Measurements of gross and net ecosystem productivity and water vapour exchange of a Pinus ponderosa ecosystem,and an evaluation of two generalized models

Net ecosystem productivity (NEP), net primary productivity (NPP), and water vapour exchange of a mature Pinus ponderosa forest (44°30′ N, 121°37′ W) growing in a region subject to summer drought were investigated along with canopy assimilation and respiratory fluxes. This paper describes seasonal and annual variation in these factors, and the evaluation of two generalized models of carbon and water balance (PnET‐II and 3‐PG) with a combination of traditional measurements of NPP, respiration and water stress, and eddy covariance measurements of above‐and below‐canopy CO₂ and water vapour exchange. The objective was to evaluate the models using two years of traditional and eddy covariance measurements, and to use the models to help interpret the relative importance of processes controlling carbon and water vapour exchange in a water‐limited pine ecosystem throughout the year. PnET‐II is a monthly time‐step model that is driven by nitrogen availability through foliar N concentration, and 3‐PG is a monthly time‐step quantum‐efficiency model constrained by extreme temperatures, drought, and vapour pressure deficits. Both models require few parameters and have the potential to be applied at the watershed to regional scale. There was 2/3 less rainfall in 1997 than in 1996, providing a challenge to modelling the water balance, and consequently the carbon balance, when driving the models with the two years of climate data, sequentially. Soil fertility was not a key factor in modelling processes at this site because other environmental factors limited photosynthesis and restricted projected leaf area index to ～1.6. Seasonally, GEP and LE were overestimated in early summer and underestimated through the rest of the year. The model predictions of annual GEP, NEP and water vapour exchange were within 1–39% of flux measurements, with greater disparity in 1997 because soil water never fully recharged. The results suggest that generalized models can provide insights to constraints on productivity on an annual basis, using a minimum of site data.     

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.     

Carbon sequestration and ecosystem respiration for 4 years in a Scots pine forest

The net exchange of CO₂ (NEE) between a Scots pine (Pinus sylvestris L.) forest ecosystem in eastern Finland and the atmosphere was measured continuously by the eddy covariance (EC) technique over 4 years (1999–2002). The annual temperature coefficient (Q₁₀) of ecosystem respiration (R) for these years, respectively, was 2.32, 2.66, 2.73 and 2.69. The light‐saturated rate of photosynthesis (A_max) was highest in July or August, with an annual average A_max of 10.9, 14.6, 15.3 and 17.1 μmol m^?2 s^?1 in the 4 years, respectively. There was obvious seasonality in NEE, R and gross primary production (GPP), exhibiting a similar pattern to photosynthetically active radiation (PAR) and air temperature. The integrated daily NEE ranged from 2.59 to ?4.97 g C m^?2 day^?1 in 1999, from 2.70 to ?4.72 in 2000, from 2.61 to ?4.71 in 2001 and from 5.27 to ?4.88 in 2002. The maximum net C uptake occurred in July, with the exception of 2000, when it was in June. The interannual variation in ecosystem C flux was pronounced. The length of the growing season, based on net C uptake, was 179, 170, 175 and 176 days in 1999–2002, respectively, and annual net C sequestration was 152, 101, 172 and 205 g C m^?2 yr^?1. It is estimated that ecosystem respiration contributed 615, 591, 752 and 879 g C m^?2 yr^?1 to the NEE in these years, leading to an annual GPP of ?768, ?692, ?924 and ?1084 g C m^?2 yr^?1. It is concluded that temperature and PAR were the main determinants of the ecosystem CO₂ flux. Interannual variations in net C sequestration are predominantly controlled by average air temperature and integrated radiation in spring and summer. Four years of EC data indicate that boreal Scots pine forest ecosystem in eastern Finland acts as a relatively powerful carbon sink. Carbon sequestration may benefit from warmer climatic conditions.     

Long-term impact of a stand-replacing fire on ecosystem CO2 exchange of a ponderosa pine forest

Ponderosa pine (Pinus ponderosa) forests of the southwestern United States are a mosaic of stands where undisturbed forests are carbon sinks, and stands recovering from wildfires may be sources of carbon to the atmosphere for decades after the fire. However, the relative magnitude of these sinks and sources has never been directly measured in this region, limiting our understanding of the role of fire in regional and US carbon budgets. We used the eddy covariance technique to measure the CO₂ exchange of two forest sites, one burned by fire in 1996, and an unburned forest. The fire was a high‐intensity stand‐replacing burn that killed all trees. Ten years after the fire, the burned site was still a source of CO₂ to the atmosphere [109±6 (SEM) g C m^?2 yr^?1], whereas the unburned site was a sink (?164±23 g C m^?2 yr^?1). The fire reduced total carbon storage and shifted ecosystem carbon allocation from the forest floor and living biomass to necromass. Annual ecosystem respiration was lower at the burned site (480±5 g C m^?2 yr^?1) than at the unburned site (710±54 g C m^?2 yr^?1), but the difference in gross primary production was even larger (372±13 g C m^?2 yr^?1 at the burned site and 858±37 g C m^?2 yr^?1at the unburned site). Water availability controlled carbon flux in the warm season at both sites, and the burned site was a source of carbon in all months, even during the summer, when wet and warm conditions favored respiration more than photosynthesis. Our study shows that carbon losses following stand‐replacing fires in ponderosa pine forests can persist for decades due to slow recovery of the gross primary production. Because fire exclusion is becoming increasingly difficult in dry western forests, a large US forest carbon sink could shift to a decadal‐scale carbon source.     

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.     

Seasonal and annual stem respiration of Scots pine trees under boreal conditions

BACKGROUND AND AIMS: Stem respiration of trees is a major, but poorly assessed component of the carbon balance of forests, and important for geo-chemistry. Measurements are required under naturally changing seasonal conditions in different years. Therefore, intra- and inter-annual carbon fluxes of stems in forests were measured continuously from April to November in three consecutive years. METHODS: Stem respiratory CO2 fluxes of 50-year-old Scots pine (Pinus sylvestris) trees were continuously measured with a CO2 analyser, and, concomitantly, stem circumference, stem and air temperature and other environmental factors and photosynthesis, were also measured automatically. KEY RESULTS: There were diurnal, seasonal and inter-annual changes in stem respiration, which peaked at 1600 h during the day and was highest in July. The temperature coefficient of stem respiration (Q10) was greater during the growing season than when growth was slow or had stopped, and more sensitive to temperature in the growing season. The annual Q10 remained relatively constant at about 2 over the three years, while respiration at a reference temperature of 15 degrees C (R15) was higher in the growing than in the non-growing season (1.09 compared with 0.78 micromol m(-2) stem surface s(-1)), but was similar between the years. Maintenance respiration was 76 %, 82 % and 80 % of the total respiration of 17.46, 17.26 and 19.35 mol m2 stem surface in 2001, 2002 and 2003, respectively. The annual total stem respiration of the stand per unit ground area was 75.97 gC m(-2) in 2001 and 74.28 gC m(-2) in 2002. CONCLUSIONS: Stem respiration is an important component in the annual carbon balance of a Scots pine stand, contributing 9 % to total carbon loss from the ecosystem and consuming about 8 % of the carbon of the ecosystem gross primary production. Stem (or air) temperature was the most important predictor of stem carbon flux. The magnitude of stem respiration is modified by photosynthesis and tree growth. Solar radiation indirectly affects stem respiration through its effect on photosynthesis.     

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.     

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.     

Carbon sequestration in boreal jack pine stands following harvesting

A large area of boreal jack pine (Pinus banksiana Lamb.) forest in Canada is recovering from clear‐cut harvesting, and the carbon (C) balance of these regenerating forests remains uncertain. Net ecosystem CO₂ exchange was measured using the eddy‐covariance technique at four jack pine sites representing different stages of stand development: three postharvest sites (HJP02, HJP94, and HJP75) and one preharvest site (OJP). The four sites, located in the southern Canadian boreal forest, Saskatchewan, Canada, are typical of low productivity jack pine stands and were 2, 10, 29, and 90 years old in 2004, respectively. Mean annual net ecosystem production (NEP) for 2004 and 2005 was ?137±11, 19±16, 73±28, and 22±30 g C m^?2 yr^?1 at HJP02, HJP94, HJP75 and OJP, respectively, showing the postharvest jack pine stands to be moderate C sources immediately after harvesting, weak sinks at 10 years, moderate C sinks at 30 years, then weak C sinks at 90 years. Mean annual gross ecosystem photosynthesis (GEP) for the 2 years was 96±10, 347±20, 576±34, and 583±35 g C m^?2 yr^?1 at HJP02, HJP94, HJP75, and OJP, respectively. The ratio of annual ecosystem respiration (R) to annual GEP was 2.51±0.15, 0.95±0.04, 0.87±0.03, and 0.96±0.03. Seasonally, NEP peaked in May or June at all four sites but GEP and R were highest in July. R at a reference soil temperature of 10 °C, ecosystem quantum yield and photosynthetic capacity were lowest for the 2‐year‐old stand. R was most sensitive to soil temperature for the 90‐year‐old stand. The primary source of variability in NEP over the course of succession of the jack pine ecosystem following harvesting was stand age due to the changes in leaf area index. Intersite variability in GEP and R was an order of magnitude greater than interannual variability at OJP. For both young and old stands, GEP had greater interannual variability than R and played a more important role than R in interannual variation in NEP. Based on year‐round flux measurements from 2000 to 2005, the 10‐year stand had larger interannual variability in GEP and R than the 90‐year stand. Interannual variability in NEP was driven primarily by early‐growing‐season temperature and growing‐season length. Photosynthesis played a dominant role in the rapid rise in NEP early in stand development. Late in stand development, however, the subtle decrease in NEP resulted primarily from increasing respiration.     

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.     

On the separation of net ecosystem exchange into assimilation and ecosystem respiration: review and improved algorithm

This paper discusses the advantages and disadvantages of the different methods that separate net ecosystem exchange (NEE) into its major components, gross ecosystem carbon uptake (GEP) and ecosystem respiration (R_eco). In particular, we analyse the effect of the extrapolation of night‐time values of ecosystem respiration into the daytime; this is usually done with a temperature response function that is derived from long‐term data sets. For this analysis, we used 16 one‐year‐long data sets of carbon dioxide exchange measurements from European and US‐American eddy covariance networks. These sites span from the boreal to Mediterranean climates, and include deciduous and evergreen forest, scrubland and crop ecosystems. We show that the temperature sensitivity of R_eco, derived from long‐term (annual) data sets, does not reflect the short‐term temperature sensitivity that is effective when extrapolating from night‐ to daytime. Specifically, in summer active ecosystems the long‐term temperature sensitivity exceeds the short‐term sensitivity. Thus, in those ecosystems, the application of a long‐term temperature sensitivity to the extrapolation of respiration from night to day leads to a systematic overestimation of ecosystem respiration from half‐hourly to annual time‐scales, which can reach >25% for an annual budget and which consequently affects estimates of GEP. Conversely, in summer passive (Mediterranean) ecosystems, the long‐term temperature sensitivity is lower than the short‐term temperature sensitivity resulting in underestimation of annual sums of respiration. We introduce a new generic algorithm that derives a short‐term temperature sensitivity of R_eco from eddy covariance data that applies this to the extrapolation from night‐ to daytime, and that further performs a filling of data gaps that exploits both, the covariance between fluxes and meteorological drivers and the temporal structure of the fluxes. While this algorithm should give less biased estimates of GEP and R_eco, we discuss the remaining biases and recommend that eddy covariance measurements are still backed by ancillary flux measurements that can reduce the uncertainties inherent in the eddy covariance data.     

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.     

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.     

Soil pCO2, soil respiration,and root activity in CO2-fumigated and nitrogen-fertilized ponderosa pine

The purpose of this paper is to describe the effects of CO₂ and N treatments on soil pCO₂, calculated CO₂ efflux, root biomass and soil carbon in open-top chambers planted with Pinus ponderosa seedlings. Based upon the literature, it was hypothesized that both elevated CO₂ and N would cause increased root biomass which would in turn cause increases in both total soil CO₂ efflux and microbial respiration. This hypothesis was only supported in part: both CO₂ and N treatments caused significant increases in root biomass, soil pCO₂, and calculated CO₂ efflux, but there were no differences in soil microbial respiration measured in the laboratory. Both correlative and quantitative comparisons of CO₂ efflux rates indicated that microbial respiration contributes little to total soil CO₂ efflux in the field. Measurements of soil pCO₂ and calculated CO₂ efflux provided inexpensive, non-invasive, and relatively sensitive indices of belowground response to CO₂ and N treatments.     

Seasonal and interannual variation in carbon dioxide exchange and carbon balance in a northern temperate grassland

Net ecosystem carbon dioxide (CO₂) exchange (NEE) was measured in a northern temperate grassland near Lethbridge, Alberta, Canada for three growing seasons using the eddy covariance technique. The study objectives were to document how NEE and its major component processes—gross photosynthesis (GPP) and total ecosystem respiration (TER)—vary seasonally and interannually, and to examine how environmental and physiological factors influence the annual C budget. The greatest difference among the three study years was the amount of precipitation received. The annual precipitation for 1998 (481.7 mm) was significantly above the 1971–2000 mean (± SD, 377.9 ± 97.0 mm) for Lethbridge, whereas 1999 (341.3 mm) was close to average, and 2000 (275.5 mm) was significantly below average. The high precipitation and soil moisture in 1998 allowed a much higher GPP and an extended period of net carbon gain relative to 1999 and 2000. In 1998, the peak NEE was a gain of 5 g C m^?2 d^?1 (day 173). Peak NEE was lower and also occurred earlier in the year on days 161 (3.2 g C m^?2 d^?1) and 141 (2.4 g C m^?2 d^?1) in 1999 and 2000, respectively. Change in soil moisture was the most important ecological factor controlling C gain in this grassland ecosystem. Soil moisture content was positively correlated with leaf area index (LAI). Gross photosynthesis was strongly correlated with changes in both LAI and canopy nitrogen (N) content. Maximum GPP (A_max: value calculated from a rectangular hyperbola fitted to the relationship between GPP and incident photosynthetic photon flux density (PPFD)) was 27.5, 12.9 and 8.6 µmol m^?2 s^?1 during 1998, 1999 and 2000, respectively. The apparent quantum yield also differed among years at the time of peak photosynthetic activity, with calculated values of 0.0254, 0.018 and 0.018 during 1998, 1999 and 2000, respectively. The ecosystem accumulated a total of 111.9 g C m^?2 from the time the eddy covariance measurements were initiated in June 1998 until the end of December 2000, with most of that C gained during 1998. There was a net uptake of almost 21 g C m^?2 in 1999, whereas a net loss of 18 g C m^?2 was observed in 2000. The net uptake of C during 1999 was the combined result of slightly higher GPP (287.2 vs. 272.3 g C m^?2 year^?1) and lower TER (266.6 vs. 290.4 g C m^?2 year^?1) than occurred in 2000.     

Seasonal net carbon dioxide exchange of a beech forest with the atmosphere

The seasonal carbon dioxide exchange of a beech forest of Central Italy was studied by means of the eddy covariance technique. Additional measurements of biomass respiration with cuvettes and relationship of carbon dioxide exchanges with temperature and light were used to interpolate missing data during the dormant and part of the growing season. The net ecosystem production of the forest equals 472 g C m^?2 y^?1 while the gross ecosystem production 1016 g C m^?2 y^?1 and respiration 544 g C m^?2 y^?1. These estimates are compared with the net primary production determined by direct biomass sampling which amounts to 802 g C m^?2 y^?1.     

Latitudinal patterns of magnitude and interannual variability in net ecosystem exchange regulated by biological and environmental variables

Over the last two and half decades, strong evidence showed that the terrestrial ecosystems are acting as a net sink for atmospheric carbon. However the spatial and temporal patterns of variation in the sink are not well known. In this study, we examined latitudinal patterns of interannual variability (IAV) in net ecosystem exchange (NEE) of CO₂ based on 163 site-years of eddy covariance data, from 39 northern-hemisphere research sites located at latitudes ranging from ∼29°N to ∼64°N. We computed the standard deviation of annual NEE integrals at individual sites to represent absolute interannual variability (AIAV), and the corresponding coefficient of variation as a measure of relative interannual variability (RIAV). Our results showed decreased trends of annual NEE with increasing latitude for both deciduous broadleaf forests and evergreen needleleaf forests. Gross primary production (GPP) explained a significant proportion of the spatial variation of NEE across evergreen needleleaf forests, whereas, across deciduous broadleaf forests, it is ecosystem respiration (Re). In addition, AIAV in GPP and Re increased significantly with latitude in deciduous broadleaf forests, but AIAV in GPP decreased significantly with latitude in evergreen needleleaf forests. Furthermore, RIAV in NEE, GPP, and Re appeared to increase significantly with latitude in deciduous broadleaf forests, but not in evergreen needleleaf forests. Correlation analyses showed air temperature was the primary environmental factor that determined RIAV of NEE in deciduous broadleaf forest across the North American sites, and none of the chosen climatic factors could explain RIAV of NEE in evergreen needleleaf forests. Mean annual NEE significantly increased with latitude in grasslands. Precipitation was dominant environmental factor for the spatial variation of magnitude and IAV in GPP and Re in grasslands.     

Influence of physiological phenology on the seasonal pattern of ecosystem respiration in deciduous forests

Understanding the environmental and biotic drivers of respiration at the ecosystem level is a prerequisite to further improve scenarios of the global carbon cycle. In this study we investigated the relevance of physiological phenology, defined as seasonal changes in plant physiological properties, for explaining the temporal dynamics of ecosystem respiration (R_ECO) in deciduous forests. Previous studies showed that empirical R_ECO models can be substantially improved by considering the biotic dependency of R_ECO on the short‐term productivity (e.g., daily gross primary production, GPP) in addition to the well‐known environmental controls of temperature and water availability. Here, we use a model‐data integration approach to investigate the added value of physiological phenology, represented by the first temporal derivative of GPP, or alternatively of the fraction of absorbed photosynthetically active radiation, for modeling R_ECO at 19 deciduous broadleaved forests in the FLUXNET La Thuile database. The new data‐oriented semiempirical model leads to an 8% decrease in root mean square error (RMSE) and a 6% increase in the modeling efficiency (EF) of modeled R_ECO when compared to a version of the model that does not consider the physiological phenology. The reduction of the model‐observation bias occurred mainly at the monthly time scale, and in spring and summer, while a smaller reduction was observed at the annual time scale. The proposed approach did not improve the model performance at several sites, and we identified as potential causes the plant canopy heterogeneity and the use of air temperature as a driver of ecosystem respiration instead of soil temperature. However, in the majority of sites the model‐error remained unchanged regardless of the driving temperature. Overall, our results point toward the potential for improving current approaches for modeling R_ECO in deciduous forests by including the phenological cycle of the canopy.