Net ecosystem production of a Douglas-fir stand for 3 years following clearcut harvesting

To investigate the variations in annual and seasonal net ecosystem production (F_NEP) during the development of a young forest, 3 years of continuous eddy covariance measurements of carbon dioxide (CO₂) fluxes were collected following clearcut harvesting and replanting of a coastal Douglas‐fir stand on the east coast of Vancouver Island, BC, Canada. The impact of changing weather and stand structure on F_NEP was examined by developing relationships between F_NEP and variables such as light, temperature, soil moisture, and leaf area index (LAI). In all 3 years, the stand was a large source of CO₂ (620, 520, and 600 g C m^?2 yr^?1 in the first, second, and third years, respectively). During this period, the growth of pioneer and understory species resulted in an increase in maximum growing season LAI from 0.2 in the year the seedlings were planted to 2.5 in the third year. The associated increase in annual gross ecosystem production (P=F_NEP?R_e, where R_e is ecosystem respiration) from 220 g C m^?2 yr^?1 in the first year to 640 g C m^?2 yr^?1 in the third year was exceeded by an increase in annual R_e from 840 to 1240 g C m^?2 yr^?1. Seasonal and interannual variations in daytime F_NEP and P were well described by variations in photosynthetically active radiation, temperature, and changes in LAI. Night‐time measurements of R_e exponentially increased with 2 cm soil temperature with an average Q₁₀ of 2 (relative increase in R_e for a 10°C increase in temperature) and R₁₀ (R_e at 10°C) that increased from 2.1 in the first year to 2.5 in the second year to 3.2 μmol m^?2 s^?1 in the third year. Although the re‐establishment of vegetation in this stand had a major impact on both P and R_e, interannual variations in weather also affected annual F_NEP. Drought, in the summer of the third year, resulted in early senescence and reduced both P and R_e. This resulted in more C being lost from the stand in the third year after harvesting than in the second year.     

Climatic controls on the carbon and water balances of a boreal aspen forest, 1994–2003

The carbon and water budgets of boreal and temperate broadleaf forests are sensitive to interannual climatic variability and are likely to respond to climate change. This study analyses 9 years of eddy‐covariance data from the Boreal Ecosystem Research and Monitoring Sites (BERMS) Southern Old Aspen site in central Saskatchewan, Canada and characterizes the primary climatic controls on evapotranspiration, net ecosystem production (F_NEP), gross ecosystem photosynthesis (P) and ecosystem respiration (R). The study period was dominated by two climatic extremes: extreme warm and cool springs, which produced marked contrasts in the canopy duration, and a severe, 3‐year drought. Annual F_NEP varied among years from 55 to 367 g C m⁻² (mean 172, SD 94). Interannual variability in F_NEP was controlled primarily by factors that affected the R/P ratio, which varied between 0.74 and 0.96 (mean 0.87, SD 0.06). Canopy duration enhanced P and F_NEP with no apparent effect on R. The fraction of annual photosynthetically active radiation (PAR) that was absorbed by the canopy foliage varied from 38% in late leaf‐emergence years to 51% in early leaf‐emergence years. Photosynthetic light‐use efficiency (mean 0.0275, SD 0.026 mol C mol⁻¹ photons) was relatively constant during nondrought years but declined with drought intensity to a minimum of 0.0228 mol C mol⁻¹ photons during the most severe drought year. The impact of drought on F_NEP varied with drought intensity. Years of mild‐to‐moderate drought suppressed R while having little effect on P, so that F_NEP was enhanced. Years of severe drought suppressed both R and P, causing either little change or a subtle reduction in F_NEP. The analysis produced new insights into the dominance of canopy duration as the most important biophysical control on F_NEP. The results suggested a simple conceptual model for annual F_NEP in boreal deciduous forests. When water is not limiting, annual P is controlled by canopy duration via its influence on absorbed PAR at constant light‐use efficiency. Water stress suppresses P, by reducing light‐use efficiency, and R, by limiting growth and/or suppressing microbial respiration. The high photosynthetic light‐use efficiency showed this site to be a highly productive boreal deciduous forest, with properties similar to many temperate deciduous forests.     

Respiratory carbon use and carbon storage in mid‐rotation loblolly pine (Pinus taeda L.) plantations: the effect of site resources on the stand carbon balance

We used estimates of autotrophic respiration (R_A), net primary productivity (NPP) and soil CO₂ evolution (S_ff), to develop component carbon budgets for 12‐year‐old loblolly pine plantations during the fifth year of a fertilization and irrigation experiment. Annual carbon use in R_A was 7.5, 9.0, 15.0, and 15.1 Mg C ha^?1 in control (C), irrigated (I), fertilized (F) and irrigated and fertilized (IF) treatments, respectively. Foliage, fine root and perennial woody tissue (stem, branch, coarse and taproot) respiration accounted for, respectively, 37%, 24%, and 39% of R_A in C and I treatments and 38%, 12% and 50% of R_A in F and IF treatments. Annual gross primary production (GPP=NPP+R_A) ranged from 13.1 to 26.6 Mg C ha^?1. The I, F, and IF treatments resulted in a 21, 94, and 103% increase in GPP, respectively, compared to the C treatment. Despite large treatment differences in NPP, R_A, and carbon allocation, carbon use efficiency (CUE=NPP/GPP) averaged 0.42 and was unaffected by manipulating site resources. Ecosystem respiration (R_E), the sum of S_ff, and above ground R_A, ranged from 12.8 to 20.2 Mg C ha^?1 yr^?1. S_ff contributed the largest proportion of R_E, but the relative importance of S_ff decreased from 0.63 in C treatments to 0.47 in IF treatments because of increased aboveground R_A. Aboveground woody tissue R_A was 15% of R_E in C and I treatments compared to 25% of R_E in F and IF treatments. Net ecosystem productivity (NEP=GPP‐R_E) was roughly 0 in the C and I treatments and 6.4 Mg C ha^?1 yr^?1 in F and IF treatments, indicating that non‐fertilized treatments were neither a source nor a sink for atmospheric carbon while fertilized treatments were carbon sinks. In these young stands, NEP is tightly linked to NPP; increased ecosystem carbon storage results mainly from an increase in foliage and perennial woody biomass.     

Effects of logging on carbon dynamics of a jack pine forest in Saskatchewan,Canada

We calculated carbon budgets for a chronosequence of harvested jack pine (Pinus banksiana Lamb.) stands (0‐, 5‐, 10‐, and～29‐year‐old) and a～79‐year‐old stand that originated after wildfire. We measured total ecosystem C content (TEC), above‐, and belowground net primary productivity (NPP) for each stand. All values are reported in order for the 0‐, 5‐, 10‐, 29‐, and 79‐year‐old stands, respectively, for May 1999 through April 2000. Total annual NPP (NPP_T) for the stands (Mg C ha^?1 yr^?1±1 SD) was 0.9±0.3, 1.3±0.1, 2.7±0.6, 3.5±0.3, and 1.7±0.4. We correlated periodic soil surface CO₂ fluxes (R_S) with soil temperature to model annual R_S for the stands (Mg C ha^?1 yr^?1±1 SD) as 4.4±0.1, 2.4±0.0, 3.3±0.1, 5.7±0.3, and 3.2±0.2. We estimated net ecosystem productivity (NEP) as NPP_T minus R_H (where R_H was calculated using a Monte Carlo approach as coarse woody debris respiration plus 30–70% of total annual R_S). Excluding C losses during wood processing, NEP (Mg C ha^?1 yr^?1±1 SD) for the stands was estimated to be ?1.9±0.7, ?0.4±0.6, 0.4±0.9, 0.4±1.0, and ?0.2±0.7 (negative values indicate net sources to the atmosphere.) We also calculated NEP values from the changes in TEC among stands. Only the 0‐year‐old stand showed significantly different NEP between the two methods, suggesting a possible mismatch for the chronosequence. The spatial and methodological uncertainties allow us to say little for certain except that the stand becomes a source of C to the atmosphere following logging.     

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.     

Net ecosystem productivity of boreal jack pine stands regenerating from clearcutting under current and future climates

Life cycle analysis of climate and disturbance effects on forest net ecosystem productivity (NEP) is necessary to assess changes in forest carbon (C) stocks under current or future climates. Ecosystem models used in such assessments need to undergo well-constrained tests of their hypotheses for climate and disturbance effects on the processes that determine CO₂ exchange between forests and the atmosphere. We tested the ability of the model ecosys to simulate diurnal changes in CO₂ fluxes under changing air temperatures (T_a) and soil water contents during forest regeneration with eddy covariance measurements over boreal jack pine (Pinus banksiana) stands along a postclearcut chronosequence. Model hypotheses for hydraulic and nutrient constraints on CO₂ fixation allowed ecosys to simulate the recovery of C cycling during the transition of boreal jack pine stands from C sources following clearcutting (NEP from −150 to −200 g C m⁻² yr⁻¹) to C sinks at maturity (NEP from 20 to 80 g C m⁻² yr⁻¹) with large interannual variability. Over a 126-year logging cycle, annualized NEP, C harvest, and net biome productivity (NBP=NEP–harvest removals) of boreal jack pine averaged 47, 33 and 14 g C m⁻² yr⁻¹. Under an IPCC SRES climate change scenario, rising T_a exacerbated hydraulic constraints that adversely affected NEP of boreal jack pine after 75 years. These adverse effects were avoided in the model by replacing the boreal jack pine ecotype with one adapted to warmer T_a. This replacement raised annualized NEP, C harvest, and NBP to 81, 56 and 25 g C m⁻² yr⁻¹ during a 126-year logging cycle under the same climate change scenario.     

Modeling to discern nitrogen fertilization impacts on carbon sequestration in a Pacific Northwest Douglas‐fir forest in the first‐postfertilization year

This study investigated how nitrogen (N) fertilization with 200 kg N ha^?1 of urea affected ecosystem carbon (C) sequestration in the first‐postfertilization year in a Pacific Northwest Douglas‐fir (Pseudotsuga menziesii) stand on the basis of multiyear eddy‐covariance (EC) and soil‐chamber measurements before and after fertilization in combination with ecosystem modeling. The approach uses a data‐model fusion technique which encompasses both model parameter optimization and data assimilation and minimizes the effects of interannual climatic perturbations and focuses on the biotic and abiotic factors controlling seasonal C fluxes using a prefertilization 9‐year‐long time series of EC data (1998–2006). A process‐based ecosystem model was optimized using the half‐hourly data measured during 1998–2005, and the optimized model was validated using measurements made in 2006 and further applied to predict C fluxes for 2007 assuming the stand was not fertilized. The N fertilization effects on C sequestration were then obtained as differences between modeled (unfertilized stand) and EC or soil‐chamber measured (fertilized stand) C component fluxes. Results indicate that annual net ecosystem productivity in the first‐post‐N fertilization year increased by～83%, from 302 ± 19 to 552 ± 36 g m^?2 yr^?1, which resulted primarily from an increase in annual gross primary productivity of～8%, from 1938 ± 22 to 2095 ± 29 g m^?2 yr^?1 concurrent with a decrease in annual ecosystem respiration (R_e) of～5.7%, from 1636 ± 17 to 1543 ± 31 g m^?2 yr^?1. Moreover, with respect to respiration, model results showed that the fertilizer‐induced reduction in R_e (～93 g m^?2 yr^?1) principally resulted from the decrease in soil respiration R_s (～62 g m^?2 yr^?1).     

The legacy of harvest and fire on ecosystem carbon storage in a north temperate forest

Forest harvesting and wildfire were widespread in the upper Great Lakes region of North America during the early 20th century. We examined how long this legacy of disturbance constrains forest carbon (C) storage rates by quantifying C pools and fluxes after harvest and fire in a mixed deciduous forest chronosequence in northern lower Michigan, USA. Study plots ranged in age from 6 to 68 years and were created following experimental clear‐cut harvesting and fire disturbance. Annual C storage was estimated biometrically from measurements of wood, leaf, fine root, and woody debris mass, mass losses to herbivory, soil C content, and soil respiration. Maximum annual C storage in stands that were disturbed by harvest and fire twice was 26% less than a reference stand receiving the same disturbance only once. The mechanism for this reduction in annual C storage was a long‐lasting decrease in site quality that endured over the 62‐year timeframe examined. However, during regrowth the harvested and burned forest rapidly became a net C sink, storing 0.53 Mg C ha⁻¹ yr⁻¹ after 6 years. Maximum net ecosystem production (1.35 Mg C ha⁻¹ yr⁻¹) and annual C increment (0.95 Mg C ha⁻¹ yr⁻¹) were recorded in the 24‐ and 50‐year‐old stands, respectively. Net primary production averaged 5.19 Mg C ha⁻¹ yr⁻¹ in experimental stands, increasing by < 10% from 6 to 50 years. Soil heterotrophic respiration was more variable across stand ages, ranging from 3.85 Mg C ha⁻¹ yr⁻¹ in the 6‐year‐old stand to 4.56 Mg C ha⁻¹ yr⁻¹ in the 68‐year‐old stand. These results suggest that harvesting and fire disturbances broadly distributed across the region decades ago caused changes in site quality and successional status that continue to limit forest C storage rates.     

Carbon losses due to soil warming: Do autotrophic and heterotrophic soil respiration respond equally?

Global warming has the potential to increase soil respiration (R_S), one of the major fluxes in the global carbon (C) cycle. R_S consists of an autotrophic (R_A) and a heterotrophic (R_H) component. We combined a soil warming experiment with a trenching experiment to assess how R_S, R_A, and R_H are affected. The experiment was conducted in a mature forest dominated by Norway spruce. The site is located in the Austrian Alps on dolomitic bedrock. We warmed the soil of undisturbed and trenched plots by means of heating cables 4 °C above ambient during the snow‐free seasons of 2005 and 2006. Soil warming increased the CO₂ efflux from control plots (R_S) by ∼45% during 2005 and ∼47% during 2006. The CO₂ efflux from trenched plots (R_H) increased by ∼39% during 2005 and ∼45% during 2006. Similar responses of R_S and R_H indicated that the autotrophic and heterotrophic components of R_S responded equally to the temperature increase. Thirty‐five to forty percent or 1 t C ha⁻¹ yr⁻¹ of the overall annual increase in R_S (2.8 t C ha⁻¹ yr⁻¹) was autotrophic. The remaining, heterotrophic part of soil respiration (1.8 t C ha⁻¹ yr⁻¹), represented the warming‐induced C loss from the soil. The autotrophic component showed a distinct seasonal pattern. Contribution of R_A to R_S was highest during summer. Seasonally derived Q₁₀ values reflected this pattern and were correspondingly high (5.3–9.3). The autotrophic CO₂ efflux increase due to the 4 °C warming implied a Q₁₀ of 2.9. Hence, seasonally derived Q₁₀ of R_A did not solely reflect the seasonal soil temperature development.     

Effect of stand age on greenhouse gas fluxes from a Sitka spruce [Picea sitchensis (Bong.) Carr.] chronosequence on a peaty gley soil

The influence of forest stand age in a Picea sitchensis plantation on (1) soil fluxes of three greenhouse gases (GHGs – CO₂, CH₄ and N₂O) and (2) overall net ecosystem global warming potential (GWP), was investigated in a 2‐year study. The objective was to isolate the effect of forest stand age on soil edaphic characteristics (temperature, water table and volumetric moisture) and the consequent influence of these characteristics on the GHG fluxes. Fluxes were measured in a chronosequence in Harwood, England, with sites comprising 30‐ and 20‐year‐old second rotation forest and a site clearfelled (CF) some 18 months before measurement. Adjoining unforested grassland (UN) acted as a control. Comparisons were made between flux data, soil temperature and moisture data and, at the 30‐year‐old and CF sites, eddy covariance data for net ecosystem carbon (C) exchange (NEE). The main findings were: firstly, integrated CO₂ efflux was the dominant influence on the GHG budget, contributing 93–94% of the total GHG flux across the chronosequence compared with 6–7% from CH₄ and N₂O combined. Secondly, there were clear links between the trends in edaphic factors as the forest matured, or after clearfelling, and the emission of GHGs. In the chronosequence sites, annual fluxes of CO₂ were lower at the 20‐year‐old (20y) site than at the 30‐year‐old (30y) and CF sites, with soil temperature the dominant control. CH₄ efflux was highest at the CF site, with peak flux 491±54.5 μg m⁻² h⁻¹ and maximum annual flux 18.0±1.1 kg CH₄ ha⁻¹ yr⁻¹. No consistent uptake of CH₄ was noted at any site. A linear relationship was found between log CH₄ flux and the closeness of the water table to the soil surface across all sites. N₂O efflux was highest in the 30y site, reaching 108±38.3 μg N₂O‐N m⁻² h⁻¹ (171 μg N₂O m⁻² h⁻¹) in midsummer and a maximum annual flux of 4.7±1.2 kg N₂O ha⁻¹ yr⁻¹ in 2001. Automatic chamber data showed a positive exponential relationship between N₂O flux and soil temperature at this site. The relationship between N₂O emission and soil volumetric moisture indicated an optimum moisture content for N₂O flux of 40–50% by volume. The relationship between C : N ratio data and integrated N₂O flux was consistent with a pattern previously noted across temperate and boreal forest soils.     

Changes in carbon storage and fluxes in a chronosequence of ponderosa pine

Forest development following stand‐replacing disturbance influences a variety of ecosystem processes including carbon exchange with the atmosphere. On a series of ponderosa pine (Pinius ponderosa var. Laws.) stands ranging from 9 to> 300 years in central Oregon, USA, we used biological measurements to estimate carbon storage in vegetation and soil pools, net primary productivity (NPP) and net ecosystem productivity (NEP) to examine variation with stand age. Measurements were made on plots representing four age classes with three replications: initiation (I, 9–23 years), young (Y, 56–89 years), mature (M, 95–106 years), and old (O, 190–316 years) stands typical of the forest type in the region. Net ecosystem productivity was lowest in the I stands (?124 g C m^?2 yr^?1), moderate in Y stands (118 g C m^?2 yr^?1), highest in M stands (170 g C m^?2 yr^?1), and low in the O stands (35 g C m^?2 yr^?1). Net primary productivity followed similar trends, but did not decline as much in the O stands. The ratio of fine root to foliage carbon was highest in the I stands, which is likely necessary for establishment in the semiarid environment, where forests are subject to drought during the growing season (300–800 mm precipitation per year). Carbon storage in live mass was the highest in the O stands (mean 17.6 kg C m^?2). Total ecosystem carbon storage and the fraction of ecosystem carbon in aboveground wood mass increased rapidly until 150–200 years, and did not decline in older stands. Forest inventory data on 950 ponderosa pine plots in Oregon show that the greatest proportion of plots exist in stands ～ 100 years old, indicating that a majority of stands are approaching maximum carbon storage and net carbon uptake. Our data suggests that NEP averages ～ 70 g C m^?2 year^?1 for ponderosa pine forests in Oregon. About 85% of the total carbon storage in biomass on the survey plots exists in stands greater than 100 years, which has implications for managing forests for carbon sequestration. To investigate variation in carbon storage and fluxes with disturbance, simulation with process models requires a dynamic parameterization for biomass allocation that depends on stand age, and should include a representation of competition between multiple plant functional types for space, water, and nutrients.     

Measurement and modelling of CO2 flux from a drained fen peatland cultivated with reed canary grass and spring barley

Cultivation of bioenergy crops has been suggested as a promising option for reduction of greenhouse gas (GHG) emissions from arable organic soils (Histosols). Here, we report the annual net ecosystem exchange (NEE) fluxes of CO₂ as measured with a dynamic closed chamber method at a drained fen peatland grown with reed canary grass (RCG) and spring barley (SB) in a plot experiment (n = 3 for each cropping system). The CO₂ flux was partitioned into gross photosynthesis (GP) and ecosystem respiration (R_E). For the data analysis, simple yet useful GP and R_E models were developed which introduce plot‐scale ratio vegetation index as an active vegetation proxy. The GP model captures the effect of temperature and vegetation status, and the R_E model estimates the proportion of foliar biomass dependent respiration (R_fb) in the total R_E. Annual R_E was 1887 ± 7 (mean ± standard error, n = 3) and 1288 ± 19 g CO₂‐C m^?2 in RCG and SB plots, respectively, with R_fb accounting for 32 and 22% respectively. Total estimated annual GP was ?1818 ± 42 and ?1329 ± 66 g CO₂‐C m^?2 in RCG and SB plots leading to a NEE of 69 ± 36 g CO₂‐C m^?2 yr^?1 in RCG plots (i.e., a weak net source) and ?41 ± 47 g CO₂‐C m^?2 yr^?1 in SB plots (i.e., a weak net sink). Standard errors related to spatial variation were small (as shown above), but more significant uncertainties were related to the modelling approach for establishment of annual budgets. In conclusion, the bioenergy cropping system was not more favourable than the food cropping system when looking at the atmospheric CO₂ emissions during cultivation. However, in a broader GHG life‐cycle perspective, the lower fertilizer N input and the higher biomass yield in bioenergy cropping systems could be beneficial.     

Global patterns and predictors of stem CO2 efflux in forest ecosystems

Stem CO₂ efflux (E_S) plays an important role in the carbon balance of forest ecosystems. However, its primary controls at the global scale are poorly understood and observation‐based global estimates are lacking. We synthesized data from 121 published studies across global forest ecosystems and examined the relationships between annual E_S and biotic and abiotic factors at individual, biome, and global scales, and developed a global gridded estimate of annual E_S. We tested the following hypotheses: (1) Leaf area index (LAI) will be highly correlated with annual E_S at biome and global scales; (2) there will be parallel patterns in stem and root CO₂ effluxes (R_A) in all forests; (3) annual E_S will decline with forest age; and (4) LAI coupled with mean annual temperature (MAT) and mean annual precipitation (MAP) will be sufficient to predict annual E_S across forests in different regions. Positive linear relationships were found between E_S and LAI, as well as gross primary production (GPP), net primary production (NPP), wood NPP, soil CO₂ efflux (R_S), and R_A. Annual E_S was correlated with R_A in temperate forests after controlling for GPP and MAT, suggesting other additional factors contributed to the relationship. Annual E_S tended to decrease with stand age. Leaf area index, MAT and MAP, predicted 74% of variation in E_S at global scales. Our statistical model estimated a global annual E_S of 6.7 ± 1.1 Pg C yr⁻¹ over the period of 2000–2012 with little interannual variability. Modeled mean annual E_S was 71 ± 43, 270 ± 103, and 420 ± 134 g C m² yr⁻¹ for boreal, temperate, and tropical forests, respectively. We recommend that future studies report E_S at a standardized constant temperature, incorporate more manipulative treatments, such as fertilization and drought, and whenever possible, simultaneously measure both aboveground and belowground CO₂ fluxes.     

Impact of severe dry season on net ecosystem exchange in the Neotropical rainforest of French Guiana

The lack of information on the ways seasonal drought modifies the CO₂ exchange between Neotropical rainforest ecosystems and the atmosphere and the resulting carbon balance hinders our ability to precisely predict how these ecosystems will respond as global environmental changes force them to face increasingly contrasting conditions in the future. To address this issue, seasonal variations in daily net ecosystem productivity (NEP_d) and two main components of this productivity, daily total ecosystem respiration (R_Ed) and daily gross ecosystem productivity (GEP_d), were estimated over 2 years at a flux tower site in French Guiana, South America (5 °16′54″N, 52 °54′44″W). We compared seasonal variations between wet and dry periods and between dry periods of contrasting levels of intensity (i.e. mild vs. severe) during equivalent 93‐day periods. During the wet periods, the ecosystem was almost in balance with the atmosphere (storage of 9.0 g C m^?2). Seasonal dry periods, regardless of their severity, are associated with higher incident radiation and lower R_Ed combined with reduced soil respiration associated with low soil water availability. During the mild dry period, as is normally the case in this region, the amount of carbon stored in the ecosystem was 32.7 g C m^?2. Severe drought conditions resulted in even lower R_Ed, whereas the photosynthetic activity was only moderately reduced and no change in canopy structure was observed. Thus, the severe dry period was characterized by greater carbon storage (64.6 g C m^?2), emphasizing that environmental conditions, such as during a severe drought, modify the CO₂ exchange between Neotropical rainforest ecosystems and the atmosphere and potentially the resulting carbon balance.     

Net ecosystem carbon dioxide exchange over grazed steppe in central Mongolia

This paper presents results of 1 year (from March 25, 2003 to March 24, 2004, 366 days) of continuous measurements of net ecosystem CO₂ exchange (NEE) above a steppe in Mongolia using the eddy covariance technique. The steppe, typical of central Mongolia, is dominated by C₃ plants adapted to the continental climate. The following two questions are addressed: (1) how do NEE and its components: gross ecosystem production (GEP) and total ecosystem respiration (R_eco) vary seasonally? (2) how do NEE, GEP, and R_eco respond to biotic and abiotic factors? The hourly minimal NEE and the hourly maximal R_eco were −3.6 and 1.2 μmol m⁻² s⁻¹, respectively (negative values denoting net carbon uptake by the canopy from the atmosphere). Peak daily sums of NEE, GEP, and R_eco were −2.3, 3.5, and 1.5 g C m⁻² day⁻¹, respectively. The annual sums of GEP, R_eco, and NEE were 179, 138, and −41 g C m⁻², respectively. The carbon removal by sheep was estimated to range between 10 and 82 g C m⁻² yr⁻¹ using four different approaches. Including these estimates in the overall carbon budget yielded net ecosystem productivity of −23 to +20 g C m⁻² yr⁻¹. Thus, within the remaining experimental uncertainty the carbon budget at this steppe site can be considered to be balanced. For the growing period (from April 23 to October 21, 2003), 26% and 53% of the variation in daily NEE and GEP, respectively, could be explained by the changes in leaf area index. Seasonality of GEP, R_eco, and NEE was closely associated with precipitation, especially in the peak growing season when GEP and R_eco were largest. Water stress was observed in late July to early August, which switched the steppe from a carbon sink to a carbon source. For the entire growing period, the light response curves of daytime NEE showed a rather low apparent quantum yield (α=−0.0047 μmol CO₂ μmol⁻¹ photons of photosynthetically active radiation). However, the α values varied with air temperature (T_a), vapor pressure deficit, and soil water content.     

Accelerating regrowth of temperate‐maritime forests due to environmental change

To understand how environmental changes have influenced forest productivity, stemwood biomass (B) dynamics were analyzed at 1267 permanent inventory plots, covering a combined 209 ha area of unmanaged temperate‐maritime forest in southwest British Columbia, Canada. Net stemwood production (ΔB) was derived from periodic remeasurements of B collected over a 40‐year measurement period (1959–1998) in stands ranging from 20 to 150 years old. Comparison between the integrated age response of net stemwood production, ΔB(A), and the age response of stemwood biomass, B(A), suggested a 58 ± 11% increase in ΔB between the first 40 years of the chronosequence period (1859–1898) and the measurement period. To estimate extrinsic forcing on ΔB, several different candidate models were developed to remove variation explained by intrinsic factors. All models exhibited temporal bias, with positive trends in (observed minus predicted) residual ΔB ranging between of 0.40 and 0.64% yr^?1. Applying the same methods to stemwood growth (G) indicated residual increases ranging from 0.43 and 0.67% yr^?1. Higher trend estimates corresponded with models that included site index (SI) as a predictor, which may reflect exaggeration of the age‐decline in SI tables. Choosing a model that excluded SI, suggested that ΔB increased by 0.40 ± 0.18% yr^?1, while G increased by 0.43 ± 0.12% yr^?1 over the measurement period. Residual G was significantly correlated with atmospheric carbon dioxide (CO₂), temperature (T), and climate moisture index (CMI). However, models driven with climate and CO₂, alone, could not simultaneously explain long‐term and measurement‐period trends without additional representation of indirect effects, perhaps reflecting compound interest on direct physiological responses to environmental change. Evidence of accelerating forest regrowth highlights the value of permanent inventories to detect and understand systematic changes in forest productivity caused by environmental change.     

Interannual variation in soil CO2 efflux and the response of root respiration to climate and canopy gas exchange in mature ponderosa pine

We examined a 6‐year record of automated chamber‐based soil CO₂ efflux (F_s) and the underlying processes in relation to climate and canopy gas exchange at an AmeriFlux site in a seasonally drought‐stressed pine forest. Interannual variability of F_s was large (CV=17%) with a range of 427 g C m^?2 yr^?1 around a mean annual F_s of 811 g C m^?2 yr^?1. On average, 76% of the variation of daily mean F_s could be quantified using an empirical model with year‐specific basal respiration rate that was a linear function of tree basal area increment (BAI) and modulated by a common response to soil temperature and moisture. Interannual variability in F_s could be attributed almost equally to interannual variability in BAI (a proxy for above‐ground productivity) and interannual variability in soil climate. Seasonal total F_s was twice as sensitive to soil moisture variability during the summer months compared with temperature variability during the same period and almost insensitive to the natural range of interannual variability in spring temperatures. A strong seasonality in both root respiration (R_r) and heterotrophic respiration (R_h) was observed with the fraction attributed to R_r steadily increasing from 18% in mid‐March to 50% in early June through early July before dropping rapidly to 10% of F_s by mid‐August. The seasonal pattern in R_r (10‐day averages) was strongly linearly correlated with tree transpiration (r²=0.90, P<0.01) as measured using sap flux techniques and gross ecosystem productivity (GEP, r²=0.83, P<0.01) measured by the eddy‐covariance approach. R_r increased by 0.43 g C m^?2 day^?1 for every 1 g C m^?2 day^?1 increase in GEP. The strong linear correlation of R_r to seasonal changes in GEP and transpiration combined with longer‐term interannual variability in the base rate of F_s, as a linear function of BAI (r²=0.64, P=0.06), provides compelling justification for including canopy processes in future models of F_s.     

Carbon storage and fluxes in ponderosa pine forests at different developmental stages

We compared carbon storage and fluxes in young and old ponderosa pine stands in Oregon, including plant and soil storage, net primary productivity, respiration fluxes, eddy flux estimates of net ecosystem exchange (NEE), and Biome‐BGC simulations of fluxes. The young forest (Y site) was previously an old‐growth ponderosa pine forest that had been clearcut in 1978, and the old forest (O site), which has never been logged, consists of two primary age classes (50 and 250 years old). Total ecosystem carbon content (vegetation, detritus and soil) of the O forest was about twice that of the Y site (21 vs. 10 kg C m^?2 ground), and significantly more of the total is stored in living vegetation at the O site (61% vs. 15%). Ecosystem respiration (R_e) was higher at the O site (1014 vs. 835 g C m^?2 year^?1), and it was largely from soils at both sites (77% of R_e). The biological data show that above‐ground net primary productivity (ANPP), NPP and net ecosystem production (NEP) were greater at the O site than the Y site. Monte Carlo estimates of NEP show that the young site is a source of CO₂ to the atmosphere, and is significantly lower than NEP(O) by c. 100 g C m^?2 year^?1. Eddy covariance measurements also show that the O site was a stronger sink for CO₂ than the Y site. Across a 15‐km swath in the region, ANPP ranged from 76 g C m^?2 year^?1 at the Y site to 236 g C m^?2 year^?1 (overall mean 158 ± 14 g C m^?2 year^?1). The lowest ANPP values were for the youngest and oldest stands, but there was a large range of ANPP for mature stands. Carbon, water and nitrogen cycle simulations with the Biome‐BGC model suggest that disturbance type and frequency, time since disturbance, age‐dependent changes in below‐ground allocation, and increasing atmospheric concentration of CO₂ all exert significant control on the net ecosystem exchange of carbon at the two sites. Model estimates of major carbon flux components agree with budget‐based observations to within ± 20%, with larger differences for NEP and for several storage terms. Simulations showed the period of regrowth required to replace carbon lost during and after a stand‐replacing fire (O) or a clearcut (Y) to be between 50 and 100 years. In both cases, simulations showed a shift from net carbon source to net sink (on an annual basis) 10–20 years after disturbance. These results suggest that the net ecosystem production of young stands may be low because heterotrophic respiration, particularly from soils, is higher than the NPP of the regrowth. The amount of carbon stored in long‐term pools (biomass and soils) in addition to short‐term fluxes has important implications for management of forests in the Pacific North‐west for carbon sequestration.     

Soil CO<Subscript>2</Subscript> efflux in loblolly pine (<Emphasis Type="Italic">Pinus taeda</Emphasis> L.) plantations on the Virginia Piedmont and South Carolina Coastal Plain over a rotation-length chronosequence

We measured soil surface CO₂ efflux (F_s) in loblolly pine stands (Pinus taeda L.) located on the Virginia Piedmont (VA) and South Carolina Coastal Plain (SC) in efforts to assess the impact climate, productivity, and cultural practices have on F_s in the managed loblolly pine ecosystem. The effect of stand age on F_s was examined using a replicated chronosequence approach in which stands ranging from 1 to 25 years since planting were investigated. Soil CO₂ efflux was measured on both VA and SC sites for over a year using a closed dynamic system. Multiple linear regression was used to evaluate F_s correlates and examine the relationship between candidate explanatory variables and F_s. Soil temperature (top 10 cm) was the major correlate with F_s on both locations. We observed a positive age effect on F_s in VA stands and no relationship between age and F_s in SC stands. Annual soil C efflux declined with stand age in SC due to both reductions in soil temperatures as crown closure occurs and a diminishing heterotrophic C substrate pool. Annual estimated efflux ranges from 16.7 to 13.2 Mg C ha⁻¹ for 1 and 20-year-old stands, respectively. In contrast, annual soil C efflux increased with age in VA stands as a result of the positive relationship between stand age and F_s, which appears to be related to an increase in the contribution of root respiration to total F_s over time. In VA stands, efflux estimates range from 7.6 to 12.3 Mg C ha⁻¹ for 1 and 20-year-old stands, respectively. These results demonstrate the need to further consider the impact forest management and within-region variability have on soil C efflux over time when estimating C budgets.     

Estimates of soil respiration and net primary production of three forests at different succession stages in South China

Soil respiration (heterotropic and autotropic respiration, R_g) and aboveground litter fall carbon were measured at three forests at different succession (early, middle and advanced) stages in Dinghushan Biosphere Reserve, Southern China. It was found that the soil respiration increases exponentially with soil temperature at 5 cm depth (T_s) according to the relation R_g=a exp(bT_s), and the more advanced forest community during succession has a higher value of a because of higher litter carbon input than the forests at early or middle succession stages. It was also found that the monthly soil respiration is linearly correlated with the aboveground litter carbon input of the previous month. Using measurements of aboveground litter and soil respiration, the net primary productions (NPPs) of three forests were estimated using nonlinear inversion. They are 475, 678 and 1148 g C m^?2 yr^?1 for the Masson pine forest (MPF), coniferous and broad‐leaf mixed forest (MF) and subtropical monsoon evergreen broad‐leaf forest (MEBF), respectively, in year 2003/2004, of which 54%, 37% and 62% are belowground NPP for those three respective forests if no change in live plant biomass is assumed. After taking account of the decrease in live plant biomass, we estimated the NPP of the subtropical MEBF is 970 g C m^?2 yr^?1 in year 2003/2004. Total amount of carbon allocated below ground for plant roots is 388 g C m^?2 yr^?1 for the MPF, 504 g C m^?2 yr^?1 for the coniferous and broad‐leaf MF and 1254 g C m^?2 yr^?1 for the subtropical MEBF in 2003/2004. Our results support the hypothesis that the amount of carbon allocation belowground increases during forest succession.