Contributions of climate, leaf area index and leaf physiology to variation in gross primary production of six coniferous forests across Europe: a model-based analysis.

Gross primary production (GPP) is the primary source of all carbon fluxes in the ecosystem. Understanding variation in this flux is vital to understanding variation in the carbon sink of forest ecosystems, and this would serve as input to forest production models. Using GPP derived from eddy-covariance (EC) measurements, it is now possible to determine the most important factor to scale GPP across sites. We use long-term EC measurements for six coniferous forest stands in Europe, for a total of 25 site-years, located on a gradient between southern France and northern Finland. Eddy-derived GPP varied threefold across the six sites, peak ecosystem leaf area index (LAI) (all-sided) varied from 4 to 22 m(2) m(-2) and mean annual temperature varied from -1 to 13 degrees C. A process-based model operating at a half-hourly time-step was parameterized with available information for each site, and explained 71-96% in variation between daily totals of GPP within site-years and 62% of annual total GPP across site-years. Using the parameterized model, we performed two simulation experiments: weather datasets were interchanged between sites, so that the model was used to predict GPP at some site using data from either a different year or a different site. The resulting bias in GPP prediction was related to several aggregated weather variables and was found to be closely related to the change in the effective temperature sum or mean annual temperature. High R(2)s resulted even when using weather datasets from unrelated sites, providing a cautionary note on the interpretation of R(2) in model comparisons. A second experiment interchanged stand-structure information between sites, and the resulting bias was strongly related to the difference in LAI, or the difference in integrated absorbed light. Across the six sites, variation in mean annual temperature had more effect on simulated GPP than the variation in LAI, but both were important determinants of GPP. A sensitivity analysis of leaf physiology parameters showed that the quantum yield was the most influential parameter on annual GPP, followed by a parameter controlling the seasonality of photosynthesis and photosynthetic capacity. Overall, the results are promising for the development of a parsimonious model of GPP.

Flux and concentration footprint modelling: state of the art.

Since early 1990s, the development of footprint models has been rapid with presently four different approaches being available: (i) analytical models, (ii) Lagrangian stochastic particle dispersion models, (iii) large-eddy simulations, and (iv) closure models. Parameterizations of some of these approaches have been developed, simplifying the original algorithms for use in practical applications. The paper provides a review of the footprint modelling. It also discusses our present understanding of the theoretical background, the most successful modelling approaches, as well as the usage and benefits of the footprint concept as it relates to flux measurements. There has recently been a trend emerging in modelling the behavior of the footprint functions using a less idealized, more realistic description of inhomogeneities, vegetation structure and topography, ultimately for reactive compounds. The estimation of footprints for application in the real world, complete with a multitude of interesting gaseous and particulate substances, remains a complex problem.

A model of bubble growth leading to xylem conduit embolism.

The dynamics of a gas bubble inside a water conduit after a cavitation event was modeled. A distinction was made between a typical angiosperm conduit with a homogeneous pit membrane and a typical gymnosperm conduit with a torus-margo pit membrane structure. For conduits with torus-margo type pits pit membrane deflection was also modeled and pit aspiration, the displacement of the pit membrane to the low pressure side of the pit chamber, was found to be possible while the emboli was still small. Concurrent with pit aspiration, the high resistance to water flow out of the conduit through the cell walls or aspirated pits will make the embolism process slow. In case of no pit aspiration and always for conduits with homogeneous pit membranes, embolism growth is more rapid but still much slower than bubble growth in bulk water under similar water tension. The time needed for the embolism to fill a whole conduit was found to be dependent on pit and cell wall conductance, conduit radius, xylem water tension, pressure rise in adjacent conduits due to water freed from the embolising conduit, and the rigidity and structure of the pits in the case of margo-torus type pit membrane. The water pressure in the conduit hosting the bubble was found to occur almost immediately after bubble induction inside a conduit, creating a sudden tension release in the conduit, which can be detected by acoustic and ultra-acoustic monitoring of xylem cavitation.

Challenges in quantifying biosphere-atmosphere exchange of nitrogen species.

Recent research in nitrogen exchange with the atmosphere has separated research communities according to N form. The integrated perspective needed to quantify the net effect of N on greenhouse-gas balance is being addressed by the NitroEurope Integrated Project (NEU). Recent advances have depended on improved methodologies, while ongoing challenges include gas-aerosol interactions, organic nitrogen and N(2) fluxes. The NEU strategy applies a 3-tier Flux Network together with a Manipulation Network of global-change experiments, linked by common protocols to facilitate model application. Substantial progress has been made in modelling N fluxes, especially for N(2)O, NO and bi-directional NH(3) exchange. Landscape analysis represents an emerging challenge to address the spatial interactions between farms, fields, ecosystems, catchments and air dispersion/deposition. European up-scaling of N fluxes is highly uncertain and a key priority is for better data on agricultural practices. Finally, attention is needed to develop N flux verification procedures to assess compliance with international protocols.

Europe-wide reduction in primary productivity caused by the heat and drought in 2003.

Future climate warming is expected to enhance plant growth in temperate ecosystems and to increase carbon sequestration. But although severe regional heatwaves may become more frequent in a changing climate, their impact on terrestrial carbon cycling is unclear. Here we report measurements of ecosystem carbon dioxide fluxes, remotely sensed radiation absorbed by plants, and country-level crop yields taken during the European heatwave in 2003. We use a terrestrial biosphere simulation model to assess continental-scale changes in primary productivity during 2003, and their consequences for the net carbon balance. We estimate a 30 per cent reduction in gross primary productivity over Europe, which resulted in a strong anomalous net source of carbon dioxide (0.5 Pg C yr(-1)) to the atmosphere and reversed the effect of four years of net ecosystem carbon sequestration. Our results suggest that productivity reduction in eastern and western Europe can be explained by rainfall deficit and extreme summer heat, respectively. We also find that ecosystem respiration decreased together with gross primary productivity, rather than accelerating with the temperature rise. Model results, corroborated by historical records of crop yields, suggest that such a reduction in Europe's primary productivity is unprecedented during the last century. An increase in future drought events could turn temperate ecosystems into carbon sources, contributing to positive carbon-climate feedbacks already anticipated in the tropics and at high latitudes.

Field measurements of ultrasonic acoustic emissions and stem diameter variations. New insight into the relationship between xylem tensions and embolism.

We examined interrelated xylem water tensions and embolism dynamics under field conditions by simultaneously monitoring ultra-acoustic emissions and changes in stem xylem diameter. Variation in stem xylem diameter was measured with linear displacement transducers to estimate variation in sap tension. Measured ultrasonic acoustic emissions coincided well with changes in xylem diameter, indicating that individual peaks in embolism occurred simultaneously with peaks in water tension. The good time resolution between measurements makes this method especially suitable for observing embolism dynamics on a short timescale. Longer lasting measurements can also be made to monitor inter-daily patterns in water tension and embolism because the techniques are non-destructive. Ultra-acoustic emissions occurred mainly during periods of decreasing stem xylem diameter, i.e., increasing water tension, when the water tension was high enough. Embolism also occurred during periods of increasing xylem diameter, i.e., decreasing water tension, but the number of embolizing conduits under these conditions was small.

Relationships between embolism, stem water tension, and diameter changes.

A model for embolism in the sapflow process was developed, in which embolism is described as a physical process linked to real physical properties of the conduits and the thermodynamic state of water. Different mechanisms leading to embolism and their effect on water relations and especially diurnal diameter changes in a tree were examined. The mechanisms of heterogeneous nucleation, air-seeding, and bubble growth have been considered. The significance of embolism has been revealed here by examining diameter changes, which is an easily measurable quantity under field conditions. The most fundamental effects of embolism on sapflow are decrease in permeability and release of water from embolizing conduits to the transpiration stream. These can be indirectly detected by observing diameter changes. If possible changes in elasticity are not accounted for, embolism generally tends to enhance the amplitude of the diurnal diameter changes due to reduced permeability and increased tensions. In the case of reduced elasticity, embolism gives rise to smaller amplitudes of diameter changes.

Tree stem diameter variations and transpiration in Scots pine: an analysis using a dynamic sap flow model.

A dynamic model for simulating water flow in a Scots pine (Pinus sylvestris L.) tree was developed. The model is based on the cohesion theory and the assumption that fluctuating water tension driven by transpiration, together with the elasticity of wood tissue, causes variations in the diameter of a tree stem and branches. The change in xylem diameter can be linked to water tension in accordance with Hookeâ s law. The model was tested against field measurements of the diurnal xylem diameter change at different heights in a 37-year-old Scots pine at Hyytiälä, southern Finland (61 degrees 51' N, 24 degrees 17' E, 181 m a.s.l.). Shoot transpiration and soil water potential were input data for the model. The biomechanical and hydraulic properties of wood and fine root hydraulic conductance were estimated from simulated and measured stem diameter changes during the course of 1 day. The estimated parameters attained values similar to literature values. The ratios of estimated parameters to literature values ranged from 0.5 to 0.9. The model predictions (stem diameters at several heights) were in close agreement with the measurements for a period of 6 days. The time lag between changes in transpiration rate and in sap flow rate at the base of the tree was about half an hour. The analysis showed that 40% of the resistance between the soil and the top of the tree was located in the rhizosphere. Modeling the water tension gradient and consequent woody diameter changes offer a convenient means of studying the link between wood hydraulic conductivity and control of transpiration.

Respiration as the main determinant of carbon balance in European forests.

Carbon exchange between the terrestrial biosphere and the atmosphere is one of the key processes that need to be assessed in the context of the Kyoto Protocol. Several studies suggest that the terrestrial biosphere is gaining carbon, but these estimates are obtained primarily by indirect methods, and the factors that control terrestrial carbon exchange, its magnitude and primary locations, are under debate. Here we present data of net ecosystem carbon exchange, collected between 1996 and 1998 from 15 European forests, which confirm that many European forest ecosystems act as carbon sinks. The annual carbon balances range from an uptake of 6.6 tonnes of carbon per hectare per year to a release of nearly 1 t C ha(-1) yr(-1), with a large variability between forests. The data show a significant increase of carbon uptake with decreasing latitude, whereas the gross primary production seems to be largely independent of latitude. Our observations indicate that, in general, ecosystem respiration determines net ecosystem carbon exchange. Also, for an accurate assessment of the carbon balance in a particular forest ecosystem, remote sensing of the normalized difference vegetation index or estimates based on forest inventories may not be sufficient.

A three-dimensional stomatal CO2 exchange model including gaseous phase and leaf mesophyll separated by irregular interface

A three-dimensional model was constructed for studying the diffusion of CO2 molecules from air into stomatal air spaces and further into leaf mesophyll cells. The model covers two different phases, mesophyll and air, with appropriate boundary conditions. Sinks for CO2 molecules were obtained from a well-known biochemical photosynthesis model parametrized for Scots pine. The effects of the varying stomatal geometry on the CO2 flux and intercellular-air-space (IAS) and mesophyll concentrations were examined. The net CO2 flux is sensitive to the size of the IAS-mesophyll interface when the mesophyll transport coefficient is low, equalling the diffusivity of CO2 in water. The simulations have revealed an optimum between the volume of the assimilating mesophyll and the air-mesophyll interface. The solubility of CO2 molecules into water in the cell surfaces, which depends on temperature and pH, had a clear effect on the flux. Increasing pH from the value indicated by the physical solubility (pH reverse similar6) leads to an increase in the temperature maximum for CO2 flux into higher temperatures and to unrealistically high CO2 concentrates in the mesophyll. Copyright 1999 Academic Press

On the concept of leaf boundary layer resistance for forced convection

The definition of leaf boundary layer resistance is reconsidered in respect of the three-dimensional diffusion-controlled mass transport region just above the leaf surface. Due to the existence of this superstomatal air layer, the conventional convective boundary layer is not in direct contact with the surface. Thus, in terms of plant physiology, the diffusive "end correction" to the stomatal resistance should be included in the boundary layer resistance. This is true for laminar as well as turbulent flows. When the surface mole fraction of an exchanged gas is estimated using the boundary layer resistance ignoring the diffusive term may lead to a noticeable error. The self-consistent approach is used to clarify the problems of the boundary layer formation and stomatal interference. If the correction is taken into account, the boundary layer resistance becomes dependent also on stomatal shape and distribution on the leaf. The traditional semiempirical formula corrected by the superstomatal diffusion is applied in numerical calculations. In estimates of the water vapour mole fraction on the surface of a transpiring leaf the relative error ranges from insignificant (quiescent air, large leaf and large stomatal pores) to 20 % (low humidity, strong wind, small leaf and small elliptic pores). The boundary layer resistance can decrease by a factor of 3 when the semiaxis lengths of the stomata increase from 1 and 0.5 &mgr;m to 10 and 5 &mgr;m. The effective thickness of the superstomatal air layer is maximally several millimetres (small stomatal surface concentration and small pores). Copyright 1998 Academic Press Limited