Uncoupling of microbial CO2 production and release in frozen soil and its implications for field studies of arctic C cycling

Knowledge of seasonal trends and controls of soil CO₂ emissions to the atmosphere is important for simulating atmospheric CO₂ concentrations and for understanding and predicting the global carbon cycle. This is particularly the case for high arctic soils subject to extreme fluctuating environmental conditions. Based on field measurements of soil CO₂ efflux, temperature, water content, pore gas composition in soil and frozen cores as well as detailed temperature experiments performed in the laboratory, we evaluated seasonal controls of CO₂ effluxes from a well-drained tundra heath site in NE-Greenland. During the growing season, near-surface temperatures correlated well with observed CO₂ effluxes (r²>0.9). However, during intensive thawing of near-surface layers we observed up to 1.5-fold higher effluxes than expected due to temperature alone. These high rates were consistent with high CO₂ concentrations in frozen soil (>10% CO₂) and suggested a spring burst event during soil thawing and a corresponding trapping of produced CO₂ during winter. Laboratory experiments revealed that microbial soil respiration continued down to a least −18 °C and that up to 80% of the produced CO₂ was trapped in soil at temperatures between 0 and −9 °C. The trapping of CO₂ in frozen soil was positively correlated with soil moisture (r²=0.85) and led to an abrupt change of the temperature sensitivity (Q₁₀) observed for soil CO₂ release at 0 °C with Q₁₀ values below 0 °C being up to 100-fold higher than above 0 °C. The results of sub-zero CO₂ production allowed us to predict the microbial soil respiration throughout the year and to evaluate to what extent burst events during thawing can be explained by the release of CO₂ being produced and trapped during winter. Taking only the upper 20 cm of the soil into account, winter soil respiration accounted for about 40% of the annual soil respiration. At least 14% of the winter CO₂ production was trapped during the winter 2000-2001 and observed to be released upon thawing. Thus, the site-specific winter soil respiration is an important part of the annual C cycle and CO₂ trapping should be accounted for in future field and modelling studies of soil respiration dynamics in arctic ecosystems. In conclusion, we have discovered a soil moisture dependent uncoupling of CO₂ production and release in frozen soils with important implications for future field studies of Arctic C cycling.     

Microbial activity in soils frozen to below −39 °C

Recent research on life in extreme environments has shown that some microorganisms metabolize at extremely low temperatures in Arctic and Antarctic ice and permafrost. Here, we present kinetic data on CO₂ and ¹⁴CO₂ release from intact and ¹⁴C-glucose amended tundra soils (Barrow, Alaska) incubated for up to a year at 0 to −39°C. The rate of CO₂ production declined exponentially with temperature but it remained positive and measurable, e.g. 2-7 ng CO₂-C cm⁻³ soil d⁻¹, at −39 °C. The variation of CO₂ release rate (v) was adequately explained by the double exponential dependence on temperature (T) and unfrozen water content (W) (r²>0.98): v=A exp(λT+kW) and where A, λ and k are constants. The rate of ¹⁴CO₂ release from added glucose declined more steeply with cooling as compared with the release of total CO₂, indicating that (a) there could be some abiotic component in the measured flux of CO₂ or (b) endogenous respiration is more cold-resistant than substrate-induced respiration. The respiration activity was completely eliminated by soil sterilization (1 h, 121 °C), stimulated by the addition of oxidizable substrate (glucose, yeast extract), and reduced by the addition of acetate, which inhibits microbial processes in acidic soils (pH 3-5). The tundra soil from Barrow displayed higher below-zero activity than boreal soils from West Siberia and Sweden. The permafrost soils (20-30 cm) were more active than the samples from seasonally frozen topsoil (0-10 cm, Barrow). Finding measurable respiration to −39 °C is significant for determining, understanding, and predicting current and future CO₂ emission to the atmosphere and for understanding the low temperature limits of microbial activity on the Earth and on other planets.     

Measurement of soil CO2 efflux using soda lime absorption: both quantitative and reliable

Measurement of soil CO₂ efflux using a non-flow-through steady-state (NFT-SS) chamber with alkali absorption of CO₂ by soda lime was tested and compared with a flow-through non-steady-state (FT-NSS) IRGA method to assess suitability of using soda lime for field monitoring over large spatial scales and integrated over a day. Potential errors and artifacts associated with the soda lime chamber method were investigated and improvements made. The following issues relating to quantification and reliable measurement of soil CO₂ efflux were evaluated: (i) absorption capacity of the soda lime, (ii) additional and thus artifactual absorption of CO₂ by soda lime during the experimental procedure, (iii) variation in the CO₂ concentration inside the chamber headspace, and (iv) effects of chamber closure on soil CO₂ efflux. Soil CO₂ efflux, as measured using soda lime (with a range of quantities: 50, 100, and 200 g per 0.082 m² ground area enclosed in chamber), was compared with transient IRGA measurements as a reference method that is based on well-established physical principles, using several forms of spatial and temporal comparisons. Natural variation in efflux rates ranged from 2 to 5.5 g C m⁻² day⁻¹ between different chambers and over different days. A comparison of the IRGA-based assay with measurement based on soda lime yielded an overall correlation coefficient of 0.82. The slope of the regression line was not significantly different from the 1:1 line, and the intercept was not significantly different from the origin. This result indicated that measurement of CO₂ efflux by soda lime absorption was quantitatively similar and unbiased in relation to the reference method. The soda lime method can be a highly practical method for field measurements if implemented with due care (in terms of drying and weighing soda lime, and in minimizing leakages), and validated for specific field conditions. A detailed protocol is presented for use of the soda lime method for measurement of CO₂ efflux from field soils.     

Variation of soil respiration at three spatial scales: Components within measurements, intra-site variation and patterns on the landscape

Soil respiration is an important component of terrestrial carbon cycling and can be influenced by many factors that vary spatially. This research aims to determine the extent and causes of spatial variation of soil respiration, and to quantify the importance of scale on measuring and modeling soil respiration within and among common forests of Northern Wisconsin. The potential sources of variation were examined at three scales: [1] variation among the litter, root, and bulk soil respiration components within individual 0.1 m measurement collars, [2] variation between individual soil respiration measurements within a site (<1 m to 10 m), and [3] variation on the landscape caused by topographic influence (100 m to 1000 m). Soil respiration was measured over a two-year period at 12 plots that included four forest types. Root exclusion collars were installed at a subset of the sites, and periodic removal of the litter layer allowed litter and bulk soil contributions to be estimated by subtraction. Soil respiration was also measured at fixed locations in six northern hardwood sites and two aspen sites to examine the stability of variation between individual measurements. These study sites were added to an existing data set where soil respiration was measured in a random, rotating, systematic clustering which allowed the examination of spatial variability from scales of <1 m to 100+ m. The combined data set for this area was also used to examine the influence of topography on soil respiration at scales of over 1000 m by using a temperature and moisture driven soil respiration model and a 4 km² digital elevation model (DEM) to model soil moisture. Results indicate that, although variation of soil respiration and soil moisture is greatest at scales of 100 m or more, variation from locations 1 m or less can be large (standard deviation during summer period of 1.58 and 1.28 μmol CO₂ m⁻² s⁻¹, respectively). At the smallest of scales, the individual contributions of the bulk soil, the roots, and the litter mat changed greatly throughout the season and between forest types, although the data were highly variable within any given site. For scales of 1-10 m, variation between individual measurements could be explained by positive relationships between forest floor mass, root mass, carbon and nitrogen pools, or root nitrogen concentration. Lastly, topography strongly influenced soil moisture and soil properties, and created spatial patterns of soil respiration which changed greatly during a drought event. Integrating soil fluxes over a 4 km² region using an elevation dependent soil respiration model resulted in a drought induced reduction of peak summer flux rates by 37.5%, versus a 31.3% when only plot level data was used. The trends at these important scales may help explain some inter-annual and spatial variability of the net ecosystem exchange of carbon.     

Developmental patterns of lamellae and stroma fractions of chloroplasts in rice plants

On examining the changes in lamellae and stroma nitrogen during leaf development, it is demonstrated that the lamellae and stroma fractions ofrice chloroplasts develop in quite different ways. In the case of stroma, the stroma materials existing in the leaf section which has just emerged from a leaf sheath are quite limited and the major part of this fraction is derived from the successive protein synthesis, i.e., the synthesis of this fraction was markedly increased during leaf expansion. This developmental pattern of the stroma coincided with the changes in the high-molecular-weight water soluble leaf protein, which seemed to be mainly composed of Fraction I protein. A rapid increase in stroma nitrogen was found to be a major cause for an increase in the leaf nitrogen content during leaf development.

On the other hand, the developmental pattern of the lamellae fraction was characterized by the fact that a considerable amount of this fraction had already been prepared when a leaf emerged from a leaf sheath and thereafter, no outstanding increase was seen compared to that of the stroma. This developmental pattern of the lamellae fraction resulted in a lowering of the proportion of lamellae nitrogen to the total leaf nitrogen during leaf development.

A great change in the lamellae-stroma composition of chloroplasts was observed. The proportion of stroma nitrogen to the total chloroplast nitrogen tended to increase as a leaf develops. Since the developmental stage varied according to the regions of a leaf, variation of the lamellaestroma composition was seen even within a leaf, i.e., the proportion of stroma nitrogen increased from base to tip.

In order to compare the synthetic rate of chlorophyll with those of the stroma and lamellae fractions, the changes in the ratios of stroma nitrogen/chlorophyll and lamellae nitrogen/chlorophyll were examined. The lamellae nitrogen/chlorophyll ratio decreased as a leaf developed, whereas the stroma nitrogen/chlorophyll ratio increased. Then the synthetic rates of these fractions during leaf development turned out to be of the same order as the stroma fraction, chlorophyll, lamellae fraction.     

黄土区刺槐林土壤含水量变化对土壤呼吸强度的影响

通过室内培养实验来评估土壤含水量的变化对土壤枯落物层、不同深度土壤层及DOC淋失后的土壤呼吸的影响.采集安塞纸坊沟31a刺槐林土样及林下混合枯落物,通过碱液吸收法测定100％,20％和2％含水量条件下3个深度土样(20,40和60 cm);去除DOC土样(仅100％含水量条件下);3种处理枯落物混合土样(林下混合枯落物、刺槐枯落物和草本类枯落物)培养过程中CO2的累计释放量.结果表明,100％和20％含水量条件下各深度土壤CO2释放量为20 cm土样＞60 cm土样＞40 cm土样;20 cm土样去除DOC后CO2释放量明显减少,40 cm明显增加,60 cm没有明显变化;混合枯落物土样在l00％含水量条件下CO2释放量最高;20％和2％含水量条件下刺槐枯落物CO2释放量明显大于草类,而100％含水量条件下草类枯落物略大于刺槐枯落物.研究证明土壤含水量对SOC组分含量和枯落物种类不同的土壤层呼吸强度存在差异性影响,强降水对DOC的淋失可造成表层土壤呼吸的减弱.     

Field measurement of carbon dioxide evolution from soil by a flow-through chamber method using a portable photosynthesis meter

Extract

Since a rise in atmospheric carbon dioxide (CO₂) concentration is expected to lead to global warming, it is important to quantify the global carbon circulation. The CO₂ evolution rate from soil has usually been measured by one of three methods: 1) CO₂ absorption (Anderson 1982), where the evolved CO₂ is absorbed in an alkali solution and the content subsequently determined, 2) closed chamber (Rolston 1986) in which the CO₂ evolution rate is calculated from the increase of the CO₂ concentration in a closed chamber covering the soil surface, and 3) flow-through chamber (Rolston 1986) in which a fixed rate of ambient air is pumped through an open chamber and the difference in the. CO₂ concentration between the inlet and the outlet is measured. Although the CO₂ absorption method is very simple in terms of apparatus and procedure, the determined CO₂ evolution rate tends to be underestimated in cases where the evolved CO₂ is not fully absorbed in the alkali solution (Ewel et al. 1987; Sakamoto and Yoshida 1988), or overestimated in cases where the CO₂ concentration in the chamber is too low to stimulate microbial activity (Koizumi et al. 1991; Nakadai et al. 1993), In the closed chamber method, when the gas concentration in the chamber is higher than that of the ambient air, gas diffusion from the soil to the atmosphere is restricted (Denmead 1978). At this point, the flow-through chamber method seems to be most suitable for measuring the CO₂ evolution rate, because the rate is determined under nearly natural conditions. However, this method has a disadvantage in that the apparatus is composed of an infra-red CO₂ analyzer, air pumps, mass flow meters, a recorder, and other items, which are too large, heavy, and complex to use in the field (Freijer and Bouten 1991). Hence, in spite of the above limitations, most of the studies on CO₂ evolution in situ have been carried out using the CO₂ absorption method (Kowalenko et al. 1978; Seto et al. 1978a, b; Ewel et al 1981, 1987; Gupta and Singh 1981; Reinke et al. 1981; Edwards and Ros-Todd 1983; Grahammer et al. 1991) or the closed chamber method (Naganawa et al. 1989; Mariko et al. 1994). The flow-through chamber method has been used only at sites where electric power supply and other types of equipment were available (Mathes and Schriefer 1985; Ewel et al. 1987; Nakadai et al. 1993). In the present report a flow-through chamber method using a portable CO₂ analyzer system was examined, for the determination of CO₂ evolution from soil without an electric power supply or other special equipment.     

A new technique to measure soil CO2 efflux at constant CO2 concentration

A new principle for measuring soil CO₂ efflux at constant ambient concentration is introduced. The measuring principle relies on the continuous absorption of CO₂ within the system to achieve a constant CO₂ concentration inside the soil chamber at ambient level, thus balancing the amount of CO₂ entering the soil chamber by diffusion from the soil. We report results that show reliable soil CO₂ efflux measurements with the new system. The novel measuring principle does not disturb the natural gradient of CO₂ within the soil, while allowing for continuous capture of the CO₂ released from the soil. It therefore holds great potential for application in simultaneous measurements of soil CO₂ efflux and its δ¹³C, since both variables show sensitivity to a distortion of the soil CO₂ profile commonly found in conventional chamber techniques.     

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.     

Theoretical model of the abiotic component of soil CO2 tracer efflux in C pulse-labeling experiments on plant-soil systems

For measurement of the time lag between photosynthesis and CO₂ efflux from soil, the carbon isotope pulse-labeling technique is considered as the most suitable. However, an interference from the abiotic tracer CO₂ component is identified as a key difficulty for obtaining accurate results with this technique. Guidelines on how to reduce this interference are therefore urgently needed. The flux of abiotic ¹³CO₂ tracer into soil during the labeling stage, and its return to atmosphere during the monitoring stage was modeled numerically, and the labeling stage also analytically. The controls of the abiotic interference were investigated using these models. The amount of the abiotic tracer component and the time distribution of its rate of return to the atmosphere, were predicted by these models. The main model parameters were D_m (=the ratio between the soil ¹³CO₂ diffusivity and the retardation factor), and the ¹³CO₂ concentration at the soil-atmosphere interface during the labeling stage (S₁₃), while background ¹³CO₂ soil production parameters were unnecessary. The presented models guide the selection of experimental parameters for minimization of the abiotic interference. With parameterization for a particular case, the present numerical model provides a preliminary order-of-magnitude estimate of the abiotic component, which would indicate if this interference is of significance.     

CO2浓度倍增对冬小麦生长发育产量形成及发芽率的影响

利用OTC-1型开顶式气室进行了CO₂浓度倍增对冬小麦影响的诊断试验,结果表明,CO₂浓度倍增对冬小麦生长发育、叶面积变化、生物量及产量形成等影响显著,且均为正效应。     

Effects of water amendment on basal and substrate-induced respiration rates of mineral soils

Summary We studied the effects of amending soils with different volumes of water or glucose solution on respiration rates measured as CO₂ evolution. Basal respiration was not significantly affected by the volume of water amendment, but substrate-induced respiration in static soil solutions was significantly reduced by increasing water contents. Inhibition of substrate-induced respiration was removed by continuously agitating the incubation vessels. Estimates of substrate-induced respiration rates for six soils differed markedly, depending on whether the vessels were stationary or agitated during the incubation. Agitation allowed increased discrimination between substrate-induced respiration rates for the soils, while static incubation only differentiated the soil with the highest substrate-induced respiration rate from the other soils.