Michael Bahn

Climate extremes and the carbon cycle

The terrestrial biosphere is a key component of the global carbon cycle and its carbon balance is strongly influenced by climate. Continuing environmental changes are thought to increase global terrestrial carbon uptake. But evidence is mounting that climate extremes such as droughts or storms can lead to a decrease in regional ecosystem carbon stocks and therefore have the potential to negate an expected increase in terrestrial carbon uptake. Here we explore the mechanisms and impacts of climate extremes on the terrestrial carbon cycle, and propose a pathway to improve our understanding of present and future impacts of climate extremes on the terrestrial carbon budget.

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.

Does photosynthesis affect grassland soil‐respired CO2 and its carbon isotope composition on a diurnal timescale?

Soil respiration is the largest flux of carbon (C) from terrestrial ecosystems to the atmosphere. Here, we tested the hypothesis that photosynthesis affects the diurnal pattern of grassland soil-respired CO(2) and its C isotope composition (delta(13)C(SR)). A combined shading and pulse-labelling experiment was carried out in a mountain grassland. delta(13)C(SR) was monitored at a high time resolution with a tunable diode laser absorption spectrometer. In unlabelled plots a diurnal pattern of delta(13)C(SR) was observed, which was not explained by soil temperature, moisture or flux rates and contained a component that was also independent of assimilate supply. In labelled plots delta(13)C(SR) reflected a rapid transfer and respiratory use of freshly plant-assimilated C and a diurnal shift in the predominant respiratory C source from recent (i.e. at least 1 d old) to fresh (i.e. photoassimilates produced on the same day). We conclude that in grasslands the plant-derived substrates used for soil respiratory processes vary during the day, and that photosynthesis provides an important and immediate C source. These findings indicate a tight coupling in the plant-soil system and the importance of plant metabolism for soil CO(2) fluxes.

Effects of climate extremes on the terrestrial carbon cycle: concepts, processes and potential future impacts

Extreme droughts, heat waves, frosts, precipitation, wind storms and other climate extremes may impact the structure, composition and functioning of terrestrial ecosystems, and thus carbon cycling and its feedbacks to the climate system. Yet, the interconnected avenues through which climate extremes drive ecological and physiological processes and alter the carbon balance are poorly understood. Here, we review the literature on carbon cycle relevant responses of ecosystems to extreme climatic events. Given that impacts of climate extremes are considered disturbances, we assume the respective general disturbance-induced mechanisms and processes to also operate in an extreme context. The paucity of well-defined studies currently renders a quantitative meta-analysis impossible, but permits us to develop a deductive framework for identifying the main mechanisms (and coupling thereof) through which climate extremes may act on the carbon cycle. We find that ecosystem responses can exceed the duration of the climate impacts via lagged effects on the carbon cycle. The expected regional impacts of future climate extremes will depend on changes in the probability and severity of their occurrence, on the compound effects and timing of different climate extremes, and on the vulnerability of each land-cover type modulated by management. Although processes and sensitivities differ among biomes, based on expert opinion, we expect forests to exhibit the largest net effect of extremes due to their large carbon pools and fluxes, potentially large indirect and lagged impacts, and long recovery time to regain previous stocks. At the global scale, we presume that droughts have the strongest and most widespread effects on terrestrial carbon cycling. Comparing impacts of climate extremes identified via remote sensing vs. ground-based observational case studies reveals that many regions in the (sub-)tropics are understudied. Hence, regional investigations are needed to allow a global upscaling of the impacts of climate extremes on global carbon–climate feedbacks.

Seasonal and inter-annual variability of the net ecosystem CO2 exchange of a temperate mountain grassland : Effects of weather and management

The role and relative importance of climate and cutting for the seasonal and inter-annual variability of the net ecosystem CO2 (NEE) of a temperate mountain grassland was investigated. Eddy covariance CO2 flux data and associated measurements of the green area index and the major environmental driving forces acquired during 2001-2006 at the study site Neustift (Austria) were analyzed. Driven by three cutting events per year which kept the investigated grassland in a stage of vigorous growth, the seasonal variability of NEE was primarily modulated by gross primary productivity (GPP). The role of environmental parameters in modulating the seasonal variability of NEE was obscured by the strong response of GPP to changes in the amount of green area, as well as the cutting-mediated decoupling of phenological development and the seasonal course of climate drivers. None of the climate and management metrics examined was able to explain the inter-annual variability of annual NEE. This is thought to result from (1) a high covariance between GPP and ecosystem respiration (Reco) at the annual time scale which results in a comparatively small inter-annual variation of NEE, (2) compensating effects between carbon exchange during and outside the management period, and (3) changes in the biotic response to rather than the climate variables per se. GPP was more important in modulating inter-annual variations in NEE in spring and before the first and second cut, while Reco explained a larger fraction of the inter-annual variability of NEE during the remaining, in particular the post-cut, periods.

Pulse-labelling trees to study carbon allocation dynamics: a review of methods, current knowledge and future prospects

Pulse-labelling of trees with stable or radioactive carbon (C) isotopes offers the unique opportunity to trace the fate of labelled CO(2) into the tree and its release to the soil and the atmosphere. Thus, pulse-labelling enables the quantification of C partitioning in forests and the assessment of the role of partitioning in tree growth, resource acquisition and C sequestration. However, this is associated with challenges as regards the choice of a tracer, the methods of tracing labelled C in tree and soil compartments and the quantitative analysis of C dynamics. Based on data from 47 studies, the rate of transfer differs between broadleaved and coniferous species and decreases as temperature and soil water content decrease. Labelled C is rapidly transferred belowground-within a few days or less-and this transfer is slowed down by drought. Half-lives of labelled C in phloem sap (transfer pool) and in mature leaves (source organs) are short, while those of sink organs (growing tissues, seasonal storage) are longer. (13)C measurements in respiratory efflux at high temporal resolution provide the best estimate of the mean residence times of C in respiratory substrate pools, and the best basis for compartmental modelling. Seasonal C dynamics and allocation patterns indicate that sink strength variations are important drivers for C fluxes. We propose a conceptual model for temperate and boreal trees, which considers the use of recently assimilated C versus stored C. We recommend best practices for designing and analysing pulse-labelling experiments, and identify several topics which we consider of prime importance for future research on C allocation in trees: (i) whole-tree C source-sink relations, (ii) C allocation to secondary metabolism, (iii) responses to environmental change, (iv) effects of seasonality versus phenology in and across biomes, and (v) carbon-nitrogen interactions. Substantial progress is expected from emerging technologies, but the largest challenge remains to carry out in situ whole-tree labelling experiments on mature trees to improve our understanding of the environmental and physiological controls on C allocation.

Relative contributions of plant traits and soil microbial properties to mountain grassland ecosystem services.

Summary 1. Plant functional diversity and soil microbial community composition are tightly coupled. Changes in these interactions may influence ecosystem functioning. Links between plant functional diversity, soil microbial communities and ecosystem functioning have been demonstrated in experiments using plant monocultures and mixtures, using broad plant and microbial functional groups, but have not been examined in diverse natural plant communities. 2. We quantified the relative effects of plant and microbial functional properties on key ecosystem functions. We measured plant functional diversity, soil microbial community composition and parameters associated with nitrogen (N) cycling and key nutrient cycling processes at three grassland sites in different parts of Europe. Because plant structure and function strongly influence soil microbial communities, we determined relationships between ecosystem properties, plant traits and soil community characteristics following a sequential approach in which plant traits were fitted first, followed by the additional effects of soil micro-organisms. 3. We identified a continuum from standing green biomass and standing litter, linked mostly with plant traits, to potential N mineralization and potential leaching of soil inorganic N, linked mostly with microbial properties. Plant and microbial functional parameters were equally important in explaining % organic matter content in soil. A parallel continuum ran from plant height, linked with above-ground biomass, to plant quality effects captured by the leaf economics spectrum, which were linked with the recycling of carbon (C) and N. 4. More exploitative species (higher specific leaf area, leaf N concentrations and lower leaf dry matter content) and taller swards, along with soil microbial communities dominated by bacteria, with rapid microbial activities, were linked with greater fodder production, but poor C and N retention. Conversely, dominance by conservative species (with opposite traits) and soil microbial communities dominated by fungi, and bacteria with slow activities, were usually linked with low production, but greater soil C storage and N retention. 5. Synthesis – Grassland production, C sequestration and soil N retention are jointly related to plant and microbial functional traits. Managing grasslands for selected, or multiple, ecosystem services will thus require a consideration of the joint effects of plant and soil communities. Further understanding of the mechanisms that link plant and microbial functional traits is essential to achieve this.

Redefinition and global estimation of basal ecosystem respiration rate

Basal ecosystem respiration rate (BR), the ecosystem respiration rate at a given temperature, is a common and important parameter in empirical models for quantifying ecosystem respiration (ER) globally. Numerous studies have indicated that BR varies in space. However, many empirical ER models still use a global constant BR largely due to the lack of a functional description for BR. In this study, we redefined BR to be ecosystem respiration rate at the mean annual temperature. To test the validity of this concept, we conducted a synthesis analysis using 276 site-years of eddy covariance data, from 79 research sites located at latitudes ranging from similar to 3 degrees S to similar to 70 degrees N. Results showed that mean annual ER rate closely matches ER rate at mean annual temperature. Incorporation of site-specific BR into global ER model substantially improved simulated ER compared to an invariant BR at all sites. These results confirm that ER at the mean annual temperature can be considered as BR in empirical models. A strong correlation was found between the mean annual ER and mean annual gross primary production (GPP). Consequently, GPP, which is typically more accurately modeled, can be used to estimate BR. A light use efficiency GPP model (i.e., EC-LUE) was applied to estimate global GPP, BR and ER with input data from MERRA (Modern Era Retrospective-Analysis for Research and Applications) and MODIS (Moderate resolution Imaging Spectroradiometer). The global ER was 103 Pg C yr (-1), with the highest respiration rate over tropical forests and the lowest value in dry and high-latitude areas.

Linking plant and ecosystem functional biogeography

Significance This article defines ecosystem functional properties, which can be derived from long-term observations of gas and energy exchange between ecosystems and the atmosphere, and shows that variations of those cannot be easily explained by classical climatological or biogeographical approaches such as plant functional types. Instead, we argue that plant traits have the potential to explain this variation, and we call for a stronger integration of research communities dedicated to plant traits and to ecosystem–atmosphere exchange. Classical biogeographical observations suggest that ecosystems are strongly shaped by climatic constraints in terms of their structure and function. On the other hand, vegetation function feeds back on the climate system via biosphere–atmosphere exchange of matter and energy. Ecosystem-level observations of this exchange reveal very large functional biogeographical variation of climate-relevant ecosystem functional properties related to carbon and water cycles. This variation is explained insufficiently by climate control and a classical plant functional type classification approach. For example, correlations between seasonal carbon-use efficiency and climate or environmental variables remain below 0.6, leaving almost 70% of variance unexplained. We suggest that a substantial part of this unexplained variation of ecosystem functional properties is related to variations in plant and microbial traits. Therefore, to progress with global functional biogeography, we should seek to understand the link between organismic traits and flux-derived ecosystem properties at ecosystem observation sites and the spatial variation of vegetation traits given geoecological covariates. This understanding can be fostered by synergistic use of both data-driven and theory-driven ecological as well as biophysical approaches.

Looking deeper into the soil: biophysical controls and seasonal lags of soil CO2 production and efflux.

We seek to understand how biophysical factors such as soil temperature (Ts), soil moisture (theta), and gross primary production (GPP) influence CO2 fluxes across terrestrial ecosystems. Recent advancements in automated measurements and remote-sensing approaches have provided time series in which lags and relationships among variables can be explored. The purpose of this study is to present new applications of continuous measurements of soil CO2 efflux (F0) and soil CO2 concentrations measurements. Here we explore how variation in Ts, theta, and GPP (derived from NASAs moderate-resolution imaging spectroradiometer [MODIS]) influence F0 and soil CO2 production (Ps). We focused on seasonal variation and used continuous measurements at a daily timescale across four vegetation types at 13 study sites to quantify: (1) differences in seasonal lags between soil CO2 fluxes and Ts, theta, and GPP and (2) interactions and relationships between CO2 fluxes with Ts, theta, and GPP. Mean annual Ts did not explain annual F0 and Ps among vegetation types, but GPP explained 73% and 30% of the variation, respectively. We found evidence that lags between soil CO2 fluxes and Ts or GPP provide insights into the role of plant phenology and information relevant about possible timing of controls of autotrophic and heterotrophic processes. The influences of biophysical factors that regulate daily F0 and Ps are different among vegetation types, but GPP is a dominant variable for explaining soil CO2 fluxes. The emergence of long-term automated soil CO2 flux measurement networks provides a unique opportunity for extended investigations into F0 and Ps processes in the near future.