ALBERT

Notes: We conducted ecosystem carbon and water vapour exchange studies in an old-growth Pinus ponderosa forest in the Pacific North-west region of the United States. The canopy is heterogeneous, with tall multiaged trees and an open, clumped canopy with low leaf area. Carbon assimilation can occur throughout relatively mild winters, although night frosts can temporarily halt the process and physiological factors limit its efficiency. In contrast, carbon assimilation is often limited in the ‘growing season’ by stomatal closure associated with high evaporative demand (D) and soil water deficits. All of these factors present a challenge to effectively modelling ecosystem processes. Our objective was to generate an understanding of the controls on ecosystem processes across seasonal and annual cycles from a combination of fine-scale process modelling, ecophysiological measurements, and carbon and water vapour fluxes measured by the eddy covariance method. Flux measurements showed that 50% and 70% of the annual carbon uptake occurred outside the ‘growing season’ (defined as bud break to senescence, ∼ days 125–275) in 1996 and 1997. On a daily basis in summer, net ecosystem productivity (NEP) was low when D and soil water deficits were large. Whole ecosystem water vapour fluxes (LE) increased from spring to summer (1.0–1.9 mm d−1) as conducting leaf area increased by 30% and as evaporative demand increased, while evaporation from the soil surface became a smaller portion of total LE as soil water deficits increased. The models underestimated soil evaporation, particularly following rain. In the SPA model, varying the temperature optimum for photosynthesis seasonally resulted in overestimation of carbon uptake in winter and spring, showing that in coniferous forests, assumptions about temperature optima are clearly important. Daily estimates of soil surface CO2 flux from measurements and site meteorological data demonstrated that modelling of soil CO2 flux based on an Arrhenius-type equation in CANPOND overestimated CO2 respired from the soil during drought and when temperatures were low.

Notes: Synthesis of results from several Arctic and boreal research programmes provides evidence for the strong role of high-latitude ecosystems in the climate system. Average surface air temperature has increased 0.3 °C per decade during the twentieth century in the western North American Arctic and boreal forest zones. Precipitation has also increased, but changes in soil moisture are uncertain. Disturbance rates have increased in the boreal forest; for example, there has been a doubling of the area burned in North America in the past 20 years. The disturbance regime in tundra may not have changed. Tundra has a 3–6-fold higher winter albedo than boreal forest, but summer albedo and energy partitioning differ more strongly among ecosystems within either tundra or boreal forest than between these two biomes. This indicates a need to improve our understanding of vegetation dynamics within, as well as between, biomes. If regional surface warming were to continue, changes in albedo and energy absorption would likely act as a positive feedback to regional warming due to earlier melting of snow and, over the long term, the northward movement of treeline. Surface drying and a change in dominance from mosses to vascular plants would also enhance sensible heat flux and regional warming in tundra. In the boreal forest of western North America, deciduous forests have twice the albedo of conifer forests in both winter and summer, 50–80% higher evapotranspiration, and therefore only 30–50% of the sensible heat flux of conifers in summer. Therefore, a warming-induced increase in fire frequency that increased the proportion of deciduous forests in the landscape, would act as a negative feedback to regional warming.Changes in thermokarst and the aerial extent of wetlands, lakes, and ponds would alter high-latitude methane flux. There is currently a wide discrepancy among estimates of the size and direction of CO2 flux between high-latitude ecosystems and the atmosphere. These discrepancies relate more strongly to the approach and assumptions for extrapolation than to inconsistencies in the underlying data. Inverse modelling from atmospheric CO2 concentrations suggests that high latitudes are neutral or net sinks for atmospheric CO2, whereas field measurements suggest that high latitudes are neutral or a net CO2 source. Both approaches rely on assumptions that are difficult to verify. The most parsimonious explanation of the available data is that drying in tundra and disturbance in boreal forest enhance CO2 efflux. Nevertheless, many areas of both tundra and boreal forests remain net sinks due to regional variation in climate and local variation in topographically determined soil moisture. Improved understanding of the role of high-latitude ecosystems in the climate system requires a concerted research effort that focuses on geographical variation in the processes controlling land–atmosphere exchange, species composition, and ecosystem structure. Future studies must be conducted over a long enough time-period to detect and quantify ecosystem feedbacks.

Notes: Forests in the south-eastern United States experienced a prolonged dry spell and above-normal temperatures during the 1995 growing season. During this episode, nearly continuous, eddy covariance measurements of carbon dioxide and water vapour fluxes were acquired over a temperate, hardwood forest. These data are used to examine how environmental factors and accumulating soil moisture deficits affected the diurnal pattern and magnitude of canopy-scale carbon dioxide and water vapour fluxes. The field data are also used to test an integrative leaf-to-canopy scaling model (CANOAK), which uses micrometeorological and physiological theory, to calculate mass and energy fluxes. When soil moisture was ample in the spring, peak rates of net ecosystem CO2 exchange (NF) occurred around midday and exceeded 20 μmol m−2 s−1. Rates of NK were near optimal when air temperature ranged between 22 and 25°C. The accumulation of soil moisture deficits and a co-occurrence of high temperatures caused peak rates of daytime carbon dioxide uptake to occur earlier in the morning. High air temperatures and soil moisture deficits were also correlated with a dramatic reduction in the magnitude of NE. On average, the magnitude of NE decreased from 20 to 7 μmol m−2 s−1 as air temperature increased from 24 to 30°C and the soil dried. The CANAOK model yielded accurate estimates of canopy-scale carbon dioxide and water vapour fluxes when the forest had an ample supply of soil moisture. During the drought and heat spell, a cumulative drought index was needed to adjust the proportionality constant of the stomatal conductance model to yield accurate estimates of canopy CO2 exchange. The adoption of the drought index also enabled the CANOAK model to give improved estimates of evaporation until midday. On the other hand, the scheme failed to yield accurate estimates of evaporation during the afternoon.

Notes: Fluxes and concentrations of carbon dioxide and 13CO2 provide information about ecosystem physiological processes and their response to environmental variation. The biophysical model, CANOAK, was adapted to compute concentration profiles and fluxes of 13CO2 within and above a temperate deciduous forest (Walker Branch Watershed, Tennessee, USA). Modifications to the model are described and the ability of the new model (CANISOTOPE) to simulate concentration profiles of 13CO2, its flux density across the canopy–atmosphere interface and leaf-level photosynthetic discrimination against 13CO2 is demonstrated by comparison with field measurements. The model was used to investigate several aspects of carbon isotope exchange between a forest ecosystem and the atmosphere. During the 1998 growing season, the mean photosynthetic discrimination against 13CO2, by the deciduous forest canopy (Δcanopy), was computed to be 22·4‰, but it varied between 18 and 27‰. On a diurnal basis, the greatest discrimination occurred during the early morning and late afternoon. On a seasonal time scale, the greatest diurnal range in Δcanopy occurred early and late in the growing season. Diurnal and seasonal variations in Δcanopy resulted from a strong dependence of Δcanopy on photosynthetically active radiation and vapour pressure deficit of air. Model calculations also revealed that the relationship between canopy-scale water use efficiency (CO2 assimilation/transpiration) and Δcanopy was positive due to complex feedbacks among fluxes, leaf temperature and vapour pressure deficit, a finding that is counter to what is predicted for leaves exposed to well-mixed environments.

Notes: Forest biomes are major reserves for terrestrial carbon, and major components of global primary productivity. The carbon balance of forests is determined by a number of component processes of carbon acquisition and carbon loss, and a small shift in the magnitude of these processes would have a large impact on the global carbon cycle. In this paper, we discuss the climatic influences on the carbon dynamics of boreal, temperate and tropical forests by presenting a new synthesis of micrometeorological, ecophysiological and forestry data, concentrating on three case-study sites. Historical changes in the carbon balance of each biome are also reviewed, and the evidence for a carbon sink in each forest biome and its likely behaviour under future global change are discussed. We conclude that there have been significant advances in determining the carbon balance of forests, but there are still critical uncertainties remaining, particularly in the behaviour of soil carbon stocks.

Notes: In order to parametrize a leaf submodel of a canopy level gas-exchange model, a series of photosynthesis and stomatal conductance measurements were made on leaves of white oak (Quercus alba L.) and red maple (Acer rubrum L.) in a mature deciduous forest near Oak Ridge, TN. Gas-exchange characteristics of sun leaves growing at the top of a 30 m canopy and of shade leaves growing at a depth of 3–4 m from the top of the canopy were determined. Measured rates of net photosynthesis at a leaf temperature of 30°C and saturating photosynthetic photon flux density, expressed on a leaf area basis, were significantly lower (P = 0.01; n = 8) in shade leaves (7.9μmol m−2 s−1) than in sun leaves (11–5μmol m−2 s−1). Specific leaf area increased significantly with depth in the canopy, and when photosynthesis rates were expressed on a dry mass basis, they were not significantly different for shade and sun leaves. The percentage leaf nitrogen did not vary significantly with height in the canopy; thus, rates expressed on a per unit nitrogen basis were also not significantly different in shade and sun leaves. A widely used model integrating photosynthesis and stomatal conductance was parametrized independently for sun and shade leaves, enabling us to model successfully diurnal variations in photosynthesis and evapotranspiration of both classes of leaves. Key photosynthesis model parameters were found to scale with leaf nitrogen levels. The leaf model parametrizations were then incorporated into a canopy-scale gas-exchange model that is discussed and tested in a companion paper (Baldocchi & Harley 1995, Plant, Cell and Environment 18, 1157–1173).

Notes: The scaling of CO2 and water vapour transfer from leaf to canopy dimensions was achieved by integrating mechanistic models for physiological (photosynthesis, stomatal conductance and soil/root and bole respiration) and micrometeorological (radiative transfer, turbulent transfer and surface energy exchanges) processes. The main objectives of this paper are to describe a canopy photosynthesis and evaporation model for a temperate broadleaf forest and to test it against field measurements. The other goal of this paper is to use the validated model to address some contemporary ecological and physiological questions concerning the transfer of carbon and water between forest canopies and the atmosphere. In particular, we examine the role of simple versus complex radiative transfer models and the effect of environmental (solar radiation and CO2) and ecophysiological (photo-synthetic capacity) variables on canopy-scale carbon and water vapour fluxes.