Is Net Ecosystem Production Equal to Ecosystem Carbon Accumulation?

Net ecosystem production (NEP), defined as the difference between gross primary production and total ecosystem respiration, represents the total amount of organic carbon in an ecosystem available for storage, export as organic carbon, or nonbiological oxidation to carbon dioxide through fire or ultraviolet oxidation. In some of the recent literature, especially that on terrestrial ecosystems, NEP has been redefined as the rate of organic carbon accumulation in the system. Here we argue that retaining the original definition maintains the conceptual coherence between NEP and net primary production and that it is congruous with the widely accepted definitions of ecosystem autotrophy and heterotrophy. Careful evaluation of NEP highlights the various potential fates of nonrespired carbon in an ecosystem.     

Terrestrial Subsidies of Organic Carbon Support Net Ecosystem Production in Temporary Forest Ponds: Evidence from an Ecosystem Experiment

Recent research suggests that secondary production in aquatic systems can be driven by inputs of energy from terrestrial sources. Temporary forest ponds appear to be unproductive ecosystems that are reliant upon allochthonous inputs of energy to support secondary production, but the functioning of these systems has not been well quantified. To assess the metabolic state of this type of ecosystem as well as to quantify the importance of terrestrial subsidies of carbon to ecosystem function, we conducted an experiment in which we manipulated the amount of leaf litter in ponds. Litter was either removed or removed and replaced (that is, control) from the dry basins of ponds immediately after leaf abscission. Once the ponds filled, we monitored net ecosystem production (NEP) on a biweekly basis from 9 April to 27 May 2002. All ponds were consistently net heterotrophic; however, NEP was significantly less negative in removal ponds. Furthermore, removal ponds also had lower levels of respiration (R) and higher dissolved oxygen levels than control ponds. The removal of litter had no effect on gross primary production, indicating that the difference in NEP between treatments was driven by the change in R. Therefore, it appears that terrestrial inputs of organic carbon support heterotrophic respiration in these ponds, and that the endogenous production of carbon is insufficient to support secondary production.     

Production,Respiration, and Overall Carbon Balance in an Old-growth <Emphasis Type="Italic">Pseudotsuga</Emphasis>-<Emphasis Type="Italic">Tsuga</Emphasis> Forest Ecosystem

Ground-based measurements of stores, growth, mortality, litterfall, respiration, and decomposition were conducted in an old-growth forest at Wind River Experimental Forest, Washington, USA. These measurements were used to estimate gross primary production (GPP) and net primary production (NPP); autotrophic respiration (R_a) and heterotrophic (R_h) respiration; and net ecosystem production (NEP). Monte Carlo methods were used to calculate uncertainty (expressed as ± 2 standard deviations of 200–400 calculations). Live carbon (C) stores were 39,800 g C m^–2 (34,800–44,800 g C m^–2). The store of C in detritus and mineral soil was 22,092 g C m^–2 (20,600–23,600 g C m^–2), and the total C stores were 61,899 g C m^–2 (56,600–67,700 g C m^–2). Total NPP was 597 g C m^–2 y^–1 (453 to 741 g C m^–2 y^–1). R_a was 1309 g C m^–2 y^–1 (845–1773 g C m^–2 y^–1), indicating a GPP of 1906 g C m^–2 y^–1 (1444–2368 g C m^–2 y^–1). R_h, including the respiration of heart rots in tree boles, was 577 g C m^–2 y^–1 (479–675 g C m^–2 y^–1). Long-term NEP was estimated to be +20 g C m^–2 y^–1 (–116 to +156 g C m^–2 y^–1), indicating this stand might be a small sink. These estimates contrast with the larger sink estimated at the same site using eddy-flux methods. Several hypotheses to explain this discrepancy were explored, including (a) undetected biomass increases, (b) underestimates of NPP, (c) unmeasured losses, and (d) a temporal mismatch between the two sets of measurements. The last hypothesis appears the most likely.     

Above- and Belowground Net Primary Production in a Temperate Mixed Deciduous Forest

Our current ability to detect and predict changes in forest ecosystem productivity is constrained by several limitations. These include a poor understanding of belowground productivity, the short duration of most analyses, and a need for greater examination of species- or community-specific variability in productivity studies. We quantified aboveground net primary productivity (ANPP) over 3 years (1999–2001), and both belowground NPP (BNPP) and total NPP over 2 years (2000–2001) in both mesic and xeric site community types of the mixed mesophytic forest of southeastern Kentucky to examine landscape variability in productivity and its relation with soil resource [water and nitrogen (N)] availability. Across sites, ANPP was significantly correlated with N availability (R² = 0.58, P = 0.028) while BNPP was best predicted by soil moisture content (R² = 0.72, P = 0.008). Because of these offsetting patterns, total NPP was unrelated to either soil resource. Interannual variability in growing season precipitation during the study resulted in a 50% decline in mesic site litter production, possibly due to a lag effect following a moderate drought year in 1999. As a result, ANPP in mesic sites declined 27% in 2000 compared to 1999, while xeric sites had no aboveground production differences related to precipitation variability. If global climate change produces more frequent occurrences of drought, then the response of mesic sites to prolonged moisture deficiency and the consequences of shifting carbon (C) allocation on C storage will become important questions.     

Annual balances and extended seasonal modelling of carbon fluxes from a temperate fen cropped to festulolium and tall fescue under two‐cut and three‐cut harvesting regimes

This study reports the annual carbon balance of a drained riparian fen under two‐cut or three‐cut managements of festulolium and tall fescue. CO₂ fluxes measured with closed chambers were partitioned into gross primary production (GPP) and ecosystem respiration (ER) for modelling according to environmental factors (light and temperature) and canopy reflectance (ratio vegetation index, RVI). Methodological assessments were made of (i) GPP models with or without temperature functions (Ft) to adjust GPP constraints imposed by low temperature (<10 °C) and (ii) ER models with RVI or GPP parameters as biomass proxies. The sensitivity of the models was also tested on partial datasets including only alternate measurement campaigns and on datasets only from the crop growing period. Use of Ft in GPP models effectively corrected GPP overestimation in cold periods, and this approach was used throughout. Annual fluxes obtained with ER models including RVI or GPP parameters were similar, and also annual GPP and ER fluxes obtained with full and partial datasets were similar. Annual CO₂ fluxes and biomass yield were not significantly different in the crop/management combinations although the individual collars (n = 12) showed some variations in GPP (?1818 to ?2409 g CO₂‐C m^?2), ER (1071 to 1738 g CO₂‐C m^?2), net ecosystem exchange (NEE, ?669 to ?949 g CO₂‐C m^?2) and biomass yield (556 to 1044 g CO₂‐C m^?2). Net ecosystem carbon balance (NECB), as the sum of NEE and biomass carbon export, was only slightly negative to positive in all crop/management combinations. NECBs, interpreted as emission factors, tended to favour the least biomass producing systems as the best management options in relation to climate saving carbon balances. Yet, considering the down‐stream advantages of biomass for fossil fuel replacement, yield‐scaled carbon fluxes are suggested to be given additional considerations for comparison of management options in terms of atmospheric impact.     

Postfire carbon pools and fluxes in semiarid ponderosa pine in Central Oregon

Forest fire dramatically affects the carbon storage and underlying mechanisms that control the carbon balance of recovering ecosystems. In western North America where fire extent has increased in recent years, we measured carbon pools and fluxes in moderately and severely burned forest stands 2 years after a fire to determine the controls on net ecosystem productivity (NEP) and make comparisons with unburned stands in the same region. Total ecosystem carbon in soil and live and dead pools in the burned stands was on average 66% that of unburned stands (11.0 and 16.5 kg C m⁻², respectively, P<0.01). Soil carbon accounted for 56% and 43% of the carbon pools in burned and unburned stands. NEP was significantly lower in severely burned compared with unburned stands (P<0.01) with an increasing trend from −125±44 g C m⁻² yr⁻¹ (±1 SD) in severely burned stands (stand replacing fire), to −38±96 and +50±47 g C m⁻² yr⁻¹ in moderately burned and unburned stands, respectively. Fire of moderate severity killed 82% of trees <20 cm in diameter (diameter at 1.3 m height, DBH); however, this size class only contributed 22% of prefire estimates of bole wood production. Larger trees (> 20 cm DBH) suffered only 34% mortality under moderate severity fire and contributed to 91% of postfire bole wood production. Growth rates of trees that survived the fire were comparable with their prefire rates. Net primary production NPP (g C m⁻² yr⁻¹, ±1 SD) of severely burned stands was 47% of unburned stands (167±76, 346±148, respectively, P<0.05), with forb and grass aboveground NPP accounting for 74% and 4% of total aboveground NPP, respectively. Based on continuous seasonal measurements of soil respiration in a severely burned stand, in areas kept free of ground vegetation, soil heterotrophic respiration accounted for 56% of total soil CO₂ efflux, comparable with the values of 54% and 49% previously reported for two of the unburned forest stands. Estimates of total ecosystem heterotrophic respiration (R_h) were not significantly different between stand types 2 years after fire. The ratio NPP/R_h averaged 0.55, 0.85 and 1.21 in the severely burned, moderately burned and unburned stands, respectively. Annual soil CO₂ efflux was linearly related to aboveground net primary productivity (ANPP) with an increase in soil CO₂ efflux of 1.48 g C yr⁻¹ for every 1 g increase in ANPP (P<0.01, r²= 0.76). There was no significant difference in this relationship between the recently burned and unburned stands. Contrary to expectations that the magnitude of NEP 2 years postfire would be principally driven by the sudden increase in detrital pools and increased rates of R_h, the data suggest NPP was more important in determining postfire NEP.     

An initial intercomparison of micrometeorological and ecological inventory estimates of carbon exchange in a mid-latitude deciduous forest

The role of mid‐latitude forests in the sequestration of carbon (C) is of interest to an increasing number of scientists and policy‐makers alike. Net CO₂ exchange can be estimated on an annual basis, using eddy‐covariance techniques or from ecological inventories of C fluxes to and from a forest. Here we present an intercomparison of annual estimates of C exchange in a mixed hardwood forest in the Morgan‐Monroe State Forest, Indiana, USA for two years, 1998 and 1999. Based on eddy‐covariance measurements made at 1.8 times canopy height from a tower, C uptake by the forest was 237 and 287 g C m^?2 y^?1 for 1998 and 1999, respectively. For the same time period, biometric and ecophysiological measures and modelled estimates of all significant carbon fluxes within deciduous forests were made, including: change in living biomass, aboveground and belowground detritus production, foliage consumption, and forest floor and soil respiration. Using this ecological inventory method for these same two time periods, C uptake was estimated to be 271 and 377 g C m^?2 y^?1, which are 14.3% and 31.4% larger, respectively, than the tower‐based values. The relative change between this method's annual estimates is consistent with that of the eddy‐covariance based values. Our results indicate that the difference in annual C exchange rates was due to reduced heterotrophic soil respiration in 1999.     

Primary production and carbon allocation in relation to nutrient supply in a tropical experimental forest

Nutrient supply commonly limits aboveground plant productivity in forests, but the effects of an altered nutrient supply on gross primary production (GPP) and patterns of carbon (C) allocation remain poorly characterized. Increased nutrient supply may lead to a higher aboveground net primary production (ANPP), but a lower total belowground carbon allocation (TBCA), with little change in either aboveground plant respiration (APR) or GPP. Alternatively, increases in nutrient supply may increase GPP, with the quantity of GPP allocated aboveground increasing more steeply than the quantity of GPP allocated belowground. To examine the effects of an elevated nutrient supply on the C allocation patterns in forests, we determined whole‐ecosystem C budgets in unfertilized plots of Eucalyptus saligna and in adjacent plots receiving regular additions of 65 kg N ha^?1, 31 kg P ha^?1, 46 kg K ha^?1, and macro‐ and micronutrients. We measured the absolute flux of C allocated to the components of GPP (ANPP, TBCA and APR), as well as the fraction of GPP allocated to these components. Fertilization dramatically increased GPP. Averaged over 3 years, GPP in the fertilized plots was 34% higher than that in the unfertilized controls (3.95 vs. 2.95 kg C m^?2 yr^?1). Fertilization‐related increases in GPP were allocated entirely aboveground – ANPP was 85% higher and APR was 57% higher in the fertilized than in the control plots, while TBCA did not differ significantly between treatments. Carbon use efficiency (NPP/GPP) was slightly higher in the fertilized (0.53) compared with the control plots (0.51). Overall, fertilization increased ANPP and APR, and these increases were related to a greater GPP and an increase in the fraction of GPP allocated aboveground.     

Using Light-Use and Production Efficiency Models to Predict Photosynthesis and Net Carbon Exchange During Forest Canopy Disturbance

Vegetation growth models are used with remotely sensed and meteorological data to monitor terrestrial carbon dynamics at a range of spatial and temporal scales. Many of these models are based on a light-use efficiency equation and two-component model of whole-plant growth and maintenance respiration that have been parameterized for distinct vegetation types and biomes. This study was designed to assess the robustness of these parameters for predicting interannual plant growth and carbon exchange, and more specifically to address inconsistencies that may arise during forest disturbances and the loss of canopy foliage. A model based on the MODIS MOD17 algorithm was parameterized for a mature upland hardwood forest by inverting CO₂ flux tower observations during years when the canopy was not disturbed. This model was used to make predictions during a year when the canopy was 37% defoliated by forest tent caterpillars. Predictions improved after algorithms were modified to scale for the effects of diffuse radiation and loss of leaf area. Photosynthesis and respiration model parameters were found to be robust at daily and annual time scales regardless of canopy disturbance, and differences between modeled net ecosystem production and tower net ecosystem exchange were only approximately 2 g C m⁻² d⁻¹ and less than 23 g C m⁻² y⁻¹. Canopy disturbance events such as insect defoliations are common in temperate forests of North America, and failure to account for cyclical outbreaks of forest tent caterpillars in this stand could add an uncertainty of approximately 4–13% in long-term predictions of carbon sequestration.     

Age-Dependent Changes in Ecosystem Carbon Fluxes in Managed Forests in Northern Wisconsin,USA

The age-dependent variability of ecosystem carbon (C) fluxes was assessed by measuring the net ecosystem exchange of C (NEE) in five managed forest stands in northern Wisconsin, USA. The study sites ranged in age from 3-year-old clearcut to mature stands (65 years). All stands, except the clearcut, accumulated C over the study period from May to October 2002. Seasonal NEE estimates were −655 ± 17.5 g C m^–2 in the mature hardwood (MHW), −648 ± 16.8 in the mature red pine (MRP), −195 ± 15.6 in the pine barrens (PB), +128 ± 17.1 in the young hardwood clearcut (YHW), and −313 ± 14.6 in the young red pine (YRP). The age-dependent differences were similar in the hardwood and conifer forests. Even though PB was not part of either the hardwood or conifer chronosequence, and had a different disturbance agent, it still fits the same general age relationship. Higher ecosystem respiration (ER) in the young than in the mature stands was the combined result of earlier soil warming in spring, and higher temperature and greater biological activity in summer, as indicated by temperature-normalized respiration rates. The fire-generated PB had lower ER than the harvest-generated YHW and YRP, where high ER was sustained partly on account of logging residue. During the main growing season, the equivalent of 31 (MHW), 48 (MRP), 68 (PB), 114 (YHW) and 71% (YRP) of daily gross ecosystem production (GEP) was released in ER during the same day. The lower ER:GEP ratio in the mature stands was driven by greater age-dependent changes in ER than GEP. The magnitude of the increase in ER:GEP ratio in spring and fall was interpreted as the extent of the decoupling of ER and GEP. Decoupling (sustained high ER despite decreasing GEP) was observed in YHW, PB and MHW, whereas in coniferous stands (MRP and YRP) the stable ER:GEP ratio suggested preferential use of new photosynthates in ER. The results indicate that a great part of the variation in landscape-level C fluxes can be accounted for by mean stand age and associated parameters, which highlights the need to consider this source of heterogeneity in regional C balance estimates.     

Net Ecosystem Exchange of Carbon dioxide in a Temperate Poor Fen: a Comparison of Automated and Manual Chamber Techniques

We used five analytical approaches to compare net ecosystem exchange (NEE) of carbon dioxide (CO₂) from automated and manual static chambers in a peatland, and found the methods comparable. Once per week we sampled manually from 10 collars with a closed chamber system using a LiCor 6200 portable photosynthesis system, and simulated four photosynthetically active radiation (PAR) levels using shrouds. Ten automated chambers sampled CO₂ flux every 3 h with a LiCor 6252 infrared gas analyzer. Results of the five comparisons showed (1) NEE measurements made from May to August, 2001 by the manual and automated chambers had similar ranges: −10.8 to 12.7 μmol CO₂ m⁻² s⁻¹ and −17.2 to 13.1 μmol CO₂ m⁻² s⁻¹, respectively. (2) When sorted into four PAR regimes and adjusted for temperature (respiration was measured under different temperature regimes), mean NEE did not differ significantly between the chambers (p < 0.05). (3) Chambers were not significantly different in regression of ln( − respiration) on temperature. (4) But differences were found in the PAR vs. NEE relationship with manual chambers providing higher maximum gross photosynthesis estimates (GP_max), and slower uptake of CO₂ at low PAR (α) even after temperature adjustment. (5) Due to the high variability in chamber characteristics, we developed an equation that includes foliar biomass, water table, temperature, and PAR, to more directly compare automated and manual NEE. Comparing fitted parameters did not identify new differences between the chambers. These complementary chamber techniques offer a unique opportunity to assess the variability and uncertainty in CO₂ flux measurements.     

Seasonal and interannual variations in carbon dioxide exchange over a cropland in the North China Plain

In China, croplands account for a relatively large form of vegetation cover. Quantifying carbon dioxide exchange and understanding the environmental controls on carbon fluxes over croplands are critical in understanding regional carbon budgets and ecosystem behaviors. In this study, the net ecosystem exchange (NEE) at a winter wheat/summer maize rotation cropping site, representative of the main cropping system in the North China Plain, was continuously measured using the eddy covariance technique from 2005 to 2009. In order to interpret the abiotic factors regulating NEE, NEE was partitioned into gross primary production (GPP) and ecosystem respiration (R_eco). Daytime R_eco was extrapolated from the relationship between nighttime NEE and soil temperature under high turbulent conditions. GPP was then estimated by subtracting daytime NEE from the daytime estimates of R_eco. Results show that the seasonal patterns of the temperature responses of R_eco and light‐response parameters are closely related to the crop phenology. Daily R_eco was highly dependent on both daily GPP and air temperature. Interannual variability showed that GPP and R_eco were mainly controlled by temperature. Water availability also exerted a limit on R_eco. The annual NEE was ?585 and ?533 g C m^?2 for two seasons of 2006–2007 and 2007–2008, respectively, and the wheat field absorbed more carbon than the maize field. Thus, we concluded that this cropland was a strong carbon sink. However, when the grain harvest was taken into account, the wheat field was diminished into a weak carbon sink, whereas the maize field was converted into a weak carbon source. The observations showed that severe drought occurring during winter did not reduce wheat yield (or integrated NEE) when sufficient irrigation was carried out during spring.     

The annual carbon budget of a French pine forest (Pinus pinaster) following harvest

Eddy covariance measurements of net ecosystem exchange (NEE) of carbon dioxide and sensible and latent heat have operated since clear felling of a 50‐year old maritime pine stand in Les Landes, in Southwestern France. Turbulent fluxes from the closed‐path system are computed via different methodologies, including those recommended from EUROFLUX (Adv. Ecol. Res. 30 (2000) 113; Agric. Forest Meteorol. 107 (2001a, b) 43 and 71), and sensitivity analysis demonstrates the merit of post‐processing for accurate flux calculation. Footprint modeling, energy balance closure, and empirical modeling corroborate the eddy flux measurements, indicating best reliability in the daytime. The ecosystem, a net source of atmospheric CO₂, is capable of fixing carbon during fair weather during any season due to the abundance of re‐growing species (mostly grass), formerly from the understorey. Annual carbon loss of 200–340 g m^?2 depends on the period chosen, with inter‐annual variability evident during the 18‐month measurement period and apparently related to available light. Empirical models, with weekly photosynthetic parameters corresponding to seasonal vegetation and respiration depending on soil temperature, fit the data well and allow partitioning of annual NEE into GPP and TER components. Comparison with a similar nearby mature forest (Agric. Forest Meteorol. 108 (2001) 183) indicates that clear‐cutting reduces GPP by two thirds but TER by only one third, transforming a strong forest sink into a source of CO₂. Likewise, the loss of 50% of evapotranspiration (by the trees) leads to increased temperatures and thus reduced net radiation (by one third), and a 50% increase in sensible heat loss by the clear cut.     

Ecosystem responses to water and nitrogen amendment in a California grassland

The world's ecosystems are experiencing simultaneous changes in the supply of multiple limiting resources. Two of these, water and nitrogen (N) can strongly limit grassland production and can affect community composition and biogeochemical cycles in different ways. Grassland ecosystems in California may be particularly vulnerable to current and predicted changes in precipitation and N deposition, and ecosystem responses to potential interactive effects of water and N are not well understood. Here, we show strong colimitation of plant production resulting from factorial addition of water and N. In addition, water and N addition in combination led to increased dominance of the two most abundant grass species, while N addition regardless of water availability led to decreased species diversity. Late season carbon (C) flux response to water addition depended on N. Only plots that received additional water, but not N, still showed net ecosystem C uptake at the end of the experiment. Our results suggest that grassland ecosystem response to N deposition will be strongly dependent on future precipitation patterns.     

Impacts of Elevated Carbon Dioxide and Temperature on a Boreal Forest Ecosystem (CLIMEX Project)

To evaluate the effects of climate change on boreal forest ecosystems, both atmospheric CO₂ (to 560 ppmv) and air temperature (by 3°–5°C above ambient) were increased at a forested headwater catchment in southern Norway. The entire catchment (860 m²) is enclosed within a transparent greenhouse, and the upper 20% of the catchment area is partitioned such that it receives no climate treatment and serves as an untreated control. Both the control and treatment areas inside the greenhouse receive deacidified rain. Within 3 years, soil nitrogen (N) mineralization has increased and the growing season has been prolonged relative to the control area. This has helped to sustain an increase in plant growth relative to the control and has also promoted increased N export in stream water. Photosynthetic capacity and carbon–nitrogen ratio of new leaves of most plant species did not change. While the ecosystem now loses N, the long-term fate of soil N is a key uncertainty in predicting the future response of boreal ecosystems to climate change. Received 18 November 1997; accepted 13 April 1998.     

Fine-root respiration in a loblolly pine (Pinus taeda L.) forest exposed to elevated CO2 and N fertilization

Forest ecosystems release large amounts of carbon to the atmosphere from fine-root respiration (R(r)), but the control of this flux and its temperature sensitivity (Q(10)) are poorly understood. We attempted to: (1) identify the factors limiting this flux using additions of glucose and an electron transport uncoupler (carbonyl cyanide m-chlorophenylhydrazone); and (2) improve yearly estimates of R(r) by directly measuring its Q(10)in situ using temperature-controlled cuvettes buried around intact, attached roots. The proximal limits of R(r) of loblolly pine (Pinus taeda L.) trees exposed to free-air CO(2) enrichment (FACE) and N fertilization were seasonally variable; enzyme capacity limited R(r) in the winter, and a combination of substrate supply and adenylate availability limited R(r) in summer months. The limiting factors of R(r) were not affected by elevated CO(2) or N fertilization. Elevated CO(2 )increased annual stand-level R(r) by 34% whereas the combination of elevated CO(2) and N fertilization reduced R(r) by 40%. Measurements of in situ R(r) with high temporal resolution detected diel patterns that were correlated with canopy photosynthesis with a lag of 1 d or less as measured by eddy covariance, indicating a dynamic link between canopy photosynthesis and root respiration. These results suggest that R(r) is coupled to daily canopy photosynthesis and increases with carbon allocation below ground.     

Sources of Variability in Net Primary Production Predictions at a Regional Scale: A Comparison Using PnET-II and TEM 4.0 in Northeastern US Forests

Because model predictions at continental and global scales are necessarily based on broad characterizations of vegetation, soils, and climate, estimates of carbon stocks and fluxes made by global terrestrial biosphere models may not be accurate for every region. At the regional scale, we suggest that attention can be focused more clearly on understanding the relative strengths of predicted net primary productivity (NPP) limitation by energy, water, and nutrients. We evaluate the sources of variability among model predictions of NPP with a regional-scale comparison between estimates made by PnET-II (a forest ecosystem process model previously applied to the northeastern region) and TEM 4.0 (a terrestrial biosphere model typically applied to the globe) for the northeastern US. When the same climate, vegetation, and soil data sets were used to drive both models, regional average NPP predictions made by PnET-II and TEM were remarkably similar, and at the biome level, model predictions agreed fairly well with NPP estimates developed from field measurements. However, TEM 4.0 predictions were more sensitive to regional variations in temperature as a result of feedbacks between temperature and belowground N availability. In PnET-II, the direct link between transpiration and photosynthesis caused substantial water stress in hardwood and pine forest types with increases in solar radiation; predicted water stress was relieved substantially when soil water holding capacity (WHC) was increased. Increasing soil WHC had little effect on TEM 4.0 predictions because soil water storage was already sufficient to meet plant demand with baseline WHC values, and because predicted N availability under baseline conditions in this region was not limited by water. Because NPP predictions were closely keyed to forest cover type, the relative coverage of low- versus high-productivity forests at both fine and coarse resolutions was an important determinant of regional NPP predictions. Therefore, changes in grid cell size and differences in the methods used to aggregate from fine to coarse resolution were important to NPP predictions insofar as they changed the relative proportions of forest cover. We suggest that because the small patches of high-elevation spruce-fir forest in this region are substantially less productive than forests in the remainder of the region, more accurate NPP predictions will result if models applied to this region use land cover input data sets that retain as much fine-resolution forest type variability as possible. The differences among model responses to variations in climate and soil WHC data sets suggest that the models will respond quite differently to scenarios of future climate. A better understanding of the dynamic interactions between water stress, N availability, and forest productivity in this region will enable models to make more accurate predictions of future carbon stocks and fluxes. Received 19 June 1998; accepted 25 June 1999.     

Partitioning of the net CO2 exchange using an automated chamber system reveals plant phenology as key control of production and respiration fluxes in a boreal peatland

The net ecosystem CO₂ exchange (NEE) drives the carbon (C) sink–source strength of northern peatlands. Since NEE represents a balance between various production and respiration fluxes, accurate predictions of its response to global changes require an in depth understanding of these underlying processes. Currently, however, detailed information of the temporal dynamics as well as the separate biotic and abiotic controls of the NEE component fluxes is lacking in peatland ecosystems. In this study, we address this knowledge gap by using an automated chamber system established across natural and trenching/vegetation removal plots to partition NEE into its production (i.e., gross and net primary production; GPP and NPP) and respiration (i.e., ecosystem, heterotrophic and autotrophic respiration; ER, Rh and Ra) fluxes in a boreal peatland in northern Sweden. Our results showed that daily NEE patterns were driven by GPP while variations in ER were governed by Ra rather than Rh. Moreover, we observed pronounced seasonal shifts in the Ra/Rh and above/belowground NPP ratios throughout the main phenological phases. Generalized linear model analysis revealed that the greenness index derived from digital images (as a proxy for plant phenology) was the strongest control of NEE, GPP and NPP while explaining considerable fractions also in the variations of ER and Ra. In addition, our data exposed greater temperature sensitivity of NPP compared to Rh resulting in enhanced C sequestration with increasing temperature. Overall, our study suggests that the temporal patterns in NEE and its component fluxes are tightly coupled to vegetation dynamics in boreal peatlands and thus challenges previous studies that commonly identify abiotic factors as key drivers. These findings further emphasize the need for integrating detailed information on plant phenology into process‐based models to improve predictions of global change impacts on the peatland C cycle.     

Landscape Patterns of Sapling Density,Leaf Area,and Aboveground Net Primary Production in Postfire Lodgepole Pine Forests,Yellowstone National Park (USA)

Causes and implications of spatial variability in postfire tree density and understory plant cover for patterns of aboveground net primary production (ANPP) and leaf area index (LAI) were examined in ninety 11-year-old lodgepole pine (Pinus contorta var. latifolia Engelm.) stands across the landscape of Yellowstone National Park (YNP), Wyoming, USA. Field studies and aerial photography were used to address three questions: (1) What is the range and spatial pattern of lodgepole pine sapling density across the burned Yellowstone landscape and what factors best explain this variability? (2) How do ANPP and LAI vary across the landscape and is their variation explained by abiotic factors, sapling density, or both? (3) What is the predicted spatial pattern of ANPP and LAI across the burned Yellowstone landscape? Stand density spanned six orders of magnitude, ranging from zero to 535,000 saplings ha^?1, and it decreased with increasing elevation and with increasing distance from unburned forest (r ²?=?0.37). Postfire densities mapped from 1:30,000 aerial photography revealed that 66% of the burned area had densities less than 5000 saplings ha^?1 and approximately 25% had densities greater than 10,000 saplings ha^?1; stand density varied spatially in a fine-grained mosaic. New allometric equations were developed to predict aboveground biomass, ANPP, and LAI of lodgepole pine saplings and the 25 most common herbaceous and shrub species in the burned forests. These allometrics were then used with field data on sapling size, sapling density, and percent cover of graminoid, forb, and shrub species to compute stand-level ANPP and LAI. Total ANPP averaged 2.8 Mg ha^?1y^?1 but ranged from 0.04 to 15.12 Mg ha^?1y^?1. Total LAI averaged 0.80 m² m^?2 and ranged from 0.01 to 6.87 m² m^?2. Variation in ANPP and LAI was explained by both sapling density and abiotic factors (elevation and soil class) (ANOVA, r ²?=?0.80); abiotic variables explained 51%–54% of this variation. The proportion of total ANPP contributed by herbaceous plants and shrubs declined sharply with increasing sapling density (r ²?=?0.72) and increased with elevation (r ²?=?0.36). However, total herbaceous productivity was always less than 2.7 Mg ha^?1 y^?1, and herbaceous productivity did not compensate for tree production when trees were sparse. When extrapolated to the landscape, 68% of the burned landscape was characterized by ANPP values less than 2.0 Mg ha^?1y^?1, 22% by values ranging from 2 to 4 Mg ha^?1y^?1, and the remaining 10% by values greater than 4 Mg ha^?1y^?1. The spatial patterns of ANPP and LAI were less heterogeneous than patterns of sapling density but still showed fine-grained variation in rates. For some ecosystem processes, postfire spatial heterogeneity within a successional stage may be similar in magnitude to the temporal variation observed through succession.