Elevated CO2 concentration and nitrogen fertilisation effects on N2O and CH4 fluxes and biomass production of Phleum pratense on farmed peat soil

The effects of elevated CO₂ supply on N₂O and CH₄ fluxes and biomass production of Phleum pratense were studied in a greenhouse experiment. Three sets of 12 farmed peat soil mesocosms (10 cm dia, 47 cm long) sown with P. pratense and equally distributed in four thermo-controlled greenhouses were fertilised with a commercial fertiliser in order to add 2, 6 or 10 g N m⁻². In two of the greenhouses, CO₂ concentration was kept at atmospheric concentration (360 μmol mol⁻¹) and in the other two at doubled concentration (720 μmol mol⁻¹). Soil temperature was kept at 15 °C and air temperature at 20 °C. Natural lighting was supported by artificial light and deionized water was used to regulate soil moisture. Forage was harvested and the plants fertilised three times during the basic experiment, followed by an extra fertilisations and harvests. At the end of the experiment CH₄ production and CH₄ oxidation potentials were determined; roots were collected and the biomass was determined. From the three first harvests the amount of total N in the aboveground biomass was determined. N₂O and CH₄ exchange was monitored using a closed chamber technique and a gas chromatograph. The highest N₂O fluxes (on average, 255 μg N₂O m⁻² h⁻¹ during period IV) occurred just after fertilisation at high water contents, and especially at the beginning of the growing season (on average, 490 μg N₂O m⁻² h⁻¹ during period I) when the competition of vegetation for N was low. CH₄ fluxes were negligible throughout the experiment, and for all treatments the production and oxidation potentials of CH₄ were inconsequential. Especially at the highest rates of fertilisation, the elevated supply of CO₂ increased above- and below-ground biomass production, but both at the highest and lowest rates of fertilisation, decreased the total amount of N in the aboveground dry biomass. N₂O fluxes tended to be higher under doubled CO₂ concentrations, indicating that increasing atmospheric CO₂ concentration may affect N and C dynamics in farmed peat soil.     

Methane flux characteristics in forest soils under an East Asian monsoon climate

Methane (CH₄) uptake by soil can possibly be suppressed more in regions with heavy summer precipitation, such as those under the East Asian monsoon climate, as compared to that in regions with a dry summer. In order to determine how precipitation patterns affect seasonal and spatial variations in CH₄ fluxes in temperate forest soils, such fluxes and selected environmental variables were measured on different parts of a hill slope in a cypress forest in central Japan. On the upper and middle parts of the slope, CH₄ uptake was observed throughout the year, and the uptake rates increased slightly with soil temperature and decreased with soil water content. The CH₄ flux predicted using data for the middle and upper parts of the slope ranged from −1.12 to −0.83 kg-CH₄ ha⁻¹ y⁻¹ (i.e. CH₄ uptake by soil) and from −2.30 to −2.04 kg-CH₄ ha⁻¹ y⁻¹, respectively. In contrast, in the relatively wet lower part of the slope near an in-stream wetland, large CH₄ emissions (>2 mg-CH₄ m⁻¹ d⁻¹) were observed during the rainy summer. In this wetter plot, the soil functioned as a net annual CH₄ source in a rainy year. Hence the variation in CH₄ flux with a change in soil water conditions and soil temperature on the lower part of the slope contrasted to that on the upper and middle parts of the slope. The predicted CH₄ flux for this lower plot ranged from −0.45 kg-CH₄ ha⁻¹ y⁻¹ in a dry year to 1.80 kg-CH₄ ha⁻¹ y⁻¹ in a rainy year. Our results suggest that consideration of the soil water conditions across a watershed is important for estimating the CH₄ budgets for entire forest watershed, particularly in regions subject to a wet summer.     

Short-term effects of clearfelling on soil CO2, CH4, and N2O fluxes in a Sitka spruce plantation

We examined the effects of forest clearfelling on the fluxes of soil CO₂, CH₄, and N₂O in a Sitka spruce (Picea sitchensis (Bong.) Carr.) plantation on an organic-rich peaty gley soil, in Northern England. Soil CO₂, CH₄, N₂O as well as environmental factors such as soil temperature, soil water content, and depth to the water table were recorded in two mature stands for one growing season, at the end of which one of the two stands was felled and one was left as control. Monitoring of the same parameters continued thereafter for a second growing season. For the first 10 months after clearfelling, there was a significant decrease in soil CO₂ efflux, with an average efflux rate of 4.0 g m⁻² d⁻¹ in the mature stand (40-year) and 2.7 g m⁻² d⁻¹ in clearfelled site (CF). Clearfelling turned the soil from a sink (−0.37 mg m⁻² d⁻¹) for CH₄ to a net source (2.01 mg m⁻² d⁻¹). For the same period, soil N₂O fluxes averaged 0.57 mg m⁻² d⁻¹ in the CF and 0.23 mg m⁻² d⁻¹ in the 40-year stand. Clearfelling affected environmental factors and lead to higher daily soil temperatures during the summer period, while it caused an increase in the soil water content and a rise in the water table depth. Despite clearfelling, CO₂ remained the dominant greenhouse gas in terms of its greenhouse warming potential.     

CH4 oxidation and emissions of CH4 and N2O from Lolium perenne swards under elevated atmospheric CO2

Emissions of N₂O and CH₄ and CH₄ oxidation rates were measured from Lolium perenne swards in a short-term study under ambient (36 Pa) and elevated (60 Pa) atmospheric CO₂ at the Free Air Carbon dioxide Enrichment experiment, Eschikon, Switzerland. Elevated pCO₂ increased (P<0.05) N₂O emissions from high N fertilised (11.2 g N m⁻²) swards by 69%, but had no significant effect on net emissions of CH₄. Application of ¹³C-CH₄ (11 μl l⁻¹; 11 at.% excess ¹³C) to closed chamber headspaces in microplots enabled determination of rates of ¹³C-CH₄ oxidation even when net CH₄ fluxes from main plots were positive. We found a significant interaction between fertiliser application rate and atmospheric pCO₂ on ¹³C-CH₄ oxidation rates that was attributed to differences in gross nitrification rates and C and N availability. CH₄ oxidation was slower and thought to be temporarily inhibited in the high N ambient pCO₂ sward. The most rapid CH₄ oxidation of 14.6 μg ¹³C-CH₄ m⁻² h⁻¹ was measured in the high fertilised elevated pCO₂ sward, and we concluded that either elevated pCO₂ had a stimulatory effect on CH₄ oxidation or inhibition of oxidation following fertiliser application was lowered under elevated pCO₂. Application of ¹⁴NH₄¹⁵NO₃ and ¹⁵NH₄¹⁵NO₃ (10 at.% excess ¹⁵N) to different replicates enabled determination of the respective contributions of nitrification and denitrification to N₂O emissions. Inhibition of CH₄ oxidation in the high fertilised ambient pCO₂ sward, due to competition between NH₃ and CH₄ for methane monooxygenase enzymes or toxic effects of NH₂OH or NO₂⁻ produced during nitrification, was hypothesised to increase gross nitrification (12.0 mg N kg dry soil⁻¹) and N₂O emissions during nitrification (327 mg ¹⁵N-N₂O m⁻² over 11 d). Our results indicate that increasing atmospheric concentrations of CO₂ may increase emissions of N₂O by denitrification, lower nitrification rates and either increase or decrease the ability of soil to act as a sink for atmospheric CH₄ depending on fertiliser management.     

Methane uptake in soils from Pinus radiata plantations, a reverting shrubland and adjacent pastures: Effects of land-use change, and soil texture, water and mineral nitrogen

Afforestation and reforestation of pastures are key land-use changes in New Zealand that help sequester carbon (C) to offset its carbon dioxide (CO₂) emissions under the Kyoto Protocol. However, relatively little attention has been given so far to associated changes in trace gas fluxes. Here, we measure methane (CH₄) fluxes and CO₂ production, as well as microbial C, nitrogen (N) and mineral-N, in intact, gradually dried (ca. 2 months at 20 °C) cores of a volcanic soil and a heavier textured, non-volcanic soil collected within plantations of Pinus radiata D. Don (pine) and adjacent permanent pastures. CH₄ fluxes and CO₂ production were also measured in cores of another volcanic soil under reverting shrubland (mainly Kunzea var. ericoides (A. Rich) J. Thompson) and an adjacent pasture. CH₄ uptake in the pine and shrubland cores of the volcanic soils at field capacity averaged about 35 and 14 μg CH₄-C m⁻² h⁻¹, respectively, and was significantly higher than in the pasture cores (about 21 and 6 μg CH₄-C m⁻² h⁻¹, respectively). In the non-volcanic soil, however, CH₄-C uptake was similar in most cores of the pine and pasture soils, averaging about 7-9 μg m⁻² h⁻¹, except in very wet samples. In contrast, rates of CO₂ production and microbial C and N concentrations were significantly lower under pine than under pasture. In the air-dry cores, microbial C and N had declined in the volcanic soil, but not in the non-volcanic soil; ammonium-N, and especially nitrate-N, had increased significantly in all samples. CH₄ uptake was, with few exceptions, not significantly influenced by initial concentrations of ammonium-N or nitrate-N, nor by their changes on air-drying. A combination of phospholipid fatty acid (PLFA) and stable isotope probing (SIP) analyses of only the pine and pasture soils showed that different methanotrophic communities were probably active in soils under the different vegetations. The C18 PLFAs (type II methanotrophs) predominated under pine and C16 PLFAs (type I methanotrophs) predominated under pasture. Overall, vegetation, soil texture, and water-filled pore space influenced CH₄-C uptake more than did soil mineral-N concentrations.     

Temporal change in spatial variability of soil respiration on a slope of Japanese cedar (Cryptomeria japonica D. Don) forest

Although information regarding the spatial variability of soil respiration is important for understanding carbon cycling and developing a suitable sampling design for estimating average soil respiration, it remains relatively understudied compared to temporal changes. In this study, soil respiration was measured at 35 locations by season on a slope of Japanese cedar forest in order to examine temporal changes in the spatial distribution of soil respiration. Spatial variability of soil respiration varied between seasons, with the highest coefficient variation in winter (42%) and lowest in summer (26%). Semivariogram analysis and kriged maps revealed different patterns of spatial distribution in each season. Factors affecting the spatial variability were relief index (autumn), soil hardness of the A layer (winter), soil hardness at 50 cm depth (spring) and the altitude and relief index (summer). Annual soil respiration (average: 39 mol m⁻² y⁻¹) varied from 26 mol m⁻² y⁻¹ to 55 mol m⁻² y⁻¹ between the 35 locations and was higher in the upper part of the slope and lower in the lower part. The average Q₁₀ value was 2.3, varying from 1.3 to 3.0 among the locations. These findings suggest that insufficient information on the spatial variability of soil respiration and imbalanced sampling could bias estimates of current and future carbon budgets.     

Seasonal variation and fire effects on CH4, N2O and CO2 exchange in savanna soils of northern Australia

Tropical savanna ecosystems are a major contributor to global CO₂, CH₄ and N₂O greenhouse gas exchange. Savanna fire events represent large, discrete C emissions but the importance of ongoing soil-atmosphere gas exchange is less well understood. Seasonal rainfall and fire events are likely to impact upon savanna soil microbial processes involved in N₂O and CH₄ exchange. We measured soil CO₂, CH₄ and N₂O fluxes in savanna woodland (Eucalyptus tetrodonta/Eucalyptus miniata trees above sorghum grass) at Howard Springs, Australia over a 16 month period from October 2007 to January 2009 using manual chambers and a field-based gas chromatograph connected to automated chambers. The effect of fire on soil gas exchange was investigated through two controlled burns and protected unburnt areas. Fire is a frequent natural and management action in these savanna (every 1-2 years). There was no seasonal change and no fire effect upon soil N₂O exchange. Soil N₂O fluxes were very low, generally between −1.0 and 1.0 μg N m⁻² h⁻¹, and often below the minimum detection limit. There was an increase in soil NH₄⁺ in the months after the 2008 fire event, but no change in soil NO₃⁻. There was considerable nitrification in the early wet season but minimal nitrification at all other times.Savanna soil was generally a net CH₄ sink that equated to between −2.0 and −1.6 kg CH₄ ha⁻¹ y⁻¹ with no clear seasonal pattern in response to changing soil moisture conditions. Irrigation in the dry season significantly reduced soil gas diffusion and as a consequence soil CH₄ uptake. There were short periods of soil CH₄ emission, up to 20 μg C m⁻² h⁻¹, likely to have been caused by termite activity in, or beneath, automated chambers. Soil CO₂ fluxes showed a strong bimodal seasonal pattern, increasing fivefold from the dry into the wet season. Soil moisture showed a weak relationship with soil CH₄ fluxes, but a much stronger relationship with soil CO₂ fluxes, explaining up to 70% of the variation in unburnt treatments. Australian savanna soils are a small N₂O source, and possibly even a sink. Annual soil CH₄ flux measurements suggest that the 1.9 million km² of Australian savanna soils may provide a C sink of between −7.7 and −9.4 Tg CO₂-e per year. This sink estimate would offset potentially 10% of Australian transport related CO₂-e emissions. This CH₄ sink estimate does not include concurrent CH₄ emissions from termite mounds or ephemeral wetlands in Australian savannas.     

Influence of warming and enchytraeid activities on soil CO2 and CH4 fluxes

To determine the sum of ‘direct’ and ‘indirect’ effects of climatic change on enchytraeid activity and C fluxes from an organic soil we assessed the influence of temperature (4, 10 and 15 °C incubations) on enchytraeid populations and soil CO₂ and CH₄ fluxes over 116 days. Moisture was maintained at 60% of soil dry weight during the experimental period and measurements of enchytraeid biomass and numbers, and CO₂ and CH₄ fluxes were made after 3, 16, 33, 44, 65, 86 and 116 days. Enchytraeid population numbers and biomass increased in all temperature treatments with the greatest increase produced at 15 °C (to over threefold initial values by day 86). Results also showed that enchytraeid activity increased CO₂ fluxes by 10.7±4.5, 3.4±4.0 and 26.8±2.6% in 4, 10 and 15 °C treatments, respectively, with the greatest CO₂ production observed at 15 °C for the entire 116 day incubation period (P<0.05). The soil respiratory quotient analyses at lower temperatures (i.e. 4-10 °C) gave a Q₁₀ of 1.7 and 1.9 with and without enchytraeids, respectively. At temperatures above 10 °C (i.e. 10-15 °C) Q₁₀ significantly increased (P<0.01) and was 25% greater in the presence of enchytraeids (Q₁₀=3.4) than without (Q₁₀=2.6). In contrast to CO₂ production, no significant relationships were observed between net CH₄ fluxes and temperature and only time showed a significant effect on CH₄ production (P<0.01).Total soil CO₂ production was positively linked with enchytraeid biomass and mean soil CO₂-C production was 77.01±6.05 CO₂-C μg mg enchytraeid tissue⁻¹ day⁻¹ irrespective of temperature treatment. This positive relationship was used to build a two step regression model to estimate the effects of temperature on enchytraeid biomass and soil CO₂ respiration in the field. Predictions of potential CO₂ production were made using enchytraeid biomass data obtained in the field from two upland grassland sites (Sourhope and Great Dun Fell at the Moor House Nature Reserve, both in the UK). The findings of this work suggest that a 5 °C increase in atmospheric temperature above mean ambient temperature could have the potential to produce a significant increase in enchytraeid biomass resulting in a near twofold increase in soil CO₂ release from both soil types. The interaction between temperature and soil biology will clearly be an important determinant of soil respiration responses to global warming.     

Lead tolerance in Aporrectodea rosea earthworms from a clay pigeon shooting site

Lead tolerance in individuals of the earthworm species Aporrectodea rosea collected from a clay pigeon shooting site was investigated. Lead concentrations in the shooting site soil and the un-shot control site were 6410±2250 and 296±98 mg_Pb kg⁻¹ dry weight, respectively. Of these concentrations 1050±240 and 12±9 mg_Pb kg⁻¹ dry weight were suggested to be available, using ammonium acetate (1 M), respectively. With respect to earthworm body burdens of lead the shooting site earthworms had a body burden of 6.1±1.2 mg_Pb g⁻¹ dry weight while the uncontaminated site earthworms had almost a 1000-times lower body burden of 7.1±9.0 μg_Pb g⁻¹ dry weight. Lead tolerance was assessed in uncontaminated soil that had been augmented with lead, using lead nitrate solutions, to obtain lead concentrations in soil of 0.5, 5 and 50 mg_Pb kg⁻¹ dry weight. Earthworms were exposed for 28 days during which time a semi-qualitative assessment was made of their condition. Results showed no decrease in condition in the shooting site earthworms with increasing exposure time or concentration. In contrast, earthworms collected from an uncontaminated site showed a significant (p<0.05) decrease in condition when exposed to lead concentrations above, and including, a concentration of 5 mg kg⁻¹ dry weight soil. These results suggested lead tolerance in the shooting site earthworms.     

Pathways and dynamics of NO3 and NH4 applied in a mountain Picea abies forest and in a nearby meadow in central Switzerland

To evaluate the pathways and dynamics of inorganic nitrogen (N) deposition in previously N-limited ecosystems, field additions of ¹⁵N tracers were conducted in two mountain ecosystems, a forest dominated by Norway spruce (Picea abies) and a nearby meadow, at the Alptal research site in central Switzerland. This site is moderately impacted by N from agricultural and combustion sources, with a bulk atmospheric deposition of 12 kg N ha⁻¹ y⁻¹ equally divided between NH₄⁺ and NO₃⁻. Pulses of ¹⁵NH₄⁺ and ¹⁵NO₃⁻ were applied separately as tracers on plots of 2.25 m². Several ecosystem pools were sampled at short to longer-term intervals (from a few hours to 1 year), above and belowground biomass (excluding trees), litter layer, soil LF horizon (approx. 5-0 cm), A horizon (approx. 0-5 cm) and gleyic B horizon (5-20 cm). Furthermore, extractable inorganic N, and microbial N pools were analysed in the LF and A horizons. Tracer recovery patterns were quite similar in both ecosystems, with most of the tracer retained in the soil pool. At the short-term (up to 1 week), up to 16% of both tracers remained extractable or entered the microbial biomass. However, up to 30% of the added ¹⁵NO₃⁻ was immobilised just after 1 h, and probably chemically bound to soil organic matter. 16% of the NH₄⁺ tracer was also immobilised within hours, but it is not clear how much was bound to soil organic matter or fixed between layers of illite-type clay. While the extractable and microbial pools lost ¹⁵N over time, a long-term increase in ¹⁵N was measured in the roots. Otherwise, differences in recovery a few hours after labelling and 1 year later were surprisingly small. Overall, more NO₃⁻ tracer than NH₄⁺ tracer was recovered in the soil. This was due to a strong aboveground uptake of the deposited NH₄⁺ by the ground vegetation, especially by mosses.     

Seasonal changes in the spatial structures of N2O, CO2, and CH4 fluxes from Acacia mangium plantation soils in Indonesia

We evaluated the spatial structures of nitrous oxide (N₂O), carbon dioxide (CO₂), and methane (CH₄) fluxes in an Acacia mangium plantation stand in Sumatra, Indonesia, in drier (August) and wetter (March) seasons. A 60 × 100-m plot was established in an A. mangium plantation that included different topographical elements of the upper plateau, lower plateau, upper slope and foot slope. The plot was divided into 10 × 10-m grids and gas fluxes and soil properties were measured at 77 grid points at 10-m intervals within the plot. Spatial structures of the gas fluxes and soil properties were identified using geostatistical analyses. Averaged N₂O and CO₂ fluxes in the wetter season (1.85 mg N m⁻² d⁻¹ and 4.29 g C m⁻² d⁻¹, respectively) were significantly higher than those in the drier season (0.55 mg N m⁻² d⁻¹ and 2.73 g C m⁻² d⁻¹, respectively) and averaged CH₄ uptake rates in the drier season (−0.62 mg C m⁻² d⁻¹) were higher than those in the wetter season (−0.24 mg C m⁻² d⁻¹). These values of N₂O fluxes in A. mangium soils were higher than those reported for natural forest soils in Sumatra, while CO₂ and CH₄ fluxes were in the range of fluxes reported for natural forest soils. Seasonal differences in these gas fluxes appears to be controlled by soil water content and substrate availability due to differing precipitation and mineralization of litter between seasons. N₂O fluxes had strong spatial dependence with a range of about 18 m in both the drier and wetter seasons. Topography was associated with the N₂O fluxes in the wetter season with higher and lower fluxes on the foot slope and on the upper plateau, respectively, via controlling the anaerobic-aerobic conditions in the soils. In the drier season, however, we could not find obvious topographic influences on the spatial patterns of N₂O fluxes and they may have depended on litter amount distribution. CO₂ fluxes had no spatial dependence in both seasons, but the topographic influence was significant in the drier season with lowest fluxes on the foot slope, while there was no significant difference between topographic positions in the wetter season. The distributions of litter amount and soil organic matter were possibly associated with CO₂ fluxes through their effects on microbial activities and fine root distribution in this A. mangium plantation.     

Methane oxidation in temperate soils: effects of inorganic N

Additions of inorganic nitrogen (N) to an oak soil with significant potential for methane (CH₄) oxidation resulted in differential reduction in CH₄ oxidation capacity depending on N species added. Nitrate, rather than nitrite or ammonium, proved to be the strongest inhibitor of CH₄ oxidation in oak soil. Both high (CH₄ at 10 μl l⁻¹) and low (CH₄ at 5 ml l⁻¹) affinity CH₄ oxidation in oak soil was completely inhibited at a nitrate concentration similar to that present in an alder soil from the same experimental site. The alder soil showed no capacity for low affinity CH₄ oxidation. A ‘low nitrate’ forest soil (oak) showed high affinity, low capacity CH₄ oxidation upto around 1 ml l⁻¹ CH₄, above which both high and low affinity CH₄ oxidation became apparent following a lag phase, indicating either an induced high affinity uptake mechanism or the existence of distinct low affinity and high affinity methanotroph populations. High affinity CH₄ oxidation became saturated at CH₄ concentrations >500 μl l⁻¹, while low affinity CH₄ oxidation became saturated at ∼30 ml l⁻¹ CH₄. In a ‘high nitrate’ forest soil (alder), CH₄ oxidation appeared to be due to high affinity CH₄ oxidation only and became undetectable at CH₄ concentrations >5 ml l⁻¹.     

Temporal and spatial dynamics of soil solution C and N concentrations during Lolium perenne L. sward establishment and the effects of elevated CO2 and N additions

There is now clear evidence for a prolonged increase in atmospheric CO₂ concentrations and enrichment of the biosphere with N. Understanding the fate of C in the plant-soil system under different CO₂ and N regimes is therefore of considerable importance in predicting the environmental effects of climate change and in predicting the sustainability of ecosystems. Swards of Lolium perenne were grown from seed in a Eutric Cambisol at either ambient (ca. 350 μmol mol⁻¹) or elevated (700 μmol mol⁻¹) atmospheric pCO₂ and subjected to two inorganic N fertilizer regimes (no added N and 70 kg N ha⁻¹ month⁻¹). After germination, soil solution concentrations of dissolved organic C (DOC), dissolved inorganic N (DIN), dissolved organic N (DON), phenolics and H⁺ were measured at five depths down the soil profile over 3 months. The exploration of soil layers down the soil profile by roots caused transient increases in soil solution DOC, DON and phenolic concentrations, which then subsequently returned to lower quasi-stable concentrations. In general, the addition of N tended to increase DOC and DON concentrations while exposure to elevated pCO₂ had the opposite effect. These treatment effects, however, gradually diminished over the duration of the experiment from the top of the soil profile downwards. The ambient pCO₂ plus added N regime was the only treatment to maintain a notable difference in soil solution solute concentration, relative to other treatments. This effect on soil solution chemistry appeared to be largely indirect resulting from increased plant growth and a decrease in soil moisture content. Our results show that although plant growth responses to elevated pCO₂ are critically dependent upon N availability, the organic chemistry of the soil solution is relatively insensitive to changes in plant growth once the plants have become established.     

CH4 oxidation and N2O emissions at varied soil water-filled pore spaces and headspace CH4 concentrations

Emission of N₂O and CH₄ oxidation rates were measured from soils of contrasting (30-75%) water-filled pore space (WFPS). Oxidation rates of ¹³C-CH₄ were determined after application of 10 μl ¹³C-CH₄ l⁻¹ (10 at. % excess ¹³C) to soil headspace and comparisons made with estimates from changes in net CH₄ emission in these treatments and under ambient CH₄ where no ¹³C-CH₄ had been applied. We found a significant effect of soil WFPS on ¹³C-CH₄ oxidation rates and evidence for oxidation of 2.2 μg ¹³C-CH₄ d⁻¹ occurring in the 75% WFPS soil, which may have been either aerobic oxidation occurring in aerobic microsites in this soil or anaerobic CH₄ oxidation. The lowest ¹³C-CH₄ oxidation rate was measured in the 30% WFPS soil and was attributed to inhibition of methanotroph activity in this dry soil. However, oxidation was lowest in the wetter soils when estimated from changes in concentration of ¹²⁺¹³C-CH₄. Thus, both methanogenesis and CH₄ oxidation may have been occurring simultaneously in these wet soils, indicating the advantage of using a stable isotope approach to determine oxidation rates. Application of ¹³C-CH₄ at 10 μl ¹³C-CH₄ l⁻¹ resulted in more rapid oxidation than under ambient CH₄ conditions, suggesting CH₄ oxidation in this soil was substrate limited, particularly in the wetter soils. Application of and (80 mg N kg soil⁻¹; 9.9 at.% excess ¹⁵N) to different replicates enabled determination of the respective contributions of nitrification and denitrification to N₂O emissions. The highest N₂O emission (119 μg ¹⁴⁺¹⁵N-N₂O kg soil⁻¹ over 72 h) was measured from the 75% WFPS soil and was mostly produced during denitrification (18.1 μg ¹⁵N-N₂O kg soil⁻¹; 90% of ¹⁵N-N₂O from this treatment). Strong negative correlations between ¹⁴⁺¹⁵N-N₂O emissions, denitrified ¹⁵N-N₂O emissions and ¹³C-CH₄ concentrations (r=−0.93 to −0.95, N₂O; r=−0.87 to −0.95, denitrified ¹⁵N-N₂O; P<0.05) suggest a close relationship between CH₄ oxidation and denitrification in our soil, the nature of which requires further investigation.     

Methane oxidation in boreal forest soils: kinetics and sensitivity to pH and ammonium

In soil incubation experiments we examined if there are differences in the kinetic parameters of atmospheric methane (CH₄) oxidation in soils of upland forests and forested peatlands. All soils showed net uptake of atmospheric CH₄. One of the upland forests included also managed (clear-cut with or without previous liming or N-fertilization) study plots. The CH₄ oxidation in the forested peat soil had a higher K_m (510 μl l⁻¹) and V_max (6.2 nmol CH₄ cm⁻³ h⁻¹) than the upland forest soils (K_m from 5 to 18 μl l⁻¹ and V_max from 0.15 to 1.7 nmol CH₄ cm⁻³ h⁻¹). The forest managements did not affect the K_m-values. At atmospheric CH₄ concentration, the upland forest soils had a higher CH₄ oxidation activity than the forested peat soil; at high CH₄ concentrations the reverse was true. Most of the soils oxidised CH₄ in the studied pH range from 3 to 7.5. The pH optimum for CH₄ oxidation varied from 4 to 7.5. Some of the soils had a pH optimum for CH₄ oxidation that was above their natural pH. The CH₄ oxidation in the upland forest soils and in the peat soil did not differ in their sensitivities to (NH₄)₂SO₄ or K₂SO₄ (used as a non-ammonium salt control). Inhibition of CH₄ oxidation by (NH₄)₂SO₄ resulted mainly from a general salt effect (osmotic stress) though NH₄⁺ did have some additional inhibitory properties. Both salts were better inhibitors of CH₄ oxidation than respiration. The differences in the CH₄ oxidation kinetics in the forested peat soil and in the upland forest soils reveal that there are differences in the physiologies of the CH₄ oxidisers in these soils.     

Elevation of atmospheric CO2 and N-nutritional status modify nodulation, nodule-carbon supply, and root exudation of Phaseolus vulgaris L.

Increased root exudation and a related stimulation of rhizosphere-microbial growth have been hypothesised as possible explanations for a lower nitrogen- (N-) nutritional status of plants grown under elevated atmospheric CO₂ concentrations, due to enhanced plant-microbial N competition in the rhizosphere. Leguminous plants may be able to counterbalance the enhanced N requirement by increased symbiotic N₂ fixation. Only limited information is available about the factors determining the stimulation of symbiotic N₂ fixation in response to elevated CO₂.In this study, short-term effects of elevated CO₂ on quality and quantity of root exudation, and on carbon supply to the nodules were assessed in Phaseolus vulgaris, grown in soil culture with limited (30 mg N kg⁻¹ soil) and sufficient N supply (200 mg N kg⁻¹ soil), at ambient (400 μmol mol⁻¹) and elevated (800 μmol mol⁻¹) atmospheric CO₂ concentrations.Elevated CO₂ reduced N tissue concentrations in both N treatments, accelerated the expression of N deficiency symptoms in the N-limited variant, but did not affect plant biomass production. ¹⁴CO₂ pulse-chase labelling revealed no indication for a general increase in root exudation with subsequent stimulation of rhizosphere microbial growth, resulting in increased N-competition in the rhizosphere at elevated CO₂. However, a CO₂-induced stimulation in root exudation of sugars and malate as a chemo-attractant for rhizobia was detected in 0.5-1.5 cm apical root zones as potential infection sites. Particularly in nodules, elevated CO₂ increased the accumulation of malate as a major carbon source for the microsymbiont and of malonate with essential functions for nodule development. Nodule number, biomass and the proportion of leghaemoglobin-producing nodules were also enhanced. The release of nod-gene-inducing flavonoids (genistein, daidzein and coumestrol) was stimulated under elevated CO₂, independent of the N supply, and was already detectable at early stages of seedling development at 6 days after sowing.     

Influence of water table levels on CO2 emissions in a Colorado subalpine fen: an in situ microcosm study

We quantified the relationship between water table position and CO₂ emissions by manipulating water table levels for two summers in microcosms installed in a Colorado subalpine fen. Water levels were manipulated in the microcosms by either adding water or removing water and ranged from +10 cm above the soil surface to 40 cm below the soil surface, with ambient water levels in the fen averaging +3 and +2 cm above the soil surface during 1998 and 1999, respectively. Microcosm installation had no significant effect on CO₂ efflux; the 2 year means of natural and reference CO₂ efflux were 205.4 and 213.9 mg CO₂-C m⁻² h⁻¹, respectively (p=0.80). Mean CO₂ emissions were lowest at the highest water tables (water +6 to +10 cm above the soil surface), averaging 133.8 mg CO₂-C m⁻² h⁻¹, increased to 231.3 mg CO₂-C m⁻² h⁻¹ when the water table was +1 to +5 cm above the soil surface and doubled to 453.7 mg CO₂-C m⁻² h⁻¹, when the water table was 0-5 cm below the soil surface. However, further lowering of the water table had little additional effect on CO₂ emissions, which averaged 470.3 and 401.1 mg CO₂-C m⁻² h⁻¹ when the water table was 6-10 cm, and 11-40 cm beneath the soil surface, respectively. The large increase in CO₂ emissions as we experimentally lowered the water table beneath the soil surface, coupled with no increase in CO₂ emissions as we furthered lowered water tables beneath the soil surface, suggest the presence of an easily oxidized labile carbon pool near the soil surface.     

Nitrate leaching in soil: Tracing the NO3 sources with the help of stable N and O isotopes

Legumes increase the plant-available N pool in soil, but might also increase NO₃⁻ leaching to groundwater. To minimize NO₃⁻ leaching, N-release processes and the contribution of legumes to NO₃⁻ concentrations in soil must be known. Our objectives were (1) to quantify NO₃⁻-N export to >0.3 m soil depth from three legume monocultures (Medicago x varia Martyn, Onobrychis viciifolia Scop., Lathyrus pratensis L.) and from three bare ground plots. Furthermore, we (2) tested if it is possible to apply a mixing model for NO₃⁻ in soil solution based on its dual isotope signals, and (3) estimated the contribution of legume mineralization to NO₃⁻ concentrations in soil solution under field conditions. We collected rainfall and soil solution at 0.3 m soil depth during 1 year, and determined NO₃⁻ concentrations and δ¹⁵N and δ¹⁸O of NO₃⁻ for >11.5 mg NO₃⁻-N l⁻¹. We incubated soil samples to assess potential N release by mineralization and determined δ¹⁵N and δ¹⁸O signals of NO₃⁻ derived from mineralization of non-leguminous and leguminous organic matter.Mean annual N export to >0.3 m soil depth was highest in bare ground plots (9.7 g NO₃⁻-N m⁻²; the SD reflects the spatial variation) followed by Medicago x varia monoculture (6.0 g NO₃⁻-N m⁻²). The O. viciifolia and L. pratensis monocultures had a much lower mean annual N export (0.5 and 0.3 g NO₃⁻-N m⁻²). The averaged NO₃⁻-N leaching during 70 days was not significantly different between field estimates and incubation for the Medicago x varia Martyn monoculture.The δ¹⁵N and δ¹⁸O values in NO₃⁻ of rainfall (δ¹⁵N: 3.3±0.8‰; δ¹⁸O: 30.8±4.7‰), mineralization of non-leguminous SOM (9.3±0.9‰; 6.7±0.8‰), and mineralization of leguminous SOM (1.5±0.6‰; 5.1±0.9‰) were markedly different. Applying a linear mixing model based on these three sources to δ¹⁵N and δ¹⁸O values in NO₃⁻ of soil solution during winter 2003, we calculated 18-41% to originate from rainfall, 38-57% from mineralization of non-leguminous SOM, and 18-40% from mineralization of leguminous SOM.Our results demonstrate that (1) even under legumes NO₃⁻-N leaching was reduced compared to bare ground, (2) the application of a three-end-member mixing model for NO₃⁻ based on its dual isotope signals produced plausible results and suggests that under particular circumstances such models can be used to estimate the contributions of different NO₃⁻ sources in soil solution, and (3) in the 2nd year after establishment of legumes, they contributed approximately one-fourth to NO₃⁻-N loss.     

Carbon dynamics in a temperate grassland soil after 9 years exposure to elevated CO2 (Swiss FACE)

Elevated pCO₂ increases the net primary production, C/N ratio, and C input to the soil and hence provides opportunities to sequester CO₂-C in soils to mitigate anthropogenic CO₂. The Swiss 9 y grassland FACE (free air carbon-dioxide enrichment) experiment enabled us to explore the potential of elevated pCO₂ (60 Pa), plant species (Lolium perenne L. and Trifolium repens L.) and nitrogen fertilization (140 and 540 kg ha⁻¹ y⁻¹) on carbon sequestration and mineralization by a temperate grassland soil. Use of ¹³C in combination with respired CO₂ enabled the identification of the origins of active fractions of soil organic carbon. Elevated pCO₂ had no significant effect on total soil carbon, and total soil carbon was also independent of plant species and nitrogen fertilization. However, new (FACE-derived depleted ¹³C) input of carbon into the soil in the elevated pCO₂ treatments was dependent on nitrogen fertilization and plant species. New carbon input into the top 15 cm of soil from L. perennne high nitrogen (LPH), L. perenne low nitrogen (LPL) and T. repens low nitrogen (TRL) treatments during the 9 y elevated pCO₂ experiment was 9.3±2.0, 12.1±1.8 and 6.8±2.7 Mg C ha⁻¹, respectively. Fractions of FACE-derived carbon in less protected soil particles >53 μm in size were higher than in <53 μm particles. In addition, elevated pCO₂ increased CO₂ emission over the 118 d incubation by 55, 61 and 13% from undisturbed soil from LPH, LPL and TRL treatments, respectively; but only by 13, 36, and 18%, respectively, from disturbed soil (without roots). Higher input of new carbon led to increased decomposition of older soil organic matter (priming effect), which was driven by the quantity (mainly roots) of newly input carbon (L. perenne) as well as the quality of old soil carbon (e.g. higher recalcitrance in T. repens). Based on these results, the potential of well managed and established temperate grassland soils to sequester carbon under continued increasing concentrations of atmospheric CO₂ appears to be rather limited.     

Carbon dioxide, nitrous oxide and methane dynamics in boreal organic agricultural soils with different soil characteristics

The annual carbon dioxide (CO₂), nitrous oxide (N₂O) and methane (CH₄) dynamics were measured with static chambers on two organic agricultural soils with different soil characteristics. Site 1 had a peat layer of 30 cm, with an organic matter (OM) content of 74% in the top 20 cm. Site 2 had a peat layer of 70 cm but an OM content of only 40% in the top 20 cm. On both sites there were plots under barley and grass and also plots where the vegetation was removed. All soils were net sources of CO₂ and N₂O, but they consumed atmospheric CH₄. Soils under barley had higher net CO₂ emissions (830 g CO₂-C m⁻² yr⁻¹) and N₂O emissions (848 mg N₂O-N m⁻² yr⁻¹) than those under grass (395 g CO₂-C m⁻³ yr⁻¹ and 275 mg N₂O-N m⁻² yr⁻¹). Bare soils had the highest N₂O emissions, mean 2350 mg N₂O-N m⁻² yr⁻¹. The mean CH₄ uptake rate from vegetated soils was 100 mg CH₄-C m⁻³ yr⁻¹ and from bare soils 55 mg CH₄-C m⁻² yr⁻¹. The net CO₂ emissions were higher from Site 2, which had a high peat bulk density and a low OM content derived from the addition of mineral soil to the peat during the cultivation history of that site. Despite the differences in soil characteristics, the mean N₂O emissions were similar from vegetated peat soils from both sites. However, bare soils from Site 2 with mineral soil addition had N₂O emissions of 2-9 times greater than those from Site 1. Site 1 consumed atmospheric CH₄ at a higher rate than Site 2 with additional mineral soil. N₂O emissions during winter were an important component of the N₂O budget even though they varied greatly, ranging from 2 to 99% (mean 26%) of the annual emission.