Climatic Influences on Summer Use of Winter Precipitation by Trees

Trees in seasonal climates may use water originating from both winter and summer precipitation. However, the seasonal origins of water used by trees have not been systematically studied. We used stable isotopes of water to compare the seasonal origins of water found in three common tree species across 24 Swiss forest sites sampled in two different years. Water from winter precipitation was observed in trees at most sites, even at the peak of summer, although the relative representation of seasonal sources differed by species. However, the representation of winter precipitation in trees decreased with site mean annual precipitation in both years; additionally, it was generally lower in the cooler and wetter year. Together, these relationships show that precipitation amount influenced the seasonal origin of water taken up by trees across both time and space. These results suggest higher turnover of the plant-available soil-water pool in wetter sites and wetter years

Seasonal Origins of Soil Water Used by Trees

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\u27 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

Seasonal origins of soil water used by trees

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&thinsp;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.</p

The influence of traffic and wood combustion on the stable isotopic composition of carbon monoxide

Carbon monoxide in the atmosphere is originating from various combustion and oxidation processes. Recently, the proportion of CO resulting from the combustion of wood for domestic heating may have increased due to political measures promoting this renewable energy source. Here, we used the stable isotope composition of CO (&amp;delta;&lt;sup&gt;13&lt;/sup&gt;C and &amp;delta;&lt;sup&gt;18&lt;/sup&gt;O) for the characterization of different CO sources in Switzerland, along with other indicators for traffic and wood combustion (NO&lt;sub&gt;x&lt;/sub&gt;-concentration, aerosol light absorption at different wavelengths). We assessed diurnal variations of the isotopic composition of CO at 3 sites during winter: a village site dominated by domestic heating, a site close to a motorway and a rural site. The isotope ratios of wood combustion emissions were studied at a test facility, indicating significantly lower &amp;delta;&lt;sup&gt;18&lt;/sup&gt;O of CO from wood combustion compared to traffic emissions. At the village and the motorway site, we observed very pronounced diurnal &amp;delta;&lt;sup&gt;18&lt;/sup&gt;O-variations of CO with an amplitude of up to 8&amp;permil;. Solving the isotope mass balance equation for three distinct sources (wood combustion, traffic, clean background air) resulted in diurnal patterns consistent with other indicators for wood burning and traffic. The average night-time contribution of wood-burning to total CO was 70% at the village site, 49% at the motorway site and 29% at the rural site based on the isotope mass balance. The results, however, depend strongly on the pure source isotope values, which are not very well known. We therefore additionally applied a combined CO/NO&lt;sub&gt;x&lt;/sub&gt;-isotope model for verification. Here, we separated the CO emissions into different sources based on distinct CO/NO&lt;sub&gt;x&lt;/sub&gt; emissions ratios for wood combustion and traffic, and inserted this information in the isotope mass balance equation. Accordingly, a highly significant agreement between measured and calculated &amp;delta;&lt;sup&gt;18&lt;/sup&gt;O-values of CO was found (&lt;i&gt;r&lt;/i&gt;=0.67, &lt;i&gt;p&lt;/i&gt;&lt;0.001). While different proxies for wood combustion all have their uncertainties, our results indicate that the oxygen isotope ratio of CO (but not the carbon isotope ratio) is an independent sensitive tool for source attribution studies

Technical note: On uncertainties in plant water isotopic composition following extraction by cryogenic vacuum distillation

Recent studies have challenged the interpretation of plant water isotopes obtained through cryogenic vacuum distillation (CVD) based on observations of a large 2H fractionation. These studies have hypothesized the existence of an H-atom exchange between water and organic tissue during CVD extraction with the magnitude of H exchange related to relative water content of the sample; however, clear evidence is lacking. Here, we systematically tested the uncertainties in the isotopic composition of CVD-extracted water by conducting a series of incubation and rehydration experiments using isotopically depleted water, water at natural isotope abundance, woody materials with exchangeable H, and organic materials without exchangeable H (cellulose triacetate and caffeine). We show that the offsets between hydrogen and oxygen isotope ratios and expected reference values (Δ2H and Δ18O) have inversely proportional relationships with the absolute amount of water being extracted, i.e. the lower the water amount, the higher the Δ2H and Δ18O. However, neither Δ2H nor Δ18O values, were related to sample relative water content. The Δ2H pattern was more pronounced for materials with exchangeable H atoms than with non-exchangeable H atoms. This is caused by the combined effect of H exchange during the incubation of materials in water and isotopic enrichments during evaporation and sublimation that depend on absolute water amount. The H exchange during CVD extraction itself was negligible. Despite these technical issues, we observed that the water amount-dependent patterns were much less pronounced for samples at natural isotope abundance and particularly low when sufficiently high amounts of water were extracted (&gt;600 µL). Our study provides new insights into the mechanisms causing isotope fractionation during CVD extraction of water. The methodological uncertainties can be controlled if large samples of natural isotope abundance are used in ecohydrological studies.</p

Technical Note: On Uncertainties in Plant Water Isotopic Composition Following Extraction by Cryogenic Vacuum Distillation

Recent studies have challenged the interpretation of plant water isotopes obtained through cryogenic vacuum distillation (CVD) based on observations of a large 2H fractionation. These studies have hypothesized the existence of an H-atom exchange between water and organic tissue during CVD extraction with the magnitude of H exchange related to relative water content of the sample; however, clear evidence is lacking. Here, we systematically tested the uncertainties in the isotopic composition of CVD-extracted water by conducting a series of incubation and rehydration experiments using isotopically depleted water, water at natural isotope abundance, woody materials with exchangeable H, and organic materials without exchangeable H (cellulose triacetate and caffeine). We show that the offsets between hydrogen and oxygen isotope ratios and expected reference values (Δ2H and Δ18O) have inversely proportional relationships with the absolute amount of water being extracted, i.e. the lower the water amount, the higher the Δ2H and Δ18O. However, neither Δ2H nor Δ18O values, were related to sample relative water content. The Δ2H pattern was more pronounced for materials with exchangeable H atoms than with non-exchangeable H atoms. This is caused by the combined effect of H exchange during the incubation of materials in water and isotopic enrichments during evaporation and sublimation that depend on absolute water amount. The H exchange during CVD extraction itself was negligible. Despite these technical issues, we observed that the water amount-dependent patterns were much less pronounced for samples at natural isotope abundance and particularly low when sufficiently high amounts of water were extracted (\u3e600 µL). Our study provides new insights into the mechanisms causing isotope fractionation during CVD extraction of water. The methodological uncertainties can be controlled if large samples of natural isotope abundance are used in ecohydrological studies

Spatial Variation in Throughfall, Soil, and Plant Water Isotopes in a Temperate Forest

Studies of stable isotopes of water in the environment have been fundamental to advancing our understanding of how water moves through the soil‐plant‐atmosphere continuum; however, much of this research focuses on how water isotopes vary in time, rather than in space. We examined the spatial variation in the δ18O and δ2H of throughfall and bulk soil water, as well as branch xylem and bulk leaf water of Picea abies (Norway Spruce) and Fagus sylvatica (Beech), in a 1 ha forest plot in the northern Alps of Switzerland. Means and ranges of water isotope ratios varied considerably among throughfall, soil, and xylem samples. Soil water isotope ratios were often poorly explained by soil characteristics and often not predictable from proximal samples. Branch xylem water isotope values varied less than either soil water or bulk leaf water. The isotopic range observed within an individual tree crown was often similar to that observed among different crowns. As a result of the heterogeneity in isotope ratios, inferences about the depth of plant root water uptake drawn from a two end‐member mixing model were highly sensitive to the soil sampling location. Our results clearly demonstrate that studies using water isotopes to infer root water uptake must explicitly consider how to characterize soil water, incorporating measures of both vertical and lateral variation. By accounting for this spatial variation and the processes that shape it, we can improve the application of water isotopes to studies of plant ecophysiology, ecohydrology, soil hydrology, and paleoclimatology

Impact of different nitrogen emission sources on tree physiology as assessed by a triple stable isotope approach

The importance that nitrogen (N) deposition has in driving the carbon (C) sequestration of forests has recently been investigated using both experimental and modeling approaches. Whether increased N deposition has positive or negative effects on such ecosystems depends on the status of the N and the duration of the deposition. By combining d13C, d18O, d15N and dendrochronological approaches, we analyzed the impact of two different sources of NOx emissions on two tree species, namely: a broadleaved species (Quercus cerris) that was located close to an oil refinery in Southern Italy, and a coniferous species (Picea abies) located close to a freeway in Switzerland. Variations in the ci/ca ratio and the distinction between stomatal and photosynthetic responses to NOx emissions in trees were assessed using a conceptual model, which combines d13C and d18O. d15N in leaves, needles and tree rings was found to be a bioindicator of N input from anthropogenic emissions, especially at the oil refinery site.We observed that N fertilization had a stimulatory effect on tree growth near the oil refinery, while the opposite effect was found for trees at the freeway site. Changes in the ci/ca ratio were mostly related to variations in d13C at the freeway site and, thus, were driven by photosynthesis. At the oil refinery site they were mainly related to stomatal conductance, as assessed using d18O. This study demonstrates that a single method approach does not always provide a complete picture of which physiological traits are more affected by N emissions. The triple isotope approach combined with dendrochronological analyses proved to be a very promising tool for monitoring the ecophysiological responses of trees to long-term N deposition

Updating the Dual C and O Isotope—Gas-exchange Model: A Concept to Understand Plant Responses to the Environment and Its Implications for Tree Rings

The combined study of carbon (C) and oxygen (O) isotopes in plant organic matter has emerged as a powerful tool for understanding plant functional responses to environmental change. The approach relies on established relationships between leaf gas exchange and isotopic fractionation to derive a series of model scenarios that can be used to infer changes in photosynthetic assimilation and stomatal conductance driven by changes in environmental parameters (CO2, water availability, air humidity, temperature, nutrients). We review the mechanistic basis for a conceptual model, in light of recently published research, and discuss where isotopic observations do not match our current understanding of plant physiological response to the environment. We demonstrate that (1) the model was applied successfully in many, but not all studies; (2) although originally conceived for leaf isotopes, the model has been applied extensively to tree-ring isotopes in the context of tree physiology and dendrochronology. Where isotopic observations deviate from physiologically plausible conclusions, this mismatch between gas exchange and isotope response provides valuable insights into underlying physiological processes. Overall, we found that isotope responses can be grouped into situations of increasing resource limitation versus higher resource availability. The dual-isotope model helps to interpret plant responses to a multitude of environmental factors

Ideas and perspectives : Tracing terrestrial ecosystem water fluxes using hydrogen and oxygen stable isotopes – challenges and opportunities from an interdisciplinary perspective

The authors thank Marialaura Bancheri, Michele Bottazzi, Roman Cibulka, Massimo Esposito, Alba Gallo, Cesar D. Jimenez-Rodriguez, Angelika Kuebert, Ruth Magh, Stefania Mambelli, Alessia Nannoni, Paolo Nasta, Vladimir Rosko, Andrea Rücker, Noelia Saavedra Berlanga, Martin Šanda, and Anna Scaini for their contributions during the discussion at the workshop “Isotope-based studies of water partitioning and plant–soil interactions in forested and agricultural environments”. The authors also thank “Villa Montepaldi” and the University of Florence for the access to the workshop location, and the municipality of San Casciano in Val di Pesa for logistical support. The authors thank the Department of Innovation, Research and University of the Autonomous Province of Bozen/Bolzano for covering the Open Access publication costs. Last, but not least, the authors wish to thank Matthias Sprenger, Stephen Good, and J. Renée Brooks, as well as the Editor David R. Bowling, whose constructive reviews greatly improved this manuscript.Peer reviewedPublisher PD