Maren Dubbert

Oxygen isotope signatures of transpired water vapor: the role of isotopic non‐steady‐state transpiration under natural conditions

The oxygen isotope signature of water is a powerful tracer of water movement from plants to the global scale. However, little is known about the short-term variability of oxygen isotopes leaving the ecosystem via transpiration, as high-frequency measurements are lacking. A laser spectrometer was coupled to a gas-exchange chamber directly estimating branch-level fluxes in order to evaluate the short-term variability of the isotopic composition of transpiration (δE ) and to investigate the role of isotopic non-steady-state transpiration under natural conditions in cork-oak trees (Quercus suber) during distinct Mediterranean seasons. The measured δ(18) O of transpiration (δE ) deviated from isotopic steady state throughout most of the day even when leaf water at the evaporating sites was near isotopic steady state. High agreement was found between estimated and modeled δE values assuming non-steady-state enrichment of leaf water. Isoforcing, that is, the influence of the transpirational δ(18) O flux on atmospheric values, deviated from steady-state calculations but daily means were similar between steady state and non-steady state. However, strong daytime isoforcing on the atmosphere implies that short-term variations in δE are likely to have consequences for large-scale applications, for example, partitioning of ecosystem fluxes or satellite-based applications.

Stable oxygen isotope and flux partitioning demonstrates understory of an oak savanna contributes up to half of ecosystem carbon and water exchange

Semi-arid ecosystems contribute about 40% to global net primary production (GPP) even though water is a major factor limiting carbon uptake. Evapotranspiration (ET) accounts for up to 95% of the water loss and in addition, vegetation can also mitigate drought effects by altering soil water distribution. Hence, partitioning of carbon and water fluxes between the soil and vegetation components is crucial to gain mechanistic understanding of vegetation effects on carbon and water cycling. However, the possible impact of herbaceous vegetation in savanna type ecosystems is often overlooked. Therefore, we aimed at quantifying understory vegetation effects on the water balance and productivity of a Mediterranean oak savanna. ET and net ecosystem CO2 exchange (NEE) were partitioned based on flux and stable oxygen isotope measurements and also rain infiltration was estimated. The understory vegetation contributed importantly to total ecosystem ET and GPP with a maximum of 43 and 51%, respectively. It reached water-use efficiencies (WUE; ratio of carbon gain by water loss) similar to cork-oak trees. The understory vegetation inhibited soil evaporation (E) and, although E was large during wet periods, it did not diminish WUE during water-limited times. The understory strongly increased soil water infiltration, specifically following major rain events. At the same time, the understory itself was vulnerable to drought, which led to an earlier senescence of the understory growing under trees as compared to open areas, due to competition for water. Thus, beneficial understory effects are dominant and contribute to the resilience of this ecosystem. At the same time the vulnerability of the understory to drought suggests that future climate change scenarios for the Mediterranean basin threaten understory development. This in turn will very likely diminish beneficial understory effects like infiltration and ground water recharge and therefore ecosystem resilience to drought.

Resolving rapid dynamics of soil–plant–atmosphere interactions

Water is the main driver of ecosystem productivity in most terrestrial ecosystems worldwide. The predicted increase in rainfall variability and extreme climatic events under future climate conditions are therefore anticipated to strongly affect plant and ecosystem functioning. As 55–70% of terrestrial evapotranspirational water loss is directly controlled by plants (Schlesinger & Jasechko, 2014), transpiration comprises the largestwater flux from Earth’s continents, creating a dominant force in the global water cycle. Plant transpiration is controlled by the availability of soil water and root uptake from different soil depths, transport velocities in the xylem, and transpirational water loss through stomates during photosynthesis driven by the atmospheric demand (leaf-to-air water vapour deficit), and the species-specific sensitivity of stomatal control of water loss. In turn, plants modulate the available water resources in ecosystems by modifying interception, rain water infiltration, soil evaporation and groundwater recharge (e.g. Rodriguez-Iturbe, 2000). Different species with diverse rooting systems and depths have different effects on soil water infiltration, as well as different capacities to utilize various soil water pools (Jackson et al., 2000). Since the 1980s, experimental evidence has ascertained the pivotal role of plant roots for soil water redistribution, such as hydraulic lift of deep water sources into shallower and dryer soil layers (Caldwell, 1987), thereby promoting or buffering competition for this highly valuable resource within plant communities. In spite of these insights, quantification of dynamic soil–vegetation feedbacks within the water cycle remains amajor challenge. Progress has partially been hampered by the fact that blue (abiotic, e.g. stream, runoff) and green (biotic, e.g. vegetation water losses) water flows were mostly investigated separately in different disciplines.Given that extreme events such as floods and droughts are predicted to increase in frequency formany regions, dynamic species-specific responses in root water uptake to changing available water pools play a pivotal role in the understanding of the ecosystem water balance and functioning. In this regard, more interdisciplinary approaches, bridging hydrology, ecophysiology and atmospheric science are needed. In this issue of New Phytologist, Volkmann et al. (pp. 839–849) were able to investigate this hidden part of the soil–vegetation interaction. Volkmann et al. quantified rapid dynamics of rainwater infiltration within the rooting zone of mixed andmonoculture tree saplings. The consecutive responses in plant root water uptake and utilization of a rain pulse revealed high species-specificities not in line with the distribution of roots within the soil profile. Furthermore, a novel dimension of temporal and spatial resolution in soil water flow processes was achieved, utilizing newly developed stable isotope-based methods.

Soil water availability and capacity of nitrogen accumulation influence variations of intrinsic water use efficiency in rice

Leaf intrinsic water use efficiency (WUEi) coupling maximum assimilation rate (Amax) and transpirable water lost via stomatal conductance (gsc) has been gaining increasing concern in sustainable crop production. Factors that influence leaf Amax and WUEi in rice (Oryza sativa L. cv Unkang) at flooding and rainfed conditions were evaluated. Positive correlations for leaf nitrogen content (Nm) and maximum carboxylation rate (Vcmax), for nitrogen allocation in Rubisco enzymes and mesophyll conductance (gm) were evident independent of cropping cultures. Rainfed rice exhibited enriched canopy leaf average Nm resulting in higher Amax, partially supporting improved leaf WUEi. Maximum WUEi (up to 0.14 μmol mmol(-1)) recorded in rainfed rice under drought conditions resulted from increasing gm/gsc ratio while at cost of significant decline in Amax due to hydraulically constrained gsc. Amax sensitivity related to gsc which was regulated by plant hydraulic conductance. WUEi was tightly correlated to Vcmax/gsc and gm/gsc ratios across the paddy and rainfed not to light environment, morphological and physiological traits, highlighting enhance capacity of Nm accumulation in rainfed rice with gsc at moderately high level similar to paddy rice facilitate optimization in Amax and WUEi while, is challenged by drought-vulnerable plant hydraulic conductance.

Impact of Leaf Traits on Temporal Dynamics of Transpired Oxygen Isotope Signatures and Its Impact on Atmospheric Vapor

Oxygen isotope signatures of transpiration (δE) are powerful tracers of water movement from plant to global scale. However, a mechanistic understanding of how leaf morphological/physiological traits effect δE is missing. A laser spectrometer was coupled to a leaf-level gas-exchange system to measure fluxes and isotopic signatures of plant transpiration under controlled conditions in seven distinct species (Fagus sylvatica, Pinus sylvestris, Acacia longifolia, Quercus suber, Coffea arabica, Plantago lanceolata, Oxalis triangularis). We analyzed the role of stomatal conductance (gs) and leaf water content (W) on the temporal dynamics of δE following changes in relative humidity (rH). Changes in rH were applied from 60 to 30% and from 30 to 60%, which is probably more than covering the maximum step changes occurring under natural conditions. Further, the impact of gs and W on isotopic non-steady state isofluxes was analyzed. Following changes in rH, temporal development of δE was well described by a one-pool modeling approach for most species. Isofluxes of δE were dominantly driven by stomatal control on E, particularly for the initial period of 30 min following a step change. Hence, the deviation of isofluxes from isotopic steady state can be large, even though plants transpire near to isotopic steady state. Notably, not only transpiration rate and stomatal conductance, but also the leaf traits stomatal density (as a measure of gmax) and leaf water content are significantly related to the time constant (τ) and non-steady-state isofluxes. This might provide an easy-to-access means of a priori assumptions for the impact of isotopic non-steady-state transpiration in various ecosystems. We discuss the implications of our results from leaf to ecosystem scale.

Increased Soil Frost Versus Summer Drought as Drivers of Plant Biomass Responses To Reduced Precipitation: Results from A Globally-Coordinated Field Experiment

Reduced precipitation treatments often are used in field experiments to explore the effects of drought on plant productivity and species composition. However, in seasonally snow-covered regions reduced precipitation also reduces snow cover, which can increase soil frost depth, decrease minimum soil temperatures and increase soil freeze–thaw cycles. Therefore, in addition to the effects of reduced precipitation on plants via drought, freezing damage to overwintering plant tissues at or below the soil surface could further affect plant productivity and relative species abundances during the growing season. We examined the effects of both reduced rainfall (via rain-out shelters) and reduced snow cover (via snow removal) at 13 sites globally (primarily grasslands) within the framework of the International Drought Experiment, a coordinated distributed experiment. Plant cover was estimated at the species level, and aboveground biomass was quantified at the functional group level. Among sites, we observed a negative correlation between the snow removal effect on minimum soil temperature and plant biomass production the next growing season. Three sites exhibited significant rain-out shelter effects on plant productivity, but there was no correlation among sites between the rain-out shelter effect on minimum soil moisture and plant biomass. There was no interaction between snow removal and rain-out shelters for plant biomass, although these two factors only exhibited significant effects simultaneously for a single site. Overall, our results reveal that reduced snowfall, when it decreases minimum soil temperatures, can be an important component of the total effect of reduced precipitation on plant productivity.

Sauna, sweat and science – quantifying the proportion of condensation water versus sweat using a stable water isotope (2H/1H and 18O/16O) tracer experiment

Most visitors of a sauna appreciate the heat pulse that is perceived when water is poured on the stones of a sauna stove. However, probably only few bathers are aware that this pleasant heat pulse is caused by latent heat being released onto our skin due to condensation of water vapour. In order to quantify the proportion of condensation water versus sweat to dripping water of test persons we conducted sauna experiments using isotopically labelled (δ18O and δ2H) thrown water as tracer. This allows differentiating between ‘pure sweat’ and ‘condensation water’. Two ways of isotope mass balance calculations were applied and yielded similar results for both water isotopes. Accordingly, condensation contributed considerably to dripping water with mean proportions of 52 ± 12 and 54 ± 7 % in a sauna experiment in winter semester 2011/12 and 30 ± 13 and 33 ± 6 % in a sauna experiment in winter semester 2012/13, respectively, depending on the way of calculating the isotope mass balance. It can be concluded from the results of our dual isotope labelling sauna experiment that it is not all about sweat in the sauna.

Quantifying differences in water and carbon cycling between paddy and rainfed rice (Oryza sativa L.) by flux partitioning

Agricultural crops play an important role in the global carbon and water cycle. Global climate change scenarios predict enhanced water scarcity and altered precipitation pattern in many parts of the world. Hence, a mechanistic understanding of water fluxes, productivity and water use efficiency of cultivated crops is of major importance, i.e. to adapt management practices. We compared water and carbon fluxes of paddy and rainfed rice by canopy scale gas exchange measurements, crop growth, daily evapotranspiration, transpiration and carbon flux modeling. Throughout a monsoon rice growing season, soil evaporation in paddy rice contributed strongly to evapotranspiration (96.6% to 43.3% from initial growth to fully developed canopy and amounted to 57.9% of total water losses over the growing seasons. Evaporation of rainfed rice was significantly lower (by 65% on average) particularly before canopy closure. Water use efficiency (WUE) was significantly higher in rainfed rice both from an agronomic (WUEagro, i.e. grain yield per evapotranspiration) and ecosystem (WUEeco, i.e. gross primary production per evapotranspiration) perspective. However, our results also show that higher WUE in rainfed rice comes at the expense of higher respiration losses compared to paddy rice (26% higher on average). Hence, suggestions on water management depend on the regional water availability (i.e. Mediterranean vs. Monsoon climate) and the balance between higher respiratory losses versus a potential reduction in CH4 and other greenhouse gas emissions. Our results suggest that a shift from rainfed/unsaturated soil to waterlogged paddy conditions after closure of the rice canopy might be a good compromise towards a sustainable use of water while preserving grain yield, particularly for water-limited production areas.

Water fluxes mediated by vegetation: emerging isotopic insights at the soil and atmosphere interfaces

Plants mediate water fluxes within the soil-vegetation-atmosphere continuum. This water transfer in soils, through plants, into the atmosphere can be effectively traced by stable isotopologues of water. However, rapid dynamic processes have only recently gained attention, such as adaptations in root water uptake depths (within hours to days) or the imprint of transpirational fluxes on atmospheric moisture, particularly promoted by the development of real-time in-situ water vapour stable isotope observation techniques. We focus on open questions and emerging insights at the soil-plant and plant-atmosphere interfaces, as we believe that these are the controlling factors for ecosystem water cycling. At both interfaces, complex pictures of interacting ecophysiological and hydrological processes emerge: root water uptake dynamics depend on both spatiotemporal variations in water availability and species-specific regulation of adaptive root conductivity within the rooting system by, for example, modulating soil-root conductivity in response to water and nutrient demands. Similarly, plant water transport and losses are a fine-tuned interplay between species-specific structural and functional strategies of water use and atmospheric processes. We propose that only by explicitly merging insights from distinct disciplines - for example, hydrology, plant physiology and atmospheric sciences - will we gain a holistic picture of the impact of vegetation on processes governing the soil-plant-atmosphere continuum.

Terpenoid Emissions of Two Mediterranean Woody Species in Response to Drought Stress

Drought is a major environmental constrain affecting plant performance and survival, particularly in Mediterranean ecosystems. Terpenoids may play a protective role under these conditions, however, observations of drought effects on plant terpenoid emissions are controversial ranging from decreased emissions to unaffected or increased release of terpenoids. In the present study we investigated terpenoid emissions of cork oak (Quercus suber) and gum rockrose (Cistus ladanifer) in response to summer drought stress in 2017. Pre-dawn leaf water potential (ΨPD) decreased from -0.64 to -1.72 MPa in Q. suber and from -1.69 to -4.05 MPa in C. ladanifer, indicating a transition from mild to severe drought along summer. Total terpenoid emissions decreased with drought, but differed significantly between species (p < 0.001) and in response to ΨPD, air temperature and assimilation rates. C. ladanifer emitted a large variety of >75 compounds comprising monoterpenes, sesquiterpenes and even diterpenes, which strongly decreased from 1.37 ± 0.23 μg g-1h-1 to 0.40 ± 0.08 μg g-1h-1 (p < 0.001) in response to drought. Total emission rates were positively correlated to air temperature (p < 0.001). C. ladanifer behavior points toward terpenoid leaf storage depletion and reduced substrate availability for terpenoid synthesis with increasing drought, most likely accelerated by high air temperatures. Q. suber emitted mainly monoterpenes and emissions declined significantly from June (0.50 ± 0.08 μg g-1h-1) to August (0.29 ± 0.02 μg g-1h-1) (p < 0.01). Emission rates were weakly correlated with net assimilation rates (R2 = 0.19, p < 0.001), but did not respond strongly to ΨPD and air temperature. Early onset of drought in 2017 most likely reduced plant metabolism in Q. suber, resulting in diminished, but stable terpenoid fluxes. Calculation of standard emission factors (at 30°C) revealed contrasting emission patterns of decreasing, unaffected, or increasing fluxes of single terpenoid compounds. Unaffected or drought-enhanced emissions of compounds such as α-pinene, camphene or manoyl oxide may point toward a specific role of these terpenoids in abiotic stress adaptation. In conclusion, these results suggest a strong negative, but species- and compound-specific effect of severe drought on terpenoid fluxes in Mediterranean ecosystems.