土壤呼吸作用时空动态变化及其影响机制研究与展望

测定不同陆地生态系统土壤呼吸速率及其时空波动, 阐明其影响因子, 对于全球碳素平衡预算和全球变化潜在效应估计是最为基本的数据。然而, 有关土壤呼吸作用变异性及其影响因素的知识仍存在局限性, 一些关键的过程和机制还有待阐明。该文综述了近年来土壤呼吸作用时空动态规律、影响机制和模拟方面的研究进展, 指出环境因子和生物因子共同驱动着土壤呼吸作用的时间动态变化; 土壤呼吸作用在不同时间尺度上还具有明显的空间异质性, 这主要是植被覆盖、根系分布、主要的环境因素和土壤特性空间分布的异质性造成的。生物因子是影响土壤呼吸作用时空动态变化的主要因素之一。然而, 目前所使用的土壤呼吸作用经验模型通常利用土壤温度、土壤湿度或者两者的交互作用模拟土壤呼吸作用动态变化, 但没有考虑生物因子的影响, 这可能会导致明显的偏差和错误。因此, 为了精确估算土壤呼吸作用, 必须解决土壤呼吸作用小尺度上的空间变异性; 加强不同时间尺度上生物要素对土壤呼吸作用动态变化的影响研究; 除了气候因子外, 土壤呼吸作用经验模型应该纳入生物因子等其它影响因素作为变量, 用以提高模型模拟的正确性和准确性。     

中国农田生态系统土壤呼吸作用研究与展望

 农田生态系统是陆地生态系统的重要组成部分,占全球陆地面积的10.5%,其CO₂排放量占人为温室气体排放量的21%～25%;由于农田生态系统 受到强烈的人为干扰,因此农田生态系统土壤呼吸作用及其影响因素对准确评估陆地生态系统碳收支具有重要的意义。中国是个有悠久历史的 农业大国,不仅为农田土壤作用的研究提供了天然的实验室, 而且中国农田土壤呼吸作用的研究对全球的碳循环研究及碳收支准确评估具有非 常重要的示范与指导意义。该文综述了近10年来中国农田生态系统土壤呼吸作用研究进展,指出水热因子、作物生物学特性和农业管理活动是 造成中国农田生态系统土壤呼吸作用时空变异 的主要因素;作物根系呼吸作用占土壤作用的比例在13%～77.2%之间,存在极大的不确定性;合 理施肥、秸秆还田和免耕有助于农田生态系统土壤呼吸作用减排和固碳。指出了中国农田生态系统土壤呼吸作用拟重点加强不同区域典型农田 生态系统土壤呼吸作用的比较、空间异质性、影响因素模拟及减排对策等方面的研究。     

土壤微生物与根系呼吸作用影响因子分析

土壤呼吸作用作为陆地生态系统碳循环的重要组成部分,是当前碳循环研究中的热点问题.对于土壤呼吸作用主要组成部分土壤微生物呼吸作用和根系呼吸作用影响因子的研究,有助于准确地评估全球碳收支.本文从气候、土壤、植被及地表覆被物、大气CO₂浓度、人为干扰等方面综述了土壤微生物呼吸作用和根系呼吸作用的主导影响因子,指出这些影响因子不仅直接或间接地影响土壤微生物呼吸作用和根系呼吸作用,而且它们之间相互作用、相互影响,且各影响因子的地位和作用会随时空尺度变化发生相应改变.在此基础上,论文提出了未来土壤呼吸作用的研究重点.     

玉米农田土壤呼吸作用的空间异质性及其根系呼吸作用的贡献

基于4月底到9月底东北地区玉米农田土壤呼吸作用全生长季的观测,阐明了土壤呼吸作用的空间异质性特征,综合分析了水热因子、土壤性质、根系生物量及其测定位置对土壤呼吸作用空间异质性的影响,并对生长季中根系呼吸作用占土壤呼吸作用的比例进行了估算。结果表明,在植株尺度上,土壤呼吸作用存在着明显的空间异质性,较高的土壤呼吸速率通常出现在靠近玉米植株的地方。根系生物量的分布格局是影响土壤呼吸作用空间异质性的关键因素。在空间尺度上,土壤呼吸作用与根系生物量呈显著的线性关系,而土壤湿度、土壤有机质、全氮和碳氮比对土壤呼吸作用空间异质性的影响并不显著。通过建立土壤呼吸作用与玉米根系生物量的回归方程,对根系呼吸作用占土壤呼吸作用的比例进行了间接估算。玉米生长季中,根系呼吸作用占土壤呼吸作用的比例在43．1％～63．6％之间波动,均值为54．5％。     

林火对大兴安岭落叶松林土壤性质的短期与长期影响

林火是北方针叶林的一种重要组分,是促进森林生态系统养分循环的重要方式。为探讨林火对土壤的影响,以大兴安岭呼中自然保护区2010和2000年火烧迹地为研究对象,并选择附近未过火区作为对照样地,测定分析了受林火影响的土壤理化及微生物指标。结果表明:与对照区相比,火后1年,兴安落叶松林土壤性质发生了显著变化,即土壤水分、有机层厚度、C/N均显著减小,而土壤pH值、铵氮和硝氮含量则显著增加;同时土壤微生物生物量碳氮含量显著减少;火后11年,土壤理化性质基本与对照间无显著差异,但土壤微生物量仍显著低于对照区。这表明火后土壤的理化性质恢复较快,而火对微生物的影响较为持久。本研究将为火后森林土壤养分的管理提供重要科学依据。     

玉米地土壤呼吸作用对土壤温度和生物因子协同作用的响应

基于2005年玉米(Zea mays)生长季土壤呼吸作用及其影响因子的动态观测资料,分析了玉米地土壤呼吸作用的日和季动态及其对土壤温度和生物因子协同作用的响应。结果表明,玉米地土壤呼吸作用的日变化为不对称的单峰型,其最小值和最大值分别出现在6∶00~7∶00和13∶00左右;玉米生长季中,土壤呼吸速率波动较大,其均值为3.16 μmol CO₂·m^-2·s^-1,最大值为4.87 μmol CO₂·m^-2·s^-1,出现在7月28日,最小值为1.32 μmol CO₂·m^-2·s^-1,出现在5月4日。在土壤呼吸作用日变化中,土壤呼吸速率(SR)与10 cm深度土壤温度(T)呈显著的线性关系:SR=αT+β。在整个生长季节,玉米净初级生产力(NPP)与直线斜率(α)呈显著正相关,生物量(B)也明显影响直线的截距(β)。基于此,建立了玉米地土壤呼吸作用动态模型SR=(aNPP+b)T+cB²+dB+e。土壤呼吸作用季节变化的大部分(97%)可以由土壤温度、NPP和生物量的季节变化来解释。当仅考虑土壤温度对土壤呼吸作用的影响时,指数方程会过大或过小地估计了土壤呼吸强度。该文的结果强调了生物因子在土壤呼吸作用季节变化中的重要作用,同时指出土壤呼吸作用模型不仅要考虑土壤温度的影响,在生物因子影响土壤呼吸作用的温度敏感性时,还应该把生物因子纳入模型。     

土壤呼吸作用普适性评估模型构建的设想

土壤呼吸作用是陆地碳循环中的重要组成部分, 其评估的准确性直接影响到陆地碳源/汇的准确评估. 对国内外土壤呼吸作用最新研究成果的综述表明, 目前土壤呼吸作用研究主要集中于时间变异性及其影响因子方面, 没有研究同一生态系统内部及不同类型生态系统土壤呼吸作用的空间变异性及其影响因素; 土壤呼吸作用估算模型大多亦只考虑了水热因子的影响, 没有发展一个适于不同的时间与空间尺度的、耦合水热-生物-土壤养分综合影响的土壤呼吸作用普适性模型. 为此, 本研究从影响土壤呼吸作用的时间和空间异质性方面提出了构建土壤呼吸作用普适性评估模型的思想与基本框架, 以及未来的研究重点, 以增进对土壤呼吸作用的理解, 提高土壤呼吸作用估算的准确性.     

大兴安岭兴安落叶松(Larix gmelinii)天然林分级木转换特征

通过调查样地,作树干解析,分析了不同结构兴安落叶松天然林分级木（优势木、平均木和被压木）转换特征。研究表明：（1）不同结构的兴安落叶松天然林分级木转换年龄、方向和转换率均不同。兴安落叶松分级木转换率29.4%。分级木中,优势木、平均木和被压木转换率分别35.3%、41.2%、11.8%。分级木转换中,优势木与平均木相互转换比例较高,优势木转平均木占83.3%,平均木转优势木占85.7%;优势木向被压木转换比例仅为16.7%;被压木不能转换成优势木,只能转换成平均木,被压木中无转换占88.2%,在森林经营和抚育采伐中应考虑伐除这些被压木。（2）在林分年龄36～65a范围内,随着林分年龄增大,其转换率呈增加趋势。林分年龄30～39a、50～59a和60～69a时,其转换率分别0、33.3%和46.7%。（3）随着林分密度增加,分级木转换率呈增高趋势。当林分密度小于2500株.hm^-2时,主要于优势木与平均木间转换。当林分密度大于2500株.hm^-2时,才出现其它分级木与被压木相互转换现象。（4）不同林型分级木转换率和转换方向不同。草类-落叶松和杜香-落叶松林分级木转换率分别50%和9.5%。（5）不同水平格局林分分级木转换率不同。聚集分布和随机分布时,其转换率分别61.1%和13.3%。     

大兴安岭北部天然针叶林土壤氮矿化特征

采用顶盖埋管法对大兴安岭地区天然针叶林(樟子松林、樟子松-兴安落叶松混交林和兴安落叶松林)土壤铵态氮(NH~+_4-N)、硝态氮(NO~-_3-N)、净氮矿化速率进行研究,并探索土壤理化性质与氮矿化之间的相关性,为大兴安岭地区森林生态系统土壤养分管理及森林经营提供帮助。结果表明:观测期内(5—10月)3种林型土壤无机氮变化范围为31.51—70.42 mg/kg,以NH~+_4-N形式存在为主,占比达90%以上,且与纯林相比混交林土壤无机氮含量较高。3种林型土壤净氮矿化、净氨化、净硝化速率月变化趋势呈V型,7、8月表现为负值,其他月份为正值。净氮矿化速率变化范围樟子松林为-0.54—1.28 mg kg~(-1) d~(-1)、樟子松-兴安落叶松混交林为-0.13—0.55 mg kg~(-1) d~(-1)、兴安落叶松林为-0.80—1.05 mg kg~(-1) d~(-1)。土壤净氨化过程在土壤氮矿化中占主要地位,占比达60%以上。3种林型土壤净氮矿化、净氨化及净硝化速率垂直差异显著,0—10 cm土层矿化作用明显高于10—20 cm土层(P0.05)。土壤氮矿化速率与土壤含水量、土壤有机碳含量、土壤C/N、枯落物全氮含量和枯落物C/N均存在显著相关性。不同类型的森林土壤及枯落物的质量也存在差异,进而影响土壤氮矿化特征。     

长白山红松针阔混交林与开垦农田土壤呼吸作用比较

利用静态箱式法测定长白山红松(Pinus koraiensis)针阔混交林及其开垦农田的土壤呼吸作用。结果表明,两者土壤呼吸作用的日动态和季节动态均主要受温度影响,农田土壤呼吸作用的日变化极值出现时间较林地提前,最大值出现在12∶00左右,比林地提前6 h左右,最小值在凌晨5∶00左右,早于林地2~3 h;在生长季,土壤呼吸速率与10 cm土壤含水量关系不显著,而与土壤5 cm温度呈显著的指数关系;农田土壤温度高于林地,但在整个生长季(5~9月)林地土壤释放CO₂量(2 674.4 g·m^-2)约为农田(1 285.3 g·m^-2)的2倍;观测期间,农田土壤呼吸速率占林地的比例范围在23.4%~76.3%之间,说明土壤呼吸作用还受不同土地利用方式下植被类型等的影响。农田和红松针阔混交林土壤呼吸作用的Q₁₀值分别为3.07和2.92,农田土壤呼吸作用的Q₁₀ 值估计可能偏大。森林转变为农田后,环境、生物因子以及土壤养分含量和物理性质发生改变,共同影响土壤呼吸作用的强度和动态特征。     

Fertilization of boreal forest reduces both autotrophic and heterotrophic soil respiration

The boreal forest is expected to experience the greatest warming of all forest biomes, raising concerns that some of the large quantities of soil carbon in these systems may be added to the atmosphere as CO₂. However, nitrogen deposition or fertilization has the potential to increase boreal forest production and retard the decomposition of soil organic matter, hence increasing both tree stand and soil C storage. The major contributors to soil‐surface CO₂ effluxes are autotrophic and heterotrophic respiration. To evaluate the effect of nutrient additions on the relative contributions from autotrophic and heterotrophic respiration, a large‐scale girdling experiment was performed in a long‐term nutrient optimization experiment in a 40‐year‐old stand of Norway spruce in northern Sweden. Trees on three nonfertilized plots and three fertilized plots were girdled in early summer 2002, and three nonfertilized and three fertilized plots were used as control plots. Each plot was 0.1 ha and contained around 230 trees. Soil‐surface CO₂ fluxes, soil moisture, and soil temperature were monitored in both girdled and nongirdled plots. In late July, the time of the seasonal maximum in soil‐surface CO₂ efflux, the total soil‐CO₂ efflux in nongirdled plots was 40% lower in the fertilized than in the nonfertilized plots, while the efflux in girdled fertilized and nonfertilized plots was 50% and 60% lower, respectively, than in the corresponding nongirdled controls. We attribute these reductions to losses of the autotrophic component of the total soil‐surface CO₂ efflux. The estimates of autotrophic respiration are conservative as root starch reserves were depleted more rapidly in roots of girdled than in nongirdled trees. Thus, heterotrophic activity was overestimated. Calculated on a unit area basis, both the heterotrophic and autotrophic soil respiration was significantly lower in fertilized plots, which is especially noteworthy given that aboveground production was around three times higher in fertilized than in nonfertilized plots.     

Partitioning sources of soil respiration in boreal black spruce forest using radiocarbon

Separating ecosystem and soil respiration into autotrophic and heterotrophic component sources is necessary for understanding how the net ecosystem exchange of carbon (C) will respond to current and future changes in climate and vegetation. Here, we use an isotope mass balance method based on radiocarbon to partition respiration sources in three mature black spruce forest stands in Alaska. Radiocarbon (Δ¹⁴C) signatures of respired C reflect the age of substrate C and can be used to differentiate source pools within ecosystems. Recently‐fixed C that fuels plant or microbial metabolism has Δ¹⁴C values close to that of current atmospheric CO₂, while C respired from litter and soil organic matter decomposition will reflect the longer residence time of C in plant and soil C pools. Contrary to our expectations, the Δ¹⁴C of C respired by recently excised black spruce roots averaged 14‰ greater than expected for recently fixed photosynthetic products, indicating that some portion of the C fueling root metabolism was derived from C storage pools with turnover times of at least several years. The Δ¹⁴C values of C respired by heterotrophs in laboratory incubations of soil organic matter averaged 60‰ higher than the contemporary atmosphere Δ¹⁴CO₂, indicating that the major contributors to decomposition are derived from a combination of sources consistent with a mean residence time of up to a decade. Comparing autotrophic and heterotrophic Δ¹⁴C end members with measurements of the Δ¹⁴C of total soil respiration, we calculated that 47–63% of soil CO₂ emissions were derived from heterotrophic respiration across all three sites. Our limited temporal sampling also observed no significant differences in the partitioning of soil respiration in the early season compared with the late season. Future work is needed to address the reasons for high Δ¹⁴C values in root respiration and issues of whether this method fully captures the contribution of rhizosphere respiration.     

Microbial activity and soil respiration under nitrogen addition in Alaskan boreal forest

Climate warming could increase rates of soil organic matter turnover and nutrient mineralization, particularly in northern high‐latitude ecosystems. However, the effects of increasing nutrient availability on microbial processes in these ecosystems are poorly understood. To determine how soil microbes respond to nutrient enrichment, we measured microbial biomass, extracellular enzyme activities, soil respiration, and the community composition of active fungi in nitrogen (N) fertilized soils of a boreal forest in central Alaska. We predicted that N addition would suppress fungal activity relative to bacteria, but stimulate carbon (C)‐degrading enzyme activities and soil respiration. Instead, we found no evidence for a suppression of fungal activity, although fungal sporocarp production declined significantly, and the relative abundance of two fungal taxa changed dramatically with N fertilization. Microbial biomass as measured by chloroform fumigation did not respond to fertilization, nor did the ratio of fungi : bacteria as measured by quantitative polymerase chain reaction. However, microbial biomass C : N ratios narrowed significantly from 16.0 ± 1.4 to 5.2 ± 0.3 with fertilization. N fertilization significantly increased the activity of a cellulose‐degrading enzyme and suppressed the activities of protein‐ and chitin‐degrading enzymes but had no effect on soil respiration rates or ¹⁴C signatures. These results indicate that N fertilization alters microbial community composition and allocation to extracellular enzyme production without affecting soil respiration. Thus, our results do not provide evidence for strong microbial feedbacks to the boreal C cycle under climate warming or N addition. However, organic N cycling may decline due to a reduction in the activity of enzymes that target nitrogenous compounds.     

Winter soil respiration during soil-freezing process in a boreal forest in Northeast China

Aims Boreal forest is the largest and contains the most soil carbon among global terrestrial biomes. Soil respiration during the prolonged winter period may play an important role in the carbon cycles in boreal forests. This study aims to explore the characteristics of winter soil respiration in the boreal forest and to show how it is regulated by environmental factors, such as soil temperature, soil moisture and snowpack.Methods Soil respiration in an old-growth larch forest (Larix gmelinii Ruppr.) in Northeast China was intensively measured during the winter soil-freezing process in 2011 using an automated soil CO₂ flux system. The effects of soil temperature, soil moisture and thin snowpack on soil respiration and its temperature sensitivity were investigated.Important findings Total soil respiration and heterotrophic respiration both showed a declining trend during the observation period, and no significant difference was found between soil respiration and heterotrophic respiration until the snowpack exceeded 20cm. Soil respiration was exponentially correlated with soil temperature and its temperature sensitivity (Q 10 value) for the entire measurement duration was 10.5. Snow depth and soil moisture both showed positive effects on the temperature sensitivity of soil respiration. Based on the change in the Q 10 value, we proposed a 'freeze–thaw critical point' hypothesis, which states that the Q 10 value above freeze–thaw critical point is much higher than that below it (16.0 vs. 3.5), and this was probably regulated by the abrupt change in soil water availability during the soil-freezing process. Our findings suggest interactive effects of multiple environmental factors on winter soil respiration and recommend adopting the freeze–thaw critical point to model soil respiration in a changing winter climate.     

Changing sources of soil respiration with time since fire in a boreal forest

Radiocarbon signatures (Δ¹⁴C) of carbon dioxide (CO₂) provide a measure of the age of C being decomposed by microbes or respired by living plants. Over a 2‐year period, we measured Δ¹⁴C of soil respiration and soil CO₂ in boreal forest sites in Canada, which varied primarily in the amount of time since the last stand‐replacing fire. Comparing bulk respiration Δ¹⁴C with Δ¹⁴C of CO₂ evolved in incubations of heterotrophic (decomposing organic horizons) and autotrophic (root and moss) components allowed us to estimate the relative contributions of O horizon decomposition vs. plant sources. Although soil respiration fluxes did not vary greatly, differences in Δ¹⁴C of respired CO₂ indicated marked variation in respiration sources in space and time. The ¹⁴C signature of respired CO₂ respired from O horizon decomposition depended on the age of C substrates. These varied with time since fire, but consistently had Δ¹⁴C greater (averaging ～120‰) than autotrophic respiration. The Δ¹⁴C of autotrophically respired CO₂ in young stands equaled those expected for recent photosynthetic products (70‰ in 2003, 64‰ in 2004). CO₂ respired by black spruce roots in stands >40 years old had Δ¹⁴C up to 30‰ higher than recent photosynthates, indicating a significant contribution of C stored at least several years in plants. Decomposition of O horizon organic matter made up 20% or less of soil respiration in the younger (<40 years since fire) stands, increasing to ～50% in mature stands. This is a minimum for total heterotrophic contribution, since mineral soil CO₂ had Δ¹⁴C close to or less than those we have assigned to autotrophic respiration. Decomposition of old organic matter in mineral soils clearly contributed to soil respiration in younger stands in 2003, a very dry year, when Δ¹⁴C of soil respiration in younger successional stands dropped below those of the atmospheric CO₂.     

Representative estimates of soil and ecosystem respiration in an old beech forest

Respiration has been proposed to be the main determinant of the carbon balance in European forests and is thus essential for our understanding of the carbon cycle. However, the choice of experimental design strongly affects estimates of annual respiration and of the contribution of soil respiration to total ecosystem respiration. In a detailed study of ecosystem and soil respiration fluxes in an old unmanaged deciduous forest in Central Germany over 3 years (2000–2002), we combined soil chamber and eddy covariance measurements to obtain a comprehensive picture of respiration in this forest. The closed portable chambers offered to investigate spatial variability of soil respiration and its controls while the eddy covariance system offered continuous measurements of ecosystem respiration. Over the year, both fluxes were mainly correlated with temperature. However, when soil moisture sank below 23 vol.% in the upper 6 cm, water limitations also became apparent. The temporal resolution of the eddy covariance system revealed that relatively high respiration rates occurred during budbreak due to increased metabolic activity and after leaf fall because of increased decomposition. Spatial variability in soil respiration rates was large and correlated with fine root biomass (r ² = 0.56) resulting in estimates of annual efflux varying across plots from 730 to 1,258 (mean 898) g C m⁻² year⁻¹. Power function calculations showed that achieving a precision in the soil respiration estimate of 20% of the full population mean at a confidence level of 95%, requires about eight sampling locations. Our results can be used as guidelines to improve the representativeness of soil respiration measurements by nested sampling designs, being applied in long-term and large-scale carbon sequestration projects such as FLUXNET and CarboEurope.     

Feedback from soil inorganic nitrogen on soil organic matter mineralisation and growth in a boreal forest ecosystem

Current nitrogen (N) deposition rates are considerably higher than during pre-industrial times and the growing interest in forest fertilisation requires better understanding of how the N and carbon (C) cycles interact. This study is based on experimental data showing how Scots pine (Pinus sylvestris L.) forests respond to single or consecutive pulse doses of N. The data were used to support the implementation of a dynamic feedback mechanism in the Q model, allowing for changes in soil N availability to regulate the rate of decomposer efficiency. Simulations of the long-term effects of slowly increasing N deposition with and without dynamic decomposer efficiency were then compared. Both versions of the model accurately predicted the response of tree growth to N fertilisation. Slowly increasing inputs of N over a century in the modified version acted on the inputs and outputs of soil C in opposing ways: (a) rate of litter input slowed down because more N was retained in the soil and thus not available for tree growth; (b) rate of C output, through soil heterotrophic respiration, was also gradually reduced due to increasing decomposer efficiency, although not enough to sufficiently balance the reduced litter input. Accurate prediction of the amount of added N retained in the ecosystem seems to be one of the key issues for estimating enhanced C sequestration.     

A distinct seasonal pattern of the ratio of soil respiration to total ecosystem respiration in a spruce-dominated forest

Annual budgets and fitted temperature response curves for soil respiration and ecosystem respiration provide useful information for partitioning annual carbon budgets of ecosystems, but they may not adequately reveal seasonal variation in the ratios of these two fluxes. Soil respiration (R_s) typically contributes 30–80% of annual total ecosystem respiration (R_eco) in forests, but the temporal variation of these ratios across seasons has not been investigated. The objective of this study was to investigate seasonal variation in the R_s/R_eco ratio in a mature forest dominated by conifers at Howland, ME, USA. We used chamber measurements of R_s and tower‐based eddy covariance measurements of R_eco. The R_s/R_eco ratio reached a minimum of about 0.45 in the early spring, gradually increased through the late spring and early summer, leveled off at about 0.65 for the summer, and then increased again to about 0.8 in the autumn. A spring pulse of aboveground respiration presumably causes the springtime minimum in this ratio. Soil respiration ‘catches up’ as the soils warm and as root growth presumably accelerates in the late spring, causing the R_s/R_eco ratios to increase. The summertime plateau of R_s/R_eco ratios is consistent with summer drought suppressing R_s that would otherwise be increasing, based on increasing soil temperature alone, thus causing the R_s/R_eco ratios to not increase as soils continue to warm. Declining air temperatures and litter fall apparently contribute to increased R_s/R_eco ratios in the autumn. Differences in phenology of growth of aboveground and belowground plant tissues, mobilization and use of stored substrates within woody plants, seasonal variation in photosynthate and litter substrates, and lags between temperature changes of air and soil contribute to a distinct seasonal pattern of R_s/R_eco ratios.     

森林土壤呼吸研究进展

各种测量森林土壤呼吸的方法都存在不足,红外CO₂分析仪法是目前最理想的方法;土壤CO₂通量模型的优点是考虑了土壤呼吸生物和物理学过程;一般情况下,温度和湿度与森林土壤呼吸呈正相关关系,火烧、采伐和施肥等营林活动对土壤呼吸的影响有很大的不确定性;森林土壤呼吸与植被、微生物生物量的关系,以及土壤呼吸的空间变异规律已成为近年来的研究热点.最后提出了森林土壤呼吸研究中存在的一些问题及今后的发展方向.