Rüdiger Grote

The long way down—are carbon and oxygen isotope signals in the tree ring uncoupled from canopy physiological processes?

The carbon (δ(13)C) and oxygen (δ(18)O) stable isotope composition is widely used to obtain information on the linkages between environmental drivers and tree physiology over various time scales. The tree-ring archive can especially be exploited to reconstruct inter- and intra-annual variation of both climate and physiology. There is, however, a lack of information on the processes potentially affecting δ(13)C and δ(18)O on their way from assimilation in the leaf to the tree ring. As a consequence, the aim of this study was to trace the isotope signals in European beech (Fagus sylvatica L.) from leaf water (δ(18)O) and leaf assimilates (δ(13)C and δ(18)O) to tree-ring wood via phloem-transported compounds over a whole growing season. Phloem and leaf samples for δ(13)C and δ(18)O analyses as well as soil water, xylem water, leaf water and atmospheric water vapour samples for δ(18)O analysis were taken approximately every 2 weeks during the growing season of 2007. The δ(13)C and δ(18)O samples from the tree rings were dated intra-annually by monitoring the tree growth with dendrometers. δ(18)O in the phloem organic matter and tree-ring whole wood was not positively related to leaf water evaporative enrichment and δ(18)O of canopy organic matter pools. This finding implies a partial uncoupling of the tree-ring oxygen isotopic signal from canopy physiology. At the same time, internal carbon storage and remobilization physiology most likely prevented δ(13)C in tree-ring whole wood from being closely related to intra-annual variation in environmental drivers. Taking into account the post-photosynthetic isotope fractionation processes resulting in alterations of δ(13)C and δ(18)O not only in the tree ring but also in phloem carbohydrates, as well as the intra-annual timing of changes in the tree internal physiology, might help to better understand the meaning of the tree-ring isotope signal not only intra- but also inter-annually.

Modeling the isoprene emission rate from leaves

The leaves of many plants emit isoprene (2-methyl-1,3-butadiene) to the atmosphere, a process which has important ramifications for global and regional atmospheric chemistry. Quantitation of leaf isoprene emission and its response to environmental variation are described by empirically derived equations that replicate observed patterns, but have been linked only in some cases to known biochemical and physiological processes. Furthermore, models have been proposed from several independent laboratories, providing multiple approaches for prediction of emissions, but with little detail provided as to how contrasting models are related. In this review we provide an analysis as to how the most commonly used models have been validated, or not, on the basis of known biochemical and physiological processes. We also discuss the multiple approaches that have been used for modeling isoprene emission rate with an emphasis on identifying commonalities and contrasts among models, we correct some mathematical errors that have been propagated through the models, and we note previously unrecognized covariances within processes of the models. We come to the conclusion that the state of isoprene emission modeling remains highly empirical. Where possible, we identify gaps in our knowledge that have prevented us from achieving a greater mechanistic foundation for the models, and we discuss the insight and data that must be gained to fill those gaps.

LandscapeDNDC: a process model for simulation of biosphere–atmosphere–hydrosphere exchange processes at site and regional scale

We present a new model system, which facilitates scaling of ecosystem processes from the site to regional simulation domains. The new framework LandscapeDNDC—partly based on the biogeochemical site scale model DNDC—inherits a series of new features with regard to process descriptions, model structure and data I/O functionality. LandscapeDNDC incorporates different vegetation types and management systems for simulating carbon, nitrogen and water related biosphere–atmosphere–hydrosphere fluxes in forest, arable and grassland ecosystems and allows the dynamic simulation of land use changes. The modeling concept divides ecosystems into six substates (canopy air chemistry, microclimate, physiology, water cycle, vegetation structure, and soil biogeochemistry) and provides alternative modules dealing with these substates. The model can be applied on the site scale, as well as for three-dimensional regional simulations. For regional applications LandscapeDNDC integrates all grid cells synchronously forward in time. This allows easy coupling to other spatially distributed models (e.g. for hydrology or atmospheric chemistry) and efficient two-way exchange of states. This paper describes the fundamental design concept of the model and its object-oriented software implementation. Two example applications are presented. First, calculation of a nitrous oxide emission inventory from agricultural soils for the State of Saxaony (Germany), including data preprocessing of the regional model input data. The computational effort for the LandscapeDNDC preprocessing and simulation could be speed up by a factor of almost 100 compared to the approach using the original DNDC version 9.3. Calculated N2O emissions for Saxony with LandscapeDNDC (2693 t N2O–N/a) were compared with the original DNDC model (2725 t N2O–N/a), the IPCC Tier I methodology (1107 t N2O–N/a), and the German National Inventory Report (equal to IPCC Tier II, 2100 t N2O–N/a). The second example illustrates the capabilities of LandscapeDNDC for building a fully coupled three-dimensional model system on the landscape scale. Therefore we coupled the biogeochemical and plant growth calculations to a hydrological transport model and demonstrate the transport of nitrogen along a virtual hillslope and associated formation of indirect nitrous oxide emissions.

Leaf-Level Models of Constitutive and Stress-Driven Volatile Organic Compound Emissions

This chapter provides a review of past and contemporary leaf-level emission algorithms that have been and currently are in use for modelling the emissions of biogenic volatile organic compounds (BVOCs) from plants. The chapter starts with a brief overview about historical efforts and elaborates on processes that describe the direct emission responses to environmental factors such as temperature and light. These phenomenological descriptions have been widely and successfully used in emission models at scales ranging from the leaf to the globe. However, while the models provide tractable mathematical functions that link environmental drivers and emission rates, and as such can be easily incorporated in higher scale predictive models, they do not provide the mechanistic context required to describe interactions among drivers and indirect influences on interactions such as those due to acclimation, accumulated stress and ontogeny. Following a discussion of these issues and the limitations they impose on the current state of model-based prognoses of BVOC emissions, we describe in some detail the knowledge gaps that need to be filled in order to move BVOC emission models into forms that are more directly coupled to physiological processes.

Competitive strategies in adult beech and spruce: space-related foliar carbon investment versus carbon gain

In Central Europe, Fagus sylvatica and Picea abies represent contrasting extremes in foliage type, crown structure and length of growing season. In order to examine the competitive strategies of these two co-occurring species, we tested the following hypotheses: (1) the space occupied by the foliage of sun branches is characterized by greater foliar mass investment compared to shade branches, (2) the carbon (C) gain per unit of occupied space is greater in sun than in shade branches, and (3) annual C and water costs of the foliage for sustaining the occupied space are low, wherever C gain per unit of occupied space is low. These were investigated in a mature forest in Southern Germany. The examination was based on the annual assessment of space-related resource investments and gains of the foliage. The foliated space around branches was regarded as the relevant volume with respect to aboveground resource availability. Occupied crown space per standing foliage mass was higher in shade compared to sun branches of beech, whereas no difference existed in crown volume per foliage mass between sun and shade branches of spruce (hypothesis 1 accepted for beech but rejected for spruce). However, beech occupied more space per foliage mass than spruce. The C gain per occupied crown volume was greater in sun than in shade branches (hypothesis 2 accepted) but did not differ between species. The amount of occupied space per respiratory and transpiratory costs did not differ between species or between sun and shade branches. In beech and spruce, the proportion of foliage investment in the annual C balance of sun and shade branches remained rather stable, whereas respiratory costs distinctly increased in shade foliage. Hence, shade branches were costly structures to occupy space, achieving only low and even negative C balances (rejection of hypothesis 3), which conflicts with the claimed C autonomy of branches. Our findings suggest that competitiveness is determined by the standing foliage mass and the annual branch volume increment rather than annual investments in foliage. Expressing competitiveness in terms of space-related resource investments versus returns, as demonstrated here, has the potential of promoting mechanistic understanding of plant–plant interactions.

Modelling the drought impact on monoterpene fluxes from an evergreen Mediterranean forest canopy

In many ecosystems drought cycles are common during the growing season but their impact on volatile monoterpene emissions is unclear. Therefore, we aimed to develop and evaluate a process-based modelling approach to explore the explanatory power of likely mechanisms. The biochemically based isoprene and monoterpene emission model SIM-BIM2 has been modified and linked to a canopy model and a soil water balance model. Simulations are carried out for Quercus ilex forest sites and results are compared to measured soil water, photosynthesis, terpene-synthase activity, and monoterpene emission rates. Finally, the coupled model system is used to estimate the annual drought impact on photosynthesis and emission. The combined and adjusted vegetation model was able to simulate photosynthesis and monoterpene emission under dry and irrigated conditions with an R2 of 0.74 and 0.52, respectively. We estimated an annual reduction of monoterpene emission of 67% for the extended and severe drought period in 2006 in the investigated Mediterranean ecosystem. It is concluded that process-based ecosystem models can provide a useful tool to investigate the involved mechanisms and to quantify the importance of specific environmental constraints.

Isoprene emission‐free poplars – a chance to reduce the impact from poplar plantations on the atmosphere

• Depending on the atmospheric composition, isoprene emissions from plants can have a severe impact on air quality and regional climate. For the plant itself, isoprene can enhance stress tolerance and also interfere with the attraction of herbivores and parasitoids. • Here, we tested the growth performance and fitness of Populus × canescens in which isoprene emission had been knocked down by RNA interference technology (PcISPS-RNAi plants) for two growing seasons under outdoor conditions. • Neither the growth nor biomass yield of the PcISPS-RNAi poplars was impaired, and they were even temporarily enhanced compared with control poplars. Modelling of the annual carbon balances revealed a reduced carbon loss of 2.2% of the total gross primary production by the absence of isoprene emission, and a 6.9% enhanced net growth of PcISPS-RNAi poplars. However, the knock down in isoprene emission resulted in reduced susceptibility to fungal infection, whereas the attractiveness for herbivores was enhanced. • The present study promises potential for the use of non- or low-isoprene-emitting poplars for more sustainable and environmentally friendly biomass production, as reducing isoprene emission will presumably have positive effects on regional climate and air quality.

Simulating mycorrhiza contribution to forest C- and N cycling-the MYCOFON model

Although mycorrhiza has been identified to be of major importance for plant nutrition and ecosystem stability, existing C- and N- simulation models on the ecosystem scale do not explicitly consider the feedbacks between ectomycorrhizal fungi and plants. We present a simple dynamic feedback model which allows estimating the main C- and N- flows between ectomycorrhizal fungi and tree roots in order to test the sensitivity of the system fungus-tree to environmental parameters and to assess the fungal contribution to plant N nutrition. Sensitivity tests carried out showed that the model responses to variations of model parameters, particularly with regard to N availability, are in agreement with published results from field and laboratory studies. However, there are still some processes and parameters which are not well constrained. Fungal N uptake rates and the ratio between mycelium, hartig net, and mantle biomass are parameters which significantly affect model results but for which published data are scarce or missing. Nevertheless, the model is already providing a platform to test our understanding of the importance of mycorrhiza for forest stand nutrition. Future coupling to a mechanistic ecosystem model will allow simulating the importance of mycorrhization for e.g. stand growth and C and N retention.

Biomass production potential from Populus short rotation systems in Romania

The aim of this study was to assess the potential of biomass production by short rotation poplar in Romania without constraining agricultural food production. Located in the eastern part of Europe, Romania provides substantial land resources suitable for bioenergy production. The process‐oriented biogeochemical model Landscape DNDC was used in conjunction with the forest‐growth model PSIM to simulate the yield of poplar grown in short‐rotation coppice at different sites in Romania. The model was validated on five sites with different climatic conditions in Central Europe. Using regional site conditions, with climatic parameters and organic carbon content in soil being the most important, the biomass production potential of poplar plantations was simulated for agricultural areas across Romania.

Modelling of microbial carbon and nitrogen turnover in soil with special emphasis on N-trace gases emission

We present a new model unifying state-of-the-art descriptions of microbial processes for denitrification, nitrification and decomposition of soil organic matter. The model is of medium complexity, filling a gap between simplistic model approaches with low predictive power and complex models, which are difficult to verify experimentally. The model Microbial Carbon and Nitrogen Turnover in soils (MiCNiT) is written in Ansi C++ and embedded into a modelling framework (MoBiLE) that provides initial conditions and accompanying ecosystem processes such as N uptake by plants, litterfall, soil water and soil temperature with established model approaches. The MiCNiT model explicitly calculates decomposition, dynamics of microbial biomass, denitrification, autotrophic and heterotrophic nitrification, applying the microbial activity concept, as well as transport of gases and solutes between anaerobic and aerobic soil fractions and through the soil profile. The model was tested against N2O and CO2 emission as well as C and N pool data from the Höglwald Forest, Germany. Due to a detailed description of the soil biochemistry and gaseous transfers, MiCNiT is capable of simulating soil air NO, N2O and N2 concentrations and the net exchange of these gases at the soil-atmosphere interface, including a possible net uptake of N2O by soils.