Abstract. Isotope pool dilution studies are increasingly reported in the soils and ecology literature as a means of measuring gross rates of nitrogen (N) mineralization, nitrification, and inorganic N assimilation in soils. We assembled data on soil characteristics and gross rates from 100 studies conducted in forest, shrubland, grassland, and agricultural systems to answer the following questions: What factors appear to be the major drivers for production and consumption of inorganic N as measured by isotope dilution studies? Do rates or the relationships between drivers and rates differ among ecosystem types? Across a wide range of ecosystems, gross N mineralization is positively correlated with microbial biomass and soil C and N concentrations, while soil C:N ratio exerts a negative effect on N mineralization only after adjusting for differences in soil C. Nitrification is a log-linear function of N mineralization, increasing rapidly at low mineralization rates but changing only slightly at high mineralization rates. In contrast, NH 4 ϩ assimilation by soil microbes increases nearly linearly over the full range of mineralization rates. As a result, nitrification is proportionately more important as a fate for NH 4 ϩ at low mineralization rates than at high mineralization rates. Gross nitrification rates show no relationship to soil pH, with some of the fastest nitrification rates occurring below pH 5 in soils with high N mineralization rates. Differences in soil organic matter (SOM) composition and concentration among ecosystem types influence the production and fate of mineralized N. Soil organic matter from grasslands appears to be inherently more productive of ammonium than SOM from wooded sites, and SOM from deciduous forests is more so than SOM in coniferous forests, but differences appear to result primarily from differing C:N ratios of organic matter. Because of the central importance of SOM characteristics and concentrations in regulating rates, soil organic matter depletion in agricultural systems appears to be an important determinant of gross process rates and the proportion of NH 4 ϩ that is nitrified. Addition of 15 N appears to stimulate NH 4 ϩ consumption more than NO 3 Ϫ consumption processes; however, the magnitude of the stimulation may provide useful information regarding the factors limiting microbial N transformations.

Ecologists have long been intrigued by the ways co-occurring species divide limiting resources. Such resource partitioning, or niche differentiation, may promote species diversity by reducing competition. Although resource partitioning is an important determinant of species diversity and composition in animal communities, its importance in structuring plant communities has been difficult to resolve. This is due mainly to difficulties in studying how plants compete for below-ground resources. Here we provide evidence from a 15N-tracer field experiment showing that plant species in a nitrogen-limited, arctic tundra community were differentiated in timing, depth and chemical form of nitrogen uptake, and that species dominance was strongly correlated with uptake of the most available soil nitrogen forms. That is, the most productive species used the most abundant nitrogen forms, and less productive species used less abundant forms. To our knowledge, this is the first documentation that the composition of a plant community is related to partitioning of differentially available forms of a single limiting resource.

Recent projections of climatic change have focused a great deal of scientific and public attention on patterns of carbon (C) cycling as well as its controls, particularly the factors that determine whether an ecosystem is a net source or sink of atmospheric carbon dioxide (CO 2 ). Net ecosystem production (NEP), a central concept in C-cycling research, has been used by scientists to represent two different concepts. We propose that NEP be restricted to just one of its two original definitions-the imbalance between gross primary production (GPP) and ecosystem respiration (ER). We further propose that a new term-net ecosystem carbon balance (NECB)-be applied to the net rate of C accumulation in (or loss from [negative sign]) ecosystems. Net ecosystem carbon balance differs from NEP when C fluxes other than C fixation and respiration occur, or when inorganic C enters or leaves in dissolved form. These fluxes include the leaching loss or lateral transfer of C from the ecosystem; the emission of volatile organic C, methane, and carbon monoxide; and the release of soot and CO 2 from fire. Carbon fluxes in addition to NEP are particularly important determinants of NECB over long time scales. However, even over short time scales, they are important in ecosystems such as streams, estuaries, wetlands, and cities. Recent technological advances have led to a diversity of approaches to the measurement of C fluxes at different temporal and spatial scales. These approaches frequently capture different components of NEP or NECB and can therefore be compared across scales only by carefully specifying the 1041 fluxes included in the measurements. By explicitly identifying the fluxes that comprise NECB and other components of the C cycle, such as net ecosystem exchange (NEE) and net biome production (NBP), we can provide a less ambiguous framework for understanding and communicating recent changes in the global C cycle.

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We use a mechanistically based ecosystem simulation model to describe and analyze the spatial and temporal patterns of terrestrial net primary productivity (NPP) in South America. The Terrestrial Ecosystem Model (TEM) is designed to predict major carbon and nitrogen fluxes and pool sizes in terrestrial ecosystems at continental to global scales. Information from intensively studies field sites is used in combination with continental-scale information on climate, soils, and vegetation to estimate NPP in each of 5888 non-wetland, 0.5° latitude °0.5° longitude grid cells in South America, at monthly time steps. Preliminary analyses are presented for the scenario of natural vegetation throughout the continent, as a prelude to evaluating human impacts on terrestrial NPP. The potential annual NPP of South America is estimated to be 12.5 Pg/yr of carbon (26.3 Pg/yr of organic matter) in a non-wetland area of 17.0 ° 10 km . More than 50% of this production occurs in the tropical and subtropical evergreen forest region. Six independent model runs, each based on an independently derived set of model parameters, generated mean annual NPP estimates for the tropical evergreen forest region ranging from 900 to 1510 g°m °yr of carbon, with an overall mean of 1170 g°m °yr . Coefficients of variation in estimated annual NPP averaged 20% for any specific location in the evergreen forests, which is probably within the confidence limits of extant NPP measurements. Predicted rates of mean annual NPP in other types of vegetation ranged from 95 g°m °yr in arid shrublands to 930 g°m ?yr in savannas, and were within the ranges measured in empirical studies. The spatial distribution of predicted NPP was directly compared with estimates made using the Miami mode of Lieth (1975). Overall, TEM predictions were °10% lower than those of the Miami model, but the two models agreed closely on the spatial patterns of NPP in south America. Unlike previous models, however, TEM estimates NPP monthly, allowing for the evaluation of seasonal phenomena. This is an important step toward integration of ecosystem models with remotely sensed information, global climate models, and atmospheric transport models, all of which are evaluated at comparable spatial and temporal scales. Seasonal patterns of NPP in South America are correlated with moisture availability in most vegetation types, but are strongly influenced by seasonal differences in cloudiness in the tropical evergreen forests. On an annual basis, moisture availability was the factor that was correlated most strongly with annual NPP in South America, but differences were again observed among vegetation types. These results allow for the investigation and analysis of climatic controls over NPP at continental scales, within and among vegetation types, and within years. Further model validation is needed. Nevertheless, the ability to investigate NPP-environment interactions with a high spatial and temporal resolution at continental scales should prove useful if not essential for rigorous analysis of th...

Quercus-Acer (oak-maple) st'.111d in c~ntral l\ l~1ssac~1u selts, US/\. The key hy pothesis J!OVe rnmg the b1oloA1cal component of the model is Urnt stomata! conductance (g,.) is var ied s o that d ail y carbon uptake per unit of foliar nit r oj:!en is maximized wit hin the limitations of ca1101>Y water avr1ilability. T he h ydraulic system is modelled as :m a nalo~ue to simple electr ical circuits in parallel. includinJ! a separate soil hydraulic resistance. plant resistance and plant capacitance for each c:m op y layer. Stomata! openin(! is initially controlled to conserve plant water stores and delay the o nset or water stress. Stomata! clos ure at :i threshold minimum l~1f water potential prevents xyle m cavitation and contr ols the maximum r:itc of wate r llux through the h ydraulic system. We show a strong correlat ion between predicted hourly C0 2 exchange rate (r2 = 0·86) and LE (r 2 = 0·87) with independent whole-forest measurements made by the eddy correlation method during the s ummer of 1992. Our theoretical derivation s hows that observed relationships between C0 2 assimilation a nd LE nux can he explained on the basis of s tomata! behaviour 01>timizing c:irhon gain, and provides mt exi>licit link hetwecn canopy structure, soil pro1>erties. atmos pheric conditions and stomatal conductance.Ke\'-111ords: Q11erc11s r11hm: l\cl'r rulnw11: soil-pla11t-a1r~10spherc coniinuum model: photosynlhcsis: plant hydr:1ulic conductance: s1o ma1al conduc tance.

Summary 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 qu...

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