Precipitation pulse use by an invasive woody legume: the role of soil texture and pulse size

Plant metabolic activity in arid and semi-arid environments is largely tied to episodic precipitation events or “pulses”. The ability of plants to take up and utilize rain pulses during the growing season in these water-limited ecosystems is determined in part by pulse timing, intensity and amount, and by hydrological properties of the soil that translate precipitation into plant-available soil moisture. We assessed the sensitivity of an invasive woody plant, velvet mesquite (Prosopis velutina Woot.), to large (35 mm) and small (10 mm) isotopically labeled irrigation pulses on two contrasting soil textures (sandy-loam vs. loamy-clay) in semi-desert grassland in southeastern Arizona, USA. Predawn leaf water potential (Ψ_pd), the isotopic abundance of deuterium in stem water (δD), the abundance of ¹³C in soluble leaf sugar (δ¹³C), and percent volumetric soil water content (θ_v) were measured prior to irrigation and repeatedly for 2 weeks following irrigation. Plant water potential and the percent of pulse water present in the stem xylem indicated that although mesquite trees on both coarse- and fine-textured soils quickly responded to the large irrigation pulse, the magnitude and duration of this response substantially differed between soil textures. After reaching a maximum 4 days after the irrigation, the fraction of pulse water in stem xylem decreased more rapidly on the loamy-clay soil than the sandy-loam soil. Similarly, on both soil textures mesquite significantly responded to the 10-mm pulse. However, the magnitude of this response was substantially greater for mesquite on the sandy-loam soil compared to loamy-clay soil. The relationship between Ψ_pd and δ¹³C of leaf-soluble carbohydrates over the pulse period did not differ between plants at the two sites, indicating that differences in photosynthetic response of mesquite trees to the moisture pulses was a function of soil water availability within the rooting zone rather than differences in plant biochemical or physiological constraints. Patterns of resource acquisition by mesquite during the dynamic wetting–drying cycle following rainfall pulses is controlled by a complex interaction between pulse size and soil hydraulic properties. A better understanding of how this interaction affects plant water availability and photosynthetic response is needed to predict how grassland structure and function will respond to climate change.     

Production and population ecology ofPhyllospadix iwatensis Makino. I. Leaf growth and biomass in an intertidal zone

In an intertidal zone on Choshi coast, Japan,Phyllospadix iwatensis Makino emerges at daytime in spring and summer, while at night time in winter. The plants therefore experience seasonally different stresses caused by emergence, for example, intense light, ultraviolet rays, extreme temperature and desiccation, all of which the plants are unable to avoid during daytime emergence. Seasonal changes in the biomass and LAI suggest that the optimum periods for growth ofP. iwatensis would be in March when the emergent period is short or nil and light availability is high while water temperature is not too low. Dense foliage and low canopy height ofP. iwatensis in the intertidal zone relieve the plant from the stresses in emergent periods and from the disturbance caused by strong water movement in some coastal areas with active wave action.     

Hydraulic redistribution in a stand of <Emphasis Type="Italic">Artemisia tridentata</Emphasis>: evaluation of benefits to transpiration assessed with a simulation model

The significance of soil water redistribution facilitated by roots (an extension of "hydraulic lift", here termed hydraulic redistribution) was assessed for a stand of Artemisia tridentata using measurements and a simulation model. The model incorporated water movement within the soil via unsaturated flow and hydraulic redistribution and soil water loss from transpiration. The model used Buckingham-Darcy's law for unsaturated flow while hydraulic redistribution was developed as a function of the distribution of active roots, root conductance for water, and relative soil-root (rhizosphere) conductance for water. Simulations were conducted to compare model predictions with time courses of soil water potential at several depths, and to evaluate the importance of root distribution, soil hydraulic conductance and root xylem conductance on transpiration rates and the dynamics of soil water. The model was able to effectively predict soil water potential during a summer drying cycle, and the rapid redistribution of water down to 1.5 m into the soil column after rainfall events. Results of simulations indicated that hydraulic redistribution could increase whole canopy transpiration over a 100-day drying cycle. While the increase was only 3.5% over the entire 100-day period, hydraulic redistribution increased transpiration up to 20.5% for some days. The presence of high soil water content within the lower rooting zone appears to be necessary for sizeable increases in transpiration due to hydraulic redistribution. Simulation results also indicated that root distributions with roots concentrated in shallow soil layers experienced the greatest increase in transpiration due to hydraulic redistribution. This redistribution had much less effect on transpiration with more uniform root distributions, higher soil hydraulic conductivity and lower root conductivity. Simulation results indicated that redistribution of water by roots can be an important component in soil water dynamics, and the model presented here provides a useful approach to incorporating hydraulic redistribution into larger models of soil processes.     

Increased drought tolerance of mycorrhizal onion plants caused by improved phosphorus nutrition

Onion plants (Allium cepa L, cv. Downing Yellow Globe) grown in pots and infected by the mycorrhizal fungusGlomus etunicatus Becker and Gerdemann were more drought tolerant than were non-mycorrhizal individials when exposed to several periods of soil water stress separated by periods of high water supply, as shown by greater fresh and dry weights and higher tissue phosphorus levels in the mycorrhizal plants. The tissues of stressed, non-mycorrhizal plants were deficient in P, despite the fact that only non-mycorrhizal plants were fertilized with high levels of P (26 mg P per 440 g soil). Differences in plant water relations (leaf water potentials or transpiration rates) and changes in soil P levels which may have affected plant growth were investigated, and discounted as factors important for the results. The P nutrition of plants has been implicated in the ability of plants to tolerate drought and it was concluded that the ability of the mycorrhizal fungus to maintain adequate P nutrition in the onions during soil water stress was a major factor in the improved drought tolerance. Infection of the root by the fungus was found not to be affected by water stress or P fertilization but fungal reproduction, as determined by spore numbers in the soil, was decreased by water stress and by P fertilization.Michigan Agricultural Experiment Station Article No. 10050     

A multi-scale perspective of water pulses in dryland ecosystems: climatology and ecohydrology of the western USA

In dryland ecosystems, the timing and magnitude of precipitation pulses drive many key ecological processes, notably soil water availability for plants and soil microbiota. Plant available water has frequently been viewed simply as incoming precipitation, yet processes at larger scales drive precipitation pulses, and the subsequent transformation of precipitation pulses to plant available water are complex. We provide an overview of the factors that influence the spatial and temporal availability of water to plants and soil biota using examples from western USA drylands. Large spatial- and temporal-scale drivers of regional precipitation patterns include the position of the jet streams and frontal boundaries, the North American Monsoon, El Niño Southern Oscillation events, and the Pacific Decadal Oscillation. Topography and orography modify the patterns set up by the larger-scale drivers, resulting in regional patterns (10²–10⁶ km²) of precipitation magnitude, timing, and variation. Together, the large-scale and regional drivers impose important pulsed patterns on long-term precipitation trends at landscape scales, in which most site precipitation is received as small events (<5 mm) and with most of the intervals between events being short (<10 days). The drivers also influence the translation of precipitation events into available water via linkages between soil water content and components of the water budget, including interception, infiltration and runoff, soil evaporation, plant water use and hydraulic redistribution, and seepage below the rooting zone. Soil water content varies not only vertically with depth but also horizontally beneath versus between plants and/or soil crusts in ways that are ecologically important to different plant and crust types. We highlight the importance of considering larger-scale drivers, and their effects on regional patterns; small, frequent precipitation events; and spatio-temporal heterogeneity in soil water content in translating from climatology to precipitation pulses to the dryland ecohydrology of water availability for plants and soil biota.     

The combined effects of a long‐term experimental drought and an extreme drought on the use of plant‐water sources in a Mediterranean forest

Vegetation in water‐limited ecosystems relies strongly on access to deep water reserves to withstand dry periods. Most of these ecosystems have shallow soils over deep groundwater reserves. Understanding the functioning and functional plasticity of species‐specific root systems and the patterns of or differences in the use of water sources under more frequent or intense droughts is therefore necessary to properly predict the responses of seasonally dry ecosystems to future climate. We used stable isotopes to investigate the seasonal patterns of water uptake by a sclerophyll forest on sloped terrain with shallow soils. We assessed the effect of a long‐term experimental drought (12 years) and the added impact of an extreme natural drought that produced widespread tree mortality and crown defoliation. The dominant species, Quercus ilex, Arbutus unedo and Phillyrea latifolia, all have dimorphic root systems enabling them to access different water sources in space and time. The plants extracted water mainly from the soil in the cold and wet seasons but increased their use of groundwater during the summer drought. Interestingly, the plants subjected to the long‐term experimental drought shifted water uptake toward deeper (10–35 cm) soil layers during the wet season and reduced groundwater uptake in summer, indicating plasticity in the functional distribution of fine roots that dampened the effect of our experimental drought over the long term. An extreme drought in 2011, however, further reduced the contribution of deep soil layers and groundwater to transpiration, which resulted in greater crown defoliation in the drought‐affected plants. This study suggests that extreme droughts aggravate moderate but persistent drier conditions (simulated by our manipulation) and may lead to the depletion of water from groundwater reservoirs and weathered bedrock, threatening the preservation of these Mediterranean ecosystems in their current structures and compositions.     

Shoot dieback during prolonged drought in Ceanothus (Rhamnaceae) chaparral of California: a possible case of hydraulic failure

Progressive diebacks of outer canopy branchlets of Ceanothus crassifolius were repeatedly observed after rainless periods up to 9 mo in duration in the Santa Monica Mountains of southern California. Mean xylem pressures of branchlets near the end of drought were as low as -11.2 MPa (N = 22) with a mean of about 60 dead branchlets per shrub. Inoculation (N = 15) with three species of fungi previously isolated from the same population of C. crassifolius did not promote dieback, suggesting that the observed decline was not fungal induced, as had been proposed. Further, at least 50% of healthy-appearing twigs, without symptoms of dieback, contained isolatible endophytic fungi. We used a centrifugal force method to determine the range of xylem pressure causing cavitation (vulnerability curves) for branchlets (N = 12) and roots (N = 16). We combined vulnerability curves with soil texture data (N = 6) into a water transport model that estimated the critical values (P(Lcrit)) of leaf xylem pressure associated with the loss of water from soil to foliage. Maximum P(Lcrit) was between -10 and -11 MPa and within the range of minimum measured xylem pressures of branchlets during drought and dieback. Branchlet dieback correlated with seasonal declines in xylem pressure in concert with declining safety margins from hydraulic failure. Symptoms of dieback were duplicated in the field by partially severing stem xylem that normally supplied branchlets with water. Taken together, these results indicate that loss of hydraulic conductance to foliage was the probable cause of the observed dieback in C. crassifolius. Partial dieback of peripheral branchlets, and its attendant reduction in evaporative surface area, may be a last-resort mechanism for whole-plant water conservation and drought survival in this species.     

Plant morphology and root hydraulics are altered by nutrient deficiency in Pistacia lentiscus (L.)

The plants in arid and semiarid areas are often limited by water and nutrients. Morpho-functional adjustments to improve nutrient capture may have important implications on plant water balance, and on plant capacity to withstand drought. Several studies have shown that N and P deficiencies may decrease plant hydraulic conductance. Surprisingly, studies on the implications of nutrient limitations on water use in xerophytes are scarce. We have evaluated the effects of strong reductions in nitrogen and phosphorus availability on morphological traits and hydraulic conductance in seedlings of a common Mediterranean shrub, Pistacia lentiscus L.. Nitrogen deficiency resulted in a decrease in aboveground biomass accumulation, but it did not affect belowground biomass accumulation or root morphology. Phosphorus-deficient plants showed a decrease in leaf area, but no changes in aboveground biomass. Root length, root surface area, and specific root length were higher in phosphorus-deficient plants than in control plants. Nitrogen and phosphorus deficiency reduced both root hydraulic conductance and root hydraulic conductance scaled by total root surface area. On the other hand, nutrient limitations did not significantly affect root conductance per unit of foliar surface area. Thus, adaptation to low nutrient availability did not affect seedling capacity for maintaining water supply to leaves. The implications for drought resistance and survival during seedling establishment in semi-arid environments are discussed.     

Plant and soil nitrogen dynamics in California annual grassland

Seasonal changes in soil water and nitrogen availability were related to the phenology and growth of plants in California annual grassland. Plant accumulation of nitrogen was mainly confined to two short periods of the year: fall and early spring. At these times, plants were in the vegetative growth phase, roots were growing rapidly and soil moisture was high. During these periods, soil nitrate was low or depleted. High flux of nitrogen in this ecosystem, however, is indicated by the rapid disappearance of the previous year's detrital material, high microbial biomass, and high mineralizable nitrogen and nitrification potential.At the end of the summer drought, significant amounts of the previous year's detrital material had disappeared, chloroform-labile N (expressed as microbial biomass N) was at its seasonal maximum, and soil inorganic nitrogen pools were high. This suggests inorganic nitrogen flux during the drought period. The drought escaper life history characteristics of annual grasses in California annual grassland, however, may prevent plants from utilizing available nitrogen during a large part of the year.     

Root deployment and shoot growth for two desert species in response to soil rockiness

Soil texture, as well as the presence of rocks, can determine the water status, growth, and distribution of plants in arid environments. The effects of soil rockiness and soil particle size distribution on shoot and root growth, root system size, rooting depth, and water relations were therefore investigated for the Crassulacean acid metabolism leaf succulent Agave deserti and the C(4) bunchgrass Pleuraphis rigida after precipitation events during the summer and winter/spring rainfall periods in the northwestern Sonoran Desert. The soils at the field site varied from sandy (<3% rocks by volume) to rocky (up to 35% rocks), with greater water availability at higher water potentials for sandy than for rocky soils. Although A. deserti was absent from the sandiest sites, its shoot and root growth during both rainfall periods were greatest in comparatively sandier sites and decreased as the soil rock content increased. Furthermore, A. deserti had twofold greater root surface area, root?:?leaf area ratio, and mean rooting depth at sandier than at rocky sites. As for A. deserti, shoot growth was greater for P. rigida at the sandier sites than at the rockier sites, even though its root surface area and mean rooting depth did not vary significantly. After early spring rainfall events, the leaf water potential for A. deserti did not differ between rocky and sandy sites, but transpiration rates were almost twofold greater at rocky than at sandy sites. During the same period, P. rigida had lower leaf water potentials and 25% lower transpiration rates at rocky than at sandy sites. The greater variability in the deployment of the root systems of A. deserti in response to soil rockiness may reflect its evergreen habit and slower growth, which allow it to endure periods of lower water availability than does P. rigida, whose leaves die during drought.     

Protaphorura tricampata, a euedaphic and highly permeable springtail that can sustain activity by osmoregulation during extreme drought

We have investigated drought physiology of soil dwelling springtails since water availability is a key environmental factor governing their performance, and predictions of climate change suggest increased frequency and intensity of summer droughts. Here we show in field and laboratory experiments that the typical euedaphic springtail, Protaphorura tricampata, can survive extreme drought and remain active in soils where the water potential is much lower than equivalent to normal osmotic pressure of springtails. Euedaphic springtails (i.e. species living in deeper soil layers) have an extraordinary ability to up-regulate osmotic pressure of body fluids and prevent water loss in soils where the water potential has dropped to well below the permanent wilting percentage of plants. The ability to regulate osmotic pressure of body fluids is based on accumulation of compatible osmolytes such as sugars and free amino acids. Alanine was the most important osmolyte in P. tricampata and accumulated to concentrations of about 300 μmol g⁻¹ dry weight. It is suggested that alanine also serves as a non-toxic storage of ammonia during drought periods where the normal urine production is hampered. The results presented here show, contrary to convention, that high cuticular permeability is not necessarily accompanied by poor drought tolerance, and is not a good predictor of drought susceptibility.     

Direct nocturnal water transfer from oaks to their mycorrhizal symbionts during severe soil drying

Symbiotic mycorrhizal fungi play an important role in the absorption of soil nutrients and water by most plants. It has been suggested that hydraulically lifted water might maintain the integrity of the external mycorrhizal mycelium during drought. We tested this hypothesis in the obligately mycorrhizal species, coast live oak (Quercus agrifolia), using a microcosm system that separated the effects of hydraulic lift in roots from those in the external mycelium. Mycorrhizal oak seedlings were established in microcosms comprising three discrete compartments for (1) upper roots, (2) tap roots, and (3) external fungal mycelium. Eight months after planting, a drought treatment was initiated: irrigation to the upper root and fungal chambers was terminated and only irrigation to the taproot compartment was maintained. After 3, 12, 30, 50, 70 and 80 days of drought, tracers were injected into the taproot compartment at dusk. At dawn the following morning, mycorrhizal hyphae (EM and AM) and spores (AM) in upper root and fungal compartments were extensively labeled with the tracers. In contrast, no labeling was observed when tracers were injected into the taproot compartment during daytime. Nocturnal water translocation from plant to mycorrhizal fungi occurred in association with hydraulic lift. Saprotrophic/parasitic fungi in the microcosms were not labeled, suggesting a direct water transfer from plants to their mycorrhizal mutualists and not to other fungi in the soil. Even after prolonged drought (70-80 days), mycorrhizal hyphae persisted in soils with water potential values as low as -20 MPa. Maintaining mycorrhizal activity through direct water translocation could potentially improve the nutrient status of deep-rooted plants during periods when the fertile upper soil is dry.     

Reappraisal of drying and wetting effects on C and N mineralization and fluxes in soils

In the next decades, many soils will be subjected to increased drying/wetting cycles or modified water availability considering predicted global changes in precipitation and evapotranspiration. These changes may affect the turnover of C and N in soils, but the direction of changes is still unclear. The aim of the review is the evaluation of involved mechanisms, the intensity, duration and frequency of drying and wetting for the mineralization and fluxes of C and N in terrestrial soils. Controversial study results require a reappraisal of the present understanding that wetting of dry soils induces significant losses of soil C and N. The generally observed pulse in net C and N mineralization following wetting of dry soil (hereafter wetting pulse) is short‐lived and often exceeds the mineralization rate of a respective moist control. Accumulated microbial and plant necromass, lysis of live microbial cells, release of compatible solutes and exposure of previously protected organic matter may explain the additional mineralization during wetting of soils. Frequent drying and wetting diminishes the wetting pulse due to limitation of the accessible organic matter pool. Despite wetting pulses, cumulative C and N mineralization (defined here as total net mineralization during drying and wetting) are mostly smaller compared with soil with optimum moisture, indicating that wetting pulses cannot compensate for small mineralization rates during drought periods. Cumulative mineralization is linked to the intensity and duration of drying, the amount and distribution of precipitation, temperature, hydrophobicity and the accessible pool of organic substrates. Wetting pulses may have a significant impact on C and N mineralization or flux rates in arid and semiarid regions but have less impact in humid and subhumid regions on annual time scales. Organic matter stocks are progressively preserved with increasing duration and intensity of drought periods; however, fires enhance the risk of organic matter losses under dry conditions. Hydrophobicity of organic surfaces is an important mechanism that reduces C and N mineralization in topsoils after precipitation. Hence, mineralization in forest soils with hydrophobic organic horizons is presumably stronger limited than in grassland or farmland soils. Even in humid regions, suboptimal water potentials often restrict microbial activity in topsoils during growing seasons. Increasing summer droughts will likely reduce the mineralization and fluxes of C and N whereas increasing summer precipitation could enhance the losses of C and N from soils.     

Seasonal variation and annual fluctuation in granite outcrop plant communities

Permanent quadrats in granite outcrop plant communities allowed us to monitor seasonal variation and annual fluctuation in community structure. Seasonal species turn-over was significant in communities on shallow soil, but not in communities on deeper soil where seasonal dominance shifts were common. Exceptional meteorological events appeared to mediate phenomena of competitive release in some island communities. A decrease in the abundance of Arenaria uniflora in Lichen-annual island communities, following a spring drought, was correlated with an increase in the abundance of Sedum smallii, a shallower-soil species. Richness in Annual-perennial island communities was higher in spring 1985 than in 1984 or 1986, and this occurred as the dominant species, Senecio tomentosus, temporarily declined in importance following a severe drought in late summer 1984. Significant annual fluctuation in the cover of Viguiera porteri could also be related to variations in the summer precipitation regime. Overall, plant responses to drought were individualistic and depended largely on the timing of these meteorological events in relation to the life-stages and/or the physiological status of the plants.Abbreviations AP = Annual-perennial island communities - HST = Herb-shrub-tree island communities - LA = Lichen-annual island communities - SS = Sedum smallii island communities     

Recycling of nitrogen in herbivore feces: plant recovery,herbivore assimilation,soil retention,and leaching losses

Herbivores directly and indirectly affect ecosystem functioning in forests. Feces deposition is a direct effect that supplies ephemeral N pulses to soils. Herbivore-mediated changes in plant N allocation and uptake are indirect effects that can also influence soil N availability. These effects may interact if defoliation influences the ability of plants to recover fecal N, and this may affect subsequent generations of herbivores. We added ¹⁵N-enriched insect feces (frass) to a series of replicated red oak, Quercus rubra, mesocosms that had been damaged experimentally and then followed the frass N over the course of 2 years. In the first season, some frass N was mineralized in the soil and leached in organic form from the mesocosms within 1 week of deposition. Within 1 month, frass N had been acquired by the oaks and enriched the foliage; late-season herbivores assimilated the frass N within the same growing season. In the second season, herbivore damage from the previous year lowered total leaf N contents and ¹⁵N recovered in the foliage. A subsequent cohort of early-season herbivores fed on this foliage consequently derived less of their N from the previous year’s frass, and feral leaf rollers colonized fewer of these saplings. The 0- to 5-cm soil fraction was the largest N sink measured, and 42% of the frass N was recovered in the soil. The results demonstrate that: (1) some frass N can be recycled rapidly into foliage and assimilated by successive cohorts of herbivore within the same season; (2) damage can affect N allocation in the following year’s foliage, influencing N availability to and host selection by herbivores; and (3) leaching losses occur soon after deposition but are buffered by soil pools, which are the largest sinks for frass N.     

Ecohydrology of Dry Regions: Storage versus Pulse Soil Water Dynamics

Although arid and semiarid regions are defined by low precipitation, the seasonal timing of temperature and precipitation can influence net primary production and plant functional type composition. The importance of precipitation seasonality is evident in semiarid areas of the western U.S., which comprise the Intermountain (IM) zone, a region that receives important winter precipitation and is dominated by woody plants and the Great Plains (GP), a region that receives primarily summer precipitation and is dominated by perennial grasses. Although these general relationships are well recognized, specific differences in water cycling between these regions have not been well characterized. We used a daily time step soil water simulation model and twenty sites from each region to analyze differences in soil water dynamics and ecosystem water balance. IM soil water patterns are characterized by storage of water during fall, winter, and spring resulting in relatively reliable available water during spring and early summer, particularly in deep soil layers. By contrast, GP soil water patterns are driven by pulse precipitation events during the warm season, resulting in fluctuating water availability in all soil layers. These contrasting patterns of soil water—storage versus pulse dynamics—explain important differences between the two regions. Notably, the storage dynamics of the IN sites increases water availability in deep soil layers, favoring the deeper rooted woody plants in that region, whereas the pulse dynamics of the Great Plains sites provide water primarily in surface layers, favoring the shallow-rooted grasses in that region. In addition, because water received when plants are either not active or only partially so is more vulnerable to evaporation and sublimation than water delivered during the growing season, IM ecosystems use a smaller fraction of precipitation for transpiration (47%) than GP ecosystems (49%). Recognizing the pulse-storage dichotomy in soil water regimes between the IM and GP regions may be useful for understanding the potential influence of climate changes on soil water patterns and resulting dominant plant functional groups in both regions.     

Importance of permafrost as a source of water for plants in east Siberian taiga

Stable oxygen isotope ratios of plant water (sap water) were observed at Spasskaya Pad experimental forest near Yakutsk, Russia in 1997–1999. The ¹⁸O of sap water in larch trees (Larix gmelinii) decreased soon after leaf unfolding every year, indicating that snowmelt water was used in the beginning of summer. During mid to late summer, a clear difference in the water source used by plants was observed between wet summers and severe drought summers. The ¹⁸O values of water in larch trees were high (–17.8 to –16.1) in August 1999 (wet summer), but low (–20.4 to –19.7) in August 1998 (drought summer). These results indicated that plants used rainwater during a wet summer, but meltwater from permafrost was used by plants during a drought summer. One important role of permafrost is to provide a direct source of water for plants in a severe drought summer; another role is to keep surplus water in the soil until the next summer. If this permafrost system is disturbed by future global warming, unique monotypic stands of deciduous larch trees in east Siberia might be seriously damaged in a severe drought summer.     

Species-specific,seasonal, inter-annual,and historically-accumulated changes in foliar terpene emission rates in <Emphasis Type="Italic">Phillyrea latifolia</Emphasis> and <Emphasis Type="Italic">Quercus ilex</Emphasis> submitted to rain exclusion in the Prades Mountains (Catalonia)

Mediterranean vegetation emits large amounts of terpenes. We aimed to study the effects of the decreases in soil water availability forecast for the next decades by global circulation models and ecophysiological models on the terpene emissions by two widely distributed Mediterranean woody species, Phillyrea latifolia L. and Quercus ilex L. We subjected holm oak forest plots to an experimental soil drought of ca. 20% decrease in soil moisture by partial rainfall exclusion and runoff exclusion. We measured the emission rates throughout the seasons for two years with contrasting precipitation and soil moisture (16.6% average in 2003 vs. 6.4% as average in 2005). Among the detected volatile terpenes, only α-pinene and limonene were present in detectable quantities in all of the studied periods. Total terpene emitted ranged from practically zero (spring 2003) to 3.6 and 58.3 μg/(g dry wt h) (winter 2005 and summer 2003 for P. latifolia and Q. ilex, respectively). A clear seasonality was found in the emission rates (they were the highest in summer in both species) and also in the qualitative composition of the emission mix. Maximum emissions of α-pinene occurred in spring and maximum emissions of limonene in winter. Neither the inter-annual differences in water availability nor the rain exclusion treatment significantly affected the emissions in P. latifolia, but Q. ilex showed by 17% lower emissions during the drier second year of study, 2005, but more than two- and threefold increases with the drought treatment in summer 2003 and in summer 2005, respectively, showing historical accumulated effects. These results, which show increased monoterpene emission under the moderate drought produced by the treatment and decreased emission under the severe second year drought, and a much higher sensitivity to drought in Q. ilex than in P. latifolia, are useful in understanding the behavior of plant volatiles under Mediterranean conditions and in modeling future emission under changing climate conditions. They show that the usage of current models could lead to under- and overestimations of the emission under summer dry conditions, because most current algorithms are based on light and temperature only.     

Do hydraulic redistribution and nocturnal transpiration facilitate nutrient acquisition in Aspalathus linearis?

The significance of soil water redistribution by roots and nocturnal transpiration for nutrient acquisition were assessed for deep-rooted 3-year-old leguminous Aspalathus linearis shrubs of the Cape Floristic Region (South Africa). We hypothesised that hydraulic redistribution and nocturnal transpiration facilitate nutrient acquisition by releasing moisture in shallow soil to enable acquisition of shallow-soil nutrients during the summer drought periods and by driving water fluxes from deep to shallow soil powering mass-flow nutrient acquisition, respectively. A. linearis was supplied with sub-surface (1-m-deep) irrigation rates of 0, 2 or 4 L day^?1 plant^?1. Some plants were unfertilized, whilst others were surface- or deep-fertilized (1 m depth) with Na¹⁵NO₃ and CaP/FePO₄. We also supplied deuterium oxide (²H₂O) at 1 m depth at dusk and measured its predawn redistribution to shallow soil and plant stems. Hydraulic redistribution of deep water was substantial across all treatments, accounting for 34–72 % of surface-soil predawn moisture. Fourteen days after fertilization, the surface-fertilized plants exhibited increased hydraulic redistribution and increased ¹⁵N and P acquisition with higher rates of deep-irrigation. Deep-fertilization also increased hydraulic redistribution to surface soils, although these plants additionally accumulated ²H₂O in their stem tissue overnight, probably due to nocturnal transpiration. Plants engaged in nocturnal transpiration also increased ¹⁵N and P acquisition from deep fertilizer sources. Thus, both nocturnal transpiration and hydraulic redistribution increased acquisition of shallow soil N and P, possibly through a combination of increased nutrient availability and mobility.     

Stability of above-ground and below-ground processes to extreme drought in model grassland ecosystems: Interactions with plant species diversity and soil nitrogen availability

Extreme drought events have the potential to cause dramatic changes in ecosystem structure and function, but the controls upon ecosystem stability to drought remain poorly understood. Here we used model systems of two commonly occurring, temperate grassland communities to investigate the short-term interactive effects of a simulated 100-year summer drought event, soil nitrogen (N) availability and plant species diversity (low/high) on key ecosystem processes related to carbon (C) and N cycling. Whole ecosystem CO₂ fluxes and leaching losses were recorded during drought and post-rewetting. Litter decomposition and C/N stocks in vegetation, soil and soil microbes were assessed 4 weeks after the end of drought. Experimental drought caused strong reductions in ecosystem respiration and net ecosystem CO₂ exchange, but ecosystem fluxes recovered rapidly following rewetting irrespective of N and species diversity. As expected, root C stocks and litter decomposition were adversely affected by drought across all N and plant diversity treatments. In contrast, drought increased soil water retention, organic nutrient leaching losses and soil fertility. Drought responses of above-ground vegetation C stocks varied depending on plant diversity, with greater stability of above-ground vegetation C to drought in the high versus low diversity treatment. This positive effect of high plant diversity on above-ground vegetation C stability coincided with a decrease in the stability of microbial biomass C. Unlike species diversity, soil N availability had limited effects on the stability of ecosystem processes to extreme drought. Overall, our findings indicate that extreme drought events promote post-drought soil nutrient retention and soil fertility, with cascading effects on ecosystem C fixation rates. Data on above-ground ecosystem processes underline the importance of species diversity for grassland function in a changing environment. Furthermore, our results suggest that plant–soil interactions play a key role for the short-term stability of above-ground vegetation C storage to extreme drought events.