sterols in Z. album, which are widely known as powerful antioxidants and used in various industrial fields.

The ZxNHX gene encoding tonoplast Na(+)/H(+) antiporter from the xerophyte Zygophyllum xanthoxylum plays important roles in response to salt and drought.

Sodium (Na(+)) has been found to play important roles in the adaptation of xerophytic species to drought conditions. The tonoplast Na(+)/H(+) antiporter (NHX) proved to be involved in the compartmentalization of Na(+) into vacuoles from the cytosol. In this study, a gene (ZxNHX) encoding tonoplast Na(+)/H(+) antiporter was isolated and characterized in Zygophyllum xanthoxylum, a succulent xerophyte growing in desert areas of northwest China. The results revealed that ZxNHX consisted of 532 amino acid residues with a conserved binding domain ((78)LFFIYLLPPI(87)) for amiloride and shared high similarity (73-81%) with the identified tonoplast Na(+)/H(+) antiporters in other plant species. Semi-quantitative RT-PCR analysis showed that the mRNA level of ZxNHX was significantly higher in the leaf than in stem or root. The transcript abundance of ZxNHX in Z. xanthoxylum subjected to salt (5-150 mM NaCl) or drought (50-15% of field water capacity (FWC)) was 1.4-8.4 times or 2.3-4.4 times that of plants grown in the absence of NaCl or 70% of FWC, respectively. Leaf Na(+) concentration in plants exposed to salt or drought was 1.7-5.2 times or 1.5-2.2 times that of corresponding control plants, respectively. It is clear that there is a positive correlation between up-regulation of ZxNHX and accumulation of Na(+) in Z. xanthoxylum exposed to salt or drought. Furthermore, Z. xanthoxylum accumulated larger amounts of Na(+) than K(+) in the leaf under drought conditions, even in low salt soil. In summary, our results suggest that ZxNHX encodes a tonoplast Na(+)/H(+) antiporter and plays important roles in Na(+) accumulation and homeostasis of Z. xanthoxylum under salt and drought conditions.

Amiloride reduces sodium transport and accumulation in the succulent xerophyte Zygophyllum xanthoxylum under salt conditions.

To investigate the effects of amiloride, which is an inhibitor of Na(+)/H(+) antiporter (NHX), on Na(+) accumulation and transport at the whole plant level, in this study, 3-week-old Zygophyllum xanthoxylum plants were exposed to 25, 50 and 100 mM NaCl with or without 0.5 mM amiloride. After 72 h of treatment, dry weight, tissue water content, Na(+) and K(+) concentrations, and transcript levels of ZxNHX were determined. The results indicated that, under salt conditions, leaf Na(+) concentration, total Na(+) quantity and root net Na(+) uptake rate in plants treated with amiloride are significantly lower than those in control plants. Amiloride remarkably increased Na(+) proportion in stem and decreased Na(+) proportion in leaf of Z. xanthoxylum. Furthermore, our results showed that the transcript levels of ZxNHX are down-regulated by amiloride. It is clear that the inhibition of vacuolar Na(+)/H(+) antiporter by amiloride could disrupt Na(+) accumulation of leaf, and reduce Na(+) uptake by root and Na(+) transport from stem to leaf, thus resulting in the growth inhibition of Z. xanthoxylum exposed to salt.

Histone modifications associated with drought tolerance in the desert plant Zygophyllum dumosum Boiss.

Zygophyllum dumosum Boiss. is a perennial Saharo-Arabian phytogeographical element and a dominant shrub on the rocky limestone southeast-facing slopes of the Negev desert. The plant is highly active during the winter, and semideciduous during the dry summer, i.e., it sheds its leaflets, while leaving the thick, fleshy petiole green and rather active during the dry season. Being resistant to extreme perennial drought, Z. dumosum appears to provide an intriguing model plant for studying epigenetic mechanisms associated with drought tolerance in natural habitats. The transition from the wet to the dry season was accompanied by a significant decrease in nuclear size and with posttranslational modifications of histone H3 N-terminal tail. Dimethylation of H3 at lysine 4 (H3K4)--a modification associated with active gene expression--was found to be high during the wet season but gradually diminished on progression to the dry season. Unexpectedly, H3K9 di- and trimethylation as well as H3K27 di- and trimethylation could not be detected in Z. dumosum; H3K9 monomethylation appears to be prominent in Z. dumosum during the wet but not during the dry season. Contrary to Z. dumosum, H3K9 dimethylation was detected in other desert plants, including Artemisia sieberi, Anabasis articulata and Haloxylon scoparium. Taken together, our results demonstrate dynamic genome organization and unique pattern of histone H3 methylation displayed by Z. dumosum, which could have an adaptive value in variable environments of the Negev desert.

[Water parameters of desert xeric shrubs in west Erdos region].

By using PV technique, this paper studied the turgor pressure (psi P), cell elastic modulus (epsilon), and relative cell volume (RCV) of super xerophytes Potaninia mongolica, Reaumuria soongorica, Tetraena mongolica and Zygophyllum xanthoxylon in west Alashan, with the relationships among the parameters analyzed. The results showed that R. soongorica had the strongest ability to maintain maximum turgor pressure (a = 2.4593). The four plants maintained their turgor pressure by different ways, i.e., P. mongolica maintained it by elastic adjustment (epsilon max = 8.4005 MPa), R. soongorica by osmotic adjustment (psi pi100 = -3.1302 MPa; psi0 = -3.5074 MPa), T. mongolica by both osmotic and elastic adjustment, and Z. xanthoxylon by osmotic adjustment, which had weak adjustment ability. The cell wall of P. mongolica was soft and highly elastic, benefiting to the water absorption by root and stem and to the fast water transmission. T. mongolica also had relatively soft and high elastic cell wall, and its psi P, and epsilon changed slowly with decreasing RCV, suggesting that this plant had strong ability of holding water and resisting dehydration.

Dynamic responses of photosystem II in the Namib Desert shrub, Zygophyllum prismatocarpum, during and after foliar deposition of limestone dust.

The effects of limestone dust deposition on vegetation in desert ecosystems have not yet been reported. We investigated these effects in a succulent shrub from the Namib Desert at a limestone quarry near Skorpion Zinc mine (Namibia). Effects of limestone dust were determined in Zygophyllum prismatocarpum (dollar bush) plants with heavy, moderate and no visible foliar dust cover by means of chlorophyll a fluorescence measurements. Limestone dust deposition decreased overall plant performance through loss of chlorophyll content, inhibition of CO(2) assimilation, uncoupling of the oxygen-evolving complex and decreased electron transport. Importantly, dynamic recovery occurred after termination of limestone extraction at the quarry. Recovery was accelerated by rainfall, mainly because of dust removal from leaves and stimulation of new growth. These results indicate that limestone dust has severe effects on photosynthesis in desert shrubs, but that recovery is possible and that, in arid environments, this process is modulated by rainfall.