植物源和土壤有机质源土壤呼吸组分的拆分原理、方法与应用进展

区分土壤呼吸组分并揭示其与环境因素的相关关系,对于准确评估土壤碳过程及其环境影响机制至关重要。根据底物来源和作用机制的差异,土壤呼吸主要包括根系呼吸、根际微生物呼吸、凋落物分解、自然条件下和激发效应下土壤有机质（SOM）分解。现有土壤呼吸组分拆分方法可以分为基于植物源CO₂测定或土壤有机质源CO₂测定的差分拆分方法,以及基于土壤呼吸组分同位素信号差异的拆分方法。土壤呼吸组分拆分研究可以解决不同土壤呼吸组分对环境变化的响应机制、植物光合碳输入与地下土壤呼吸组分的交互作用、土壤呼吸组分变化对土壤碳库周转的影响机制等科学问题,但其理论假设、观测技术方法、潜在的误差来源等仍需要继续关注并系统研究。     

火干扰对森林生态系统土壤呼吸组分的影响研究进展

土壤呼吸是陆地生态系统与大气碳交换的主要方式,主要分为自养呼吸和异养呼吸。土壤呼吸不仅是森林生态系统碳循环过程的关键环节,也是森林生态系统能量流动和物质循环的重要生态过程。火作为森林生态系统中一个重要的生态因子,可以在短时间内对土壤呼吸组分造成巨大的影响。火干扰对土壤呼吸组分的影响与火烧强度、火烧频率、火烧持续时间以及火后恢复等因子有关,通过影响植被的根系与组成、微生物群落数量与结构,凋落物的数量以及生态系统的环境和小气候等,进而对土壤呼吸产生影响。火干扰对土壤呼吸影响整体表现为火烧后土壤呼吸速率下降,在几个月至几年内恢复到火烧前水平,之后火继续对土壤呼吸产生影响长达数年至数十年。通过描述火烧强度、火烧频率以及火后恢复时间,阐述火干扰对土壤呼吸组分的直接影响,以及通过火后环境对土壤呼吸组分产生的间接影响,来揭示火干扰对森林生态系统土壤呼吸组分的影响。同时针对火干扰对土壤呼吸组分的影响进行以下3个方面的研究展望：（1）火后产生的黑碳对土壤呼吸组分的影响;（2）火后植被恢复对土壤呼吸组分产生的影响;（3）火后土壤呼吸组分的长期变化规律。     

鸡公山典型落叶阔叶林土壤呼吸对食叶虫灾爆发的响应

土壤呼吸是陆地生态系统碳循环的关键环节之一。随着极端气候事件的频发,森林虫害的发生频率和强度也趋于增加,森林虫害爆发已经是影响森林生态系统碳循环过程的一种重要的自然干扰。气候过渡带典型森林生态系统虫灾的爆发是否会影响土壤的碳排放过程目前仍不清楚。本研究利用鸡公山地区麻栎-枫香混交林大规模爆发食叶性害虫的机会,比较虫灾爆发当年(2014)与正常年份(2015)的土壤碳排放通量,以阐明森林虫灾爆发对土壤碳排放通量的影响。结果表明:虫灾爆发当年7、8、9、10月份土壤平均温度比正常年份相应各月份分别高0.26、0.51、0.83、0.07℃,土壤呼吸分别显著提高了129.9%、77.1%、61.6%和58.9%。虫灾爆发年份生长季的平均土壤呼吸为3.55μmol m~(-2)s~(-1),比正常年份(2.77μmol m~(-2)s~(-1))高36.2%;生长季期间的平均土壤异养呼吸比正常年份增加了29.7%。该研究表明森林食叶虫害的爆发至少在短期内可导致森林土壤碳排放量呈显著的增加趋势,近而对森林生态系统土壤碳库积累产生重要影响。因此,充分认识病虫害对森林生态系统的干扰和影响,将有助于陆地生态系统碳循环的准确估算和模拟。     

东北东部森林生态系统土壤呼吸组分的分离量化

对森林生态系统的土壤呼吸组分进行分离和量化,确定不同组分CO2释放速率的控制因子,是估测局域和区域森林生态系统碳平衡研究中必不可少的内容。采用挖壕法和红外气体分析法测定无根和有根样地的土壤表面CO2通量(RS),确定东北东部6种典型森林生态系统RS中异养呼吸(RH)和根系自养呼吸(RA)的贡献量及其影响因子。具体研究目标包括:(1)量化各种生态系统的RH及其与主要环境影响因子的关系;(2)量化各种生态系统RS中根系呼吸贡献率(RC)的季节动态;(3)比较6种森林生态系统RH和RA的年通量。土壤温度、土壤含水量及其交互作用显著地影响森林生态系统的RH(R2=0.465~0.788),但其影响程度因森林生态系统类型而异。硬阔叶林和落叶松人工林的RH主要受土壤温度控制,其他生态系统RH受土壤温度和含水量的联合影响。各个森林生态系统类型的RC变化范围依次为:硬阔叶林32.40%~51.44%;杨桦林39.72%~46.65%;杂木林17.94%~47.74%;蒙古栎林34.31%~37.36%;红松人工林33.78%~37.02%;落叶松人工林14.39%~35.75%。每个生态系统类型RH年通量都显著高于RA年通量,其变化范围分别为337~540 gC.m-2.a-1和88~331 gC.m-2.a-1。不同生态系统间的RH和RA也存在着显著性差异。     

青藏高原东缘亚高山针叶林人工恢复过程中的土壤呼吸特征

采用动态密闭气室红外CO₂分析法,对青藏高原东缘云杉人工林的土壤呼吸进行连续定位测定,并用挖壕沟法区分土壤自养呼吸和异养呼吸.结果表明:4种云杉林的土壤呼吸速率与土壤5 cm层温度有显著的正指数关系,与土壤含水量的相关性不显著.4种云杉林土壤呼吸年通量在792.08～1070.20 g C·m^-2·a^-1,大小依次为：天然云杉林＞22年生云杉人工林＞65年生云杉人工林＞35年生云杉人工林,随着人工林的恢复呈先降低后升高的趋势.在森林恢复过程中,人工云杉土壤自养和异养呼吸年通量均先减少后增加, 在253.36～357.05 g C·m^-2·a^-1和538.69～703.82 g C·m^-2·a^-1范围变化.22年生、35年生、65年生云杉人工林和天然云杉林非生长季 （2007-11-2008-03）和生长季（2008-04-2008-10）的Q₁₀值分别为：4.59、6.54、4.77、3.18和4.17、4.66、3.11、2.74.除22年生云杉人工林,Q₁₀值随云杉林的恢复更新而逐渐降低, 且非生长季节Q₁₀值均明显高于生长季节.     

Partitioning soil respiration of temperate forest ecosystems in northeastern China

Quantifying soil respiration components and their relations to environmental controls are essential to estimate both local and regional carbon (C) budgets of forest ecosystems. In this study, we used the trenching-plot and infrared gas exchange analyzer approaches to determine heterotrophic (R_H) and autotrophic respiration (R_A) in the soil surface CO₂ flux for six major temperate forest ecosystems in northeastern China. The ecosystems were: Mongolian oak forest (dominated by Quercus mongolica), aspen-birch forest (dominated by Populous davidiana and Betula platyphylla), mixed wood forest (composed of P. davidiana, B. platyphylla, Fraxinus mandshurica, Tilia amurensis, Acer amono, etc.), hardwood forest (dominated by F. mandshurica, Juglans mandshurica, and Phellodendron amurense), Korean pine (Pinus koraiensis), and Dahurian larch (Larix gmelinii) plantations, representing the typical secondary forest ecosystems in this region. Our specific objectives were to: (1) quantify R_H and its relationship with the environmental factors of the forest ecosystems, (2) characterize seasonal dynamics in the contribution of root respiration to total soil surface CO₂ flux (RC), and (3) compare annual CO₂ fluxes from R_H and R_A among the six forest ecosystems. Soil temperature, water content, and their interactions significantly affected R_H in the ecosystems and accounted for 46.5%–78.8% variations in R_H. However, the environmental controlling factors of R_H varied with ecosystem types: soil temperature in hardwood and Dahurian larch forest ecosystems, soil temperature, and water content in the others. The RC for hardwood, poplar-birch, mixed wood, Mongolian oak, Korean pine, and Dahurian larch forest ecosystems varied between 32.40%–51.44%, 39.72%–46.65%, 17.94%–47.74%, 34.31%–37.36%, 33.78%–37.02%, and 14.39%–35.75%, respectively. The annual CO₂ fluxes from R_H were significantly greater than those from R_A for all the ecosystems, ranging from 337–540 g Cm^-2a^-1 and 88‐331 gCm^-2a^-1 for R_H and R_A, respectively. The annual CO₂ fluxes from R_H and R_A differed significantly among the six forest ecosystems.     

北亚热带-南暖温带过渡区典型森林生态系统土壤呼吸及其组分分离

为阐明北亚热带．南暖温带过渡区典型森林生态系统土壤呼吸与其组分的碳排放速率及其对土壤水热变化的响应规律,本研究用壕沟断根法布设了土壤呼吸组分分离试验,并对土壤温湿度与呼吸速率进行了一年的观测。统计分析结果表明：土壤呼吸及其组分的呼吸速率在夏秋季较高、春冬季较低;土壤温度低于15℃时,呼吸速率的季节性变化主要受控于土壤温度;土壤温度高于15℃,而含水量低于0．20kg·kg^-1时,含水量对呼吸速率有明显的抑制作用;当土壤温湿度分别高于15℃与0．20kg·kg^-1,呼吸速率同时受到土壤温湿度的影响;土壤温湿度分别能解释呼吸速率季节性变化的80．36％～94．94％与7．20％～48．45％,温度的影响高于含水量;5种类型中土壤呼吸、自养与异养呼吸的Q10值变化范围分别为2．30～2．44、2．49～2．82与2．09～2．35,每个类型中自养呼吸的温度敏感性均为最高,其次为土壤呼吸,异养呼吸最低;锐齿栎幼林、锐齿栎老林、华山松与短柄袍针阔混交林、千金榆与短柄袍阔叶混交林及栓皮栎林自养呼吸日贡献率的变化范围分别为35．19％～57．73％、28．73％～49．24％、28．67％～49．82％、24．24％～41．70％与30．07％～46．22％,土壤呼吸的年排放量分别为1105．15gC·m^-2·a^-1、779．12gC·m^-2·a^-1、821．23gC·m^-2·a^-1、912．19gC·m^-2·a^-1与899．50gC·m^-2·a^-1,其中自养呼吸的年贡献率分别为52．89％、39．77％、44．17％、38．15％与43．26％,若考虑断根样方内细根分解的影响,则自养呼吸的年贡献率分别为65．56％、47．95％、53．80％、46．83％与53．86％;5个林分间的土壤呼吸速率、异养呼吸速率没有显著差异（P〉0．05）,而自养呼吸速率存在显著差异（P〈0．05）,类型间活细根生物量的差异解释了自养呼吸速率差异的94．71％。     

Partitioning the components of soil respiration: a research challenge

Little is known about the respiratory components of CO₂ emitted from soils and attaining a reliable quantification of the contribution of root respiration remains one of the major challenges facing ecosystem research. Resolving this would provide major advances in our ability to predict ecosystem responses to climate change. The merits and technical and theoretical difficulties associated with different approaches adopted for partitioning respiration components are discussed here. The way forward is suggested to be the development of non-invasive regression analysis validated by stable isotope approaches to increase the sensitivity of model functions to include components of rhizosphere microbial activity, changing root biomass and the dynamics of a wide range of soil C pools. Section Editor: A. Hodge     

降雨量改变对常绿阔叶林干旱和湿润季节土壤呼吸的影响

通过野外原位试验,研究降雨量改变对华西雨屏区常绿阔叶林干旱和湿润季节土壤呼吸速率的影响。采用LI-8100土壤碳通量分析系统(LI-COR Inc.,USA)测定干旱和湿润季节对照(CK)、增雨10%(LA)、增雨5%(TA)、减雨10%(LR)、减雨20%(MR)、减雨50%(HR)6个处理水平的土壤呼吸速率,并通过回归方程分析温度和湿度与土壤呼吸速率间的关系。结果表明:湿润季节土壤呼吸速率高于干旱季节,HR处理对干旱季节土壤呼吸速率影响较大,而LA处理对湿润季节土壤呼吸速率的影响较大。TA和LR处理使土壤呼吸的温度敏感性增加,而HR、LA和MR处理使土壤呼吸的温度敏感性降低,干旱季节Q10值高于湿润季节。各处理湿润季节土壤微生物量碳氮含量显著高于干旱季节,HR、MR和LA处理减少土壤微生物生物量碳、氮的含量,而TA和LR处理增加土壤微生物生物量碳、氮的含量。与湿润季节相比,干旱季节土壤水分对土壤呼吸速率的影响较大;而与土壤温度相比,土壤水分对土壤呼吸速率的影响较小。在降雨量改变的背景下,华西雨屏区常绿阔叶林无论是干旱还是湿润季节,适当增雨和减雨都会促进土壤呼吸速率,而较高量的增雨和减雨会抑制土壤呼吸速率。     

Evaluation of soil respiration and soil CO2 concentration in a lowland moist forest in Panama

Soil gas exchange was investigated in a lowland moist forest in Panama. Soil water table level and soil redox potentials indicate that the soils are not waterlogged. Substantial microspatial variation exists for soil respiration and soil CO₂ concentration. During the rainy season, soil CO₂ at 40 cm below the surface accumulates to 2.3%–4.6% and is correlated with rainfall during the previous two weeks. Temporal changes in soil CO₂ are rapid, large and share similar trends between sampling points. Possible effects of soil CO₂ changes on plant growth or phenology are discussed.     

土壤各组分呼吸区分方法研究进展

土壤呼吸分为自养型呼吸(根呼吸)和异养型呼吸(微生物和动物呼吸),区分各组分呼吸可了解在全球变化条件下土壤碳循环和碳平衡的动态。本文综述了3种主要区分自养呼吸和异养呼吸的方法:①组分法;②根去除术;③同位素法。其中同位素法对根和土壤的影响最小,是最可靠的一种方法;综合各方面考虑,根去除法是最切实可行的方法。     

Effect of elevated atmospheric CO2 concentration on soil and root respiration in winter wheat by using a respiration partitioning chamber

Soil respiration in a cropland is the sum of heterotrophic (mainly microorganisms) and autotrophic (root) respiration. The contribution of both these types to soil respiration needs to be understood to evaluate the effects of environmental change on soil carbon cycling and sequestration. In this paper, the effects of free-air CO₂ enrichment (FACE) on hetero- and autotrophic respiration in a wheat field were differentiated and evaluated by a novel split-root growth and gas collection system. Elevated atmospheric pCO₂ of approximately 200 μmol mol⁻¹ above the ambient pCO₂ significantly increased soil respiration by 15.1 and 14.8% at high nitrogen (HN) and low nitrogen (LN) application rates, respectively. The effect of elevated atmospheric pCO₂ on root respiration was not consistent across the wheat growth stages. Elevated pCO₂ significantly increased and decreased root respiration at the booting-heading stage (middle stage) and the late-filling stage (late stage), respectively, in HN and LN treatments; however, no significant effect was found at the jointing stage (early stage). Thus, the effect of increased pCO₂ on cumulative root respiration for the entire wheat growing season was not significant. Cumulative root respiration accounted for approximately 25–30% of cumulative soil respiration in the entire wheat growing season. Consequently, cumulative microbial respiration (soil respiration minus root respiration) increased by 22.5 and 21.1% due to elevated pCO₂ in HN and LN, respectively. High nitrogen application significantly increased root respiration at the late stage under both elevated pCO₂ and ambient pCO₂; however, no significant effects were found on cumulative soil respiration, root respiration, and microbial respiration. These findings suggest that heterotrophic respiration, which is influenced by increased substrate supplies from the plant to the soil, is the key process to determine C emission from agro-ecosystems with regard to future scenarios of enriched pCO₂.     

三源区分土壤呼吸组分研究

三源区分土壤呼吸组分是指将土壤呼吸区分为纯根呼吸、根际微生物呼吸和土壤有机质呼吸3个部分。土壤有机质呼吸、纯根呼吸和根际微生物呼吸是3种不同的生物学过程,这3种呼吸对环境变化具有不同的响应机制。区分土壤呼吸中由根系引起的自养和异养呼吸组分的研究对定量评价陆地生态系统碳平衡具有重要的意义。论述了三源区分土壤呼吸组分的意义、方法和应用,分析了不同条件下土壤呼吸组分区分的研究结果。实验室纯根和根际微生物呼吸占根源呼吸比重约为45%和55%;野外条件下约为60%和40%。最后对本研究未来的发展方向进行了展望。     

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.     

模拟氮沉降和降雨对华西雨屏区常绿阔叶林土壤呼吸的影响

从2013年12月至2014年11月,通过野外原位试验,对华西雨屏区常绿阔叶林进行了模拟氮沉降和降雨试验,采用LI-8100土壤碳通量分析系统(LI-COR Inc.,USA)测定了对照(CK)、氮沉降(N)、减雨(R)、增雨(W)、氮沉降+减雨(NR)、氮沉降+增雨(NW)6个处理水平的土壤呼吸速率,并通过回归方程分析了温度和湿度与土壤呼吸速率间的关系。结果表明:(1)氮沉降和增雨抑制了常绿阔叶林土壤呼吸速率,减雨促进了常绿阔叶林土壤呼吸速率。(2)减雨使华西雨屏区常绿阔叶林土壤呼吸年通量增加了258 g/m~2,而模拟氮沉降和增雨使华西雨屏区常绿阔叶林土壤呼吸年通量分别减少了321g/m~2和406g/m~2。(3)减雨增加了土壤呼吸的温度敏感性,模拟氮沉降和增雨降低了土壤呼吸的温度敏感性。(4)模拟温度和湿度与土壤呼吸速率间回归方程分析表明,土壤水分对土壤呼吸速率的影响较小。(5)模拟氮沉降和增雨处理减少土壤微生物生物量碳、氮的含量,减雨处理增加了土壤微生物生物量碳、氮的含量。(6)模拟氮沉降和降雨对华西雨屏区土壤CO_2释放的影响未表现出明显的交互作用。     

Separating root and soil microbial contributions to soil respiration: A review of methods and observations

Forest soil respiration is the sum of heterotrophic (microbes, soil fauna) and autotrophic (root) respiration. The contribution of each group needs to be understood to evaluate implications of environmental change on soil carbon cycling and sequestration. Three primary methods have been used to distinguish hetero- versus autotrophic soil respiration including: integration of components contributing to in situ forest soil CO₂ efflux (i.e., litter, roots, soil), comparison of soils with and without root exclusion, and application of stable or radioactive isotope methods. Each approach has advantages and disadvantages, but isotope based methods provide quantitative answers with the least amount of disturbance to the soil and roots. Published data from all methods indicate that root/rhizosphere respiration can account for as little as 10 percent to greater than 90 percent of total in situ soil respiration depending on vegetation type and season of the year. Studies which have integrated percent root contribution to total soil respiration throughout an entire year or growing season show mean values of 45.8 and 60.4 percent for forest and nonforest vegetation, respectively. Such average annual values must be extrapolated with caution, however, because the root contribution to total soil respiration is commonly higher during the growing season and lower during the dormant periods of the year.     

中国森林土壤呼吸模式

通过收集国内62个森林样地的土壤呼吸及相关因子数据,分析中国森林土壤呼吸模式.结果表明,中国森林土壤呼吸年通量与年均气温、年均降水量、年凋落物量和年地上净生产力均呈显著的线性正相关,土壤呼吸的Q10则与年均气温和年均降水量均呈显著的负相关.根系呼吸、枯枝落叶层呼吸与土壤呼吸间均呈显著线性正相关;土壤异养呼吸和枯枝落叶层呼吸与年凋落物量呈显著正相关;土壤异养呼吸与自养呼吸间呈显著的线性正相关.根系呼吸、枯枝落叶层呼吸、矿质土壤呼吸占土壤呼吸的比例均值分别为34.7%、20.2%和50.2%.矿质土壤呼吸所占比例与气温和降水量呈显著负相关,而异养呼吸所占比例则与降水量呈显著负相关.根系呼吸所占比例与根系呼吸之间呈渐近线关系(渐近值为45.9%).     

Partitioning of soil respiration components and evaluating the mycorrhizal contribution to soil respiration in a semiarid grassland

土壤呼吸组分的区分对于理解地下碳循环过程非常重要。而菌根真菌在地下碳循环过程中扮演着重要的角色, 但是有关菌根呼吸在草原生态系统中的研究相对较少。该研究在内蒙古半干旱草原应用深浅环网孔法, 结合浅环、深环(排除根系)和一个带有40 μm孔径窗口的土壤环(排除根系但是有菌根菌丝体)将根和菌丝物理分离, 来区分不同的呼吸组分。结果表明: 异养呼吸对总呼吸的贡献比例为51%, 根呼吸的贡献比例为26%, 菌根呼吸的贡献比例为23%, 菌根呼吸的比例3年变化范围为21%-26%。与国内外研究相比, 此方法提供了一个相对稳定的菌根呼吸测量精度范围, 在草原生态系统中切实可行。对菌根呼吸的准确定量将有助于预测草原生态系统土壤碳释放过程对未来气候变化的响应。     

西双版纳热带季节雨林与橡胶林土壤呼吸的季节变化

采用挖壕沟法与红外气体分析法,研究了西双版纳热带季节雨林和人工橡胶林内土壤呼吸包括根系呼吸、异养呼吸的干湿季动态变化.结果表明：季节雨林内土壤呼吸和异养呼吸速率均显著大于橡胶林(P＜0.01),但根系呼吸差异不显著;土壤温湿度是呼吸速率变化的主要影响因子,季节雨林和橡胶林内土壤呼吸和异养呼吸速率均为雨季＞干热季＞雾凉季,但季节雨林内根系呼吸为雨季＞雾凉季＞干热季,而橡胶林内为雾凉季＞雨季＞干热季;季节雨林内根系呼吸对土壤呼吸的贡献率(29%)小于橡胶林(42%,P＜0.01),而季节雨林内异养呼吸对土壤呼吸的贡献率为71%、橡胶林为58%;当5 cm土壤温度在12 ℃～32 ℃范围内变化时,季节雨林内土壤呼吸及根系呼吸、异养呼吸的Q₁₀值均大于橡胶林,且异养呼吸的Q₁₀值最大而根系呼吸的Q₁₀值最小.     

半干旱草原土壤呼吸组分区分与菌根呼吸的贡献

土壤呼吸组分的区分对于理解地下碳循环过程非常重要。而菌根真菌在地下碳循环过程中扮演着重要的角色, 但是有关菌根呼吸在草原生态系统中的研究相对较少。该研究在内蒙古半干旱草原应用深浅环网孔法, 结合浅环、深环(排除根系)和一个带有40 μm孔径窗口的土壤环(排除根系但是有菌根菌丝体)将根和菌丝物理分离, 来区分不同的呼吸组分。结果表明: 异养呼吸对总呼吸的贡献比例为51%, 根呼吸的贡献比例为26%, 菌根呼吸的贡献比例为23%, 菌根呼吸的比例3年变化范围为21%-26%。与国内外研究相比, 此方法提供了一个相对稳定的菌根呼吸测量精度范围, 在草原生态系统中切实可行。对菌根呼吸的准确定量将有助于预测草原生态系统土壤碳释放过程对未来气候变化的响应。