Publications

Drought stress causes multiple feedback responses in plants. These responses span from stomata closure and enzymatic downregulation of photosynthetic activity to structural adjustments of xylem biomass and leaf area. Some of these processes are not easily reversible and may persist long after the stress has ended. Despite a multitude of hydraulic model approaches, simulation models still widely lack an integrative mechanistic description of how this sequence of physiological to structural tree responses may be realized that is also simple enough to be generally applicable. Here, we suggest an integrative, sequential approach to simulate drought stress responses. First, decreasing plant water potential triggers stomatal closure alongside a downregulation of photosynthetic performance, thereby effectively slowing down further desiccation. A second protective mechanism is introduced by increasing the soil-root resistance, represented by a disconnection of fine roots after a threshold soil water potential has been reached. Further decreases in plant water potential due to residual transpiration and loss of internal stem water storage consistently lead to a loss of hydraulic functioning, which is reflected in sapwood loss and foliage senescence. This new model functionality has been used to investigate the responses of tree hydraulics, carbon uptake, and transpiration to soil and atmospheric drought in an extremely dry Aleppo pine (Pinus halepensis Mill.) plantation. Using the hypothesis of a sequential triggering of stress-mitigating responses, the model was able to reflect carbon uptake and transpiration patterns under varying soil water supply and atmospheric demand conditions - especially during summer - and respond realistically regarding medium-term responses, such as leaf and sapwood senescence. We could show that the observed avoidance strategy was only achieved when the model accounted for very early photosynthesis downregulation, and the relatively high measured plant water potentials were well reproduced with a root-soil disconnection strategy that started before major xylem conductance losses occurred. Residual canopy conductance was found to be pivotal in explaining dehydration and transpiration patterns during summer, but it also disclosed the fact that explaining the water balance in the driest periods requires water supply from stem water and deep soil layers. In agreement with the high drought resistance observed at the site, our model indicated little loss of hydraulic functioning in Aleppo pine, despite the intensive seasonal summer drought.

Climate change is often associated with increasing vapour pressure deficit (VPD) and changes in soil moisture (SM). While atmospheric and soil drying often co-occur, their differential effects on plant functioning and productivity remain uncertain. We investigated the divergent effects and underlying mechanisms of soil and atmospheric drought based on continuous, in situ measurements of branch gas exchange with automated chambers in a mature semiarid Aleppo pine forest. We investigated the response of control trees exposed to combined soil-atmospheric drought (low SM, high VPD) during the rainless Mediterranean summer and that of trees experimentally unconstrained by soil dryness (high SM; using supplementary dry season water supply) but subjected to atmospheric drought (high VPD). During the seasonal dry period, branch conductance (gbr), transpiration rate (E) and net photosynthesis (Anet) decreased in low-SM trees but greatly increased in high-SM trees. The response of E and gbr to the massive rise in VPD (to 7 kPa) was negative in low-SM trees and positive in high-SM trees. These observations were consistent with predictions based on a simple plant hydraulic model showing the importance of plant water potential in the gbr and E response to VPD. These results demonstrate that avoiding drought on the supply side (SM) and relying on plant hydraulic regulation constrains the effects of atmospheric drought (VPD) as a stressor on canopy gas exchange in mature pine trees under field conditions.Under high evaporative demand conditions, mature Aleppo pine trees demonstrated increased transpiration and photosynthesis in response to supplementary irrigation. Stomatal conductance exhibited hypo-sensitivity to increasing water demand (high vapour pressure deficit) but was sensitive to a decrease in water supply, expressed as soil water content or plant water potential. Our findings highlight that water supply, rather than water demand, is the primary limiting factor for transpiration in Aleppo pine trees.

Suppression of carbon emissions through photovoltaic (PV) energy and carbon sequestration through afforestation provides complementary climate change mitigation (CCM) strategies. However, a quantification of the "break-even time"(BET) required to offset the warming impacts of the reduced surface reflectivity of incoming solar radiation (albedo effect) is needed, though seldom accounted for in CCM strategies. Here, we quantify the CCM potential of PV fields and afforestation, considering atmospheric carbon reductions, solar panel life cycle analysis (LCA), surface energy balance, and land area required across different climatic zones, with a focus on drylands, which offer the main remaining land area reserves for forestation aiming climate change mitigation (Rohatyn S, Yakir D, Rotenberg E, Carmel Y. Limited climate change mitigation potential through forestation of the vast dryland regions. 2022. Science 377:1436-1439). Results indicate a BET of PV fields of -2.5 years but >50× longer for dryland afforestation, even though the latter is more efficient at surface heat dissipation and local surface cooling. Furthermore, PV is -100× more efficient in atmospheric carbon mitigation. While the relative efficiency of afforestation compared with PV fields significantly increases in more mesic climates, PV field BET is still -20× faster than in afforestation, and land area required greatly exceeds availability for tree planting in a sufficient scale. Although this analysis focusing purely on the climatic radiative forcing perspective quantified an unambiguous advantage for the PV strategy over afforestation, both approaches must be combined and complementary, depending on climate zone, since forests provide crucial ecosystem, climate regulation, and even social services.

Drought stress is imposing multiple feedback responses in plants. These responses span from stomata closure and enzymatic downregulation of photosynthetic activity to structural adjustments in leaf area. Some of these processes are not easily reversible and may persist long after the stress ended. Unfortunately, simulation models widely lack an integrative mechanistic description on how this sequence of tree physiological to structural responses occur.Here, we suggest an integrative approach to simulate drought stress responses. Firstly, a decreasing plant water potential triggers stomatal closure alongside a downregulation of photosynthetic performance. This is followed by a disconnection of roots and soil and the reliance on internal stem water storage or water uptake from deep soil layers. Consistently, loss in hydraulic functioning is reflected in sapwood loss of functionality and foliage senescence. This new model functionality has been used to investigate responses of tree hydraulics, carbon uptake and transpiration to soil- and atmospheric drought in an extremely dry Aleppo pine (Pinus halepensis L.) plantation.Using the hypothesis of a sequential triggering of stress-mitigating responses, the model was able to reflect the carbon uptake and transpiration patterns under varying soil water supply and atmospheric demand especially during summer and responded realistically regarding medium-term responses such as leaf and sapwood senescence. In agreement with the high drought resistance observed at the site our model indicated little loss of hydraulic functioning in Aleppo pine, despite the intensive seasonal summer drought.

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 BallBerry 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 BallBerry slope of 7.1 without using transpiration measurements. Sensitivity tests reveal that LRU is more sensitive to photosynthetic capacity than to the BallBerry 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.

Remote sensing (RS) for vegetation monitoring can involve mixed pixels with contributions from vegetation and background surfaces, causing biases in signals and their interpretations, especially in low-density forests. In a case study in the semi-arid Yatir forest in Israel, we observed a mismatch between satellite (Landsat 8 surface product) and tower-based (Skye sensor) multispectral data and contrasting seasonal cycles in near-infrared (NIR) reflectance. We tested the hypothesis that this mismatch was due to the different fractional contributions of the various surface components and their unique reflectance. Employing an unmanned aerial vehicle (UAV), we obtained high-resolution multispectral images over selected forest plots and estimated the fraction, reflectance, and seasonal cycle of the three main surface components (canopy, shade, and sunlit soil). We determined that the Landsat 8 data were dominated by soil signals (70%), while the tower-based data were dominated by canopy signals (95%). We then developed a procedure to resolve the canopy (i.e., tree foliage) normalized difference vegetation index (NDVI) from the mixed satellite data. The retrieved and corrected canopy-only data resolved the original mismatch and indicated that the spatial variations in Landsat 8 NDVI were due to differences in stand density, while the canopy-only NDVI was spatially uniform, providing confidence in the local flux tower measurements.

Global warming and drying trends, as well as the increase in frequency and intensity of droughts, may have unprecedented impacts on various forest ecosystems. We assessed the role of internal water storage (WS) in drought resistance of mature pine trees in the semi-arid Yatir Forest. Transpiration (T), soil moisture and sap flow (SF) were measured continuously, accompanied by periodical measurements of leaf and branch water potential (ψleaf) and water content (WC). The data were used to parameterize a tree hydraulics model to examine the impact of WS capacitance on the tree water relations. The results of the continuous measurements showed a 5-h time lag between T and SF in the dry season, which peaked in the early morning and early afternoon, respectively. A good fit between model results and observations was only obtained when the empirically estimated WS capacitance was included in the model. Without WS during the dry season, ψleaf would drop below a threshold known to cause hydraulic failure and cessation of gas exchange in the studied tree species. Our results indicate that tree WS capacitance is a key drought resistance trait that could enhance tree survival in a drying climate, contributing up to 45% of the total daily transpiration during the dry season.

The drier climates predicted for many regions will result in reduced evaporative cooling, leading to leaf heat stress and enhanced mortality. The extent to which nonevaporative cooling can contribute to plant resilience under these increasingly stressful conditions is not well known at present. Using a novel, high accuracy infrared system for the continuous measurement of leaf temperature in mature trees under field conditions, we assessed leaf-to-air temperature differences (ΔT_leafair) of pine needles during drought. On mid-summer days, ΔT_leafair remained leafair was weakly related to variations in the radiation load and mean wind speed in the lower part of the canopy, but was dependent on canopy structure and within-canopy turbulence that enhanced the H. Nonevaporative cooling is demonstrated as an effective cooling mechanism in needle-leaf trees which can be a critical factor in forest resistance to drying climates. The generation of a large H at the leaf scale provides a basis for the development of the previously identified canopy-scale convector effect.

Drought can cause tree mortality through hydraulic failure and carbon starvation. To prevent excess water loss, plants typically close their stomata before massive embolism formation occurs. However, unregulated water loss through leaf cuticles and bark continues after stomatal closure. Here, we studied the diurnal and seasonal dynamics of bark transpiration and how it is affected by tree water availability. We measured continuously for six months water loss and CO2 efflux from branch segments and needle-bearing shoots in Pinus halepensis growing in a control and an irrigation plot in a semi-arid forest in Israel. Our aim was to find out how much passive bark transpiration is affected by tree water status in comparison with shoot transpiration and bark CO2 emission that involve active plant processes, and what is the role of bark transpiration in total tree water use during dry summer conditions. Maximum daily water loss rate per bark area was 0.03-0.14 mmol m(-2) s(-1), which was typically ~76% of the shoot transpiration rate (on leaf area basis) but could even surpass the shoot transpiration rate during the highest evaporative demand in the control plot. Irrigation did not affect bark transpiration rate. Bark transpiration was estimated to account for 64-78% of total water loss in drought-stressed trees, but only for 6-11% of the irrigated trees, due to differences in stomatal control between the treatments. Water uptake through bark was observed during most nights, but it was not high enough to replenish the lost water during the day. Unlike bark transpiration, branch CO2 efflux decreased during drought due to decreased metabolic activity. Our results demonstrate that although bark transpiration represents a small fraction of the total water loss through transpiration from foliage in non-stressed trees, it may have a large impact during drought.

The leaf economics spectrum^1,2 and the global spectrum of plant forms and functions³ revealed fundamental axes of variation in plant traits, which represent different ecological strategies that are shaped by the evolutionary development of plant species². Ecosystem functions depend on environmental conditions and the traits of species that comprise the ecological communities⁴. However, the axes of variation of ecosystem functions are largely unknown, which limits our understanding of how ecosystems respond as a whole to anthropogenic drivers, climate and environmental variability^4,5. Here we derive a set of ecosystem functions⁶ from a dataset of surface gas exchange measurements across major terrestrial biomes. We find that most of the variability within ecosystem functions (71.8%) is captured by three key axes. The first axis reflects maximum ecosystem productivity and is mostly explained by vegetation structure. The second axis reflects ecosystem water-use strategies and is jointly explained by variation in vegetation height and climate. The third axis, which represents ecosystem carbon-use efficiency, features a gradient related to aridity, and is explained primarily by variation in vegetation structure. We show that two state-of-the-art land surface models reproduce the first and most important axis of ecosystem functions. However, the models tend to simulate more strongly correlated functions than those observed, which limits their ability to accurately predict the full range of responses to environmental changes in carbon, water and energy cycling in terrestrial ecosystems^7,8.

The chapter describes a long-term (20002019) perspective of carbon and energy fluxes from the leaf to the whole ecosystem scale in a semi-arid Aleppo pine forest plantation (the Yatir Forest) in the northern edge of the Negev desert (Israel), using the eddy covariance approach combined with meteorological and supplemental small-scale measurements. The site is characterized by a long dry season (over 7 months) with mean annual precipitation of 285 mm. The forest was a carbon sink during the wet season from December to April, reaching a maximum Net Ecosystem Production (NEP) rate of 4 gC m−2 d−1 in March, and a carbon source in JuneOctober, with a mean summer value of ~ −1 gC m−2 d−1. The carbon inventory showed that during 20012016 Yatir Forest accumulated 145 ± 26 gC m−2 year−1, ~71% of which was stored in the soil. By sequestering carbon, the forest has a cooling effect on the earths surface. The solar radiation burden over the forest was high, with an annual average flux of 240 Wm−2, while the forest albedo (the reflected solar radiation) was 0.12 and significantly lower than that of the surrounding desert, which was 0.25. Heat exchange with the atmosphere was characterized by high sensible heat flux (H) representing 5090% of the net absorbed radiation flux (Rn). The net absorbed Rn by the forest ecosystem was 67% higher than in the surrounding desert; the additional absorbed radiation has a warming effect on the surface. Our campaign-based measurements in another Aleppo pine forest in northern Israel with 755 mm annual precipitation (the Birya Forest, 190 Km north of Yatir) showed that this forest was a carbon sink both in the wet and dry seasons with mean NEP ~5 and ~3 gC m−2 d−1, respectively. The vegetation in the ecosystems around Birya Forest was much more developed than the desert vegetation in Yatir, thus the albedo effect and the thermal radiation suppression were lower. The net radiation load difference between Birya Forest and its surroundings was one-third lower than between Yatir and its surrounding desert. This case study show that pine forests can adjust to conditions at the dry, semi-arid, timberline and store significant amounts of carbon below ground with long residence time. Its large effects on the surface energy budget can result with local warming, but this can change considerably across relatively small geographical scale.

Climate change will impact forest productivity worldwide. Forecasting the magnitude of such impact, with multiple environmental stressors changing simultaneously, is only possible with the help of process-based models. In order to assess their performance, such models require careful evaluation against measurements. However, direct comparison of model outputs against observational data is often not reliable, as models may provide the right answers due to the wrong reasons. This would severely hinder forecasting abilities under unprecedented climate conditions. Here, we present a methodology for model assessment, which supplements the traditional output-to-observation model validation. It evaluates model performance through its ability to reproduce observed seasonal changes of the most limiting environmental driver (MLED) for a given process, here daily gross primary productivity (GPP). We analyzed seasonal changes of the MLED for GPP in two contrasting pine forests, the Mediterranean Pinus halepensis Mill. Yatir (Israel) and the boreal Pinus sylvestris L. Hyytiälä (Finland) from three years of eddy-covariance flux data. Then, we simulated the same period with a state-of-the-art process-based simulation model (LandscapeDNDC). Finally, we assessed if the model was able to reproduce both GPP observations and MLED seasonality. We found that the model reproduced the seasonality of GPP in both stands, but it was slightly overestimated without site-specific fine-tuning. Interestingly, although LandscapeDNDC properly captured the main MLED in Hyytiälä (temperature) and in Yatir (soil water availability), it failed to reproduce high-temperature and high-vapor pressure limitations of GPP in Yatir during spring and summer. We deduced that the most likely reason for this divergence is an incomplete description of stomatal behavior. In summary, this study validates the MLED approach as a model evaluation tool, and opens up new possibilities for model improvement.

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 0.941.07). 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.

Drought-induced productivity reductions and tree mortality have been increasing in recent decades in forests around the globe. Prescribed reduction in stand density, i.e. thinning, has been proposed as a management tool to improve forest sustainability in face of a warmer, drier future. Thinning should potentially reduce net stand water use and improve water-availability for remaining trees, thus reducing their subsequent drought vulnerability. However, few studies have directly measured these effects.In 2009 we established a large-scale thinning experiment in a semi-arid, 40-years-old pine afforestation. Study plots (70 x 70 m) were thinned to 100,200, and 300 trees ha(-1), and compared with unthinned control plots (210-400 trees ha(-1)), each at five replications. Stem and needle growth, and needle gas exchange were measured along 3-9 consecutive years at seasonal to annual temporal resolution. Measurements at the tree-scale were further up-scaled using both simple upscaling relationships and using an ecosystem model of coupled carbon, energy and hydrology (Regional Hydro Ecologic Simulation System, RHESSys).At the needle scale, photosynthesis was 70% higher at the 100 trees ha(-1) than at 300 trees ha(-1), whereas transpiration was merely 10% higher. Consequently, stem and needle growth increased by 100% and 20%, respectively. For most parameters, there was little change between 200 and 100 trees ha(-1). Applying RHESSys at the stand-scale, these effects on tree physiology translated into 35% reduction in CO2 uptake and a 47% reduction in tree water-use, which was compensated for by increased evaporation from exposed soil.Our long-term measurements at the dry timberline highlight the role of thinning in enhancing the activity and growth of remaining trees, with increased water-use efficiency. Unexpectedly, this density reduction was associated with a relatively small decrease in forest carbon uptake. Light availability was a limiting factor in the higher density plots, even in our light-abundant forest.

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

Efficiency and energetics of the turbulent transport in the canopy sublayer of a semi-arid pine forest and the atmospheric surface layer of a sparse desert-like shrubland are investigated from ultrasonic anemometer measurements during the summer dry season. The results show that an increased sensible heat flux over the forest canopy is generated by more energetic turbulence at small and large scales, but the transport process itself is equally efficient compared to the neighbouring shrubland. In contrast, the turbulent momentum flux over the forest canopy is caused by more efficient and energetic momentum transport at large scales. The more energetic turbulence appears to reduce the aerodynamic resistance to heat transfer of the semi-arid forest, which enables the increased sensible heat flux at a lower surface temperature. The results help explain the observed differences in diurnal variation of statistical moments of turbulent quantities that are caused by the interaction between radiative forcing, the background wind and the different turbulence production regimes of the forest and the shrubland. Lastly, the results also explain the observed cooler nighttime temperatures and quicker formation of a residual layer at the forest site.

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.

Carbonyl sulfide (COS) is a tracer of ecosystem photosynthesis that can advance carbon cycle research from leaf to global scales; however, a range of newly reported caveats related to sink/source strength of various ecosystem components hinder its application. Using comprehensive eddy-covariance and chamber measurements, we systematically measure ecosystem contributions from leaf, stem, soil, and litter and were able to close the ecosystem COS budget. The relative contributions of nonphotosynthetic components to the overall canopy-scale flux are relatively small (~4% during peak activity season) and can be independently estimated based on their responses to temperature and humidity. Converting COS to photosynthetic CO₂ fluxes based on the leaf relative uptake of COS/CO₂, faces challenges due to observed daily and seasonal changes. Yet, this ratio converges around a constant value (~1.6), and the variations, dominated by light intensity, were found unimportant on a flux-weighted daily time-scale, indicating a mean ratio of daytime gross-to-net primary productivity of ~2 in our ecosystem. The seasonal changes in the leaf relative uptake ratio may indicate a reduction in mesophyll conductance in winter, and COS-derived canopy conductance permitted canopy temperature estimate consistent with radiative skin temperature. These results support the feasibility of using COS as a powerful and much-needed means of assessing ecosystem function and its response to change.

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.

Estimations of ecosystem-level evapotranspiration (ET) and CO 2 uptake in water-limited environments are scarce and scaling up ground-level measurements is not straightforward. A biophysical approach was previously proposed for ecosystem-level assessment relying on vegetation index and meteorological data (RS-Met) in temperate Mediterranean ecosystems. However, these RS-Met models have not been tested yet in extreme high-energy water-limited ecosystems that suffer from continuous stress conditions. Owing to the lack of ET and CO 2 flux estimations in the Eastern Mediterranean, we examined the RS-Met approach using a newly developed mobile lab system and the single active Fluxnet station operating in this region, in seven forest and non-forest sites across a climatic transect in Israel (280-770 mm y −1 ). The RS-Met models were used with and without the addition of a seasonal drought stress factor (f DS ), which was based on daily rainfall, temperature and radiation data.Results show that the RS-Met models with the inclusion of the f DS were significantly improved compared to the non-f DS models (r=0.64-0.91 compared to 0.05-0.80; P=0.06 and r=0.72-0.92 compared to r=0.56-0.90; P

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 (20062010) in pine forests distributed along a rainfall gradient (P = 280820 mm yr⁻¹) in the Eastern Mediterranean (aridity factor = 0.170.56). 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. In the context of predicted climate change, such adjustments are key factors in sustaining forested catchments in water-limited regions.

This is a protocol to evaluate gross primary productivity (GPP) of a forest stand based on the measurements of trees sap flow (SF), 13C derived water use efficiency (WUE), and meteorological (met) data. GPP was calculated from WUE and stomatal conductance (gs), the later obtained from SF up-scaled from sampled trees to stand level on a daily time-scale and met data. WUE is obtained from 13C measurements in dated tree-ring wood and/or foliage samples. This protocol is based on the recently published study of Klein et al., 2016.

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 23-fold higher A and g s but with much more variability between seasons, especially for Q. ithaburensis with A and g s that decreased 1030-fold from spring to summer. Terpene emission rates for pine were not different regionally in spring but they were 58-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

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 (Ψ_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.

Understanding the processes that control the terrestrial exchange of carbon is critical for assessing atmospheric CO₂ budgets. Carbonyl sulfide (COS) is taken up by vegetation during photosynthesis following a pathway that mirrors CO₂ but has a small or nonexistent emission component, providing a possible tracer for gross primary production. Field measurements of COS and CO₂ mixing ratios were made in forest, senescent grassland, and riparian ecosystems using a laser absorption spectrometer installed in a mobile trailer. Measurements of leaf fluxes with a branch-bag gas-exchange system were made across species from 10 genera of trees, and soil fluxes were measured with a flow-through chamber. These data show (1) the existence of a narrow normalized daytime uptake ratio of COS to CO₂ across vascular plant species of 1.7, providing critical information for the application of COS to estimate photosynthetic CO₂ fluxes and (2) a temperature-dependent normalized uptake ratio of COS to CO₂ from soils. Significant nighttime uptake of COS was observed in broad-leafed species and revealed active stomatal opening prior to sunrise. Continuous high-resolution joint measurements of COS and CO₂ concentrations in the boundary layer are used here alongside the flux measurements to partition the influence that leaf and soil fluxes and entrainment of air from above have on the surface carbon budget. The results provide a number of critical constraints on the processes that control surface COS exchange, which can be used to diagnose the robustness of global models that are beginning to use COS to constrain terrestrial carbon exchange. Key Points Carbonyl sulfide can be measured in situ using a laser absorption spectrometer Ratio of COS to CO2 fluxes from soils and plants converge on a normalized value Soil and plant CO2 fluxes can be partitioned using ambient COS and CO2 conc

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 (300treesha^-1) in Israel's semiarid northern Negev, during three consecutive years (starting 2003/4) in which rainfall (R) was 231, 334 and 224mm, 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 (T) ranged from 0.1 to 1.6mmd^-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) ≤0.15m³m^-3, the primary driver of T was SWC; regression of T on SWC yielded highly significant quadratic relationships indicating little response below SWC of approximately 0.12m³m^-3, and a steep linear response above it. For SWC >0.15m³m^-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 (215d), on average, cumulative ET (201mm) accounted for 0.76 R (R=263mm) and cumulative T (116mm; range 103-126mm) - an independently estimated component of cumulative ET - accounted for 0.45 R. Cumulative E was 70mm. On an annual basis, total evapotranspiration losses were approximately equal to R, with 58% exiting the system via the trees and 39% via soil and undergrowth vegetation. Water balance data combined with assumptions regarding tree minimum transpiration led to a first approximation of sustainable forest density. This approach indicated that the Yatir forest should be thinned to stands of 250 or 190treesha^-1 in order to remain sustainable under annual rainfall regimes of 200 or 150mm, respectively.

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.

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 nighttime 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 CO ₂ 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 CO ₂ 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 \mbox{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 12mL septum-capped vials for 0.5-4h before aliquots of incubation air were injected into 3.7mL 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±0.01atm; storing the gas samples at -17°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 10-31, with precision generally better than ±0.5. 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°C; difference in minimum temperatures of 4.6°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 ∼0.8, rural/remote ∼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 ¹³C 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 environmental conditions may mask dramatic shifts in respiratory functioning. Overall, our findings indicating greater seedling growth and establishment at a critical regeneration phase of forest development may have important implications for the ecology of urban forests as well as the predicted growth of the terrestrial biosphere in temperate regions in response to climate change.

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 NO _x 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 ¹³C and ¹⁸O 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 (~0.04°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 ¹³C concentration in atmospheric CO₂ has been declining over the past 150 years as large quantities of ¹³C-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 ¹³C 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 ¹³C 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. ¹³C analyses of paper from two European and two American publications showed, on average, a - 1.65 1.00 trend between 1880 and 2000, compared with - 1.45 and - 1.57 for air and tree-ring analyses, respectively. The greater decrease in plant-derived ¹³C 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 ∼1 (±0.25) 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.

We report the first investigation of changes in electron partitioning via the alternative respiratory pathway (AP) and alternative oxidase (AOX) protein abundance in field-grown plants and their role in seasonal acclimation of respiration. We sampled two alpine grasses native to New Zealand, Chionochloa rubra and Chionochloa pallens, from field sites of different altitudes, over 1yr and also intensively over a 2-wk period. In both species, respiration acclimated to seasonal changes in temperature through changes in basal capacity (R₁₀) but not temperature sensitivity (E₀). In C. pallens, acclimation of respiration may be associated with a higher AOX cytochrome c oxidase (COX) protein abundance ratio. Oxygen isotope discrimination (D), which reflects relative changes in AP electron partitioning, correlated positively with daily integrated photosynthetically active radiation (PAR) in both species over seasonal timescales. Respiratory parameters, the AOX:COX protein ratio and D were stable over a 2-wk period, during which significant temperature changes were experienced in the field. We conclude that respiration in Chionochloa spp. acclimates strongly to seasonal, but not to short-term, temperature variation. Alternative oxidase appears to be involved in the plant response to both seasonal changes in temperature and daily changes in light, highlighting the complexity of the function of AOX in the field.

The distribution of precipitation inputs into different hydrological components of water-limited forest ecosystems determines water availability to trees and consequently forest productivity. We constructed a complete hydrological budget of a semi-arid pine forest (285 mm annual precipitation) by directly measuring its main components: precipitation (P), soil water content, evapotranspiration (ET, eddy covariance), tree transpiration (sap flux), soil evaporation (soil chambers), and intercepted precipitation (calculated). Our results indicated that on average for the 4-year study period, ET accounted for 94% of P, varying between 100% when P 300 mm (with indications for losses to subsurface flow and soil moisture storage in wetter years). Direct measurements of the components of the ET flux demonstrated that both transpiration and soil evaporation were significant in this dry forest (45% and 36% of ET, respectively). Comparison between ecosystem ET (eddy covariance measurements) and the sum of its measured components showed good agreement on annual scales, but up to 30% discrepancies (in both directions) on shorter timescales. The pulsed storm pattern, characteristics of semi-arid climates, was sufficient to maintain the topsoil layer wet during the whole wet season. Only less often and intensive storms resulted in infiltration to the root zone, increasing water availability for uptake by deeper roots. Our results indicate that climate change predictions that link reduced precipitation with increased storm intensity may have a smaller effect on water availability to forest ecosystems than reduced precipitation alone, which could help forests' survival and maintain productivity even under drier conditions.

P>Carbonyl sulfide (COS) exchange in C-3 leaves is linked to that of CO2, providing a basis for the use of COS as a powerful tracer of gross CO2 fluxes between plants and the atmosphere, a critical element in understanding the response of the land biosphere to global change. Here, we carried out controlled leaf-scale gas-exchange measurements of COS and CO2 in representative C-3 plants under a range of light intensities, relative humidities and temperatures, CO2 and COS concentrations, and following abscisic acid treatments. No 'respiration-like' emission of COS or detectable compensation point, and no cross-inhibition effects between COS and CO2 were observed. The mean ratio of COS to CO2 assimilation flux rates, As/Ac, was c. 1.4 pmol mu mol-1 and the leaf relative uptake (assimilation normalized to ambient concentrations, (As/Ac)(C(a)c/C(a)s)) was 1.6-1.7 across species and conditions, with significant deviations under certain conditions. Stomatal conductance was enhanced by increasing COS, which was possibly mediated by hydrogen sulfide (H2S) produced from COS hydrolysis, and a correlation was observed between As and leaf discrimination against C18OO. The results provide systematic and quantitative information necessary for the use of COS in photosynthesis and carbon-cycle research on the physiological to global scales.

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 uncatalyzed rate, consistent with typical population sizes for CAcontaining soil microorganisms. Including accelerated soil hydration in global model simulations modifies contributions from soil and foliage to the global CO¹⁸O 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 CO ₂ gross fluxes obtained by using the δ¹⁸O of atmospheric CO₂ by 50%.

This paper presents a study of the isotopic composition of dissolved inorganic carbon (DIC) stable and, radioactive, in pore water of the unsaturated zone (USZ) above the coastal aquifer of Israel The carbon. content and its isotopic composition in the gas and solid phases of the USZ are also presented. In the soil gas, large quantities of CO2 with quite modern C-14 activity were measured along the section (0.15 to 2.7% and 97 to 109 pMC respectively). In the inorganic fraction of the sediments, the C-14 activity between 2.5 and 13 m was 33 to 1.5 pMC. In the organic fraction the 14C activity between 7 and 9 m was 40 to 33 pMC. In the pore water, the high values of tritium and C-14 at depths of similar to 15-18 m have been attributed to the thermonuclear contamination of the 1960s. A significant decrease with depth in the DIC of pore water, from 23 to 4 mmol C/L, along with a decrease with depth in dissolved inorganic C-14, from 100 pMC near the surface to 72 pMC at depth of 20 m were observed. The delta C-13 values in the DIC are similar to the values in the inorganic sediment (similar to -10 parts per thousand). A first order reaction was applied to estimate the yearly rates of DIC loss by net precipitation (3.2%/year) and of C-14 activity (dpm/L) loss from the DIC by gross precipitation from the DIC on the sediment, (4.4%/year). From the gradient of dissolved inorganic C-14, the initial level at the bottom of the USZ, which is the groundwater table of the coastal aquifer of Israel, was estimated to be 0.54 of contemporaneous atmospheric value of C-14 when the rain fell on the ground. This value can be used for improved dating of groundwater with C-14 in the coastal aquifer of Israel. (C) 2009 Elsevier B.V. All rights reserved.

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 F_s 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 Q₁₀ = 1.18 when soil moisture was kept high by a supplement summer irrigation treatment. Interannual variations in F₅ 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 ∼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 ∼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%.

Biological soil crusts (BSC) contribute significantly to the soil surface cover in many dryland ecosystems. A mixed type of BSC, which consists of cyanobacteria, mosses and cyanolichens, constitutes more than 60% of ground cover in the semiarid grass-shrub steppe at Sayeret Shaked in the northern Negev Desert, Israel. This study aimed at parameterizing the carbon sink capacity of well-developed BSC in undisturbed steppe systems. Mobile enclosures on permanent soil borne collars were used to investigate BSC-related CO₂ fluxes in situ and with natural moisture supply during 10 two-day field campaigns within seven months from fall 2001 to summer 2002. Highest BSC-related CO ₂ deposition between ĝ\u20ac"11.31 and ĝ\u20ac"17.56 mmol m⁻² per 15 h was found with BSC activated from rain and dew during the peak of the winter rain season. Net CO₂ deposition by BSC was calculated to compensate 120%, ĝ\u20ac"26%, and less than 3% of the concurrent soil CO₂ efflux from Novemberĝ\u20ac"January, Februaryĝ\u20ac"May and Novemberĝ\u20ac"May, respectively. Thus, BSC effectively compensated soil CO₂ effluxes when CO₂ uptake by vascular vegetation was probably at its low point. Nighttime respiratory emission reduced daily BSC-related CO₂ deposition within the period Novemberĝ\u20ac"January by 11ĝ\u20ac"123% and on average by 27%. The analysis of CO₂ fluxes and water inputs from the various sources showed that the bulk of BSC-related CO₂ deposition occurs during periods with frequent rain events and subsequent condensation from water accumulated in the upper soil layers. Significant BSC activity on days without detectable atmospheric water supply emphasized the importance of high soil moisture contents as additional water source for soil-dwelling BSC, whereas activity upon dew formation at low soil water contents was not of major importance for BSC-related CO₂ deposition. However, dew may still be important in attaining a pre-activated status during the transition from a long "summer" anabiosis towards the first winter rain.

This study explored possible advantages conferred by the phase shift between leaf phenology and photosynthesis seasonality in a semi-arid Pinus halepensis forest system, not seen in temperate sites. Leaf-scale measurements of gas exchange, nitrogen and phenology were used on daily, seasonal and annual time-scales. Peak photosynthesis was in late winter, when high soil moisture, mild temperatures and low leaf vapour pressure deficit (D_L) allowed high rates associated with high water- and nitrogen-use efficiencies. Self-sustained new needle growth through the dry and hot summer maximized photosynthesis in the following wet season, without straining carbon storage. Low rates of water loss were associated with increasing sensitivity of stomatal conductance (g_s) to soil moisture below a relative extractable water (REW) of 0.4, and decreased g_s sensitivity to D_L below REW of approx. 0.2. This response was captured by the modified Ball-Berry (Leuning) model. While most physiological parameters and responses measured were typical of temperate pines, the photosynthesis- phenological phasing contributed to high productivity under warm-dry conditions. This contrasts with reported effects of short-term periodical droughts and could lead to different predictions of the effect of warming and drying climate on pine forest productivity.

To identify factors that influence the relatively high productivity of a semi-arid pine afforestation system in southern Israel, we investigated inorganic nitrogen deposition and mineralization for more than 2 years. To this end, we measured bulk and dry deposition, in situ N-mineralization over the seasonal cycle, and the potential activity of nitrifying microorganisms by soil slurry incubations. There was a small increase in bulk N deposition in the forest, compared with shrubland, but no change in dry deposition. An unexpected rapid increase in nitrite concentration in the forest soil was observed after soil rewetting by the first winter rains, which could not be explained by deposition. This was accompanied by a decrease in ammonium and only a slight increase in nitrate concentrations. Only a small increase in nitrite and a rapid increase in nitrate concentration in the mineral soil were observed in the surrounding shrubland. Soil slurry incubations from the forest sites exhibited significant delay in nitrite, compared with nitrate accumulation (up to 50 h under lab conditions) in samples taken in the dry season, but not in the wet season. This indicated different rates of ammonium and nitrite oxidation that are most likely linked to differential activation of different microbial populations after the summer stress. The initial oxidation process of ammonia to nitrate, upon soil rewetting in semi-arid environments, appears to occur as a partially uncoupled two-step process, as opposed to a rapid continuous one in wetter environments. This may have implications for the synchronization of nitrate availability to plants and therefore for high forest productivity and nitrogen use efficiency. Forest productivity in the semi-arid regions, in turn, is becoming increasingly more important with persistent predictions of warming and drying trends over the entire Mediterranean basin and other regions.

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 m² 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 (R² = 0.79, p

Vacuum distillation is shown to be useful for the quantitative extraction of dissolved inorganic carbon (as CO₂) and water from sediments of the unsaturated zone in the Coastal Aquifer of Israel. Several tests of vacuum extractions from tap water and sediments are presented, including standard addition, which show that the distillation procedure is quantitative, with minimal or no carbon isotope fractionation. The optimal temperature of the sediment during the extraction was also defined. Examples of vacuum extractions of sediments are shown.

The isotopic composition of atmospheric O₂ depends on the rates of oxygen cycling in photosynthesis, respiration, photochemical reactions in the stratosphere and on δ¹⁷O and δ¹⁸O of ocean and leaf water. While most of the factors affecting δ¹⁷O and δ¹⁸O of air O₂ have been studied extensively in recent years, δ¹⁷O of leaf water-the substrate for all terrestrial photosynthesis-remained unknown. In order to understand the isotopic composition of atmospheric O₂ 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 δ¹⁷O and δ¹⁸O 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(δ¹⁷O + 1) vs. ln(δ¹⁸O + 1) plots are characterized by very high precision (∼0.001) despite of relatively large differences between duplicates in both δ¹⁷O and δ¹⁸O (0.02-0.05). This is so because the errors in δ¹⁸O and δ¹⁷O 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 × 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.

This paper reports on ranges of carbon dioxide (CO₂) 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 on 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 dunes 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 CO₂ fluxes on 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 ₂ exchange measurements, linear mixture model, and NDVI values is suitable for monitoring BSC's productivity in arid regions.

Variation in the C¹⁸OO content of atmospheric CO ₂ (δ¹⁸O_a) can be used to distinguish photosynthesis from soil respiration, which is based on carbonic anhydrase (CA)-catalyzed ¹⁸O exchange between CO₂ and ¹⁸O-enriched leaf water (δ¹⁸O_w). Here we tested the hypothesis that mean leaf δ¹⁸O_w and assimilation rates can be used to estimate whole-leaf C¹⁸OO 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 δ¹⁸O_w (7 to 21), δ¹⁸O and δ¹³C of leaf organic matter (25 to 30 and -12.8 to -13.2, respectively), and substomatal CO₂ concentrations (intercellular CO₂ concentrations, c_i, at the leaf center were ∼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 ∼10%. Conversely, use of measured parameters from a leaf segment could produce large errors in predicting leaf-integrated C¹⁸OO fluxes.

The use of the ¹³C : ¹²C isotopic ratio (δ¹³C) of leaf-respired CO₂ to trace carbon fluxes in plants and ecosystems is limited by little information on temporal variations in δ¹³C of leaf dark-respired CO₂ (δ¹³C_r) under field conditions. Here, we explored variability in δ¹³C_r and its relationship to key respiratory substrates from collections of leaf dark-respired CO ₂, carbohydrate extractions and gas exchange measurements over 24-h periods in two Quercus canopies. Throughout both canopies, δ¹³C_r became progressively ¹³C-enriched during the photoperiod, by up to 7, then ¹³C-depleted at night relative to the photoperiod. This cycle could not be reconciled with δ¹³C of soluble sugars (δ¹³C_ss), starch (δ¹³C_st), lipids (δ¹³C _l), cellulose (δ¹³C_c) or with calculated photosynthetic discrimination (Δ). However, photoperiod progressive enrichment in δ¹³C_r was correlated with cumulative carbon assimilation (r² = 0.91). We concluded that there is considerable short-term variation in δ¹³C_r in forest canopies, that it is consistent with current hypotheses for ¹³C fractionation during leaf respiration, that leaf carbohydrates cannot be used as surrogates for δ¹³C_r, and that diel changes in leaf carbohydrate status could be used to predict changes in δ ¹³C_r empirically.

Associations between delta C-13 values and leaf gas exchanges and tree-ring or needle growth, used in ecophysiological compositions, can be complex depending on the relative timing of CO2 uptake and subsequent redistribution and allocation of carbon to needle and stem components. For palaeoenvironmental and dendroecological studies it is often interpreted in terms of a simple model of delta C-13 fractionation in C-3 plants. However, in spite of potential complicating factors, few studies have actually examined these relationships in mature trees over inter- and intra-annual time-scales. Here, we present results from a 4 years study that investigated the links between variations in leaf gas-exchange properties, growth, and dated delta C-13 values along the needles and across tree rings of Aleppo pine trees growing in a semi-arid region under natural conditions or with supplemental summer irrigation. Sub-sections of tissue across annual rings and along needles, for which time of formation was resolved from growth rate analyses, showed rapid growth and 6 C responses to changing environmental conditions. Seasonal cycles of growth and delta C-13 (Up to similar to 4%.) significantly correlated (P

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 benefit native perennial shrub re-generation in the intact ecosystem. Taken together, the results of our study indicate that vegetation transition from shrublands to post-fire successional plant communities may (1) decrease water recharge in winters leading to lower water storage in winter and spring and potentially reduce groundwater recharge; (2) decrease the amount of plant-available soil water in the root zone during phenologically important times of the years; and (3) reduce the lateral patchiness in surface soil water, which may impede the re-establishment of sagebrush or other patch-dependent native perennials.

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 the whole 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.

The ¹⁸O content of atmospheric O₂ is an important tracer for past changes in the biosphere. Its quantitative use depends on knowledge of the discrimination against ¹⁸O associated with the various O₂ consumption processes. Here we evaluated, for the first time, the in situ ¹⁸O discrimination associated with soil respiration in natural ecosystems. The discrimination was estimated from the measured [O₂] and δ¹⁸O of O₂ in the soil-air. The discriminations that were found are 10.1 ± 1.5, 17.8 ± 1.0, and 22.5 ± 3.6, for tropical, temperate, and boreal forests, respectively, 17.9 ± 2.5 for Mediterranean woodland, and 15.4 ± 1:6 for tropical shrub land. Current understanding of the isotopic composition of atmospheric O₂ is based on the assumption that the magnitude of the fractionation in soil respiration is identical to that of dark respiration through the cytochrome pathway alone (∼18). The discrimination we found in the tropical sites is significantly lower, and is explained by slow diffusion in soil aggregates and root tissues that limits the O₂ concentration in the consumption sites. The high discrimination in the boreal sites may be the result of high engagement of the alternative oxidase pathway (AOX), which has high discrimination associated with it (∼27 . The intermediate discrimination (∼18) in the temperate and Mediterranean sites can be explained by the opposing effects of AOX and diffusion limitation that cancel out. Since soil respiration is a major component of the global oxygen uptake, the contribution of large variations in the discrimination, observed here, to the global Dole Effect should be considered in global scale studies.

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 (δ¹³C) measurements in myrtle (Myrtus communis), buckthorn (Rhamnus alaternus), and velvet bean (Mucuna pruriens) showed that only 72% to 91% of the variations in the δ¹³C values of fixed carbon were reflected in the δ¹³C 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 δ¹³C 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 ¹³C relative to mean photosynthetically fixed carbon by 4 to 11. Variable ¹³C 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.

The oxygen-18 (O-18) content of atmospheric carbon dioxide (CO2) is an important indicator of CO2 uptake on land. It has generally been assumed that during photosynthesis, oxygen in CO2 reaches isotopic equilibrium with oxygen in O-18-enriched water in leaves. We show, however, large differences in the activity of carbonic anhydrase (which catalyzes CO2 hydration and O-18 exchange in leaves) among major plant groups that cause variations in the extent of O-18 equilibrium (theta (eq)). A clear distinction in theta (eq) between C-3 trees and shrubs, and C-4 grasses makes atmospheric (COO)-O-18 a potentially sensitive indicator to changes in C-3 and C-4 productivity. We estimate a global mean theta (eq) value of similar to0.8, which reasonably reconciles inconsistencies between O-18 budgets of atmospheric O-2 (Dole effect) and CO2.

Stable isotopes are a powerful research tool in environmental sciences and their use in ecosystem research is increasing. In this review we introduce and discuss the relevant details underlying the use of carbon and oxygen isotopic compositions in ecosystem gas exchange research. The current use and potential developments of stable isotope measurements together with concentration and flux measurements of CO₂ and water vapor are emphasized. For these applications it is critical to know the isotopic identity of specific ecosystem components such as the isotopic composition of CO₂, organic matter, liquid water, and water vapor, as well as the associated isotopic fractionations, in the soil-plant-atmosphere system. Combining stable isotopes and concentration measurements is very effective through the use of 'Keeling plots.' This approach allows the identification of the isotopic composition and the contribution of ecosystem, or ecosystem components, to the exchange fluxes with the atmosphere. It also allows the estimation of net ecosystem discrimination and soil disequilibrium effects. Recent modifications of the Keeling plot approach permit examination of CO₂ recycling in ecosystems. Combining stable isotopes with dynamic flux measurements requires precision in isotopic sampling and analysis, which is currently at the limit of detection. Combined with the micrometeorological gradient approach (applicable mostly in grasslands and crop fields), stable isotope measurements allow separation of net CO₂ exchange into photosynthetic and soil respiration components, and the evapotranspiration flux into soil evaporation and leaf transpiration. Similar applications in conjunction with eddy correlation techniques (applicable to forests, in addition to grasslands and crop fields) are more demanding, but can potentially be applied in combination with the Keeling plot relationship. The advance and potential in using stable isotope measurements should make their use a standard component in the limited arsenal of ecosystem-scale research tools.

Measurements of ¹⁸O in atmospheric CO₂ can be used to trace gross photosynthetic and respiratory CO₂ fluxes between the atmosphere and the terrestrial biosphere. However, this requires knowledge of the ¹⁸O signatures attributable to the fluxes from soil and leaves. Newly developed methods were employed to measure the ¹⁸O of soil-respired CO₂ and depth profiles of near-surface soil CO₂, in order to evaluate the factors influencing isotopic soil-atmosphere CO₂ exchange. The ¹⁸O of soil-respired CO₂ varied predominantly as a function of the ¹⁸O of soil water which, in turn, changed with soil drying and with seasonal variations in source water. The ¹⁸O of soil-respired CO₂ corresponds to full isotopic equilibrium with soil water at a depth ranging between 5 and 15 cm. The ¹⁸O of respired CO₂, in reality, results from a weighted average of partial equilibria over a range of depths. Soil water isotopic enrichment of up to 10 in the top 5 cm did not appear to strongly influence the isotopic composition of the respired CO₂. We demonstrate that during measurements 'invasion' of atmospheric CO₂ (the diffusion of ambient CO₂ into the soil, followed by partial equilibration and retrodiffusion) must be considered to accurately calculate the ¹⁸O of the soil-respired CO₂. The impact of invasion in natural settings is also considered. We also have determined the effective kinetic fractionation of CO₂ diffusion out of the soil to be 7.2 ± 0.3. High-resolution (1 cm) depth profiles of ¹⁸O of near-surface (top 10 cm) soil CO₂ were carried out by gas chromatography-isotope ratio mass spectrometry (GC-IRMS). This novel technique allowed us to observe the competitive diffusion-equilibration process near the soil surface and to test simulations by a diffusion and equilibration model of the soil CO₂¹⁸O content.

CO₂ profiles obtained along a 30-m-thick unsaturated zone under land irrigated with sewage effluents show two production regions: a seasonal one in the root zone and another, at steady state, near the water table (29 m). On an annual basis the CO₂ flux from the deep source toward the atmosphere (6.3 g C m^-2 yr^-1) is balanced by a similar influx of soluble organic carbon (SOC) from sewage effluents. The δ¹³C values of soil CO₂ indicate that CO₂ is produced from plant material in the root zone and from biodegradation of total sedimentary organic carbon in the capillary fringe. High CO₂ concentration in the capillary fringe (up to 2%) is likely to reflect a decrease in diffusivity relative to the unsaturated zone due to increase in both water content and tortuosity induced by the capillary fringe structure. The long residence time of SOC and CO₂ in the unsaturated zone (29 years at the study site) suggests that the unsaturated zone of deep aquifers may have a significant storage capacity for carbon and may act as a temporary carbon sink.

Keywords: PROTEIN

The ¹³C/¹²C and ¹⁸O/¹⁶O 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 δ¹⁸O of the wood cellulose was strongly similar to that observed in leaf water over a diurnal cycle. Most of the data for δ¹³C and all of the data for δ¹⁸O could be fitted to two independent linear equations that, combined, allowed the reconstruction of RH and the δ¹⁸O 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 δ¹⁸O 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 clear 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 mixing patterns of water from the water-storage multiple-epidermis (ME) and the spongy parenchyma (SP) of leaves of Peperomia congesta HBK were studied by following variations in the natural abundance of deuterium. The ²H/¹H ratios of water from the two tissues were different throughout the diel cycle and were always lower in the ME. Withholding water from the potted plants resulted in complete isotopic homogeneity of leaf water within 3 days. Isotopic homogeneity and the extent of ²H enrichment of leaf water were constant for the duration of the 3-week stress period while the weight of the ME differentially and continuously decreased during the same period. Reinstating irrigation resulted in the rapid restoration of pre-stress ME weight, and a simultaneous sharp drop in the ²H/¹H ratio of water in both the ME and SP. The proportion of recharge by soil water was similar whether calculated based on changes in tissue weight or on ²H/¹H ratios. In contrast to the rapid recovery in water content, isotopic enrichment and restoration of isotopic heterogeneity were gradual over several days. The results indicate that under favorable conditions water, and not only solutes, stored in the ME does not mix well with water in other tissues, but becomes a more integral part of total leaf water under water stress conditions. In recovery from water stress, recharge with soil water is rapid (hours) while communication with the external environment, as reflected by isotopic enrichment and restoration of isotopic heterogeneity in leaf water, is gradual (days). The results demonstrate the potential of using stable isotopes at the natural abundance level to study the dynamics of water movement within plants.