The turnover of carbon pools contributing to soil CO2 and soil respiration in a temperate forest exposed to elevated CO2 concentration

Soil carbon is returned to the atmosphere through the process of soil respiration, which represents one of the largest fluxes in the terrestrial C cycle. The effects of climate change on the components of soil respiration can affect the sink or source capacity of ecosystems for atmospheric carbon, but no current techniques can unambiguously separate soil respiration into its components. Long‐term free air CO₂ enrichment (FACE) experiments provide a unique opportunity to study soil C dynamics because the CO₂ used for fumigation has a distinct isotopic signature and serves as a continuous label at the ecosystem level. We used the ¹³C tracer at the Duke Forest FACE site to follow the disappearance of C fixed before fumigation began in 1996 (pretreatment C) from soil CO₂ and soil‐respired CO₂, as an index of belowground C dynamics during the first 8 years of the experiment. The decay of pretreatment C as detected in the isotopic composition of soil‐respired CO₂ and soil CO₂ at 15, 30, 70, and 200 cm soil depth was best described by a model having one to three exponential pools within the soil system. The majority of soil‐respired CO₂ (71%) originated in soil C pools with a turnover time of about 35 days. About 55%, 50%, and 68% of soil CO₂ at 15, 30, and 70 cm, respectively, originated in soil pools with turnover times of less than 1 year. The rest of soil CO₂ and soil‐respired CO₂ originated in soil pools that turn over at decadal time scales. Our results suggest that a large fraction of the C returned to the atmosphere through soil respiration results from dynamic soil C pools that cannot be easily detected in traditionally defined soil organic matter standing stocks. Fast oxidation of labile C substrates may prevent increases in soil C accumulation in forests exposed to elevated [CO₂] and may consequently result in shorter ecosystem C residence times.     

The response of heterotrophic CO2 flux to soil warming

In a forest ecosystem at steady state, net carbon (C) assimilation by plants and C loss through soil and litter decomposition by heterotrophic organisms are balanced. However, a perturbation to the system, such as increased mean soil temperature, will lead to faster decay, enhancing CO₂ release from decomposers, and thus upsetting the balance. Recent in situ experiments have indicated that the stimulation of soil respiration following a step increase in annual average soil temperature declines over time. One possible explanation for this decline may be changes in substrate availability. This hypothesis is examined by using the ecosystem model G'DAY, which simulates C and nitrogen (N) dynamics in plants and soil. We applied the model to observations from a soil‐warming experiment in a Norway spruce (Picea abies (L.) Karst.) stand by simulating a step increase of soil temperature. The model provided a good qualitative reproduction of the observed reduction of heterotrophic respiration (R_h) under sustained warming. The simulations showed how the combined effects of faster turnover and reduced substrate availability lead to a transient increase of R_h. The simulated annual increase in R_h from soil was 60% in the first year after perturbation but decreased to 30% after a decade. One conclusion from the analysis of the simulations is that R_h can decrease even though the temperature response function for decomposition remains unchanged. G'DAY suggests that acclimation of R_h to soil warming is partly an effect of substrate depletion of labile C pools during the first decade of warming as a result of accelerated rates of mineralization. The response is attributed mainly to changing levels of C in pools with short time constants, reflecting the importance of high‐quality soil C fractions. Changes of the structure or physiology of the decomposer community were not invoked. Therefore, it becomes a question of definition whether the simulated dynamics of the declining response of CO₂ release to the warming should be named acclimation or seen as a natural part of the system dynamics.     

Response of soil surface CO2 flux in a boreal forest to ecosystem warming

Soil surface carbon dioxide (CO₂) flux (R_S) was measured for 2 years at the Boreal Soil and Air Warming Experiment site near Thompson, MB, Canada. The experimental design was a complete random block design that consisted of four replicate blocks, with each block containing a 15 m × 15 m control and heated plot. Black spruce [Picea mariana (Mill.) BSP] was the overstory species and Epilobium angustifolium was the dominant understory. Soil temperature was maintained (～5 °C) above the control soil temperature using electric cables inside water filled polyethylene tubing for each heated plot. Air inside a 7.3‐m‐diameter chamber, centered in the soil warming plot, contained approximately nine black spruce trees was heated ～5 °C above control ambient air temperature allowing for the testing of soil‐only warming and soil+air warming. Soil surface CO₂ flux (R_S) was positively correlated (P < 0.0001) to soil temperature at 10 cm depth. Soil surface CO₂ flux (R_S) was 24% greater in the soil‐only warming than the control in 2004, but was only 11% greater in 2005, while R_S in the soil+air warming treatments was 31% less than the control in 2004 and 23% less in 2005. Live fine root mass (< 2 mm diameter) was less in the heated than control treatments in 2004 and statistically less (P < 0.01) in 2005. Similar root mass between the two heated treatments suggests that different heating methods (soil‐only vs. soil+air warming) can affect the rate of decomposition.     

Water availability controls microbial temperature responses in frozen soil CO2 production

Soil processes in high-latitude regions during winter are important contributors to global carbon circulation, but our understanding of the mechanisms controlling these processes is poor and observed temperature response coefficients of CO₂ production in frozen soils deviate markedly from thermodynamically predicted responses (sometimes by several orders of magnitude). We investigated the temperature response of CO₂ production in 23 unfrozen and frozen surface soil samples from various types of boreal forests and peatland ecosystems and also measured changes in water content in them after freezing. We demonstrate that deviations in temperature responses at subzero temperatures primarily emanates from water deficiency caused by freezing of the soil water, and that the amount of unfrozen water is mainly determined by the quality of the soil organic matter, which is linked to the vegetation cover. Factoring out the contribution of water limitation to the CO₂ temperature responses yields response coefficients that agree well with expectations based on thermodynamic theory concerning biochemical temperature responses. This partitioning between a pure temperature response and the effect of water availability on the response of soil CO₂ production at low temperatures is crucial for a thorough understanding of low-temperature soil processes and for accurate predictions of C-balances in northern terrestrial ecosystems.     

Vertical partitioning of CO2 production within a temperate forest soil

The major driving factors of soil CO₂ production – substrate supply, temperature, and water content – vary vertically within the soil profile, with the greatest temporal variations of these factors usually near the soil surface. Several studies have demonstrated that wetting and drying of the organic horizon contributes to temporal variation in summertime soil CO₂ efflux in forests, but this contribution is difficult to quantify. The objectives of this study were to partition CO₂ production vertically in a mixed hardwood stand of the Harvard Forest, Massachusetts, USA, and then to use that partitioning to evaluate how the relative contributions of CO₂ production by genetic soil horizon vary seasonally and interannually. We measured surface CO₂ efflux and vertical soil profiles of CO₂ concentration, temperature, water content, and soil physical characteristics. These data were applied to a model of effective diffusivity to estimate CO₂ flux at the top of each genetic soil horizon and the production within each horizon. A sensitivity analysis revealed sources of uncertainty when applying a diffusivity model to a rocky soil with large spatial heterogeneity, especially estimates of bulk density and volumetric water content and matching measurements of profiles and surface fluxes. We conservatively estimate that the O horizon contributed 40–48% of the total annual soil CO₂ efflux. Although the temperature sensitivity of CO₂ production varied across soil horizons, the partitioning of CO₂ production by horizon did not improve the overall prediction of surface CO₂ effluxes based on temperature functions. However, vertical partitioning revealed that water content covaried with CO₂ production only in the O horizon. Large interannual variations in estimates of O horizon CO₂ production indicate that this layer could be an important transient interannual source or sink of ecosystem C.     

The response of soil CO2 flux to changes in atmospheric CO2, nitrogen supply and plant diversity

We measured soil CO₂ flux over 19 sampling periods that spanned two growing seasons in a grassland Free Air Carbon dioxide Enrichment (FACE) experiment that factorially manipulated three major anthropogenic global changes: atmospheric carbon dioxide (CO₂) concentration, nitrogen (N) supply, and plant species richness. On average, over two growing seasons, elevated atmospheric CO₂ and N fertilization increased soil CO₂ flux by 0.57 µmol m^?2 s^?1 (13% increase) and 0.37 µmol m^?2 s^?1 (8% increase) above average control soil CO₂ flux, respectively. Decreases in planted diversity from 16 to 9, 4 and 1 species decreased soil CO₂ flux by 0.23, 0.41 and 1.09 µmol m^?2 s^?1 (5%, 8% and 21% decreases), respectively. There were no statistically significant pairwise interactions among the three treatments. During 19 sampling periods that spanned two growing seasons, elevated atmospheric CO₂ increased soil CO₂ flux most when soil moisture was low and soils were warm. Effects on soil CO₂ flux due to fertilization with N and decreases in diversity were greatest at the times of the year when soils were warm, although there were no significant correlations between these effects and soil moisture. Of the treatments, only the N and diversity treatments were correlated over time; neither were correlated with the CO₂ effect. Models of soil CO₂ flux will need to incorporate ecosystem CO₂ and N availability, as well as ecosystem plant diversity, and incorporate different environmental factors when determining the magnitude of the CO₂, N and diversity effects on soil CO₂ flux.     

Trends and methodological impacts in soil CO2 efflux partitioning: A metaanalytical review

Partitioning soil carbon dioxide (CO₂) efflux (R_S) into autotrophic (R_A; including plant roots and closely associated organisms) and heterotrophic (R_H) components has received considerable attention, as differential responses of these components to environmental change have profound implications for the soil and ecosystem C balance. The increasing number of partitioning studies allows a more detailed analysis of experimental constraints than was previously possible. We present results of an exhaustive literature search of partitioning studies and analyse global trends in flux partitioning between biomes and ecosystem types by means of a metaanalysis. Across all data, an overall decline in the R_H/R_S ratio for increasing annual R_S fluxes emerged. For forest ecosystems, boreal coniferous sites showed significantly higher (P<0.05) R_H/R_S ratios than temperate sites, while both temperate or tropical deciduous forests did not differ in ratios from any of the other forest types. While chronosequence studies report consistent declines in the R_H/R_S ratio with age, no difference could be detected for different age groups in the global data set. Different methodologies showed generally good agreement if the range of R_S under which they had been measured was considered, with the exception of studies estimating R_H by means of root mass regressions against R_S, which resulted in consistently lower R_H/R_S estimates out of all methods included. Additionally, the time step over which fluxes were partitioned did not affect R_H/R_S ratios consistently. To put results into context, we review the most common techniques and point out the likely sources of errors associated with them. In order to improve soil CO₂ efflux partitioning in future experiments, we include methodological recommendations, and also highlight the potential interactions between soil components that may be overlooked as a consequence of the partitioning process itself.     

Forest soil CO2 flux: uncovering the contribution and environmental responses of ectomycorrhizas

Forests play a critical role in the global carbon cycle, being considered an important and continuing carbon sink. However, the response of carbon sequestration in forests to global climate change remains a major uncertainty, with a particularly poor understanding of the origins and environmental responses of soil CO₂ efflux. For example, despite their large biomass, the contribution of ectomycorrhizal (EM) fungi to forest soil CO₂ efflux and responses to changes in environmental drivers has, to date, not been quantified in the field. Their activity is often simplistically included in the ‘autotrophic’ root respiration term. We set up a multiplexed continuous soil respiration measurement system in a young Lodgepole pine forest, using a mycorrhizal mesh collar design, to monitor the three main soil CO₂ efflux components: root, extraradical mycorrhizal hyphal, and soil heterotrophic respiration. Mycorrhizal hyphal respiration increased during the first month after collar insertion and thereafter remained remarkably stable. During autumn the soil CO₂ flux components could be divided into ∼60% soil heterotrophic, ∼25% EM hyphal, and ∼15% root fluxes. Thus the extraradical EM mycelium can contribute substantially more to soil CO₂ flux than do roots. While EM hyphal respiration responded strongly to reductions in soil moisture and appeared to be highly dependent on assimilate supply, it did not responded directly to changes in soil temperature. It was mainly the soil heterotrophic flux component that caused the commonly observed exponential relationship with temperature. Our results strongly suggest that accurate modelling of soil respiration, particularly in forest ecosystems, needs to explicitly consider the mycorrhizal mycelium and its dynamic response to specific environmental factors. Moreover, we propose that in forest ecosystems the mycorrhizal CO₂ flux component represents an overflow ‘CO₂ tap’ through which surplus plant carbon may be returned directly to the atmosphere, thus limiting expected carbon sequestration from trees under elevated CO₂.     

Soil CO2 efflux in a boreal pine forest under atmospheric CO2 enrichment and air warming

The response of forest soil CO₂ efflux to the elevation of two climatic factors, the atmospheric concentration of CO₂ (↑CO₂ of 700 μmol mol⁻¹) and air temperature (↑ T with average annual increase of 5°C), and their combination (↑CO₂+↑ T ) was investigated in a 4-year, full-factorial field experiment consisting of closed chambers built around 20-year-old Scots pines ( Pinus sylvestris L.) in the boreal zone of Finland. Mean soil CO₂ efflux in May–October increased with elevated CO₂ by 23–37%, with elevated temperature by 27–43%, and with the combined treatment by 35–59%. Temperature elevation was a significant factor in the combined 4-year efflux data, whereas the effect of elevated CO₂ was not as evident. Elevated temperature had the most pronounced impact early and late in the season, while the influence of elevated CO₂ alone was especially notable late in the season. Needle area was found to be a significant predictor of soil CO₂ efflux, particularly in August, a month of high root growth, thus supporting the assumption of a close link between whole-tree physiology and soil CO₂ emissions. The decrease in the temperature sensitivity of soil CO₂ efflux observed in the elevated temperature treatments in the second year nevertheless suggests the existence of soil response mechanisms that may be independent of the assimilating component of the forest ecosystem. In conclusion, elevated atmospheric CO₂ and air temperature consistently increased forest soil CO₂ efflux over the 4-year period, their combined effect being additive, with no apparent interaction.     

Effects of elevated atmospheric CO2 on soil microbiota in calcareous grassland

Microbial responses to three years of CO₂ enrichment (600 μL L^–1) in the field were investigated in calcareous grassland. Microbial biomass carbon (C) and soil organic C and nitrogen (N) were not significantly influenced by elevated CO₂. Microbial C:N ratios significantly decreased under elevated CO₂ (– 15%, P = 0.01) and microbial N increased by + 18% (P = 0.04). Soil basal respiration was significantly increased on one out of 7 sampling dates (+ 14%, P = 0.03; December of the third year of treatment), whereas the metabolic quotient for CO₂ (qCO₂ = basal respiration/microbial C) did not exhibit any significant differences between CO₂ treatments. Also no responses of microbial activity and biomass were found in a complementary greenhouse study where intact grassland turfs taken from the field site were factorially treated with elevated CO₂ and phosphorus (P) fertilizer (1 g P m^–2 y^–1). Previously reported C balance calculations showed that in the ecosystem investigated growing season soil C inputs were strongly enhanced under elevated CO₂. It is hypothesized that the absence of microbial responses to these enhanced soil C fluxes originated from mineral nutrient limitations of microbial processes. Laboratory incubations showed that short-term microbial growth (one week) was strongly limited by N availability, whereas P was not limiting in this soil. The absence of large effects of elevated CO₂ on microbial activity or biomass in such nutrient-poor natural ecosystems is in marked contrast to previously published large and short-term microbial responses to CO₂ enrichment which were found in fertilized or disturbed systems. It is speculated that the absence of such responses in undisturbed natural ecosystems in which mineral nutrient cycles have equilibrated over longer periods of time is caused by mineral nutrient limitations which are ineffective in disturbed or fertilized systems and that therefore microbial responses to elevated CO₂ must be studied in natural, undisturbed systems.     

Stimulation of microbial extracellular enzyme activities by elevated CO2 depends on soil aggregate size

Increased belowground carbon (C) transfer by plant roots at elevated CO₂ may change properties of the microbial community in the rhizosphere. Previous investigations that focused on total soil organic C or total microbial C showed contrasting results: small increase, small decrease or no changes. We evaluated the effect of 5 years of elevated CO₂ (550 ppm) on four extracellular enzymes: β‐glucosidase, chitinase, phosphatase, and sulfatase. We expected microorganisms to be differently localized in aggregates of various sizes and, therefore analyzed microbial biomass (C_mic by SIR) and enzyme activities in three aggregate‐size classes: large macro‐ (> 2 mm), small macro‐ (0.25–2 mm), and microaggregates (< 0.25 mm). To estimate the potential enzyme production, we activated microorganisms by substrate (glucose and nutrients) amendment. Although C_total and C_mic as well as the activities of β‐glucosidase, phosphatase, and sulfatase were unaffected in bulk soil and in aggregate‐size classes by elevated CO₂, significant changes were observed in potential enzyme production after substrate amendment. After adding glucose, enzyme activities under elevated CO₂ were 1.2–1.9‐fold higher than under ambient CO₂. This indicates the increased activity of microorganisms, which leads to accelerated C turnover in soil under elevated CO₂. Significantly higher chitinase activity in bulk soil and in large macroaggregates under elevated CO₂ revealed an increased contribution of fungi to turnover processes. At the same time, less chitinase activity in microaggregates underlined microaggregate stability and the difficulties for fungal hyphae penetrating them. We conclude that quantitative and qualitative changes of C input by plants into the soil at elevated CO₂ affect microbial community functioning, but not its total content. Future studies should therefore focus more on the changes of functions and activities, but less on the pools.     

Interactions of tropospheric CO2 and O3 enrichments and moisture variations on microbial biomass and respiration in soil

Soil microbial biomass C (C_mic) is a sensitive indicator of trends in organic matter dynamics in terrestrial ecosystems. This study was conducted to determine the effects of tropospheric CO₂ or O₃ enrichments and moisture variations on total soil organic C (C_org), mineralizable C fraction (C_Min), C_mic, maintenance respiratory (qCO₂) or C_mic death (qD) quotients, and their relationship with basal respiration (BR) rates and field respiration (FR) fluxes in wheat‐soybean agroecosystems. Wheat (Triticum aestivum L.) and soybean (Glycine max. L. Merr) plants were grown to maturity in 3‐m dia open‐top field chambers and exposed to charcoal‐filtered (CF) air at 350 μL CO₂ L^?1; CF air + 150 μL CO₂ L^?1; nonfiltered (NF) air + 35 nL O₃ L^?1; and NF air + 35 nL O₃ L^?1 + 150 μL CO₂ L^?1 at optimum (? 0.05 MPa) and restricted soil moisture (? 1.0 ± 0.05 MPa) regimes. The + 150 μL CO₂ L^?1 additions were 18 h d^?1 and the + 35 nL O₃ L^?1 treatments were 7 h d^?1 from April until late October. While C_org did not vary consistently, C_Min, C_mic and C_mic fractions increased in soils under tropospheric CO₂ enrichment (500 μL CO₂ L^?1) and decreased under high O₃ exposures (55 ± 6 nL O₃ L^?1 for wheat; 60 ± 5 nL O₃ L^?1 for soybean) compared to the CF treatments (25 ± 5 nL O₃ L^?1). The qCO₂ or qD quotients of C_mic were also significantly decreased in soils under high CO₂ but increased under high O₃ exposures compared to the CF control. The BR rates did not vary consistently but they were higher in well‐watered soils. The FR fluxes were lower under high O₃ exposures compared to soils under the CF control. An increase in C_mic or C_mic fractions and decrease in qCO₂ or qD observed under high CO₂ treatment suggest that these soils were acting as C sinks whereas, reductions in C_mic or C_mic fractions and increase in qCO₂ or qD in soils under elevated tropospheric O₃ exposures suggest the soils were serving as a source of CO₂.     

Impact of elevated atmospheric CO2 concentration on soil microbial biomass and activity in a complex, weedy field model ecosystem

Although soil organisms play an essential role in the cycling of elements in terrestrial ecosystems, little is known of the impact of increasing atmospheric CO₂ concentrations on soil microbial processes. We determined microbial biomass and activity in the soil of multitrophic model ecosystems housed in the Ecotron (NERC Centre for Population Biology, Ascot, UK) under two atmospheric CO₂ concentrations (ambient vs. ambient + 200 ppm). The model communities consist of four annual plant species which naturally co-occur in weedy fields and disturbed ground throughout southern England, together with their herbivores, parasitoids and soil biota. At the end of two experimental runs lasting 9 and 4.5 months, respectively, root dry weight and quality showed contradictory responses to elevated CO₂ concentrations, probably as a consequence of the different time-periods (and hence number of plant generations) in the two experiments. Despite significant root responses no differences in microbial biomass could be detected. Effects of CO₂ concentration on microbial activity were also negligible. Specific enzymes (protease and xylanase) showed a significant decrease in activity in one of the experimental runs. This could be related to the higher C:N ratio of root tissue. We compare the results with data from the literature and conclude that the response of complex communities cannot be predicted on the basis of oversimplified experimental set-ups.     

Effects of an induced drought on soil carbon dioxide (CO2) efflux and soil CO2 production in an Eastern Amazonian rainforest, Brazil

In the next few decades, climate of the Amazon basin is expected to change, as a result of deforestation and rising temperatures, which may lead to feedback mechanisms in carbon (C) cycling that are presently unknown. Here, we report how a throughfall exclusion (TFE) experiment affected soil carbon dioxide (CO₂) production in a deeply weathered sandy Oxisol of Caxiuanã (Eastern Amazon). Over the course of 2 years, we measured soil CO₂ efflux and soil CO₂ concentrations, soil temperature and moisture in pits down to 3 m depth. Over a period of 2 years, TFE reduced on average soil CO₂ efflux from 4.3±0.1 μmol CO₂ m⁻² s⁻¹ (control) to 3.2±0.1 μmol CO₂ m⁻² s⁻¹ (TFE). The contribution of the subsoil (below 0.5 m depth) to the total soil CO₂ production was higher in the TFE plot (28%) compared with the control plot (17%), and it did not differ between years. We distinguished three phases of drying after the TFE was started. The first phase was characterized by a translocation of water uptake (and accompanying root activity) to deeper layers and not enough water stress to affect microbial activity and/or total root respiration. During the second phase a reduction in total soil CO₂ efflux in the TFE plot was related to a reduction of soil and litter decomposers activity. The third phase of drying, characterized by a continuing decrease in soil CO₂ production was dominated by a water stress‐induced decrease in total root respiration. Our results contrast to results of a drought experiment on clay Oxisols, which may be related to differences in soil water retention characteristics and depth of rooting zone. These results show that large differences exist in drought sensitivity among Amazonian forest ecosystems, which primarily seem to be affected by the combined effects of texture (affecting water holding capacity) and depth of rooting zone.     

Hierarchical saturation of soil carbon pools near a natural CO2 spring

Soil has been identified as a possible carbon (C) sink to mitigate increasing atmospheric CO₂ concentration. However, several recent studies have suggested that the potential of soil to sequester C is limited and that soil may become saturated with C under increasing CO₂ levels. To test this concept of soil C saturation, we studied a gley and organic soil at a grassland site near a natural CO₂ spring. Total and aggregate‐associated soil organic C (SOC) concentration showed a significant increase with atmospheric CO₂ concentration. An asymptotic function showed a better fit of SOC and aggregation with CO₂ level than a linear model. There was a shift in allocation of total C from smaller size fractions to the largest aggregate fraction with increasing CO₂ concentration. Litter inputs appeared to be positively related to CO₂ concentration. Based on modeled function parameters and the observed shift in the allocation of the soil C from small to large aggregate‐size classes, we postulate that there is a hierarchy in C saturation across different SOC pools. We conclude that the asymptotic response of SOC concentration at higher CO₂ levels indicates saturation of soil C pools, likely because of a limit to physical protection of SOC.     

Impacts of Hurricane Frances on Florida scrub-oak ecosystem processes: defoliation, net CO2 exchange and interactions with elevated CO2

Hurricane disturbances have profound impacts on ecosystem structure and function, yet their effects on ecosystem CO₂ exchange have not been reported. In September 2004, our research site on a fire‐regenerated scrub‐oak ecosystem in central Florida was struck by Hurricane Frances with sustained winds of 113 km h⁻¹ and wind gusts as high as 152 km h⁻¹. We quantified the hurricane damage on this ecosystem resulting from defoliation: we measured net ecosystem CO₂ exchange, the damage and recovery of leaf area, and determined whether growth in elevated carbon dioxide concentration in the atmosphere (C_a) altered this disturbance. The hurricane decreased leaf area index (LAI) by 21%, which was equal to 60% of seasonal variation in canopy growth during the previous 3 years, but stem damage was negligible. The reduction in LAI led to a 22% decline in gross primary production (GPP) and a 25% decline in ecosystem respiration (R_e). The compensatory declines in GPP and R_e resulted in no significant change in net ecosystem production (NEP). Refoliation began within a month after the hurricane, although this period was out of phase with the regular foliation period, and recovered 20% of the defoliation loss within 2.5 months. Full recovery of LAI, ecosystem CO₂ assimilation, and ecosystem respiration did not occur until the next growing season. Plants exposed to elevated C_a did not sustain greater damage, nor did they recover faster than plants grown under ambient C_a. Thus, our results indicate that hurricanes capable of causing significant defoliation with negligible damage to stems have negligible effects on NEP under current or future CO₂‐enriched environment.     

Seasonal patterns of soil CO2 efflux and soil air CO2 concentration in a Scots pine forest: comparison of two chamber techniques

A non‐vented non‐steady state flow‐through chamber and a non‐vented non‐steady state non‐flow‐through chamber technique were used to measure CO₂ efflux of a young Scots pine forest on a fertile till soil in southern Finland. Soil temperature, soil moisture and soil CO₂ concentration were measured concurrently with CO₂ efflux for two and a half successive years. The CO₂ efflux showed a seasonal pattern, effluxes ranging from low 0.0–0.1 g CO₂ m ^{? 2} h ^{? 1} in winter to peak values of 2.3 g CO₂ m ^{? 2} h ^{? 1} occurring in late June and in July. The daily average effluxes in July measured by flow through chambers were 1.23 and 0.98 g CO₂ m ^{? 2} h ^{? 1} in 1998 and 1999, respectively. The annual accumulated CO₂ efflux was 3117 and 3326 g CO₂ m ^{? 2} in 1998 and 1999, respectively. The spatial variation in CO₂ efflux was high (CV 0.18–0.45) and increased with increasing efflux. Soil air CO₂ concentration showed similar seasonal pattern the peak concentrations occurring in July–August. The CO₂ concentrations ranged from 580 to 780 µ mol mol ^{? 1} in the humus layer to 13 620–14 470 µ mol mol ^{? 1} in the C‐horizon. In winter the soil air CO₂ concentrations were lower, especially in deeper soil layers. Drought decreased CO₂ efflux and soil air CO₂ concentration. The in situ comparison on forest soil between the chamber methods showed the non‐flow‐through chamber to give ～～50% lower efflux values than that of the flow‐through chamber. When calibrated against known CO₂ efflux ranging from 0.4 to 0.8 g CO₂ m ^{? 2} h ^{? 1} generated with a diffusion box method developed by Widén and Lindroth [Acta Universitatis Agriculturae Suecia Silvestria, 2001], the flow‐through chamber gave equal effluxes at the lower end of the calibration range, but overestimated high effluxes by 20%. Non‐flow‐through chamber underestimated the CO₂ efflux by 30%.