John Tenhunen

Energy balance closure at FLUXNET sites

A comprehensive evaluation of energy balance closure is performed across 22 sites and 50 site-years in FLUXNET, a network of eddy covariance sites measuring long-term carbon and energy fluxes in contrasting ecosystems and climates. Energy balance closure was evaluated by statistical regression of turbulent energy fluxes (sensible and latent heat (LE)) against available energy (net radiation, less the energy stored) and by solving for the energy balance ratio, the ratio of turbulent energy fluxes to available energy. These methods indicate a general lack of closure at most sites, with a mean imbalance in the order of 20%. The imbalance was prevalent in all measured vegetation types and in climates ranging from Mediterranean to temperate and arctic. There were no clear differences between sites using open and closed path infrared gas analyzers. At a majority of sites closure improved with turbulent intensity (friction velocity), but lack of total closure was still prevalent under most conditions. The imbalance was greatest during nocturnal periods. The results suggest that estimates of the scalar turbulent fluxes of sensible and LE are underestimated and/or that available energy is overestimated. The implications on interpreting long-term CO2 fluxes at FLUXNET sites depends on whether the imbalance results primarily from general errors associated

Energy Partitioning Between Latent And Sensible Heat Flux During The Warm Season At Fluxnet Sites

The warm season (mid-June through late August) partitioning between sensible (H) and latent (LE) heat flux, or the Bowen ratio (beta=H/LE), was investigated at 27 sites over 66 site years within the international network of eddy covariance sites (FLUXNET). Variability in beta across ecosystems and climates was analyzed by quantifying general climatic and surface characteristics that control flux partitioning. The climatic control on beta was quantified using the climatological resistance (R-i), which is proportional to the ratio of vapor pressure deficit (difference between saturation vapor pressure and atmospheric vapor pressure) to net radiation (large values of R-i decrease beta). The control of flux partitioning by the vegetation and underlying surface was quantified by computing the surface resistance to water vapor transport (R-c, with large values tending to increase beta). There was a considerable range in flux partitioning characteristics (R-c, R-i and beta) among sites, but it was possible to define some general differences between vegetation types and climates. Deciduous forest sites and the agricultural site had the lowest values of R-c and beta (0.25-0.50). Coniferous forests typically had a larger R-c and higher beta (typically between 0.50 and 1.00 but also much larger). However, there was notable variability in R-c and R-i between coniferous site years, most notably differences between oceanic and continental climates and sites with a distinct dry summer season (Mediterranean climate). Sites with Mediterranean climates generally had the highest net radiation, R-c, R-i, and beta. There was considerable variability in beta between grassland site years, primarily the result of interannual differences in soil water content and R-c

Explaining temporal variation in soil CO2 efflux in a mature spruce forest in Southern Germany

An open dynamic chamber system was used to measure the soil CO2 efflux intensively and continuously throughout a growing season in a mature spruce forest (Picea abies) in Southern Germany. The resulting data set contained a large amount of temporally highly resolved information on the variation in soil CO2 efflux together with environmental variables. Based on this background, the dependencies of the soil CO2 efflux rate on the controlling environmental factors were analysed in-depth. Of the abiotic factors, soil temperature alone explained 72% of the variation in the efflux rate, and including soil water content (SWC) as an additional variable increased the explained variance to about 83%. Between April and December, average rates ranged from 0.43 to 5.15 mmol CO2 m 22 s 21 (in November and July, respectively) with diurnal variations of up to 50% throughout the experiment. The variability in wind speed above the forest floor influenced the CO2 efflux rates for measuring locations with a litter layer of relatively low bulk density (and hence relatively high proportions of pore spaces). For the temporal integration of flux rates for time scales of hours to days, however, wind velocities were of no effect, reflecting the fact that wind forcing acts on the transport, but not the production of CO2 in the soil. The variation in both the magnitude of the basal respiration rate and the temperature sensitivity throughout the growing season was only moderate (coefficient of variation of 15 and 25%, respectively). Soil water limitation of the CO2 production in the soil could be best explained by a reduction in the temperature-insensitive basal respiration rate, with no discernible effect on the temperature sensitivity. Using a soil CO2 efflux model with soil temperature and SWC as driving variables, it was possible to calculate the annual soil CO2 efflux for four consecutive years for which meteorological data were available. These simulations indicate an average efflux sum of 560 g C m 22 yr 21 (SE ¼ 22 g C m 22 yr 21 ). An alternative model derived from the same data but using temperature alone as a driver over-estimated the annual flux sum by about 7% and showed less inter-annual variability. Given a likely shift in precipitation patterns alongside temperature changes under projected global change scenarios, these results demonstrate the necessity to include soil moisture in models that calculate the evolution of CO2 from temperate forest soils. q 2003 Elsevier Ltd. All rights reserved.

Phase and amplitude of ecosystem carbon release and uptake potentials as derived from FLUXNET measurements

As length and timing of the growing season are major factors explaining differences in carbon exchange of ecosystems, we analyzed seasonal patterns of net ecosystem carbon exchange (FNEE) using eddy covariance data of the FLUXNET data base (http://www-eosdis.ornl.gov/FLUXNET). The study included boreal and temperate, deciduous and coniferous forests, Mediterranean evergreen systems, rainforest, native and managed temperate grasslands, tundra, and C3 and C4 crops. Generalization of seasonal patterns are useful for identifying functional vegetation types for global dynamic vegetation models, as well as for global inversion studies, and can help improve phenological modules in SVAT or biogeochemical models. The results of this study have important validation potential for global carbon cycle modeling. The phasing of respiratory and assimilatory capacity differed within forest types: for temperate coniferous forests seasonal uptake and release capacities are in phase, for temperate deciduous and boreal coniferous forests, release was delayed compared to uptake. According to seasonal pattern of maximum nighttime release (evaluated over 15-day periods, Fmax) the study sites can be grouped in four classes: (1) boreal and high altitude conifers and grasslands; (2) temperate deciduous and temperate conifers; (3) tundra and crops; (4) evergreen Mediterranean and tropical forests. Similar results are found for maximum daytime uptake (Fmin) and the integral net carbon flux, but temperate deciduous forests fall into class 1. For forests, seasonal amplitudes of Fmax and Fmin increased in the order tropical C3-crops>temperate deciduous forests>temperate conifers>boreal conifers>tundra ecosystems. Due to data restrictions, our analysis centered mainly on Northern Hemisphere temperate and boreal forest ecosystems. Grasslands, crops, Mediterranean ecosystems, and rainforests are under-represented, as are savanna systems, wooded grassland, shrubland, or year-round measurements in tundra systems. For regional or global estimates of carbon sequestration potentials, future investigations of eddy covariance should expand in these systems.

Coordination theory of leaf nitrogen distribution in a canopy

It has long been observed that leaf nitrogen concentrations decline with depth in closed canopies in a number of plant communities. This phenomenon is generally believed to be related to a changing radiation environment and it has been suggested by some researchers that plants allocate nitrogen in order to optimize total whole canopy photosynthesis. Although optimization theory has been successfully utilized to describe a variety of physiological and ecological phenomena, it has some shortcomings that are subject to criticism (e.g., time constraints, oversimplifications, lack of insights, etc.). In this paper we present an alternative to the optimization theory of plant canopy nitrogen distribution, which we term coordination theory. We hypothesize that plants allocate nitrogen to maintain a balance between two processes, each of which is dependent on leaf nitrogen content and each of which potentially limits photosynthesis. These two processes are defined as Wc, the Rubiscolimited rate of carboxylation, and Wj, the electron transport-limited rate of carboxylation. We suggest that plants allocate nitrogen differentially to, leaves in different canopy layers in such a way that Wc and Wj remain roughly balanced. In this scheme, the driving force for the allocation of nitrogen within a canopy is the difference between the leaf nitrogen content that is required to bring Wc and Wj into balance and the current nitrogen content. We show that the daily carbon assimilation of a canopy with a nitrogen distribution resulting from this internal coordination of Wc and Wj is very similar to that obtained using optimization theory.

Water relations, gas exchange, and growth of resprouts and mature plant shoots of "Arbutus unedo L." and "Quercus ilex L."

Resprout and mature plant shoot growth, leaf water status and gas exchange behavior, tissue nutrient content, flowering, and production were studied for co-occurring shallow-rooted (Arbutus unedo L.) and deeprooted (Quercus ilex L.) Mediterranean tree species at the Collserola Natural Park in Northeast Spain Resprouts showed higher growth rates than mature plant shoots. During fall, no differences in eco-physiological performance of leaves were found, but mobilization of carbohydrates from burls strongly stimulated growth of fall resprouts compared to spring resprouts, despite low exposed leaf area of the fall shoots. During summer drought, resprouts exhibited improved water status and carbon fixation compared to mature plant shoots. Shoot growth of Q. ilex was apparently extended due to deep rooting so that initial slower growth during spring and early summer as compared to A. unedo was compensated. Tissue nutrient contents varied only slightly and are postulated to be of minor importance in controlling rate of shoot growth, perhaps due to the relatively fertile soil of the site. Fall flowering appeared to inhibit fall shoot growth in A. unedo, but did not occur in Q. ilex. The results demonstrate that comparative examinations utilizing vegetation elements with differing morphological and physiological adaptations can be used to analyze relatively complex phenomena related to resprouting behavior. The studies provide an important multi-dimensional background framework for further studies of resprouting in the European Mediterranean region.

Analyzing the ecosystem carbon dynamics of four European coniferous forests using a biogeochemistry model

AbstractThis paper provides the first steps toward a regional-scale analysis of carbon (C) budgets. We explore the ability of the ecosystem model BIOME-BGC to estimate the daily and annual C dynamics of four European coniferous forests and shifts in these dynamics in response to changing environmental conditions. We estimate uncertainties in the model results that arise from incomplete knowledge of site management history (for example, successional stage of forest). These uncertainties are especially relevant in regional-scale simulations, because this type of information is difficult to obtain. Although the model predicted daily C and water fluxes reasonably well at all sites, it seemed to have a better predictive capacity for the photosynthesis-related processes than for respiration. Leaf area index (LAI) was modeled accurately at two sites but overestimated at two others (as a result of poor long-term climate drivers and uncertainties in model parameterization). The overestimation of LAI (and consequently gross photosynthetic production (GPP)), in combination with reasonable estimates of the daily net ecosystem productivity (NEP) of those forests, also illustrates the problem with modeled respiration. The model results suggest that all four European forests have been net sinks of C at the rate of 100–300 gC/m2/y and that this C sequestration capacity would be 30%–70% lower without increasing nitrogen (N) deposition and carbon dioxide (CO2) concentrations. The magnitude of the forest responses was dependent not only on the rate of changes in environmental factors, but also on site-specific conditions such as climate and soil depth. We estimated that the modeled C exchange at the study sites was reduced by 50%–100% when model simulations were performed for climax forests rather than regrowing forests. The estimates of water fluxes were less sensitive to different initializations of state variables or environmental change scenarios than C fluxes.

Biospheric aspects of the hydrological cycle; preface

Abstract The Core Project Biospheric Aspects of the Hydrological Cycle (BAHC) of the International Geosphere Biosphere Programme (IGBP) addresses the biospheric aspects of the hydrological cycle through experiments and modelling of energy, water, carbon dioxide and sediment fluxes in the soil– vegetation–atmosphere system at a variety of spatial and temporal scales. Active regulation of water, energy and carbon dioxide fluxes by the vegetation make it an important factor in regulating the Earth’s hydrological cycle and in the formation of the climate. Consequently, human induced conversion of vegetation cover is an important driver for climate change. A number of recent studies, discussed in this paper, emphasise the importance of the terrestrial biosphere for the climate system. Initially, these studies demonstrate the influence of the land surface on tropical weather and climate, revealing the mechanisms, acting at various scales, that connect increasing temperatures and decreasing rainfall to large-scale deforestation and other forms of land degradation. More recently, the significance of the land surface processes for water cycle and for weather and climate in temperate and boreal zones was demonstrated. In addition the terrestrial biosphere plays a significant role in the carbon dioxide fluxes and in global carbon balance. Recent work suggests that many ecosystems both in the tropics and in temperate zones may act as a substantial sink for carbon dioxide, though the temporal variability of this sink strength is yet unclear. Further, carbon dioxide uptake and evaporation by vegetation are intrinsically coupled, leading to links and feedbacks between land surface and climate that are hardly explored yet. Earth’s vegetation cover and its changes owing to human impact have a profound influence on a lateral redistribution of water and transported constituents, such as nutrients and sediments, and acts therefore as an important moderator of Earth’s biogeochemical cycles. In the BAHC science programme, the importance of studying the influence of climate and human activities on mobilisation and river-borne transport of constituents is explicitly articulated. The terrestrial water and associated material cycles are studied as highly dynamic in space and time, and reflect a complex interplay among climatic forcing, topography, land cover and vegetation dynamics. Despite a large progress in our understanding of how the terrestrial biosphere interacts with Earth’s and climate system and with the terrestrial part of its hydrological cycle, a number of basic issues still remain unresolved. Limited to the scope of BAHC, the paper briefly assesses the present status and identifies the most important outstanding issues, which require further research. Two, arguably most important outstanding issues are identified: a limited understanding of natural variability, especially with respect to seasonal to inter-annual cycles, and of a complex ecosystem behaviour resulting from multiple feedbacks and multiple coupled biogeochemical cycles within the overall climate system. This leads to two major challenges for the future science agenda related to global change research. First, there is a need for a strong multidisciplinary integration of research efforts in both modelling and experiments, the latter extending to inter-annual timescales. Second, the ever increasing complexity in characterisation and modelling of the climate system, which is mainly owing to incorporation of the biosphere’s and human feedbacks, may call for a new approach in global change impact studies. Methodologies need to be developed to identify risks to, and vulnerability of environmental systems, taking into account all important interactions between atmospheric, ecological and hydrological processes at relevant scales. With respect to the influence of climate and human activities on mobilisation and river-borne transport of constituents, the main issues for the future are related to declining availability and quality of ground data for quantity and quality of water discharge. Such assessments as presented in this paper, in combination with community wide science evaluation, have lead to an update of the science agenda for BAHC, a summary of which is provided in the appendix.

Variability in Leaf Morphology and Chemical Composition as a Function of Canopy Light Environment in Coexisting Deciduous Trees

Morphology, chemical composition, and photosynthetic capacity of leaf laminas were investigated in Populus tremula L. and Tilia cordata Mill. along a canopy light gradient. Variables determining the thickness of boundary layer for heat and water exchange at a given wind speed—effective leaf width (Ww) and length (Wd)—scaled positively with daily integrated quantum flux density averaged over the season (Qint, mol m−2 d−1) in T. cordata, but Wd decreased and Ww was constant with increasing Qint in P. tremula, bringing about a moderately improved capacity for convective cooling at greater irradiances in the latter species. Foliar stable carbon isotope discrimination (Δ) decreased with increasing Qint, demonstrating that, possibly because of more severe foliar water stress, leaves operated at a lower intercellular CO2 concentration in the upper canopy. Further analysis of foliar characteristics provided additional evidence of the interaction between water stress and Qint. Leaf dry matter content and both components of lamina dry mass per area (MA)—lamina thickness and density (dry mass per unit volume, ρB)—increased with increasing Qint in both species. The ρB and lamina dry matter content were also positively related to lamina carbon concentration, variability in which along the canopy was related to changes in carbon‐rich lignin concentration. Since both increases in lamina density and lignin concentration improve leaf tolerance of low‐water potentials, these foliar modifications were interpreted as indicative of acclimation to enhanced water limitations in high light. For the whole material, foliar nitrogen concentrations decreased with increasing ρB, suggesting that an improvement of foliar mechanical strength may result in declining foliar assimilative potential. However, foliar photosynthetic electron transport capacity per unit area increased with increasing ρB, possibly because increases in ρB with light are not only attributable to greater cell wall lignification but also to denser packing of leaf cells, in particular, in fractional increases in palisade tissues with Qint. Because of a positive scaling of leaf thickness and density with total tree height, MA was greater in taller trees of T. cordata, foliage of which also had lower Δ and was likely to function with less open stomata. In summary, we conclude that leaf water stress, which scales with both Qint and total tree height, is a major factor altering foliage structure and assimilative capacity.

A model of the gas exchange response ofPicea abies to habitat conditions

Databases describing branch gas exchange ofPicea abies L. at two montane forest sites, Lägeren, Switzerland (National Forschungsprojekt 14 of the Schweizerische Nationalfonds) and Oberwarmensteinach, Germany (Bayerische Forschungsgruppe Forsttoxikologie), were analyzed in conjunction with a physiologically based model. Parameter estimates for describing carboxylase kinetics, electron transport, and stomatal function were derived, utilizing information from both single factor dependencies and diurnal time course measurements of gas exchange. Data subsets were used for testing the model at the branch level. Most of the observed variation in gas exchange characteristics can be explained with the model, while a number of systematic errors remain unexplained. Factors seen as contributing to the unexplained residual variation and not included in the model are light acclimation, degree of damage in adjustment to pollutant deposition, needle age, and cold stress effects. Nevertheless, a set of parameter values has been obtained for general application with spruce, e.g., for use in calculating canopy flux rates and to aid in planning of focused leaf and canopy level experiments. The value of the model for estimating fluxes between the forest and the atmosphere must be evaluated together with measurements at the stand level.