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.     

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.     

Soil CO2 flux in six monospecific forest plantations in Southern Rwanda

Forest soils contain the largest carbon stock of all terrestrial biomes and are probably the most important source of carbon dioxide (CO₂) to atmosphere. Soil CO₂ fluxes from 54 to 72-year-old monospecific stands in Rwanda were quantified from March 2006 to December 2007. The influences of soil temperature, soil water content, soil carbon (C) and nitrogen (N) stocks, soil pH, and stand characteristics on soil CO₂ flux were investigated. The mean annual soil CO₂ flux was highest under Eucalyptus saligna (3.92 μmol m⁻² s⁻¹) and lowest under Entandrophragma excelsum (3.13 μmol m⁻² s⁻¹). The seasonal variation in soil CO₂ flux from all stands followed the same trend and was highest in rainy seasons and lowest in dry seasons. Soil CO₂ flux was mainly correlated to soil water content (R² = 0.36-0.77), stand age (R² = 0.45), soil C stock (R² = 0.33), basal area (R² = 0.21), and soil temperature (R² = 0.06-0.17). The results contribute to the understanding of factors that influence soil CO₂ flux in monocultural plantations grown under the same microclimatic and soil conditions. The results can be used to construct models that predict soil CO₂ emissions in the tropics.     

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.     

Emissions of CO2, CH4 and N2O from Southern European peatlands

Peatlands play an important role in emissions of the greenhouse gases CO₂, CH₄ and N₂O, which are produced during mineralization of the peat organic matter. To examine the influence of soil type (fen, bog soil) and environmental factors (temperature, groundwater level), emission of CO₂, CH₄ and N₂O and soil temperature and groundwater level were measured weekly or biweekly in loco over a one-year period at four sites located in Ljubljana Marsh, Slovenia using the static chamber technique. The study involved two fen and two bog soils differing in organic carbon and nitrogen content, pH, bulk density, water holding capacity and groundwater level. The lowest CO₂ fluxes occurred during the winter, fluxes of N₂O were highest during summer and early spring (February, March) and fluxes of CH₄ were highest during autumn. The temporal variation in CO₂ fluxes could be explained by seasonal temperature variations, whereas CH₄ and N₂O fluxes could be correlated to groundwater level and soil carbon content. The experimental sites were net sources of measured greenhouse gases except for the drained bog site, which was a net sink of CH₄. The mean fluxes of CO₂ ranged between 139 mg m⁻² h⁻¹ in the undrained bog and 206 mg m⁻² h⁻¹ in the drained fen; mean fluxes of CH₄ were between −0.04 mg m⁻² h⁻¹ in the drained bog and 0.05 mg m⁻² h⁻¹ in the drained fen; and mean fluxes of N₂O were between 0.43 mg m⁻² h⁻¹ in the drained fen and 1.03 mg m⁻² h⁻¹ in the drained bog. These results indicate that the examined peatlands emit similar amounts of CO₂ and CH₄ to peatlands in Central and Northern Europe and significantly higher amounts of N₂O.     

Emission of N2O, N2 and CO2 from soil fertilized with nitrate: effect of compaction, soil moisture and rewetting

Soil compaction and soil moisture are important factors influencing denitrification and N₂O emission from fertilized soils. We analyzed the combined effects of these factors on the emission of N₂O, N₂ and CO₂ from undisturbed soil cores fertilized with (150 kg N ha⁻¹) in a laboratory experiment. The soil cores were collected from differently compacted areas in a potato field, i.e. the ridges (ρ_D=1.03 g cm⁻³), the interrow area (ρ_D=1.24 g cm⁻³), and the tractor compacted interrow area (ρ_D=1.64 g cm⁻³), and adjusted to constant soil moisture levels between 40 and 98% water-filled pore space (WFPS).High N₂O emissions were a result of denitrification and occurred at a WFPS≥70% in all compaction treatments. N₂ production occurred only at the highest soil moisture level (≥90% WFPS) but it was considerably smaller than the N₂O-N emission in most cases. There was no soil moisture effect on CO₂ emission from the differently compacted soils with the exception of the highest soil moisture level (98% WFPS) of the tractor-compacted soil in which soil respiration was significantly reduced. The maximum N₂O emission rates from all treatments occurred after rewetting of dry soil. This rewetting effect increased with the amount of water added. The results show the importance of increased carbon availability and associated respiratory O₂ consumption induced by soil drying and rewetting for the emissions of N₂O.     

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.     

Carbon exchange in a freshwater marsh in the Sanjiang Plain, northeastern China

Northern wetlands are critically important to global change because of their role in modulating atmospheric concentrations of greenhouse gases, especially CO₂ and CH₄. At present, continuous observations for CO₂ and CH₄ fluxes from northern wetlands in Asia are still very limited. In this paper, two growing season measurements for CO₂ flux by eddy covariance technique and CH₄ flux by static chamber technique were conducted in 2004 and 2005, at a permanently inundated marsh in the Sanjiang Plain, northeastern China. The seasonal variations of CO₂ exchange and CH₄ flux and the environmental controls on them were investigated. During the growing seasons, large variations in net ecosystem CO₂ exchange (NEE) and gross ecosystem productivity (GEP) were observed with the range of −4.0 to 2.2 (where negative exchange is a gain of carbon from the atmosphere) and 0-7.6 g C m⁻² d⁻¹, respectively. Ecosystem respiration (RE) displayed relatively smooth seasonal pattern with the range of 0.8-4.2 g C m⁻² d⁻¹. More than 70% of the total GEP was consumed by respiration, which resulted in a net CO₂ uptake of 143 ± 9.8 and 100 ± 9.2 g C m⁻² for the marsh over the growing seasons of 2004 and 2005, respectively. A significant portion of the accumulated NEE-C was lost by CH₄ emission during the growing seasons, indicating the great potential of CH₄ emission from the inundated marsh. Air temperature and leaf area index jointly affected the seasonal variation of GEP and the seasonal dynamic of RE was mainly controlled by soil temperature and leaf area index. Soil temperature also exerted the dominant influence over variation of CH₄ flux while no significant relationship was found between CH₄ emission and water table level. The close relationships between carbon fluxes and temperature can provide insights into the response of marsh carbon exchange to a changing climate. Future long term flux measurements over the freshwater marsh ecosystems are undoubtedly necessary.     

Fluxes of CO2, CH4, and N2O in an alpine meadow affected by yak excreta on the Qinghai-Tibetan plateau during summer grazing periods

To assess the impacts of yak excreta patches on greenhouse gas (GHG) fluxes in the alpine meadow of the Qinghai-Tibetan plateau, methane (CH₄), carbon dioxide (CO₂), and nitrous oxide (N₂O) fluxes were measured for the first time from experimental excreta patches placed on the meadow during the summer grazing seasons in 2005 and 2006. Dung patches were CH₄ sources (average 586 μg m⁻² h⁻¹ in 2005 and 199 μg m⁻² h⁻¹ in 2006) during the investigation period of two years, while urine patches (average −31 μg m⁻² h⁻¹ in 2005 and −33 μg m⁻² h⁻¹ in 2006) and control plots (average −28 μg m⁻² h⁻¹ in 2005 and −30 μg m⁻² h⁻¹ in 2006) consumed CH₄. The cumulative CO₂ emission for dung patches was about 36-50% higher than control plots during the experimental period in 2005 and 2006. The cumulative N₂O emissions for both urine and dung patches were 2.1-3.7 and 1.8-3.5 times greater than control plots in 2005 and 2006, respectively. Soil water-filled pore space (WFPS) explained 35% and 36% of CH₄ flux variation for urine patches and control plots, respectively. Soil temperature explained 40-75% of temporal variation of CO₂ emissions for all treatments. Temporal N₂O flux variation in urine patches (34%), dung patches (48%), and control (56%) plots was mainly driven by the simultaneous effect of soil temperature and WFPS. Although yak excreta patches significantly affected GHG fluxes, their contributions to the whole grazing alpine meadow in terms of CO₂ equivalents are limited under the moderate grazing intensity (1.45 yak ha⁻¹). However, the contributions of excreta patches to N₂O emissions are not negligible when estimating N₂O emissions in the grazing meadow. In this study, the N₂O emission factor of yak excreta patches varied with year (about 0.9-1.0%, and 0.1-0.2% in 2005 and 2006, respectively), which was lower than IPCC default value of 2%.     

Nitrogen fixation by biological soil crusts and heterotrophic bacteria in an intact Mojave Desert ecosystem with elevated CO2 and added soil carbon

Fixation of N by biological soil crusts and free-living heterotrophic soil microbes provides a significant proportion of ecosystem N in arid lands. To gain a better understanding of how elevated CO₂ may affect N₂-fixation in aridland ecosystems, we measured C₂H₂ reduction as a proxy for nitrogenase activity in biological soil crusts for 2 yr, and in soils either with or without dextrose-C additions for 1 yr, in an intact Mojave Desert ecosystem exposed to elevated CO₂. We also measured crust and soil δ¹⁵N and total N to assess changes in N sources, and δ¹³C of crusts to determine a functional shift in crust species, with elevated CO₂. The mean rate of C₂H₂ reduction by biological soil crusts was 76.9±5.6 μmol C₂H₄ m⁻² h⁻¹. There was no significant CO₂ effect, but crusts from plant interspaces showed high variability in nitrogenase activity with elevated CO₂. Additions of dextrose-C had a positive effect on rates of C₂H₂ reduction in soil. There was no elevated CO₂ effect on soil nitrogenase activity. Plant cover affected soil response to C addition, with the largest response in plant interspaces. The mean rate of C₂H₂ reduction in soils either with or without C additions were 8.5±3.6 μmol C₂H₄ m⁻² h⁻¹ and 4.8±2.1 μmol m⁻² h⁻¹, respectively. Crust and soil δ¹⁵N and δ¹³C values were not affected by CO₂ treatment, but did show an effect of cover type. Crust and soil samples in plant interspaces had the lowest values for both measurements. Analysis of soil and crust [N] and δ¹⁵N data with the Rayleigh distillation model suggests that any plant community changes with elevated CO₂ and concomitant changes in litter composition likely will overwhelm any physiological changes in N₂-fixation.     

CO2 emission in an intensively cultivated loam as affected by long-term application of organic manure and nitrogen fertilizer

A long-term field experiment was conducted to examine the influence of mineral fertilizer and organic manure on the equilibrium dynamics of soil organic C in an intensively cultivated fluvo-aquic soil in the Fengqiu State Key Agro-Ecological Experimental Station (Fengqiu county, Henan province, China) since September 1989. Soil CO₂ flux was measured during the maize and wheat growing seasons in 2002-2003 and 2004 to evaluate the response of soil respiration to additions and/or alterations in mineral fertilizer, organic manure and various environmental factors. The study included seven treatments: organic manure (OM), half-organic manure plus half-fertilizer N (NOM), fertilizer NPK (NPK), fertilizer NP (NP), fertilizer NK (NK), fertilizer PK (PK) and control (CK). Organic C in soil and the soil heavy fraction (organo-mineral complex) was increased from 4.47 to 8.61 mg C g⁻¹ and from 3.32 to 5.68 mg C g⁻¹, respectively, after the 13 yr application of organic manure. In contrast, organic C and the soil heavy fraction increased in NPK soil to only 5.41 and 4.38 mg C g⁻¹, respectively. In the CK treatment, these parameters actually decreased from the initial C concentrations (4.47 and 3.32 mg C g⁻¹) to 3.77 and 3.11 mg C g⁻¹, respectively. Therefore, organic manure efficiently elevated soil organic C. However, only 66% of the increased soil organic C was combined with clay minerals in the OM treatment. Cumulative soil CO₂ emissions from inter-row soil in the OM and NPK treatments were 228 and 188 g C m⁻² during the 2002 maize growing season, 132 and 123 g C m⁻² during the 2002/2003 wheat growing season, and 401 and 346 g C m⁻² yr⁻¹ in 2002-2003, respectively. However, during the 2004 maize growing season, cumulative soil CO₂ emissions were as high as 617 and 556 g C m⁻², respectively, due to the contribution of rhizosphere respiration. The addition of organic manure contributed to a 16% increase in soil CO₂ emission in 2002-2003 (compared to NPK), where only 27%, 36% and 24% of applied organic C was released as CO₂ during the 2002 and 2004 maize growing seasons and in 2002-2003, respectively. During the 2002/2003 wheat growing season, soil CO₂ flux was significantly affected by soil temperature below 20 °C, but by soil moisture (WFPS) during the 2004 maize growing season at soil temperatures above 18 °C. Optimum soil WFPS for soil CO₂ flux was approximately 70%. When WFPS was below 50%, it no longer had a significant impact on soil CO₂ flux during the 2002 maize growing season. This study indicates the application of organic manure composted with wheat straw may be a preferred strategy for increasing soil organic C and sequestering C in soil.     

Explaining temporal variation in soil CO2 efflux in a mature spruce forest in Southern Germany

An open dynamic chamber system was used to measure the soil CO₂ efflux intensively and continuously throughout a growing season in a mature spruce forest (Picea abies) in Southern Germany. The resulting data set contained a large amount of temporally highly resolved information on the variation in soil CO₂ efflux together with environmental variables. Based on this background, the dependencies of the soil CO₂ efflux rate on the controlling environmental factors were analysed in-depth. Of the abiotic factors, soil temperature alone explained 72% of the variation in the efflux rate, and including soil water content (SWC) as an additional variable increased the explained variance to about 83%. Between April and December, average rates ranged from 0.43 to 5.15 μmol CO₂ m⁻² s⁻¹ (in November and July, respectively) with diurnal variations of up to 50% throughout the experiment. The variability in wind speed above the forest floor influenced the CO₂ efflux rates for measuring locations with a litter layer of relatively low bulk density (and hence relatively high proportions of pore spaces). For the temporal integration of flux rates for time scales of hours to days, however, wind velocities were of no effect, reflecting the fact that wind forcing acts on the transport, but not the production of CO₂ in the soil. The variation in both the magnitude of the basal respiration rate and the temperature sensitivity throughout the growing season was only moderate (coefficient of variation of 15 and 25%, respectively). Soil water limitation of the CO₂ production in the soil could be best explained by a reduction in the temperature-insensitive basal respiration rate, with no discernible effect on the temperature sensitivity. Using a soil CO₂ efflux model with soil temperature and SWC as driving variables, it was possible to calculate the annual soil CO₂ efflux for four consecutive years for which meteorological data were available. These simulations indicate an average efflux sum of 560 g C m⁻² yr⁻¹ (SE=22 g C m⁻² yr⁻¹). An alternative model derived from the same data but using temperature alone as a driver over-estimated the annual flux sum by about 7% and showed less inter-annual variability. Given a likely shift in precipitation patterns alongside temperature changes under projected global change scenarios, these results demonstrate the necessity to include soil moisture in models that calculate the evolution of CO₂ from temperate forest soils.     

CO2 emissions from subtropical arable soils of China

CO₂ efflux plays a key role in carbon exchange between the biosphere and atmosphere, but our understanding of the mechanism controlling its temporal and spatial variations is limited. The purpose of this study is to determine annual soil CO₂ flux and assess its variations in arable subtropical soils of China in relation to soil temperature, moisture, rainfall, microbial biomass carbon (MBC) and dissolved organic carbon (DOC) using the closed chamber method. Soils were derived from three parent materials including granite (G), tertiary red sandstone (T) and quaternary red clay (Q). The experiment was conducted at the Ecological Station of Red Soil, The Chinese Academy of Sciences, in a subtropical region of China. The results showed that soil CO₂ flux had clear seasonal fluctuations with the maximum value in summer, the minimum in winter and intermediate in spring and autumn. Further, significant differences in soil CO₂ flux were found among the three red soils, generally in the order of G>T>Q. The average annual fluxes were estimated as 2.84, 2.13 and 1.41 kg CO₂ m⁻² year⁻¹ for red soils derived from G, T and Q, respectively. Soil temperature strongly affects the seasonal variability of soil CO₂ flux (85.0-88.5% of the variability), followed by DOC (55.8-84.4%) and rainfall (43.0-55.8%). The differences in soil CO₂ flux among the three red soils were partly explained by MBC (33.7-58.9% of the variability) and DOC (23.8-33.6%).     

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.     

Interannual and interseasonal soil CO2 efflux and VOC exchange rates in a Mediterranean holm oak forest in response to experimental drought

Climate models predict drier conditions in the next decades in the Mediterranean basin. Given the importance of soil CO₂ efflux in the global carbon balance and the important role of soil monoterpene and volatile organic compounds (VOCs) in soil ecology, we aimed to study the effects of the predicted drought on soil CO₂, monoterpenes and other VOC exchange rates and their seasonal and interannual variations. We decreased soil water availability in a Mediterranean holm oak forest soil by means of an experimental drought system performed since 1999 to the present. Measurements of soil gas exchange were carried out with IRGA, GC and PTR-MS techniques during two annual campaigns of contrasting precipitation. Soil respiration was twice higher the wet year than the dry year (2.27±0.26 and 1.05±0.15, respectively), and varied seasonally from 3.76±0.85 μmol m⁻² s⁻¹ in spring, to 0.13±0.01 μmol m⁻² s⁻¹ in summer. These results highlight the strong interannual and interseasonal variation in CO₂ efflux in Mediterranean ecosystems. The drought treatment produced a significant soil respiration reduction in drought plots in the wet sampling period. This reduction was even higher in wet springs (43% average reduction). These results show (1) that soil moisture is the main factor driving seasonal and interannual variations in soil respiration and (2) that the response of soil respiration to increased temperature is constrained by soil moisture. The results also show an additional control of soil CO₂ efflux by physiology and phenology of trees and animals. Soil monoterpene exchange rates ranged from −0.01 to 0.004 nmol m⁻² s⁻¹, thus the contribution of this Mediterranean holm oak forest soil to the total monoterpenes atmospheric budget seems to be very low. Responses of individual monoterpenes and VOCs to the drought treatment were different depending on the compound. This suggests that the effect of soil moisture reduction in the monoterpenes and VOC exchange rates seems to be dependent on monoterpene and VOC type. In general, soil monoterpene and other VOC exchange rates were not correlated with soil CO₂ efflux. In all cases, only a low proportion of variance was explained by the soil moisture changes, since almost all VOCs increased their emission rates in summer 2005, probably due to the effect of high soil temperature. Results indicate thus that physical and biological processes in soil are controlling soil VOC exchange but further research is needed on how these factors interact to produce the observed VOCs exchange responses.     

Fluxes of CO2, CH4 and N2O from drained organic soils in deciduous forests

We examined net greenhouse gas exchange at the soil surface in deciduous forests on soils with high organic contents. Fluxes of CO₂, CH₄ and N₂O were measured using dark static chambers for two consecutive years in three different forest types; (i) a drained and medium productivity site dominated by birch, (ii) a drained and highly productive site dominated by alder and (iii) an undrained and highly productive site dominated by alder. Although the drained sites had shallow mean groundwater tables (15 and 18 cm, respectively) their average annual rates of forest floor CO₂ release were almost twice as high compared to the undrained site (1.9±0.4 and 1.7±0.3, compared to 1.0±0.2 kg CO₂ m⁻² yr⁻¹). The average annual CH₄ emission was almost 10 times larger at the undrained site (7.6±3.1 compared to 0.9±0.5 g CH₄ m⁻² yr⁻¹ for the two drained sites). The average annual N₂O emissions at the undrained site (0.1±0.05 g N₂O m⁻² yr⁻¹) were lower than at the drained sites, and the emissions were almost five times higher at the drained alder site than at the drained birch site (0.9±0.35 compared to 0.2±0.11 g N₂O m⁻² yr⁻¹). The temporal variation in forest floor CO₂ release could be explained to a large extent by differences in groundwater table and air temperature, but little of the variation in the CH₄ and N₂O fluxes could be explained by these variables. The measured soil variables were only significant to explain for the within-site spatial variation in CH₄ and N₂O fluxes at the undrained swamp, and dark forest floor CO₂ release was not explained by these variables at any site. The between-site spatial variation was attributed to variations in drainage, groundwater level position, productivity and tree species for all three gases. The results indicate that N₂O emissions are of greater importance for the net greenhouse gas exchange at deciduous drained forest sites than at coniferous drained forest sites.     

Soil CO2 efflux in a temperate deciduous forest: Environmental drivers and component contributions

Soil CO₂ efflux is a large component of total respiration in many ecosystems. It is important to understand the environmental controls on soil CO₂ efflux, in order to evaluate potential responses of ecosystems to climate change. This study investigated the relationship between total soil CO₂ efflux and soil temperature, soil moisture and solar radiation on an interannual basis for a plot of temperate deciduous ancient semi-natural woodland at Wytham Woods in central southern England. We also aimed to quantify the contribution of soil organic matter decomposition (SOM), root-and-rhizosphere respiration, and mycorrhizal respiration components to total soil CO₂ efflux, and determine their environmental correlates. Total soil CO₂ efflux was measured regularly from April 2006 to December 2008 and found to average 4.1 Mg C ha⁻¹ yr⁻¹ in both 2007 and 2008. In addition, we applied a recently developed approach to partition the efflux into SOM, root-and-rhizosphere, and mycorrhizal components in situ using mesh bags. SOM decomposition, root-and-rhizosphere, and mycorrhizal respiration were estimated to contribute 70 ± 6%, 22 ± 6% and 8 ± 3% of total soil CO₂ efflux respectively, equating to 3.0 ± 0.3, 0.9 ± 0.2 and 0.3 ± 0.1 Mg C ha⁻¹ yr⁻¹. In order to avoid the effect of temporal correlation between variables caused by seasonality, we investigated interannual variability by examining the relationship between CO₂ flux anomalies and anomalies in environmental variables. Variation in soil temperature explained 50% of the interannual variance in soil CO₂ efflux, and soil moisture a further 18% of the residual variance. Solar radiation, as a proxy for plant photosynthesis, had no significant effect on total soil CO₂ efflux, but was positively correlated with root-and-rhizosphere respiration, and mycorrhizal respiration. The relationship between anomalies in soil CO₂ efflux and soil temperature was highly significant, with a sensitivity of 0.164 ± 0.023 μmol CO₂ m⁻² s⁻¹ °C⁻¹. For mean peak summer efflux rates (2.03 μmol CO₂ m² s⁻¹), this is equivalent to 8% per °C, or a Q₁₀ temperature sensitivity of 2.2 ± 0.2. We demonstrate the utility of an anomaly analysis approach and conclude that soil temperature is the key driver of total soil CO₂ efflux primarily through its positive relationship with SOM-decomposition rate.     

Below ground carbon turnover and greenhouse gas exchanges in a sub-arctic wetland

Here we present results from a field experiment in a sub-arctic wetland near Abisko, northern Sweden, where the permafrost is currently disintegrating with significant vegetation changes as a result. During one growing season we investigated the fluxes of CO₂ and CH₄ and how they were affected by ecosystem properties, i.e., composition of species that are currently expanding in the area (Carex rotundata, Eriophorum vaginatum and Eriophorum angustifolium), dissolved CH₄ in the pore water, substrate availability for methane producing bacteria, water table depth, active layer, temperature, etc. We found that the measured gas fluxes over the season ranged between: CH₄ 0.2 and 36.1 mg CH₄ m⁻² h⁻¹, Net Ecosystem Exchange (NEE) −1000 and 1250 mg CO₂ m⁻² h⁻¹ (negative values meaning a sink of atmospheric CO₂) and dark respiration 110 and 1700 mg CO₂ m⁻² h⁻¹. We found that NEE, photosynthetic rate and CH₄ emission were affected by the species composition. Multiple stepwise regressions indicated that the primary explanatory variables for NEE was photosynthetic rate and for respiration and photosynthesis biomass of green leaves. The primary explanatory variables for CH₄ emissions were depth of the water table, concentration of organic acid carbon and biomass of green leaves. The negative correlations between pore water concentration and emission of CH₄ and the concentrations of organic acid, amino acid and carbohydrate carbon indicated that these compounds or their fermentation by-products were substrates for CH₄ formation. Furthermore, calculation of the radiative forcing of the species expanding in the area as a direct result of permafrost degradation and a change in hydrology indicate that the studied mire may act as an increasing source of radiative forcing in future.     

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.     

Land use effects on gross nitrogen mineralization, nitrification, and N2O emissions in ephemeral wetlands

Stable ¹⁵N isotope dilution and tracer techniques were used in cultivated (C) and uncultivated (U) ephemeral wetlands in central Saskatchewan, Canada to: (1) quantify gross mineralization and nitrification rates and (2) estimate the relative proportion of N₂O emissions from these wetlands that could be attributed to denitrification versus nitrification-related processes. In-field incubation experiments were repeated in early May, mid-June and late July. Mean gross mineralization and nitrification rates (10.3 and 3.1 mg kg⁻¹ d⁻¹, respectively) did not differ between C and U wetlands on any given date. Despite these similarities, the mean NH₄⁺ pool size in the U wetlands (17.2 mg kg⁻¹) was two to three times that of the C wetlands (6.7 mg kg⁻¹) whereas the mean NO₃⁻ pool size in U wetlands (2.2 mg kg⁻¹) was less than half that of C wetlands (5.8 mg kg⁻¹). Mean N₂O emissions from the C wetlands decreased from 112.8 to 17.0 ng N₂O m² s⁻¹ from May to July, whereas mean U-wetland N₂O emissions ranged only from 31.8 to 51.1 ng N₂O m² s⁻¹ over the same period. This trend is correlated to water-filled pore space in C wetlands, demonstrating a soil moisture influence on emissions. Denitrification is generally considered the dominant emitter of N₂O under anaerobic conditions, but in the C wetlands, only 49% of the May emissions could be directly attributed to denitrification, decreasing to 29% in July. In contrast, more than 75% of the N₂O emissions from the U wetlands arose from denitrification of the soil NO₃⁻ pool throughout the season. These land use differences in emission sources and rates should be taken into consideration when planning management strategies for greenhouse gas mitigation.