Xylem Water Isotope Research Articles

Larger water use efficiency associated with greater dependence on deep water sources in hot-dry valleys with reservoirs

Identifying the carbon–water relationship of plants helps to understand their adaptive water use strategies. The relationship between WUE and the water source remains uncertain in the hot-dry valleys with reservoirs. The leaf δ13C and soil and xylem water isotopes (δ2H and δ18O) were determined to identify the WUE and water source contributions, respectively, in Leucaena leucocephala (Lam.) de Wit communities. We propose a source contributing score (SCS), calculated as a value for the contribution of water from different soil layers (a higher SCS value indicates plants use deeper water sources). The positive relationship between leaf δ13C and SCS (R2 = 0.16, P = 0.022) and the negative relationship between leaf δ13C and topsoil contribution (R2 = 0.21, P = 0.009) confirmed the close association of larger WUE with less reliance on shallow soil water. The redundancy analysis suggested a stronger connection between leaf δ13C and d-excess than leaf δ13C and topsoil moisture content, which emphasized the key role of soil moisture d-excess. Variation partitioning analysis (VPA) suggested soil properties and microclimate explain more than 89.8 % of the variations in tree water use. VPA also suggested different roles of microclimate and soil properties in explaining tree water use strategies. Findings broadened the understanding of the relationship between WUE and water sources and emphasized the key role of soil moisture d-excess in hot-dry valleys with reservoirs.

Tree‐ and stand‐scale variability of xylem water stable isotope signatures in mature beech, oak and spruce

AbstractIn ecohydrology, water isotopologues are used to assess potential sources of root water uptake by comparing xylem water signatures with source water signatures. Such comparisons are affected by the variability and uncertainty of the isotope signatures of plant water and water sources. The tree‐scale and stand‐scale variabilities of the isotope signatures in stem xylem water are often unknown but are important for sampling design and uncertainty estimation in assessing the sources of tree water uptake. Here, we quantified tree‐scale and stand‐scale variabilities of xylem water isotope signatures in beech, oak and spruce trees in a mature forest on the Swiss plateau. For stem xylem water, sub‐daily replicates and replicates in different cardinal directions showed no systematic differences, but we found systematic differences with sampling height. The observed variability of isotope signatures at different heights along the stem suggests that water residence times within trees need to be considered, along with their effects on the isotope signatures in different compartments (stem, branches, leaves). Further, concerning the hydrogen signatures, we found height‐ and species‐specific offsets (SW‐excess δ2H). Stem xylem water's tree‐scale variability was similar in magnitude to its stand‐scale variability and smaller than the variabilities in branch xylem and bulk soil water around each tree. Xylem water from stem cores close to the ground, therefore, can give a more precise estimate of the isotopic signal of the most recent root water uptake and facilitate more accurate source water attribution.

Root water uptake model shows age-related water uptake patterns of apple trees on the Chinese Loess Plateau

Study regionChangwu Tableland, Chinese Loess Plateau Study focusSoil water status and water demand are two important factors to influence root water uptake (RWU) patterns, but the mechanisms involved in deep-rooted plants remain unknown. This study addresses this knowledge gap by investigating diurnal variations in RWU patterns for two different-aged apple orchards (7 vs. 19 years) on the Chinese Loess Plateau (CLP). We achieved this by inputting measured plant and soil information into a physical-based numerical RWU model. New hydrological insights for the regionThe results showed that the 19-year-old orchard exhibits distinct characteristics compared to the 7-year-old orchard. Specifically, the 19-year-old orchard had less deep soil water content, resulting in lower matric potential and hydraulic conductance. In contrast to cropland soil water, the cumulative deep soil water (>2 m) deficits were 17.6 and 1108.3 mm in the 7-year-old and 19-year-old orchards, respectively. Despite the 19-year-old orchard having 67% of its fine roots below 5 m soil depth, their contribution ratio to total RWU was only 30–40%, indicating less available deep soil water for the older orchard.Sap flow and leaf water potential had bell-shaped diurnal variations, leading to shifting soil water contributions to RWU in the 7-year-old and 19-year-old orchards. These changes were particularly evident when soil water potential in the root zone was heterogeneous, resulting in contrasting RWU patterns during different water demand periods. Notably, the 7-year-old orchard exhibited less water contribution from deep 2–5 m layer and enriched xylem water isotopes at midday with high water demand, while the 19-year-old orchard had more water contribution from deep 5–21.6 m layer and depleted xylem water isotopes. These contrasting RWU variations shed light on the profound impact of orchard age on deep RWU patterns, especially under the varying water demand conditions after CLP afforestation.

Evaluating the bias effects of rooting depth and cryogenic vacuum extraction to quantify root water uptake patterns in deep-rooted apple trees

Accurately assessing isotope information on water sources and mixtures is essential for determining root water uptake (RWU) patterns. This study investigates the impact of rooting depth and cryogenic vacuum extraction (CVE) on the isotopic composition of deep soil water and RWU patterns in a 19-year-old apple orchard on China’s Loess Plateau. We used a Bayesian mixing model (MixSIAR) to analyze corrected isotopic data from soil and xylem water samples collected at different depths, ranging from 3 m to 21.6 m. Our findings reveal a progressive decline in the isotopic composition of deep soil water by 0.63‰ m–1, which decreased deep soil water contributions to transpiration by 1–12% at different rooting depths. The xylem and soil water isotopes became more enriched after correction, with xylem water isotopes closer to the soil water isotope line and water isotopes enriched in the shallow soil layer. Consequently, the contributions of shallow soil water to transpiration increased after xylem water isotope correction but decreased after soil water isotope correction. Monthly averages of these ratios decreased by 12.8% or increased by 8.4%, respectively. After correcting for both soil and xylem water isotopes, the average contributions of shallow soil water increased by 7.4% each month, while those of deep soil water were not consistent. Our findings suggest that rooting depth and CVE have a greater impact on seasonal contribution ratios but a comparatively milder influence on seasonal patterns. This delineation is important for accurately quantifying plant water use strategies.

Water stable isotope characteristics and water use strategies of co-occurring plants in ecological and economic forests in subtropical monsoon regions

Identifying the spatio-temporal variation in water use by co-occurring species in mixed forests can determine whether there is hydrological niche partitioning or niche overlapping, i.e., whether plant co-occurrence brings about resource complementation or interspecific competition, and provides a reference for plantation management. Water stable isotopic compositions in precipitation, soil, and four plants (xylem and leaf) in ecological and economic forests and hydrometeorological factors were continuously monitored for a year, and the water uptake by plants was determined using a Bayesian MixSIR model. The isotopic compositions of soil water and plant water in the ecological forest (Camphora camphora and Cunninghamia lanceolate) and economic forest (Castanea mollissima and Camellia pitardii) showed similar seasonal variation, and no significant difference between their mean isotopic values was found (p > 0.05). The isotopic compositions in leaf water of different plants in the two forests were controlled by xylem water isotope and meteorological factors, rather than soil moisture conditions. With decreasing soil water availability, the water sources of C. camphora, C. lanceolate, and C. pitardii changed from shallow to deep, while the water use pattern of C. mollissima was insensitive to changing soil moisture conditions. The water source overlapping between C. camphora and C. lanceolate in the ecological forest might generate water resource competition during the dry period, limiting their ecological functions. On the other hand, the water source partitioning between C. mollissima and C. pitardii in the economic forest facilitated the water resource allocation. Our study suggested that the overlapping and partitioning of hydrological niches among species in mixed forests were species-specific and provided a scientific basis for fully exploiting the ecological and economic benefits of silviculture ecosystems.

Differences between stem and branch xylem water isotope composition in four tropical tree species

AbstractStable isotope studies (δ2H and δ18O) of water within plants are providing new information on water sources, competitive interactions and water use patterns under natural conditions. This is based on the assumption that there is no fractionation at the time of water uptake or during its transport within trees. However, previous studies have found fractionation does occur in water taken up through the roots in halophytic and xerophytic plants. It is unclear how widespread such fractionation is in other species. In this study, we tested this fractionation by comparing stem and branch xylem water isotopes over the period of one day (from 8 am to 5 pm) in four wet tropical rainforest tree species (Dendrocnide photinophylla,Aphananthe philippinensis,Daphnandra repandulaandMallotus polyadenos). We found branch water isotope ratios (δ2H and δ18O) were enriched compared with stem xylem water isotopes.D. photinophyllahad a significantly different branch δ18O ratio thanA. philippinensis,D. repandulaandM. polyadenos. In contrast, there were no significant differences in stem xylem δ18O among the four observed tree species. We found clear differences in the stable isotope ratios of δ18O for stem and branch xylem water forD. photinophyllaandD. repandulaof upto −0.85% and 0.50%, respectively. Remarkable δ2H differences were also found between stem and branch xylem water isotope ratios forA. philippinensis,D. repandulaandM. polyadenos, being upto −12.14%, −16.17% and −9.65%, respectively. A dual isotope (δ2H–δ18O) plot showed branch water values were more enriched than stem xylem water values forD. photinophyllaandD. repandula, indicating a clear difference between stem and branch xylem water. This study suggests that δ2H and δ18O fractionation could be a species‐specific phenomenon in tropical trees, which has important implications for plant water source identification and evapotranspiration partitioning.

Precipitation fate and transport in a Mediterranean catchment through models calibrated on plant and stream water isotope data

Abstract. To predict hydrologic responses to inputs and perturbations, it is important to understand how precipitation is stored in catchments, released back to the atmosphere via evapotranspiration (ET), or transported to aquifers and streams. We investigated this partitioning of precipitation using stable isotopes of water (18O) at the Can Vila catchment in the Spanish Pyrenees mountains. The isotope data covered four years of measurements, comprising >550 rainfall and >980 stream water samples, capturing intra-event variations. They were complemented by fortnightly plant water isotope data sampled over eight months. The isotope data were used to quantify how long it takes for water to become evapotranspiration or to be discharged as streamflow using StorAge Selection (SAS) functions. We calibrated the SAS functions using a conventional approach fitting the model solely to stream water isotope data and a multi-objective calibration approach in which the model was simultaneously fitted to tree xylem water isotope data. Our results showed that the conventional model-fitting approach was not able to constrain the model parameters that represented the age of water supplying ET. Consequently, the ET isotope ratios simulated by the conventionally calibrated model failed to adequately simulate the observed xylem isotope ratios. However, the SAS model was capable of adequately simulating both observed stream water and xylem water isotope ratios, if those xylem water isotope observations were used in calibration (i.e., the multi-objective approach). The multi-objective calibration approach led to a more constrained parameter space facilitating parameter value identification. The model was tested on a segment of data reserved for validation showing a Kling–Gupta Efficiency of 0.72 compared to the 0.83 observed during in the calibration period. The water-age dynamics inferred from the model calibrated using the conventional approach differed substantially from those inferred from the multi-objective calibration model. The latter suggested that the median ages of water supplying evapotranspiration is much older (150–300 d) than what was suggested by the former (50–200 d). Regardless, the modeling results support recent findings in ecohydrological field studies that highlighted both subsurface heterogeneity of water storage and fluxes and the use of relatively old water by trees. We contextualized the SAS-derived water ages by also using young-water-fraction and endmember-splitting approaches, which respectively also showed the contribution of young water to streamflow was variable but sensitive to runoff rates and that ET was largely sourced by winter precipitation that must have resided in the subsurface across seasons.

Continuous in situ measurements of water stable isotopes in soils, tree trunk and root xylem: Field approval.

New methods to measure stable isotopes of soil and tree water directly in the field enable us to increase the temporal resolution of obtained data and advance our knowledge on the dynamics of soil and plant water fluxes. Only few field applications exist. However, these are needed to further improve novel methods and hence exploit their full potential. We tested the borehole equilibration method in the field and collected in situ and destructive samples of stable isotopes of soil, trunk and root xylem water over a 2.5-month experiment in a tropical dry forest under natural abundance conditions and following labelled irrigation. Water from destructive samples was extracted using cryogenic vacuum extraction. Isotope ratios were determined with IRIS instruments using cavity ring-down spectroscopy both in the field and in the laboratory. In general, timelines of both methods agreed well for both soil and xylem samples. Irrigation labelled with heavy hydrogen isotopes clearly impacted the isotope composition of soil water and one of the two studied tree species. Inter-method deviations increased in consequence of labelling, which revealed their different capabilities to cover spatial and temporal heterogeneities. We applied the novel borehole equilibration method in a remote field location. Our experiment reinforced the potential of this in situ method for measuring xylem water isotopes in both tree trunks and roots and confirmed the reliability of gas permeable soil probes. However, in situ xylem measurements should be further developed to reduce the uncertainty within the range of natural abundance and hence enable their full potential.

Temporal dynamics of tree xylem water isotopes: in situ monitoring and modeling

Abstract. We developed a setup for a fully automated, high-frequency in situ monitoring system of the stable water isotope deuterium and 18O in soil water and tree xylem. The setup was tested for 12 weeks within an isotopic labeling experiment during a large artificial sprinkling experiment including three mature European beech (Fagus sylvatica) trees. Our setup allowed for one measurement every 12–20 min, enabling us to obtain about seven measurements per day for each of our 15 in situ probes in the soil and tree xylem. While the labeling induced an abrupt step pulse in the soil water isotopic signature, it took 7 to 10 d until the isotopic signatures at the trees' stem bases reached their peak label concentrations and it took about 14 d until the isotopic signatures at 8 m stem height leveled off around the same values. During the experiment, we observed the effects of several rain events and dry periods on the xylem water isotopic signatures, which fluctuated between the measured isotopic signatures observed in the upper and lower soil horizons. In order to explain our observations, we combined an already existing root water uptake (RWU) model with a newly developed approach to simulate the propagation of isotopic signatures from the root tips to the stem base and further up along the stem. The key to a proper simulation of the observed short-term dynamics of xylem water isotopes was accounting for sap flow velocities and the flow path length distribution within the root and stem xylem. Our modeling framework allowed us to identify parameter values that relate to root depth, horizontal root distribution and wilting point. The insights gained from this study can help to improve the representation of stable water isotopes in trees within ecohydrological models and the prediction of transit time distribution and water age of transpiration fluxes.

Robust Estimation of Absorbing Root Surface Distributions From Xylem Water Isotope Compositions With an Inverse Plant Hydraulic Model

The vertical distribution of absorbing roots is one of the most influential plant traits determining plant strategy to access below ground resources. Yet little is known of natural variability in root distribution since collecting field data is challenging and labor-intensive. Studying stable water isotope compositions in plants could offer a cost-effective and practical solution to estimate the absorbing root surfaces distribution. However, such an approach requires developing realistic inverse modeling techniques that enable robust estimation of rooting distributions and associated uncertainty from xylem water isotopic composition observations. This study introduces an inverse modeling method that supports the assessment of the root allocation parameter (β) that defines the exponential vertical decay of a plants’ absorbing root surfaces distribution with soil depth. The method requires measurements obtained from xylem and soil water isotope composition, soil water potentials, and sap flow velocities when plants’ xylem water is sampled at a certain height above the rooting point. In a simulation study, we show that the approach can provide unbiased estimates of β and its associated uncertainty due to measuring errors and unmeasured environmental factors that can impact the xylem water isotopic data. We also recommend improving the accuracy and power of β estimation, highlighting the need for considering accurate soil water potential and sap flow monitoring. Finally, we apply the inverse modeling method to xylem water isotope data of lianas and trees collected in French Guiana. Our work shows that the inverse modeling procedure provides a robust analytical and statistical framework to estimate β. The method accounts for potential bias due to extraction errors and unmeasured environmental factors, which improves the viability of using stable water isotope compositions to estimate the distribution of absorbing root surfaces complementary to the assessment of relative root water uptake profiles.

Isotopic fractionation from deep roots to tall shoots: A forensic analysis of xylem water isotope composition in mature tropical savanna trees

Studies of plant water sources generally assume that xylem water integrates the isotopic composition (δ2H and δ18O) of water sources and does not fractionate during uptake or transport along the transpiration pathway. However, woody xerophytes, halophytes, and trees in mesic environments can show isotopic fractionation from source waters. Isotopic fractionation and variation in isotope composition can affect the interpretation of tree water sources, but most studies to date have been greenhouse experiments. Here we present a field-based forensic analysis of xylem water isotope composition for 12 Eucalyptus tetrodonta and Corymbia nesophila trees. We used a 25-tonne excavator to access materials from the trees' maximum rooting depth of 3 m to their highest canopies at 38 m. Substantial within-tree variation occurred in δ2H (−91.1‰ to −35.7‰ E. tetrodonta; −88.8‰ to −24.5‰ C. nesophila) and δ18O (−12.3‰ to −5.0‰ E. tetrodonta; −10.9‰ to −0.3‰ C. nesophila), with different root-to-branch isotope patterns in each species. Soil water δ2H and δ18O dual isotope slopes (7.26 E. tetrodonta, 6.66 C. nesophila) were closest to the Local Meteoric Water Line (8.4). The dual isotope slopes of the trees decreased progressively from roots (6.45 E. tetrodonta, 6.07 C. nesophila), to stems (4.61 E. tetrodonta, 5.97 C. nesophila) and branches (4.68 E. tetrodonta, 5.67 C. nesophila), indicative of fractionation along the xylem stream. Roots of both species were more enriched in 2H and 18O than soil water at all sampled depths. Bayesian mixing model analysis showed that estimated proportions of water sourced from different depths reflected the contrasting root systems of these species. Our study adds evidence of isotopic fractionation from water uptake and along the transpiration stream in mature trees in monsoonal environments, affecting the interpretation of water sources. We discuss the findings with view of interpreting aboveground xylem water isotopic composition, incorporating knowledge of root systems.

Tropical forest water source patterns revealed by stable isotopes: A preliminary analysis of 46 neighboring species

Stable isotope tracing of plant water use can illuminate plant water sources. But to date, the number of species tested at any given site has been minimal. Here, we sample 46 tropical hardwood tree species in a 0.32 ha plot with uniform soils. Soil water was characterized at 6 depths at 0.2 m intervals down to 1 m and showed simple and predictable depth patterns of δ2H and δ18O, and simple and spatially uniform isotope composition at each depth. Nevertheless, tree xylem water δ2H and δ18O showed remarkable variation covering the full range of soil composition, suggesting strong sorting and niche segregation across the small plot. Wood density, tree size and mean basal area increment together explained approximately 55% of the variance of xylem water isotope composition through principal component analysis. A Bayesian mixing model was applied to the data and showed that sampled trees were either sourcing their water from very shallow or deep soil layers, with very little contribution from the middle portion of the soil profile. The majority of the observed species relied on 0.0–0.2 m depth soil water. This layer contributed approximately 75% of the xylem water which was significantly higher than the contributions from all other depths. The contribution from shallow soil was highest for trees with high wood density, slow-growing trees and small-sized trees. Our work suggests that stable isotope tracers may aid a better understanding of tropical forest water uptake depths and their relation to tree functional traits and potential hydrological niche segregation among co-occurring tropical species.

The use of regional and alluvial groundwater by riparian trees in the wet‐dry tropics of northern Australia

AbstractRiparian trees play a critical role in the ecological function of rivers, yet are threatened by anthropogenic change to the hydrological cycle. Identifying the sources of water used by riparian trees can inform sustainable water policy. We used isotopic analysis complemented by measurements of plant water relations to assess water sources for riparian trees at two sites with contrasting hydrogeological processes; one with an alluvial aquifer overlaying an aquitard, and one where fault‐induced preferential pathways in the aquitard allowed the flow of deeper, older groundwater from a regional aquifer to the alluvium. At both sites, plant water potential, stomatal conductance, and plant water isotope composition in the xylem sap of riparian trees were collected from two landscape positions, the riverbank and floodplain. We used a Bayesian mixing model (MixSIAR) to assess differences in the proportion of water sources for sites and landscape positions. We found that xylem water isotope values differed between the two sites in line with their hydrogeological characteristics, with trees at the regional aquifer site using water sourced from the regional groundwater and trees at the site with only an alluvial aquifer present using a mixture of water sources, with no dominant source identified. Higher plant predawn water potential values at the regional site indicated greater water availability and support the inference that plants were using more groundwater at the regional site compared to the alluvial site. Trees closer to the river had higher isotope values, indicative of surficial water sources i.e. shallow soil water and river water. Our findings show that the water sources used by riparian trees reflect local hydrogeology and resource availability. Water managers should identify and protect plant water sources to ensure maintenance of riparian trees.

In situ measurements of soil and plant water isotopes: a review of approaches, practical considerations and a vision for the future

Abstract. The number of ecohydrological studies involving water stable isotope measurements has been increasing steadily due to technological (e.g., field-deployable laser spectroscopy and cheaper instruments) and methodological (i.e., tracer approaches or improvements in root water uptake models) advances in recent years. This enables researchers from a broad scientific background to incorporate water-isotope-based methods into their studies. Several isotope effects are currently not fully understood but might be essential when investigating root water uptake depths of vegetation and separating isotope processes in the soil–vegetation–atmosphere continuum. Different viewpoints exist on (i) extraction methods for soil and plant water and methodological artifacts potentially introduced by them, (ii) the pools of water (mobile vs. immobile) measured with those methods, and (iii) spatial variability and temporal dynamics of the water isotope composition of different compartments in terrestrial ecosystems. In situ methods have been proposed as an innovative and necessary way to address these issues and are required in order to disentangle isotope effects and take them into account when studying root water uptake depths of plants and for studying soil–plant–atmosphere interaction based on water stable isotopes. Herein, we review the current status of in situ measurements of water stable isotopes in soils and plants, point out current issues and highlight the potential for future research. Moreover, we put a strong focus and incorporate practical aspects into this review in order to provide a guideline for researchers with limited previous experience with in situ methods. We also include a section on opportunities for incorporating data obtained with described in situ methods into existing isotope-enabled ecohydrological models and provide examples illustrating potential benefits of doing so. Finally, we propose an integrated methodology for measuring both soil and plant water isotopes in situ when carrying out studies at the soil–vegetation–atmosphere continuum. Several authors have shown that reliable data can be generated in the field using in situ methods for measuring the soil water isotope composition. For transpiration, reliable methods also exist but are not common in ecohydrological field studies due to the required effort. Little attention has been paid to in situ xylem water isotope measurements. Research needs to focus on improving and further developing those methods. There is a need for a consistent and combined (soils and plants) methodology for ecohydrological studies. Such systems should be designed and adapted to the environment to be studied. We further conclude that many studies currently might not rely on in situ methods extensively because of the technical difficulty and existing methodological uncertainties. Future research needs to aim on developing a simplified approach that provides a reasonable trade-off between practicability and precision and accuracy.

Variation in the access to deep soil water pools explains tree-to-tree differences in drought-triggered dieback of Mediterranean oaks

Individual differences in the access to deep soil water pools may explain the differential damage among coexisting, conspecific trees as a consequence of drought-induced dieback. We addressed this issue by comparing the responses to a severe drought of three Mediterranean oak species with different drought tolerance, Quercus pubescens L. and Quercus frainetto Ten., mainly thriving at xeric and mesic sites, respectively, and Quercus cerris L., which dominates at intermediate sites. For each species, we compared coexisting declining (D) and non-declining (ND) trees. The stable isotope composition (δ2H, δ18O) of xylem and soil water was used to infer a differential use of soil water sources. We also measured tree size and radial growth to quantify the long-term divergence of wood production between D and ND trees and non-structural carbohydrates (NSCs) in sapwood to evaluate if D trees presented lower NSC values. The ND trees had access to deeper soil water than D trees except in Q. frainetto, as indicated by significantly more depleted xylem water values. However, a strong δ2H offset between soil and xylem water isotopes observed in peak summer could suggest that both tree types were not physiologically active under extreme drought conditions. Alternative processes causing deuterium fractionation, however, could not be ruled out. Tree height and recent (last 15-25years) growth rates in all species studied were lower in D than in ND trees by 22 and 44%, respectively. Lastly, there was not a consistent pattern of NSC sapwood concentration; in Q. pubescens, it was higher in ND trees while in Q. frainetto, the D trees were the ones exhibiting the higher NSC concentration. We conclude that the vulnerability to drought among conspecific Mediterranean oaks depends on the differential access to deep soil water pools, which may be related to differences in rooting depth, tree size and growth rate.

Using Water Stable Isotopes to Trace Water Sources of Three Typical Japanese Tree Species under Heavy Rainfall Conditions

Stable isotopes of xylem water (18O and D) have been successfully used to determine sources of soil water for plant transpiration, but mainly in drought-prone environments. The water uptake strategies of three representative tree species in Japan, namely cedar (Cryptomeria japonica), larch (Larix kaempferi) and beech (Fagus crenata), were investigated using δ18O and δD of water (precipitation, soil and xylem), together with wood α-cellulose δ13C and δ18O, along one growing season. The study was carried out in the research forest of Yamagata University (Shonai region), a high precipitation area in Japan, which exceeds 3000 mm per year. Precipitation water δ18O and δD increased along the summer growing season, but oxygen and hydrogen isotopic composition of soil water remained essentially unchanged. In general, xylem water isotopes of cedar and larch followed the local meteoric water line, but beech xylem water was decoupled from soil and precipitation values in July and August. For this tree species, the xylem water isotopic records were more depleted than cedar and larch xylem water isotopic values and the precipitation water isotopic records, indicating that beech used more water from soil layers during July-August than the other two species, which mainly used newly-fallen precipitation. Wood δ18O showed an opposite seasonal trend to the one found for xylem water, likely because of leaf water isotope enrichment, which was in turn controlled by seasonal transpiration rate. The higher δ13C values of cedar during summer suggested that this species had enhanced water-use efficiency during the growing season compared with the deciduous species larch and beech. Our results highlight different water use strategies among forest tree species even in areas where the annual water balance is far from limiting plant performance.

The role of deep vadose zone water in tree transpiration during drought periods in karst settings – Insights from isotopic tracing and leaf water potential

Karst environments are unusual because their dry, stony and shallow soils seem to be unfavorable to vegetation, and yet they are often covered with forests. How can trees survive in these environments? Where do they find the water that allows them to survive? This study uses midday and predawn water potentials and xylem water isotopes of branches to assess tree water status and the origin of transpired water. Monitoring was conducted during the summers of 2014 and 2015 in two dissimilar plots of Mediterranean forest located in karst environments. The results show that the three monitored tree species (Abies alba Mill, Fagus sylvatica L, and Quercus ilex L.) use deep water resources present in the karst vadose zone (unsaturated zone) more intensively during drier years. Quercus ilex, a species well- adapted to water stress, which grows at the drier site, uses the deep water resource very early in the summer season. Conversely, the two other species exploit the deep water resource only during severe drought. These results open up new perspectives to a better understanding of ecohydrological equilibrium and to improved water balance modeling in karst forest settings.

Using stable isotopes to quantify water sources for trees and shrubs in a riparian cottonwood ecosystem in flood and drought years

AbstractRiparian cottonwood forests in dry regions of western North America do not typically receive sufficient growing season precipitation to completely support their relatively high transpiration requirements. Water used in transpiration by riparian ecosystems must include alluvial groundwater or water stored in the potentially large reservoir of the unsaturated soil zone. We used the stable oxygen and hydrogen isotope composition of stem xylem water to evaluate water sources used by the dominant riparian cottonwood (Populus spp.) trees and shrubs (Shepherdia argentea and Symphoricarpos occidentalis) in Lethbridge, Alberta, during 3 years of contrasting environmental conditions. Cottonwoods did not exclusively take up alluvial groundwater but made extensive use of water sourced from the unsaturated soil zone. The oxygen and hydrogen isotope compositions of cottonwood stem water did not strongly overlap with those of alluvial groundwater, which were closely associated with the local meteoric water line. Instead, cottonwood stem water δ18O and δ2H values were located below the local meteoric water line, forming a line with a low slope that was indicative of water exposed to evaporative enrichment of heavy isotopes. In addition, cottonwood xylem water isotope compositions had negative values of deuterium excess (d‐excess) and line‐conditioned (deuterium) excess (lc‐excess), both of which provided evidence that water taken up by the cottonwoods had been exposed to fractionation during evaporation. The shrub species had lower values of d‐excess and lc‐excess than had the cottonwood trees due to shallower rooting depths, and the d‐excess values declined during the growing season, as shallow soil water that was taken up by the plants was exposed to increasing, cumulative evaporative enrichment. The apparent differences in functional rooting pattern between cottonwoods and the shrub species, strongly influenced the ratio of net photosynthesis to stomatal conductance (intrinsic water‐use efficiency), as shown by variation among species in the δ13C values of leaf tissue.

Seasonal origins of soil water used by trees

Abstract. Rain recharges soil water storages and either percolates downward into aquifers and streams or is returned to the atmosphere through evapotranspiration. Although it is commonly assumed that summer rainfall recharges plant-available water during the growing season, the seasonal origins of water used by plants have not been systematically explored. We characterize the seasonal origins of waters in soils and trees by comparing their midsummer isotopic signatures (δ2H) to seasonal isotopic cycles in precipitation, using a new seasonal origin index. Across 182 Swiss forest sites, xylem water isotopic signatures show that summer rain was not the predominant water source for midsummer transpiration in any of the three sampled tree species. Beech and oak mostly used winter precipitation, whereas spruce used water of more diverse seasonal origins. Even in the same plots, beech consistently used more winter precipitation than spruce, demonstrating consistent niche partitioning in the rhizosphere. All three species' xylem water isotopes indicate that trees used more winter precipitation in drier regions, potentially mitigating their vulnerability to summer droughts. The widespread occurrence of winter isotopic signatures in midsummer xylem implies that growing-season rainfall may have minimally recharged the soil water storages that supply tree growth, even across diverse humid climates (690–2068 mm annual precipitation). These results challenge common assumptions concerning how water flows through soils and is accessed by trees. Beyond these ecological and hydrological implications, our findings also imply that stable isotopes of δ18O and δ2H in plant tissues, which are often used in climate reconstructions, may not reflect water from growing-season climates.

Dynamics of competition over water in a mixed oak-pine Mediterranean forest: Spatio-temporal and physiological components

Understanding inter- and intra-specific plant interactions and competition over water is challenging because of the lack of effective approaches for accessing and monitoring root distribution and activity. In this context, stable isotopes are excellent eco-hydrological tracers that allow characterizing the dynamics of water uptake patterns in trees and shrubs. Here, we studied biotic interactions for water uptake between two typical Mediterranean tree species, Aleppo pine (Pinus halepensis) and holm oak (Quercus ilex), coexisting in a mixed forest. We measured stable isotope composition (δ18O and δ2H) of xylem water in all trees found in the studied stand during one growing season, covering an exceptionally long summer drought and subsequent recovery. We applied point-process statistics together with stand density information to evaluate tree-to-tree interactions for water use. In pines, we observed a clear uncoupling between soil and xylem water isotope composition after two months of persistent drought. Conversely, the isotope composition of xylem water in oaks tracked observed changes in the soil during the first two months of drought, but began to depart from soil values after three months. These results suggest that during drought the oaks were able to keep active for longer using alternative soil water sources, not available for the pines. Point-process statistics revealed more positive isotope compositions at distances below 4–6m, but only between con-specific individuals (i.e. pine-pine, oak-oak). These intra-specific responses were first seen in the pines (after two months of drought) and subsequently in oaks (after three months), coinciding with the onset of soil-xylem uncoupling for each species. On the other hand, the isotope composition of individual oaks decreased with increasing neighbor pine density, but increased in response to oak density. Conversely, the pines showed more positive values with increasing oak density. Our results suggest that the use of shallow water in oaks is limited by the presence of pines, which force them to shift to deep-soil water use, whereas pines have more restricted access to deep water in the presence of oaks, leading to more positive isotope values. According to the dynamics of interaction patterns, we conclude that inter-specific differences in pine-oak mixed forests hold two components: a static, spatial component determined by root distribution, and a dynamic, physiological component related to water uptake capacity within the soil profile.