Ansgar Kahmen

Cellulose δ18O is an index of leaf-to-air vapor pressure difference (VPD) in tropical plants

Cellulose in plants contains oxygen that derives in most cases from precipitation. Because the stable oxygen isotope composition, δ18O, of precipitation is associated with environmental conditions, cellulose δ18O should be as well. However, plant physiological models using δ18O suggest that cellulose δ18O is influenced by a complex mix of both climatic and physiological drivers. This influence complicates the interpretation of cellulose δ18O values in a paleo-context. Here, we combined empirical data analyses with mechanistic model simulations to i) quantify the impacts that the primary climatic drivers humidity (ea) and air temperature (Tair) have on cellulose δ18O values in different tropical ecosystems and ii) determine which environmental signal is dominating cellulose δ18O values. Our results revealed that ea and Tair equally influence cellulose δ18O values and that distinguishing which of these factors dominates the δ18O values of cellulose cannot be accomplished in the absence of additional environmental information. However, the individual impacts of ea and Tair on the δ18O values of cellulose can be integrated into a single index of plant-experienced atmospheric vapor demand: the leaf-to-air vapor pressure difference (VPD). We found a robust relationship between VPD and cellulose δ18O values in both empirical and modeled data in all ecosystems that we investigated. Our analysis revealed therefore that δ18O values in plant cellulose can be used as a proxy for VPD in tropical ecosystems. As VPD is an essential variable that determines the biogeochemical dynamics of ecosystems, our study has applications in ecological-, climate-, or forensic-sciences.

Stable isotopes in leaf water of terrestrial plants

Leaf water contains naturally occurring stable isotopes of oxygen and hydrogen in abundances that vary spatially and temporally. When sufficiently understood, these can be harnessed for a wide range of applications. Here, we review the current state of knowledge of stable isotope enrichment of leaf water, and its relevance for isotopic signals incorporated into plant organic matter and atmospheric gases. Models describing evaporative enrichment of leaf water have become increasingly complex over time, reflecting enhanced spatial and temporal resolution. We recommend that practitioners choose a model with a level of complexity suited to their application, and provide guidance. At the same time, there exists some lingering uncertainty about the biophysical processes relevant to patterns of isotopic enrichment in leaf water. An important goal for future research is to link observed variations in isotopic composition to specific anatomical and physiological features of leaves that reflect differences in hydraulic design. New measurement techniques are developing rapidly, enabling determinations of both transpired and leaf water δ(18) O and δ(2) H to be made more easily and at higher temporal resolution than previously possible. We expect these technological advances to spur new developments in our understanding of patterns of stable isotope fractionation in leaf water.

Convergence of soil nitrogen isotopes across global climate gradients

Quantifying global patterns of terrestrial nitrogen (N) cycling is central to predicting future patterns of primary productivity, carbon sequestration, nutrient fluxes to aquatic systems, and climate forcing. With limited direct measures of soil N cycling at the global scale, syntheses of the 15N:14N ratio of soil organic matter across climate gradients provide key insights into understanding global patterns of N cycling. In synthesizing data from over 6000 soil samples, we show strong global relationships among soil N isotopes, mean annual temperature (MAT), mean annual precipitation (MAP), and the concentrations of organic carbon and clay in soil. In both hot ecosystems and dry ecosystems, soil organic matter was more enriched in 15N than in corresponding cold ecosystems or wet ecosystems. Below a MAT of 9.8°C, soil δ15N was invariant with MAT. At the global scale, soil organic C concentrations also declined with increasing MAT and decreasing MAP. After standardizing for variation among mineral soils in soil C and clay concentrations, soil δ15N showed no consistent trends across global climate and latitudinal gradients. Our analyses could place new constraints on interpretations of patterns of ecosystem N cycling and global budgets of gaseous N loss.

No shift to a deeper water uptake depth in response to summer drought of two lowland and sub-alpine C3-grasslands in Switzerland

Temperate C3-grasslands are of high agricultural and ecological importance in Central Europe. Plant growth and consequently grassland yields depend strongly on water supply during the growing season, which is projected to change in the future. We therefore investigated the effect of summer drought on the water uptake of an intensively managed lowland and an extensively managed sub-alpine grassland in Switzerland. Summer drought was simulated by using transparent shelters. Standing above- and belowground biomass was sampled during three growing seasons. Soil and plant xylem waters were analyzed for oxygen (and hydrogen) stable isotope ratios, and the depths of plant water uptake were estimated by two different approaches: (1) linear interpolation method and (2) Bayesian calibrated mixing model. Relative to the control, aboveground biomass was reduced under drought conditions. In contrast to our expectations, lowland grassland plants subjected to summer drought were more likely (43–68xa0%) to rely on water in the topsoil (0–10xa0cm), whereas control plants relied less on the topsoil (4–37xa0%) and shifted to deeper soil layers (20–35xa0cm) during the drought period (29–48xa0%). Sub-alpine grassland plants did not differ significantly in uptake depth between drought and control plots during the drought period. Both approaches yielded similar results and showed that the drought treatment in the two grasslands did not induce a shift to deeper uptake depths, but rather continued or shifted water uptake to even more shallower soil depths. These findings illustrate the importance of shallow soil depths for plant performance under drought conditions.

Effects of leaf water evaporative 2H‐enrichment and biosynthetic fractionation on leaf wax n‐alkane δ2H values in C3 and C4 grasses

Leaf wax n-alkane δ2 H values carry important information about environmental and ecophysiological processes in plants. However, the physiological and biochemical drivers that shape leaf wax n-alkane δ2 H values are not completely understood. It is particularly unclear why n-alkanes in grasses are typically 2 H-depleted compared with plants from other taxonomic groups such as dicotyledonous plants and why C3 grasses are 2 H-depleted compared with C4 grasses. To resolve these uncertainties, we quantified the effects of leaf water evaporative 2 H-enrichment and biosynthetic hydrogen isotope fractionation on n-alkane δ2 H values for a range of C3 and C4 grasses grown in climate-controlled chambers. We found that only a fraction of leaf water evaporative 2 H-enrichment is imprinted on the leaf wax n-alkane δ2 H values in grasses. This is interesting, as previous studies have shown in dicotyledonous plants a nearly complete transfer of this 2 H-enrichment to the n-alkane δ2 H values. We thus infer that the typically observed 2 H-depletion of n-alkanes in grasses (as opposed to dicots) is because only a fraction of the leaf water evaporative 2 H-enrichment is imprinted on the δ2 H values. Our experiments also show that differences in n-alkane δ2 H values between C3 and C4 grasses are largely the result of systematic differences in biosynthetic fractionation between these two plant groups, which was on average -198‰ and-159‰ for C3 and C4 grasses, respectively.

The enigma of effective path length for (18) O enrichment in leaf water of conifers.

The Péclet correction is often used to predict leaf evaporative enrichment and requires an estimate of effective path length (L). Studies to estimate L in conifer needles have produced unexpected patterns based on Péclet theory and leaf anatomy. We exposed seedlings of six conifer species to different vapour pressure deficits (VPD) in controlled climate chambers to produce steady-state leaf water enrichment (in (18) O). We measured leaf gas exchange, stable oxygen isotopic composition (δ(18) O) of input and plant waters as well as leaf anatomical characteristics. Variation in bulk needle water δ(18) O was strongly related to VPD. Conifer needles had large amounts of water within the vascular strand that was potentially unenriched (up to 40%). Both standard Craig-Gordon and Péclet models failed to accurately predict conifer leaf water δ(18) O without taking into consideration the unenriched water in the vascular strand and variable L. Although L was linearly related to mesophyll thickness, large within-species variation prevented the development of generalizations that could allow a broader use of the Péclet effect in predictive models. Our results point to the importance of within needle water pools and isolating mechanisms that need further investigation in order to integrate Péclet corrections with two compartment leaf water concepts.

Tightly bound soil water introduces isotopic memory effects on mobile and extractable soil water pools

ABSTRACT Cryogenic vacuum extraction is the well-established method of extracting water from soil for isotopic analyses of waters moving through the soil–plant–atmosphere continuum. We investigate if soils can alter the isotopic composition of water through isotope memory effects, and determined which mechanisms are responsible for it. Soils with differing physicochemical properties were re-wetted with reference water and subsequently extracted by cryogenic water distillation. Results suggest some reference waters bind tightly to the soil and not all of this tightly bound water is removed during cryogenic vacuum extraction. Kinetic isotopic fractionation occurring when reference water binds to the soil is likely responsible for the 18O-depletion of re-extracted reference water, suggesting an enrichment of the tightly bound soil water pool. Further re-wetting of cryogenically extracted soils indicates an isotopic memory effect of tightly bound soil water on water added to the soil. The data suggest tightly bound soil water can influence the isotopic composition of mobile soil water. Findings show that soils influence the isotope composition of soil water by (i) kinetic fractionation when water is bound to the soil and (ii) equilibrium fractionation between different soil water pools. These findings could be relevant for plant water uptake investigations and complicate ecohydrological and paleohydrological studies.

Cryogenic vacuum artifacts do not affect plant water-uptake studies using stable isotope analysis

Water movement through the soil-plant-atmosphere continuum can be tracked through its hydrogen and oxygen isotopic composition. How the isotopic composition of water evolves as it moves through this continuum provides critical information on ecosystem hydrology and climate change. Cryogenic vacuum extraction is the most applied method to extract waters from soil and plant material for such studies. However, recent experiments where oven dried soils were spiked with a reference water suggest potential for these techniques to bias results. We present results from a greenhouse-based plant water uptake and cryogenic vacuum distillation experiment using Salix viminalis cuttings. We tested the capacity for cryogenic vacuum extraction to recover irrigation water and the plant-available water pool from 4 soils that were subject to continuous irrigation as would occur in nature. Results show that δ2H and δ18O values of extracted soil waters reflect those of irrigation water, but with some influence of evaporation that varied with differences in soil humic and clay content. This suggests that isotope effects observed in laboratory experiments with oven-dried soils may not be relevant under natural conditions. Cryogenic vacuum extraction also recovered xylem water δ2H values that matched the δ2H values of extracted soil water but revealed small, consistent offsets between xylem and soil water δ18O values of 0.84xa0±xa01.13‰. Although these effects may not significantly bias water isotope sourcing applications, they do result in large extrapolation errors when estimating original source water isotope composition from extracted xylem waters using the back extrapolation to the global meteoric water line method.

Low secondary leaf wax n-alkane synthesis on fully mature leaves of C3 grasses grown at controlled environmental conditions and variable humidity.

RATIONALEnLeaf wax n-alkanes are long-chained aliphatic compounds that are present in the cuticle of terrestrial plant leaves. Their δ2 H values are used for the reconstruction of past environments and for plant ecological investigations. The timing of n-alkane synthesis during leaf development and the rate of synthesis of secondary n-alkanes in fully matured leaves are still a matter of debate.nnnMETHODSnUsing a 2 H-labeling approach we estimated secondary leaf wax n-alkane synthesis rates in mature leaf blades of six C3 grass species grown in climate chambers under controlled environmental conditions.nnnRESULTSnWe found that mature grass leaves continue the synthesis of leaf wax n-alkanes after leaf maturation. The rate of secondary n-alkanes synthesis was, however, relatively low and varied in response to atmospheric humidity and among species from 0.09 to 1.09% per day.nnnCONCLUSIONSnOur investigation provides new evidence on the timing of cuticular wax synthesis in grass leaves and indicates that the majority of n-alkanes are synthesized during the initial development of the leaf. Our study will improve the interpretation of leaf wax n-alkane δ2 H values in environmental and geological studies as it suggests that secondary synthesis of leaf wax n-alkanes in grass leaves contributes only slightly to the geological record. Copyright

Oxygen isotope fractionations across individual leaf carbohydrates in grass and tree species

Almost no δ18 O data are available for leaf carbohydrates, leaving a gap in the understanding of the δ18 O relationship between leaf water and cellulose. We measured δ18 O values of bulk leaf water (δ18 OLW ) and individual leaf carbohydrates (e.g. fructose, glucose and sucrose) in grass and tree species and δ18 O of leaf cellulose in grasses. The grasses were grown under two relative humidity (rH) conditions. Sucrose was generally 18 O-enriched compared with hexoses across all species with an apparent biosynthetic fractionation factor (εbio ) of more than 27‰ relative to δ18 OLW , which might be explained by isotopic leaf water and sucrose synthesis gradients. δ18 OLW and δ18 O values of carbohydrates and cellulose in grasses were strongly related, indicating that the leaf water signal in carbohydrates was transferred to cellulose (εbio xa0=xa025.1‰). Interestingly, damping factor pex px , which reflects oxygen isotope exchange with less enriched water during cellulose synthesis, responded to rH conditions if modelled from δ18 OLW but not if modelled directly from δ18 O of individual carbohydrates. We conclude that δ18 OLW is not always a good substitute for δ18 O of synthesis water due to isotopic leaf water gradients. Thus, compound-specific δ18 O analyses of individual carbohydrates are helpful to better constrain (post-)photosynthetic isotope fractionation processes in plants.