Seasonal and annual respiration of a ponderosa pine ecosystem

The net ecosystem exchange of CO₂ between forests and the atmosphere, measured by eddy covariance, is the small difference between two large fluxes of photosynthesis and respiration. Chamber measurements of soil surface CO₂ efflux (F_s), wood respiration (F_w) and foliage respiration (F_f) help identify the contributions of these individual components to net ecosystem exchange. Models developed from the chamber data also provide independent estimates of respiration costs. We measured CO₂ efflux with chambers periodically in 1996–97 in a ponderosa pine forest in Oregon, scaled these measurements to the ecosystem, and computed annual totals for respiration by component. We also compared estimated half-hourly ecosystem respiration at night (F_nc) with eddy covariance measurements. Mean foliage respiration normalized to 10 °C was 0.20 μmol m^–2 (hemi-leaf surface area) s^–1, and reached a maximum of 0.24 μmol m^–2 HSA s^–1 between days 162 and 208. Mean wood respiration normalized to 10 °C was 5.9 μmol m^–3 sapwood s^–1, with slightly higher rates in mid-summer, when growth occurs. There was no significant difference (P > 0.10) between wood respiration of young (45 years) and old trees (250 years). Soil surface respiration normalized to 10 °C ranged from 0.7 to 3.0 μmol m^–2 (ground) s^–1 from days 23 to 329, with the lowest rates in winter and highest rates in late spring. Annual CO₂ flux from soil surface, foliage and wood was 683, 157, and 54 g C m^–2 y^–1, with soil fluxes responsible for 76% of ecosystem respiration. The ratio of net primary production to gross primary production was 0.45, consistent with values for conifer sites in Oregon and Australia, but higher than values reported for boreal coniferous forests. Below-ground carbon allocation (root turnover and respiration, estimated as F_s– litterfall carbon) consumed 61% of GPP; high ratios such as this are typical of sites with more water and nutrient constraints. The chamber estimates were moderately correlated with change in CO₂ storage in the canopy (F_stor) on calm nights (friction velocity u* < 0.25 m s^–1; R² = 0.60); F_stor was not significantly different from summed chamber estimates. On windy nights (u* > 0.25 m s^–1), the sum of turbulent flux measured above the canopy by eddy covariance and F_stor was only weakly correlated with summed chamber estimates (R² = 0.14); the eddy covariance estimates were lower than chamber estimates by 50%.     

Soil CO2 efflux in a tropical forest in the central Amazon

This study investigated the spatial and temporal variation in soil carbon dioxide (CO₂) efflux and its relationship with soil temperature, soil moisture and rainfall in a forest near Manaus, Amazonas, Brazil. The mean rate of efflux was 6.45±0.25 SE μmol CO₂ m^?2s^?1 at 25.6±0.22 SE°C (5 cm depth) ranging from 4.35 to 9.76 μmol CO₂ m^?2s^?1; diel changes in efflux were correlated with soil temperature (r²=0.60). However, the efflux response to the diel cycle in temperature was not always a clear exponential function. During period of low soil water content, temperature in deeper layers had a better relationship with CO₂ efflux than with the temperature nearer the soil surface. Soil water content may limit CO₂ production during the drying‐down period that appeared to be an important factor controlling the efflux rate (r²=0.39). On the other hand, during the rewetting period microbial activity may be the main controlling factor, which may quickly induce very high rates of efflux. The CO₂ flux chamber was adapted to mimic the effects of rainfall on soil CO₂ efflux and the results showed that efflux rates reduced 30% immediately after a rainfall event. Measurements of the CO₂ concentration gradient in the soil profile showed a buildup in the concentration of CO₂ after rain on the top soil. This higher CO₂ concentration developed shortly after rainfall when the soil pores in the upper layers were filled with water, which created a barrier for gas exchange between the soil and the atmosphere.     

Wood CO2 efflux in a primary tropical rain forest

The balance between photosynthesis and plant respiration in tropical forests may substantially affect the global carbon cycle. Woody tissue CO₂ efflux is a major component of total plant respiration, but estimates of ecosystem‐scale rates are uncertain because of poor sampling in the upper canopy and across landscapes. To overcome these problems, we used a portable scaffolding tower to measure woody tissue CO₂ efflux from ground level to the canopy top across a range of sites of varying slope and soil phosphorus content in a primary tropical rain forest in Costa Rica. The objectives of this study were to: (1) determine whether to use surface area, volume, or biomass for modeling and extrapolating wood CO₂ efflux, (2) determine if wood CO₂ efflux varied seasonally, (3) identify if wood CO₂ efflux varied by functional group, height in canopy, soil fertility, or slope, and (4) extrapolate wood CO₂ efflux to the forest. CO₂ efflux from small diameter woody tissue (<10 cm) was related to surface area, while CO₂ efflux from stems >10 cm was related to both surface area and volume. Wood CO₂ efflux showed no evidence of seasonality over 2 years. CO₂ efflux per unit wood surface area at 25° (F_A) was highest for the N‐fixing dominant tree species Pentaclethra macroloba, followed by other tree species, lianas, then palms. Small diameter F_A increased steeply with increasing height, and large diameter F_A increased with diameter. Soil phosphorus and slope had slight, but complex effects on F_A. Wood CO₂ efflux per unit ground area was 1.34±0.36 μmol m^?2 s^?1, or 508±135 g C m^?2 yr^?1. Small diameter wood, only 15% of total woody biomass, accounted for 70% of total woody tissue CO₂ efflux from the forest; while lianas, only 3% of total woody biomass, contributed one‐fourth of the total wood CO₂ efflux.     

Soil CO2 efflux measurements in a mixed forest: impact of chamber disturbances,spatial variability and seasonal evolution

A closed‐dynamic‐chamber system (CDCS) was used to measure the spatial and seasonal variability of the soil CO₂ efflux (F_s) in beech and in Douglas fir patches of the Vielsalm forest (Belgium). First the difference between natural and measured soil CO₂ efflux induced by the presence of the CDCS was studied. The impact on the measurements of the pressure difference between the outside (natural condition) and the inside of the chamber was found to be small (0.4%). The influence of wind disturbance in the closed chamber was tested by comparison with an open‐chamber system characterized by a different wind distribution. A very good correlation between the two systems was found (r² = 0.99) but the open system yielded slightly lower fluxes than the closed one (slope = 0.88 ± 0.05). A measurement procedure has been developed to minimize the effect of the other sources of perturbation. The spatial and seasonal evolution of the soil CO₂ efflux was obtained by performing regular measurements on 29 spots in the beech patch over a period of 12 months and on 24 spots in the Douglas fir patch over 8 months. For each spot, the experimental relationship between Fs and soil temperature was compared with the fitted line for an Arrhenius relationship with a soil temperature‐dependent activation energy. Soil temperature explains 73% of the seasonal variation for all the data. The spatial average of the soil CO₂ efflux at 10 °C (Fs₁₀) in the beech patch is 2.57 ± 0.41 μmol m^?2 s^?1, approximately twice the average in the Douglas fir patch recorded at 1.42 ± 0.22 μmol m^?2 s^?1. The litter fall analysis seems to indicate that soil organic matter quality and quantity may be one the reasons for this difference. Finally the annual soil CO₂ efflux was calculated for the beech and Douglas fir patches (870 ± 140 and 438 ± 68 gC m^?2 y^?1, respectively). The beech value would represent 92 ± 15% of the annual ecosystem respiration estimated from the eddy covariance measurements.     

Forest ecosystem respiration estimated from eddy covariance and chamber measurements under high turbulence and substantial tree mortality from bark beetles

Eddy covariance nighttime fluxes are uncertain due to potential measurement biases. Many studies report eddy covariance nighttime flux lower than flux from extrapolated chamber measurements, despite corrections for low turbulence. We compared eddy covariance and chamber estimates of ecosystem respiration at the GLEES Ameriflux site over seven growing seasons under high turbulence [summer night mean friction velocity (u*) = 0.7 m s^?1], during which bark beetles killed or infested 85% of the aboveground respiring biomass. Chamber‐based estimates of ecosystem respiration during the growth season, developed from foliage, wood, and soil CO₂ efflux measurements, declined 35% after 85% of the forest basal area had been killed or impaired by bark beetles (from 7.1 ± 0.22 μmol m^?2 s^?1 in 2005 to 4.6 ± 0.16 μmol m^?2 s^?1 in 2011). Soil efflux remained at ~3.3 μmol m^?2 s^?1 throughout the mortality, while the loss of live wood and foliage and their respiration drove the decline of the chamber estimate. Eddy covariance estimates of fluxes at night remained constant over the same period, ~3.0 μmol m^?2 s^?1 for both 2005 (intact forest) and 2011 (85% basal area killed or impaired). Eddy covariance fluxes were lower than chamber estimates of ecosystem respiration (60% lower in 2005, and 32% in 2011), but the mean night estimates from the two techniques were correlated within a year (r² from 0.18 to 0.60). The difference between the two techniques was not the result of inadequate turbulence, because the results were robust to a u* filter of >0.7 m s^?1. The decline in the average seasonal difference between the two techniques was strongly correlated with overstory leaf area (r² = 0.92). The discrepancy between methods of respiration estimation should be resolved to have confidence in ecosystem carbon flux estimates.     

Seasonal patterns and environmental control of carbon dioxide and water vapour exchange in an ecotonal boreal forest

Carbon dioxide, water vapour, and sensible heat fluxes were measured above and within a spruce dominated forest near the southern ecotone of the boreal forest in Maine, USA. Summer, mid-day carbon dioxide uptake was higher than at other boreal coniferous forests, averaging about – 13 μmol CO₂ m^–2 s^–1. Nocturnal summer ecosystem respiration averaged ≈ 6 μmol CO₂ m^–2 s^–1 at a mean temperature of ≈ 15 °C. Significant ecosystem C uptake began with the thawing of the soil in early April and was abruptly reduced by the first autumn frost in early October. Half-hourly forest CO₂ exchange was regulated mostly by the incident photosynthetically active photon flux density (PPFD). In addition to the threshold effects of freezing temperatures, there were seasonal effects on the inferred photosynthetic parameters of the forest canopy. The functional response of this forest to environmental variation was similar to that of other spruce forests. In contrast to reports of carbon loss from northerly boreal forest sites, in 1996 the Howland forest was a strong carbon sink, storing about 2.1 t C ha^–1.     

Scaling relationships for woody tissue respiration in two tropical rain forests

The relationship between gross primary productivity (GPP) and net primary productivity (NPP) is not fully understood. One of the uncertainties relevant to this issue is the magnitude of woody tissue respiration. Although some data exist for temperate and boreal zones, measurements of woody tissue respiration in tropical forests are sparse. We made in situ chamber measurements of woody tissue respiration in two tropical rain forests, one in the Brazilian Amazon (Reserva Jarú) and one in Central Cameroon (Mbalmayo Reserve). We made measurements on a wide range of species at each site and over a range of stem diameters from 0·02 to 1·4 m. The rate of efflux of carbon dioxide (CO₂) from bark at 25 °C, R_t, varied from 0·1 to 5·2 µmol m^?2 s^?1 across the two sites, and the efflux was related to both volume and surface area components of the measured stem sections. The temperature response in R_t was slightly higher at Jarú than at Mbalmayo, with Q₁₀ values of 1·8 (± 0·1 SE) and 1·6 (± 0·1 SE), respectively. A log–log regression showed that R_t was significantly related to stem diameter, D (P < 0·001; r² = 0·58–0·62) and was significantly higher at Mbalmayo than at Jarú (P < 0·001), but that the rate of increase in R_t with stem diameter, D, was similar between sites. At the Mbalmayo site, tree growth measurements made over a 4 month period were used to make two estimates of the maintenance (R_m) and construction (R_c) components of respiration embedded in R_t. The two methods agreed closely, suggesting that R_m was approximately 80% of R_c at this site. R_m could be strongly related to D using a sigmoidal relationship that described both surface area and volume components as sources of respiratory CO₂ (r² = 0·71). This functional model was combined with inventory, growth and climate data for the Mbalmayo site to make a first estimate of annual above‐ground woody tissue respiration, R_A, which was 257 (± 18 SE) g C m^?2 year^?1. This value corresponds to approximately 10% of GPP, slightly lower than that found for another tropical rain forest, but higher than for temperate forests. When combined with data from six other sites in tropical, temperate and boreal settings, a very strong relationship was found between R_A and leaf area index (LAI), and between R_A/GPP and LAI (P < 0·001, r² = 0·98). This indicates that R_A exerts an appreciable constraint on NPP and that this constraint varies closely with LAI across widely differing types of woody vegetation.     

Woody-tissue respiration for Simarouba amara and Minquartia guianensis,two tropical wet forest trees with different growth habits

We measured CO₂ efflux from stems of two tropical wet forest trees, both found in the canopy, but with very different growth habits. The species were Simarouba amara, a fast-growing species associated with gaps in old-growth forest and abundant in secondary forest, and Minquartia guianensis, a slow-growing species tolerant of low-light conditions in old-growth forest. Per unit of bole surface, CO₂ efflux averaged 1.24 mol m^–2 s^–1 for Simarouba and 0.83 mol m^–2s^–1 for Minquartia. CO₂ efflux was highly correlated with annual wood production (r ²=0.65), but only weakly correlated with stem diameter (r ²=0.22). We also partitioned the CO₂ efflux into the functional components of construction and maintenance respiration. Construction respiration was estimated from annual stem dry matter production and maintenance respiration by subtracting construction respiration from the instantaneous CO₂ flux. Estimated maintenance respiration was linearly related to sapwood volume (39.6 mol m^–3s^–1 at 24.6° C, r ²=0.58), with no difference in the rate for the two species. Maintenance respiration per unit of sapwood volume for these tropical wet forest trees was roughly twice that of temperate conifers. A model combining construction and maintenance respiration estimated CO₂ very well for these species (r ²=0.85). For our sample, maintenance respiration was 54% of the total CO₂ efflux for Simarouba and 82% for Minquartia. For our sample, sapwood volume averaged 23% of stem volume when weighted by tree size, or 40% with no size weighting. Using these fractions, and a published estimate of aboveground dry-matter production, we estimate the annual cost of woody tissue respiration for primary forest at La Selva to be 220 or 350 g C m^–2 year^–1, depending on the assumed sapwood volume. These costs are estimated to be less than 13% of the gross production for the forest.     

Seasonal respiration of foliage, fine roots, and woody tissues in relation to growth, tissue N, and photosynthesis

Autotrophic respiration may regulate how ecosystem productivity responds to changes in temperature, atmospheric [CO₂] and N deposition. Estimates of autotrophic respiration are difficult for forest ecosystems, because of the large amount of biomass, different metabolic rates among tissues, and seasonal variation in respiration rates. We examined spatial and seasonal patterns in autotrophic respiration in a Pinus strobus ecosystem, and hypothesized that seasonal patterns in respiration rates at a common temperature would vary with [N] for fully expanded foliage and fine roots, with photosynthesis for foliage, and with growth for woody tissues (stems, branches, and coarse roots). We also hypothesized that differences in [N] would largely explain differences in maintenance or dormant‐season respiration among tissues. For April–November, mean respiration at 15 °C varied from 1.5 to 2.8 μmol kg^?1 s^?1 for fully expanded foliage, 1.7–3.0 for growing foliage, 0.8–1.6 for fine roots, 0.6–1.1 (sapwood) for stems, 0.5–1.8 (sapwood) for branches, and 0.2–1.5 (sapwood) for coarse roots. Growing season variation in respiration for foliage produced the prior year was strongly related to [N] (r² = 0.94), but fine root respiration was not related to [N]. For current‐year needles, respiration did not covary with [N]. Night‐time foliar respiration did not vary in concert with previous‐day photosynthesis for either growing or fully expanded needles. Stem growth explained about one‐third of the seasonal variation in stem respiration (r² = 0.38), and also variation among trees (r² = 0.43). We did not determine the cause of seasonal variation in branch and coarse root respiration, but it is unlikely to be directly related to growth, as the pattern of respiration in coarse roots and branches was not synchronized with stem growth. Seasonal variations in temperature‐corrected respiration rates were not synchronized among tissues, except foliage and branches. Spatial variability in dormant‐season respiration rates was significantly related to tissue N content in foliage (r² = 0.67), stems (r² = 0.45), coarse roots (r² = 0.36), and all tissues combined (r² = 0.83), but not for fine roots and branches. Per unit N, rates for P. strobus varied from 0.22 to 3.4 μmol molN^?1 s^?1 at 15 °C, comparable to those found for other conifers. Accurate estimates of annual autotrophic respiration should reflect seasonal and spatial variation in respiration rates of individual tissues.     

Variations in leaf respiration across different seasons for Mediterranean evergreen species

Leaf respiration (R _L) of evergreen species co-occurring in the Mediterranean maquis developing along the Latium coast was analyzed. The results on the whole showed that the considered evergreen species had the same R _L trend during the year, with the lowest rates [0.83 ± 0.43 μmol(CO₂) m^?2 s^?1, mean value of the considered species] in winter, in response to low air temperatures. Higher R _L were reached in spring [2.44 ± 1.00 μmol(CO₂) m^?2 s^?1, mean value] during the favorable period, and in summer [3.17 ± 0.89 μmol(CO₂) m^?2 s^?1] during drought. The results of the regression analysis showed that 42% of R _L variations depended on mean air temperature and 13% on total monthly rainfall. Among the considered species, C. incanus, was characterized by the highest R _L in drought [4.93 ± 0.27 μmol(CO₂) m^?2 s^?1], low leaf water potential at predawn (Ψ_pd= ?1.08 ± 0.18 MPa) and midday (Ψ_md = ?2.75 ± 0.11 MPa) and low relative water content at predawn (RWC_pd = 80.5 ± 3.4%) and midday (RWC_md = 67.1 ± 4.6%). Compared to C. incanus, the sclerophyllous species (Q. ilex, P. latifolia, P. lentiscus, A. unedo) and the liana (S. aspera), had lower R _L [2.72 ± 0.66 μmol(CO₂) m^?2 s^?1, mean value of the considered species], higher RWC_pd (91.8 ± 1.8%), RWC_md (82.4 ± 3.2%), Ψ_pd (?0.65 ± 0.28 MPa) and Ψ_md (?2.85 ± 1.20 MPa) in drought. The narrow-leaved species (E. multiflora, R. officinalis, and E. arborea) were in the middle. The coefficients, proportional to the respiration increase for each 10°C rise (Q₁₀), ranging from 1.49 (E. arborea) to 1.98 (A. unedo) were indicative of the different sensitivities of the considered species to air temperature variation.     

Photosynthesis of Marine Macroalgae from Antarctica: Light and Temperature Requirements

The photosynthetic performance of macroalgae isolated in Antarctica was studied in the laboratory. Species investigated were the brown algae Himantothallus grandifolius, Desmarestia anceps, Ascoseira mirabilis, the red algae Palmaria decipiens, Iridaea cordata, Gigartina skottsbergii, and the green algae Enteromorpha bulbosa, Acrosiphonia arcta, Ulothrix subflaccida and U. implexa. Unialgal cultures of the brown and red algae were maintained at 0°C, the green algae were cultivated at 10°C. I_K values were between 18 and 53 μmol m^?2 s^?1 characteristic or low light adapted algae. Only the two Ulothrix species showed higher I_K values between 70 and 74 μmol m^?2 s^?1. Photosynthesis compensated dark respiration at very low photon fluence rates between 1.6 and 10.6 μmol m^?2 s^?1. Values of α were high: between 0.4 and 1.1 μmol O₂ g^?1 FW h^?1 (μmol m^?2 s^?1)^?1 in the brown and red algae and between 2.1 and 4.9 μmol O₂ g^?1 FW h^?1 (μmol m^?2 s^?1)^?1 in the green algal species. At 0°C P_max values of the brown and red algae ranged from 6.8 to 19.1 μmol O₂ g^?1 FW h^?1 and were similarly high or higher than those of comparable Arctic-cold temperate species. Optimum temperatures for photosynthesis were 5 to 10°C in A. mirabilis, 10°C in H. grandifolius, 15°C in G. skottsbergii and 20°C or higher in D. anceps and I. cordata. P: R ratios strongly decreased in most brown and red algae with increasing temperatures due to different Q₁₀ values for photosynthesis (1.4 to 2.5) and dark respiration (2.5 to 4.1). These features indicate considerable physiological adaptation to the prevailing low light conditions and temperatures of Antarctic waters. In this respect the lower depth distribution limits and the northern distribution boundaries of these species partly depend on the physiological properties described here.     

Fluxes of carbon, water and energy over Brazilian cerrado: an analysis using eddy covariance and stable isotopes

We present the energy and mass balance of cerrado sensu stricto (a Brazilian form of savanna), in which a mixture of shrubs, trees and grasses forms a vegetation with a leaf area index of 1·0 in the wet season and 0·4 in the dry season. In the wet season the available energy was equally dissipated between sensible heat and evaporation, but in the dry season at high irradiance the sensible heat greatly exceeded evaporation. Ecosystem surface conductance g_s in the wet season rose abruptly to 0·3 mol m^?2 s^?1 and fell gradually as the day progressed. Much of the total variation in g_s was associated with variation in the leaf-to-air vapour pressure deficit of water and the solar irradiance. In the dry season the maximal g_s values were only 0·1 mol m^?2 s^?1. Maximal net ecosystem fluxes of CO₂ in the wet and dry season were –10 and –15 μmol CO₂ m^?2 s^?1, respectively (sign convention: negative denotes fluxes from atmosphere to vegetation). The canopy was well coupled to the atmosphere, and there was rarely a significant build-up of respiratory CO₂ during the night. For observations in the wet season, the vegetation was a carbon dioxide sink, of maximal strength 0·15 mol m^?2 d^?1. However, it was a source of carbon dioxide for a brief period at the height of the dry season. Leaf carbon isotopic composition showed all the grasses except for one species to be C₄, and all the palms and woody plants to be C₃. The CO₂ coming from the soil had an isotopic composition that suggested 40% of it was of C₄ origin.     

Large seasonal variation of soil respiration in a secondary tropical moist forest in Puerto Rico

Tropical forests are the largest contributors to global emissions of carbon dioxide (CO₂) to the atmosphere via soil respiration (R_s). As such, identifying the main controls on R_s in tropical forests is essential for accurately projecting the consequences of ongoing and future global environmental changes to the global C cycle. We measured hourly R_s in a secondary tropical moist forest in Puerto Rico over a 3‐year period to (a) quantify the magnitude of R_s and (b) identify the role of climatic, substrate, and nutrient controls on the seasonality of R_s. Across 3 years of measurements, mean R_s was 7.16 ± 0.02 μmol CO₂ m^‐2 s^‐1 (or 2,710 g C m^‐2 year^‐1) and showed significant seasonal variation. Despite small month‐to‐month variation in temperature (~4°C), we found significant positive relationships between daily and monthly R_s with both air and soil temperature, highlighting the importance of temperature as a driver of R_s even in warm ecosystems, such as tropical forests. We also found a significant parabolic relationship between mean daily volumetric soil moisture and mean daily R_s, with an optimal moisture value of 0.34 m³ m^‐3. Given the relatively consistent climate at this site, the large range in mean monthly R_s (~7 μmol CO₂ m^‐2 s^‐1) was surprising and suggests that even small changes in climate can have large implications for ecosystem respiration. The strong positive relationship of R_s with temperature at monthly timescales particularly stands out, as moisture is usually considered a stronger control of R_s in tropical forests that already experience warm temperatures year‐round. Moreover, our results revealed the strong seasonality of R_s in tropical moist forests, which given its high magnitude, can represent a significant contribution to the seasonal patterns of atmospheric (CO₂) globally.     

Topographically regulated traps of dissolved organic carbon create hotspots of soil carbon dioxide efflux in forests

Soil carbon pools are an essential but poorly understood factor in heterotrophic soil respiration on forested landscapes. We hypothesized that the topographically regulated distribution of dissolved organic carbon (DOC) is the dominant factor contributing to soil CO₂ efflux. We tested this hypothesis by monitoring soil CO₂ efflux and sampling particulate and dissolved substrates (both mobile DOC in soil solution and DOC potentially sorbed onto Fe and Al oxyhydroxides) in surface (freshly fallen leaves (FFL) and forest floor) and near-surface (A-horizon or top 10 cm of peat) soils along three hillslope transects (15°, 25° and 35° slopes) that included upland (crest, shoulder, backslope, footslope, and toeslope) and wetland (periphery and central) topographic features during the snowfree season within a sugar maple forest. We observed that median snowfree season soil CO₂ efflux ranged from <1 to >5 μmol CO₂ m^?2 s^?1. Substrates in the near-surface mineral soil were most strongly related to median soil CO₂ efflux, and when combined mobile DOC and sorbed DOC together explained 78% of the heterogeneity in median soil CO₂ efflux (p < 0.001). When the carbon pool in FFL (an important source of DOC to the forest soils) was included, the explanation of variance increased to 81% (p < 0.001). Topographically regulated processes created high concentrations of mobile DOC at the footslope, and high concentrations of sorbed DOC further downslope at the toeslope, forming distinct traps of DOC that can become hotspots for soil CO₂ production. A reduction in the uncertainty of forest carbon budgets can be achieved by taking into consideration the topographic regulation of the substrates contributing to soil CO₂ efflux.     

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.     

Annual cycles of water vapour and carbon dioxide fluxes in and above a boreal aspen forest

Water vapour and CO₂ fluxes were measured using the eddy correlation method above and below the overstorey of a 21-m tall aspen stand in the boreal forest of central Saskatchewan as part of the Boreal Ecosystem-Atmosphere Study (BOREAS). Measurements were made at the 39.5-m and 4-m heights using 3-dimensional sonic anemometers (Kaijo-Denki and Solent, respectively) and closed-path gas analysers (LI-COR 6262) with 6-m and 4.7-m long heated sampling tubing, respectively. Continuous measurements were made from early October to mid-November 1993 and from early February to late-September 1994. Soil CO₂ flux (respiration) was measured using a LI-COR 6000-09 soil chamber and soil evaporation was measured using Iysimetry. The leaf area index of the aspen and hazelnut understorey reached 1.8 and 3.3, respectively. The maximum daily evapotranspiration (E) rate was 5–6 mm d^?1. Following leaf-out the hazelnut and soil accounted for 22% of the forest E. The estimated total E was 403 mm for 1994. About 88% of the precipitation in 1994 was lost as evapotranspiration. During the growing season, the magnitude of half-hourly eddy fluxes of CO₂ from the atmosphere into the forest reached 1.2 mg CO₂ m^?2 s^?1 (33 μmol C m^?2 s^?1) during the daytime. Downward eddy fluxes at the 4-m height were observed when the hazelnut was growing rapidly in June and July. Under well-ventilated night-time conditions, the eddy fluxes of CO₂ above the aspen and hazelnut, corrected for canopy storage, increased exponentially with soil temperature at the 2-cm depth. Estimates of daytime respiration rates using these relationships agreed well with soil chamber measurements. During the 1994 growing season, the cumulative net ecosystem exchange (NEE) was -3.5 t C ha^?1 y^?1 (a net gain by the system). For 1994, cumulative NEE, ecosystem respiration (R) and gross ecosystem photosynthesis (GEP = R - NEE) were estimated to be -1.3, 8.9 and 10.2 t C ha^?1 y^?1 respectively. Gross photosynthesis of the hazelnut was 32% of GEP.     

Soil-surface carbon dioxide efflux and microbial biomass in relation to tree density 13 years after a stand replacing fire in a lodgepole pine ecosystem

The effects of fire on soil‐surface carbon dioxide (CO₂) efflux, F_S, and microbial biomass carbon, C_mic, were studied in a wildland setting by examining 13‐year‐old postfire stands of lodgepole pine differing in tree density (< 500 to > 500 000 trees ha^?1) in Yellowstone National Park (YNP). In addition, young stands were compared to mature lodgepole pine stands (～110‐year‐old) in order to estimate ecosystem recovery 13 years after a stand replacing fire. Growing season F_S increased with tree density in young stands (1.0 µmol CO₂ m^?2 s^?1 in low‐density stands, 1.8 µmol CO₂ m^?2 s^?1 in moderate‐density stands and 2.1 µmol CO₂ m^?2 s^?1 in high‐density stands) and with stand age (2.7 µmol CO₂ m^?2 s^?1 in mature stands). Microbial biomass carbon in young stands did not differ with tree density and ranged from 0.2 to 0.5 mg C g^?1 dry soil over the growing season; C_mic was significantly greater in mature stands (0.5–0.8 mg C g^?1 dry soil). Soil‐surface CO₂ efflux in young stands was correlated with biotic variables (above‐ground, below‐ground and microbial biomass), but not with abiotic variables (litter and mineral soil C and N content, bulk density and soil texture). Microbial biomass carbon was correlated with below‐ground plant biomass and not with soil carbon and nitrogen, indicating that plant activity controls not only root respiration, but C_mic pools and overall F_S rates as well. These findings support recent studies that have demonstrated the prevailing importance of plants in controlling rates of F_S and suggest that decomposition of older, recalcitrant soil C pools in this ecosystem is relatively unimportant 13 years after a stand replacing fire. Our results also indicate that realistic predictions and modeling of terrestrial C cycling must account for the variability in tree density and stand age that exists across the landscape as a result of natural disturbances.     

Soil‐surface CO2 efflux and its spatial and temporal variations in a young ponderosa pine plantation in northern California

Soil‐surface CO₂ efflux and its spatial and temporal variations were examined in an 8‐y‐old ponderosa pine plantation in the Sierra Nevada Mountains in California from June 1998 to August 1999. Continuous measurements of soil CO₂ efflux, soil temperatures and moisture were conducted on two 20 × 20 m sampling plots. Microbial biomass, fine root biomass, and the physical and chemical properties of the soil were also measured at each of the 18 sampling locations on the plots. It was found that the mean soil CO₂ efflux in the plantation was 4.43 µmol m^?2 s^?1 in the growing season and 3.12 µmol m^?2 s^?1 in the nongrowing season. These values are in the upper part of the range of published soil‐surface CO₂ efflux data. The annual maximum and minimum CO₂ efflux were 5.87 and 1.67 µmol m^?2 s^?1, respectively, with the maximum occurring between the end of May and early June and the minimum in December. The diurnal fluctuation of CO₂ efflux was relatively small (< 20%) with the minimum appearing around 09.00 hours and the maximum around 14.00 hours. Using daytime measurements of soil CO₂ efflux tends to overestimate the daily mean soil CO₂ efflux by 4–6%. The measurements taken between 09.00 and 11.00 hours (local time) seem to better represent the daily mean with a reduced sampling error of 0.9–1.5%. The spatial variation of soil CO₂ efflux among the 18 sampling points was high, with a coefficient of variation of approximately 30%. Most (84%) of the spatial variation was explained by fine root biomass, microbial biomass, and soil physical and chemical properties. Although soil temperature and moisture explained most of the temporal variations (76–95%) of soil CO₂ efflux, the two variables together explained less than 34% of the spatial variation. Microbial biomass, fine root biomass, soil nitrogen content, organic matter content, and magnesium content were significantly and positively correlated with soil CO₂ efflux, whereas bulk density and pH value were negatively correlated with CO₂ efflux. The relationship between soil CO₂ efflux and soil temperature was significantly controlled by soil moisture with a Q₁₀ value of 1.4 when soil moisture was <14% and 1.8 when soil moisture was >14%. Understanding the spatial and temporal variations is essential to accurately assessment of carbon budget at whole ecosystem and landscape scales. Thus, this study bears important implications for the study of large‐scale ecosystem dynamics, particularly in response to climatic variations and management regimes.     

The contribution of bryophytes to the carbon exchange for a temperate rainforest

Bryophytes blanket the floor of temperate rainforests in New Zealand and may influence a number of important ecosystem processes, including carbon cycling. Their contribution to forest floor carbon exchange was determined in a mature, undisturbed podocarp‐broadleaved forest in New Zealand, dominated by 100–400‐year‐old rimu (Dacrydium cupressimum) trees. Eight species of mosses and 13 species of liverworts contributed to the 62% cover of the diverse forest floor community. The bryophyte community developed a relatively thin (depth <30 mm), but dense, canopy that experienced elevated CO₂ partial pressures (median 46.6 Pa immediately below the bryophyte canopy) relative to the surrounding air (median 37.6 Pa at 100 mm above the canopy). Light‐saturated rates of net CO₂ exchange from 14 microcosms collected from the forest floor were highly variable; the maximum rate of net uptake (bryophyte photosynthesis – whole‐plant respiration) per unit ground area at saturating irradiance was 1.9 μmol m^?2 s^?1 and in one microcosm, the net rate of CO₂ exchange was negative (respiration). CO₂ exchange for all microcosms was strongly dependent on water content. The average water content in the microcosms ranged from 1375% when fully saturated to 250% when air‐dried. Reduction in water content across this range resulted in an average decrease of 85% in net CO₂ uptake per unit ground area. The results from the microcosms were used in a model to estimate annual carbon exchange for the forest floor. This model incorporated hourly variability in average irradiance reaching the forest floor, water content of the bryophyte layer, and air and soil temperature. The annual net carbon uptake by forest floor bryophytes was 103 g m^?2, compared to annual carbon efflux from the forest floor (bryophyte and soil respiration) of ?1010 g m^?2. To put this in perspective of the magnitude of the components of CO₂ exchange for the forest floor, the bryophyte layer reclaimed an amount of CO₂ equivalent to only about 10% of forest floor respiration (bryophyte plus soil) or ～11% of soil respiration. The contribution of forest floor bryophytes to productivity in this temperate rainforest was much smaller than in boreal forests, possibly because of differences in species composition and environmental limitations to photosynthesis. Because of their close dependence on water table depth, the contribution of the bryophyte community to ecosystem CO₂ exchange may be highly responsive to rapid changes in climate.     

秦岭火地塘林区油松(Pinus tabulaeformis)林休眠期的土壤呼吸

林木休眠期林地土壤CO2释放是森林生态系统碳平衡关键组成部分之一.由于绝大多数森林生态系统林木休眠期土壤CO2释放过程测定困难,国内有关林木休眠期CO2释放,量化方面的研究开展较少.采用动态开路气室法对秦岭火地塘林区天然次生油松(Pinus tabulaeformis)林土壤呼吸的日变化进行了测定,分析了土壤呼吸速率(mgCO2m-2h-1)与土壤温度和体积含水率的关系,基于土壤日均呼吸速率和土壤日均温度指数方程与观测季的总天数,估算了林木休眠期林地土壤CO2释放量.结果表明:(1)研究区林地土壤呼吸速率存在较大的时、空变异.不同观测部位土壤呼吸速率的峰值出现时间各异,呼吸作用较弱的时段也不一致.同一观测部位不同观测月中,土壤日均呼吸速率变异系数分别为48.38%,82.51%和81.88%;(2)当土温＞8.5 ℃时,0～5 cm和5～10 cm土层,土壤日均温与土壤日均呼吸速率间存在极显著(p＜0.001)的指数关系,Q10分别为1.297和1.323;(3)0～5 cm和5～10 cm土层,土壤体积含水率与土壤呼吸速率间关系复杂;(4) 林木休眠期研究区林地土壤CO2释放量变化于(977.37±88.43)～(997.19±80.73) gCm-2(p=0.005)间.