Publications

Responses of terrestrial ecosystems to climate change have been explored in many regions worldwide. While continued drying and warming may alter process rates and deteriorate the state and performance of ecosystems, it could also lead to more fundamental changes in the mechanisms governing ecosystem functioning. Here we argue that climate change will induce unprecedented shifts in these mechanisms in historically wetter climatic zones, towards mechanisms currently prevalent in dry regions, which we refer to as ‘dryland mechanisms’. We discuss 12 dryland mechanisms affecting multiple processes of ecosystem functioning, including vegetation development, water flow, energy budget, carbon and nutrient cycling, plant production and organic matter decomposition. We then examine mostly rare examples of the operation of these mechanisms in non-dryland regions where they have been considered irrelevant at present. Current and future climate trends could force microclimatic conditions across thresholds and lead to the emergence of dryland mechanisms and their increasing control over ecosystem functioning in many biomes on Earth.

Carbonyl sulfide (COS) has emerged as a multi-scale tracer for terrestrial photosynthesis. To infer ecosystem-scale photosynthesis from COS fluxes often requires knowledge of leaf relative uptake (LRU), the concentration-normalized ratio between leaf COS uptake and photosynthesis. However, current mechanistic understanding of LRU variability remains inadequate for deriving robust COS-based estimates of photosynthesis. We derive a set of closed-form equations to describe LRU responses to light, humidity and CO2 based on the Ball–Berry stomatal conductance model and the biochemical model of photosynthesis. This framework reproduces observed LRU responses: decreasing LRU with increasing light or decreasing humidity; it also predicts that LRU increases with ambient CO2. By fitting the LRU equations to flux measurements on a C3 reed (Typha latifolia), we obtain physiological parameters that control LRU variability, including an estimate of the Ball–Berry slope of 7.1 without using transpiration measurements. Sensitivity tests reveal that LRU is more sensitive to photosynthetic capacity than to the Ball–Berry slope, indicating stomatal response to photosynthesis. This study presents a simple framework for interpreting observed LRU variability and upscaling LRU. The stoma-regulated LRU response to CO2 suggests that COS may offer a unique window into long-term stomatal acclimation to elevated CO2.

Temperature is a key control over biological activities from the cellular to the ecosystem scales. However, direct, high‐precision measurements of surface temperature of small objects, such as leaves, under field conditions with large variations in ambient conditions remain rare. Contact methods, such as thermocouples, are prone to large errors. The use of noncontact remote‐sensing methods, such as thermal infrared measurements, provides an ideal solution, but their accuracy has been low (c. 2°C) owing to the necessity for corrections for material emissivity and fluctuations in background radiation Lbg.A novel ‘dual‐reference’ method was developed to increase the accuracy of infrared needle‐leaf surface temperature measurements in the field. It accounts for variations in Lbg and corrects for the systematic camera offset using two reference plates.We accurately captured surface temperature and leaf‐to‐air temperature differences of needle‐leaves in a forest ecosystem with large diurnal and seasonal temperature fluctuations with an uncertainty of ± 0.23°C and ± 0.28°C, respectively.Routine high‐precision leaf temperature measurements even under harsh field conditions, such as demonstrated here, opens the way for investigating a wide range of leaf‐scale processes and their dynamics.

Drylands cover more than 40% of Earth's land surface and occur at the margin of forest distributions due to the limited availability of water for tree growth. Recent elevated temperature and low precipitation have driven greater forest declines and pulses of tree mortality on dryland sites compared to humid sites, particularly in temperate Eurasia and North America. Afforestation of dryland areas has been widely implemented and is expected to increase in many drylands globally to enhance carbon sequestration and benefits to the human environment, but the interplay of sometimes conflicting afforestation outcomes has not been formally evaluated yet. Most previous studies point to conflicts between additional forest area and water consumption, in particular water yield and soil conservation/desalinization in drylands, but were generally confined to local and regional scales. Our global synthesis demonstrates that additional tree cover can amplify water consumption through a nonlinear increase in evapotranspiration-depending on tree species, age, and structure-which will be further intensified by future climate change. In this review we identify substantial knowledge gaps in addressing the dryland afforestation dilemma, where there are trade-offs with planted forests between increased availability of some resources and benefits to human habitats versus the depletion of other resources that are required for sustainable development of drylands. Here we propose a method of addressing comprehensive vegetation carrying capacity, based on regulating the distribution and structure of forest plantations to better deal with these trade-offs in forest multifunctionality. We also recommend new priority research topics for dryland afforestation, including: responses and feedbacks of dryland forests to climate change; shifts in the ratio of ecosystem ET to tree cover; assessing the role of scale of afforestation in influencing the trade-offs of dryland afforestation; and comprehensive modeling of the multifunctionality of dryland forests, including both ecophysiological and socioeconomic aspects, under a changing climate.

•Quantifying the carbon and water economy of non-commercial urban citrus orchards can help recognize the value of such orchards in the urban settings.•A 40 years old citrus orchard within urban setting and sustained on minimal irrigation showed high water use efficiency and significant carbon storage capacity.•The water and carbon use efficiency of the urban orchard were higher than that of a similar commercial ones, in spite of 2x difference in water supply.•Low-density row-crop type orchard help minimize water and carbon losses in evaporation and respiration in the low activity open spaces.Characterizing the carbon and water economy of non-commercial urban citrus orchards can help determine their value in urban settings. This includes provisions of urban ecology and ecosystem services, such as shade, conservation of biodiversity, and carbon sequestration, under space and water limitations that may be particularly suitable in the semi-arid Mediterranean regions. We carried out canopy-scale eddy covariance (EC) measurements of net ecosystem CO2 exchange (NEE) and evapotranspiration (ET), partitioned these fluxes to their components using chamber-based soil fluxes, and combined them with carbon stock to provide a first approximation of the apparent ecosystem carbon turnover rate (τeco). The urban orchard switched from a carbon sink of 0.6 μmol m−2 s−1 to a carbon source of 0.2 μmol m−2 s−1 between winter and summer, with a first approximation of the annual carbon storage capacity of ∼75.3 g m−2 and a total carbon accumulation over its 40 year life span of ∼3014 g m−2. Carbon accumulated predominantly below ground (67 % of total), and soil CO2 effluxes showed low sensitivity to temperature (Q10 ∼1.6) and therefore also to climate warming, but also a fast C turnover rate (∼5.4 y), and therefore sensitivity to disturbances. The rates of ET increased from 0.40 to 1.25 mmol m−2 s−1 between the wet and dry seasons, and was ∼50 % of a similar nearby commercial orchard. Partitioning of the ecosystem carbon and water fluxes indicated high canopy water use efficiency (11.7 μmol CO2/mmol H2O; during the peak activity period). We demonstrate the potential of urban citrus orchards with low supplemental irrigation (50 % compared with commercial orchards in the area) to store significant amounts of carbon with high transpiration efficiencies. The results will help decision making in regard to urban tree planting and the ecological management of urban green spaces and community use in water-limited environments.

Forestation actions are a major tool for both climate-change mitigation and biodiversity conservation. We address two weaknesses in this approach: the little attention given to the negative effects of reduced albedo associated with forestation in many regions, and ignoring the potential of drylands that account for 40% of the global potential land area for forestation. We propose an approach to identify suitable land for forestation and quantify its 'net equivalent carbon stock change' over 80 years of forest lifetime (NESC), accounting for both carbon sequestration and albedo changes. We combined remote-sensing tools with data-based estimates of surface parameters and with published climate matrices, to identify suitable land for forestation actions. We then calculated the cumulative (over 80 years) 'net sequestration potential' (ΔSP), the 'emission equivalent of shortwave radiation forcing' (EESF) due to changes in surface albedo, and, in turn, the combined NESC = ΔSP−EESF, of planting forests with >40% tree-cover. Demonstrating our approach in a large climatically diverse state (Queensland), we identified 14.5 million hectares of potential forestation land in its semi-arid land and show that accounting for the EESF, reduces the climatic benefits of the ΔSP by almost 50%. Nevertheless, it results in a total NESC of 0.72 Gt C accumulated by the end of the century, and 80 years of forestation cycle. This estimated NESC is equivalent to 15% of the projected carbon emissions for the same period in Queensland, for a scenario of no change in emission rates during that period. Our approach extends restoration efforts by identifying new land for forestation and carbon sequestration but also demonstrates the importance of quantifying the climatic value of forestation in drylands.

Drought‐related tree mortality is increasing globally, but the sequence of events leading to it remains poorly understood. To identify this sequence, we used a 2016 tree mortality event in a semi‐arid pine forest where dendrometry and sap flow measurements were carried out in 31 trees, of which seven died. A comparative analysis revealed three stages leading to mortality. First, a decrease in tree diameter in all dying trees, but not in the surviving trees, 8 months “prior to the visual signs of mortality” (PVSM; e.g., near complete canopy browning). Second, a decay to near zero in the diurnal stem swelling/shrinkage dynamics, reflecting the loss of stem radial water flow in the dying trees, 6 months PVSM. Third, cessation of stem sap flow 3 months PVSM. Eventual mortality could therefore be detected long before visual signs were observed, and the three stages identified here demonstrated the differential effects of drought on stem growth, water storage capacity and soil water uptake. The results indicated that breakdown of stem radial water flow and phloem function is a critical element in defining the “point of no return” in the sequence of events leading to mortality of mature trees.

The rate of change in atmospheric CO2 is significantly affected by the terrestrial carbon sink, but the size and spatial distribution of this sink, and the extent to which it can be enhanced to mitigate climate change are highly uncertain. We combined carbon stock (CS) and eddy covariance (EC) flux measurements that were collected over a period of 15 years (2001-2016) in a 55-year-old 30 km2 pine forest growing at the semi-arid timberline (with no irrigating or fertilization). The objective was to constrain estimates of the carbon (C) storage potential in forest plantations in such semi-arid lands, which cover ~ 18% of the global land area. The forest accumulated 145-160 g C m-2 y-1 over the study period based on the EC and CS approaches, with a mean value of 152.5 ± 30.1 g C m-2 y-1 indicating 20% uncertainty in carbon uptake estimates. Current total stocks are estimated at 7,943 ± 323 g carbon m-2 and 372 g nitrogen m-2 . Carbon accumulated mostly in the soil (~71% and 29% for soil and standing biomass carbon, respectively) with long soil carbon turnover time (59 y). Regardless of un-expected disturbances beyond those already observed at the study site, the results support a considerable carbon sink potential in semi-arid soils and forest plantations, and imply that afforestation of even 10% of semi-arid land area under conditions similar to that of the study site, could sequester ~ 0.4 Pg C y-1 over several decades.

The future of forests and their productivity in dry environments will depend on both water availability through precipitation and ecosystem and plant water use characteristics. It is increasingly recognized that better understanding water use patterns and their response to change depends on our ability to partition evapotranspiration (ET). Here, we use chamber-based direct measurements of soil evaporation (Es) in a semi-arid Pinus halepensis forest to partition ET to Es and tree transpiration (Et), to assess the daily and seasonal changes and to compare annual-scale values with measurements carried out at the same site ten years earlier. The ecosystem is characterized by a high annual Es/ET ratio of 0.26, and an Et/ET of 0.63. Es diminished in the long dry season, but as much as 74 ± 5% of the residual flux was due to the re-evaporation of nighttime moisture adsorption, which may provide critical protection from soil drying. Over the 10 years observation period concurrent increase in the transpiration ratio (TR=Et/ET; +29%) and in leaf area index (LAI; +44%) were observed, with the ratio of TR/LAI remaining constant at ~0.31, and with persistently closed hydrological balance (ET/P of 0.94–1.07). The observed Et/ET values are similar to the estimated global mean values, but are attained at a much higher aridity index (5.5) than the mean one, demonstrating the potential for expanding forestation into dry regions.

Photoprotection strategies in a Pinus halepensis Mill. forest at the dry timberline that shows sustained photosynthetic activity during 6-7 month summer drought were characterized and quantified under field conditions. Measurements of chlorophyll fluorescence, leaf-level gas exchange and pigment concentrations were made in both control and summer-irrigated plots, providing the opportunity to separate the effects of atmospheric from soil water stress on the photoprotection responses. The proportion of light energy incident on the leaf surface ultimately being used for carbon assimilation was 18% under stress-free conditions (irrigated, winter), declining to 4% under maximal stress (control, summer). Allocation of absorbed light energy to photochemistry decreased from 25 to 15% (control) and from 50% to 30% (irrigated) between winter and summer, highlighting the important role of pigment-mediated energy dissipation processes. Photorespiration or other non-assimilatory electron flow accounted for 15-20% and ~10% of incident light energy during periods of high and low carbon fixation, respectively, representing a proportional increase in photochemical energy going to photorespiration in summer but a decrease in the absolute amount of photorespiratory CO loss. Resilience of the leaf photochemical apparatus was expressed in the complete recovery of photosystem II (PSII) efficiency (φPSII) and relaxation of the xanthophyll de-epoxidation state on the diurnal cycle throughout the year, and no seasonal decrease in pre-dawn maximal PSII efficiency (Fv/Fm). The response of CO assimilation and photoprotection strategies to stomatal conductance and leaf water potential appeared independent of whether stress was due to atmospheric or soil water deficits across seasons and treatments. The range of protection characteristics identified provides insights into the relatively high carbon economy under these dry conditions, conditions that are predicted for extended areas in the Mediterranean and other regions due to global climate change.

Accurate determination of infrared (IR) emissivity is important for non-contact temperature measurement and for energy balance evaluation in systems that exchange radiation. A method for accurate measurement is proposed based on active modulation of the background radiation. The hemispherical directional reflectance is measured as a proxy for directional emissivity using an IR camera and an integrating sphere, while the background radiation is modulated using an IR emitter and a mechanical shutter. Measurement of the apparent temperature observed by the camera under two different illumination conditions allows the extraction of reflectance and emissivity. The accuracy of the measurement and its sensitivity to surface properties are analyzed, showing uncertainty values as low as 0.004 in some cases. Example measurements of natural and artificial surfaces are presented. (C) 2019 Optical Society of America

Drought-related tree mortality had become a widespread phenomenon in forests around the globe. This process leading to these events and its complexity is not fully understood. Trees in the dry timberline are exposed to ongoing drought, and the available water for transpiration in the soil can determine their survival chances. Recent drought years led to 5%–10% mortality in the semi-arid pine forest of Yatir (Israel). The distribution of dead trees was, however, highly heterogeneous with parts of the forest showing >80% dead trees (D plots) and others with mostly live trees (L plots). At the tree level, visible stress was associated with low pre-dawn leaf water potential at the dry season (−2.8 MPa vs. −2.3 MPa in non-stressed trees), shorter needles (5.5 vs. 7.7 mm) and lower chlorophyll content (0.6 vs. 1 mg/g dw). Trends in tree-ring widths reflected differences in stress intensity (30% narrower rings in stressed compared with unstressed trees), which could be identified 15–20 years prior to mortality. At the plot scale, no differences in topography, soil type, tree age or stand density could explain the mortality difference between the D and L plots. It could only be explained by the higher surface rock cover and in stoniness across the soil profile in the L plots. Simple bucket model simulations using the site’s long-term hydrological data supported the idea that these differences could result in higher soil water concentration (m 3 /m 3 ) in the L plots and extend the time above wilting point by several months across the long dry season. Accounting for subsurface heterogeneity may therefore critical to assessing stand-level response to drought and projecting tree survival, and can be used in management strategies in regions undergoing drying climate trends. 

We investigate the effects of an isolated meso-γ-scale surface heterogeneity for roughness and albedo on the atmospheric boundary-layer (ABL) height, with a case study at a semi-arid forest surrounded by sparse shrubland (forest area: 28km2, forest length in the main wind direction: 7 km). Doppler lidar and ceilometer measurements at this semi-arid forest show an increase in the ABL height over the forest compared with the shrubland on four out of eight days. The differences in the ABL height between shrubland and forest are explained for all days with a model that assumes a linear growth of the internal boundary layer of the forest through the convective ABL upwind of the forest followed by a square-root growth into the stable free atmosphere. For the environmental conditions that existed during our measurements, the increase in ABL height due to large sensible heat fluxes from the forest (600Wm-2 in summer) is subdued by stable stratification in the free atmosphere above the ABL, or reduced by high wind speeds in the mixed layer.

Ecosystem evapotranspiration (ET) can approach annual precipitation (P) often leaving a residual [P-ET], referred to as an ecosystem water yield (WYe). Using a mobile lab, we estimate ET and WYe, in paired forest and nonforest (shrub or grassland) sites along the precipitation gradient (285–755 mm a
−1) in Israel. WYe was 69 mm in the dry sites and was further reduced by ∼51 mm by forestation. Both WYe and the impact of forestation increased in the wetter sites, with forestation reducing WYe by >200 mm, equivalent to ∼30% of the local P. This was associated with increase in ET by a factor of 2.2 and 1.8 in the forest and nonforest sites, respectively, along the rainfall gradient. Losses in WYe due to forestation approached a maximum of ∼200 mm above P ∼ 500 mm, but the forest WYe could vary between ∼300 mm at P = 900 mm and ∼100 mm at P = 500 mm (with equivalent change in WYe between 500 and 300 mm in the nonforest sites), reflecting the increasing “hydrological cost” associated with vegetation ET and the expected climate change in these regions. The results quantify the interactions of land use and climate on ecosystem ET, indicating that in dry climates, afforestation impact on WYe varies significantly across small spatial scales and can reduce WYe with significant impacts on local hydrology. Such impact may be diminished by management (e.g., plant species, thinning, and grazing) but should also consider the trade-offs with other ecosystem services (e.g., carbon sequestration, soil protection, and surface cooling).

The role of secondary circulations has recently been studied in the context of well-defined surface heterogeneity in a semiarid ecosystem where it was found that energy balance closure over a desert-forest system and the structure of the boundary layer was impacted by advection and flux divergence. As a part of the CliFF ("Climate feedbacks and benefits of semi-arid forests", a collaboration between KIT, Germany, and the Weizmann Institute, Israel) campaign, we studied the boundary layer dynamics and turbulent transport of energy corresponding to this effect in Yatir Forest situated in the Negev Desert in Israel. The forest surrounded by small shrubs presents a distinct feature of surface heterogeneity, allowing us to study the differences between their interactions with the atmosphere above by conducting measurements with two eddy covariance (EC) stations and two Doppler lidars. As expected, the turbulence intensity and vertical fluxes of momentum and sensible heat are found to be higher above the forest compared to the shrubland. Turbulent statistics indicative of nonlocal motions are also found to differ over the forest and shrubland and also display a strong diurnal cycle. The production of turbulent kinetic energy (TKE) over the forest is strongly mechanical, while buoyancy effects generate most of the TKE over the shrubland. Overall TKE production is much higher above the forest compared to the shrubland. The forest is also found to be more efficient in dissipating TKE. The TKE budget appears to be balanced on average both for the forest and shrubland, although the imbalance of the TKE budget, which includes the role of TKE transport, is found to be quite different in terms of diurnal cycles for the forest and shrubland. The difference in turbulent quantities and the relationships between the components of TKE budget are used to infer the characteristics of the turbulent transport of energy between the desert and the forest.

Aims: This study investigated the impact of canopy cover and seasonality on litter decay in Mediterranean pine forests to enhance climate predictions. Methods: We conducted litterbag experiments in plots of different tree densities in two Mediterranean pine forests differing in precipitation amounts. In each plot, local litter was placed in forest gaps and under tree canopies for 613 days, starting in the dry season. Results: Litter mass loss was greater in forest gaps than under tree canopies across forests and tree densities. Similarly, a reduction in tree density tended to increase mass loss. Additionally, while the decay rate slowed down from the first to the second wet season, the decay rate remained constant during the first and the second dry season, and the dry seasons contributed 30% to the overall mass loss. Conclusions: Reduction in canopy cover enhances litter decay, and the stability and magnitude of the dry season contribution to annual mass loss have the potential to control litter mass loss when accounting also for the dry periods in the wet season. Combined, the ongoing tree mortality and the predicted prolongation of dry periods due to climate change may enhance litter decay, possibly reducing ecosystem carbon stocks in drylands.

The authors suggest an approach to analyze the effects of small-scale afforestation on the surrounding climate of a large heterogenic area. While simple statistics have difficulty identifying the effect, here a well-known eigenvector technique is used to overcome several specific challenges that result from a limited research region, complex topography, and multiple atmospheric circulation patterns. This approach is applied to investigate the influence of the isolated Yatir forest, at the north edge of Israel's Negev Desert. It was found that this forest does influence the daily climate, primarily seen in the main pattern of the empirical orthogonal function (EOF) of temperature and humidity. The EOF explains 93% and 80 respectively, of the total variance in the data. Although the Yatir forest is small, it is significant in regulating the climate in the nearby surroundings, as it is located in a sharp transition area toward an arid climate. The results are presented as maps of correlation and regression between the normalized principal component time series of each pattern as well as other time series of the raw data and spatially interpolated data stations. Analysis of short-term campaign measurements around the Yatir forest supports the EOF results, and shows the forest's influence to the south, mainly during nighttime when the forest becomes cooler than its surroundings. Overall, results suggest that in areas of transition to semiarid climates, forests regulate the surrounding surface air temperature and humidity fields. Wind analysis based on a complex EOF technique reveals the pattern of the daily cycle of surface wind over the region.

More frequent and intense droughts are projected during the next century, potentially changing the hydrological balances in many forested catchments. Although the impacts of droughts on forest functionality have been vastly studied, little attention has been given to studying the effect of droughts on forest hydrology. Here, we use the Budyko framework and two recently introduced Budyko metrics (deviation and elasticity) to study the changes in the water yields (rainfall minus evapotranspiration) of forested catchments following a climatic drought (2006-2010) in pine forests distributed along a rainfall gradient (P = 280-820 mm yr(-1)) in the Eastern Mediterranean (aridity factor = 0.17-0.56). We use a satellite-based model and meteorological information to calculate the Budyko metrics. The relative water yield ranged from 48% to 8% (from the rainfall) in humid to dry forests and was mainly associated with rainfall amount (increasing with increased rainfall amount) and bedrock type (higher on hard bedrocks). Forest elasticity was larger in forests growing under drier conditions, implying that drier forests have more predictable responses to drought, according to the Budyko framework, compared to forests growing under more humid conditions. In this context, younger forests were shown more elastic than older forests. Dynamic deviation, which is defined as the water yield departure from the Budyko curve, was positive in all forests (i.e., less-than-expected water yields according to Budyko's curve), increasing with drought severity, suggesting lower hydrological resistance to drought in forests suffering from larger rainfall reductions. However, the dynamic deviation significantly decreased in forests that experienced relatively cooler conditions during the drought period. Our results suggest that forests growing under permanent dry conditions might develop a range of hydrological and eco-physiological adjustments to drought leading to higher hydrological resilience

Water stress results in a reduction of the metabolism of plants and in a reorganization of their use of resources geared to survival. In the Mediterranean region, periods of drought accompanied by high temperatures and high irradiance occur in summer. Plants have developed various mechanisms to survive in these conditions by resisting, tolerating or preventing stress. We used three typical Mediterranean tree species in Israel, Pinus halepensis L., Quercus calliprinos and Quercus ithaburensis Webb, as models for studying some of these adaptive mechanisms. We measured their photosynthetic rates (A), stomatal conductance (g s), and terpene emission rates during spring and summer in a geophysical gradient from extremely dry to mesic from Yatir (south, arid) to Birya (north, moist) with intermediate conditions in Solelim. A and g s of P. halepensis were threefold higher in Birya than in Yatir where they remained very low both seasons. Quercus species presented 2–3-fold higher A and g s but with much more variability between seasons, especially for Q. ithaburensis with A and g s that decreased 10–30-fold from spring to summer. Terpene emission rates for pine were not different regionally in spring but they were 5–8-fold higher in Birya than in Yatir in summer (P

Plant phenological development is orchestrated through subtle changes in photoperiod, temperature, soil moisture and nutrient availability. Presently, the exact timing of plant development stages and their response to climate and management practices are crudely represented in land surface models. As visual observations of phenology are laborious, there is a need to supplement long-term observations with automated techniques such as those provided by digital repeat photography at high temporal and spatial resolution. We present the first synthesis from a growing observational network of digital cameras installed on towers across Europe above deciduous and evergreen forests, grasslands and croplands, where vegetation and atmosphere CO2 fluxes are measured continuously. Using colour indices from digital images and using piecewise regression analysis of time series, we explored whether key changes in canopy phenology could be detected automatically across different land use types in the network. The piecewise regression approach could capture the start and end of the growing season, in addition to identifying striking changes in colour signals caused by flowering and management practices such as mowing. Exploring the dates of green-up and senescence of deciduous forests extracted by the piecewise regression approach against dates estimated from visual observations, we found that these phenological events could be detected adequately (RMSE

Soil respiration is the sum of respiration processes in the soil and is a major flux in the global carbon cycle. It is usually assumed that the CO2 efflux is equal to the soil respiration rate. Here we challenge this assumption by combining measurements of CO2 with high-precision measurements of O-2. These measurements were conducted on different ecosystems and soil types and included measurements of air samples taken from the soil profile of three Mediterranean sites: a temperate forest and two alpine forests. Root-free soils from the alpine sites were also incubated in the lab. We found that the ratio between the CO2 efflux and the O-2 influx (defined as apparent respiratory quotient, ARQ) was in the range of 0.14 to 1.23 and considerably deviated from the value of 0.9 +/- 0.1 expected from the elemental composition of average plants and soil organic matter. At the Mediterranean sites, these deviations are explained as a result of CO2 dissolution in the soil water and transformation to bicarbonate ions in these high-pH soils, as well as by carbonate mineral dissolution and precipitation processes. Thus, a correct estimate of the short-term, chamber-based biological respiratory flux in such soils can only be made by dividing the measured soil CO2 efflux by the average (efflux-weighted) soil profile ARQ. Applying this approach to a semiarid pine forest resulted in an estimated short-term biological respiration rate that is 3.8 times higher than the chamber-measured surface CO2. The ARQ values often observed in the more acidic soils were unexpectedly low (

Knowledge of the relationship between soil water dynamics and tree water use is critical to understanding forest response to environmental change in water-limited ecosystems. However, the dynamics in soil water availability for tree transpiration (T-t) cannot be easily deduced from conventional measurements of soil water content (SWC), notably because T-t is influenced by soil water potential ((s)) that, in turn, depends on soil characteristics. Using tree sap flow and water potential and deriving depth-dependent soil water retention curves, we quantified the transpirable soil water content' (tSWC) and its seasonal and inter-annual variations in a semi-arid Pinus halepensis forest. The results indicated that tSWC varied in time and with soil depth. Over one growing season T-t was 57% of rain and 72% of the infiltrated SWC. In early winter, T-t was exclusively supported by soil moisture at the top 10cm (tSWC=11mm), whereas in spring (tSWC>18mm) and throughout the dry season, source water for T-t shifted to 20-40cm, where the maximum fine root density occurs. Simulation with the soil-plant-atmosphere water and energy transport model MuSICA supported the idea that consistent tSWC at the 20-40cm soil layer critically depended on limited water infiltration below 40cm, because of high water retention below this depth. Quantifying tSWC is critical to the precise estimation of the onset and termination of the growing season (when tSWC>0) in this semi-arid ecosystem. Copyright (c) 2013 John Wiley & Sons, Ltd.

Laboratory studies indicate that plant respiratory efficiency may decrease in response to low nutrient availability due to increased partitioning of electrons to the energy-wasteful alternative oxidase (AOX); however, field confirmation of this hypothesis is lacking. We therefore investigated plant respiratory changes associated with succession and retrogression in soils aged from 10 to 120000 years along the Franz Josef soil chronosequence, New Zealand. Respiration rates and electron partitioning were determined based on oxygen isotopic fractionation. Leaf structural traits, foliar nutrient status, carbohydrates and species composition were measured as explanatory variables. Although soil nutrient levels and species composition varied by site along the chronosequence, foliar respiration across all sites and species corresponded strongly with leaf nitrogen concentration (r2=0.8). In contrast, electron partitioning declined with increasing nitrogen/phosphorus (r2=0.23) and AOX activity correlated with phosphorus (r2=0.64). Independently, total respiration was further associated with foliar Cu, possibly linked to its effect on AOX. Independent control of AOX and cytochrome pathway activities is also discussed. These responses of plant terminal respiratory oxidases and therefore respiratory carbon efficiency to multiple nutrient deficiencies demonstrate that modulation of respiratory metabolism may play an important role in plant responses to nutrient gradients.

Transpiration is a fundamental datum in understanding the ecophysiology of planted forests in dry regions and is central to the construction of an ecosystem-level water balance. The present aims were: (i) to measure daily transpiration in a dryland Pinus halepensis Mill. (Aleppo pine) forest and to examine its relationship to environmental conditions such as soil water content and evaporative demand; (ii) to determine the seasonal and annual water balances of the ecosystem; and (iii) to explore management implications in the context of a climate-change scenario of increasing aridity. The study was conducted in the Yatir forest (300 trees ha(-1)) in Israel's semiarid northern Negev, during three consecutive years (starting 2003/4) in which rainfall (R) was 231, 334 and 224 mm, the last designated a drought because of relatively long dry spells between major rain events. Tree transpiration was measured by the heat-pulse method and values were upscaled to the forest canopy level. Independent estimates were obtained for daily ecosystem-level evapotranspiration (ET), and soil and understory vegetation evapotranspiration (E). Daily canopy-level transpiration rate (7) ranged from 0.1 to 1.6 mm d(-1) and showed a highly dynamic and irregular pattern during the rainy season. For the two non-drought years a large part of the variation in T could be attributed to the following simple relationships. When soil water content (SWC) 0.15 m(3) m(-3), potential evapotranspiration (PET) was of paramount importance; quadratic regression of T on PET yielded highly significant relationships and explained a high proportion of the variance in T. During the wet season (215 d), on average, cumulative ET (201 mm) accounted for 0.76 R (R = 263 mm) and

CO2 exchange between terrestrial ecosystems and the atmosphere is key to understanding the feedbacks between climate change and the land surface. In regions with carbonaceous parent material, CO2 exchange patterns occur that cannot be explained by biological processes, such as disproportionate outgassing during the daytime or night-time CO2 uptake during periods when all vegetation is senescent. Neither of these phenomena can be attributed to carbonate weathering reactions, since their CO2 exchange rates are too small. Soil ventilation induced by high atmospheric turbulence is found to explain atypical CO2 exchange between carbonaceous systems and the atmosphere. However, by strongly altering subsurface CO2 concentrations, ventilation can be expected to influence carbonate weathering rates. By imposing ventilation-driven CO2 outgassing in a carbonate weathering model, we show here that carbonate geochemistry is accelerated and does play a surprisingly large role in the observed CO2 exchange pattern of a semi-arid ecosystem. We found that by rapidly depleting soil CO2 during the daytime, ventilation disturbs soil carbonate equilibria and therefore strongly magnifies daytime carbonate precipitation and associated CO2 production. At night, ventilation ceases and the depleted CO2 concentrations increase steadily. Dissolution of carbonate is now enhanced, which consumes CO2 and largely compensates for the enhanced daytime carbonate precipitation. This is why only a relatively small effect on global carbonate weathering rates is to be expected. On the short term, however, ventilation has a drastic effect on synoptic carbonate weathering rates, resulting in a pronounced diel pattern that exacerbates the non-biological behavior of soil-atmosphere CO2 exchanges in dry regions with carbonate soils.

Plants can alter rates of electron transport through the alternative oxidase (AOX) pathway in response to environmental cues, thus modulating respiratory efficiency, but the 18O discrimination method necessary for measuring electron partitioning in vivo has been restricted to laboratory settings. To overcome this limitation, we developed a field-compatible analytical method. Series of plant tissue subsamples were incubated in 12 mL septum-capped vials for 0.54 h before aliquots of incubation air were injected into 3.7 mL evacuated storage vials. Vials were stored for up to 10 months before analysis by mass spectrometry. Measurements were corrected for unavoidable contamination. Additional mathematical tools were developed for detecting and addressing non-linearity (whether intrinsic or due to contamination) in the data used to estimate discrimination values. Initial contamination in the storage vials was 0.03 +/- 0.01 atm; storing the gas samples at -17 degrees C eliminated further contamination effects over 10 months. Discrimination values obtained using our offline incubation and computation method replicated previously reported results over a range of 1031 parts per thousand, with precision generally better than +/- 0.5 parts per thousand. Our method enables large-scale investigations of plant alternative respiration along natural environmental gradients under field conditions.

Urbanization is accelerating across the globe, elevating the importance of studying urban ecology. Urban environments exhibit several factors affecting plant growth and function, including high temperatures (particularly at night), CO2 concentrations and atmospheric nitrogen deposition. We investigated the effects of urban environments on growth in Quercus rubra L. seedlings. We grew seedlings from acorns for one season at four sites along an urban-rural transect from Central Park in New York City to the Catskill Mountains in upstate New York (difference in average maximum temperatures of 2.4 degrees C; difference in minimum temperatures of 4.6 degrees C). In addition, we grew Q. rubra seedlings in growth cabinets (GCs) mimicking the seasonal differential between the city and rural sites (based on a 5-year average). In the field experiment, we found an eightfold increase in biomass in urban-grown seedlings relative to those grown at rural sites. This difference was primarily related to changes in growth allocation. Urban-grown seedlings and seedlings grown at urban temperatures in the GCs exhibited a lower root: shoot ratio (urban similar to 0.8, rural/remote similar to 1.5), reducing below-ground carbon costs associated with construction and maintenance. These urban seedlings instead allocated more growth to leaves than did rural-grown seedlings, resulting in 10-fold greater photosynthetic area but no difference in photosynthetic capacity of foliage per unit area. Seedlings grown at urban temperatures in both the field and GC experiments had higher leaf nitrogen concentrations per unit area than those grown at cooler temperatures (increases of 23% in field, 32% in GC). Lastly, we measured threefold greater C-13 enrichment of respired CO2 (relative to substrate) in urban-grown leaves than at other sites, which may suggest greater allocation of respiratory function to growth over maintenance. It also shows that lack of differences in total R flux in response to en

Nitrogen (N) and water availability are important factors affecting ecosystem productivity that can be influenced by land-use change. We hypothesized that the observed increase in carbon (C) sequestration associated with afforestation of semi-arid sparse shrubland must also be associated with an increase in N input. We tested this hypothesis by reconstructing the ecosystem N budget of two ecosystems, a semi-arid shrubland and a nearby planted pine forest, using measurements augmented with literature-based estimates. Our findings demonstrate that, contrary to our hypothesis, massive C sequestration by the pine forest could be accounted for without a change in the net N budget (i.e., neither elevated N inputs nor reduced N losses). However, in comparison to the shrubland, the forest showed an almost tripling in aboveground N use efficiency (NUE; 235 vs. 83 kg dry mass kg(-1) N) and a doubling in ecosystem level C/N ratio (16 vs. 8, for the forest and shrubland, respectively). Nitrogen cycling slowed in the forest compared to the shrubland: net N mineralization rates in soils decreased by approximately 50%, decomposition rates decreased by approximately 20%, and NOx loss decreased by approximately 64%. These adjustments in N cycling provide a possible basis for increased NUE and subsequent C sequestration without net change in the overall N budget, which should be addressed in future investigations.

Motivated by persistent predictions of warming and drying in the entire Mediterranean and other regions, we have examined the interactions of intrinsic water-use efficiency (W(i)) with environmental conditions in Pinus halepensis. We used 30-year (1974-2003) tree-ring records of basal area increment (BAI) and cellulose (13)C and (18)O composition, complemented by short-term physiological measurements, from three sites across a precipitation (P) gradient (280-700 mm) in Israel. The results show a clear trend of increasing W(i) in both the earlywood (EW) and latewood (LW) that varied in magnitude depending on site and season, with the increase ranging from ca. 5 to 20% over the study period. These W(i) trends were better correlated with the increase in atmospheric CO(2) concentration, C(a), than with the local increase in temperature (similar to 0.04 degrees C year(-1)), whereas age, height and density variations had minor effects on the long-term isotope record. There were no trends in P over time, but W(i) from EW and BAI were dependent on the interannual variations in P. From reconstructed C(i) values, we demonstrate that contrasting gas-exchange responses at opposing ends of the hydrologic gradient underlie the variation in W(i) sensitivity to C(a) between sites and seasons. Under the mild water limitations typical of the main seasonal growth period, regulation was directed at increasing C(i)/C(a) towards a homeostatic set-point observed at the most mesic site, with a decrease in the W(i) response to C(i) with increasing aridity. With more extreme drought stress, as seen in the late season at the drier sites, the response was W(i) driven, and there was an increase in the W(i) sensitivity to C(a) with aridity and a decreasing sensitivity of C(i) to C(a). The apparent C(a)-driven increases in W(i) can help to identify the adjustments to drying conditions that forest ecosystems can make in the face of predicted atmospheric change.

The C-13 concentration in atmospheric CO2 has been declining over the past 150 years as large quantities of C-13-depleted CO2 from fossil fuel burning are added to the atmosphere. Deforestation and other land use changes have also contributed to the trend. Looking at the C-13 variations in the atmosphere and in annual growth rings of trees allows us to estimate CO2 uptake by land plants and the ocean, and assess the response of plants to climate. Here I show that the effects of the declining C-13 trend in atmospheric CO2 are recorded in the isotopic composition of paper used in the printing industry, which provides a well-organized archive and integrated material derived from trees' cellulose. C-13 analyses of paper from two European and two American publications showed, on average, a -1.65 +/- 1.00 parts per thousand trend between 1880 and 2000, compared with -1.45 and -1.57 parts per thousand for air and tree-ring analyses, respectively. The greater decrease in plant-derived C-13 in the paper we tested than in the air is consistent with predicted global-scale increases in plant intrinsic water-use efficiency over the 20th century. Distinct deviations from the atmospheric trend were observed in both European and American publications immediately following the World War II period.

Effective leaf area index (LAI(e)) in the semi-arid Pinus halepensis plantation, located between arid and semi-arid climatic zones at the edge of the Negev and Judean deserts, was measured bi-annually during four years (2001-2004) and more intensively (monthly) during the following two years (2004-2006) by a number of non-contact optical devices. The measurements showed a gradual increase in LAI(e) from similar to 1 (+/- 0.25) to similar to 1.8 (+/- 0.11) during these years. All instruments, when used properly, gave similar results that were also comparable with actual leaf area index measured by litter collection and destructive sampling and allometric estimates. Because of the constraint of clear sky conditions, which limited the use of the fisheye type sensors to times of twilight, the fisheye techniques were less useful. The tracing radiation and architecture of canopies system, which includes specific treatment of two levels of clumpiness of the sparse forest stand, was used successfully for the intensive monitoring. The mean clumpiness index, 0.61, is considered representative for the specific environment. Finally, the LAI(e) measurements at the start of each season were used to constrain phenology-based estimates of annual LAI(e) development, resulting in a continuous course of LAI(e) in the forest over the five-year period. Intra-seasonal LAI(e) variation in the order of 10% of total LAI(e) predicted by the model was also observed in the intensive TRAC measurements, giving confidence in the TRAC system and indicating its sensitivity and applicability in woodlands even with low LAI(e) values. The results can be important for forest management decision support as well as for use in evaluation of remote sensing techniques for forests at the lowest range of LAI(e) values. (C) 2011 Elsevier B.V. All rights reserved.

While there is currently intense effort to examine the 13C signal of CO2 evolved in the dark, less is known on the isotope composition of day-respired CO2. This lack of knowledge stems from technical difficulties to measure the pure respiratory isotopic signal: day respiration is mixed up with photorespiration, and there is no obvious way to separate photosynthetic fractionation (pure c(i)/c(a) effect) from respiratory effect (production of CO2 with a different delta 13C value from that of net-fixed CO2) at the ecosystem level. Here, we took advantage of new simple equations, and applied them to sunflower canopies grown under low and high [CO2]. We show that whole mesocosm-respired CO2 is slightly 13C depleted in the light at the mesocosm level (by 0.2-0.8 parts per thousand), while it is slightly 13C enriched in darkness (by 1.5-3.2 parts per thousand). The turnover of the respiratory carbon pool after labelling appears similar in the light and in the dark, and accordingly, a hierarchical clustering analysis shows a close correlation between the 13C abundance in day- and night-evolved CO2. We conclude that the carbon source for respiration is similar in the dark and in the light, but the metabolic pathways associated with CO2 production may change, thereby explaining the different 12C/13C respiratory fractionations in the light and in the dark.

Forests both take up CO(2) and enhance absorption of solar radiation, with contrasting effects on global temperature. Based on a 9-year study in the forests' dry timberline, we show that substantial carbon sequestration (cooling effect) is maintained in the large dry transition zone (precipitation from 200 to 600 millimeters) by shifts in peak photosynthetic activities from summer to early spring, and this is counteracted by longwave radiation (L) suppression (warming effect), doubling the forestation shortwave (S) albedo effect. Several decades of carbon accumulation are required to balance the twofold S + L effect. Desertification over the past several decades, however, contributed negative forcing at Earth's surface equivalent to similar to 20% of the global anthropogenic CO(2) effect over the same period, moderating warming trends.

Improved global estimates of terrestrial photosynthesis and respiration are critical for predicting the rate of change in atmospheric CO2. The oxygen isotopic composition of atmospheric CO2 can be used to estimate these fluxes because oxygen isotopic exchange between CO2 and water creates distinct isotopic flux signatures. The enzyme carbonic anhydrase (CA) is known to accelerate this exchange in leaves, but the possibility of CA activity in soils is commonly neglected. Here, we report widespread accelerated soil CO2 hydration. Exchange was 10-300 times faster than the un-catalyzed rate, consistent with typical population sizes for CA-containing soil microorganisms. Including accelerated soil hydration in global model simulations modifies contributions from soil and foliage to the global (COO)-O-18 budget and eliminates persistent discrepancies existing between model and atmospheric observations. This enhanced soil hydration also increases the differences between the isotopic signatures of photosynthesis and respiration, particularly in the tropics, increasing the precision of CO2 gross fluxes obtained by using the delta O-18 of atmospheric CO2 by 50%.

Warming and drying is predicted for most of the Mediterranean and other regions, and knowledge of this effect on forest carbon dynamics cannot be easily extrapolated from temperate climates. Instead, we provide quantitative information from a 6-year study in a 40-year old pine forest at the dry "timberline" (280 mm annual rainfall) on soil CO2 efflux (F-s) and some of its controlling factors. Annual Fs was relatively low (405.9 +/- 23.8 g C m(-2) a(-1)), but within one standard deviation of a global nonlinear relationship we derived between mean annual precipitation and F-s in forests. Seasonal variations in F-s were dominated by soil temperature (with Q(10) = 2.45) in the wet season, and by soil moisture in the water-limited seasons, but not by pulse responses to precipitation. No temperature sensitivity was observed in the dry season, but there was clear evidence for down regulation of sensitivity to Q(10) = 1.18 when soil moisture was kept high by a supplement summer irrigation treatment. Interannual variations in F-s correlated linearly with cumulative rainwater availability, indicating the combined importance of both precipitation amount and its temporal distribution between the wet (and cool) season and the transitional periods characterized by high evaporative demand. Low rates of soil carbon loss combined with high belowground carbon allocation (41% of canopy CO2 uptake) might explain the high soil organic carbon accumulation and net ecosystem productivity in this dry forest. Our results indicate that F-s in pine forests may adjust to dry conditions with better carbon economy than estimated from response to episodic drought in more temperate climate.

Nitric oxide (NO) plays a central role in the formation of tropospheric ozone, hydroxyl radicals, as well as nitrous and nitric acids. There are, however, large uncertainties around estimates of global NO emissions due to the paucity of data. In particular, there is little information on the rate of NO emission and its sensitivity to processes such as land use changes in dry environments. Here we report on a two-year study on the influence of afforestation on soil NO fluxes in the semi-arid afforestation system in Southern Israel (Yatir forest, mean annual precipitation similar to 280 mm). Laboratory incubations were carried out under seasonally defined conditions of soil moisture and temperature using soils sampled in different seasons from the native shrubland (taken both under shrub canopy and in the inter-shrub areas), and from the adjacent similar to 2800 ha, 40-year-old pine afforestation site. Combining laboratory results with field measurements of soil moisture and temperature, we up-scaled soil-atmosphere NO fluxes to the ecosystem level. The different microsites differed in their annual mean NO release rates (0.04, 0.14 and 0.03 mg m(-2) d(-1) for the shrubland under and between shrubs and for the forest, respectively), and exhibited high inter-seasonal variability in NO emission rates (ranging from zero up to 0.25 mg m(-2) d(-1) in the wet and dry-rewetting seasons, respectively), as well as in temperature responses. Up-scaling results to annual and ecosystem scales indicated that afforestation of the semi-arid shrubland could reduce soil NO emission by up to 65%. (C) 2009 Elsevier Ltd. All rights reserved.

The launch of the Moderate Resolution Imaging Spectroradiometer (MODIS) onboard the Terra and Aqua satellites improved the ability to evaluate several surface biophysical parameters, including Leaf Area Index (LAI), which is provided as an operational MODIS product, available at 1-km spatial resolution and at 8-day intervals. However, for heterogeneous and sparse planted forests that are common to the semi-arid eastern Mediterranean region, the data at low spatial resolution may be significantly biased by the contribution of different background elements to the total surface reflectance received by the sensor and cannot therefore correctly reflect the real forest phenology. In the current paper the performance of the MODIS LAI product was examined over a dryland Mediterranean forest in southern Israel. The study found a significant discrepancy between ground-based and MODIS LAI datasets. In general, MODIS LAI values were c. 51% of the ground-based LAI measurements. In addition ground based LAI peaked in the summer due to the natural growth cycle of the pine trees, while MODIS values peaked in the winter. The MODIS seasonal course could be explained by the development of annuals and crypto- and micro-phytes in the understorey and the clearing areas during the mid winter months that are included in the MODIS LAI product but not in the ground based measurements. However, for that period MODIS estimates should have exceeded ground-based estimates while in fact they were still lower. The relationship between MOD12C1 Land Cover Type 3 and MOD15A2 products is discussed.

Isotopic measurements of leaf water have provided insights into a range of ecophysiological and biogeochemical processes, but require an extraction step which often constitutes the major analytical bottleneck in large-scale studies. Current standard procedures for leaf water analysis are based on cryogenic vacuum or azeotrophic distillation, and are laborious, require sophisticated distillation lines and the use of toxic materials. We report a rapid technique based on centrifugation/filtration of leaf samples pulverised in their original sampling tubes, using a specifically adapted, simple apparatus. The leaf water extracts produced are suitable for isotopic analysis via pyrolysis gas chromatography isotope ratio mass spectrometry (PYR/GC/IRMS). The new method was validated against cryogenic vacuum distillation and showed an overall accuracy of +/- 0.5 parts per thousand (nine grouped comparisons, n = 110) over a range of 21 parts per thousand. Effects due to the presence of soluble carbohydrates were near the detection limits for most samples analysed, and these effects could be corrected for (the extracted soluble organics could also be used for isotopic analysis). The extraction time for a routine eight-sample subset was reduced from 4 h (cryogenic distillation) to 45 min, limited only by the size of the centrifuge(s) used. This method provides a rapid, low-cost and reliable alternative to conventional vacuum and other distillation methods that can alleviate current restrictions on ecosystem- and global-scale studies that require high-throughput leaf water isotopic analysis. Copyright 2008 John Wiley & Sons, Ltd.

Although the isotopic composition of precipitation is widely used in global climate change studies, use of water vapour isotopes is considerably more limited. Here we present the results from 9 yr of atmospheric vapour measurements in the Eastern Mediterranean, at a site in Israel. The measurements show a strong mean seasonal cycle of about 4 parts per thousand in 180 (peaking around July). This seasonality could not be adequately explained by changes in surface interactions or in air mass trajectories, as usually invoked for variations in local precipitation. We could explain this cycle only as a combination of three components: (1) rainout effects; (2) temperature and relative humidity control of the initial vapour and (3) seasonal variations in the vertical mixing across the top of the planetary boundary layer. This last component is emphasized in the Current Study, and it was shown to be a significant factor in the seasonal cycle features. The measurements were also compared with an isotope-enabled GCM (CAM2) run, which exhibited a markedly different seasonal cycle. Such comparisons with vapour isotopes data could help in constraining models better.

Predictions of warming and drying in the Mediterranean and other regions require quantifying of such effects on ecosystem carbon dynamics and respiration. Long-term effects can only be obtained from forests in which seasonal drought is a regular feature. We carried out measurements in a semiarid Pinus halepensis (Aleppo pine) forest of aboveground respiration rates of foliage, R(f), and stem, R(t) over 3 years. Component respiration combined with ongoing biometric, net CO(2) flux [net ecosystem productivity (NEP)] and soil respiration measurements were scaled to the ecosystem level to estimate gross and net primary productivity (GPP, NPP) and carbon-use efficiency (CUE=NPP/GPP) using 6 years data. GPP, NPP and NEP were, on average, 880, 350 and 211 g C m(-2) yr(-1), respectively. The above ground respiration made up half of total ecosystem respiration but CUE remained high at 0.4. Large seasonal variations in both R(f) and R(t) were not consistently correlated with seasonal temperature trends. Seasonal adjustments of respiration were observed in both the normalized rate (R(20)) and short-term temperature sensitivity (Q(10)), resulting in low respiration rates during the hot, dry period. R(f) in fully developed needles was highest over winter-spring, and foliage R(20) was correlated with photosynthesis over the year. Needle growth occurred over summer, with respiration rates in developing needles higher than the fully developed foliage at most times. R(t) showed a distinct seasonal maximum in May irrespective of year, which was not correlated to the winter stem growth, but could be associated with phenological drivers such as carbohydrate re-mobilization and cambial activity. We show that in a semiarid pine forest photosynthesis and stem growth peak in (wet) winter and leaf growth in (dry) summer, and associated adjustments of component respiration, dominated by those in R(20), minimize annual respiratory losses. This is likely a key for maintaining high CUE and e

An approach is presented for determining leaf area index (LAI) of a forest located at the desert fringe by using high spatial resolution imagery and by implementing values from a moderate spatial but high temporal resolution sensor. A 4-m spatial resolution multi-spectral IKONOS image was acquired under clear sky conditions on March 25, 2004. Normalized differences vegetation index (NDVI) and a linear mixture model were applied to calculate fractional vegetation cover (FVC). LAI was calculated using a non-linear relationship to FVC and then compared with ground truth measurements made in ten 1000 m(2) plots Using the tracing radiation and architecture of canopies (TRAC) canopy analyzer under bright and clear sky conditions during March and April, 2004. Calculated LAI, corrected with a measured clumping index, was highly correlated with measured LAI (R-2=0.79, p

Ecosystems in dry regions are generally low in productivity and carbon (C) storage. We report, however, large increases in C sequestration following afforestation of a semi-arid shrubland with Pinus halepensis trees. Using C and nitrogen (N) inventories, based in part on site-specific allometric equations, we measured an increase in the standing ecosystem C stock from 2380 g C m(-2) in the shrubland to 5840 g C m(-2) in the forest after 35 years, with no significant change in N stocks. Carbon sequestration following afforestation was associated with increased N use efficiency as reflected by an overall increase in C/N ratio from 7.6 in the shrubland to 16.6 in the forest. The C accumulation rate in the forest was particularly high for soil organic C (SOC; increase of 1760 g C m(-2) or 50 g C m(-2) yr(-1)), which was associated with the following factors: 1) Analysis of a small (13)C signal within this pure C(3) system combined with size fractionation of soil organic matter indicated a significant addition of new SOC derived from forest vegetation (68% of total forest SOC) and a considerable portion of the old original shrubland SOC (53%) still remaining in the forest. 2) A large part of both new and old SOC appeared to be protected from decomposition as about 60% of SOC under both land-use types were in mineral-associated fractions. 3) A short-term decomposition study indicated decreased decomposition of lower-quality litter and SOC in the forest, based on reduced decay rates of up to 90% for forest compared to shrubland litter. 4) Forest soil included a significant component of live and dead roots (12% of total SOC). Our results suggest a role for increased N use efficiency, enhanced SOC protection and reduced decomposition rates in the large C sequestration potential following afforestation in semi-arid regions. These results are particularly relevant in light of persistent predictions of drying trends in the Mediterranean and other regions.

The isotopic composition of atmospheric O-2 depends on the rates of oxygen cycling in photosynthesis, respiration, photochemical reactions in the stratosphere and on delta O-17 and delta O-18 of ocean and leaf water. While most of the factors affecting delta O-17 and delta O-18 of air O-2 have been studied extensively in recent years, delta O-17 of leaf water-the substrate for all terrestrial photosynthesis-remained unknown. In order to understand the isotopic composition of atmospheric O-2 at present and in fossil air in ice cores, we studied leaf water in field experiments in Israel and in a European survey. We measured the difference in delta O-17 and delta O-18 between stem and leaf water, which is the result of isotope enrichment during transpiration. We calculated the slopes of the lines linking the isotopic compositions of stem and leaf water. The obtained slopes in ln(delta O-17 + 1) VS. ln(delta O-18 + 1) plots are characterized by very high precision (similar to 0.001) despite of relatively large differences between duplicates in both delta O-17 and delta O-18 (0.02-0.05 parts per thousand). This is so because the errors in 6180 and 617 0 are mass-dependent. The slope of the leaf transpiration process varied between 0.5111 +/- 0.0013 and 0.5204 +/- 0.0005, which is considerably smaller than the slope linking liquid water and vapor at equilibrium (0.529). We further found that the slope of the transpiration process decreases with atmospheric relative humidity (h) as 0.522-0.008 x h, for h in the range 0.3-1. This slope is neither influenced by the plant species, nor by the environmental conditions where plants grow nor does it show strong variations along long leaves. (c) 2006 Elsevier Inc. All rights reserved.

This paper reports oil ranges of carbon dioxide (CO,) activity in biological soil crusts (BSC) correlated with different ranges of the BSC's spectral reflectance throughout the Phenological cycle of the year. Methodology is based oil surface CO2 exchange measurements, ground spectral measurements, and satellite images interpretation. Thirty-nine field campaigns, each of duration of 3 days, were conducted over the Course of 2 years at a sand duties and a loess environment of the northwestern Negev desert in Israel, in order to relate the CO2 fluxes and the spectral signals to the seasonal phenology. The Normalized Difference Vegetation Index (NDVI) was derived from ground measurements of the BSC's reflectance and correlated with their CO2 exchange data. A linear mixture model, incorporating the different contributions of the sites' ground features, was calculated and compared with SPOT-HRV data. From the ground measurements, fairly good correlations were found between the NDVI and the CO, fluxes oil a seasonal scale. Hence, the NDVI successfully indicates the potential magnitude and capacity of the BSC's assimilation activity. The linear mixture model Successfully describes the phenological cycles of the BSC, annual, and perennial plants and corresponds well to the satellite data. Moreover, the model enables annual changes of the phenology cycle and the growing season length to be distinguished. Both the linear mixture model and the derived NDVI values recorded the recovery of the BSC at the beginning of the wet season before annuals had germinated. Finally, it is concluded that a combination of CO, exchange measurements, linear mixture model, and NDVI values is suitable for monitoring BSC's productivity in and regions. (c) 2005 Elsevier Inc. All rights reserved.

Variation in the (COO)-O-18 content of atmospheric CO2 (delta(18)O(a)) can be used to distinguish photosynthesis from soil respiration, which is based on carbonic anhydrase (CA)-catalyzed O-18 exchange between CO2 and O-18-enriched leaf water (delta(18)O(w)). Here we tested the hypothesis that mean leaf delta(18)O(w) and assimilation rates can used to estimate whole-leaf (COO)-O-18 flux (isoflux), ignoring intraleaf variations in CA activity and gas exchange parameters. We observed variations in CA activity along the leaf (>30% decline from the leaf center toward the leaf ends), which were only partially correlated to those in delta(18)O(w) (7 to 21 parts per thousand), delta(18)O and delta(13)C of leaf organic matter (25 to 30 parts per thousand and -12.8 to -13.2 parts per thousand, respectively), and substomatal CO2 concentrations (intercellular CO2 concentrations, c(i), at the leaf center were similar to 40% of those at the leaf tip). The combined effect of these variations produced a leaf-integrated isoflux that was different from that predicted based on bulk leaf values. However, because of canceling effects among the influencing parameters, isoflux overestimations were only similar to 10%. Conversely, use of measured parameters from a leaf segment could produce large errors in predicting leaf-integrated (COO)-O-18 fluxes.

We present carbon stable isotope, delta C-13, results from air and organic matter samples collected during 98 individual field campaigns across a network of Carboeuroflux forest sites in 2001 (14 sites) and 2002 (16 sites). Using these data, we tested the hypothesis that delta C-13 values derived from large-scale atmospheric measurements and models, which are routinely used to partition carbon fluxes between land and ocean, and potentially between respiration and photosynthesis on land, are consistent with directly measured ecosystem-scale delta C-13 values. In this framework, we also tested the potential of delta C-13 in canopy air and plant organic matter to record regional-scale ecophysiological patterns. Our network estimates for the mean delta C-13 of ecosystem respired CO2 and the related 'discrimination' of ecosystem respiration, delta(er) and Delta(er), respectively, were -25.6 +/- 1.9 parts per thousand and 17.8 +/- 2.0 parts per thousand in 2001 and -26.6 +/- 1.5 parts per thousand and 19.0 +/- 1.6 parts per thousand in 2002. The results were in close agreement with delta C-13 values derived from regional-scale atmospheric measurement programs for 2001, but less so in 2002, which had an unusual precipitation pattern. This suggests that regional-scale atmospheric sampling programs generally capture ecosystem delta C-13 signals over Europe, but may be limited in capturing some of the interannual variations. In 2001, but less so in 2002, there were discernable longitudinal and seasonal trends in delta(er). From west to east, across the network, there was a general enrichment in C-13 (similar to 3 parts per thousand and similar to 1 parts per thousand for the 2 years, respectively) consistent with increasing Gorczynski continentality index for warmer and drier conditions. In 2001 only, seasonal C-13 enrichment between July and September, followed by depletion in November (from about -26.0 parts per thousand to -24.5 parts per thousand to -30.0 parts per tho

Wild emmer wheat (Triticum turgidum spp. dicoccoides (Korn.) Thell.), the tetraploid progenitor of cultivated wheat, is a potential source for various agronomical traits, including drought resistance. The objectives of this study were to characterize (1) the genetic diversity for drought resistance in wild emmer wheat, and (2) the relationship between drought responses of the wild emmer germplasm and the ecogeographical parameters of its collection sites. A total of 110 wild emmer accessions consisting of 25 populations and three control durum wheat cultivars were examined under two irrigation regimes, well-watered ('wet') and water-limited ('dry'). Wide genetic diversity was found both between and within the wild emmer populations in most variables under each treatment. A considerable number of the wild emmer accessions exhibited an advantage in productivity (spike and total dry matter) over their cultivated counterparts. Most wild emmer wheat accessions exhibited a greater carbon isotope ratio (delta(13)C, indicating higher water-use efficiency) under the dry treatment and higher plasticity of delta(13)C relative to the cultivated controls, which may have contributed to the drought adaptations in the former. The most outstanding drought-tolerance capacity (in term of productivity under the dry treatment and susceptibility indices) was detected in wild emmer populations originated from hot dry locations. The results suggest that wild emmer has the potential to improve drought resistance in cultivated wheat.

A widespread conversion of sagebrush-dominated plant communities to post-fire communities dominated by herbaceous annual species has occurred in the Great Basin of North America's Intermountain West during the last century, mainly driven by invasions of alien species and a subsequent increase in fire frequency. Little experimental data, however, are available on the potential consequences of wildfire and post-fire plant community alteration on key ecosystem functions such as ecosystem hydrology. The objectives of our study were to (1) quantify the effects of this vegetation transformation on the spatial and temporal distribution of soil water in the rooting zone; (2) quantify the effects of wild-fire on surface soil water distribution (upper 20 cm) to infer about potential consequences on re-establishment of the native plant community; and (3) examine which processes (e.g., soil water recharge or replenishment, plant water uptake, and soil evaporation) may be responsible for observed changes soil water distribution. Continuous measurement of soil water content using a segmented time domain reflectrometry system in sixteen locations in an intact sagebrush ecosystem (eight shrub and eight inter-shrub locations) and in eight locations in an adjacent post-fire ecosystem from March 2001 to October 2002 indicated that soil water contents in the upper 75 cm of the soil were similar and very low during summers, but that substantially lower water recharge in post-fire ecosystems after large snowfalls - possibly due to decreased snow deposition and higher sublimation - caused large differences in water storage between the two ecosystems in winter. These differences declined with the onset of the vegetation period in March, probably due to higher plant water uptake in the more densely vegetated intact sagebrush ecosystem. The presence of patchiness of near-surface soil water (0-20 cm), quantified among 80 points of nested grid systems placed in each ecosystem, appears to be

Scaling is a naturally iterative and bi-directional component of problem solving in ecology and in climate science. Ecosystems and climate systems are unquestionably the sum of all their parts, to the smallest imaginable scale, in genomic processes or in the laws of fluid dynamics. However, in the process of scaling-up, for practical purposes thewhole usually has to be construed as a good deal less than this. This essay demonstrates how controlled large-scale experiments can be used to deduce key mechanisms and thereby reduce much of the detail needed for the process of scaling-up. Collection of the relevant experimental evidence depends on controlling the environment and complexity of experiments, and on applications of technologies that report on, and integrate, small-scale processes. As the role of biological feedbacks in the behavior of climate systems is better appreciated, so the need grows for experimentally based understanding of ecosystem processes. We argue that we cannot continue as we are doing, simply observing the progress of the greenhouse gas-driven experiment in global change, and modeling its future outcomes. We have to change the way we think about climate system and ecosystem science, and in the process move to experimental modes at larger scales than previously thought achievable.

Rising atmospheric CO2 concentrations may lead to increased water availability because the water use efficiency of photosynthesis (WUE) increases with CO2 in most plant species. This should allow the extension of afforestation activities into drier regions. Using eddy flux, physiological and inventory measurements we provide the first quantitative information on such potential from a 35-year old afforestation system of Aleppo pine (Pinus halepensis Mill.) at the edge of the Negev desert. This 2800 ha arid-land forest contains 6.5 +/- 1.2 kg C m(-2) , and continues to accumulate 0.13-0.24 kg C m(-2) yr(-1) . The CO2 uptake is highest during the winter, out of phase with most northern hemispheric forest activity. This seasonal offset offers low latitude forests similar to10 ppm higher CO2 concentrations than that available to higher latitude forests during the productive season, in addition to the 30% increase in mean atmospheric CO2 concentrations since the 1850s. Expanding afforestation efforts into drier regions may be significant for C sequestration and associated benefits (restoration of degraded land, reducing runoff, erosion and soil compaction, improving wildlife) because of the large spatial scale of the regions potentially involved (ca. 2 x 10(9) ha of global shrub-land and C4 grassland). Quantitative information on forest activities under dry conditions may also become relevant to regions predicted to undergo increasing aridity.

Isoprene emission from leaves is dynamically coupled to photosynthesis through the use of primary and recent photosynthate in the chloroplast. However, natural abundance carbon isotope composition (delta(13)C) measurements in myrtle (Myrtus communis), buckthorn (Rhamnus alaternus), and velvet bean (Mucuna pruriens) showed that only 72% to 91% of the variations in the delta(13)C values of fixed carbon were reflected in the delta(13)C values of concurrently emitted isoprene. The results indicated that 9% to 28% carbon was contributed from alternative, slow turnover, carbon source(s). This contribution increased when photosynthesis was inhibited by CO2-free air. The observed variations in the delta(13)C of isoprene under ambient and CO2-free air were consistent with contributions to isoprene synthesis in the chloroplast from pyruvate associated with cytosolic Glc metabolism. Irrespective of alternative carbon source(s), isoprene was depleted in C-13 relative to mean photosynthetically fixed carbon by 4%. to 11%.. Variable C-13 discrimination, its increase by partially inhibiting isoprene synthesis with fosmidomicin, and the associated accumulation of pyruvate suggested that the main isotopic discrimination step was the deoxyxylulose-5-phosphate synthase reaction.

[1] Photosynthesis and respiration impart distinct isotopic signatures to the atmosphere that are used to constrain global carbon source/sink estimates and partition ecosystem fluxes. Increasingly, the "Keeling plot'' method is being used to determine the carbon isotope composition of ecosystem respiration (delta(13)C(R)) in order to better understand the processes controlling ecosystem isotope discrimination. In this paper we synthesize emergent patterns in delta(13)C(R) by analyzing 146 Keeling plots constructed at 33 sites across North and South America. In order to interpret results from disparate studies, we discuss the assumptions underlying the Keeling plot method and recommend standardized methods for determining delta(13)C(R). These include the use of regression calculations that account for error in the x variable, and constraining estimates of delta(13)C(R) to nighttime periods. We then recalculate delta(13)C(R) uniformly for all sites. We found a high degree of temporal and spatial variability in C-3 ecosystems, with individual observations ranging from -19.0 to -32.6parts per thousand. Mean C-3 ecosystem discrimination was 18.3parts per thousand. Precipitation was a major driver of both temporal and spatial variability of delta(13)C(R), suggesting (1) a large influence of recently fixed carbon on ecosystem respiration and (2) a significant effect of previous climatic effects on delta(13)CR. These results illustrate the importance of water availability as a key control on atmospheric (CO2)-C-13 and highlight the potential of delta(13)C(R) as a useful tool for integrating environmental effects on dynamic canopy and ecosystem processes.

Isoprene (2-methyl-1,3-butadiene) protection against effects of singlet oxygen was investigated in Myrtus communis and Rhamnus alaternus. In M. communis, singlet oxygen produced in the leaves by Rose Bengal (RB) led to a 65% decrease in net assimilation rates within 3 h, whereas isoprene emission rates showed either a 30% decrease at ambient CO(2) concentrations or a 70% increase under high CO(2). In both cases, these changes led to an increase in calculated internal isoprene concentrations. The isoprene protection effect was directly demonstrated by fumigation of young (non-emitting) leaves, treated with RB or bromoxynil (simulating photoinhibition). There was 42% and 29% reduction in the damage to net assimilation compared with non-fumigated leaves for RB or bromoxynil, respectively. In R. alaternus, similar effects of RB on net assimilation were observed, and additional fluorescence measurements showed a significantly smaller decrease in F(v)/F(m) in isoprene-fumigated young leaves treated with RB (from 0.78 to 0.52), compared with non-fumigated leaves (from 0.77 to 0.27). The internal isoprene concentrations used in this study and possible rate of (1)O(2) production in leaves indicate that the protective effects observed should be beneficial also under natural conditions.

The interaction of genotype with environment is of primary importance in many aspects of genomic research and is a special priority in the study of major crops grown in a wide range of environments. Water deficit, the major factor limiting plant growth and crop productivity worldwide, is expected to increase with the spread of arid lands. In genetically equivalent cotton populations grown under well-watered arid water-limited conditions (the latter is responsible for yield reduction of similar to 50% relative to well-watered conditions), productivity arid quality were shown to be partly accounted for by different quantitative trait loci (QTLs), indicating that adaptation to both arid and favorable conditions can be combined in the same genotype. QTL mapping was also used to test the association between productivity and quality under water deficit with a Suite of traits often found to differ between genotypes adapted to arid versus well-watered conditions. In this study, only reduced plant osmotic potential was clearly implicated in improved cotton productivity under arid conditions. Genomic tools and approaches may expedite breeding of genotypes that respond favorably to specific environments, help test roles of additional physiological factors, and guide the isolation of genes that protect crop performance under arid conditions toward improved adaptation of crops to arid cultivation.

The use of stable isotopes to estimate evapotranspiration (ET) fluxes from vegetated areas is increasing. By complementing conventional net flux measurements (gradient or eddy correlation techniques), isotope analyses can allow partitioning ET between its gross components, soil evaporation and leaf transpiration. Isotopic analyses of atmospheric water vapour above canopies can also constrain, or provide alternatives for estimating ET. A brief discussion of the isotope approach is aimed at highlighting some of the uncertainties that require further research. We also demonstrate first, the application of combined concentration and isotopic gradient analysis of atmospheric water vapour above crop fields in order to estimate ET fluxes and its gross components (soil evaporation was estimated at 1.5-3.5% of mid-day ET flux in a mature wheat field). Second, we demonstrate the potential in monitoring delta(ss)- delta(L), the difference between predicted and measured leaf water delta(18)O values, as an indicator of seasonal variations in canopy-scale transpiration in a desert ecosystem (linear correlation between this indicator and conventional ET measurements was observed). Improving our analytical capabilities for high-precision isotopic analysis of very small water vapour samples was a limiting factor in the above applications and a method for pyrolysis and on-line O-18 analysis of 0.2-2 mu L water samples is described. Copyright (C) 2000 John Wiley & Sons, Ltd.

The effects of sugar concentration in the medium on the autotrophic/heterotrophic carbon metabolism were studied in tissue cultured potato plants. The weight of plants grown in the light on media containing 3 % sucrose was significantly higher than that of plants grown on 8 % sucrose media. Similarly, the weight increased when plants were transferred from 8 % to 3 % sucrose and decreased in the reverse case. In contrast, under dark conditions, a higher weight was observed in plants grown on higher sucrose concentrat ion. The uptake of C-14-sucrose from the medium was higher when the sugar's concentration was 8% as compared to 3 %, indicating a higher proportion of heterotrophic carbon in the former. These results were further supported by the natural abundance of C-13/C-12 ratios in the plant tissue. Analyses of the C-13/C-12 in the plants, sugars and the air CO2, indicated that 90 % of the tissue carbon was of heterotrophic origin in light-grown plants on 8 % sucrose. Only 50 % of the tissue carbon was of heterotrophic origin when the plants were grown on 2 % glucose medium. These results provide evidence that a high sugar concentration in the medium significantly reduced the contribution of autotrophic CO2 assimilation to the accumulation of dry weight. The possible mechanisms inhibiting CO2 assimilation in potato plants grown in vitro, under conditions of high sugar concentrations (or low osmotic potential) are discussed.

Cotton (Gossypium spp.) is often exposed to drought, which adversely affects both yield and quality. Improved water-use efficiency (WUE = total dry matter produced or yield harvested / water used) is expected to reduce these adverse effects. Genetic variability in WUE and its association with photosynthetic rate and carbon isotope ratio (C-13/C-12) in cotton are reported in this paper. WUE of six cotton cultivars-G. hirsutum L., G. barnbadense L., and an interspecific F-1 hybrid (G. hirsutum X G. barbadense, ISH), was examined under two irrigation regimes in two field trials. The greatest WUE was obtained by two G. hirsutum cultivars (2.55 g dry matter or 1.12 g seed-cotton L-1 H2O); the ISH obtained similar or somewhat lower values, and two G. barbadense cultivars and one G. hirsutum cultivar exhibited the lowest values (2.1 g dry matter or 0.8 to 0.85 g seed-cotton L-1 H2O). These results indicate that different cotton cultivars may have evolved different environmental adaptations that affect their WUE. Photosynthetic rate was correlated with WUE in only a few cases, emphasizing the limitation of this parameter as a basis for estimating crop WUE. Under both trials, WUE was positively correlated with carbon isotope ratio, indicating the potential of this technique as a selection criterion for improving cotton WUE.

The C-13/C-12 and O-18/O-16 ratios of stem cellulose of Tamarix jordanis (a tree common in wadis of arid regions) increased with decreasing relative humidity (RH) in individual trees growing along a climatic gradient in Israel. The response to RH observed in the delta(18)O of the wood cellulose was strongly similar to that observed in leaf water over a diurnal cycle. Most of the data for delta(13)C and all of the data for delta(18)O could be fitted to two independent linear equations that, combined, allowed the reconstruction of RH and the delta(18)O of source water from the isotopic composition of ancient T. jordanis wood previously reported from the ancient fortress of Masada. Since the Roman period, RH at Masada decreased by about 17%, while the delta(18)O value of local groundwater remained similar to present-day values, suggesting that changing atmospheric circulation has played a role in climate change in the Middle East over the past two millennia.

A strong correlation is observed between an El Nino index (anomalies in tropical Pacific sea surface temperature) and rainfall in the Judean foothills near Jerusalem over the past 20 years. These relationships clearly influenced the growth of local pine trees, as reflected in the width of their annual tree rings. The ability to predict El Nino events about a year in advance lend a special significance to relationships reported here for ecology, agriculture and water management in this climatic transition zone. To help explain the observed, long-range teleconnection we propose a possible mechanism based on a newly identified direct cloud connection between equatorial Africa (more directly affected by El Nino) and the Southeastern Mediterranean shoreland. The penetration and contribution of the moisture current from equatorial Africa to this region may depend on a shift in the usual rain generating moisture currents to southwesterly trajectories (passing over north Africa). The occurrence of such shifts is supported by the observed decrease in the mean O-18 content of the local precipitation during El Nino winters.

The contribution of the cyanide-resistant, alternative pathway to plant mitochondrial electron transport has been studied using a modified aqueous phase on-line mass spectrometry-gas chromatography system. This technique permits direct measurement of the partitioning of electrons between the cytochrome and alternative pathways in the absence of added inhibitors. We demonstrate that in mitochondria isolated from soybean (Glycine max L. cv Ransom) cotyledons, the alternative pathway contributes significantly to oxygen uptake under state 4 conditions, when succinate is used as a substrate. However, when NADH is the substrate, addition of pyruvate, an allosteric activator of the alternative pathway, is required to achieve the same level of alternative pathway activity. Under state 3 conditions, when the reduction state of the ubiquinone pool is low, the addition of pyruvate allows the alternative pathway to compete with the cytochrome pathway for electrons from the ubiquinone pool when the cytochrome pathway is not saturated. These results provide direct experimental verification of the kinetics consequences of pyruvate addition on the partitioning of electron flow between the two respiratory pathways. This distribution of electrons between the two unsaturated pathways could not be measured using;conventional oxygen electrode methods and illustrates a dear advantage of the mass spectrometry technique. These results have significant ramifications for studies of plant respiration using the oxygen electrode, particularly those studies involving intact tissues.

THE level of the Dead Sea, the lowest (about -400 m) and one of the most salty (salinity about 340 g l(-1))lakes on earth, is lowering at a rate of approximately 0.5 m annually owing to extensive exploitation of its main perennial tributary (the Jordan River) and the extreme aridity of the region (annual precipitation is about 60 mm)(1). Consequently, new hypersaline sea shores are exposed, forming a unique, originally sterile ecosystem. The first plants invade these newly exposed shores after several years while soil water salinity is still extremely high, Here we use stable isotopes of oxygen and hydrogen to show that a variety of such perennial pioneer plants are able to make use of occasional floodwater which is distinct from the bulk of the hypersaline soil water found in their root zone. Our results provide new insight into the ways in which plants can invade extremely hostile environments and extend their ecological limits of distribution.

The isotopic ratios C-13/C-12 and O-18/O-16 of cellulose from tamarix trees which were used by the Roman army as a groundwork of the siege-rampart of Masada (AD 70-73) were compared with ratios measured in present-day tamarix trees growing in the Masada region and in central Israel. The ancient tamarix cellulose is depleted in both C-13 and O-18 compared to cellulose from trees growing in the Masada region today. Similar trends were observed on comparing modern tamarix trees growing in the Negev Desert with those growing in the temperate climate of central Israel. Considering the factors that can contribute to the observed changes in isotopic composition, we conclude that the ancient trees enjoyed less arid environmental conditions during their growth compared to contemporary trees in this desert region. This report demonstrates the potential in using combined O-18 and C-13 analyses of archeological plant material as independent indication of regional climatic change in desert areas (where conventional isotopic analyses, such as in tree rings, are impractical).