Carbon fluxes in soil food webs of increasing complexity revealed by C labelling and C natural abundance

Soil food webs are mainly based on three primary carbon (C) sources: root exudates, litter, and recalcitrant soil organic matter (SOM). These C sources vary in their availability and accessibility to soil organisms, which could lead to different pathways in soil food webs. The presence of three C isotopes (¹²C, ¹³C and ¹⁴C) offers an unique opportunity to investigate all three C sources simultaneously. In a microcosm experiment we studied the effect of food web complexity on the utilization of the three carbon sources. We choose an incomplete three factorial design with (i) living plants, (ii) litter and (iii) food web complexity. The most complex food web consisted of autochthonous microorganisms, nematodes, collembola, predatory mites, endogeic and anecic earthworms. We traced C from all three sources in soil, in CO₂ efflux and in individual organism groups by using maize grown on soil developed under C₃ vegetation and application of ¹⁴C labelled ryegrass shoots as a litter layer. The presence of living plants had a much greater effect on C pathways than food web complexity. Litter decomposition, measured as ¹⁴CO₂ efflux, was decreased in the presence of living plants from 71% to 33%. However, living plants increased the incorporation of litter C into microbial biomass and arrested carbon in the litter layer and in the upper soil layer. The only significant effect of food web complexity was on the litter C distribution in the soil layers. In treatments with fungivorous microarthropods (Collembola) the incorporation of litter carbon into mineral soil was reduced. Root exudates as C source were passed through rhizosphere microorganisms to the predator level (at least to the third trophic level). We conclude that living plants strongly affected C flows, directly by being a source of additional C, and indirectly by modifying the existing C flows within the food web including CO₂ efflux from the soil and litter decomposition.     

The Impact of Diesel Oil Pollution on the Hydrophobicity and CO<Subscript>2</Subscript> Efflux of Forest Soils

The contamination of soil with petroleum products is a major environmental problem. Petroleum products are common soil contaminants as a result of human activities, and they are causing substantial changes in the biological (particularly microbiological) processes, chemical composition, structure and physical properties of soil. The main objective of this study was to assess the impact of soil moisture on CO₂ efflux from diesel-contaminated albic podzol soils. Two contamination treatments (3000 and 9000 mg of diesel oil per kg of soil) were prepared for four horizons from two forest study sites with different initial levels of soil water repellency. CO₂ emissions were measured using a portable infrared gas analyser (LCpro+, ADC BioScientific, UK) while the soil samples were drying under laboratory conditions (from saturation to air-dry). The assessment of soil water repellency was performed using the water drop penetration time test. An analysis of variance (ANVOA) was conducted for the CO₂ efflux data. The obtained results show that CO₂ efflux from diesel-contaminated soils is higher than efflux from uncontaminated soils. The initially water-repellent soils were found to have a bigger CO₂ efflux. The non-linear relationship between soil moisture content and CO₂ efflux only existed for the upper soil horizons, while for deeper soil horizons, the efflux is practically independent of soil moisture content. The contamination of soil by diesel leads to increased soil water repellency.     

Carbon dioxide emissions of soils under pure and mixed stands of beech and spruce, affected by decomposing foliage litter mixtures

Soil respiration is the largest terrestrial source of CO₂ to the atmosphere. In forests, roughly half of the soil respiration is autotrophic (mainly root respiration) while the remainder is heterotrophic, originating from decomposition of soil organic matter. Decomposition is an important process for cycling of nutrients in forest ecosystems. Hence, tree species induced changes may have a great impact on atmospheric CO₂ concentrations. Since studies on the combined effects of beech-spruce mixtures are very rare, we firstly measured CO₂ emission rates in three adjacent stands of pure spruce (Picea abies), mixed spruce-beech and pure beech (Fagus sylvatica) on three base-rich sites (Flysch) and three base-poor sites (Molasse; yielding a total of 18 stands) during two summer periods using the closed chamber method. CO₂ emissions were higher on the well-aerated sandy soils on Molasse than on the clayey soils on Flysch, characterized by frequent water logging. Mean CO₂ effluxes increased from spruce (41) over the mixed (55) to the beech (59) stands on Molasse, while tree species effects were lower on Flysch (30-35, mixed > beech = spruce; all data in mg CO₂-C m⁻² h⁻¹). Secondly, we studied decomposition after fourfold litter manipulations at the 6 mixed species stands: the O_i - and O_e horizons were removed and replaced by additions of beech -, spruce - and mixed litter of the adjacent pure stands of known chemical quality and one zero addition (blank) in open rings (20 cm inner diameter), which were covered with meshes to exclude fresh litter fall. Mass loss within two years amounted to 61-68% on Flysch and 36-44% on Molasse, indicating non-additive mixed species effects (mixed litter showed highest mass loss). However, base cation release showed a linear response, increasing from the spruce - over the mixed - to the beech litter. The differences in N release (immobilization) resulted in a characteristic converging trend in C/N ratios for all litter compositions on both bedrocks during decomposition. In the summers 2006 and 2007 we measured CO₂ efflux from these manipulated areas (a closed chamber fits exactly over such a ring) as field indicator of the microbial activity. Net fluxes (subtracting the so-called blank values) are considered an indicator of litter induced changes only and increased on both bedrocks from the spruce - over the mixed - to the beech litter. According to these measurements, decomposing litter contributed between 22-32% (Flysch) and 11-28% (Molasse) to total soil respiration, strengthening its role within the global carbon cycle.     

Application of wood ash accelerates soil respiration and tree growth on drained peatland

Forested peatlands contain large pools of terrestrial carbon. As well as drainage, forest management such as fertilizer application can affect these pools. We studied the effect of wood ash (application rates 0, 5 and 15 t ha^?1) on the heterotrophic soil respiration (CO₂ efflux), cellulose decomposition, soil nutrients, biomass production and amount of C accumulated in a tree stand on a pine‐dominated drained mire in central Finland. The ash was spread 13 years before the respiration measurements. The annual CO₂ efflux was statistically modelled using soil temperature as the driving variable. Wood ash application increased the amounts of mineral nutrients of peat substantially and increased soil pH in the uppermost 10 cm layer by 1.5–2 pH units. In the surface peat, the decomposition rate of cellulose in the ash plots was roughly double that in control plots. Annual CO₂ efflux was least on the unfertilized site, 238 g CO₂‐C m^?2 year^?1. The use of wood ash nearly doubled CO₂ efflux to 420–475 g CO₂‐Cm^?2 year^?1 on plots fertilized with 5–15 t ha^?1 of ash, respectively. Furthermore, ash treatments resulted also in increased stand growth, and during the measurement year, the growing stand on ash plots accumulated carbon 11–12 times faster than the control plot. The difference between peat C emission and amount of C sequestered by trees on the ash plots was 43–58 g C m^?2, while on the control plot it was 204 g C m^?2. Our conclusion is that adding wood ash as a fertilizer increases more C sequestration in the tree stand than C efflux from the peat.     

Warming and elevated CO2 combine to increase microbial mineralisation of soil organic matter

The net annual exchange of carbon between the atmosphere and terrestrial ecosystems is of prime importance in determining the concentration of CO₂ ([CO₂]) in the atmosphere and consequently future climate. Carbon loss occurs primarily through soil respiration; it is known that respiration is sensitive to the global changes in [CO₂] and temperature, suggesting that the net carbon balance may change in the future. However, field manipulations of temperature and [CO₂] alter many important environmental factors so it is unclear how much of the observed alterations in soil respiration is due to changes of microbial function itself instead of changes to the physical and chemical environment. Here we focus on resolving the importance of changes in the microbial community in response to warming and elevated [CO₂] on carbon mineralisation, something not possible in field measurements. We took plant material and soil inocula from a long running experiment where native grassland had been exposed to both warming and elevated CO₂ and constructed a reciprocal transplant experiment. We found that the rate of decomposition (heterotrophic respiration) was strongly determined by the origin of the microbial community. The combined warming + elevated CO₂ treatment produced a soil community that gave respiration rates 30% higher when provided with shoot litter and 70% for root litter than elevated CO₂ treatment alone, with the treatment source of the litter being unimportant. Warming, especially in the presence of elevated CO₂, increased the size of the apparent labile carbon pool when either C₃ or C₄ litter was added. Thus, the metabolic activity of the soil community was affected by the combination of warming and elevated CO₂ such that it had an increased ability to mineralise added organic matter, regardless of its source. Therefore, soil C efflux may be substantially increased in a warmer, high CO₂ world. Current ecosystem models mostly drive heterotrophic respiration from plant litter quality, soil moisture and temperature but our findings suggest equal attention will need to be paid to capturing microbial processes if we are to accurately project the future C balance of terrestrial ecosystems and quantify the feedback effect on atmospheric concentrations of CO₂.     

Forest Litters as a Link in the Carbon Cycle in Coniferous–Broadleaved Forests of the Southern Far East of Russia

Forest litter as a component of the carbon cycle in pine–broadleaved forests of different ages was characterized. Field studies of the forest site of the Primorskaya State Agricultural Academy in the south of the Sikhote Alin Range continued for three years. Multiple sampling of forest litter and plant litter fall and measurements of the CO₂ emission from the litter and underlying soil horizons were performed on test plots. The maximum litter pool (14.44 ± 0.86 t/ha) was found in the mature stand of Pinus koraiensis; the minimum litter pool (11.52 ± 0.65 t/ha), in the 80-year-old stand. The carbon stock in the litters amounted to 3.7% of the phytomass carbon. The rate of carbon turnover in the forest litters was relatively low in comparison with that in other regions: the ratio of carbon pools in the litter horizons and annual plant falloff reached 3.5. The winter season provided about 10–25% of the annual plant litter fall. The data obtained in this study describe a part of the carbon cycle and contribute to our understanding of the ecosystem function of climate regulation by valuable forest massifs in the south of the Far East of Russia. In this region, the decomposition of forest litters generates the CO₂ flux amounting up to 16% of the total CO₂ emission from the soil.     

Influence of temperature on CO2 evolution,microbial biomass C and metabolic quotient during the decomposition of two humic forest horizons

The decomposition of the litter layer and the humic mineral horizon from a beech forest site was studied at temperatures of 5, 12, and 22°C for both substrates and additionally at 32°C for beech litter. Weight losses, basal and substrate-induced CO₂ production, and the extractable biomass C were monitored periodically during a 2-year incubation period. Weight losses and microbial activity were controlled by substrate quality and temperature. No significant differences were found between 5 and 12°C in decomposition, biomass C, and the metabolic quotient in the humic mineral horizon. The decay of beech litter and the humic mineral horizon was highest at 22°C but was faster in the litter material by a factor of 2.9 on average. In the glucose-amended samples, the relationship among the CO₂-C fluxes was 1:1:2:3 at temperatures of 5, 12, 22, and 32°C in the litter layer, and 1: 2: 2.4 at 5, 12, and 22°C in the A horizon, respectively. The microbial activity in the humic mineral horizon was only 2–11% of that in the litter layer. The level of biomass C remained constant over 1 year and no significant differences were obtained from the 12 and 22°C treatments in the litter layer.     

Adsorption complex,pH relationships in some acid mediterranean forest soil profiles

Acid soils in some mediterranean forests were investigated for the composition of the adsorption complex and the gradients in soil pH. The effective CEC (235–838 mmol_c kg^?1) and base saturation (93–98 %) are highest in ectorganic horizons. In the mineral horizons the effective CEC (23–52 mmol_c kg^?1) and base saturation (11–40 %) are much lower. The exchange complex of mineral horizons consists for 90 (AEh) to 40 percent (Bw2) of organic matter. The effective CEC of the mineral clay fraction is low (60 mmol_c kg^?1 clay). The clear trends in soil pH within the ectorganic layer of deciduous and sclerophyllous oak forests are attributed to vertical spatial separation of nitrogen mineralization (ammonification and strongly impeded nitrification) and nutrient uptake by roots (mainly NH₄). This leads to a high effective CEC in the fermentation layer and acidification of the uppermost part of the mineral soil. In contrast to the situation in temperate forests this process is impeded in mediterranean coniferous forests, where litter decomposition is extremely slow and both proton production and consumption take place in the organic rich mineral horizon.     

Factors controlling soil CO2 effluxes and the effects of rewetting on effluxes in adjacent deciduous, coniferous, and mixed forests in Korea

To better understand the factors that control forest soil CO₂ efflux and the effects of rewetting on efflux, we measured soil CO₂ efflux in adjacent deciduous, coniferous, and mixed forests in the central part of the Korean Peninsula over the course of one year. We also conducted laboratory rewetting experiments with soil collected from the three sites using three different incubation temperatures (4 °C, 10 °C, and 20 °C). Soil moisture (SM), soil organic matter (SOM), and total root mass values of the three sites were significantly different from one another; however, soil temperature (ST), observed soil CO₂ efflux and sensitivity of soil CO₂ efflux to ST (i.e., Q₁₀ = 3.7 ± 0.1) were not significantly different among the three sites. Soil temperature was a dominant control factor regulating soil CO₂ efflux during most of the year. We infer that soil CO₂ efflux was not significantly different among the sites due to similar ST and Q₁₀. Though a significant increase in soil CO₂ efflux following rewetting of dry soil was observed both in the field observations (60-170%) and laboratory incubation experiments (100-1000%), both the increased rates of soil CO₂ efflux and the magnitude of change in SM were not significantly different among the sites. The increased rates of soil CO₂ efflux following rewetting depended on the initial SM before rewetting. During drying phase after rewetting, a significant correlation between SM and soil CO₂ efflux was found, but the effect of ST on increased soil CO₂ efflux was not clear. Cumulative peak soil CO₂ efflux (11.3 ± 0.7 g CO₂ m⁻²) following rewetting in the field was not significantly different among the sites. Those evidences indicate that the observed similar rewetting effects on soil CO₂ efflux can be explained by the similar magnitude of change in SM after rewetting at the sites. We conclude that regardless of vegetation type, soil CO₂ efflux and the effect of rewetting on soil CO₂ efflux do not differ among the sites, and ST is a primary control factor for soil CO₂ efflux while SM modulates the effect of rewetting on soil CO₂ efflux. Further studies are needed to quantify and incorporate relationship of initial dryness of the soil and the frequency of the dry-wet cycle on soil CO₂ efflux into models describing carbon (C) processes in forested ecosystems.     

Clear‐cutting and slash burning effects on soil CO2 efflux partitioning in Chinese fir and evergreen broadleaved forests in subtropical China

Clear‐cutting (CC) and slash burning (SB) are common silvicultural practices in subtropical China, yet the time‐course response of soil CO₂ efflux components to such disturbance is not well understood. This study examined the effects of CC and SB on soil CO₂ efflux components in a Cunninghamia lanceolata (Lamb.) Hook (Chinese fir, CF) plantation and a secondary evergreen broadleaved forest (BF) located in Fujian Province, southeastern China. Aboveground litter removal and root trenching were used to estimate CO₂ fluxes from soil organic matter decomposition (R_SOM), litter decomposition (R_L), and autotrophic respiration by roots and mycorrhizae (R_R). These components were measured 5–7 times per month from 18 October 2001 to 25 December 2003 using soda lime absorption. We found that R_R, R_L and R_SOM were initially higher in CC and SB plots than controls in both forests, but these three component fluxes in disturbed plots all fell below those of the control 5–20 months after the disturbance. Also, Q₁₀ values of these components decreased following disturbance. The annual flux of each respiration component was greater under BF than CF. The contribution of R_R to soil CO₂ efflux in the control plots averaged 35% in CF and 46% in BF. R_SOM was the dominant component of soil CO₂ efflux in CC and SB plots, accounting for over 50%. Our results highlight the importance of temporal trends of the component fluxes following disturbance and contribute to a broader understanding of forest management effects on the soil C cycle.     

Sources of CO2 efflux from soil and review of partitioning methods

Five main biogenic sources of CO₂ efflux from soils have been distinguished and described according to their turnover rates and the mean residence time of carbon. They are root respiration, rhizomicrobial respiration, decomposition of plant residues, the priming effect induced by root exudation or by addition of plant residues, and basal respiration by microbial decomposition of soil organic matter (SOM). These sources can be grouped in several combinations to summarize CO₂ efflux from the soil including: root-derived CO₂, plant-derived CO₂, SOM-derived CO₂, rhizosphere respiration, heterotrophic microbial respiration (respiration by heterotrophs), and respiration by autotrophs. These distinctions are important because without separation of SOM-derived CO₂ from plant-derived CO₂, measurements of total soil respiration have very limited value for evaluation of the soil as a source or sink of atmospheric CO₂ and for interpreting the sources of CO₂ and the fate of carbon within soils and ecosystems. Additionally, the processes linked to the five sources of CO₂ efflux from soil have various responses to environmental variables and consequently to global warming. This review describes the basic principles and assumptions of the following methods which allow SOM-derived and root-derived CO₂ efflux to be separated under laboratory and field conditions: root exclusion techniques, shading and clipping, tree girdling, regression, component integration, excised roots and insitu root respiration; continuous and pulse labeling, ¹³C natural abundance and FACE, and radiocarbon dating and bomb-¹⁴C. A short sections cover the separation of the respiration of autotrophs and that of heterotrophs, i.e. the separation of actual root respiration from microbial respiration, as well as methods allowing the amount of CO₂ evolved by decomposition of plant residues and by priming effects to be estimated. All these methods have been evaluated according to their inherent disturbance of the ecosystem and C fluxes, and their versatility under various conditions. The shortfalls of existing approaches and the need for further development and standardization of methods are highlighted.     

Impacts of extreme winter warming events on litter decomposition in a sub-Arctic heathland

Arctic climate change is expected to lead to a greater frequency of extreme winter warming events. During these events, temperatures rapidly increase to well above 0 °C for a number of days, which can lead to snow melt at the landscape scale, loss of insulating snow cover and warming of soils. However, upon return of cold ambient temperatures, soils can freeze deeper and may experience more freeze-thaw cycles due to the absence of a buffering snow layer. Such loss of snow cover and changes in soil temperatures may be critical for litter decomposition since a stable soil microclimate during winter (facilitated by snow cover) allows activity of soil organisms. Indeed, a substantial part of fresh litter decomposition may occur in winter. However, the impacts of extreme winter warming events on soil processes such as decomposition have never before been investigated. With this study we quantify the impacts of winter warming events on fresh litter decomposition using field simulations and lab studies.Winter warming events were simulated in sub-Arctic heathland using infrared heating lamps and soil warming cables during March (typically the period of maximum snow depth) in three consecutive years of 2007, 2008, and 2009. During the winters of 2008 and 2009, simulations were also run in January (typically a period of shallow snow cover) on separate plots. The lab study included soil cores with and without fresh litter subjected to winter-warming simulations in climate chambers.Litter decomposition of common plant species was unaffected by winter warming events simulated either in the lab (litter of Betula pubescens ssp. czerepanovii), or field (litter of Vaccinium vitis-idaea, and B. pubescens ssp. czerepanovii) with the exception of Vaccinium myrtillus (a common deciduous dwarf shrub) that showed less mass loss in response to winter warming events. Soil CO₂ efflux measured in the lab study was (as expected) highly responsive to winter warming events but surprisingly fresh litter decomposition was not. Most fresh litter mass loss in the lab occurred during the first 3-4 weeks (simulating the period after litter fall).In contrast to past understanding, this suggests that winter decomposition of fresh litter is almost non-existent and observations of substantial mass loss across the cold season seen here and in other studies may result from leaching in autumn, prior to the onset of “true” winter. Further, our findings surprisingly suggest that extreme winter warming events do not affect fresh litter decomposition.     

Modeling of the thermal influence of fires on the physicochemical properties and microbial activity of litter in cryogenic soils

Periodic surface fires in the cryolithozone (the northern taiga subzone) are the main factor determining the qualitative and quantitative characteristics of the soil organic matter. The specific features of the changes in the physicochemical parameters and microbial activity of the organic horizons in the cryogenic soils under larch forests of the northern taiga after the impact of high temperatures were revealed. The temperatures of fires of different intensity were simulated in laboratory conditions. The thermal impact on the litter organic matter during the surface fires may increase the CO₂ emission from the surface of the soil in the postfire communities due to the destruction of organic compounds only for a short time. After fires of high intensity with strong mineralization of the litters, during a period of more than 1 month, the pyrogenic effect on the organic horizons of the soils under the larch forests of the cryolithozone determined the reduction of the CO₂ emissions in the freshly burned areas as compared to the intact stands.     

Inter- and intra-specific interactions of Lumbricus rubellus (Hoffmeister, 1843) and Octolasion lacteum (Örley, 1881) (Lumbricidae) and the implication for C cycling

Earthworms are important engineering species of many terrestrial ecosystems as they play a significant role in regulating C turnover. The effects of earthworms on moderating C decomposition processes differ across species and with interactions between species, which is not fully understood. We carried out an experiment to study the interactions of Lumbricus rubellus and Octolasion lacteum, and their effects on soil respiration. Laboratory mesocosms were set up using tulip poplar (Liriodendron tulipifera) leaf litter and varying densities of earthworms in single and combined species treatments. CO₂ efflux rate was used as an indicator of C decomposition rates, and measured with CO₂ sensors every five days over one month. L. rubellus induced higher leaf consumption rate and higher CO₂ efflux than O. lacteum; meanwhile O. lacteum grew more than L. rubellus. Both litter consumption rate and growth rate of earthworms decreased with increasing earthworm density. Soil CO₂ efflux increased with increasing earthworm density (from ∼1-2 μg CO₂ g⁻¹ hr⁻¹ with no earthworms to ∼ 4 μg CO₂ g⁻¹ hr⁻¹ with 8 earthworms). Combining the two species had a synergistic effect on leaf litter consumption, and neutralizing effects on soil respiration. The data suggest that the strength of intra- and inter-specific interactions among earthworm ecological groups varies at different absolute and relative densities, leading to altered leaf litter decomposition and C cycling.     

Photodegradation effects on CO2 emissions from litter and SOM and photo-facilitation of microbial decomposition in a California grassland

Decomposition of soil organic matter (SOM) and plant litter has been shown to be affected by high solar radiation; this could partly explain why biogeochemical models underestimate decomposition in arid and semi-arid ecosystems. We set out to test the effect of using traditional PVC chambers for measuring soil gas fluxes versus quartz chambers that allowed passage of light during field measurements in a dry-land field in Davis, CA. Results showed that fluxes from quartz-top chambers were on average 29% higher than from opaque chambers. We also studied the effect of solar light exposure on decomposition of native grass litter and SOM in a field experiment where plots were shaded or left exposed for 157 days during summer; litter did not seem to be affected by exposure to light. However, we concluded that SOM decomposition was affected by light exposure since shaded soil had similar respiration to sunlight-exposed soil indicating that microbial respiration occurred under the shade while photo-degradation likely occurred under the sun. Additionally, ¹⁵N-labeled grass was placed in litter bags in the field with either clear filters to allow light or aluminum covers to block light; 3-month exposure caused a change in lignin degradability as indicated by the change in the Ad/Al ratio. Incubation of that litter showed 9.3% more CO₂ produced from litter in clear and aluminum bags than unexposed litter. This showed that photo-facilitation occurred although to a small degree and was a result of light exposure and/or heat degradation. We attributed the similar respiration from clear- and aluminum-exposed litter to heat degradation of the aluminum-exposed litter. In conclusion, our results show that in hot dry ecosystems conventional PVC chambers underestimate measured CO₂ flux rates; sunlight exposure changes litter chemistry and appears to affect the degradation of soil organic matter, but the magnitude of degradation depends on an interaction of factors such as soil temperature and moisture.     

Microbial enzyme activities in leaf litter, humus and mineral soil layers of European forests

The rationale of the study was to investigate microbial activity in different soil horizons in European forests. Hence, activities of chitinase and cellulase, microbial biomass carbon (C_mic) and basal respiration were measured in litter, fragmentation, humus and mineral soil layers collected several times from various beech and spruce forests. Sites were selected to form a gradient in N availability. Analyses were also performed on beech litter from a litterbag transplant experiment. Furthermore, microbiological parameters were measured in horizons of beech and spruce chronosequence sites with different stand age in order to investigate the influence of forest rotation, and hence changes in soil organic matter (SOM) dynamics, on microbial activity. Finally in horizons of one beech forest, the seasonal variation of selected microbiological parameters was measured more intensively. β-Glucosaminidase and cellobiohydrolase activities were measured using fluorogenic 4-methylumbelliferyl substrates to estimate chitinase and cellulase activities, respectively. On a spatial scale, chitinase and cellulase activities, C_mic determined by substrate induced respiration, and basal respiration ranged from 144 to 1924 and 6-177 nmol 4-MU g⁻¹ org-C h⁻¹, 8-48 mg C g⁻¹ org-C and 11-149 μg CO₂-C g⁻¹ org-C h⁻¹, respectively; in general values were significantly lower in layers of humus and mineral soil than of litter. Chitinase activity, C_mic and basal respiration from humus and mineral soil layers, together, correlated positively, while none correlated with cellulase activity. Similarly in the litter layer, no correlations were found between the microbiological parameters. On a seasonal scale, a time lag between a burst in basal respiration rate and activities of both enzymes were observed. In general, activities of cellulase and chitinase, C_mic and basal respiration, did not change with stand age, except in the humus layer in the spruce chronosequence, where C_mic decreased with stand age. In the litter layer, cellulase activity was significantly and positively related to the C:N ratio, while only a tendency for chitinase activity was shown, indicating that enzyme activities decreased with increasing N availability. In accordance, the enzyme activities and C_mic decreased significantly with increasing chronic N deposition in the humus layer, while basal respiration only tended to decrease with increasing N deposition. In contrast, enzyme activities in beech litter from litterbags after 2 years of incubation were generally higher at sites with higher N deposition. The results show different layer-specific responses of enzyme activities to changes in N availability, indicating different impacts of N availability on decomposition of SOM and stage of litter decomposition.     

Effects of soil mesofauna and farming management on decomposition of clover litter: a microcosm experiment

The effects of soil mesofauna and different farming systems on decomposition of clover (Trifolium repens) litter were investigated in a laboratory experiment. Microcosms were incubated for 16 weeks with fine and coarse litterbags in soils from three types of management systems: fallow, integrated farming and organic farming, the latter two cropped with wheat. The effects were studied by analysing litter mass loss, C and N content, DOC, nitrate and pH in soil leachate, and CO₂ production, as well as mesofauna. Mesofauna significantly accelerated mass loss and C and N release from clover litter in all three soils. With mesofauna access, at the end of the experiment average clover mass loss was almost twice as high and clover C and N content were 60% lower than without mesofauna. Farming systems influenced the decomposition through affecting both element turnover and mesofauna. Although in the first weeks less N was leached from organic farming than from integrated farming soil, cumulative N leaching did not differ between these soils. However, more than 20% less N was leached from the fallow soil than from the field soils. CO₂ production was highest in fallow soil. Here, mesofauna had no effect on this variable. In soil with integrated farming, mesofauna reduced cumulative CO₂ production by 10% whereas in soil from organic farming it increased CO₂ production by 20%. Our data suggest that differences in C and N turnover in different management systems are strongly mediated by soil mesofauna.     

Long-term effects of seasonal and diurnal temperature fluctuations on carbon dioxide efflux from a forest soil

Temperature fluctuations are a fundamental entity of the soil environment in the temperate zone and show fast (diurnal) and slow (seasonal) dynamics. Responses of soil respiration to temperature fluctuations were investigated in a root-free soil of a mid-European beech-oak forest. First, in laboratory we analysed the efflux of CO₂ from soil microcosms exposed to seasonal (±5 °C of the annual mean) and diurnal fluctuations (±5 °C of the seasonal levels) in a two-factorial design. Second, in field microcosms we investigated effects of smoothing diurnal temperature fluctuations in soil (simulating a possible global trend) on CO₂ efflux. Third, the natural temperature regime was simulated in laboratory microcosms and their CO₂ efflux was compared to the one in the field. The experiments lasted for 1 year to differentiate seasonal and annual responses.Dynamics of CO₂ efflux, microbial basal respiration, biomass and qO₂ varied with seasonal temperature regime. However, in the laboratory the annual cumulative CO₂-C production did not differ between treatments and varied between 10.9% and 11.7% of the total microcosm C, disregarding seasonal and/or diurnal fluctuations. The similarity of cumulative C production suggests that the availability of microbially mobilisable carbon pools rather than the temperature regime limited soil respiration. Diurnal fluctuations generally did not affect CO₂ efflux and microbial activity, though winter Q₁₀ values were increased in their absence. Simulation of the natural temperature regime in the laboratory resulted in CO₂ efflux similar to field microcosms. In the field, rates of CO₂ efflux and microbial activity, seasonal and annual cumulative CO₂-C production were significantly higher at smoothed than at natural temperature conditions (annually 13.1% and 11.0% of total C was respired, respectively). Facing global climate changes the mechanisms regulating responses of soil respiration to temperature fluctuations need further investigation.     

The CLIMEX soil-heating experiment: soil response after 2 years of treatment

 Most model predictions concerning the response of boreal forest ecosystems to climate change are inferred from small-scale experiments on artificial, simplified systems. Whole-ecosystem experiments designed to validate these models are scarce. We experimentally manipulated a small forested catchment in southern Norway by increasing soil temperature (+3  °C in summer to +5  °C in winter) using heating cables installed at 1 cm depth in the litter layer. Especially nitrification in the 0 to 10-cm soil layer increased as a result of the climate manipulation. Betula litter, produced after exposing trees for 2 years to ambient and elevated CO₂ in greenhouses, was incubated for 1 year in the manipulated catchment. Exposure to elevated CO₂ did not affect the C/N ratio or decomposition of the Betula litter, but lignin content decreased by 10%. We found no effect of elevated temperature on litter decomposition, probably due to desiccation of the litter. The heating cables caused a permanent increase in soil temperature in this soil layer, but when soils were dry, the temperature difference between control and heated plots decreased with increasing distance from the cables. When soils were wet, no gradients in temperature increase occurred. Received: 25 November 1997     

The bacterial community inhabiting temperate deciduous forests is vertically stratified and undergoes seasonal dynamics

Bacterial communities living in forest soils contribute to the decomposition of organic matter and the recycling of nutrients in these ecosystems and form one of the most diverse habitats on Earth. Unfortunately, due to difficulty in culturing soil bacteria, the understanding of their ecology is still limited. In the case of temperate deciduous forests, soil microbial communities face large seasonal variations in environmental conditions, such as temperature or moisture. Moreover, the supply of nutrients also differs due to seasonal processes, such as the allocation of photosynthates into soil by the roots of primary producers or the seasonal input of fresh litter. The aim of this study was to reveal how the bacterial community responds to these seasonal processes in the litter and soil of a Quercus petraea forest. Bacterial communities from litter and from the organic and mineral horizons of soil were analyzed during the four seasons of the year by 16S rRNA gene pyrosequencing. The results revealed that the composition of the bacterial community is horizon specific. The litter horizon had a higher relative abundance of Proteobacteria and Bacteroidetes than soil, while the organic and mineral horizons had a higher abundance of Acidobacteria, Firmicutes and Actinobacteria than litter. Moreover, the bacterial community was significantly affected by seasonality in all horizons. Bacterial communities in the litter showed significant differences between the vegetation season (May and July) and the autumn and winter seasons (October, February). In mineral soil, bacterial community composition was specific in the summer, when it was significantly different from all other seasons, with a larger number of taxa described as rhizosphere and mycorrhizosphere inhabitants. The results indicate that litter decomposition is the main driver of bacterial community composition in litter horizon. In contrast to reports on fungal communities, bacterial community composition in mineral soil responds to the seasonal peaks of rhizodeposition in the summer.