Temperature dependence of soil CO2 efflux is strongly modulated by seasonal patterns of moisture availability in a Mediterranean ecosystem

Extensive research has focused on the temperature sensitivity of soil respiration. However, in Mediterranean ecosystems, soil respiration may have a pulsed response to precipitation events, especially during prolonged dry periods. Here, we investigate temporal variations in soil respiration (R_s), soil temperature (T) and soil water content (SWC) under three different land uses (a forest area, an abandoned agricultural field and a rainfed olive grove) in a dry Mediterranean area of southeast Spain, and evaluate the relative importance of soil temperature and water content as predictors of R_s. We hypothesize that soil moisture content, rather than soil temperature, becomes the major factor controlling CO₂ efflux rates in this Mediterranean ecosystem during the summer dry season. Soil CO₂ efflux was measured monthly between January 2006 and December 2007 using a portable soil respiration instrument fitted with a soil respiration chamber (LI-6400-09). Mean annual soil respiration rates were 2.06 ± 0.07, 1.71 ± 0.09, and 1.12 ± 0.12 μmol m⁻² s⁻¹ in the forest, abandoned field and olive grove, respectively. R_s was largely controlled by soil temperature above a soil water content threshold value of 10% at 0-15 cm depth for forest and olive grove, and 15% for abandoned field. However, below those thresholds R_s was controlled by soil moisture. Exponential and linear models adequately described R_s responses to environmental variables during the growing and dry seasons. Models combining abiotic (soil temperature and soil rewetting index) and biotic factors (above-ground biomass index and/or distance from the nearest tree) explained between 39 and 73% of the temporal variability of R_s in the forest and olive grove. However, in the abandoned field, a single variable - either soil temperature (growing season) or rewetting index (dry season) - was sufficient to explain between 51 and 63% of the soil CO₂ efflux. The fact that the rewetting index, rather than soil water content, became the major factor controlling soil CO₂ efflux rates during the prolonged summer drought emphasizes the need to quantify the effects of rain pulses in estimates of net annual carbon fluxes from soil in Mediterranean ecosystems.     

Subtropical plantations are large carbon sinks: Evidence from two monoculture plantations in South China

Quantifying the net carbon (C) storage of forest plantations is required to assess their potential to offset fossil fuel emissions. In this study, a biometric approach was used to estimate net ecosystem productivity (NEP) for two monoculture plantations in South China: Acacia crassicarpa and Eucalyptus urophylla. This approach was based on stand-level net primary productivity (NPP, based on direct biometric inventory) and heterotrophic respiration (R_h). In comparisons of R_h determination based on trenching vs. tree girdling, both trenching and tree girdling changed soil temperature and soil moisture relative to undisturbed control plots, and we assess the effects of corrections for disturbances of soil moisture and soil moisture on the estimation of soil CO₂ efflux partitioning. Soil microbial biomass and dissolved organic carbon were significantly lower in trenched plots than in tree girdled plots for both plantations. Annual soil CO₂ flux in trenched plots (R_h-t) was significantly lower than in tree-girdled plots (R_h-g) in both plantations. The estimates of R_h-t and R_h-g, expressed as a percentage of total soil respiration, were 58 ± 4% and 74 ± 6%, respectively, for A. crassicarpa, and 64 ± 3% and 78 ± 5%, respectively, for E. urophylla. By the end of experiment, the difference in soil CO₂ efflux between the trenched plots and tree-girdled plots had become small for both plantations. Annual R_h (mean of the annual R_h-t and R_h-g) and net primary production (NPP) were 470 ± 25 and 800 ± 118 g C m⁻² yr⁻¹, respectively, for A. crassicarpa, and 420 ± 35 and 2380 ± 187 g C m⁻² yr⁻², respectively, for E. urophylla. The two plantations in the developmental stage were large carbon sinks: NEP was 330 ± 76 C m⁻² yr⁻¹ for A. crassicarpa and 1960 ± 178 g C m⁻² yr⁻¹ for E. urophylla.     

Links between vegetation patterns, soil C and N pools and respiration rate under three different land uses in a dry Mediterranean ecosystem

Purpose

Soil respiration (R _s) is controlled by abiotic soil parameters interacting with characteristics of the vegetation and the soil microbial community. Few studies have attempted a comprehensive approach that simultaneously addresses the roles of all the major factors known to influence R _s. Our goal was to explore the links between heterogeneity in R _s, aboveground plant biomass and belowground properties in three representative land-use types in a dry Mediterranean ecosystem: (1) a 150-year-old mixed Aleppo pine-kermes oak open forest, (2) an abandoned agricultural field, which was cultivated with cereal for several years until abandonment in 1980, when establishment of typical Mediterranean shrubland vegetation started and (3) a rain-fed olive grove, which has been cultivated for 100 years.

Materials and methods

We selected two distinctive sampling periods coinciding with annual minimum or near minimum (December) and maximum (April) rates of R _s in this dry Mediterranean ecosystem. In each sampling period, R _s, temperature and moisture, aboveground plant biomass, carbon (C) and nitrogen (N) contents in both light and heavy soil organic matter fractions, extractable dissolved organic C (EDOC), as well as microbial and fine root biomass were measured within each land-use type.

Results and discussion

Across sites, R _s rates were significantly higher in April (3.07?±?0.1 μmol?m^?2?s^?1) than in December (1.30?±?0.1 μmol?m^?2?s^?1). The labile soil organic matter fractions (light fraction C and N contents, microbial biomass C and EDOC) were consistently and strongly related to one another, and to a lesser extent, to the C and N contents in the heavy fraction across sites and seasons. Linear models adequately explained a large proportion of the within-site variability in R _s (R ² values ranged from 41 to 91 % depending on land use and season) but major controls on R _s differed considerably between sites and seasons. Primary controls on spatial patterns in R _s were linked to recent plant-derived C inputs in both forest and olive grove sites. However, in the abandoned agricultural field site R _s appeared to be mainly driven by microbial activity, which could be sustained by intermediate or recalcitrant C and N pools derived from previous land use.

Conclusions

Conversion of native woodland to agricultural land and subsequent land abandonment leads to profound changes in the relationships between R _s, aboveground biomass and belowground properties in this dry Mediterranean ecosystem. While above- and belowground vegetation are the primary controls on spatial variability in labile soil C pools and R _s in the open forest and olive grove sites, a complete lack of influence of current vegetation patterns on soil C pools and respiration rates in the abandoned agricultural field was observed.     

Integrated diurnal soil respiration model during growing season of a typical temperate steppe: Effects of temperature, soil water content and biomass production

Soil respiration was measured with the enclosed chamber method in an ungrazed Leymus chinensis steppe during the growing seasons of 2001 and 2002. Soil respiration rate (R_S) was significantly influenced by air temperature (T) at the diurnal scale, and could be described by Van't Hoff's equation (R_S = R₁₀ exp(β(T − 10))). At the seasonal scale, the normalized soil respiration rate at 10 °C (R₁₀) was mainly controlled by soil water content (R² = 0.717, P < 0.001), while the sensitivity of soil respiration to temperature (Q₁₀) was partially affected by absolute growth rate (R² = 0.482, P = 0.004). Thus, soil respiration could be described as R_S = (20.015W − 84.085) (0.103AGR + 1.786)^(T−10)/10 during the growing seasons, integrating soil water content (W) and absolute growth rate (AGR) into the temperature-dependent soil respiration equation. It was validated by the observed soil respiration rates in this study (R² = 0.890, P < 0.001) and observations from near-field experiment (R² = 0.687, P = 0.011). It implied that accurately evaluating annual soil respiration should include the effects of plant biomass production and other abiotic factors besides air temperature.     

Spatial and seasonal variations in soil respiration in a temperate deciduous forest with fluctuating water table

Our objectives were to determine both spatial and temporal variations in soil respiration of a mixed deciduous forest, with soils exhibiting contrasting levels of hydromorphy. Soil respiration (R_S) showed a clear seasonal trend that reflected those of soil temperature (T_S) and soil water content (W_S), especially during summer drought. Using a bivariate model (RMSE=1.03), both optimal soil water content for soil respiration (W_SO) and soil respiration at both 10 °C and optimal soil water content (R_S10) varied among plots, ranging, respectively, from 0.25 to 0.40 and from 2.30 to 3.60 μmol m⁻² s⁻¹. Spatial variation in W_SO was related to bulk density and to topsoil N content, while spatial variation in R_S10 was related to basal area and the difference in pH measured in water or KCl suspensions. These results offer promising perspectives for spatializing ecosystem carbon budget at the regional scale.     

Competition between invasive earthworms (Amynthas corticis, Megascolecidae) and native North American millipedes (Pseudopolydesmus erasus, Polydesmidae): Effects on carbon cycling and soil structure

Invasive earthworms can have significant impacts on C dynamics through their feeding, burrowing, and casting activities, including the protection of C in microaggregates and alteration of soil respiration. European earthworm invasion is known to affect soil micro- and mesofauna, but little is known about impacts of invasive earthworms on other soil macrofauna. Asian earthworms (Amynthas spp.) are increasingly being reported in the southern Appalachian Mountains in southeastern North America. This region is home to a diverse assemblage of native millipedes, many of which share niches with earthworm species. This situation indicates potential for earthworm-millipede competition in areas subject to Amynthas invasion.In a laboratory microcosm experiment, we used two ¹³C enriched food sources (red oak, Quercus rubra, and eastern hemlock, Tsuga canadensis) to assess food preferences of millipedes (Pseudopolydesmus erasus), to determine the effects of millipedes and earthworms (Amynthas corticis) on soil structure, and to ascertain the nature and extent of the interactions between earthworms and millipedes. Millipedes consumed both litter species and preferred red oak litter over eastern hemlock litter. Mortality and growth of millipedes were not affected by earthworm presence during the course of the experiment, but millipedes assimilated much less litter-derived C when earthworms were present.Fauna and litter treatments had significant effects on soil respiration. Millipedes alone reduced CO₂ efflux from microcosms relative to no fauna controls, whereas earthworms alone and together with millipedes increased respiration, relative to the no fauna treatment. CO₂ derived from fresh litter was repressed by the presence of macrofauna. The presence of red oak litter increased CO₂ efflux considerably, compared to hemlock litter treatments.Millipedes, earthworms, and both together reduced particulate organic matter. Additionally, earthworms created significant shifts in soil aggregates from the 2000-250 and 250-53 μm fractions to the >2000 μm size class. Earthworm-induced soil aggregation was lessened in the 0-2 cm layer in the presence of millipedes. Earthworms translocated litter-derived C to soil throughout the microcosm.Our results suggest that invasion of ecosystems by A. corticis in the southern Appalachian Mountains is unlikely to be limited by litter species and these earthworms are likely to compete directly for food resources with native millipedes. Widespread invasion could cause a net loss of C due to increased respiration rates, but this may be offset by C protected in water-stable soil aggregates.     

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.     

Temperature sensitivity of soil respiration in different ecosystems in China

Understanding the sensitivity of soil respiration to temperature change and its impacting factors is an important base for accurately evaluating the response of terrestrial carbon balance to future climatic change, and thus has received much recent attention. In this study, we synthesized 161 field measurement data from 52 published papers to quantify temperature sensitivity of soil respiration in different Chinese ecosystems and its relationship with climate factors, such as temperature and precipitation. The results show that the observed Q₁₀ value (the factor by which respiration rates increase for a 10 °C increase in temperature) is strongly dependent on the soil temperature measurement depth. Generally, Q₁₀ significantly increased with the depth (0 cm, 5 cm, and 10 cm) of soil temperature measuring point. Different ecosystem types also exhibit different Q₁₀ values. In response to soil temperature at the depth of 5 cm, alpine meadow and tundra has the largest Q₁₀ value with magnitude of 3.05 ± 1.06, while the Q₁₀ value of evergreen broadleaf forests is approximately half that amount (Q₁₀ = 1.81 ± 0.43). Spatial correlation analysis also shows that the Q₁₀ value of forest ecosystems is significantly and negatively correlated with mean annual temperature (R = −0.51, P < 0.001) and mean annual precipitation (R = −0.5, P < 0.001). This result not only implies that the temperature sensitivity of soil respiration will decline under continued global warming, but also suggests that such acclimation of soil respiration to warming should be taken into account in forecasting future terrestrial carbon cycle and its feedback to climate system.     

Contribution of crop residue carbon to soil respiration at a northern Prairie site using stable isotope flux measurements

Heterotrophic respiration from agricultural soils can be differentiated as originating from microbial decomposition of recent litter inputs or crop residue carbon (CRC) and resident soil organic carbon (SOC) pools of varying age and stages of decomposition. Our objective was to determine the relative contributions of these pools to respiration in a northern agroecosystem where the non-growing season is long. A tunable diode laser trace gas analyzer was used to determine atmospheric stable C isotope ratio (δ¹³C) values and ¹²CO₂ and ¹³CO₂ fluxes over an agricultural field in the Red River Valley of southern Manitoba, Canada. Measurement campaigns were conducted in the fall of 2006 and spring of 2007 following harvest of a maize (C₄) crop from soil having SOC derived from previous C₃ crops. Stable CO₂ isotopologue gradients were measured from the center of four 200 × 200 m experimental plots, and fluxes were calculated using the aerodynamic flux gradient method. The soil in two of the experimental plots underwent intensive tillage, while the other two plots were managed using a form of reduced tillage. Approximately 70% and 20-30% of the total respiration flux originated from the maize C₄-CRC during the fall of 2006 and spring of 2007, respectively. At least 25% of the maize residue was lost to respiration during this non-growing period. No difference in the partitioning of heterotrophic respiration into that derived from CRC and SOC was detected between the intensive tillage and recently established reduced tillage treatments at the site.     

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.