华西雨屏区苦竹人工林土壤呼吸各组分特征及其温度敏感性

通过在华西雨屏区苦竹(Pleioblastus amarus)人工林内建立固定样地、定期监测等方法,研究该人工林生态系统土壤呼吸各组分特征及其温度敏感性.结果表明:2010年2月-2011年1月,苦竹林平均土壤呼吸速率为1.13 μmol·m-2·s-1,仲夏最高,深冬最低;凋落物层、无根土壤和植物根系对苦竹林土壤呼吸的贡献率分别为30.9％、20.8％和48.3％,各呼吸组分的季节动态均与土壤总呼吸类似,并与温度和凋落量等因素相关;苦竹林土壤总呼吸(RST)、凋落物层CO2排放(RSL)、无根土壤CO2排放(RSS)和植物根系呼吸(RSR)的年碳排放量分别为4.27、1.32、0.87和2.08 MgC· hm-2 ·a-1;土壤总呼吸及其各组分与凋落量呈显著正线性相关,与土壤10 cm温度和气温均呈显著正指数相关;基于土壤温度计算的RST、RSL、RSS和RSR的Q10值分别为2.90、2.28、3.09和3.19,凋落物层CO2排放的温度敏感性显著低于总呼吸和其他各组分.     

中国土壤呼吸温度敏感性空间格局的反演

土壤呼吸的温度敏感性（Q10）是模拟全球变暖与生态系统碳释放之间反馈强度的重要参数．虽然实验研究表明Q10值具有明显的空间异质性,但由于其空间分布格局的定量数据的缺乏,目前绝大多数生物地球化学模型将其简化成一个常数,并以此来预测未来的气候变化,这在一定程度上增大了模型预测的不确定性．本研究基于土壤有机碳的实测数据,并结合碳循环过程模型（CASA模型）,利用反演分析方法估算了8km空间分辨率下中国土壤呼吸温度敏感性的空间分布．结果表明,Q10值具有明显的空间异质性,且与实验方法估算的Q10值具有一致性;不同土壤类型的Q10值在1．09～2．38之间变化,其中火山灰土的Q10值最大,冷棕钙土的值最小;Q10值的空间分布与降水及土壤有机碳含量的关系密切．研究表明,该方法能有效反演Q10值的空间分布,从而有助于揭示碳循环规律并降低未来大气CO2浓度及气候变化预测的不确定性．     

土壤呼吸温度敏感性的影响因素和不确定性

土壤呼吸是陆地生态系统碳循环的重要环节之一, 其对温度升高的敏感程度在很大程度上决定着全球气候变化与碳循环之间的反馈关系。为了深刻理解地下生态过程对气候变化的响应和适应,本文综述了土壤呼吸温度敏感性（Q10）的影响因子及其内在机制,并分析了当前研究存在的不确定性。土壤生物、底物质量和底物供应显著调控着土壤呼吸的Q10值,但研究结论仍然有很大差异。温度和水分等环境因子则通过对土壤生物和底物的影响而作用于土壤呼吸的温度敏感性,一般情况下,随着温度的升高,土壤呼吸的Q10值下降;水分过高或过低时Q10值降低。另外本文从土壤温度测定深度、时空尺度、土壤呼吸不同组分温度敏感性差异、激发效应以及采用方法的不同等几方面分析了温度敏感性研究存在的不确定性。并在此基础上, 指出了未来拟重点加强的研究方向:（1）土壤呼吸不同组分温度敏感性差异的机理;（2）底物质量和底物供应对温度敏感性的交互影响;（3）生物因子对土壤呼吸温度敏感性的影响。     

施肥对油茶园土壤呼吸和异养呼吸及其温度敏感性的影响

油茶是中国南方重要的木本食用油料树种,研究施肥对油茶园土壤呼吸及其温度敏感性的影响,对于估算中国南方典型种植园林温室气体排放及其对气候变化的响应具有重要意义。设置对照(CK)、施肥(OF)、断根(CK-T)和断根施肥(OF-T)4个处理,采用静态箱-气相色谱法,通过多年观测,分析探讨施肥对油茶园土壤呼吸和异养呼吸及其温度敏感性的影响。结果表明:(1)施肥对油茶园土壤呼吸和异养呼吸无显著影响。研究期间,各处理(OF、CK、OF-T、CK-T)土壤CO_2通量依次为(77.91±2.59)、(73.71±0.97)、(66.82±1.02)mg C m~(-2)h~(-1)和(66.84±3.94)mg C m~(-2)h~(-1);(2)各处理土壤呼吸温度敏感性(Q_(10))表现为OF-T(1.96±0.01)CK-T(1.79±0.03)OF(1.77±0.01)CK(1.75±0.03),其中,OF-T处理下Q_(10)显著高于其他3个处理,即施肥显著增加了断根处理土壤呼吸Q_(10);(3)施肥显著增加了土壤表层NH_4~+-N和NO_3~--N含量,Q_(10)与土壤表层NH_4~+-N和NO_3~--N含量表现出显著的正相关关系。     

施石灰对杉木人工林土壤呼吸及其温度敏感性的影响

我国酸沉降主要分布区域与杉木人工林主要分布区域重合,石灰添加是改良酸化土壤的有效措施。为探究酸沉降背景下施石灰对土壤呼吸及其温度敏感性的影响,本研究以杉木人工林土壤为对象,在2018年6月一次性添加0、1和5 t·hm^-2的氧化钙,于2020年6月开始进行为期一年的原位土壤呼吸速率观测。结果表明：与不施石灰相比,施石灰显著提高了土壤pH值和交换性Ca²⁺含量,不同石灰施用量之间无显著差异。杉木人工林土壤呼吸及其组分具有明显的季节差异,表现为夏季最高,冬季最低,施石灰未显著改变其季节动态特征。施石灰显著降低了土壤异养呼吸速率,提高了自养呼吸速率,最终导致施石灰对土壤呼吸无显著影响。土壤呼吸月动态变化与温度月动态变化基本保持一致,土壤呼吸与土壤温度呈显著的指数关系,施石灰后土壤呼吸及自养呼吸的温度敏感性(Q₁₀)呈上升趋势,土壤异养呼吸的Q₁₀呈下降趋势。综上,施石灰提高了杉木人工林土壤自养呼吸,显著降低了土壤异养呼吸,这有利于杉木人工林土壤固碳。     

两种农田土壤不同组分呼吸及其温度敏感性

为探讨农田土壤不同组分呼吸及其对温度变化的响应,选取山东平邑旱耕土和湖南桃江水稻土为供试土壤,设置4个温度水平(5、15、25、35 ℃),对两种土壤的轻组、重组及全土进行63 d的培养试验.结果表明: 两种土壤全土的呼吸均高于轻组和重组.旱耕土重组的呼吸高于轻组,水稻土重组和轻组的呼吸在5～25 ℃温度水平下无显著差异,但35 ℃下重组高于轻组.在不同温度水平下,旱耕土轻组、重组和全土累积呼吸量分别占其初始碳的0.3%～2.8%、0.4%～3.7%和0.6%～7.0%,水稻土分别占其初始碳的0.4%～3.0%、0.3%～3.8%和0.7%～5.3%.两种土壤全土及轻、重组呼吸的温度敏感性(Q₁₀)均随温度升高和培养时间延长而降低;水稻土重组的Q₁₀高于轻组,旱耕土重组和轻组Q₁₀的差异无明显规律.在5～25 ℃温度水平下,旱耕土全土Q₁₀显著高于水稻土,但在25～35 ℃下低于水稻土.说明平邑旱耕土有机碳矿化强度高于桃江水稻土,且对温度变化的响应总体比水稻土更敏感.     

氮添加对高寒草甸土壤微生物呼吸及其温度敏感性的影响

土壤氮素的可利用性是控制土壤微生物呼吸的重要因素之一,大量研究已经表明增加土壤活性氮的含量可以降低微生物呼吸,但是土壤氮输入对土壤微生物呼吸温度敏感性的影响还不清楚。以青藏高原高寒草甸为研究对象,通过野外施氮试验和室内控制试验相结合的方式,在5℃、15℃和25℃条件下对3种施氮水平的土壤(对照,0g N m~(-2)a~(-1);低氮,5g N m~(-2)a~(-1);高氮,15g N m~(-2)a~(-1))进行培养,探讨土壤微生物呼吸及其温度敏感性对不同氮添加水平的响应情况。结果表明:(1)3个温度培养下的土壤微生物呼吸速率和累积碳释放量均随施氮量的增加而显著降低(P0.05);(2)氮添加对5℃和15℃培养条件下的微生物呼吸温度敏感性没有显著影响,但显著地增加了15℃和25℃培养条件下的微生物呼吸温度敏感性(P0.05);(3)线性相关分析表明,土壤累积碳释放量与土壤有机碳的难降解性显著负相关(P0.05),而15℃和25℃培养条件下的微生物呼吸温度敏感性与土壤有机碳的难降解性显著正相关(P0.05)。结果表明,在全球气候变暖的背景下,土壤氮输入将增加预测青藏高原高寒草甸地区土壤碳排放的不确定性。     

不同土地利用方式下土壤呼吸及其温度敏感性

采用静态箱-气相色谱法对四川盆地中部紫色土丘陵区3种土地利用方式（林地、草地和轮作旱地）土壤呼吸进行测定,结果表明,林地、草地和旱地土壤呼吸速率变化范围分别为78．63～577．97、39．28～584．18和34．48～484．65mgCO2·m^-2·h^-1,年平均土壤呼吸速率分别为264．68、242．91、182．21mgCO2·m^-2h^-1。3种土地利用方式的土壤呼吸速率季节变化趋势均呈单峰曲线,林地和草地土壤呼吸速率最大值均出现在夏末（7月底与8月初之间）,旱地土壤呼吸速率最大值出现的时间比林地和草地要早,在6月底与7月初之间;最小值均出现在12月底与翌年1月初之间。土壤温度和土壤湿度是影响本地区土壤呼吸的主要因子,双因素关系模型（R=αe^bTw^c）较好地拟合了土壤温度和土壤湿度对土壤呼吸的影响,二者共同解释了土壤呼吸变化的64％～90％。土壤呼吸的温度敏感性指数Q10值受土壤（5cm处）温度和土壤（0～10cm）湿度的影响。分析表明3种土地利用土壤的Q10值与土壤温度呈显著负相关关系,而与土壤湿度呈显著正相关关系。     

模拟氮沉降对长江滩地杨树林土壤呼吸温度敏感性的影响

研究氮沉降量增加对土壤呼吸温度敏感性的影响,对于研究土壤呼吸在气候变化中的作用有重要意义。以长江中下游滩地杨树人工林为对象,通过定位模拟氮沉降实验的方法,研究了滩地杨树人工林生态系统土壤呼吸的变化特征和土壤呼吸各组分的温度敏感性对几种氮沉降量浓度的短期响应。结果表明:(1)各处理土壤总呼吸、土壤微生物呼吸、根系呼吸与各层次土壤温度均呈显著正相关关系,和5cm层土壤温度相关性最大。5cm层土壤温度可以解释土壤总呼吸、土壤微生物呼吸和根系呼吸季节变化的比例分别为50.5%—71.0%、51.5%—73.9%、35.7%—63.2%;(2)对照组(CK,0g N m-2a-1)土壤总呼吸、土壤微生物呼吸与根呼吸的Q10值分别为2.54、2.72和1.94;(3)在各氮添加水平中,中氮水平(MN,10g N m-2a-1)促进了土壤总呼吸、土壤微生物呼吸和植物根呼吸的温度敏感性。高氮水平(HN,20g N m-2a-1)都降低了土壤总呼吸、土壤微生物呼吸和植物根呼吸的温度敏感性,低氮水平(LN,5g N m-2a-1)降低了土壤总呼吸和土壤微生物呼吸的温度敏感性,促进了根呼吸的敏感性。     

Soil respiration in six temperate forests in China

Scaling soil respiration (R_S), the major CO₂ source to the atmosphere from terrestrial ecosystems, from chamber‐based measurements to ecosystems requires studies on variations and correlations of R_S from various biomes and across geographic regions. However, few studies on R_S are available for Chinese temperate forest despite the importance of this forest in the national and global carbon budgets. In this study, we conducted 18‐month R_S measurements during 2004–2005 in six temperate forest types, representing the typical secondary forest ecosystems across various site conditions in northeastern China: Mongolian oak (Quercus mongolica Fisch.), aspen‐birch (Populous davidiana Dode and Betula platyphylla Suk.), mixed deciduous (no dominant tree species), hardwood (dominated by Fraxinus mandshurica Rupr., Juglans mandshurica Maxim., and Phellodendron amurense Rupr.) forests, Korean pine (Pinus koraiensis Sieb. et Zucc.) and Dahurian larch (Larix gmelinii Rupr.) plantations. Our specific objectives were to: (1) explore relationships of R_S against soil temperature and water content for the six forest ecosystems, (2) quantify annual soil surface CO₂ flux and its relations to belowground carbon storage, (3) examine seasonal variations in R_S and related environmental factors, and (4) quantify among‐ and within‐ecosystem variations in R_S. The R_S was positively correlated to soil temperature in all forest types, and was significantly influenced by the interactions of soil temperature and water content in the pine, larch, and mixed deciduous forests. The sensitivity of R_S to soil temperature at 10 cm depth (Q₁₀) ranged from 2.61 in the oak forest to 3.75 in the aspen‐birch forests. The Q₁₀ tended to increase with soil water content until reaching a threshold, and then decline. The annual R_S for the larch, pine, hardwood, oak, mixed deciduous, and aspen‐birch forests averaged 403, 514, 781, 785, 786, and 813 g C m^?2 yr^?1, respectively. The annual R_S of the broadleaved forests was 72% greater than that of the coniferous forests. The annual R_S was positively correlated to soil organic carbon (SOC) concentration at O horizon (R²=0.868) and total biomass of roots <0.5 cm in diameter (R²=0.748). The coefficient of variation (CV) of R_S among forest types averaged 25% across the 18‐month measurements. The CV of R_S within plots varied from 20% to 27%, significantly (P<0.001) greater than those among plots (9–15%), indicating the importance of the fine‐scaled heterogeneity in R_S. This study emphasized that variations in soil respiration and potential sampling bias should be appropriately tackled for accurate soil CO₂ flux estimates.     

Carbon limitation of soil respiration under winter snowpacks: potential feedbacks between growing season and winter carbon fluxes

A reduction in the length of the snow‐covered season in response to a warming of high‐latitude and high‐elevation ecosystems may increase soil carbon availability both through increased litter fall following longer growing seasons and by allowing early winter soil frosts that lyse plant and microbial cells. To evaluate how an increase in labile carbon during winter may affect ecosystem carbon balance we investigated the relationship between carbon availability and winter CO₂ fluxes at several locations in the Colorado Rockies. Landscape‐scale surveys of winter CO₂ fluxes from sites with different soil carbon content indicated that winter CO₂ fluxes were positively related to carbon availability and experimental additions of glucose to soil confirmed that CO₂ fluxes from snow‐covered soil at temperatures between 0 and ?3°C were carbon limited. Glucose added to snow‐covered soil increased CO₂ fluxes by 52–160% relative to control sites within 24 h and remained 62–70% higher after 30 days. Concurrently a shift in the δ¹³C values of emitted CO₂ toward the glucose value indicated preferential utilization of the added carbon confirming the presence of active heterotrophic respiration in soils at temperatures below 0°C. The sensitivity of these winter fluxes to substrate availability, coupled with predicted changes in winter snow cover, suggests that feedbacks between growing season carbon uptake and winter heterotrophic activity may have unforeseen consequences for carbon and nutrient cycling in northern forests. For example, published winter CO₂ fluxes indicate that on average 50% of growing season carbon uptake currently is respired during the winter; changes in winter CO₂ flux in response to climate change have the potential to reduce substantially the net carbon sink in these ecosystems.     

On the variability of respiration in terrestrial ecosystems: moving beyond Q10

Respiration, which is the second most important carbon flux in ecosystems following gross primary productivity, is typically represented in biogeochemical models by simple temperature dependence equations. These equations were established in the 19th century and have been modified very little since then. Recent applications of these equations to data on soil respiration have produced highly variable apparent temperature sensitivities. This paper searches for reasons for this variability, ranging from biochemical reactions to ecosystem‐scale substrate supply. For a simple membrane‐bound enzymatic system that follows Michaelis–Menten kinetics, the temperature sensitivities of maximum enzyme activity (V_max) and the half‐saturation constant that reflects the affinity of the enzyme for the substrate (K_m) can cancel each other to produce no net temperature dependence of the enzyme. Alternatively, when diffusion of substrates covaries with temperature, then the combined temperature sensitivity can be higher than that of each individual process. We also present examples to show that soluble carbon substrate supply is likely to be important at scales ranging from transport across membranes, diffusion through soil water films, allocation to aboveground and belowground plant tissues, phenological patterns of carbon allocation and growth, and intersite differences in productivity. Robust models of soil respiration will require that the direct effects of substrate supply, temperature, and desiccation stress be separated from the indirect effects of temperature and soil water content on substrate diffusion and availability. We speculate that apparent Q₁₀ values of respiration that are significantly above about 2.5 probably indicate that some unidentified process of substrate supply is confounded with observed temperature variation.     

The response of soil CO2 flux to changes in atmospheric CO2, nitrogen supply and plant diversity

We measured soil CO₂ flux over 19 sampling periods that spanned two growing seasons in a grassland Free Air Carbon dioxide Enrichment (FACE) experiment that factorially manipulated three major anthropogenic global changes: atmospheric carbon dioxide (CO₂) concentration, nitrogen (N) supply, and plant species richness. On average, over two growing seasons, elevated atmospheric CO₂ and N fertilization increased soil CO₂ flux by 0.57 µmol m^?2 s^?1 (13% increase) and 0.37 µmol m^?2 s^?1 (8% increase) above average control soil CO₂ flux, respectively. Decreases in planted diversity from 16 to 9, 4 and 1 species decreased soil CO₂ flux by 0.23, 0.41 and 1.09 µmol m^?2 s^?1 (5%, 8% and 21% decreases), respectively. There were no statistically significant pairwise interactions among the three treatments. During 19 sampling periods that spanned two growing seasons, elevated atmospheric CO₂ increased soil CO₂ flux most when soil moisture was low and soils were warm. Effects on soil CO₂ flux due to fertilization with N and decreases in diversity were greatest at the times of the year when soils were warm, although there were no significant correlations between these effects and soil moisture. Of the treatments, only the N and diversity treatments were correlated over time; neither were correlated with the CO₂ effect. Models of soil CO₂ flux will need to incorporate ecosystem CO₂ and N availability, as well as ecosystem plant diversity, and incorporate different environmental factors when determining the magnitude of the CO₂, N and diversity effects on soil CO₂ flux.     

No overall stimulation of soil respiration under mature deciduous forest trees after 7 years of CO2 enrichment

The anthropogenic rise in atmospheric CO₂ is expected to impact carbon (C) fluxes not only at ecosystem level but also at the global scale by altering C cycle processes in soils. At the Swiss Canopy Crane (SCC), we examined how 7 years of free air CO₂ enrichment (FACE) affected soil CO₂ dynamics in a ca. 100‐year‐old mixed deciduous forest. The use of ¹³C‐depleted CO₂ for canopy enrichment allowed us to trace the flow of recently fixed C. In the 7th year of growth at ～550 ppm CO₂, soil respiratory CO₂ consisted of 39% labelled C. During the growing season, soil air CO₂ concentration was significantly enhanced under CO₂‐exposed trees. However, elevated CO₂ failed to stimulate cumulative soil respiration (R_s) over the growing season. We found periodic reductions as well as increases in instantaneous rates of R_s in response to elevated CO₂, depending on soil temperature and soil volumetric water content (VWC; significant three‐way interaction). During wet periods, soil water savings under CO₂‐enriched trees led to excessive VWC (>45%) that suppressed R_s. Elevated CO₂ stimulated R_s only when VWC was ≤40% and concurrent soil temperature was high (>15 °C). Seasonal Q₁₀ estimates of R_s were significantly lower under elevated (Q₁₀=3.30) compared with ambient CO₂ (Q₁₀=3.97). However, this effect disappeared when three consecutive sampling dates of extremely high VWC were disregarded. This suggests that elevated CO₂ affected Q₁₀ mainly indirectly through changes in VWC. Fine root respiration did not differ significantly between treatments but soil microbial biomass (C_mic) increased by 14% under elevated CO₂ (marginally significant). Our findings do not indicate enhanced soil C emissions in such stands under future atmospheric CO₂. It remains to be shown whether C losses via leaching of dissolved organic or inorganic C (DOC, DIC) help to balance the C budget in this forest.     

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.     

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.