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Background To assess the extent of belowground carbon transfer to ectomycorrhizal fungi in natural forests, we used δ13C and loge C/N measurements to calculate spatial dynamics of carbon movement into ectomycorrhizal sporocarps. Methods Fourteen broad-leaved trees were labeled with 13C-depleted CO2 from 2001–2005 in Switzerland and 13C traced into ectomycorrhizal sporocarps collected at different distances. We then used stepwise regressions on patterns of δ13C and loge C/N in ectomycorrhizal sporocarps as a function of distance (zone), solar radiation, fungal genus, and association type. Results CO2-labeled trees contributed 76 ± 5%, 36 ± 6%, and 19 ± 7% of sporocarp carbon at 0–6 m, 6–12 m, and 12–18 m from labeled trees, respectively. Literature estimates of hyphal development in different taxa did not correlate with carbon acquisition patterns. After drought in 2003, sporocarp loge C/N was low in 2004 and 13C-depleted carbon from elevated CO2 trees contributed less than in other years to sporocarps. In contrast, sporocarp loge C/N peaked in 2005 and contributions from elevated CO2 trees to the 6–12 m zone increased. Therefore, carbohydrate transport belowground decreased in 2004, reflecting plant allocation priorities, and increased in 2005. Sporocarp loge C/N varied less among years under elevated CO2 than elsewhere. Conclusions These patterns indicated that 1) belowground transport was influenced by climate and plant allocation, 2) root transport rather than ectomycorrhizal transport drove carbon spatial dynamics of ectomycorrhizal fungi, and 3) elevated CO2 decreased the sensitivity of belowground allocation to climatic fluctuations, suggesting improved drought resistance in a high-CO2 world.

Under the United Nations Framework Convention on Climate Change (UNFCCC), industrialized countries and countries with economies in transition (so called Annex 1 countries) are encouraged to move towards more sophisticated approaches for national greenhouse gas reporting. To develop a model-based approach for estimating nitrous oxide (N2O) emissions from agricultural soils, model calibration is one of the first important steps. Extensive multisite field observations are necessary for this purpose, as agricultural management in Western Europe is complex (e.g., diverse crop rotations, different types of fertilizer and soil tillage). In the present study, we used ca. 24,000 daily N2O flux observations from six cropland sites, two in France and four in Switzerland, to conduct an automatic data-driven calibration of the biogeochemical model DayCent. This model is planned to be used for greenhouse gas reporting in the entire European Union as well as in Switzerland. After a site-specific calibration, a leave-one-out (LOO) cross-evaluation was conducted to assess the model’s ability to predict N2O emissions for sites it was not calibrated for. Mean observed N2O fluxes for 54 interactions of crop cycles, field studies and treatments were used to evaluate the model. The LOO cross-evaluation resulted in a R2 of 0.63 for the prediction of mean N2O fluxes per crop cycle, compared to an R2 of 0.51 obtained with default parameterization. Our results showed that the improvement in N2O predictions was associated with the adjustment of only seven parameters controlling the N cycle in soil (e.g., the maximum daily nitrification amount and the inflection point for the effect of water-filled pore space on denitrification) out of several hundred parameters. These parameters showed a wide range of values between sites, revealing an important challenge for calibration-based improvement of N2O simulations. Despite the remaining uncertainty, our model-based estimates of N2O emission per crop cycle (2.64 kg N ha-1) were clearly closer to measurements (2.67 kg N ha-1) than commonly used emission factor approaches (1.60–1.71 kg N ha-1). Based on extensive field observations, our results suggest that, after data-driven calibration of only few N cycle parameters, DayCent simulations are useful for reporting N2O emissions of complex cropland management. These model based-estimates were more accurate, because they consider key drivers that are disregarded by simpler approaches. Moving towards more complex methods of N2O reporting, is therefore expected to improve the accuracy and additionally allows to assess mitigation options., European Journal of Agronomy, 141, ISSN:1161-0301, ISSN:1873-7331

Background. The spatial extent and timing of carbon fluxes from mature trees to ectomycorrhizal fungi associated with different hosts is challenging to assess in natural forests but could provide insights into carbon dynamics of fungi differing in exploration capabilities. Methods. We analyzed carbon movement into ectomycorrhizal sporocarps at the Swiss Forest Free-Air CO2 Enrichment (FACE) site in 2010 and 2011 during continuous labeling of five mature Picea abies (L.) H. Karst. with 13C-depleted CO2. Sporocarps were collected 0-6 m, 6-12 m, 12-18 m, and > 18 m from 13C-labeled trees. We then used stepwise regressions on patterns of 13C:12C ratios (δ13C) in ectomycorrhizal sporocarps as a function of distance (zone), solar radiation, fungal genus, and association type. Results. 18%, 10%, 40%, and 32% of ectomycorrhizal sporocarps were associated with conifers, broad-leaved trees, both, or of unknown association, respectively. Conifer-associated sporocarps derived 58 ± 7%, 41 ± 10%, and 20 ± 7% of their carbon from labeled trees in the 0-6 m, 6-12 m and 12-18 m zones, respectively, whereas other sporocarps in the 0-6 m zone only derived 34 ± 4% of their carbon from labeled trees. Sporocarp δ13C correlated positively with solar radiation for the 16-19 days prior to sporocarp harvest. Conclusions. These patterns indicated that (1) carbon spread further through conifer-associated mycorrhizal networks than through other fungal networks, and (2) carbon accumulation, storage, and transfer dynamics for sporocarp formation can take several weeks. Mature conifers supplied carbon to mycorrhizal fungi up to ~15 m from host trees.

Summary The application of dynamic models to report changes in soil organic carbon (SOC) stocks, for example as part of greenhouse gas inventories, is becoming increasingly important. Most of these models rely on input data from harvest residues or decaying plant parts and also organic fertilizer, together referred to as soil carbon inputs (C). The soil C inputs from plants are derived from measured agricultural yields using allometric equations. Here we compared the results of five previously published equations. Our goal was to test whether the choice of method is critical for modelling soil C and if so, which of these equations is most suitable for Swiss conditions. For this purpose we used the five equations to calculate soil C inputs based on yield data from a Swiss long-term cropping experiment. Estimated annual soil C inputs from various crops were averaged from 28 years and four fertilizer treatments; the average was 3.6 Mg C ha−1 and covered a surprisingly large range from 2.1 to 5.3 Mg C ha−1 year−1 among the five equations. For single crop species, differences reached 6.6 Mg C ha−1 year−1. The variation between estimated soil C inputs depended on crop type and increased with yield. Simulations with the model C-TOOL showed that calculated SOC stocks were affected strongly by the choice of the allometric equation. With four equations, a decrease in SOC stocks was simulated, whereas with one equation there was no change. This considerable uncertainty in modelled soil C is attributable solely to the allometric equation used to estimate the soil C input. We identify the evaluation and selection of allometric equations and associated coefficients as critical steps when setting up a model-based soil C inventory for agricultural systems. Highlights Uncertainty analyses in soil C modelling have focused mainly on model parameters and structures. The uncertainty related to estimation of soil C inputs from crop residues was calculated. Simulated soil C stocks and stock changes depend strongly on the method of estimating C input. The method to derive soil C inputs is critical for model-based soil C inventories.

Models predicting ecosystem carbon dioxide (CO2 ) exchange under future climate change rely on relatively few real-world tests of their assumptions and outputs. Here, we demonstrate a rapid and cost-effective method to estimate CO2 exchange from intact vegetation patches under varying atmospheric CO2 concentrations. We find that net ecosystem CO2 uptake (NEE) in a boreal forest rose linearly by 4.7 ± 0.2% of the current ambient rate for every 10 ppm CO2 increase, with no detectable influence of foliar biomass, season, or nitrogen (N) fertilization. The lack of any clear short-term NEE response to fertilization in such an N-limited system is inconsistent with the instantaneous downregulation of photosynthesis formalized in many global models. Incorporating an alternative mechanism with considerable empirical support - diversion of excess carbon to storage compounds - into an existing earth system model brings the model output into closer agreement with our field measurements. A global simulation incorporating this modified model reduces a long-standing mismatch between the modeled and observed seasonal amplitude of atmospheric CO2 . Wider application of this chamber approach would provide critical data needed to further improve modeled projections of biosphere-atmosphere CO2 exchange in a changing climate.

Records of stable oxygen isotope ratios in tree rings are valuable tools to reconstruct past climatic conditions and investigate the response of trees to those conditions. So far the use of stable oxygen isotope signatures of tree rings has not been systematically evaluated in dynamic global vegetation models (DGVMs). DGVMs integrate many hydrological and physiological processes and their application could improve proxy-model comparisons and the interpretation of oxygen isotope records. Here we present an approach to simulate leaf water and stem cellulose δ18O of trees using the LPX-Bern DGVM (LPX-Bern). Our results lie within a few per mil of measured tree ring δ18O of 31 different forest stands mainly located in Europe. Temporal means over the last 5 decades as well as interannual variations for a subset of sites in Switzerland are captured. A sensitivity analysis reveals that relative humidity, temperature, and the water isotope boundary conditions have the largest influence on simulated stem cellulose δ18O, followed by all climatic factors combined, whereas increasing atmospheric CO2 and nitrogen deposition exert no impact. We conclude that simulations with LPX-Bern are useful for investigating large-scale oxygen isotope patterns of tree ring cellulose to elucidate the importance of different environmental factors on isotope variations and therefore help to reduce uncertainties in the interpretation of δ18O of tree rings.

Soil carbon sequestration (SCS) is one of the cheapest and technically least demanding carbon dioxide (CO2) removal (CDR) or negative CO2 emission technologies. For a realistic assessment of SCS, it is critical to evaluate how much carbon (C) can be stored in soil organic matter under actual agricultural practices. This includes typical crop rotations and fertilization strategies, depends on resources that are available (e.g. farmyard manure (FYM)) and are affordable for farmers. Furthermore, it is important to assess SCS based on given climatic and soil conditions. Here, we evaluate changes in soil C storage for Switzerland using data from eleven long-term field experiments on cropland and permanent grassland that include common local practices. At all sites, changes in soil organic carbon (SOC) stocks were measured in topsoil (∼0-0.2 m) in response to a total of 80 different treatments including different types of mineral or organic fertilization (e.g. FYM, slurry, peat, compost) or soil management (tillage vs. no-till). The treatments were applied to different, diverse crop rotations or grass mixtures that are representative for Switzerland. We found that topsoils lost C at an average rate of 0.29 Mg C ha−1 yr−1, although many of the investigated treatments were expected to lead to SOC increases. Based on a linear mixed effects model we showed that SOC change rates (ΔSOC) were driven by C inputs to soil (harvest residues and organic fertilizer), soil cover and initial SOC stocks. The type of land use or soil tillage had no significant effect. Our analysis suggests that current efforts to manage soils sustainably need to be intensified and complemented with further techniques if Switzerland wants to achieve the goal of the 4 per 1000 initiative.

[1] Our understanding of Earth's carbon climate system depends critically upon interactions between rising atmospheric CO2, changing land use, and nitrogen limitation on vegetation growth. Using a global land model, we show how these factors interact locally to generate the global land carbon sink over the past 200 years. Nitrogen constraints were alleviated by N2 fixation in the tropics and by atmospheric nitrogen deposition in extratropical regions. Nonlinear interactions between land use change and land carbon and nitrogen cycling originated from three major mechanisms: (i) a sink foregone that would have occurred without land use conversion; (ii) an accelerated response of secondary vegetation to CO2 and nitrogen, and (iii) a compounded clearance loss from deforestation. Over time, these nonlinear effects have become increasingly important and reduce the present-day net carbon sink by ~40% or 0.4 PgC yr−1.

Rapidly increasing atmospheric CO2 is not only changing the climate system but may also affect the biosphere directly through stimulation of plant growth and ecosystem carbon and nutrient cycling. Although forest ecosystems play a critical role in the global carbon cycle, experimental information on forest responses to rising CO2 is scarce, due to the sheer size of trees. Here, we present a synthesis of the only study world-wide where a diverse set of mature broadleaved trees growing in a natural forest has been exposed to future atmospheric CO2 levels (c. 550ppm) by free-air CO2 enrichment (FACE). We show that litter production, leaf traits and radial growth across the studied hardwood species remained unaffected by elevated CO2 over 8years. CO2 enrichment reduced tree water consumption resulting in detectable soil moisture savings. Soil air CO2 and dissolved inorganic carbon both increased suggesting enhanced below-ground activity. Carbon release to the rhizosphere and/or higher soil moisture primed nitrification and nitrate leaching under elevated CO2; however, the export of dissolved organic carbon remained unaltered.Synthesis. Our findings provide no evidence for carbon-limitation in five central European hardwood trees at current ambient CO2 concentrations. The results of this long-term study challenge the idea of a universal CO2 fertilization effect on forests, as commonly assumed in climate-carbon cycle models.

Fruiting is typically considered to massively burden the seasonal carbon budget of trees. The cost of reproduction has therefore been suggested as a proximate factor explaining observed mast-fruiting patterns. Here, we used a large-scale, continuous (13)C labeling of mature, deciduous trees in a temperate Swiss forest to investigate to what extent fruit formation in three species with masting reproduction behavior (Carpinus betulus, Fagus sylvatica, Quercus petraea) relies on the import of stored carbon reserves. Using a free-air CO2 enrichment system, we exposed trees to (13)C-depleted CO2 during 8 consecutive years. By the end of this experiment, carbon reserve pools had significantly lower δ(13)C values compared to control trees. δ(13)C analysis of new biomass during the first season after termination of the CO2 enrichment allowed us to distinguish the sources of built-in carbon (old carbon reserves vs. current assimilates). Flowers and expanding leaves carried a significant (13)C label from old carbon stores. In contrast, fruits and vegetative infructescence tissues were exclusively produced from current, unlabeled photoassimilates in all three species, including F. sylvatica, which had a strong masting season. Analyses of δ(13)C in purified starch from xylem of fruit-bearing shoots revealed a complete turn-over of starch during the season, likely due to its usage for bud break. This study is the first to directly demonstrate that fruiting is independent from old carbon reserves in masting trees, with significant implications for mechanistic models that explain mast seeding.

Emerging leaves in evergreen tree species are supplied with carbon (C) from the previous year's foliage. In deciduous trees, no older leaves are present, and the early phase of leaf development must rely on C reserves from other tissues. How soon developing leaves become autotrophic and switch from being C sinks to sources has rarely been studied in mature forest trees, and simultaneous comparisons of species are scarce. Using a canopy crane and a simple (13)CO(2)-pulse-labelling technique, we demonstrate that young leaves of mature trees in three European deciduous species (Fagus sylvatica L., Quercus petraea (Matt.) Liebl., Tilia platyphyllos Scop.) start assimilating CO(2) at a very early stage of development (10-50% expanded). One month after labelling, all leaves were still strongly (13)C enriched, suggesting that recent photosynthates had been incorporated into slow turnover pools such as cellulose or lignin and thus had contributed to leaf growth. In line with previous studies performed at the same site, we found stronger incorporation of recent photosynthates into growing tissues of T. platyphyllos compared with F. sylvatica and Q. petraea. Non-structural carbohydrate (NSC) concentrations analysed for one of the three study species (F. sylvatica) showed that sugar and starch pools rapidly increased during leaf development, suggesting that newly developed leaves soon produce more NSC than can be used for growth. In conclusion, our findings indicate that expanding leaves of mature deciduous trees become C autonomous at an early stage of development despite the presence of vast amounts of mobile carbohydrate reserves.

Physical diffusion of isotopic tracers into and out of soil pores causes considerable uncertainty for the timing and magnitude of plant belowground allocation in pulse-labelling experiments. Here, we partitioned soil CO(2) isotopic fluxes into abiotic tracer flux (physical return), heterotrophic flux, and autotrophic flux contributions following (13)CO(2) labelling of a Swedish Pinus sylvestris forest. Soil CO(2) efflux and its isotopic composition from a combination of deep and surface soil collars was monitored using a field-deployed mass spectrometer. Additionally, (13)CO(2) within the soil profile was monitored. Physical (abiotic) efflux of (13)CO(2) from soil pore spaces was found to be significant for up to 48 h after pulse labelling, and equalled the amount of biotic label flux over 6 d. Measured and modelled changes in (13)CO(2) concentration throughout the soil profile corroborated these results. Tracer return via soil CO(2) efflux correlated significantly with the proximity of collars to trees, while daily amplitudes of total flux (including heterotrophic and autotrophic sources) showed surprising time shifts compared with heterotrophic fluxes. The results show for the first time the significance of the confounding influence of physical isotopic CO(2)-tracer return from the soil matrix, calling for the inclusion of meaningful control treatments in future pulse-chase experiments.

How rapidly newly assimilated carbon (C) is invested into recalcitrant structures of forests, and how closely C pools and fluxes are tied to photosynthesis, is largely unknown. A crane and a purpose-built free-air CO2 enrichment (FACE) system permitted us to label the canopy of a mature deciduous forest with 13C-depleted CO2 for 4 yr and continuously trace the flow of recent C through the forest without disturbance. Potted C4 grasses in the canopy ('isometers') served as a reference for the C-isotope input signal. After four growing seasons, leaves were completely labelled, while newly formed wood (tree rings) still contained 9% old C. Distinct labels were found in fine roots (38%) and sporocarps of mycorrhizal fungi (62%). Soil particles attached to fine roots contained 9% new C, whereas no measurable signal was detected in bulk soil. Soil-air CO2 consisted of 35% new C, indicating that considerable amounts of assimilates were rapidly returned back to the atmosphere. These data illustrate a relatively slow dilution of old mobile C pools in trees, but a pronounced allocation of very recent assimilates to C pools of short residence times.

Symbioses between plant roots and mycorrhizal fungi are thought to enhance plant uptake of nutrients through a favourable exchange for photosynthates. Ectomycorrhizal fungi are considered to play this vital role for trees in nitrogen (N)-limited boreal forests. We followed symbiotic carbon (C)-N exchange in a large-scale boreal pine forest experiment by tracing (13) CO(2) absorbed through tree photosynthesis and (15) N injected into a soil layer in which ectomycorrhizal fungi dominate the microbial community. We detected little (15) N in tree canopies, but high levels in soil microbes and in mycorrhizal root tips, illustrating effective soil N immobilization, especially in late summer, when tree belowground C allocation was high. Additions of N fertilizer to the soil before labelling shifted the incorporation of (15) N from soil microbes and root tips to tree foliage. These results were tested in a model for C-N exchange between trees and mycorrhizal fungi, suggesting that ectomycorrhizal fungi transfer small fractions of absorbed N to trees under N-limited conditions, but larger fractions if more N is available. We suggest that greater allocation of C from trees to ectomycorrhizal fungi increases N retention in soil mycelium, driving boreal forests towards more severe N limitation at low N supply.

Fine roots play a key role in the forest carbon balance, but their carbon dynamics remain largely unknown. We pulse labelled 50 m(2) patches of young boreal forest by exposure to (13)CO(2) in early and late summer. Labelled photosynthates were traced into carbon compounds of1 and 1-3 mm diameter roots (fine roots), and into bulk tissue of these and first-order roots (root tips). Root tips were the most strongly labelled size class. Carbon allocation to all size classes was higher in late than in early summer; mean residence times (MRTs) in starch increased from 4 to 11 months. In structural compounds, MRTs were 0.8 yr in tips and 1.8 yr in fine roots. The MRT of carbon in sugars was in the range of days. Functional differences within the fine root population were indicated by carbon allocation patterns and residence times. Pronounced allocation of recent carbon and higher turnover rates in tips are associated with their role in nutrient and water acquisition. In fine roots, longer MRTs but high allocation to sugars and starch reflect their role in structural support and storage. Accounting for heterogeneity in carbon residence times will improve and most probably reduce the estimates of fine root production.

*The flux of carbon from tree photosynthesis through roots to ectomycorrhizal (ECM) fungi and other soil organisms is assumed to vary with season and with edaphic factors such as nitrogen availability, but these effects have not been quantified directly in the field. *To address this deficiency, we conducted high temporal-resolution tracing of (13)C from canopy photosynthesis to different groups of soil organisms in a young boreal Pinus sylvestris forest. *There was a 500% higher below-ground allocation of plant C in the late (August) season compared with the early season (June). Labelled C was primarily found in fungal fatty acid biomarkers (and rarely in bacterial biomarkers), and in Collembola, but not in Acari and Enchytraeidae. The production of sporocarps of ECM fungi was totally dependent on allocation of recent photosynthate in the late season. There was no short-term (2 wk) effect of additions of N to the soil, but after 1 yr, there was a 60% reduction of below-ground C allocation to soil biota. *Thus, organisms in forest soils, and their roles in ecosystem functions, appear highly sensitive to plant physiological responses to two major aspects of global change: changes in seasonal weather patterns and N eutrophication.

Physical diffusion of isotopic tracers into and out of soil pores causes considerable uncertainty for the timing and magnitude of plant belowground allocation in pulse-labelling experiments. Here, we partitioned soil CO(2) isotopic fluxes into abiotic tracer flux (physical return), heterotrophic flux, and autotrophic flux contributions following (13)CO(2) labelling of a Swedish Pinus sylvestris forest. Soil CO(2) efflux and its isotopic composition from a combination of deep and surface soil collars was monitored using a field-deployed mass spectrometer. Additionally, (13)CO(2) within the soil profile was monitored. Physical (abiotic) efflux of (13)CO(2) from soil pore spaces was found to be significant for up to 48 h after pulse labelling, and equalled the amount of biotic label flux over 6 d. Measured and modelled changes in (13)CO(2) concentration throughout the soil profile corroborated these results. Tracer return via soil CO(2) efflux correlated significantly with the proximity of collars to trees, while daily amplitudes of total flux (including heterotrophic and autotrophic sources) showed surprising time shifts compared with heterotrophic fluxes. The results show for the first time the significance of the confounding influence of physical isotopic CO(2)-tracer return from the soil matrix, calling for the inclusion of meaningful control treatments in future pulse-chase experiments.

Stable C isotope signals in plant tissues became a key tool in explaining growth responses to the environment. The technique is based on the fundamental assumption that the isotopic composition of a given unit of tissue (e.g. a tree ring) reflects the specific C uptake conditions in the leaf at a given time. Beyond the methodological implications of any deviation from this assumption, it is of physiological interest whether new C is transferred directly from sources (a photosynthesizing leaf) to structural sinks (e.g. adjacent stem tissue), or inherently passes through existing (mobile) C pools, which may be of variable (older) age. Here, we explore the fate of (13)C-labelled photosynthates in the crowns of a 30-35 m tall, mixed forest using a canopy crane. In all nine study species labelled C reached woody tissue within 2-9 h after labelling. Four months later, very small signals were left in branch wood of Tilia suggesting that low mixing of new, labelled C with old C had taken place. In contrast, signals in Fagus and Quercus had increased, indicating more intense mixing. This species-specific mixing of new with old C pools is likely to mask year- or season-specific linkages between tree ring formation and climate and has considerable implications for climate reconstruction using stable isotopes as proxies for past climatic conditions.

The pathways of currently fixed carbon in fruit bearing branchlets were investigated in two temperate forest tree species (CARPINUS BETULUS and FAGUS SYLVATICA), which differ in texture of their vegetative infructescence tissues (leaf-like in CARPINUS vs. woody in FAGUS). During late spring, (13)C pulse-labelling was conducted on girdled, defoliated, girdled plus defoliated and untreated fruiting branchlets of mature trees IN SITU, to assess changes in C relations in response to the introduced C source-sink imbalances. At harvest in early August, 75 - 100 % of the recovered (13)C label was bound to infructescences (either fruits or vegetative infructescence tissue), revealing them as the prime C sinks for current photoassimilates. Leaves on girdled branchlets were not stronger labelled than on ungirdled ones in both species, indicating no upregulation of the leaves' photosynthetic capacity in response to the prevention of phloemic transport, which was also supported by measurements of light saturated photosynthesis. In contrast, (13)C labels tended to be higher after complete defoliation in the vegetative infructescence tissues of CARPINUS, suggesting enhanced net photosynthesis of green infructescence parts as compensation for the loss of regular leaves. The total labelling-derived (13)C content of whole infructescences was very similar between foliated and defoliated CARPINUS branchlets. Cupulae of FAGUS, on the other hand, remained almost unlabelled on defoliated branchlets, indicating the photosynthetic inactivity of this woody infructescence tissue. Consequently, CARPINUS still produced relatively high fruit masses on girdled plus defoliated branchlets, while in FAGUS fruit development ceased almost completely at this most severe treatment. Our results highlight that green vegetative infructescence tissue assimilates substantial amounts of C and can partly substitute regular leaves as C sources for successful fruit development.

Whether rising atmospheric carbon dioxide (CO 2 ) concentrations will cause forests to grow faster and store more carbon is an open question. Using free air CO 2 release in combination with a canopy crane, we found an immediate and sustained enhancement of carbon flux through 35-meter-tall temperate forest trees when exposed to elevated CO 2 . However, there was no overall stimulation in stem growth and leaf litter production after 4 years. Photosynthetic capacity was not reduced, leaf chemistry changes were minor, and tree species differed in their responses. Although growing vigorously, these trees did not accrete more biomass carbon in stems in response to elevated CO 2 , thus challenging projections of growth responses derived from tests with smaller trees.

Stomatal conductance (g s) of mature trees exposed to elevated CO2 concentrations was examined in a diverse deciduous forest stand in NW Switzerland. Measurements of g s were carried out on upper canopy foliage before noon, over four growing seasons, including an exceptionally dry summer (2003). Across all species reductions in stomatal conductance were smaller than 25% most likely around 10%, with much variation among species and trees. Given the large heterogeneity in light conditions within a tree crown, this signal was not statistically significant, but the responses within species were surprisingly consistent throughout the study period. Except during a severe drought, stomatal conductance was always lower in trees of Carpinus betulus exposed to elevated CO2 compared to Carpinus trees in ambient air, but the difference was only statistically significant on 2 out of 15 days. In contrast, stomatal responses in Fagus sylvatica and Quercus petraea varied around zero with no consistent trend in relation to CO2 treatment. During the 2003 drought in the third treatment year, the CO2 effect became reversed in Carpinus, resulting in higher g s in trees exposed to elevated CO2 compared to control trees, most likely due to better water supply because of the previous soil water savings. This was supported by less negative predawn leaf water potential in CO2 enriched Carpinus trees, indicating an improved water status. These findings illustrate (1) smaller than expected CO2-effects on stomata of mature deciduous forest trees, and (2) the possibility of soil moisture feedback on canopy water relations under elevated CO2.

Aims Predictions of medium to long-term changes in soil organic carbon (SOC) stocks involve simulation models with plant carbon (C) inputs estimated from crop yields using crop harvest indices and allometric scaling involving fixed root-to-shoot ratios. The choice of allometric approach and initial distribution of SOC among model pools is crucial for model estimates. Methods We evaluated the impact of a modified allometric approach for grass leys and initial model set-up using the C-TOOL model. Four European agricultural long-term experiments were considered including three experiments with different frequency of grass leys in the crop rotation and one permanent grassland. Results The modified allometric function for ley relied on fixed below-ground C input regardless of mineral fertilizer inputs and modified pool initialization involved available site history. Including available, but insufficient, pre-experiment history to adjust the initial set-up of model SOC pools did not improve to the C-TOOL simulations. Changing the allometric approach for ley from fixed shoot-to-root ratios to fixed below-ground C input decreased the soil C input dramatically and improved the C-TOOL simulation of SOC stocks for fertilized treatments in all experiments when combined with standard model set-up. For unfertilized treatments, however, the efficiency of the standard allometric function was superior to the modified one. Conclusions Our results challenge the prevalent use of allometric approaches based on fixed root-to-shoot in SOC simulation models. The potential of modifying the initial distribution of SOC among model pools by accounting for historic land use remains in the dark.