Matthias Saurer

Linking stable oxygen and carbon isotopes with stomatal conductance and photosynthetic capacity: a conceptual model

Abstract Based on measurements of δ18O and δ13C in organic matter of C3-plants, we have developed a conceptual model that gives insight into the relationship between stomatal conductance (gl) and photosynthetic capacity (Amax) resulting from differing environmental constraints and plant-internal factors. This is a semi-quantitative approach to describing the long-term effects of environmental factors on CO2 and H2O gas exchange, whereby we estimate the intercellular CO2 concentration (ci) from δ13C and the air humidity from δ18O. Assuming that air humidity is an important factor influencing gl, the model allows us to distinguish whether differences in ci are caused by a response of gl or of Amax. As an application of the model we evaluated the isotope data from three species in plots differing in intensity of land use (hay meadows and abandoned areas) at three sites along a south north transect in the Eastern Alps. We found three different δ18O–δ13C response patterns in native and planted grassland species (cultivated in the greenhouse). After preliminary confirmation by gas-exchange measurements we conclude that the proposed model is a promising tool for deriving carbon water relations in different functional groups from δ18O and δ13C isotope data.

Reducing uncertainties in δ13C analysis of tree rings: Pooling, milling, and cellulose extraction

Recent developments of on-line methods have provided another boost to the determination of stable isotope ratios in organic material. Along with a significant increase in sample throughput, the sample sizes decrease, both of which are necessary conditions to acquire long time series from limited wood amounts. In view of this new technique we reconsidered the most important factors influencing the measured isotopic signature which are (1) pooling, (2) homogeneity, and (3) cellulose extraction. In most cases, pooling (i.e., mixing wood of the same year from different trees) can be made in a simple way by mixing the whole wood available because mass-weighted and unweighted isotope measurements were the same within the error. More attention must be paid in homogenizing the sample. Theoretical considerations underpinned by experimental results suggest a fineness of 0.15 mm (115 mesh) if cellulose is extracted and 0.1 mm (165 mesh) for direct wood analysis. Many of previous studies did not achieve this fineness. We find that wood is as good a climate proxy as cellulose. This is shown by comparing correlations of wood and corresponding cellulose isotope values with meteorological data, which are identical within the uncertainty.

Estimating the uptake of traffic-derived NO2 from 15N abundance in Norway spruce needles

Abstract The 15N ratio of nitrogen oxides (NOx) emitted from vehicles, measured in the air adjacent to a highway in the Swiss Middle Land, was very high [δ15N(NO2) = +5.7‰]. This high 15N abundance was used to estimate long-term NO2 dry deposition into a forest ecosystem by measuring δ15N in the needles and the soil of potted and autochthonous spruce trees [Picea abies (L.) Karst] exposed to NO2 in a transect orthogonal to the highway. δ15N in the current-year needles of potted trees was 2.0‰ higher than that of the control after 4 months of exposure close to the highway, suggesting a 25% contribution to the N-nutrition of these needles. Needle fall into the pots was prevented by grids placed above the soil, while the continuous decomposition of needle litter below the autochthonous trees over previous years has increased δ15N values in the soil, resulting in parallel gradients of δ15N in soil and needles with distance from the highway. Estimates of NO2 uptake into needles obtained from the δ15N data were significantly correlated with the inputs calculated with a shoot gas exchange model based on a parameterisation widely used in deposition modelling. Therefore, we provide an indication of estimated N inputs to forest ecosystems via dry deposition of NO2 at the receptor level under field conditions.

Temporal dynamics of the carbon isotope composition in a Pinus sylvestris stand: from newly assimilated organic carbon to respired carbon dioxide

The 13C isotopic signature (C stable isotope ratio; δ13C) of CO2 respired from forest ecosystems and their particular compartments are known to be influenced by temporal changes in environmental conditions affecting C isotope fractionation during photosynthesis. Whereas most studies have assessed temporal variation in δ13C of ecosystem-respired CO2 on a day-to-day scale, not much information is available on its diel dynamics. We investigated environmental and physiological controls over potential temporal changes in δ13C of respired CO2 by following the short-term dynamics of the 13C signature from newly assimilated organic matter pools in the needles, via phloem-transported organic matter in twigs and trunks, to trunk-, soil- and ecosystem-respired CO2. We found a strong 24-h periodicity in δ13C of organic matter in leaf and twig phloem sap, which was strongly dampened as carbohydrates were transported down the trunk. Periodicity reappeared in the δ13C of trunk-respired CO2, which seemed to originate from apparent respiratory fractionation rather than from changes in δ13C of the organic substrate. The diel patterns of δ13C in soil-respired CO2 are partly explained by soil temperature and moisture and are probably due to changes in the relative contribution of heterotrophic and autotrophic CO2 fluxes to total soil efflux in response to environmental conditions. Our study shows that direct relations between δ13C of recent assimilates and respired CO2 may not be present on a diel time scale, and other factors lead to short-term variations in δ13C of ecosystem-emitted CO2. On the one hand, these variations complicate ecosystem CO2 flux partitioning, but on the other hand they provide new insights into metabolic processes underlying respiratory CO2 emission.

Stable carbon isotopes in tree rings of beech: climatic versus site-related influences

Abstract Stable carbon isotopes in tree rings are a promising tool in palaeoclimate research, provided attempts are made to disentangle climatic from local effects (e.g. soil properties, competition, light). The 13C/12C variations in cellulose of tree rings of beech (Fagus sylvatica) were determined at several sites in the Swiss Central Plateau covering the last 50 years. We chose sites which differ in moisture conditions and sampled cores from four to six trees per site. The mean 13C/12C series from the different dry sites (distant by up to 40 km) are closely interrelated suggesting a common external cause. Correlation analysis with climate data proved the total precipitation in the months May, June and July to have the strongest effect on the carbon isotopes (r =  – 0.73). This result is in agreement with the commonly used model which relates the isotope discrimination to the water use efficiency. On the other hand, the isotope series of the wet sites are not as well correlated to the climate. At two of the sites (a dry and a humid) tree ring width suddenly increased. We used this effect as a test-case to study the influence of local growth conditions on the climate-isotope relationship.

Seasonal growth, δ13C in leaves and stem, and phloem structure of birch (Betula pendula) under low ozone concentrations

SummaryThe growth of potted birch cuttings (one clone of Betula pendula) was studied under low O3 concentrations (0, 0.050, 0.075, 0.100 μl l-1) throughout an entire growing season. With increasing O3 dose, 20–50% of all leaves formed were prematurely shed, while 40–70% of the remaining foliage displayed advanced discoloration by the end of the season. Ozonation affected the S, P and N concentration of leaves and increased δ13C in leaves and stem, while the CO2 assimilation rate declined with increasing CO2 concentration in mesophyll intercellulars. While whole-plant production correlated negatively with the O3 dose, ozone increased the specific leaf weight (i.e. leaf weight/leaf area, SLW) but decreased the ratios of stem weight/stem length and root/shoot biomass. Neither the latter ratio nor SLW changed in experimentally defoliated control plants, whereas in ozonated plants starch accumulated along leaf veins and phloem tissue was deformed in the leaf petioles and the stem. Only in early summer was the relative growth rate higher in the ozonated than in the control plants. The ratio of whole-plant biomass production versus total foliage area formed was lowered under O3 stress. However, when relating biomass to the actual foliage area present due to leaf loss, this ratio did not differ between treatments. Similarly the ratio of actual foliage area versus basal stem area in cross-section did not differ. Overall, whole-plant production was strongly determined by O3-caused changes in crown structure and began to be limited at O3 doses (approximately 180 μl l-1 h) similar to those of rural sites in Central Europe.

Elevated CO2 increases tree‐level intrinsic water use efficiency: insights from carbon and oxygen isotope analyses in tree rings across three forest FACE sites

Elevated CO₂ increases intrinsic water use efficiency (WUE(i) ) of forests, but the magnitude of this effect and its interaction with climate is still poorly understood. We combined tree ring analysis with isotope measurements at three Free Air CO₂ Enrichment (FACE, POP-EUROFACE, in Italy; Duke FACE in North Carolina and ORNL in Tennessee, USA) sites, to cover the entire life of the trees. We used δ¹³C to assess carbon isotope discrimination and changes in water-use efficiency, while direct CO₂ effects on stomatal conductance were explored using δ¹⁸O as a proxy. Across all the sites, elevated CO₂ increased ¹³C-derived water-use efficiency on average by 73% for Liquidambar styraciflua, 77% for Pinus taeda and 75% for Populus sp., but through different ecophysiological mechanisms. Our findings provide a robust means of predicting water-use efficiency responses from a variety of tree species exposed to variable environmental conditions over time, and species-specific relationships that can help modelling elevated CO₂ and climate impacts on forest productivity, carbon and water balances.

Drought response of five conifer species under contrasting water availability suggests high vulnerability of Norway spruce and European larch

The ability of tree species to cope with anticipated decrease in water availability is still poorly understood. We evaluated the potential of Norway spruce, Scots pine, European larch, black pine, and Douglas-fir to withstand drought in a drier future climate by analyzing their past growth and physiological responses at a xeric and a mesic site in Central Europe using dendroecological methods. Earlywood, latewood, and total ring width, as well as the δ(13) C and δ(18) O in early- and latewood were measured and statistically related to a multiscalar soil water deficit index from 1961 to 2009. At the xeric site, δ(13) C values of all species were strongly linked to water deficits that lasted longer than 11 months, indicating a long-term cumulative effect on the carbon pool. Trees at the xeric site were particularly sensitive to soil water recharge in the preceding autumn and early spring. The native species European larch and Norway spruce, growing close to their dry distribution limit at the xeric site, were found to be the most vulnerable species to soil water deficits. At the mesic site, summer water availability was critical for all species, whereas water availability prior to the growing season was less important. Trees at the mesic were more vulnerable to water deficits of shorter duration than the xeric site. We conclude that if summers become drier, trees growing on mesic sites will undergo significant growth reductions, whereas at their dry distribution limit in the Alps, tree growth of the highly sensitive spruce and larch may collapse, likely inducing dieback and compromising the provision of ecosystem services. However, the magnitude of these changes will be mediated strongly by soil water recharge in winter and thus water availability at the beginning of the growing season.

The relationship between the stable carbon isotope composition of needle bulk material, starch, and tree rings in Picea abies

We investigated the relationship between the δ13C signal in current-year and 1-year-old needle bulk material, starch extracts, and early- or late-wood in mature spruce trees (Picea abies) to identify the modifying influence of climatic conditions on the different δ13C signals. Seasonal patterns of δ13C were determined in total bulk needle material from 1998 to 2000, and in acid soluble starch extracts in 1999 and 2000, and δ13C values of early- and late-wood were measured for the years 1991–2000. δ13C of bulk needle material was most enriched in spring with a trend towards depletion in the course of the season. Current-year needles showed a more distinct seasonal pattern in δ13C compared to 1-year-old needles. Seasonal trends in bulk material and starch were similar, but the highly enriched signal in spring could not be fully explained by the influence of the δ13C values of starch (weighted with the corresponding starch amounts). δ13C of starch in 1-year-old needles, and to a lesser extent of current-year needles, correlated with δ13C of early-wood, indicating a transfer of the isotopic signal. In addition, early-wood δ13C corresponded weakly to winter precipitation. In the summer, δ13C of total bulk needle material and starch showed no relation to the late-wood δ13C signature. Late-wood δ13C, however, related to global radiation, relative humidity and temperature, with more enriched values corresponding to warmer and drier conditions. We conclude that the signature of early-wood is determined more by biochemical fractionation, e.g. during starch formation, than by climatic conditions, which exert only a minor influence and are reflected in the isotopic signal of late-wood.

Spatial variability and temporal trends in water‐use efficiency of European forests

The increasing carbon dioxide (CO2 ) concentration in the atmosphere in combination with climatic changes throughout the last century are likely to have had a profound effect on the physiology of trees: altering the carbon and water fluxes passing through the stomatal pores. However, the magnitude and spatial patterns of such changes in natural forests remain highly uncertain. Here, stable carbon isotope ratios from a network of 35 tree-ring sites located across Europe are investigated to determine the intrinsic water-use efficiency (iWUE), the ratio of photosynthesis to stomatal conductance from 1901 to 2000. The results were compared with simulations of a dynamic vegetation model (LPX-Bern 1.0) that integrates numerous ecosystem and land-atmosphere exchange processes in a theoretical framework. The spatial pattern of tree-ring derived iWUE of the investigated coniferous and deciduous species and the model results agreed significantly with a clear south-to-north gradient, as well as a general increase in iWUE over the 20th century. The magnitude of the iWUE increase was not spatially uniform, with the strongest increase observed and modelled for temperate forests in Central Europe, a region where summer soil-water availability decreased over the last century. We were able to demonstrate that the combined effects of increasing CO2 and climate change leading to soil drying have resulted in an accelerated increase in iWUE. These findings will help to reduce uncertainties in the land surface schemes of global climate models, where vegetation-climate feedbacks are currently still poorly constrained by observational data.