Annual soil respiration in broadleaf forests of northern Wisconsin: influence of moisture and site biological,chemical, and physical characteristics

Soil temperature and moisture influence soil respiration at a range of temporal and spatial scales. Although soil temperature and moisture may be seasonally correlated, intra and inter-annual variations in soil moisture do occur. There are few direct observations of the influence of local variation in species composition or other stand/site characteristics on seasonal and annual variations in soil moisture, and on cumulative annual soil carbon release. Soil climate and soil respiration from twelve sites in five different forest types were monitored over a 2-year period (1998–1999). Also measured were stand age, species composition, basal area, litter inputs, total above-ground wood production, leaf area index, forest floor mass, coarse and fine root mass, forest floor carbon and nitrogen concentration, root carbon and nitrogen concentration, soil carbon and nitrogen concentration, coarse fraction mass and volume, and soil texture. General soil respiration models were developed using soil temperature, daily soil moisture, and various site/soil characteristics. Of the site/soil characteristics, above-ground production, soil texture, roots + forest floor mass, roots + forest floor carbon:nitrogen, and soil carbon:nitrogen were significant predictors of soil respiration when used alone in respiration models; all of these site variables were weakly to moderately correlated with mean site soil moisture. Daily soil climate data were used to estimate the annual release of carbon (C) from soil respiration for the period 1998–1999. Mean annual soil temperature did not differ between the 2 years but mean annual soil moisture was approximately 9% lower in 1998 due to a summer drought. Soil C respired during 1998 ranged from 8.57 to 11.43 Mg C ha⁻¹ yr⁻¹ while the same sites released 10.13 and 13.57 Mg C ha⁻¹ yr⁻¹ in 1999; inter-annual differences of 15.41 and 15.73%, respectively. Among the 12 sites studied, we calculated that the depression of soil respiration linked to the drought caused annual differences of soil respiration from 11.00 to 15.78%. Annual estimates of respired soil C decreased with increasing site mean soil moisture. Similarly, the difference of respired carbon between the drought and the non-drought years generally decreased with increasing site mean soil moisture.     

Seasonal patterns in soil surface CO<Subscript>2</Subscript> flux under snow cover in 50 and 300 year old subalpine forests

Soil CO₂ flux can contribute as much as 60–80% of total ecosystem respiration in forests. Although considerable research has focused on quantifying this flux during the growing season, comparatively little effort has focused on non-growing season fluxes. We measured soil CO₂ efflux through snow in 50 and ~300 year old subalpine forest stands near Fraser CO. Our objectives were to quantify seasonal patterns in wintertime soil CO₂ flux; determine if differences in soil CO₂ flux between the two forest ages during the growing season persist during winter; and to quantify the sample size necessary to discern treatment differences. Soil CO₂ flux during the 2002–2003 and 2003–2004 snow season averaged 0.31 and 0.35 μmols m⁻² s⁻¹ for the young and old forests respectively; similar to the relative difference observed during summer. There was a significant seasonal pattern of soil CO₂ flux during the winter with fluxes averaging 0.22 μmols m⁻² s⁻¹ in December and January and increasing to an average of 0.61 μmols m⁻² s⁻¹ in May. Within-plot variability for measurements used in calculating flux was low. The coefficients of variation (CV) for CO₂ concentration, snowpack density, and snow depth were 17, 8 and 14%, respectively, yielding a CV for flux measurements within-plot of 29%. A within plot CV of 29% requires 8 sub-samples per plot to estimate the mean flux with a standard error of ±10% of the mean. Variability in CO₂ flux estimates among plots (size = 400 m²) was similar to that within plot and was also low (CV = ~28%). With a CV of 28% among plots, ten plots per treatment would have a 50% probability of detecting a 25% difference in treatment means for α = 0.05.     

小兴安岭5种林型土壤呼吸时空变异

原始阔叶红松林、谷地云冷杉林、阔叶红松择伐林、次生白桦林、人工落叶松林是小兴安岭乃至东北地区的重要森林类型。采用红外气体分析法比较测定了这几种森林类型的土壤呼吸及其相关环境因子,分析探讨了这几种森林类型土壤呼吸的时空变异。结果表明:各林型土壤呼吸与5 cm深土壤温度(T5)呈显著的指数相关,并且土壤呼吸与土壤温度、土壤湿度及其相互作用的回归模型可以解释各林型土壤呼吸约71%的季节变异。生长季平均土壤呼吸速率为次生白桦林(3.59μmolCO.2m-.2s-1)>谷地云冷杉林(3.52μmolCO.2m-.2s-1)>阔叶红松择伐林(3.44μmolCO.2m-.2s-1)>原始阔叶红松林(2.58μmolCO.2m-.2s-1)>人工落叶松林(2.29μmolCO.2m-.2s-1),说明土壤呼吸对原始阔叶红松林人为干扰的响应是不同的。各林型Q10值介于1.84(人工落叶松林)—2.32(次生白桦林)之间。在整个生长季,各林型之间土壤呼吸的变异系数变化幅度为19.74%—37.39%,而各林型内土壤环间其变化幅度为32.13%—60.20%,显著大于样地间的变化幅度14.28%—35.70%(P<0.001),说明土壤呼吸在细微尺度上的差异更大。土壤湿度可以解释各林型(阔叶红松林除外)内部土壤呼吸15.8%—33.5%的空间异质性。     

Spatial–temporal variation in soil respiration in an oak–grass savanna ecosystem in California and its partitioning into autotrophic and heterotrophic components

The spatial upscaling of soil respiration from field measurements to ecosystem levels will be biased without studying its spatial variation. We took advantage of the unique spatial gradients of an oak–grass savanna ecosystem in California, with widely spaced oak trees overlying a grass layer, to study the spatial variation in soil respiration and to use these natural gradients to partition soil respiration according to its autotrophic and heterotrophic components. We measured soil respiration along a 42.5 m transect between two oak trees in 2001 and 2002, and found that soil respiration under tree canopies decreased with distance from its base. In the open area, tree roots have no influence on soil respiration. Seasonally, soil respiration increased in spring until late April, and decreased in summer following the decrease in soil moisture content, despite the further increase in soil temperature. Soil respiration significantly increased following the rain events in autumn. During the grass growing season between November and mid-May, the average of CO₂ efflux under trees was 2.29 μmol m⁻² s⁻¹, while CO₂ efflux from the open area was 1.40 μmol m⁻² s⁻¹. We deduced that oak root respiration averaged as 0.89 μmol m⁻² s⁻¹, accounting for 39% of total soil respiration (oak root + grass root + microbes). During the dry season between mid-May and October, the average of CO₂ efflux under trees was 0.87 μmol m⁻² s⁻¹, while CO₂ efflux from the open areas was 0.51 μmol m⁻² s⁻¹. Oak root respiration was 0.36 μmol m⁻² s⁻¹, accounting for 41% of total soil respiration (oak root + microbes). The seasonal pattern of soil CO₂ efflux under trees and in open areas was simulated by a bi-variable model driven by soil temperature and moisture. The diurnal pattern was influenced by tree physiology as well. Based on the spatial gradient of soil respiration, spatial analysis of crown closure and the simulation model, we spatially and temporally upscaled chamber measurements to the ecosystem scale. We estimated that the cumulative soil respiration in 2002 was 394 gC m⁻² year⁻¹ in the open area and 616 gC m⁻² year⁻¹ under trees with a site-average of 488 gC m⁻² year⁻¹.     

Nutrient return with leaf litter fall in Fagus sylvatica forests across a soil fertility gradient

Leaf litter fall is an important nutrient flux in temperature deciduous forests which supplies a large part of the rapidly mineralisable nutrient fraction to the soil. This study investigates nutrient return with leaf litter fall in 36 old-growth forest stands of Fagus sylvatica across a broad gradient of soil fertility covering 9 mesozoic and kaenozoic parent material types (three limestones, two sandstones, two clay stones, one sand and one loess substrate). Study objectives were to analyse (i) the dependency of leaf litter nutrient concentrations on soil fertility, and (ii) the relationship between soil fertility and nutrient return with leaf litter at the stand level. Beech stands on the nine parent material types produced similar annual leaf litter masses irrespective of soil fertility or acidity. Leaf litter from the nine parent materials showed only minor variation with respect to N and K concentrations (factors of 1.5 and 1.4), moderate variation for Ca, Mg and P concentrations (factors of 2.2 to 2.9), and high variation for Al and Mn concentrations (factors of 6.7 and 10.5). Consequently, annual nutrient return with litter fall (leaf litter mass x litter nutrient concentration) was more similar among the parent materials for N (165–273 mmol m⁻² yr1) ⁻¹ and K (16–30 mm m⁻² yr⁻¹) than for Ca, P, Mg, Mn and Al. A possible explanation is increased N deposition in recent time. According to a correlation analysis, return rates of N, P, K and Mg (but not Ca) were independent of the pool size of the respective nutrient in the soil. N return rate was neither influenced by the soil pools of Nt, plant- available P (Pa) or exchangeable Ca, K and Mg, nor by soil acidity or the exchangeable Al pool. P return, in contrast, showed a negative relation to soil fertility. We hypothesize that nutrient fluxes with leaf litter fall do not necessarily reduce the fitness of tree populations as has been postulated from a tree-centred view. Rather, we suggest that nutrient fluxes with litter fall can increase, instead of decrease, plant fitness by improving nutrient availability in the densely rooted topsoil which reduces the roots’ carbon and nutrient costs of nutrient acquisition.     

Dynamics of carbon stocks in soils and detritus across chronosequences of different forest types in the Pacific Northwest, USA

We investigated variation in carbon stock in soils and detritus (forest floor and woody debris) in chronosequences that represent the range of forest types in the US Pacific Northwest. Stands range in age from <13 to >600 years. Soil carbon, to a depth of 100 cm, was highest in coastal Sitka spruce/western hemlock forests (36±10 kg C m^?2) and lowest in semiarid ponderosa pine forests (7±10 kg C m^?2). Forests distributed across the Cascade Mountains had intermediate values between 10 and 25 kg C m^?2. Soil carbon stocks were best described as a linear function of net primary productivity (r²=0.52), annual precipitation (r²=0.51), and a power function of forest floor mean residence time (r²=0.67). The highest rates of soil and detritus carbon turnover were recorded on mesic sites of Douglas‐fir/western hemlock forests in the Cascade Mountains with lower rates in wetter and drier habitats, similar to the pattern of site productivity. The relative contribution of soil and detritus carbon to total ecosystem carbon decreased as a negative exponential function of stand age to a value of ～35% between 150 and 200 years across the forest types. These age‐dependent trends in the portioning of carbon between biomass and necromass were not different among forest types. Model estimates of soil carbon storage based on decomposition of legacy carbon and carbon accumulation following stand‐replacing disturbance showed that soil carbon storage reached an asymptote between 150 and 200 years, which has significant implications to modeling carbon dynamics of the temperate coniferous forests following a stand‐replacing disturbance.     

Carbon storage in forest soil of Finland. 1. Effect of thermoclimate

A total of 30 coniferous forest sites representing two productivityclasses, forest types, were investigated on a temperature gradient(effective temperature sum using +5^°C threshold 800–1300degree-days and annual mean temperature –0.6–+3.9^°C) inFinland for studying the effect of thermoclimate on the soil C storage.Other soil forming factors were standardized within the forest types sothat the variation in the soil C density could be related to temperature.According to the applied regression model, the C density of the 0–1 mmineral soil layer increased 0.266 kg m^–2 for every 100 degree-dayincrease in the temperature sum, and the layer contained 57% and28% more C under the warmest conditions of the gradient comparedto the coolest in the less and more productive forest type, respectively.Accordingly, this soil layer was estimated to contain 23 more C ina new equilibrium with a 4^°C higher annual meantemperature in Finland. The C density of the organic layer was notassociated with temperature. Both soil layers contained more C at thesites of the more productive forest type, and the forest type explained36% and 70% of the variation in the C density of the organic and 0–1m layers, respectively. Within the forest types, the temperature sumaccounted for 33–41% of the variation in the 0–1 m layer. Theseresults suggest that site productivity is a cause for the large variation inthe soil C density within the boreal zone, and relating the soil C densityto site productivity and temperature would help to estimate the soil Creserves more accurately in the boreal zone.     

神农架海拔梯度上4种典型森林的土壤呼吸组分及其对温度的敏感性

量化森林土壤呼吸及其组分对温度的响应对准确评估未来气候变化背景下陆地生态系统的碳平衡极其重要。该文通过对神农架海拔梯度上常绿阔叶林、常绿落叶阔叶混交林、落叶阔叶林以及亚高山针叶林4种典型森林土壤呼吸的研究发现: 4种森林类型的年平均土壤呼吸速率和年平均异养呼吸速率分别为1.63、1.79、1.74、1.35 μmol CO₂·m^-2·s^-1和1.13、1.12、1.12、0.80 μmol CO₂·m^-2·s^-1。该地区的土壤呼吸及其组分呈现出明显的季节动态, 夏季最高, 冬季最低。4种森林类型中, 阔叶林的土壤呼吸显著高于针叶林, 但阔叶林之间的土壤呼吸差异不显著。土壤温度是影响土壤呼吸及其组分的主要因素, 二者呈显著的指数关系; 土壤含水量与土壤呼吸之间没有显著的相关关系。4种典型森林土壤呼吸的Q₁₀值分别为2.38、2.68、2.99和4.24, 随海拔的升高土壤呼吸对温度的敏感性增强, Q₁₀值随海拔的升高而增加。     

Response of soil respiration to simulated N deposition in a disturbed and a rehabilitated tropical forest in southern China

Responses of soil respiration (CO₂ emission) to simulated N deposition were studied in a disturbed (reforested forest with previous understory and litter harvesting) and a rehabilitated (reforested forest with no understory and litter harvesting) tropical forest in southern China from October 2005 to September 2006. The objectives of the study were to test the following hypotheses: (1) soil respiration is higher in rehabilitated forest than in disturbed forest; (2) soil respiration in both rehabilitated and disturbed tropical forests is stimulated by N additions; and (3) soil respiration is more sensitive to N addition in disturbed forest than in rehabilitated forest due to relatively low soil nutrient status in the former, resulting from different previous human disturbance. Static chamber and gas chromatography techniques were employed to quantify the soil respiration, following different N treatments (Control, no N addition; Low-N, 5 g N m⁻² year⁻¹; Medium-N, 10 g N m⁻² year⁻¹), which had been applied continuously for 26 months before the respiration measurement. Results showed that soil respiration exhibited a strong seasonal pattern, with the highest rates observed in the hot and wet growing season (April–September) and the lowest rates in winter (December–February) in both rehabilitated and disturbed forests. Soil respiration rates exhibited significant positive exponential relationship with soil temperature and significant positive linear relationship with soil moisture. Soil respiration was also significantly higher in the rehabilitated forest than in the disturbed forest. Annual mean soil respiration rate in the rehabilitated forest was 20% lower in low-N plots (71 ± 4 mg CO₂-C m⁻² h⁻¹) and 10% lower in medium-N plots (80 ± 4 mg CO₂-C m⁻² h⁻¹) than in the control plots (89 ± 5 mg CO₂-C m⁻² h⁻¹), and the differences between the control and low-N or medium-N treatments were statistically significant. In disturbed forest, annual mean soil respiration rate was 5% lower in low-N plots (63 ± 3 mg CO₂-C m⁻² h⁻¹) and 8% lower in medium-N plots (61 ± 3 mg CO₂-C m⁻² h⁻¹) than in the control plots (66 ± 4 mg CO₂-C m⁻² h⁻¹), but the differences among treatments were not significant. The depressed effects of experimental N deposition occurred mostly in the hot and wet growing season. Our results suggest that response of soil respiration to elevated N deposition in the reforested tropical forests may vary depending on the status of human disturbance. Responsible Editor: Hans Lambers.     

Supply-side controls on soil respiration among Oregon forests

To test the hypothesis that variation in soil respiration is related to plant production across a diverse forested landscape, we compared annual soil respiration rates with net primary production and the subsequent allocation of carbon to various ecosystem pools, including leaves, fine roots, forests floor, and mineral soil for 36 independent plots arranged as three replicates of four age classes in three climatically distinct forest types. Across all plots, annual soil respiration was not correlated with aboveground net primary production (R²=0.06, P>0.1) but it was moderately correlated with belowground net primary production (R²=0.46, P<0.001). Despite the wide range in temperature and precipitation regimes experienced by these forests, all exhibited similar soil respiration per unit live fine root biomass, with about 5 g of carbon respired each year per 1 g of fine root carbon (R²=0.45, P<0.001). Annual soil respiration was only weakly correlated with dead carbon pools such as forest floor and mineral soil carbon (R²=0.14 and 0.12, respectively). Trends between soil respiration, production, and root mass among age classes within forest type were inconsistent and do not always reflect cross‐site trends. These results are consistent with a growing appreciation that soil respiration is strongly influenced by the supply of carbohydrates to roots and the rhizosphere, and that some regional patterns of soil respiration may depend more on belowground carbon allocation than the abiotic constraints imposed on subsequent metabolism.     

Quantitative evaluation of atmospheric CO2 sink into forest soils from the tropics to the boreal zone during the past three decades

A model of soil carbon cycling in forest ecosystems was applied to predict the soil carbon balance in nine forest ecosystems from the tropics to the boreal zone during the past three decades (1965–95). The parameters of carbon flows and initial conditions of carbon pools were decided based on data obtained in each forest stand. Assumptions for model calculation were: (i) primary production (i.e. litterfall and root turnover rates) increased with increasing CO₂ concentrations in the atmosphere (10% per 40 p.p.m. CO₂); and (ii) temperature increased by 0.6°C per 100 years, but precipitation changed little. The simulation employed a daily time step and used daily air temperature and precipitation observed near each forest stand over an average year during the last decade. The model calculations suggest that the accumulation of total soil carbon increased 8.5–10.4 tC (ton of carbon) ha^–1 in broad-leaved forests from the tropics to the cool-temperate zone during the past three decades, but the amount of soil carbon (3.0–8.4 tC ha^–1) increased much less in needle forests from the subtropical to boreal zones during the same period. There is a linear relationship between the increasing rate of soil carbon stock during the past three decades (1965–95) in forest stands concerned (R_MS, % per 30 years) and annual mean temperature of their soils (T₀,°C), as: R_MS = 0.34T₀ + 4.1. Based on the data of carbon stock in forest soil in each climate zone reported, the global sink of atmospheric CO₂ into forest soil was roughly estimated to be 42 GtC (billion tons of carbon) per 30 years, which was 1.4 GtC year^–1 on average over the past three decades.     

Soil respiration of forest ecosystems in Japan and global implications

Within terrestrial ecosystems, soil respiration is one of the largest carbon flux components. We discuss the factors controlling soil respiration, while focusing on research conducted at the Takayama Experimental Site. Soil respiration was affected by soil temperature, soil moisture, rainfall events, typhoons, and root respiration. We consider the temporal and spatial variability of soil respiration at the Takayama Experimental Site and review the variability of annual soil respiration in Japanese forests. In the 26 compiled studies, the values of annual soil respiration ranged from 203 to 1,290 g C m⁻² year⁻¹, with a mean value of 669 g C m⁻² year⁻¹ (SD=264, CV=40). We note the need for more studies and data synthesis for the accurate prediction of soil respiration and soil carbon dynamics in Japanese forests. Finally, several methods for measuring soil respiration rates are compared and the implications of soil respiration rates for global climate change are discussed.     

Impact of forest conversion to agriculture on carbon and nitrogen mineralization in subarctic Alaska

Land-use change is likely to be a major component of global change at high latitudes, potentially causing significant alterations in soil C and N cycling. We addressed the biogeochemical impacts of land-use change in fully replicated black spruce forests and agricultural fields of different ages (following deforestation) and under different management regimes in interior Alaska. Change from forests to cultivated fields increased summer temperatures in surface soil layers by 4–5 °C, and lengthened the season of biological activity by two to three weeks. Decomposition of a common substrate (oat stubble) was enhanced by 25% in fields compared to forests after litter bags were buried for one year. In-situ net N mineralization rates in site-specific soil were similar in forests and fields during summer, but during winter, forests were the only sites where net N immobilization occurred. Field age and management had a significant impact on C and N mineralization. Rates of annual decomposition, soil respiration and summer net N mineralization tended to be lower in young than in old fields and higher in fallow than in planted young fields. To identify the major environmental factors controlling C and N mineralization, soil temperature, moisture and N availability were studied. Decomposition and net N mineralization seemed to be mainly driven by availability of inorganic N. Soil temperature played a role only when comparing forests and fields, but not in field-to-field differences. Results from soil respiration measurements in fields confirmed low sensitivity of heterotrophic respiration, and thus decomposition to temperature. In addition, both soil respiration and net N mineralization were limited by low soil water contents. Our study showed that (1) C and N mineralization are enhanced by forest clearing in subarctic soils, and (2) N availability is more important than soil temperature in controling C and N mineralization following forest clearing. Projecting the biogeochemical impacts of land-use change at high latitudes requires an improved understanding of its interactions with other factors of global change, such as changing climate and N deposition.     

Fine Root Production and Turnover in a Norway Spruce Stand in Northern Sweden: Effects of Nitrogen and Water Manipulation

Fine root length production, biomass production, and turnover in forest floor and mineral soil (0–30 cm) layers were studied in relation to irrigated (I) and irrigated-fertilized (IL) treatments in a Norway spruce stand in northern Sweden over a 2-year period. Fine roots (<1 mm) of both spruce and understory vegetation were studied. Minirhizotrons were used to estimate fine root length production and turnover, and soil cores were used to estimate standing biomass. Turnover was estimated as both the inverse of root longevity (RTL) and the ratio of annual root length production to observed root length (RTR). RTR values of spruce roots in the forest floor in I and IL plots were 0.6 and 0.5 y⁻¹, respectively, whereas the corresponding values for RTL were 0.8 and 0.9 y⁻¹. In mineral soil, corresponding values for I, IL, and control (C) plots were 1.2, 1.2, and 0.9 y⁻¹ (RTR) and 0.9, 1.1, and 1 y⁻¹ (RTL). RTR and RTL values of understory vegetation roots were 1 and 1.1 y⁻¹, respectively. Spruce root length production in both the forest floor and the mineral soil in I plots was higher than in IL plots. The IL-treated plots gave the highest estimates of spruce fine root biomass production in the forest floor, but, for the mineral soil, the estimates obtained for the I plots were the highest. The understory vegetation fine root production in the I and IL plots was similar for both the forest floor and the mineral soil and higher (for both layers) than in C plots. Nitrogen (N) turnover in the forest floor and mineral soil layers (summed) via spruce roots in IL, I, and C plots amounted to 2.4, 2.1, and 1.3 g N m⁻² y⁻¹, and the corresponding values for field vegetation roots were 0.6, 0.5, and 0.3 g N m⁻² y⁻¹. It was concluded that fertilization increases standing root biomass, root production, and N turnover of spruce roots in both the forest floor and mineral soil. Data on understory vegetation roots are required for estimating carbon budgets in model studies.     

Allocation of carbon in a mature eucalypt forest and some effects of soil phosphorus availability

Pools and annual fluxes of carbon (C) were estimated for a mature Eucalyptus pauciflora (snowgum) forest with and without phosphorus (P) fertilizer addition to determine the effect of soil P availability on allocation of C in the stand. Aboveground biomass was estimated from allometric equations relating stem and branch diameters of individual trees to their biomass. Biomass production was calculated from annual increments in tree diameters and measurements of litterfall. Maintenance and construction respiration were calculated for each component using equations given by Ryan (1991a). Total belowground C flux was estimated from measurements of annual soil CO₂ efflux less the C content of annual litterfall (assuming forest floor and soil C were at approximate steady state for the year that soil CO₂ efflux was measured). The total C content of the standing biomass of the unfertilized stand was 138 t ha^-1, with approximately 80% aboveground and 20% belowground. Forest floor C was 8.5 t ha^-1. Soil C content (0–1 m) was 369 t ha^-1 representing 70% of the total C pool in the ecosystem. Total gross annual C flux aboveground (biomass increment plus litterfall plus respiration) was 11.9 t ha^-1 and gross flux belowground (coarse root increment plus fine root production plus root respiration) was 5.1 t ha^-1. Total annual soil efflux was 7.1 t ha^-1, of which 2.5 t ha^-1 (35%) was contributed by litter decomposition.The short-term effect of changing the availability of P compared with C on allocation to aboveground versus belowground processes was estimated by comparing fertilized and unfertilized stands during the year after treatment. In the P-fertilized stand annual wood biomass increment increased by 30%, there was no evidence of change in canopy biomass, and belowground C allocation decreased by 19% relative to the unfertilized stand. Total annual C flux was 16.97 and 16.75 t ha^-1 yr^-1 and the ratio of below- to aboveground C allocation was 0.43 and 0.35 in the unfertilized and P-fertilized stands, respectively. Therefore, the major response of the forest stand to increased soil P availability appeared to be a shift in C allocation; with little change in total productivity. These results emphasise that both growth rate and allocation need to be estimated to predict changes in fluxes and storage of C in forests that may occur in response to disturbance or climate change.     

Pine Forest Floor Carbon Accumulation in Response to N and PK Additions: Bomb <Superscript>14</Superscript>C Modelling and Respiration Studies

The addition of nitrogen via deposition alters the carbon balance of temperate forest ecosystems by affecting both production and decomposition rates. The effects of 20 years of nitrogen (N) and phosphorus and potassium (PK) additions were studied in a 40-year-old pine stand in northern Sweden. Carbon fluxes of the forest floor were reconstructed using a combination of data on soil ¹⁴C, tree growth, and litter decomposition. N-only additions caused an increase in needle litterfall, whereas both N and PK additions reduced long-term decomposition rates. Soil respiration measurements showed a 40% reduction in soil respiration for treated compared to control plots. The average age of forest floor carbon was 17 years. Predictions of future soil carbon storage indicate an increase of around 100% in the next 100 years for the N plots and 200% for the NPK plots. As much as 70% of the increase in soil carbon was attributed to the decreased decomposition rate, whereas only 20% was attributable to increased litter production. A reduction in decomposition was observed at a rate of N addition of 30 kg C ha^–1 y^–1, which is not an uncommon rate of N deposition in central Europe. A model based on the continuous-quality decomposition theory was applied to interpret decomposer and substrate parameters. The most likely explanations for the decreased decomposition rate were a fertilizer-induced increase in decomposer efficiency (production-to-assimilation ratio), a more rapid rate of decrease in litter quality, and a decrease in decomposer basic growth rate.     

Representative estimates of soil and ecosystem respiration in an old beech forest

Respiration has been proposed to be the main determinant of the carbon balance in European forests and is thus essential for our understanding of the carbon cycle. However, the choice of experimental design strongly affects estimates of annual respiration and of the contribution of soil respiration to total ecosystem respiration. In a detailed study of ecosystem and soil respiration fluxes in an old unmanaged deciduous forest in Central Germany over 3 years (2000–2002), we combined soil chamber and eddy covariance measurements to obtain a comprehensive picture of respiration in this forest. The closed portable chambers offered to investigate spatial variability of soil respiration and its controls while the eddy covariance system offered continuous measurements of ecosystem respiration. Over the year, both fluxes were mainly correlated with temperature. However, when soil moisture sank below 23 vol.% in the upper 6 cm, water limitations also became apparent. The temporal resolution of the eddy covariance system revealed that relatively high respiration rates occurred during budbreak due to increased metabolic activity and after leaf fall because of increased decomposition. Spatial variability in soil respiration rates was large and correlated with fine root biomass (r ² = 0.56) resulting in estimates of annual efflux varying across plots from 730 to 1,258 (mean 898) g C m⁻² year⁻¹. Power function calculations showed that achieving a precision in the soil respiration estimate of 20% of the full population mean at a confidence level of 95%, requires about eight sampling locations. Our results can be used as guidelines to improve the representativeness of soil respiration measurements by nested sampling designs, being applied in long-term and large-scale carbon sequestration projects such as FLUXNET and CarboEurope.     

The soil organic carbon in particle-size separates under different regrowth forest stands of north eastern Costa Rica

Despite the importance of the secondary forest (SF) in tropical areas, few studies have quantified the soil organic carbon (SOC) pool in Costa Rica. Most of the studies conducted to date in this country have focused mainly on changes in the soil C pool following conversion of forests to pastures, which is the predominant land use in the tropics. The aim of this study was to measure SOC concentration and pool in particle-size fractions down to 50 cm depth in four SF stands regenerating from different intensities of prior land use in loamy sand and sandy loam soils of northeast Costa Rica: (i) a gallery forest (GF), (ii) a 15-year-old SF enriched with commercially planted native trees (15SF), (iii) a 25-year-old SF (25SF), and (iv) an abandoned Theobromma cacao plantation >60 years old (60SF). Additional objectives were (1) to determine the relationship of SOC concentration with selected physical and chemical soil properties, and (2) to establish the key determinants of the depth distribution of SOC in order to identify meaningful trends in the SOC pool. The SOC pool was highest under the 60SF (221.4 Mg C ha⁻¹) followed by the 15SF (212.1 Mg C ha⁻¹), the 25SF (195.9 Mg C ha⁻¹) and the lowest in the GF (183.5 Mg C ha⁻¹). The SOC concentration decreased significantly from 59.7 to 94.1 g kg⁻¹ in the 0–10 cm layer down to 31.0 to 45.5 g kg⁻¹ in the 40–50 cm layer in all forest stands. The fine silt + clay fraction contained the highest values of SOC concentration in all forest stands. Soil texture and the age of the SF were identified as the main factors that explained the variability in SOC. The age of SF stand influenced the distribution of size class aggregates and SOC.     

Contribution of aboveground litter,belowground litter,and rhizosphere respiration to total soil CO<Subscript>2</Subscript> efflux in an old growth coniferous forest

In an old growth coniferous forest located in the central Cascade Mountains, Oregon, we added or removed aboveground litter and terminated live root activity by trenching to determine sources of soil respiration. Annual soil efflux from control plots ranged from 727 g C m⁻² year⁻¹ in 2002 to 841 g C m⁻² year⁻¹ in 2003. We used aboveground litter inputs (149.6 g C m⁻² year⁻¹) and differences in soil CO₂ effluxes among treatment plots to calculate contributions to total soil efflux by roots and associated rhizosphere organisms and by heterotrophic decomposition of organic matter derived from aboveground and belowground litter. On average, root and rhizospheric respiration (R_r) contributed 23%, aboveground litter decomposition contributed 19%, and belowground litter decomposition contributed 58% to total soil CO₂ efflux, respectively. These values fall within the range of values reported elsewhere, although our estimate of belowground litter contribution is higher than many published estimates, which we argue is a reflection of the high degree of mycorrhizal association and low nutrient status of this ecosystem. Additionally, we found that measured fluxes from plots with doubled needle litter led to an additional 186 g C m⁻² year⁻¹ beyond that expected based on the amount of additional carbon added; this represents a priming effect of 187%, or a 34% increase in the total carbon flux from the plots. This finding has strong implications for soil C storage, showing that it is inaccurate to assume that increases in net primary productivity will translate simply and directly into additional belowground storage.     

Contemporary carbon stocks of mineral forest soils in the Swiss Alps

Soil organic carbon (SOC) has been identified as the main globalterrestrial carbon reservoir, but considerable uncertainty remains as toregional SOC variability and the distribution of C between vegetationand soil. We used gridded forest soil data (8–km × 8–km)representative of Swiss forests in terms of climate and forest typedistribution to analyse spatial patterns of mineral SOC stocks alonggradients in the European Alps for the year 1993. At stand level, meanSOC stocks of 98 t C ha^–1 (N = 168,coefficient of variation: 70%) were obtained for the entiremineral soil profile, 76 t C ha^–1 (N =137, CV: 50%) in 0–30 cm topsoil, and 62 t Cha^–1 (N = 156, CV: 46%) in0–20 cm topsoil. Extrapolating to national scale, we calculatedcontemporary SOC stocks of 110 Tg C (entire mineral soil, standarderror: 6 Tg C), 87 Tg C (0–30 cm topsoil, standarderror: 3.5 Tg C) and 70 Tg C (0–20 cm topsoil, standarderror: 2.5 Tg C) for mineral soils of accessible Swiss forests(1.1399 Mha). According to our estimate, the 0–20 cm layers ofmineral forest soils in Switzerland store about half of the Csequestered by forest trees (136 Tg C) and more than five times morethan organic horizons (13.2 Tg C).At stand level, regression analyses on the entire data set yielded nostrong climatic or topographic signature for forest SOC stocks in top(0–20 cm) and entire mineral soils across the Alps, despite thewide range of values of site parameters. Similarly, geostatisticalanalyses revealed no clear spatial trends for SOC in Switzerland at thescale of sampling. Using subsets, biotic, abiotic controls andcategorial variables (forest type, region) explained nearly 60%of the SOC variability in topsoil mineral layers (0–20 cm) forbroadleaf stands (N = 56), but only little of thevariability in needleleaf stands (N = 91,R ² = 0.23 for topsoil layers).Considerable uncertainties remain in assessments of SOC stocks, due tounquantified errors in soil density and rock fraction, lack of data onwithin-site SOC variability and missing or poorly quantifiedenvironmental control parameters. Considering further spatial SOCvariability, replicate pointwise soil sampling at 8–km × 8–kmresolution without organic horizons will thus hardly allow to detectchanges in SOC stocks in strongly heterogeneous mountain landscapes.