Carbon losses due to soil warming: Do autotrophic and heterotrophic soil respiration respond equally?

Global warming has the potential to increase soil respiration (R_S), one of the major fluxes in the global carbon (C) cycle. R_S consists of an autotrophic (R_A) and a heterotrophic (R_H) component. We combined a soil warming experiment with a trenching experiment to assess how R_S, R_A, and R_H are affected. The experiment was conducted in a mature forest dominated by Norway spruce. The site is located in the Austrian Alps on dolomitic bedrock. We warmed the soil of undisturbed and trenched plots by means of heating cables 4 °C above ambient during the snow‐free seasons of 2005 and 2006. Soil warming increased the CO₂ efflux from control plots (R_S) by ∼45% during 2005 and ∼47% during 2006. The CO₂ efflux from trenched plots (R_H) increased by ∼39% during 2005 and ∼45% during 2006. Similar responses of R_S and R_H indicated that the autotrophic and heterotrophic components of R_S responded equally to the temperature increase. Thirty‐five to forty percent or 1 t C ha⁻¹ yr⁻¹ of the overall annual increase in R_S (2.8 t C ha⁻¹ yr⁻¹) was autotrophic. The remaining, heterotrophic part of soil respiration (1.8 t C ha⁻¹ yr⁻¹), represented the warming‐induced C loss from the soil. The autotrophic component showed a distinct seasonal pattern. Contribution of R_A to R_S was highest during summer. Seasonally derived Q₁₀ values reflected this pattern and were correspondingly high (5.3–9.3). The autotrophic CO₂ efflux increase due to the 4 °C warming implied a Q₁₀ of 2.9. Hence, seasonally derived Q₁₀ of R_A did not solely reflect the seasonal soil temperature development.     

Trends and methodological impacts in soil CO2 efflux partitioning: A metaanalytical review

Partitioning soil carbon dioxide (CO₂) efflux (R_S) into autotrophic (R_A; including plant roots and closely associated organisms) and heterotrophic (R_H) components has received considerable attention, as differential responses of these components to environmental change have profound implications for the soil and ecosystem C balance. The increasing number of partitioning studies allows a more detailed analysis of experimental constraints than was previously possible. We present results of an exhaustive literature search of partitioning studies and analyse global trends in flux partitioning between biomes and ecosystem types by means of a metaanalysis. Across all data, an overall decline in the R_H/R_S ratio for increasing annual R_S fluxes emerged. For forest ecosystems, boreal coniferous sites showed significantly higher (P<0.05) R_H/R_S ratios than temperate sites, while both temperate or tropical deciduous forests did not differ in ratios from any of the other forest types. While chronosequence studies report consistent declines in the R_H/R_S ratio with age, no difference could be detected for different age groups in the global data set. Different methodologies showed generally good agreement if the range of R_S under which they had been measured was considered, with the exception of studies estimating R_H by means of root mass regressions against R_S, which resulted in consistently lower R_H/R_S estimates out of all methods included. Additionally, the time step over which fluxes were partitioned did not affect R_H/R_S ratios consistently. To put results into context, we review the most common techniques and point out the likely sources of errors associated with them. In order to improve soil CO₂ efflux partitioning in future experiments, we include methodological recommendations, and also highlight the potential interactions between soil components that may be overlooked as a consequence of the partitioning process itself.     

Experimental forest soil warming: response of autotrophic and heterotrophic soil respiration to a short-term 10°C temperature rise

We warmed the top soil of a mature coniferous forest stand by means of heating cables on control and trenched plots within 24 h by 10°C at 1 cm soil depth (9°C at 5 cm depth) and measured the effect on the autotrophic (R_A) and heterotrophic (R_H) component of total soil CO₂ efflux (R_S). The short time frame of warming enabled us to exclude confounding fluctuations in soil moisture and carbon (C) flow from the canopy. The results of the field study were backed up by a lab soil incubation experiment. During the first 12 h of warming, R_A strongly responded to soil warming; The Q ₁₀ values were 5.61 and 6.29 for 1 and 5 cm soil depth temperature. The Q ₁₀ values for R_A were almost twice as high as the Q ₁₀ values of R_H (3.04 and 3.53). Q ₁₀ values above 5 are above reasonable plant physiological values for root respiration. We see interactions of roots, mycorrhizae and heterotrophic microbes, combined with fast substrate supply to the rhizosphere as an explanation for the high short-term temperature response of R_A. When calculated over the whole duration (24 h) of the field soil-warming experiment, temperature sensitivities of R_A and R_H were similar (no significant difference at P < 0.05); Q ₁₀ values were 3.16 and 3.96 for R_A and 2.94 and 3.35 for R_H calculated with soil temperatures at 1 and 5 cm soil depth, respectively. Laboratory incubation showed that different soil moisture contents of trenched and control plots affected rates of R_H, but did not affect the temperature sensitivity of R_H. We conclude that a single parameter is sufficient to describe the temperature sensitivity of R_S in soil C models which operate on larger temporal and spatial scales. The strong short-term response of R_A may be of relevance in soils suspected to experience increasingly strong diurnal temperature variations.     

Impacts of an anomalously warm year on soil CO2 efflux in experimentally manipulated tallgrass prairie ecosystems

Modeling analyses suggest that an increase in growth rate of atmospheric CO₂ concentrations during an anomalously warm year may be caused by a decrease in net ecosystem production (NEP) in response to increased heterotrophic respiration (R_h). To test this hypothesis, 12 intact soil monoliths were excavated from a tallgrass prairie site near Purcell, Oklahoma, USA and divided among four large dynamic flux chambers (Ecologically Controlled Enclosed Lysimeter Laboratories (EcoCELLs)). During the first year, all four EcoCELLs were subjected to Oklahoma air temperatures. During the second year, air temperature in two EcoCELLs was increased by 4°C throughout the year to simulate anomalously warm conditions. This paper reports on the effect of warming on soil CO₂ efflux, representing the sum of autotrophic respiration (R_a) and R_h. During the pretreatment year, weekly average soil CO₂ efflux was similar in all EcoCELLs. During the late spring, summer and early fall of the treatment year, however, soil CO₂ efflux was significantly lower in the warmed EcoCELLs. In general, soil CO₂ efflux was correlated with soil temperature and to a lesser extent with moisture. A combined temperature and moisture regression explained 64% of the observed variation in soil CO₂ efflux. Soil CO₂ efflux correlated well with a net primary production (NPP) weighted greenness index derived from digital photographs. Although separate relationships for control and warmed EcoCELLs showed better correlations, one single relationship explained close to 70% of the variation in soil CO₂ efflux across treatments and years. A strong correlation between soil CO₂ efflux and canopy development and the lack of initial response to warming indicate that soil CO₂ efflux is dominated by R_a. This study showed that a decrease in soil CO₂ efflux in response to a warm year was most likely dominated by a decrease in R_a instead of an increase in R_h.     

Responses of dryland soil respiration and soil carbon pool size to abrupt vs. gradual and individual vs. combined changes in soil temperature, precipitation, and atmospheric [CO2]: a simulation analysis

With the large extent and great amount of soil carbon (C) storage, drylands play an important role in terrestrial C balance and feedbacks to climate change. Yet, how dryland soils respond to gradual and concomitant changes in multiple global change drivers [e.g., temperature (T_s), precipitation (Ppt), and atmospheric [CO₂] (CO₂)] has rarely been studied. We used a process‐based ecosystem model patch arid land simulator to simulate dryland soil respiration (R_s) and C pool size (C_s) changes to abrupt vs. gradual and single vs. combined alterations in T_s, Ppt and CO₂ at multiple treatment levels. Results showed that abrupt perturbations generally resulted in larger R_s and had longer differentiated impacts than did gradual perturbations. R_s was stimulated by increases in T_s, Ppt, and CO₂ in a nonlinear fashion (e.g., parabolically or asymptotically) but suppressed by Ppt reduction. Warming mainly stimulated heterotrophic R_s (i.e., R_h) whereas Ppt and CO₂ influenced autotrophic R_s (i.e., R_a). The combined effects of warming, Ppt, and CO₂ were nonadditive of primary single‐factor effects as a result of substantial interactions among these factors. Warming amplified the effects of both Ppt addition and CO₂ elevation whereas Ppt addition and CO₂ elevation counteracted with each other. Precipitation reduction either magnified or suppressed warming and CO₂ effects, depending on the magnitude of factor's alteration and the components of R_s (R_a or R_h) being examined. Overall, Ppt had dominant influence on dryland R_s and C_s over T_s and CO₂. Increasing Ppt individually or in combination with T_s and CO₂ benefited soil C sequestration. We therefore suggested that global change experimental studies for dryland ecosystems should focus more on the effects of precipitation regime changes and the combined effects of Ppt with other global change factors (e.g., T_s, CO₂, and N deposition).     

Effects of soil moisture on the temperature sensitivity of heterotrophic respiration vary seasonally in an old‐field climate change experiment

Microbial decomposition of soil organic matter produces a major flux of CO₂ from terrestrial ecosystems and can act as a feedback to climate change. Although climate‐carbon models suggest that warming will accelerate the release of CO₂ from soils, the magnitude of this feedback is uncertain, mostly due to uncertainty in the temperature sensitivity of soil organic matter decomposition. We examined how warming and altered precipitation affected the rate and temperature sensitivity of heterotrophic respiration (R_h) at the Boston‐Area Climate Experiment, in Massachusetts, USA. We measured R_h inside deep collars that excluded plant roots and litter inputs. In this mesic ecosystem, R_h responded strongly to precipitation. Drought reduced R_h, both annually and during the growing season. Warming increased R_h only in early spring. During the summer, when R_h was highest, we found evidence of threshold, hysteretic responses to soil moisture: R_h decreased sharply when volumetric soil moisture dropped below ~15% or exceeded ~26%, but R_h increased more gradually when soil moisture rose from the lower threshold. The effect of climate treatments on the temperature sensitivity of R_h depended on the season. Apparent Q₁₀ decreased with high warming (~3.5 °C) in spring and fall. Presumably due to limiting soil moisture, warming and precipitation treatments did not affect apparent Q₁₀ in summer. Drought decreased apparent Q₁₀ in fall compared to ambient and wet precipitation treatments. To our knowledge, this is the first field study to examine the response of R_h and its temperature sensitivity to the combined effects of warming and altered precipitation. Our results highlight the complex responses of R_h to soil moisture, and to our knowledge identify for the first time the seasonal variation in the temperature sensitivity of microbial respiration in the field. We emphasize the importance of adequately simulating responses such as these when modeling trajectories of soil carbon stocks under climate change scenarios.     

A global relationship between the heterotrophic and autotrophic components of soil respiration?

Soil surface CO₂ flux (R_S) is overwhelmingly the product of respiration by roots (autotrophic respiration, R_A) and soil organisms (heterotrophic respiration, R_H). Many studies have attempted to partition R_S into these two components, with highly variable results. This study analyzes published data encompassing 54 forest sites and shows that R_A and R_H are each strongly (R²>0.8) correlated to annual R_S across a wide range of forest ecosystems. Monte Carlo simulation showed that these correlations were significantly stronger than any correlation introduced as an artefact of measurement method. Biome type, measurement method, mean annual temperature, soil drainage, and leaf habit were not significant. For sites with available data, there was a significant (R²=0.56) correlation between total detritus input and R_H, while R_A was unrelated to net primary production. We discuss why R_A and R_H might be related to each other on large scales, as both ultimately depend on forest carbon balance and photosynthate supply. Limited data suggest that these or similar relationships have broad applicability in other ecosystem types. Site‐specific measurements are always more desirable than the application of inferred broad relationships, but belowground measurements are difficult and expensive, while measuring R_S is straightforward and commonly done. Thus the relationships presented here provide a useful method that can help constrain estimates of terrestrial carbon budgets.     

Wood CO2 efflux in a primary tropical rain forest

The balance between photosynthesis and plant respiration in tropical forests may substantially affect the global carbon cycle. Woody tissue CO₂ efflux is a major component of total plant respiration, but estimates of ecosystem‐scale rates are uncertain because of poor sampling in the upper canopy and across landscapes. To overcome these problems, we used a portable scaffolding tower to measure woody tissue CO₂ efflux from ground level to the canopy top across a range of sites of varying slope and soil phosphorus content in a primary tropical rain forest in Costa Rica. The objectives of this study were to: (1) determine whether to use surface area, volume, or biomass for modeling and extrapolating wood CO₂ efflux, (2) determine if wood CO₂ efflux varied seasonally, (3) identify if wood CO₂ efflux varied by functional group, height in canopy, soil fertility, or slope, and (4) extrapolate wood CO₂ efflux to the forest. CO₂ efflux from small diameter woody tissue (<10 cm) was related to surface area, while CO₂ efflux from stems >10 cm was related to both surface area and volume. Wood CO₂ efflux showed no evidence of seasonality over 2 years. CO₂ efflux per unit wood surface area at 25° (F_A) was highest for the N‐fixing dominant tree species Pentaclethra macroloba, followed by other tree species, lianas, then palms. Small diameter F_A increased steeply with increasing height, and large diameter F_A increased with diameter. Soil phosphorus and slope had slight, but complex effects on F_A. Wood CO₂ efflux per unit ground area was 1.34±0.36 μmol m^?2 s^?1, or 508±135 g C m^?2 yr^?1. Small diameter wood, only 15% of total woody biomass, accounted for 70% of total woody tissue CO₂ efflux from the forest; while lianas, only 3% of total woody biomass, contributed one‐fourth of the total wood CO₂ efflux.     

Precipitation variability and fire influence the temporal dynamics of soil CO2 efflux in an arid grassland

Climate models suggest that extreme rainfall events will become more common with increased atmospheric warming. Consequently, changes in the size and frequency of rainfall will influence biophysical drivers that regulate the strength and timing of soil CO₂ efflux – a major source of terrestrial carbon flux. We used a rainfall manipulation experiment during the summer monsoon season (July–September) to vary both the size and frequency of precipitation in an arid grassland 2 years before and 2 years after a lightning‐caused wildfire. Soil CO₂ efflux rates were always higher under increased rainfall event size than under increased rainfall event frequency, or ambient precipitation. Although fire reduced soil CO₂ efflux rates by nearly 70%, the overall responses to rainfall variability were consistent before and after the fire. The overall sensitivity of soil CO₂ efflux to temperature (Q₁₀) converged to 1.4, but this value differed somewhat among treatments especially before the fire. Changes in rainfall patterns resulted in differences in the periodicity of soil CO₂ efflux with strong signals at 1, 8, and 30 days. Increased rainfall event size enhanced the synchrony between photosynthetically active radiation and soil CO₂ efflux over the growing season before and after fire, suggesting a change in the temporal availability of substrate pools that regulate the temporal dynamics and magnitude of soil CO₂ efflux. We conclude that arid grasslands are capable of rapidly increasing and maintaining high soil CO₂ efflux rates in response to increased rainfall event size more than increased rainfall event frequency both before and after a fire. Therefore, the amount and pattern of multiple rain pulses over the growing season are crucial for understanding CO₂ dynamics in burned and unburned water‐limited ecosystems.     

Apparent thermal acclimation of soil heterotrophic respiration mainly mediated by substrate availability

Multiple lines of existing evidence suggest that increasing CO₂ emission from soils in response to rising temperature could accelerate global warming. However, in experimental studies, the initial positive response of soil heterotrophic respiration (R_H) to warming often weakens over time (referred to apparent thermal acclimation). If the decreased R_H is driven by thermal adaptation of soil microbial community, the potential for soil carbon (C) losses would be reduced substantially. In the meanwhile, the response could equally be caused by substrate depletion, and would then reflect the gradual loss of soil C. To address uncertainties regarding the causes of apparent thermal acclimation, we carried out sterilization and inoculation experiments using the soil samples from an alpine meadow with 6 years of warming and nitrogen (N) addition. We demonstrate that substrate depletion, rather than microbial adaptation, determined the response of R_H to long-term warming. Furthermore, N addition appeared to alleviate the apparent acclimation of R_H to warming. Our study provides strong empirical support for substrate availability being the cause of the apparent acclimation of soil microbial respiration to temperature. Thus, these mechanistic insights could facilitate efforts of biogeochemical modeling to accurately project soil C stocks in the future climate.     

Indirect partitioning of soil respiration in a series of evergreen forest ecosystems

A simple estimation of heterotrophic respiration can be obtained analytically as the y-intercept of the linear regression between soil-surface CO₂ efflux and root biomass. In the present study, a development of this indirect methodology is presented by taking into consideration both the temporal variation and the spatial heterogeneity of heterotrophic respiration. For this purpose, soil CO₂ efflux, soil carbon content and main stand characteristics were estimated in seven evergreen forest ecosystems along an elevation gradient ranging from 250 to 1740 m. For each site and for each sampling date the measured soil CO₂ efflux (R _S) was predicted with the model R _S = a × S _C + b × R _D ± ε, where S _C is soil carbon content per unit area to a depth of 30 cm and R _D is the root density of the 2–5 mm root class. Regressions with statistically significant a and b coefficients allowed the indirect separation of the two components of soil CO₂ efflux. Considering that the different sampling dates were characterized by different soil temperature, it was possible to investigate the temporal and thermal dependency of autotrophic and heterotrophic respiration. It was estimated that annual autotrophic respiration accounts for 16–58% of total soil CO₂ efflux in the seven different evergreen ecosystems. In addition, our observations show a decrease of annual autotrophic respiration at increasing availability of soil nitrogen. Section Editor: A. Hodge     

Coupling between carbon cycling and climate in a high-elevation,subalpine forest: a model-data fusion analysis

Fundamental questions exist about the effects of climate on terrestrial net ecosystem CO₂ exchange (NEE), despite a rapidly growing body of flux observations. One strategy to clarify ecosystem climate–carbon interactions is to partition NEE into its component fluxes, gross ecosystem CO₂ exchange (GEE) and ecosystem respiration (R _E), and evaluate the responses to climate of each component flux. We separated observed NEE into optimized estimates of GEE and R _E using an ecosystem process model combined with 6 years of continuous flux data from the Niwot Ridge AmeriFlux site. In order to gain further insight into the processes underlying NEE, we partitioned R _E into its components: heterotrophic (R _H) and autotrophic (R _A) respiration. We were successful in separating GEE and R _E, but less successful in accurately partitioning R _E into R _A and R _H. Our failure in the latter was due to a lack of adequate contrasts in the assimilated data set to distinguish between R _A and R _H. We performed most model runs at a twice-daily time step. Optimizing on daily-aggregated data severely degraded the model’s ability to separate GEE and R _E. However, we gained little benefit from using a half-hourly time step. The model-data fusion showed that most of the interannual variability in NEE was due to variability in GEE, and not R _E. In contrast to several previous studies in other ecosystems, we found that longer growing seasons at Niwot Ridge were correlated with less net CO₂ uptake, due to a decrease of available snow-melt water during the late springtime photosynthetic period. Warmer springtime temperatures resulted in increased net CO₂ uptake only if adequate moisture was available; when warmer springtime conditions led into mid-summer drought, the annual net uptake declined.     

Elevated CO2 does not affect stem CO2 efflux nor stem respiration in a dry Eucalyptus woodland,but it shifts the vertical gradient in xylem [CO2]

To quantify stem respiration (R_S) under elevated CO₂ (eCO₂), stem CO₂ efflux (E_A) and CO₂ flux through the xylem (F_T) should be accounted for, because part of respired CO₂ is transported upwards with the sap solution. However, previous studies have used E_A as a proxy of R_S, which could lead to equivocal conclusions. Here, to test the effect of eCO₂ on R_S, both E_A and F_T were measured in a free‐air CO₂ enrichment experiment located in a mature Eucalyptus native forest. Drought stress substantially reduced E_A and R_S, which were unaffected by eCO₂, likely as a consequence of its neutral effect on stem growth in this phosphorus‐limited site. However, xylem CO₂ concentration measured near the stem base was higher under eCO₂, and decreased along the stem resulting in a negative contribution of F_T to R_S, whereas the contribution of F_T to R_S under ambient CO₂ was positive. Negative F_T indicates net efflux of CO₂ respired below the monitored stem segment, likely coming from the roots. Our results highlight the role of nutrient availability on the dependency of R_S on eCO₂ and suggest stimulated root respiration under eCO₂ that may shift vertical gradients in xylem [CO₂] confounding the interpretation of E_A measurements.     

中国南亚热带森林不同演替阶段土壤呼吸的分离量化

量化森林土壤呼吸(R_S)及其组分对准确地评估森林土壤碳吸存极其重要。该文以鼎湖山南亚热带季风常绿阔叶林及其演替系列针阔叶混交林和马尾松(Pinus massoniana)林为研究对象, 采用挖壕沟法结合静态气室CO₂测定法对这3种林分类型的R_S进行分离量化。结果表明: 鼎湖山3种森林演替系列上的森林R_S及其组分(自养呼吸R_A、异养呼吸R_H)均呈现出明显的季节动态, 表现为夏季最高、冬季最低的格局。在呼吸总量上, 季风常绿阔叶林显著高于针阔叶混交林和马尾松林, 但混交林与马尾松林之间差异不显著; R_A除季风常绿阔叶林显著大于针阔叶混交林外, 其余林分之间差异不显著; 对于R_H来说, 3个林分之间均无显著差异。随着森林正向演替的进行, 由马尾松林至针阔叶混交林至季风常绿阔叶林, R_A对土壤总呼吸的年平均贡献率分别为(39.48 ± 15.49)%、(33.29 ± 17.19)%和(44.52 ± 10.67)%, 3个林分之间差异不显著。方差分析结果表明, 土壤温度是影响R_S及其组分的主要环境因子, 温度与R_S及其组分呈显著的指数关系; 土壤含水量对R_S的影响不显著, 甚至表现为轻微的抑制现象, 但未达到显著性水平。对温度敏感性指标Q₁₀值的分析表明, 3个林分均为R_A的温度敏感性最大, R_H的温度敏感性最小。     

Rhizospheric and heterotrophic components of soil respiration in six Chinese temperate forests

Partitioning soil respiration (R_S) into heterotrophic (R_H) and rhizospheric (R_R) components is an important step for understanding and modeling carbon cycling in forest ecosystems, but few studies on R_R and R_H exist in Chinese temperate forests. In this study, we used a trenching plot approach to partition R_S in six temperate forests in northeastern China. Our specific objectives were to (1) examine seasonal patterns of soil surface CO₂ fluxes from trenched (R_T) and untrenched plots (R_UT) of these forests; (2) quantify annual fluxes of R_S components and their relative contributions in the forest ecosystems; and (3) examine effects of plot trenching on measurements of R_S and related environmental factors. The R_T maximized in early growing season, but the difference between R_UT and R_T peaked in later summer. The annual fluxes of R_H and R_R varied with forest types. The estimated values of R_H for the Korean pine (Pinus koraiensis Sieb. et Zucc.), Dahurian larch (Larix gmelinii Rupr.), aspen‐birch (Populous davidiana Dode and Betula platyphylla Suk.), hardwood (Fraxinus mandshurica Rupr., Juglans mandshurica Maxim. and Phellodendron amurense Rupr.), Mongolian oak (Quercus mongolica Fisch.) and mixed deciduous (no dominant tree species) forests averaged 89, 196, 187, 245, 261 and 301 g C m⁻² yr⁻¹, respectively; those of R_R averaged 424, 209, 628, 538, 524 and 483 g C m⁻² yr⁻¹, correspondingly; calculated contribution of R_R to R_S (RC) varied from 52% in the larch forest to 83% in the pine forest. The annual flux of R_R was strongly correlated to biomass of roots <0.5 cm in diameter, while that of R_H was weakly correlated to soil organic carbon concentration at A horizon. We concluded that vegetation type and associated carbon metabolisms of temperate forests should be considered in assessing and modeling R_S components. The significant impacts of changed soil physical environments and substrate availability by plot trenching should be appropriately tackled in analyzing and interpreting measurements of R_S components.     

Soil respiration under climate change: prolonged summer drought offsets soil warming effects

Climate change may considerably impact the carbon (C) dynamics and C stocks of forest soils. To assess the combined effects of warming and reduced precipitation on soil CO₂ efflux, we conducted a two‐way factorial manipulation experiment (4 °C soil warming + throughfall exclusion) in a temperate spruce forest from 2008 until 2010. Soil was warmed by heating cables throughout the growing seasons. Soil drought was simulated by throughfall exclusions with three 100 m² roofs during 25 days in July/August 2008 and 2009. Soil warming permanently increased the CO₂ efflux from soil, whereas throughfall exclusion led to a sharp decrease in soil CO₂ efflux (45% and 50% reduction during roof installation in 2008 and 2009, respectively). In 2008, CO₂ efflux did not recover after natural rewetting and remained lowered until autumn. In 2009, CO₂ efflux recovered shortly after rewetting, but relapsed again for several weeks. Drought offset the increase in soil CO₂ efflux by warming in 2008 (growing season CO₂ efflux in t C ha^?1: control: 7.1 ± 1.0; warmed: 9.5 ± 1.7; warmed + roof: 7.4 ± 0.3; roof: 5.9 ± 0.4) and in 2009 (control: 7.6 ± 0.8; warmed + roof: 8.3 ± 1.0). Throughfall exclusion mainly affected the organic layer and the top 5 cm of the mineral soil. Radiocarbon data suggest that heterotrophic and autotrophic respiration were affected to the same extent by soil warming and drying. Microbial biomass in the mineral soil (0–5 cm) was not affected by the treatments. Our results suggest that warming causes significant C losses from the soil as long as precipitation patterns remain steady at our site. If summer droughts become more severe in the future, warming induced C losses will likely be offset by reduced soil CO₂ efflux during and after summer drought.     

Inhibition of whole plant respiration by elevated CO2 as modified by growth temperature

Plants of alfalfa (Medicago sativa) and orchard grass (Dactylus glomerata) were grown in controlled environment chambers at two CO₂ concentrations (350 and 700 μmol mol^-1) and 4 constant day/night growth temperatures of 15, 20, 25 and 30°C for 50–90 days to determine changes in growth and whole plant CO₂ efflux (dark respiration). To facilitate comparisons with other studies, respiration data were expressed on the basis of leaf area, dry weight and protein. Growth at elevated CO₂ increased total plant biomass at all temperatures relative to ambient CO₂, but the relative enhancement declined (P≤0.05) as temperature increased. Whole plant respiration (R_d) at elevated CO₂ declined at 15 and 20°C in D. glomerata on an area, weight or protein basis and in M. sativa on a weight or protein basis when compared to ambient CO₂. Separation of R_d into respiration required for growth (R_g) and maintenance (R_m) showed a significant effect of elevated CO₂ on both components. R_m was reduced in both species but only at lower temperatures (15°C in M. sativa and 15 and 20°C in D. glomerata). The effect on R_m could not be accounted for by protein content in either species. R_g was also reduced with elevated CO₂; however no particular effect of temperature was observed, i. e. R_g was reduced at 20, 25 and 30°C in M. sativa and at 15 and 25°C in D. glomerata. For the two perennial species used in the present study, the data suggest that both R_g and R_m can be reduced by anticipated increases in atmospheric CO₂; however, CO₂ inhibition of total plant respiration may decline as a function of increasing temperature     

Intraseasonal Variation in Water and Carbon Dioxide Flux Components in a Semiarid Riparian Woodland

Abstract We investigated how the distribution of precipitation over a growing season influences the coupling of carbon and water cycle components in a semiarid floodplain woodland dominated by the deep-rooted velvet mesquite (Prosopis velutina). Gross ecosystem production (GEP) and ecosystem respiration (R _eco) were frequently uncoupled because of their different sensitivities to growing season rainfall. Soon after the first monsoon rains, R _eco was high and was not proportional to slight increases in GEP. During the wettest month of the growing season (July), the system experienced a net carbon loss equivalent to 46% of the carbon accumulated over the 6-month study period (114 g C m⁻²; May–October). It appears that a large CO₂ efflux and a rapid water loss following precipitation early in the growing season and a later CO₂ gain is a defining characteristic of seasonally dry ecosystems. The relative contribution of plant transpiration (T) to total evapotranspiration (ET) (T/ET) was 0.90 for the entire growing season, with T/ET reaching a value of 1 during dry conditions and dropping to as low as 0.65 when the soil surface was wet. The evaporation fraction (E) was equivalent to 31% of the precipitation received during the study period (253 mm) whereas trees and understory vegetation transpired 38 and 31%, respectively, of this water source. The water-use efficiency of the vegetation (GEP/T) was higher later in the growing season when the C4 grassy understory was fully developed. The influence of rain on net ecosystem production (NEP) can be interpreted as the proportion of precipitation that is transpired by the plant community; the water-use efficiency of the vegetation and the precipitation fraction that is lost by evaporation.     

Responses of Soil,Heterotrophic, and Autotrophic Respiration to Experimental Open-Field Soil Warming in a Cool-Temperate Deciduous Forest

How global warming will affect soil respiration (R _S) and its source components is poorly understood despite its importance for accurate prediction of global carbon (C) cycles. We examined the responses of R _S, heterotrophic respiration (R _H), autotrophic respiration (R _A), nitrogen (N) availability, and fine-root biomass to increased temperature in an open-field soil warming experiment. The experiment was conducted in a cool-temperate deciduous forest ecosystem in northern Japan. As this forest is subjected to strong temporal variation in temperature, on scales ranging from daily to seasonal, we also investigated the temporal variation in the effects of soil warming on R _S, R _H, and R _A. Soil temperature was continuously elevated by about 4.0°C from 2007 to 2014 using heating wires buried in the soil, and we measured soil respiratory processes in all four seasons from 2012 to 2014. Soil warming increased annual R _S by 32–45%, but the magnitude of the increase was different between the components: R _H and R _A were also stimulated, and increased by 39–41 and 17–18%, respectively. Soil N availability during the growing season and fine-root biomass were not remarkably affected by the warming treatment. We found that the warming effects varied seasonally. R _H increased significantly throughout the year, but the warming effect showed remarkable seasonal differences, with the maximum stimulation in the spring. This suggests that warmer spring temperature will produce a greater increase in CO₂ release than warmer summer temperatures. In addition, we found that soil warming reduced the temperature sensitivity (Q ₁₀) of R _S. Although the Q ₁₀ of both R _H and R _A tended to be reduced, the decrease in the Q ₁₀ of R _S was caused mainly by a decrease in the response of R _A to warming. These long-term results indicate that a balance between the rapid and large response of soil microbes and the acclimation of plant roots both play important roles in determining the response of R _S to soil warming, and must be carefully considered to predict the responses of soil C dynamics under future temperature conditions.     

The response of soil respiration to precipitation change is asymmetric and differs between grasslands and forests

Intensification of the Earth's hydrological cycle amplifies the interannual variability of precipitation, which will significantly impact the terrestrial carbon (C) cycle. However, it is still unknown whether previously observed relationship between soil respiration (R_s) and precipitation remains applicable under extreme precipitation change. By analyzing the observations from a much larger dataset of field experiments (248 published papers including 151 grassland studies and 97 forest studies) across a wider range of precipitation manipulation than previous studies, we found that the relationship of R_s response with precipitation change was highly nonlinear or asymmetric, and differed significantly between grasslands and forests, between moderate and extreme precipitation changes. Response of R_s to precipitation change was negatively asymmetric (concave‐down) in grasslands, and double‐asymmetric in forests with a positive asymmetry (concave‐up) under moderate precipitation changes and a negative asymmetry (concave‐down) under extreme precipitation changes. In grasslands, the negative asymmetry in R_s response was attributed to the higher sensitivities of soil moisture, microbial and root activities to decreased precipitation (DPPT) than to increased precipitation (IPPT). In forests, the positive asymmetry was predominantly driven by the significant increase in microbial respiration under moderate IPPT, while the negative asymmetry was caused by the reductions in root biomass and respiration under extreme DPPT. The different asymmetric responses of R_s between grasslands and forests will greatly improve our ability to forecast the C cycle consequences of increased precipitation variability. Specifically, the negative asymmetry of R_s response under extreme precipitation change suggests that the soil C efflux will decrease across grasslands and forests under future precipitation regime with more wet and dry extremes.