Components of forest soil CO2 efflux estimated from Δ14C values of soil organic matter

Aims

The partitioning of the total soil CO₂ efflux into its two main components: respiration from roots (and root-associated organisms) and microbial respiration (by means of soil organic matter (SOM) and litter decomposition), is a major need in soil carbon dynamics studies in order to understand if a soil is a net sink or source of carbon.

Methods

The heterotrophic component of the CO₂ efflux was estimated for 11 forest sites as the ratio between the carbon stocks of different SOM pools and previously published (Δ¹⁴C derived) turnover times. The autotrophic component, including root and root-associated respiration, was calculated by subtracting the heterotrophic component from total soil chamber measured CO₂ efflux.

Results

Results suggested that, on average, 50.4 % of total soil CO₂ efflux was derived from the respiration of the living roots, 42.4 % from decomposition of the litter layers and less than 10 % from decomposition of belowground SOM.

Conclusions

The Δ¹⁴C method proved to be an efficient tool by which to partition soil CO₂ efflux and quantify the contribution of the different components of soil respiration. However the average calculated heterotrophic respiration was statistically lower compared with two previous studies dealing with soil CO₂ efflux partitioning (one performed in the same study area; the other a meta-analysis of soil respiration partitioning). These differences were probably due to the heterogeneity of the SOM fraction and to a sub-optimal choice of the litter sampling period.     

Fertilization of boreal forest reduces both autotrophic and heterotrophic soil respiration

The boreal forest is expected to experience the greatest warming of all forest biomes, raising concerns that some of the large quantities of soil carbon in these systems may be added to the atmosphere as CO₂. However, nitrogen deposition or fertilization has the potential to increase boreal forest production and retard the decomposition of soil organic matter, hence increasing both tree stand and soil C storage. The major contributors to soil‐surface CO₂ effluxes are autotrophic and heterotrophic respiration. To evaluate the effect of nutrient additions on the relative contributions from autotrophic and heterotrophic respiration, a large‐scale girdling experiment was performed in a long‐term nutrient optimization experiment in a 40‐year‐old stand of Norway spruce in northern Sweden. Trees on three nonfertilized plots and three fertilized plots were girdled in early summer 2002, and three nonfertilized and three fertilized plots were used as control plots. Each plot was 0.1 ha and contained around 230 trees. Soil‐surface CO₂ fluxes, soil moisture, and soil temperature were monitored in both girdled and nongirdled plots. In late July, the time of the seasonal maximum in soil‐surface CO₂ efflux, the total soil‐CO₂ efflux in nongirdled plots was 40% lower in the fertilized than in the nonfertilized plots, while the efflux in girdled fertilized and nonfertilized plots was 50% and 60% lower, respectively, than in the corresponding nongirdled controls. We attribute these reductions to losses of the autotrophic component of the total soil‐surface CO₂ efflux. The estimates of autotrophic respiration are conservative as root starch reserves were depleted more rapidly in roots of girdled than in nongirdled trees. Thus, heterotrophic activity was overestimated. Calculated on a unit area basis, both the heterotrophic and autotrophic soil respiration was significantly lower in fertilized plots, which is especially noteworthy given that aboveground production was around three times higher in fertilized than in nonfertilized plots.     

Tree-girdling to separate root and heterotrophic respiration in two Eucalyptus stands in Brazil

The release of carbon as CO₂ from belowground processes accounts for about 70% of total ecosystem respiration. Insights about factors controlling soil CO₂ efflux are constrained by the challenge of apportioning sources of CO₂ between autotrophic tree roots (and mycorrhizal fungi) and heterotrophic microorganisms. In some temperate conifer forests, the reduction in soil CO₂ efflux after girdling (phloem removal) has been used to separate these sources. Girdling stops the flow of carbohydrates to the belowground portion of the ecosystem, which should slow respiration by roots and mycorrhizae while heterotrophic respiration should remain constant or be enhanced by the decomposition of newly dead roots. Therefore, the reduction in CO₂ efflux after girdling should be a conservative estimate of the belowground flux of C from trees. We tested this approach in two tropical Eucalyptus plantations. Tree canopies remained intact for more than 3 months after girdling, showing no reduction in light interception. The reduction in soil CO₂ efflux averaged 16–24% for the 3-month period after girdling. The reduction in CO₂ efflux was similar for plots with one half of the trees girdled and those with all of the trees girdled. Girdling did not reduce live fine root biomass for at least 5 months after treatment, indicating that large reserves of carbohydrates in the root systems of Eucalyptus trees maintained the roots and root respiration. Our results suggest that the girdling approach is unlikely to provide useful insights into the contribution of tree roots and heterotrophs to soil CO₂ efflux in this type of forest ecosystem.     

Simulating soil N2O emissions and heterotrophic CO2 respiration in arable systems using FASSET and MoBiLE-DNDC

Modelling of soil emissions of nitrous oxide (N₂O) and carbon dioxide (CO₂) is complicated by complex interactions between processes and factors influencing their production, consumption and transport. In this study N₂O emissions and heterotrophic CO₂ respiration were simulated from soils under winter wheat grown in three different organic and one inorganic fertilizer-based cropping system using two different models, i.e., MoBiLE-DNDC and FASSET. The two models were generally capable of simulating most seasonal trends of measured soil heterotrophic CO₂ respiration and N₂O emissions. Annual soil heterotrophic CO₂ respiration was underestimated by both models in all systems (about 10?C30% by FASSET and 10?C40% by MoBiLE-DNDC). Both models overestimated annual N₂O emissions in all systems (about 10?C580% by FASSET and 20?C50% by MoBiLE-DNDC). In addition, both models had some problems in simulating soil mineral nitrogen, which seemed to originate from deficiencies in simulating degradation of soil organic matter, incorporated residues of catch crops and organic fertilizers. To improve the performance of the models, organic matter decomposition parameters need to be revised.     

Environmental Effects over the First 2½ Rotation Periods of a Fertilised Poplar Short Rotation Coppice

A short rotation coppice (SRC) with poplar was established in a randomised fertilisation experiment on sandy loam soil in Potsdam (Northeast Germany). The main objective of this study was to assess if negative environmental effects as nitrogen leaching and greenhouse gas emissions are enhanced by mineral nitrogen (N) fertiliser applied to poplar at rates of 0, 50 and 75 kg N ha^?1 year^?1 and how these effects are influenced by tree age with increasing number of rotation periods and cycles of organic matter decomposition and tree growth after each harvesting event. Between 2008 and 2012, the leaching of nitrate (NO₃ ^?) was monitored with self-integrating accumulators over 6-month periods and the emissions of the greenhouse gases (GHG) nitrous oxide (N₂O) and carbon dioxide (CO₂) were determined in closed gas chambers. During the first 4 years of the poplar SRC, most nitrogen was lost through NO₃ ^? leaching from the main root zone; however, there was no significant relationship to the rate of N fertilisation. On average, 5.8 kg N ha^?1 year^?1 (13.0 kg CO_2equ) was leached from the root zone. Nitrogen leaching rates decreased in the course of the 4-year study parallel to an increase of the fine root biomass and the degree of mycorrhization. In contrast to N leaching, the loss of nitrogen by N₂O emissions from the soil was very low with an average of 0.61 kg N ha^?1 year^?1 (182 kg CO_2equ) and were also not affected by N fertilisation over the whole study period. Real CO₂ emissions from the poplar soil were two orders of magnitude higher ranging between 15,122 and 19,091 kg CO₂ ha^?1 year^?1 and followed the rotation period with enhanced emission rates in the years of harvest. As key-factors for NO₃ ^? leaching and N₂O emissions, the time after planting and after harvest and the rotation period have been identified by a mixed effects model.     

Estimation of autotrophic and heterotrophic components of soil respiration by trenching is sensitive to corrections for root decomposition and changes in soil water content

This study aims to assess the effects of corrections for disturbances such as an increased amount of dead roots and an increase in volumetric soil water content on the calculation of soil CO₂ efflux partitioning. Soil CO₂ efflux, soil temperature and superficial soil water content were monitored in two young beech sites (H1 and H2) during a trenching experiment. Trenching induced a significant input of dead root mass that participated in soil CO₂ efflux and reduced the soil dissolved organic carbon content, while it increased superficial soil water content within the trenched plot. Annual soil CO₂ efflux in control plots was 528 g C m⁻² year⁻¹ at H1 and 527 g C m⁻² year⁻¹ at H2. The annual soil CO₂ efflux in trenched plots was 353 g C m⁻² year⁻¹ at H1 and 425 g C m⁻² year⁻¹ at H2. By taking into account annual CO₂ efflux from decaying trenched roots, the autotrophic contribution to total soil CO₂ efflux reached 69% at H1 and 54% at H2. The partitioning calculation was highly sensitive to the initial root mass estimated within the trenched plots. Uncertainties in the remaining root mass, the fraction of root C that is incorporated into soil organic matter during root decomposition, and the root decomposition rate constant had a limited impact on the partitioning calculation. Corrections for differences in superficial soil water content had a significant impact on annual respired CO₂ despite a limited effect on partitioning.     

Root exclusion through trenching does not affect the isotopic composition of soil CO2 efflux

Disentangling the autotrophic and heterotrophic components of soil CO₂ efflux is critical to understanding the role of soil system in terrestrial carbon (C) cycling. In this study, we combined a stable C-isotope natural abundance approach with the trenched plot method to determine if root exclusion significantly affected the isotopic composition (δ¹³C) of soil CO₂ efflux (R_S). This study was performed in different forest ecosystems: a tropical rainforest and two temperate broadleaved forests, where trenched plots had previously been installed. At each site, R_S and its δ¹³C (δ¹³C_Rs) tended to be lower in trenched plots than in control plots. Contrary to R_S, δ¹³C_Rs differences were not significant. This observation is consistent with the small differences in δ¹³C measured on organic matter from root, litter and soil. The lack of an effect on δ¹³C_Rs by root exclusion could be from the small difference in δ¹³C between autotrophic and heterotrophic soil respirations, but further investigations are needed because of potential artefacts associated with the root exclusion technique.     

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.     

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.     

Tropical tree species composition affects the oxidation of dissolved organic matter from litter

Plant species effects on soil nutrient availability are relatively well documented, but the effects of species differences in litter chemistry on soil carbon cycling are less well understood, especially in the species-rich tropics. In many wet tropical forest ecosystems, leaching of dissolved organic matter (DOM) from the litter layer accounts for a significant proportion of litter mass loss during decomposition. Here we investigated how tree species differences in soluble dissolved organic C (DOC) and nutrients affected soil CO₂ fluxes in laboratory incubations. We leached DOM from freshly fallen litter of six canopy tree species collected from a tropical rain forest in Costa Rica and measured C-mineralization. We found significant differences in litter solubility and nutrient availability. Following DOM additions to soil, rates of heterotrophic respiration varied by as much as an order of magnitude between species, and overall differences in total soil CO₂ efflux varied by more than four-fold. Variation in the carbon: phosphorus ratio accounted for 51% of the variation in total CO₂ flux between species. These results suggest that tropical tree species composition may influence soil C storage and mineralization via inter-specific variation in plant litter chemistry.     

Elevated CO2 and temperature impacts on different components of soil CO2 efflux in Douglas-fir terracosms

Although numerous studies indicate that increasing atmospheric CO₂ or temperature stimulate soil CO₂ efflux, few data are available on the responses of three major components of soil respiration [i.e. rhizosphere respiration (root and root exudates), litter decomposition, and oxidation of soil organic matter] to different CO₂ and temperature conditions. In this study, we applied a dual stable isotope approach to investigate the impact of elevated CO₂ and elevated temperature on these components of soil CO₂ efflux in Douglas-fir terracosms. We measured both soil CO₂ efflux rates and the ¹³C and ¹⁸O isotopic compositions of soil CO₂ efflux in 12 sun-lit and environmentally controlled terracosms with 4-year-old Douglas fir seedlings and reconstructed forest soils under two CO₂ concentrations (ambient and 200 ppmv above ambient) and two air temperature regimes (ambient and 4 °C above ambient). The stable isotope data were used to estimate the relative contributions of different components to the overall soil CO₂ efflux. In most cases, litter decomposition was the dominant component of soil CO₂ efflux in this system, followed by rhizosphere respiration and soil organic matter oxidation. Both elevated atmospheric CO₂ concentration and elevated temperature stimulated rhizosphere respiration and litter decomposition. The oxidation of soil organic matter was stimulated only by increasing temperature. Release of newly fixed carbon as root respiration was the most responsive to elevated CO₂, while soil organic matter decomposition was most responsive to increasing temperature. Although some assumptions associated with this new method need to be further validated, application of this dual-isotope approach can provide new insights into the responses of soil carbon dynamics in forest ecosystems to future climate changes.     

Model for rhizodeposition and CO2 efflux from planted soil and its validation by 14C pulse labelling of ryegrass

A model for rhizodeposition and root respiration was developed and parameterised based on ¹⁴C pulse labelling of Lolium perenne. The plants were grown in a two-compartment chamber on a loamy Haplic Luvisol under controlled laboratory conditions. The dynamics of ¹⁴CO₂ efflux from the soil and ¹⁴C content in shoots, roots, micro-organisms, dissolved organic carbon (DOC) and soil were measured during the first 11 days after labelling. Modelled parameters were estimated by fitting on measured ¹⁴C dynamics in the different pools. The model and the measured ¹⁴C dynamics in all pools corresponded well (r ²=0.977). The model describes well ¹⁴CO₂ efflux from the soil and ¹⁴C dynamics in shoots, roots and soil, but predicts unsatisfactorily the ¹⁴C content in micro-organisms and DOC. The model also allows for division of the total ¹⁴CO₂ efflux from the soil in ¹⁴CO₂ derived from root respiration and ¹⁴CO₂ derived from rhizomicrobial respiration by use of exudates and root residues. Root respiration and rhizomicrobial respiration amounted for 7.6% and 6.0% of total assimilated C, respectively, which accounts for 56% and 44% of root-derived ¹⁴CO₂ efflux from the soil planted with 43-day-old Lolium perenne, respectively. The sensitivity analysis has shown that root respiration rate affected the curve of ¹⁴CO₂ efflux from the soil mainly during the first day after labelling. The changes in the exudation rate influenced the ¹⁴CO₂ efflux later than first 24 h after labelling.     

Partitioning sources of soil respiration in boreal black spruce forest using radiocarbon

Separating ecosystem and soil respiration into autotrophic and heterotrophic component sources is necessary for understanding how the net ecosystem exchange of carbon (C) will respond to current and future changes in climate and vegetation. Here, we use an isotope mass balance method based on radiocarbon to partition respiration sources in three mature black spruce forest stands in Alaska. Radiocarbon (Δ¹⁴C) signatures of respired C reflect the age of substrate C and can be used to differentiate source pools within ecosystems. Recently‐fixed C that fuels plant or microbial metabolism has Δ¹⁴C values close to that of current atmospheric CO₂, while C respired from litter and soil organic matter decomposition will reflect the longer residence time of C in plant and soil C pools. Contrary to our expectations, the Δ¹⁴C of C respired by recently excised black spruce roots averaged 14‰ greater than expected for recently fixed photosynthetic products, indicating that some portion of the C fueling root metabolism was derived from C storage pools with turnover times of at least several years. The Δ¹⁴C values of C respired by heterotrophs in laboratory incubations of soil organic matter averaged 60‰ higher than the contemporary atmosphere Δ¹⁴CO₂, indicating that the major contributors to decomposition are derived from a combination of sources consistent with a mean residence time of up to a decade. Comparing autotrophic and heterotrophic Δ¹⁴C end members with measurements of the Δ¹⁴C of total soil respiration, we calculated that 47–63% of soil CO₂ emissions were derived from heterotrophic respiration across all three sites. Our limited temporal sampling also observed no significant differences in the partitioning of soil respiration in the early season compared with the late season. Future work is needed to address the reasons for high Δ¹⁴C values in root respiration and issues of whether this method fully captures the contribution of rhizosphere respiration.     

The influence of temperature and labile C substrates on heterotrophic respiration in response to elevated CO2 and nitrogen fertilization

Important effects of elevated [CO₂] on SOM are expected as a consequence of increased labile organic substrates derived from plants. The present study tests the hypotheses that, under elevated [CO₂]: 1) soil heterotrophic respiration will increase due to roots-microbes-soil interactions; 2) the increased labile C will boost soil heterotrophic respiration, depending on N availability; 3) the temperature sensitivity of soil respiration will change, depending on nitrogen inputs and plant activity. To test these hypotheses, we measured the heterotrophic respiration of intact soil cores collected in a poplar plantation exposed to elevated [CO₂] and two nitrogen inputs, at different temperatures. Additional physical (water content, root biomass) and biochemical parameters (microbial biomass, labile C) were determined on the same samples. The soil samples were collected at the POP-EuroFACE experimental site (Italy), where a Populus x euramericana plantation was exposed for 6 years to 550 ppm [CO₂] (Free Air CO₂ Enrichment) at two different nitrogen inputs (none or 290 kg ha^?1). The higher heterotrophic respiration under elevated [CO₂] (+30% on average) was driven by the larger pool of soil labile C (+57% on average). The temperature sensitivity of soil respiration was unaffected by elevated [CO₂], but was positively affected by N fertilization. Our results indicate that only a fraction of the extra carbon fixed by photosynthesis in elevated [CO₂] will contribute to enhanced carbon storage into the soil because of the contemporary stimulation of soil heterotrophic respiration. At the same time, the fraction remaining in the soil will enhance the pool of soil labile C.     

Impact of grazing intensity on seasonal variations in soil organic carbon and soil CO2 efflux in two semiarid grasslands in southern Botswana

Biological soil crusts (BSCs) are an important source of organic carbon, and affect a range of ecosystem functions in arid and semiarid environments. Yet the impact of grazing disturbance on crust properties and soil CO₂ efflux remain poorly studied, particularly in African ecosystems. The effects of burial under wind-blown sand, disaggregation and removal of BSCs on seasonal variations in soil CO₂ efflux, soil organic carbon, chlorophyll a and scytonemin were investigated at two sites in the Kalahari of southern Botswana. Field experiments were employed to isolate CO₂ efflux originating from BSCs in order to estimate the C exchange within the crust. Organic carbon was not evenly distributed through the soil profile but concentrated in the BSC. Soil CO₂ efflux was higher in Kalahari Sand than in calcrete soils, but rates varied significantly with seasonal changes in moisture and temperature. BSCs at both sites were a small net sink of C to the soil. Soil CO₂ efflux was significantly higher in sand soils where the BSC was removed, and on calcrete where the BSC was buried under sand. The BSC removal and burial under sand also significantly reduced chlorophyll a, organic carbon and scytonemin. Disaggregation of the soil crust, however, led to increases in chlorophyll a and organic carbon. The data confirm the importance of BSCs for C cycling in drylands and indicate intensive grazing, which destroys BSCs through trampling and burial, will adversely affect C sequestration and storage. Managed grazing, where soil surfaces are only lightly disturbed, would help maintain a positive carbon balance in African drylands.     

13C isotope fractionation during rhizosphere respiration of C3 and C4 plants

Stable carbon isotopes are used extensively to partition total soil CO₂ efflux into root-derived rhizosphere respiration or autotrophic respiration and soil-derived heterotrophic respiration. However, it remains unclear whether CO₂ from rhizosphere respiration has the same δ¹³C value as root biomass. Here we investigated the magnitude of ¹³C isotope fractionation during rhizosphere respiration relative to root biomass in six plant species. Plants were grown in a carbon-free sand-perlite medium inoculated with microorganisms from a farm soil for 62 days inside a greenhouse. We measured the δ¹³C value of rhizosphere respiration using a closed-circulation 48-hour CO₂ trapping method during 40~42 and 60~62 days after sowing. We found a consistent depletion in ¹³C (0.9~1.7‰) of CO₂ from rhizosphere respiration relative to root biomass in three C₃ species (Glycine max L. Merr., Helianthus annuus L. and Triticum aestivum L.), but a relatively large depletion in ¹³C (3.7~7.0‰) in three C₄ species (Amaranthus tricolor L., Sorghum bicolor (L.) Moench and Zea mays L. ssp. mays). Overall, our results indicate that CO₂ from rhizosphere respiration is more ¹³C-depleted than root biomass. Therefore, accounting for this ¹³C fractionation is required for accurately partitioning total soil CO₂ efflux into root-derived and soil-derived components using natural abundance stable carbon isotope methods.     

Radiocarbon based assessment of soil organic matter contribution to soil respiration in a pine stand of the Campine region, Belgium

The contribution of decomposing soil organic carbon (SOC) to total annual soil respiration (SR) was evaluated by radiocarbon measurements at a Scots pine stand growing on a plaggen soil in the Belgian Campine region. Two approaches were used to estimate the contribution of different C pools to SR. In the first approach, the variations in ¹⁴C content of soil CO₂ efflux were monitored during one year (2003) and compared to the atmospheric and SOC ¹⁴C signatures to determine the contribution of ??fast?? (root respiration and fast decomposing SOC) and ??slow?? cycling C pools to total SR. In the second approach an estimate of the total heterotrophic soil respiration (Rh), comprising the slow cycling C and the heterotrophic part of the fast-cycling C pools, was derived applying a box model based on the amount of the bulk SOC pool and its ¹⁴C-derived mean residence time (MRT). The quantification of the Rh and the decomposition rate of the slow-cycling SOC allows to indirectly determining the contribution of the heterotrophic C that decompose within a year. Measurements of total SR performed in the field allowed assessing the contribution of the different C pools to total soil C efflux. On an annual basis, the fast-cycling C was the main contributor to SR, about 85%, while the contribution of the slow-cycling C (with MRT >1 yr) to total SR was 15%. Total annual Rh was 36% of total SR, which is in the lower range reported for temperate coniferous forests. The comparison of Rh with other estimates for the same site (47?C50% of total SR) suggest a possible underestimation of the C flux from the mineral soil. In fact, the ??very old?? C contained in the plaggen horizon strongly affects the signature of the mostly young C leaving the soil. In conclusion, our results indicate that the contribution of SOC decomposition to total soil CO₂ flux in this forest is less than 40%, and at least half of it comes from organic compounds less than 1 year old.     

Indirect partitioning of soil respiration in a series of evergreen forest ecosystems

A simple estimation of heterotrophic respiration can be obtained analytically as the y-intercept of the linear regression between soil-surface CO₂ efflux and root biomass. In the present study, a development of this indirect methodology is presented by taking into consideration both the temporal variation and the spatial heterogeneity of heterotrophic respiration. For this purpose, soil CO₂ efflux, soil carbon content and main stand characteristics were estimated in seven evergreen forest ecosystems along an elevation gradient ranging from 250 to 1740 m. For each site and for each sampling date the measured soil CO₂ efflux (R _S) was predicted with the model R _S = a × S _C + b × R _D ± ε, where S _C is soil carbon content per unit area to a depth of 30 cm and R _D is the root density of the 2–5 mm root class. Regressions with statistically significant a and b coefficients allowed the indirect separation of the two components of soil CO₂ efflux. Considering that the different sampling dates were characterized by different soil temperature, it was possible to investigate the temporal and thermal dependency of autotrophic and heterotrophic respiration. It was estimated that annual autotrophic respiration accounts for 16–58% of total soil CO₂ efflux in the seven different evergreen ecosystems. In addition, our observations show a decrease of annual autotrophic respiration at increasing availability of soil nitrogen. Section Editor: A. Hodge     

Tree root and soil heterotrophic respiration as revealed by girdling of boreal Scots pine forest: extending observations beyond the first year

Limitations in available techniques to separate autotrophic (root) and soil heterotrophic respiration have hampered the understanding of forest C cycling. The former is here defined as respiration by roots, their associated mycorrhizal fungi and other micro‐organisms in the rhizosphere directly dependent on labile C compounds leaked from roots. In order to separate the autotrophic and heterotrophic components of soil respiration, all Scots pine trees in 900 m² plots were girdled to instantaneously terminate the supply of current photosynthates from the tree canopy to roots. Högberg et al. (Nature 411, 789–792, 2001) reported that autotrophic activity contributed up to 56% of total soil respiration during the first summer of this experiment. They also found that mobilization of stored starch (and likely also sugars) in roots after girdling caused an increased apparent heterotrophic respiration on girdled plots. Herein a transient increase in the δ¹³C of soil CO₂ efflux after girdling, thought to be due to decomposition of ¹³C‐enriched ectomycorrhizal mycelium and root starch and sugar reserves, is reported. In the second year after girdling, when starch reserves of girdled tree roots were exhausted, calculated root respiration increased up to 65% of total soil CO₂ efflux. It is suggested that this estimate of its contribution to soil respiration is more precise than the previous based on one year of observation. Heterotrophic respiration declined in response to a 20‐day‐long 6 °C decline in soil temperature during the second summer, whereas root respiration did not decline. This did not support the idea that root respiration should be more sensitive to variations in soil temperature. It is suggested that above‐ground photosynthetic activity and allocation patterns of recent photosynthates to roots should be considered in models of responses of forest C balances to global climate change.     

Carbon sink reduction by fruit removal triggers respiration but not nitrous oxide emissions from the root zone of cucumber

Root exudation of organic carbon (C) is generally believed to be the cause of positive effects of root activity on nitrous oxide (N₂O) emissions. We tested the effects of root exudation in an actual soil–plant system on N₂O emissions while excluding most other potential factors. The C source/sink ratio in cucumber was changed by removing fruits to increase root exudation. Root‐zone emissions of carbon dioxide (CO₂) and N₂O were monitored in complete stands of adult plants in a greenhouse. Whereas CO₂ emissions rapidly increased as a result of fruit removal the N₂O emissions were completely unaffected. After cutting the shoots CO₂ emissions decreased within 2 weeks in both the fruit removal treatment and the control to a value significantly lower than that before the start of the treatments. However, N₂O emissions immediately exhibited a short peak, which was significantly higher in the fruit removal treatment compared to the control. Thereafter N₂O emissions in both treatments remained on the same level but considerably higher than before shoot cutting. We concluded that in a well‐aerated root zone, a root exudation pulse does not necessarily increase N₂O emissions, because C substrates are quickly respired by microorganisms before they can support heterotrophic denitrification. The results further indicate the significance of dying/dead roots for the creation of denitrificaton hot‐spots, which likely result from providing C substrates as well as poorly aerated habitats.