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.     

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.     

Optimum temperatures for soil respiration along a semi-arid elevation gradient in southern California

Soil respiration (R) has not been adequately studied at temperatures above 35 °C, which are common temperatures for soils in the southwestern United States and may be important for C dynamics in semi-arid regions. While frequently excluded from ecosystem models or set to 35 °C, the optimum temperature for soil R is poorly understood. Optimum temperatures are likely controlled by substrate availability, soil moisture content, and previous climate. To quantify the optimal temperature for soil R and hypothesized relationships, we collected soils from beneath and between plant canopies at three sites along a semi-arid elevation gradient. Processed soil samples were incubated at three soil moisture contents and soil R was measured at 6 temperatures, successively (25–55 °C). From these data, an activation energy for reaction kinetics and deactivation energy for enzyme functionality model was used to generate soil R curves from which two parameters were derived: R_max, the maximum rate of soil R and T_opt, the optimum temperature for soil R. R_max was significantly greater for soils at the highest elevation and at medium soil moisture content. T_opt was greater than 35 °C at all locations. In addition, T_opt was significantly greater for soils with greater amounts of SOM but not significantly different along the elevation gradient or at different moisture contents. These results support inclusion of much higher optimum temperatures than currently used in many ecosystem and land-surface models and provide support for explaining variation in T_opt as regulated by substrate quantity within a site and general insensitivity across climate differences.     

Responses of soil respiration and its components to drought stress

Purpose

Climate change is likely to increase both intensity and frequency of drought stress. The responses of soil respiration (R _s) and its components (root respiration, R _r; mycorrhizal respiration, R _m; and heterotrophic respiration, R _h) to drought stress can be different. This work aims to review the recent and current literature about the variations in R _s during the period of drought stress, to explore potential coupling processes and mechanisms between R _s and driving factors in the context of global climate change.

Results and discussion

The sensitivity of soil respiration and its components to drought stress depended on the ecosystems and seasonality. Drought stress depressed R _s in mesic and xeric ecosystems, while it stimulated R _s in hydric ecosystems. The reductions in supply and availability of substrate decreased both auto- and heterotrophic respirations, leading to the temporal decoupling of root and mycorrhizal respiration from canopy photosynthesis as well as C allocation. Drought stress also reduced the diffusion of soluble C substrate, and activities of extracellular enzymes, consequently, limited microbial activity and reduced soil organic matter decomposition. Drought stress altered Q ₁₀ values and broke the coupling between temperature and soil respiration. Under drought stress conditions, R _m is generally less sensitive to temperature than R _r and R _h. Elevated CO₂ concentration alleviated the negative effect of drought stress on soil respiration, principally due to the promotion of plant C assimilation subsequently, which increased substrate supply for respiration in both roots and soil microorganisms. Additionally, rewetting stimulated soil respiration dramatically in most cases, except for soil that experienced extreme drought stress periods. The legacy of drought stress can also regulate the response of soil respiration rate to rewetting events in terrestrial ecosystems through changing abiotic drivers and microbial community structure.

Conclusions and perspectives

There is increasing evidence that drought stress can result in the decoupling of the above- and belowground processes, which are associated with soil respiration. However, studies on the variation in rates of soil respiration and its components under different intensities and frequencies of drought stress over the ecosystems should be reinforced. Meanwhile, molecular phylogenetics and functional genomics should be applied to link microbial ecology to the process of R _s. In addition, we should quantify the relationship between soil respiration and global change parameters (such as warming and elevated [CO₂]) under drought stress. Models simulating the rates of soil respiration and its components under global climate change and drought stress should also be developed.     

Study of soil respiration and fruit quality of table grape (Vitis vinifera L.) in response to different soil water content in a greenhouse

ABSTRACT

To understand the response of grape (Hutai No.8) quality and soil respiration (R_s) to different soil relative water contents (SRWCs), this study was designed with three soil moisture levels (A: 80–95%, B: 60–75%, and C: 40–55% of SRWC) for grape cultivation. Meanwhile, environmental factors, including air temperature (T_a), air relative humidity, and light intensity, were also recorded. The results showed the following: (1) Through the comprehensive analysis of fruit quality by the method of subordinate function, we concluded that the optimum soil moisture treatment was 60–75% SRWC, and the soluble sugars, proanthocyanidin, and resveratrol were most abundant. In addition, vitamin C (Vc) content was the largest under C treatment. (2) Photosynthetic characteristic under high soil moisture was better than those under low soil moisture condition during grape coloring periods, and it was largest under A treatment in 2015. R_s rate was in accordance with the trend of grape photosynthesis. High soil moisture could accelerate the photosynthetic rate of grape leaves and increase R_s. (3) Correlation analysis showed that higher soil moisture and air humidity and lower soil temperature (T_s) and T_a could promote the accumulation of more nutrients in grape berries; it also could increase photosynthetic rate and R_s during grape coloring periods. In conclusion, 60–75% SRWC was the optimum soil moisture condition, which could improve the nutrient contents and accumulate more bioactive substances. Of course, keeping a lower T_s and T_a, as well as higher air humidity, was also necessary.

Abbreviations: SRWC: soil relative water content; A, 90-95% SRWC; B, 70-75% SRWC; C, 40-55% SRWC; Rs: soil respiration; Ta: air temperature; Ts: soil temperature; OPC: proanthocyanidin; TSS: total soluble solids.     

Temperature sensitivity of soil respiration is affected by prevailing climatic conditions and soil organic carbon content: A trans-China based case study

Understanding the spatial variation of temperature sensitivity (i.e. Q₁₀) of soil respiration (R_s) and its controlling factors, is critical to improve the precision of carbon budget estimations at regional scales. In this study, data from 2-3 continuous years of R_s measurements over 15 ecosystems of ChinaFLUX were summarized to analyze the response of R_s to soil temperature. Moreover, we improved our dataset by collecting previously published Q₁₀ values from 34 ecosystems in China. The ecosystems studied were located in the main climatic zones of China, spanning from alpine via temperate to tropical. Spatial variations of Q₁₀ and its controlling factors were analyzed. The results showed that soil temperature at a 5 cm depth satisfactorily explained the seasonal variations in R_s of the 15 ChinaFLUX ecosystems (R² varying from 0.37 to 0.83). Based on the overall data, the Q₁₀ values of R_s in China ranged from 1.28 to 4.75. The spatial variations in Q₁₀ were primarily determined by soil temperature during measurement periods, soil organic carbon (SOC) content, and ecosystem type. Ecosystems in colder regions and with higher SOC content had relatively higher Q₁₀ values. Moreover, ecosystems of different vegetation types showed different Q₁₀ values. A temperature- and SOC-dependent function for Q₁₀ is suggested, which could be a valuable reference for improving the regional-scale models of R_s and ecosystem carbon cycles.     

Influence of temperature and drought on seasonal and interannual variations of soil, bole and ecosystem respiration in a boreal aspen stand

Continuous half-hourly measurements of soil (R_s) and bole respiration (R_b), as well as whole-ecosystem CO₂ exchange, were made with a non steady-state automated chamber system and with the eddy covariance (EC) technique, respectively, in a mature trembling aspen stand between January 2001 and December 2003. Our main objective was to investigate the influence of long-term variations of environmental and biological variables on component-specific and whole-ecosystem respiration (R_e) processes. During the study period, the stand was exposed to severe drought conditions that affected much of the western plains of North America. Over the 3 years, daily mean R_s varied from a minimum of 0.1 μmol m⁻² s⁻¹ during winter to a maximum of 9.2 μmol m⁻² s⁻¹ in mid-summer. Seasonal variations of R_s were highly correlated with variations of soil temperature (T_s) and water content (θ) in the surface soil layers. Both variables explained 96, 95 and 90% of the variance in daily mean R_s from 2001 to 2003. Aspen daily mean R_b varied from negligible during winter to a maximum of 2.5 μmol m⁻² bark s⁻¹ (2.2 μmol m⁻² ground s⁻¹) during the growing season. Maximum R_b occurred at the end of the aspen radial growth increment and leaf emergence period during each year. This was 2 months before the peak in bole temperature (T_b) in 2001 and 2003. Nonetheless, R_b was highly correlated with T_b and this variable explained 77, 87 and 62% of the variance in R_b in the respective years. Partitioning of R_b between its maintenance (R_bm) and growth (R_bg) components using the mature tissue method showed that daily mean R_bg occurred at the same time as aspen radial growth increment during each growing season. This method led, however, to systematic over- and underestimations of R_bm and R_bg, respectively, during each year. Annual totals of R_s, R_b and estimated foliage respiration (R_f) from hazelnut and aspen trees were, on average, 829, 159 and 202 g C m⁻² year⁻¹, respectively, over the 3 years. These totals corresponded to 70, 14 and 16%, respectively, of scaled-up respiration estimates of R_e from chamber measurements. Scaled R_e estimates were 25% higher (1190 g C m⁻² year⁻¹) than the annual totals of R_e obtained from EC (949 g C m⁻² year⁻¹). The independent effects of temperature and drought on annual totals of R_e and its components were difficult to separate because the two variables co-varied during the 3 years. However, recalculation of annual totals of R_s to remove the limitations imposed by low θ, suggests that drought played a more important role than temperature in explaining interannual variations of R_s and R_e.     

Fire interacts with season to influence soil respiration in tropical savannas

Soil respiration (R_s) is the second-largest source of CO₂ to the atmosphere in terrestrial systems. In tropical savannas seasonal moisture availability and frequent fires drive ecosystem dynamics and may have a considerable impact on soil carbon (C) cycling, including R_s. In order to test the effect of fire on soil C cycling we measured R_s in annually burnt and unburnt plots in wet and dry seasons at a long-term fire experiment established in savanna woodlands of northern Australia. There was a significant interaction between season and fire, with highest rates of daily R_s (722 mmol CO₂ m⁻² d⁻¹) observed in the wet season on unburnt, leaf litter patches. The three fold higher R_s rate on unburnt plots in the wet season was due to greater root-derived respiration (R_root: 356 mmol CO₂ m⁻² d⁻¹), while smaller changes to soil-derived respiration (R_soil: 51 mmol CO₂ m⁻² d⁻¹) were simply the result of C moving through decomposition rather than combustion pathways. Relationships between instantaneous R_s and soil temperature showed hysteresis with variable direction, suggesting that season and fire treatment also influence the soil depth at which CO₂ is produced. We suggest that (1) changes to fire regimes, through active management or climate change, in tropical savannas could have an impact on R_s, and (2) the direct effect of fire on soil C cycling is limited to the removal of aboveground litter inputs.     

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.     

Three scales of temporal resolution from automated soil respiration measurements

Soil respiration (R_s) is a combination of autotrophic and heterotrophic respiration, but it is often modeled as a single efflux process, influenced by environmental variables similarly across all time scales. Continued progress in understanding sources of variation in soil CO₂ efflux will require development of R_s models that incorporate environmental influences at multiple time scales. Coherence analysis, which requires high temporal frequency data on R_s and related environmental variables, permits examination of covariation between R_s and the factors that influence it at varying temporal frequencies, thus isolating the factors important at each time scale. Automated R_s measurements, along with air, soil temperature and moisture were collected at half hour intervals at a temperate forest at Harvard Forest, MA in 2003 and a boreal transition forest at the Howland Forest, ME in 2005. As in other temperate and boreal forests, seasonal variation in R_s was strongly correlated with soil temperature. The organic and mineral layer water contents were significantly related to R_s at synoptic time scales of 2–3 days to weeks, representing the wetting and drying of the soils as weather patterns move across the region. Post-wetting pulses of R_s were correlated with the amount of precipitation and the magnitude of the change from pre-wet-up moisture content to peak moisture content of the organic horizon during the precipitation events. Although soil temperature at 8–10 cm depth and R_s showed strong coherence at a 24-h interval, calculated diel Q₁₀ values for R_s were unreasonably high (6–74) during all months for the evergreen forest and during the growing season for the deciduous forest, suggesting that other factors that covary with soil temperature, such as canopy assimilatory processes, may also influence the diel amplitude of R_s. Lower diel Q₁₀ values were obtained based on soil temperature measured at shallower depths or with air temperature, but the fit was poorer and a lag was needed to improve the fit (peak R_s followed peak air temperature by several hours), suggesting a role for delayed substrate supply from aboveground processes to affect diel patterns of R_s. High frequency automated R_s datasets afford the opportunity to disentangle the temporal scales at which environmental factors, such as seasonal temperature and phenology, synoptic weather events and soil moisture, and diel variation in temperature and photosynthesis, affect soil respiration processes.     

Temporal and spatial variability of soil respiration in a boreal mixedwood forest

Temporal and spatial variability of soil respiration (R_s) was measured and analyzed in a 74-year-old, mixedwood, boreal forest in Ontario, Canada, over a period of 2 years (August 2003–July 2005). The ranges of R_s measured during the two study years were 0.5–6.9 μmol CO₂ m⁻² s⁻¹ for 2003–2004 (Year 1) and 0.4–6.8 μmol CO₂ m⁻² s⁻¹ for 2004–2005 (Year 2). Mean annual R_s for the stand was the same for both years, 2.7 μmol CO₂ m⁻² s⁻¹. Temporal variability of R_s was controlled mainly by soil temperature (T_s), but soil moisture had a confounding effect on T_s. Annual estimates of total soil CO₂ emissions at the site, calculated using a simple empirical R_s–T_s relationship, showed that R_s can account for about 88 ± 27% of total annual ecosystem respiration at the site. The majority of soil CO₂ emissions came from the upper 12 to 20 cm organic LFH (litter–fibric–humic) soil layer. The degree of spatial variability in R_s, along the measured transect, was seasonal and followed the seasonal trend of mean R_s: increasing through the growing season and converging to a minimum in winter (coefficient of variation (CV) ranged from 4 to 74% in Year 1 and 4 to 62% in Year 2). Spatial variability in R_s was found to be negatively related to spatial variability in the C:N ratio of the LHF layer at the site. Spatial variability in R_s was also found to depend on forest tree species composition within the stand. R_s was about 15% higher in a broadleaf deciduous tree patch compared to evergreen coniferous area. However, the difference was not always significant (at 95% CI). In general, R_s in the mixedwood patch, having both deciduous and coniferous species, was dominated by broadleaf trees, reflecting changing physiological controls on R_s with seasons. Our results highlight the importance of discerning soil CO₂ emissions at a variety of spatial and temporal scales. They also suggest including the LFH soil layer and allowing for seasonal variability in CO₂ production within that layer, when modeling soil respiration in forest ecosystems.     

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.     

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.     

Winter soil CO2 efflux and its contribution to annual soil respiration in different ecosystems of a forest-steppe ecotone, north China

Most soil respiration measurements are conducted during the growing season. In tundra and boreal forest ecosystems, cumulative winter soil CO₂ fluxes are reported to be a significant component of their annual carbon budgets. However, little information on winter soil CO₂ efflux is known from mid-latitude ecosystems. Therefore, comparing measurements of soil respiration taken annually versus during the growing season will improve the accuracy of ecosystem carbon budgets and the response of soil CO₂ efflux to climate changes. In this study we measured winter soil CO₂ efflux and its contribution to annual soil respiration for seven ecosystems (three forests: Pinus sylvestris var. mongolica plantation, Larix principis-rupprechtii plantation and Betula platyphylla forest; two shrubs: Rosa bella and Malus baccata; and two meadow grasslands) in a forest-steppe ecotone, north China. Overall mean winter and growing season soil CO₂ effluxes were 0.15-0.26 μmol m⁻² s⁻¹ and 2.65-4.61 μmol m⁻² s⁻¹, respectively, with significant differences in the growing season among the different ecosystems. Annual Q₁₀ (increased soil respiration rate per 10 °C increase in temperature) was generally higher than the growing season Q₁₀. Soil water content accounted for 84% of the variations in growing season Q₁₀ and soil temperature range explained 88% of the variation in annual Q₁₀. Soil organic carbon density to 30 cm depth was a good surrogate for SR₁₀ (basal soil respiration at a reference temperature of 10 °C). Annual soil CO₂ efflux ranged from 394.76 g C m⁻² to 973.18 g C m⁻² using observed ecosystem-specific response equations between soil respiration and soil temperature. Estimates ranged from 424.90 g C m⁻² to 784.73 g C m⁻² by interpolating measured soil respiration between sampling dates for every day of the year and then computing the sum to obtain the annual value. The contributions of winter soil CO₂ efflux to annual soil respiration were 3.48-7.30% and 4.92-7.83% using interpolated and modeled methods, respectively. Our results indicate that in mid-latitude ecosystems, soil CO₂ efflux continues throughout the winter and winter soil respiration is an important component of annual CO₂ efflux.     

Modeling soil respiration based on carbon, nitrogen, and root mass across diverse Great Lake forests

The variability in the net ecosystem exchange of carbon (NEE) is a major source of uncertainty in quantifying global carbon budget and atmospheric CO₂. Soil respiration, which is a large component of NEE, could be strongly influential to NEE variability. Vegetation type, landscape position, and site history can influence soil properties and therefore drive the microbial and root production of soil CO₂. This study measured soil respiration and soil chemical, biological and physical properties on various types of temperate forest stands in Northern Wisconsin (USA), which included ash elm, aspen, northern hardwood, red pine forest types, clear-cuts, and wetland edges. Soil respiration at each of the 19 locations was measured six times during 1 year from early June to mid-November. These data were combined with two additional data sets from the same landscape that represent two smaller spatial scales. Large spatial variation of soil respiration occurred within and among each forest type, which appeared to be from differences in soil moisture, root mass and the ratio of soil carbon to soil nitrogen (C:N). A soil climate driven model was developed that contained quadratic functions for root mass and the ratio of soil carbon to soil nitrogen. The data from the large range of forest types and site conditions indicated that the range of root mass and C:N on the landscape was also large, and that trends between C:N, root mass, and soil respiration were not linear as previously reported, but rather curvilinear. It should be noted this function appeared to level off and decline at C:N larger than 25, approximately the value where microbial nitrogen immobilization limits free soil nitrogen. Weak but significant relationships between soil water and soil C:N, and between soil C:N and root mass were observed indicating an interrelatedness of (1) topographically induced hydrologic patterns and soil chemistry, and (2) soil chemistry and root production. Future models of soil respiration should address multiple spatial and temporal factors as well as their co-dependence.     

Fluxes of CO2 above a plantation of Eucalyptus in southeast Brazil

Eddy-covariance measurements of net ecosystem exchange of CO₂ (NEE) and estimates of gross ecosystem productivity (GEP) and ecosystem respiration (R_E) were obtained in a 2-4 year old Eucalyptus plantation during two years with very different winter rainfall. In the first (drier) year the annual NEE, GEP and R_E were lower than the sums in the second (normal) year, and conversely the total respiratory costs of assimilated carbon were higher in the dry year than in the normal year.Although the net primary production (NPP) in the first year was 23% lower than that of the second year, the decrease in the carbon use efficiency (CUE = NPP/GEP) was 11% and autotrophic respiration utilized more resources in the first, dry year than in the second, normal year. The time variations in NEE were followed by NPP, because in these young Eucalyptus plantations NEE is very largely dominated by NPP, and heterotrophic respiration plays only a relatively minor role.During the dry season a pronounced hysteresis was observed in the relationship between NEE and photosynthetically active radiation, and NEE fluxes were inversely proportional to humidity saturation deficit values greater than 0.8 kPa. Nighttime fluxes of CO₂ during calm conditions when the friction velocity (u_*) was below the threshold (0.25 m s⁻¹) were estimated based on a Q₁₀ temperature-dependence relationship adjusted separately for different classes of soil moisture content, which regulated the temperature sensitivity of ecosystem respiration.     

On the 'temperature sensitivity' of soil respiration: Can we use the immeasurable to predict the unknown?

The temperature dependence of soil respiration (R_S) is widely used as a key characteristic of soils or organic matter fractions within soils, and in the context of global climatic change is often applied to infer likely responses of R_S to warmer future conditions. However, the way in which these temperature dependencies are calculated, interpreted and implemented in ecosystem models requires careful consideration of possible artefacts and assumptions. We argue that more conceptual clarity in the reported relationships is needed to obtain meaningful meta-analyses and better constrained parameters informing ecosystem models. Our critical assessment of common methodologies shows that it is impossible to measure actual temperature response of R_S, and that a range of confounding effects creates the observed apparent temperature relations reported in the literature. Thus, any measureable temperature response function will likely fail to predict effects of climate change on R_s. For improving our understanding of R_S in changing environments we need a better integration of the relationships between substrate supply and the soil biota, and of their long-term responses to changes in abiotic soil conditions. This is best achieved by experiments combining isotopic techniques and ecosystem manipulations, which allow a disentangling of abiotic and biotic factors underlying the temperature response of soil CO₂ efflux.     

Interpreting the dependence of soil respiration on soil temperature and water content in a boreal aspen stand

Continuous half-hourly measurements of soil CO₂ efflux made between January and December 2001 in a mature trembling aspen stand located at the southern edge of the boreal forest in Canada were used to investigate the seasonal and diurnal dependence of soil respiration (R_s) on soil temperature (T_s) and water content (θ). Daily mean R_s varied from a minimum of 0.1 μmol m⁻² s⁻¹ in February to a maximum of 9.2 μmol m⁻² s⁻¹ in mid-July. Daily mean T_s at the 2-cm depth was the primary variable accounting for the temporal variation of R_s and no differences between Arrhenius and Q₁₀ response functions were found to describe the seasonal relationship. R_s at 10 °C (R_s10) and the temperature sensitivity of R_s (Q_10Rs) calculated at the seasonal time scale were 3.8 μmol m⁻² s⁻¹ and 3.8, respectively. Temperature normalization of daily mean R_s (R_sN) revealed that θ in the 0–15 cm soil layer was the secondary variable accounting for the temporal variation of R_s during the growing season. Daily R_sN showed two distinctive phases with respect to soil water field capacity in the 0–15 cm layer (θ_fc, 0.30 m³ m⁻³): (1) R_sN was strongly reduced when θ decreased below θ_fc, which reflected a reduction in microbial decomposition, and (2) R_sN slightly decreased when θ increased above θ_fc, which reflected a restriction of CO₂ or O₂ transport in the soil profile.Diurnal variations of half-hourly R_s were usually out of phase with T_s at the 2-cm depth, which resulted in strong diurnal hysteresis between the two variables. Daily nighttime R_s10 and Q_10Rs parameters calculated from half-hourly nighttime measurements of R_s and T_s at the 2-cm depth (when there was steady cooling of the soil) varied greatly during the growing season and ranged from 6.8 to 1.6 μmol m⁻² s⁻¹ and 5.5 to 1.3, respectively. On average, daily nighttime R_s10 (4.5 μmol m⁻² s⁻¹) and Q_10Rs (2.8) were higher and lower, respectively, than the values obtained from the seasonal relationship. Seasonal variations of these daily parameters were highly correlated with variations of θ in the 0–15 cm soil layer, with a tendency of low R_s10 and Q_10Rs values at low θ. Overall, the use of seasonal R_s10 and Q_10Rs parameters led to an overestimation of daily ranges of half-hourly R_s (ΔR_s) during drought conditions, which supported findings that the short-term temperature sensitivity of R_s was lower during periods of low θ. The use of daily nighttime R_s10 and Q_10Rs parameters greatly helped at simulating ΔR_s during these periods but did not improve the estimation of half-hourly R_s throughout the year as it could not account for the diurnal hysteresis effect.     

Soil CO2 efflux vs. soil respiration: Implications for flux models

In the long term, all CO₂ produced in the soil must be emitted by the surface and soil CO₂ efflux (FCO₂) must correspond to soil respiration (R_soil). In the short term, however, the efflux can deviate from the instantaneous soil respiration, if the amount of CO₂ stored in the soil pore-space (SCO₂) is changing. We measured FCO₂ continuously for one year using an automated chamber system. Simultaneously, vertical soil profiles of CO₂ concentration, moisture, and temperature were measured in order to assess the changes in the amount of CO₂ stored in the soil. R_soil was calculated as the sum of the rate of change of the CO₂ storage over time and FCO₂. The experiment was split into a warm and a cold season. The dependency of soil respiration and soil efflux on soil temperature and on soil moisture was analyzed separately. Only the moisture-driven model of the warm season was significantly different for FCO₂ and R_soil. At our site, a moisture-driven soil-respiration model derived from CO₂ efflux data would underestimate the importance of soil moisture. This effect can be attributed to a temporary storage of CO₂ in the soil pore-space after rainfalls where up to 40% of the respired CO₂ were stored.     

Litter contribution to diurnal and annual soil respiration in a tropical montane cloud forest

Respiration of CO₂ from soils (R_s) is a major component of the carbon cycle of ecosystems, but understanding is still poor of both the relative contributions of different respiratory sources to R_s, and the environmental factors that drive diurnal variations in R_s. We measured total and litter-free R_s at half-hourly intervals over full 24 h periods, and thereafter twice a month for 10 months in a tropical montane cloud forest (TMCF) in Peru. Total R_s declined by about 61% during the night as a result of variations in respiration rate in the litter, which were partly correlated with the soil surface air temperature. Most of the diurnal variation of R_s in this TMCF appears to be driven by respiration in the litter layer, which contributed 37% to the total soil CO₂ efflux. Total R_s rates at this particular site would have been overestimated by 60% if derived from daytime measurements that had not been corrected for diurnal variations in R_s.