Temperature‐sensitive biochemical 18^{18}O‐fractionation and humidity‐dependent attenuation factor are needed to predict δ 18^{18}O of cellulose from leaf water in a grassland ecosystem

We explore here our mechanistic understanding of the environmental and physiological processes that determine the oxygen isotope composition of leaf cellulose (δ$^{18}$O$_{cellulose}$) in a drought‐prone, temperate grassland ecosystem. A new allocation‐and‐growth model was designed and added to an $^{18}$O‐enabled soil–vegetation–atmosphere transfer model (MuSICA) to predict seasonal (April–October) and multi‐annual (2007–2012) variation of δ$^{18}$O$_{cellulose}$ and $^{18}$O‐enrichment of leaf cellulose (Δ$^{18}$O$_{cellulose}$) based on the Barbour–Farquhar model. Modelled δ$^{18}$O$_{cellulose}$ agreed best with observations when integrated over c. 400 growing‐degree‐days, similar to the average leaf lifespan observed at the site. Over the integration time, air temperature ranged from 7 to 22°C and midday relative humidity from 47 to 73%. Model agreement with observations of δ$^{18}$O$_{cellulose}$ (R$^{2}$ = 0.57) and Δ$^{18}$O$_{cellulose}$ (R$^{2}$ = 0.74), and their negative relationship with canopy conductance, was improved significantly when both the biochemical $^{18}$O‐fractionation between water and substrate for cellulose synthesis (ε$_{bio}$, range 26–30‰) was temperature‐sensitive, as previously reported for aquatic plants and heterotrophically grown wheat seedlings, and the proportion of oxygen in cellulose reflecting leaf water $^{18}$O‐enrichment (1 – p$_{ex}$p$_{x}$, range 0.23–0.63) was dependent on air relative humidity, as observed in independent controlled experiments with grasses. Understanding physiological information in δ$^{18}$O$_{cellulose}$ requires quantitative knowledge of climatic effects on p$_{ex}$p$_{x}$ and ε$_{bio}$

The 18O ecohydrology of a grassland ecosystem - predictions and observations

This research has been supported by the Deutsche Forschungsgemeinschaft (grant no. SCHN 557/9-1), the Agence Nationale de la Recherche (grant no. ANR-13-BS06-0005), and the European Commission (grant no. SOLCA 338264). This work was supported by the German Research Foundation (DFG) and the Technical University of Munich (TUM) in the framework of the Open Access Publishing Program.Peer reviewedPublisher PD

Do 2H and 18O in leaf water reflect environmental drivers differently?

We compiled hydrogen and oxygen stable isotope compositions (δ H and δ O) of leaf water from multiple biomes to examine variations with environmental drivers. Leaf water δ H was more closely correlated with δ H of xylem water or atmospheric vapour, whereas leaf water δ O was more closely correlated with air relative humidity. This resulted from the larger proportional range for δ H of meteoric waters relative to the extent of leaf water evaporative enrichment compared with δ O. We next expressed leaf water as isotopic enrichment above xylem water (Δ H and Δ O) to remove the impact of xylem water isotopic variation. For Δ H, leaf water still correlated with atmospheric vapour, whereas Δ O showed no such correlation. This was explained by covariance between air relative humidity and the Δ O of atmospheric vapour. This is consistent with a previously observed diurnal correlation between air relative humidity and the deuterium excess of atmospheric vapour across a range of ecosystems. We conclude that H and O in leaf water do indeed reflect the balance of environmental drivers differently; our results have implications for understanding isotopic effects associated with water cycling in terrestrial ecosystems and for inferring environmental change from isotopic biomarkers that act as proxies for leaf water

18 O enrichment of sucrose and photosynthetic and nonphotosynthetic leaf water in a C 3 grass—atmospheric drivers and physiological relations

The 18O enrichment (Δ18O) of leaf water affects the Δ18O of photosynthetic products such as sucrose, generating an isotopic archive of plant function and past climate. However, uncertainty remains as to whether leaf water compartmentation between photosynthetic and nonphotosynthetic tissue affects the relationship between Δ18O of bulk leaf water (Δ18OLW) and leaf sucrose (Δ18OSucrose). We grew Lolium perenne (a C3 grass) in mesocosm-scale, replicated experiments with daytime relative humidity (50% or 75%) and CO2 level (200, 400 or 800 μmol mol−1) as factors, and determined Δ18OLW, Δ18OSucrose and morphophysiological leaf parameters, including transpiration (Eleaf), stomatal conductance (gs) and mesophyll conductance to CO2 (gm). The Δ18O of photosynthetic medium water (Δ18OSSW) was estimated from Δ18OSucrose and the equilibrium fractionation between water and carbonyl groups (εbio). Δ18OSSW was well predicted by theoretical estimates of leaf water at the evaporative site (Δ18Oe) with adjustments that correlated with gas exchange parameters (gs or total conductance to CO2). Isotopic mass balance and published work indicated that nonphotosynthetic tissue water was a large fraction (~0.53) of bulk leaf water. Δ18OLW was a poor proxy for Δ18OSucrose, mainly due to opposite Δ18O responses of nonphotosynthetic tissue water (Δ18Onon-SSW) relative to Δ18OSSW, driven by atmospheric conditions

Do 2H and 18O in leaf water reflect environmental drivers differently?

We compiled hydrogen and oxygen stable isotope compositions (δ2 H and δ18 O) of leaf water from multiple biomes to examine variations with environmental drivers. Leaf water δ2 H was more closely correlated with δ2 H of xylem water or atmospheric vapour, whereas leaf water δ18 O was more closely correlated with air relative humidity. This resulted from the larger proportional range for δ2 H of meteoric waters relative to the extent of leaf water evaporative enrichment compared with δ18 O. We next expressed leaf water as isotopic enrichment above xylem water (Δ2 H and Δ18 O) to remove the impact of xylem water isotopic variation. For Δ2 H, leaf water still correlated with atmospheric vapour, whereas Δ18 O showed no such correlation. This was explained by covariance between air relative humidity and the Δ18 O of atmospheric vapour. This is consistent with a previously observed diurnal correlation between air relative humidity and the deuterium excess of atmospheric vapour across a range of ecosystems. We conclude that 2 H and 18 O in leaf water do indeed reflect the balance of environmental drivers differently; our results have implications for understanding isotopic effects associated with water cycling in terrestrial ecosystems and for inferring environmental change from isotopic biomarkers that act as proxies for leaf water