Terrestrial ecosystems sequester 2.1 Pg of atmospheric carbon annually. A large amount of the terrestrial sink is realized by forests. However, considerable uncertainties remain regarding the fate of this carbon over both short and long timescales. Relevant data to address these uncertainties are being collected at many sites around the world, but syntheses of these data are still sparse. To facilitate future synthesis activities, we have assembled a comprehensive global database for forest ecosystems, which includes carbon budget variables (fluxes and stocks), ecosystem traits (e.g. leaf area index, age), as well as ancillary site information such as management regime, climate, and soil characteristics. This publicly available database can be used to quantify global, regional or biome-specific carbon budgets; to re-examine established relationships; to test emerging hypotheses about ecosystem functioning [e.g. a constant net ecosystem production (NEP) to gross primary production (GPP) ratio]; and as benchmarks for model evaluations. In this paper, we present the first analysis of this database. We discuss the climatic influences on GPP, net primary production (NPP) and NEP and present the CO 2 balances for boreal, temperate, and tropical forest biomes based on micrometeorological, ecophysiological, and biometric flux and inventory estimates. Globally, GPP of forests benefited from higher temperatures and precipitation whereas NPP saturated above either a threshold of 1500 mm precipitation or a mean annual temperature of 10 1C. The global pattern in NEP was insensitive to climate and is hypothesized to be mainly determined by nonclimatic conditions such as successional stage, management, site history, and site disturbance. In all biomes, closing the CO 2 balance required the introduction of substantial biome-specific closure terms. Nonclosure was taken as an indication that respiratory processes, advection, and non-CO 2 carbon fluxes are not presently being adequately accounted for. Nomenclauture:DOC 5 dissolved organic carbon; fNPP 5 foliage component of NPP; GPP 5 gross primary production (GPP40 denotes photosynthetic uptake); mNPP 5 missing component of NPP;NBP 5 net biome production (NBP40 denotes biome uptake); NECB 5 net ecosystem carbon balance (NECB40 denotes ecosystem uptake); NEE 5 net ecosystem exchange (NEE40 denotes ecosystem uptake); NEP 5 net ecosystem production (NEP40 denotes ecosystem uptake); NPP 5 net primary production (NPP40 denotes ecosystem uptake); R a 5 autotrophic respiration (R a 40 denotes respiratory losses); R e 5 ecosystem respiration (R e 40 denotes respiratory losses); R h 5 heterotrophic respiration (R h 40 denotes respiratory losses); rNPP 5 root component of NPP;R s 5 soil respiration (R s 40 denotes respiratory losses); VOC 5 volatile organic compounds; wNPP 5 wood component of NPP

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This paper develops a statistical model for daily gross primary production (GPP) in boreal and temperate coniferous forests. The model applies the light use efficiency (LUE) approach, which estimates the conversion efficiency of daily absorbed photosynthetically active radiation (APAR) into daily GPP as a product of potential LUE and modifying factors. The latter were derived from daily total APAR and daily mean temperature, vapour pressure deficit (VPD) and soil water content (SWC). Modelling data came from five European eddy covariance measurement towers over 2-8 years. The model was tested against independent data from two AmeriFlux stations. The model with the APAR, temperature and VPD modifiers worked well in almost all the site-year combinations, but the SWC modifier only improved the fit in few cases. Geographical variation was found in the modifiers and potential LUE in site-specific models. When a model was fitted to pooled data, differences between sites could be explained by potential LUE, leaf area and environmental conditions. The test against the AmeriFlux data corroborated this finding. The potential LUE varied from 1.9 to 3.1 g C MJ À1 , and a weak correlation was found between foliar nitrogen concentration and potential LUE. Some year-to-year variation remained which could be captured by neither the pooled nor the site-specific models.

Disturbance of ecosystems is a major factor in regional carbon budgets, and it is believed to be partly responsible for the large inter-annual variability of the terrestrial part of the carbon balance. Forest fires have so far been considered as the most important disturbance but also other forms of disturbance such as insect outbreaks or wind-throw might contribute significantly to the largely unexplained inter-annual variability, at least in specific regions. The effect of wind-throw has not yet been estimated because of lack of data on how carbon fluxes are affected. The Gudrun storm, which hit Sweden in January 2005, resulted in ca. 66 million m 3 of wind-thrown stem wood on an area of ca. 272 000 ha. Using a model (BIOME-BGC) calibrated to CO 2 flux measurements at two sites, the annual net ecosystem productivity during the first year after the storm was estimated to be in the range À897 to À1259 g C m À2 yr À1 . This is a much higher loss compared with harvested (clear-cut) forests in Europe, which ranged between ca. À420 and À100 g m À2 yr À1 . The reduction in the carbon sink scaled to the whole wind-thrown area was estimated at ca. 3 million tons C during the first year. By historical data on windthrow in Europe combined with modelling, we estimated that the large Lothar storm in 1999 reduced the European carbon balance by ca. 16 million tons C, this is ca. 30% of the net biome production in Europe. We conclude that the impact of increased forest damage by more frequent storms in future climate change scenarios must be considered and that intermittent large wind-throw events may explain a part of the large inter-annual variability in the terrestrial carbon sink.

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Widespread tree species must show physiological and structural plasticity to deal with contrasting water balance conditions. To investigate these plasticity mechanisms, a meta-analysis of Pinus sylvestris L. sap flow and its response to environmental variables was conducted using datasets from across its whole geographical range. For each site, a Jarvis-type, multiplicative model was used to fit the relationship between sap flow and photosynthetically active radiation, vapour pressure deficit (D) and soil moisture deficit (SMD); and a logarithmic function was used to characterize the response of stomatal conductance (G(s)) to D. The fitted parameters of those models were regressed against climatic variables to study the acclimation of Scots pine to dry/warm conditions. The absolute value of sap flow and its sensitivity to D and SMD increased with the average summer evaporative demand. However, relative sensitivity of G(s) to D (m/G (s,ref), where m is the slope and G(s,ref) is reference G(s) at D = 1 kPa) did not increase with evaporative demand across populations, and transpiration per unit leaf area at a given D increased accordingly in drier/warmer climates. This physiological plasticity was linked to the previously reported climate- and size-related structural acclimation of leaf to sapwood area ratios. G (s,ref), and its absolute sensitivity to D(m), tended to decrease with age/height of the trees as previously reported for other pine species. It is unclear why Scots pines have higher transpiration rates at drier/warmer sites, at the expense of lower water-use efficiency. In any case, our results suggest that these structural adjustments may not be enough to prevent lower xylem tensions at the driest sites.

Abstract.Temperature measurements in trunks and branches in a mature ca. 100 years-old mixed pine and spruce forest in central Sweden were used to estimate the heat storage in the tree biomass. The estimated heat flux in the sample trees and data on biomass distributions were used to scale up to stand level biomass heat fluxes. The rate of change of sensible and latent heat storage in the air layer below the level of the flux measurements was estimated from air temperature and humidity profile measurements and soil heat flux was estimated from heat flux plates and soil temperature measurements. The fluxes of sensible and latent heat from the forest were measured with an eddy covariance system in a tower. The analysis was made for a two-month period in summer of 1995. The tree biomass heat flux was the largest of the estimated storage components and varied between 40 and −35 W m −2 on summer days with nice weather. Averaged over two months the diurnal maximum of total heat storage was 45 W m −2 and the minimum was −35 W m −2 . The soil heat flux and the sensible heat storage in air were out of phase with the biomass flux and they reached maximum values that were about 75% of the maximum of the tree biomass heat storage. The energy balance closure improved significantly when the total heat storage was added to the turbulent fluxes. The slope of a regression line with sum of fluxes and storage as independent and net radiation as dependent variable, increased from 0.86 to 0.95 for half-hourly data and the scatter was also reduced. The most significant finding was, however, that during nights with strongly stable conditions when the sensible heat flux dropped to nearly zero, the total storage matched the net radiation very well. Another interesting result was that the mean energy imbalance started to increase when the Richardson number became more negative than ca. −0.1. In fact, the largest energy deficit occurred at Correspondence to: A. Lindroth (anders.lindroth@nateko.lu.se) maximum instability. Our conclusion is that eddy covariance measurements can function well during stable conditions but that the functioning under strong instabilities might be a so far unforeseen problem.

Abstract. The DO3SE (Deposition of O3 for Stomatal Exchange) model is an established tool for estimating ozone (O3) deposition, stomatal flux and impacts to a variety of vegetation types across Europe. It has been embedded within the EMEP (European Monitoring and Evaluation Programme) photochemical model to provide a policy tool capable of relating the flux-based risk of vegetation damage to O3 precursor emission scenarios for use in policy formulation. A key limitation of regional flux-based risk assessments has been the assumption that soil water deficits are not limiting O3 flux due to the unavailability of evaluated methods for modelling soil water deficits and their influence on stomatal conductance (gsto), and subsequent O3 flux. This paper describes the development and evaluation of a method to estimate soil moisture status and its influence on gsto for a variety of forest tree species. This DO3SE soil moisture module uses the Penman-Monteith energy balance method to drive water cycling through the soil-plant-atmosphere system and empirical data describing gsto relationships with pre-dawn leaf water status to estimate the biological control of transpiration. We trial four different methods to estimate this biological control of the transpiration stream, which vary from simple methods that relate soil water content or potential directly to gsto, to more complex methods that incorporate hydraulic resistance and plant capacitance that control water flow through the plant system. These methods are evaluated against field data describing a variety of soil water variables, gsto and transpiration data for Norway spruce (Picea abies), Scots pine (Pinus sylvestris), birch (Betula pendula), aspen (Populus tremuloides), beech (Fagus sylvatica) and holm oak (Quercus ilex) collected from ten sites across Europe and North America. Modelled estimates of these variables show consistency with observed data when applying the simple empirical methods, with the timing and magnitude of soil drying events being captured well across all sites and reductions in transpiration with the onset of drought being predicted with reasonable accuracy. The more complex methods, which incorporate hydraulic resistance and plant capacitance, perform less well, with predicted drying cycles consistently underestimating the rate and magnitude of water loss from the soil. A sensitivity analysis showed that model performance was strongly dependent upon the local parameterisation of key model drivers such as the maximum gsto, soil texture, root depth and leaf area index. The results suggest that the simple modelling methods that relate gsto directly to soil water content and potential provide adequate estimates of soil moisture and influence on gsto such that they are suitable to be used to assess the potential risk posed by O3 to forest trees across Europe.