Alleviation of Zn toxicity by low water availability.

Heavy metal contamination and drought are expected to increase in large areas worldwide. However, their combined effect on plant performance has been scantly analyzed. This study examines the effect of Zn supply at different water availabilities on morpho-physiological traits of Quercus suber L. in order to analyze the combined effects of both stresses. Seedlings were treated with four levels of zinc from 3 to 150 µM and exposed to low watering (LW) or high watering (HW) frequency in hydroponic culture, using a growth chamber. Under both watering regimes, Zn concentration in leaves and roots increased with Zn increment in nutrient solution. Nevertheless, at the highest Zn doses, Zn tissue concentrations were almost twice in HW than in LW seedlings. Functional traits as leaf photosynthetic rate and root hydraulic conductivity, and morphological traits as root length and root biomass decreased significantly in response to Zn supply. Auxin levels increased with Zn concentrations, suggesting the involvement of this phytohormone in the seedling response to this element. LW seedlings exposed to 150 µM Zn showed higher root length and root biomass than HW seedlings exposed to the same Zn dose. Our results suggest that low water availability could mitigate Zn toxicity by limiting internal accumulation. Morphological traits involved in the response to both stresses probably contributed to this response.

Root architecture and hydraulic conductance in nutrient deprived Pistacia lentiscus L. seedlings.

Plants respond to low nutrient availability by modifying root morphology and root system topology. Root responses to nitrogen (N) and phosphorus (P) limitation may affect plant capacity to withstand water stress. But studies on the effect of nutrient availability on plant ability to uptake and transport water are scarce. In this study, we assess the effect of nitrogen and phosphorus limitation on root morphology and root system topology in Pistacia lentiscus L seedlings, a common Mediterranean shrub, and relate these changes to hydraulic conductivity of the whole root system. Nitrogen and phosphorus deprivation had no effect on root biomass, but root systems were more branched in nutrient limited seedlings. Total root length was higher in seedlings subjected to phosphorus deprivation. Root hydraulic conductance decreased in nutrient-deprived seedlings, and was related to the number of root junctions but not to other architectural traits. Our study shows that changes in nutrient availability affect seedling water use by modifying root architecture. Changes in nutrient availability should be taken into account when evaluating seedling response to drought.

Ecology and functional roles of biological soil crusts in semi-arid ecosystems of Spain.

Biological soil crusts (BSCs), composed of lichens, cyanobacteria, mosses, liverworts and microorganisms, are key biotic components of arid and semi-arid ecosystems worldwide. Despite they are widespread in Spain, these organisms have been historically understudied in this country. This trend is beginning to change as a recent wave of research has been identifying BSCs as a model ecological system. Many studies and research projects carried out in Spain have explored the role of BSCs on water, carbon and nitrogen fluxes, the interactions between BSCs and vascular plants, their dynamics after disturbances, and their response to global change, among other topics. In this article we review the growing body of research on BSCs available from semi-arid areas of Spain, highlighting its importance for increasing our knowledge on this group of organisms. We also discuss how it is breaking new ground in emerging research areas on the ecology of BSCs, and how it can be use to guide management and restoration efforts. Finally, we provide directions for future research on the ecology of BSCs in Spain and abroad.

Sensitivity to zinc of Mediterranean woody species important for restoration.

Heavy metals have increased in natural woodlands and shrublands over the last several decades as a consequence of anthropogenic activities. However, our knowledge of the effects of these elements on woody species is scarce. In this study, we examined the responses of six Mediterranean woody species to increasing levels of zinc in hydroponic culture and discussed the possible implications for the restoration of contaminated sites. The species used, Pinus pinea L., Pinus pinaster Ait., Pinus halepensis Mill., Tetraclinis articulata (Vahl) Mast., Rhamnus alaternus L. and Quercus suber L. represent a climatic gradient from dry sub-humid to semi-arid conditions. Zinc concentrations in shoots ranged from 53 microg g(-1) in Q. suber to 382 microg g(-1) in T. articulata and were well below the levels found in roots. Zinc inhibited root elongation and root biomass and changed the root length distribution per diameter class, but the magnitude of the effects was species-specific. Only P. halepensis and Q. suber showed toxicity symptoms in aboveground parts. Species more characteristic from xeric environments (T. articulata, R. alaternus and P. halepensis) were more sensitive to zinc than species from mesic environments (Q. suber, P. pinaster and P. pinea). According to the Zn responses and bioaccumulation, Q. suber P. pinea and P. halepensis are the best candidates for field trials to test the value of woody species to restore contaminated sites. None of the species tested seemed suitable for phytoremediation.

Degradation of organic pollutants in Mediterranean forest soils amended with sewage sludge.

The degradation of two groups of organic pollutants in three different Mediterranean forest soils amended with sewage sludge was studied for nine months. The sewage sludge produced by a domestic water treatment plant was applied to soils developed from limestone, marl and sandstone, showing contrasting alkalinity and texture. The compounds analysed were: linear alkylbenzene sulphonates (LAS) with a 10-13 carbon alkylic chain, and nonylphenolic compounds, including nonylphenol (NP) and nonylphenol ethoxylates with one and two ethoxy groups (NP1EO+NP2EO). These compounds were studied because they frequently exceed the limits proposed for sludge application to land in Europe. After nine months, LAS decomposition was 86-96%, and NP+NP1EO+NP2EO decomposition was 61-84%, which can be considered high. Temporal trends in LAS and NP+NP1EO+NP2EO decomposition were similar, and the concentrations of both types of compounds were highly correlated. The decomposition rates were higher in the period of 6-9 months (summer period) than in the period 0-6 months (winter+spring period) for total LAS and NP+NP1EO+NP2EO. Differences in decay rates with regard to soil type were not significant. The average values of decay rates found are similar to those observed in agricultural soils.

Sensitivity of Mediterranean woody seedlings to copper, nickel and zinc.

The restoration of heavy metal contaminated areas requires information on the response of native plant species to these contaminants. The sensitivity of most Mediterranean woody species to heavy metals has not been established, and little is known about phytotoxic thresholds and environmental risks. We have evaluated the response of four plant species commonly used in ecological restoration, Pinus halepensis, Pistacia lentiscus, Juniperus oxycedrus, and Rhamnus alaternus, grown in nutrient solutions containing a range of copper, nickel and zinc concentrations. Seedlings of these species were exposed to 0.048, 1 and 4 microM of Cu; 0, 25 and 50 microM of Ni; and 0.073, 25 and 100 microM of Zn in a hydroponic silica sand culture for 12 weeks. For all four species, the heavy metal concentration increased in plants as the solution concentration increased and was always higher in roots than in shoots. Pinus halepensis and P. lentiscus showed a higher capacity to accumulate metals in roots than J. oxycedrus and R. alaternus, while the allocation to shoots was considerably higher in the latter two. Intermediate heavy-metal doses enhanced biomass accumulation, whereas the highest doses resulted in reductions in biomass. Decreases in shoot biomass occurred at internal concentrations ranging from 25 to 128 microg g-1 of Zn, and 1.7 to 4.1 microg g( -1) of Cu. Nickel phytoxicity could not be established within the range of doses used. Rhamnus alaternus and J. oxycedrus showed higher sensitivity to Cu and Zn than P. halepensis and, especially, P. lentiscus. Contrasted responses to heavy metals must be taken into account when using Mediterranean woody species for the restoration of heavy metal contaminated sites.

Response of Pinus halepensis Mill. seedlings to biosolids enriched with Cu, Ni and Zn in three Mediterranean forest soils.

We investigated the response of Pinus halepensis seedlings to the application of biosolids enriched with Cu, Ni and Zn on three Mediterranean forest soils under semiarid conditions. One-year-old seedlings were planted in lysimeters on soils developed from marl, limestone and sandstone which were left unamended, amended with biosolids, or amended with biosolids enriched in Cu, Ni and Zn. Enriched biosolids increased plant heavy metal concentration, but always below phytotoxic levels. Seedlings receiving unenriched biosolids showed a weak reduction in Cu and Zn concentration in needles, negatively affecting physiological status during drought. This effect was alleviated by the application of enriched sludge. Sewage sludge with relatively high levels of Cu, Zn and Ni had minor effects on plant performance on our experimental conditions. Results suggest that micronutrient limitations in these soils may be alleviated by the application of biosolids with a higher Cu, Zn and Ni content than those established by current regulations.

Watering, fertilization, and slurry inoculation promote recovery of biological crust function in degraded soils.

Biological soil crusts are very sensitive to human-induced disturbances and are in a degraded state in many areas throughout their range. Given their importance in the functioning of arid and semiarid ecosystems, restoring these crusts may contribute to the recovery of ecosystem functionality in degraded areas. We conducted a factorial microcosm experiment to evaluate the effects of inoculation type (discrete fragments vs slurry), fertilization (control vs addition of composted sewage sludge), and watering frequency (two vs five times per week) on the cyanobacterial composition, nitrogen fixation, chlorophyll content, and net CO2 exchange rate of biological soil crusts inoculated on a semiarid degraded soil from SE Spain. Six months after the inoculation, the highest rates of nitrogen fixation and chlorophyll a content were found when the biological crusts were inoculated as slurry, composted sewage sludge was added, and the microcosms were watered five times per week. Net CO2 exchange rate increased when biological crusts were inoculated as slurry and the microcosms were watered five times per week. Denaturing gradient gel electrophoresis fingerprints and phylogenetic analyses indicated that most of the cyanobacterial species already present in the inoculated crust had the capability to spread and colonize the surface of the surrounding soil. These analyses showed that cyanobacterial communities were less diverse when the microcosms were watered five times per week, and that watering frequency (followed in importance by the addition of composted sewage sludge and inoculation type) was the treatment that most strongly influenced their composition. Our results suggest that the inoculation of biological soil crusts in the form of slurry combined with the addition of composted sewage sludge could be a suitable technique to accelerate the recovery of the composition and functioning of biological soil crusts in drylands.

Do positive interactions increase with abiotic stress? A test from a semi-arid steppe.

Theoretical models predict that the relative importance of facilitation and competition may vary inversely across gradients of abiotic stress. However, these predictions have not been thoroughly tested in the field, especially in semi-arid environments. In this study, we evaluated how the net effect of the tussock grass Stipa tenacissima on the shrub Pistacia lentiscus varied across a gradient of abiotic stress in semi-arid Mediterranean steppes. We fitted the relationship between accumulated rainfall and the relative neighbour index (our measures of abiotic stress and of the net effect of S. tenacissima on P. lentiscus, respectively), which varied across this gradient, to a quadratic model. Competitive interactions dominated at both extremes of the gradient. Our results do not support established theory. Instead, they suggest that a shift from facilitation to competition under high abiotic stress conditions is likely to occur when the levels of the most limiting resource are so low that the benefits provided by the facilitator cannot overcome its own resource uptake.