Interpreting,measuring, and modeling soil respiration

This paper reviews the role of soil respiration in determining ecosystem carbon balance, and the conceptual basis for measuring and modeling soil respiration. We developed it to provide background and context for this special issue on soil respiration and to synthesize the presentations and discussions at the workshop. Soil respiration is the largest component of ecosystem respiration. Because autotrophic and heterotrophic activity belowground is controlled by substrate availability, soil respiration is strongly linked to plant metabolism, photosynthesis and litterfall. This link dominates both base rates and short-term fluctuations in soil respiration and suggests many roles for soil respiration as an indicator of ecosystem metabolism. However, the strong links between above and belowground processes complicate using soil respiration to understand changes in ecosystem carbon storage. Root and associated mycorrhizal respiration produce roughly half of soil respiration, with much of the remainder derived from decomposition of recently produced root and leaf litter. Changes in the carbon stored in the soil generally contribute little to soil respiration, but these changes, together with shifts in plant carbon allocation, determine ecosystem carbon storage belowground and its exchange with the atmosphere. Identifying the small signal from changes in large, slow carbon pools in flux dominated by decomposition of recent material and autotrophic and mycorrhizal respiration is a significant challenge. A mechanistic understanding of the belowground carbon cycle and of the response of different components to the environment will aid in identifying this signal. Our workshop identified information needs to help build that understanding: (1) the mechanisms that control the coupling of canopy and belowground processes; (2) the responses of root and heterotrophic respiration to environment; (3) plant carbon allocation patterns, particularly in different forest developmental stages, and in response to treatments (warming, CO₂, nitrogen additions); and (4) coupling measurements of soil respiration with aboveground processes and changes in soil carbon. Multi-factor experiments need to be sufficiently long to allow the systems to adjust to the treatments. New technologies will be necessary to reduce uncertainty in estimates of carbon allocation, soil carbon pool sizes, and different responses of roots and microbes to environmental conditions.     

川中丘陵区冬水田种植模式转旱作土壤呼吸组分特征及碳平衡

冬水田-水稻是川中丘陵区传统的稻田种植模式,冬水田种植模式转变是实现多熟种植及机械化的重要途径。为探究冬水田-水稻种植模式转旱作过程中作物季及休闲期土壤呼吸速率及其组分构成,试验设置冬水田-水稻转旱作(FTD)、冬水田-水稻(FR)和冬闲田-玉米(FM)3种不同种植模式,采用根排除法和静态明箱-气相色谱法原位取样测定作物季及季后休闲期土壤呼吸及其组分,并通过测算净生态系统生产力(NEP)进而判断冬水田-水稻转旱作过程的农田系统碳汇强度。结果表明:(1)FTD显著提高了土壤总呼吸速率及其自养和异养呼吸速率,从而提高了其累积排放量(P<0.05)。与FR相比,FTD的土壤总呼吸及其自养和异养呼吸的累积排放量分别提高了13.14倍、11.32倍和15.56倍(P<0.05);与FM相比,FTD的土壤总呼吸及其自养和异养呼吸的累积排放量分别提高了70.56%、40.83%和115.47%(P<0.05)。(2)与FR和FM相比,FTD均降低了土壤呼吸及其组分的温度敏感性(Q₁₀),且土壤总呼吸的温度敏感性介于异养呼吸和自养呼吸之间。(3)FR,FM和FTD的净生态系统生产力(NEP)均为正值,其数值分别为7911.66 kg/hm²,5667.89 kg/hm²和1583.46 kg/hm²,均表现为大气CO₂的碳汇,但与FR与FM相比,FTD显著降低了其净生态系统生产力,呈现出较弱的碳汇。     

Tree-girdling to separate root and heterotrophic respiration in two Eucalyptus stands in Brazil

The release of carbon as CO₂ from belowground processes accounts for about 70% of total ecosystem respiration. Insights about factors controlling soil CO₂ efflux are constrained by the challenge of apportioning sources of CO₂ between autotrophic tree roots (and mycorrhizal fungi) and heterotrophic microorganisms. In some temperate conifer forests, the reduction in soil CO₂ efflux after girdling (phloem removal) has been used to separate these sources. Girdling stops the flow of carbohydrates to the belowground portion of the ecosystem, which should slow respiration by roots and mycorrhizae while heterotrophic respiration should remain constant or be enhanced by the decomposition of newly dead roots. Therefore, the reduction in CO₂ efflux after girdling should be a conservative estimate of the belowground flux of C from trees. We tested this approach in two tropical Eucalyptus plantations. Tree canopies remained intact for more than 3 months after girdling, showing no reduction in light interception. The reduction in soil CO₂ efflux averaged 16–24% for the 3-month period after girdling. The reduction in CO₂ efflux was similar for plots with one half of the trees girdled and those with all of the trees girdled. Girdling did not reduce live fine root biomass for at least 5 months after treatment, indicating that large reserves of carbohydrates in the root systems of Eucalyptus trees maintained the roots and root respiration. Our results suggest that the girdling approach is unlikely to provide useful insights into the contribution of tree roots and heterotrophs to soil CO₂ efflux in this type of forest ecosystem.     

Net ecosystem exchange modifies the relationship between the autotrophic and heterotrophic components of soil respiration with abiotic factors in prairie grasslands

We investigated the relationships of net ecosystem carbon exchange (NEE), soil temperature, and moisture with soil respiration rate and its components at a grassland ecosystem. Stable carbon isotopes were used to separate soil respiration into autotrophic and heterotrophic components within an eddy covariance footprint during the 2008 and 2009 growing seasons. After correction for self‐correlation, rates of soil respiration and its autotrophic and heterotrophic components for both years were found to be strongly influenced by variations in daytime NEE – the amount of C retained in the ecosystem during the daytime, as derived from NEE measurements when photosynthetically active radiation was above 0 μmol m^?2 s^?1. The time scale for correlation of variations in daytime NEE with fluctuations in respiration was longer for heterotrophic respiration (36–42 days) than for autotrophic respiration (4–6 days). In addition to daytime NEE, autotrophic respiration was also sensitive to soil moisture but not soil temperature. In contrast, heterotrophic respiration from soils was sensitive to changes in soil temperature, soil moisture, and daytime NEE. Our results show that – as for forests – plant activity is an important driver of both components of soil respiration in this tallgrass prairie grassland ecosystem. Heterotrophic respiration had a slower coupling with plant activity than did autotrophic respiration. Our findings suggest that the frequently observed variations in the sensitivity of soil respiration to temperature or moisture may stem from variations in the proportions of autotrophic and heterotrophic components of soil respiration. Rates of photosynthesis at seasonal time scales should also be considered as a driver of both autotrophic and heterotrophic soil respiration for ecosystem flux modeling.     

Transpiration alters the contribution of autotrophic and heterotrophic components of soil CO2 efflux

? An unbiased partitioning of autotrophic and heterotrophic components of soil CO(2) efflux is important to estimate forest carbon budgets and soil carbon sequestration. The contribution of autotrophic sources to soil CO(2) efflux (F(A)) may be underestimated during the daytime as a result of internal transport of CO(2) produced by root respiration through the transpiration stream. ? Here, we tested the hypothesis that carbon isotope composition of soil CO(2) efflux (δ(FS)) in a Eucalyptus plantation grown on a C(4) soil is enriched during the daytime, which will indicate a decrease in F(A) during the periods of high transpiration. ? Mean δ(FS) of soil CO(2) efflux decreased to -25.7‰ during the night and increased to -24.7‰ between 11:00 and 15:00 h when the xylem sap flux density was at its maximum. ? Our results indicate a decrease in the contribution of root respiration to soil CO(2) efflux during the day that may be interpreted as a departure of root-produced CO(2) in the transpiration stream, leading to a 17% underestimation of autotrophic contribution to soil CO(2) efflux on a daily timescale.     

Substrate availability regulates the suppressive effects of Canada goldenrod invasion on soil respiration

基质有效性调节加拿大一枝黄花入侵对土壤呼吸的抑制作用 外来植物入侵不仅会降低河边近岸湿地生态系统植被多样性,而且会改变湿地生态系统的地下碳过程。外来入侵植物加拿大一枝黄花(Solidago canadensis L.)已广泛入侵我国东南部地区,但加拿大一枝黄花入侵对入侵地生态系统地下土壤碳循环过程的影响却知之甚少。本研究通过野外原位观测实验和温室模拟入侵实验,探究外来植物加拿大一枝黄花入侵对入侵地土壤呼吸的影响规律及其驱动因素。野 外原位观测实验开展于2018年7月21日至12月15日,期间每周测定样地土壤呼吸。温室模拟入侵实验开展于2019年7月15日至12月15日,期间每月1日与15日上午测定土壤呼吸、自养呼吸和异养呼吸。土壤呼吸、自养呼吸和异养呼吸通过静态箱结合深埋根系隔离法测定。野外原位观测实验和温室模拟入侵实验结果均显示,加拿大一枝黄花的入侵降低了土壤二氧化碳的排放通量。加拿大一枝黄花入侵对土壤呼吸的抑制作用可能归因于其入侵引起的土壤可利用底物质量与数量的变化,表明外来入侵植物加拿大一枝黄花可通过改变植物释放基质以及与本地植物和/或土壤微生物争夺土壤有效基质而影响土壤碳循环。这些研究结果对于评估外来入侵植物对入侵地地下碳动态的影响以及对全球变暖的贡献具有重要意义。     

杉木林年龄序列地下碳分配变化

  森林地下碳分配在森林碳平衡和碳吸存中具有重要作用, 而揭示人工林生长过程中地下碳分配变化对于人工林碳汇估算和碳汇管理等有重要意义。通过采用年龄序列方法研究了杉木(Cunninghamia lanceolata)林生长过程中地下碳分配变化特点。年龄序列为福建省南平7 a生(幼龄林)、16 a生(中龄林)、21 a生(近熟林)、41 a生(成熟林)和88 a生(老龄林)的杉木林。细根净生产力测定采用连续土芯法, 根系呼吸测定采用壕沟法, 生物量增量测定采用异速生长方程, 地上年凋落物量采用凋落物收集框测定。结果表明: 杉木林细根净生产力在中龄林前没有显著差异, 维持在较高水平; 但此后则显著下降。细根净生产力/地上凋落物量比值随林龄增加而显著下降。老龄林的根系呼吸显著低于其它林龄林分, 根系呼吸与细根生物量间呈显著线性相关。中龄林和近成熟林的地下碳分配(Total belouground carbon allocation, TBCA)显著高于幼龄林和成熟林, 而老龄林的则最低。中龄林、近成熟林和成熟林的地上部分净生产力/TBCA比值显著高于幼龄林和老龄林, 而杉木林的根系碳利用效率(RCUE)则呈现出随林龄增加而降低的趋势。     

Components of forest soil CO2 efflux estimated from Δ14C values of soil organic matter

Aims

The partitioning of the total soil CO₂ efflux into its two main components: respiration from roots (and root-associated organisms) and microbial respiration (by means of soil organic matter (SOM) and litter decomposition), is a major need in soil carbon dynamics studies in order to understand if a soil is a net sink or source of carbon.

Methods

The heterotrophic component of the CO₂ efflux was estimated for 11 forest sites as the ratio between the carbon stocks of different SOM pools and previously published (Δ¹⁴C derived) turnover times. The autotrophic component, including root and root-associated respiration, was calculated by subtracting the heterotrophic component from total soil chamber measured CO₂ efflux.

Results

Results suggested that, on average, 50.4 % of total soil CO₂ efflux was derived from the respiration of the living roots, 42.4 % from decomposition of the litter layers and less than 10 % from decomposition of belowground SOM.

Conclusions

The Δ¹⁴C method proved to be an efficient tool by which to partition soil CO₂ efflux and quantify the contribution of the different components of soil respiration. However the average calculated heterotrophic respiration was statistically lower compared with two previous studies dealing with soil CO₂ efflux partitioning (one performed in the same study area; the other a meta-analysis of soil respiration partitioning). These differences were probably due to the heterogeneity of the SOM fraction and to a sub-optimal choice of the litter sampling period.     

Impacts of nitrogen fertilisation and coppicing on total and heterotrophic soil CO2 efflux in a short rotation poplar plantation

Short rotation forests can serve as sources of renewable energy and possibly for soil C storage. However, the high frequency of management practices and the fertilisation could reduce C storage into the soil, by increasing CO₂ emissions and annulling the potential of C sequestration. The objectives of this work were to evaluate the impacts of coppicing and fertilisation on total soil CO₂ efflux, soil heterotrophic processes and consequent changes of soil C storage in a short rotation poplar plantation. Field soil CO₂ efflux, heterotrophic soil CO₂ efflux and soil organic C were compared before and after coppicing. Temporal dynamics of fine root biomass and water-soluble carbon after coppicing were also analysed. Coppicing increased total soil CO₂ efflux by more than 50%, while heterotrophic soil CO₂ efflux remained unchanged. Nevertheless, an increase in total organic carbon was observed as a result of above and belowground litter inputs, as well as root re-growth and exudation. This trend was more evident in fertilised soils due to lower heterotrophic and autotrophic soil CO₂ effluxes. Fertilisation can reduce the increase of CO₂ emissions after coppicing. Although soil organic C storage increased, the accumulation of labile fractions may trigger microbial respiration in the following years.     

Mycorrhizal Status Influences the Rate but not the Temperature Sensitivity of Soil Respiration

Mycorrhizal fungi, which can produce a large portion of total soil respiration, respond strongly to global changes such as elevated CO₂, N-deposition, and land-use change. Predictions of future ecosystem C sequestration hinge on respiration budgets, but the mycorrhizal influence on total soil respiration remains unknown. In this study, sunflowers (Helianthus annuus) were subjected to various mycorrhizal treatments, and their root and soil systems were enclosed in chambers that continuously monitored belowground (root + mycorrhizal + heterotrophic) CO₂ production during plant growth, death, and decomposition. Rhizocosms with high mycorrhizal colonization exhibited higher soil respiration rates as plants matured, an increase that was in proportion to the mycorrhizal stimulation of plant growth. Living mycorrhizal plants behaved like nonmycorrhizal ones in that total rhizocosm respiration had the same relationship to plant mass and the same temperature sensitivity as nonmycorrhizal plants. Upon removal of the shoots though, mycorrhizal plants exhibited the largest relative reduction in respiration resulting in a unique relationship of soil respiration with plant mass. The mycorrhizal influence on heterotrophic respiration merits as much attention from experimenters and modelers as the mycorrhizal contribution to autotrophic respiration.     

Soil Respiration and Belowground Carbon Allocation in Mangrove Forests

Mangrove forests cover large areas of tropical and subtropical coastlines. They provide a wide range of ecosystem services that includes carbon storage in above- and below ground biomass and in soils. Carbon dioxide (CO₂) emissions from soil, or soil respiration is important in the global carbon budget and is sensitive to increasing global temperature. To understand the magnitude of mangrove soil respiration and the influence of forest structure and temperature on the variation in mangrove soil respiration I assessed soil respiration at eleven mangrove sites, ranging from latitude 27°N to 37°S. Mangrove soil respiration was similar to those observed for terrestrial forest soils. Soil respiration was correlated with leaf area index (LAI) and aboveground net primary production (litterfall), which should aid scaling up to regional and global estimates of soil respiration. Using a carbon balance model, total belowground carbon allocation (TBCA) per unit litterfall was similar in tall mangrove forests as observed in terrestrial forests, but in scrub mangrove forests TBCA per unit litter fall was greater than in terrestrial forests, suggesting mangroves allocate a large proportion of their fixed carbon below ground under unfavorable environmental conditions. The response of soil respiration to soil temperature was not a linear function of temperature. At temperatures below 26°C Q10 of mangrove soil respiration was 2.6, similar to that reported for terrestrial forest soils. However in scrub forests soil respiration declined with increasing soil temperature, largely because of reduced canopy cover and enhanced activity of photosynthetic benthic microbial communities.     

Decadal warming causes a consistent and persistent shift from heterotrophic to autotrophic respiration in contrasting permafrost ecosystems

Soil carbon in permafrost ecosystems has the potential to become a major positive feedback to climate change if permafrost thaw increases heterotrophic decomposition. However, warming can also stimulate autotrophic production leading to increased ecosystem carbon storage—a negative climate change feedback. Few studies partitioning ecosystem respiration examine decadal warming effects or compare responses among ecosystems. Here, we first examined how 11 years of warming during different seasons affected autotrophic and heterotrophic respiration in a bryophyte‐dominated peatland in Abisko, Sweden. We used natural abundance radiocarbon to partition ecosystem respiration into autotrophic respiration, associated with production, and heterotrophic decomposition. Summertime warming decreased the age of carbon respired by the ecosystem due to increased proportional contributions from autotrophic and young soil respiration and decreased proportional contributions from old soil. Summertime warming's large effect was due to not only warmer air temperatures during the growing season, but also to warmer deep soils year‐round. Second, we compared ecosystem respiration responses between two contrasting ecosystems, the Abisko peatland and a tussock‐dominated tundra in Healy, Alaska. Each ecosystem had two different timescales of warming (<5 years and over a decade). Despite the Abisko peatland having greater ecosystem respiration and larger contributions from heterotrophic respiration than the Healy tundra, both systems responded consistently to short‐ and long‐term warming with increased respiration, increased autotrophic contributions to ecosystem respiration, and increased ratios of autotrophic to heterotrophic respiration. We did not detect an increase in old soil carbon losses with warming at either site. If increased autotrophic respiration is balanced by increased primary production, as is the case in the Healy tundra, warming will not cause these ecosystems to become growing season carbon sources. Warming instead causes a persistent shift from heterotrophic to more autotrophic control of the growing season carbon cycle in these carbon‐rich permafrost ecosystems.     

What controls variation in carbon use efficiency among Amazonian tropical forests?

Why do some forests produce biomass more efficiently than others? Variations in Carbon Use Efficiency (CUE: total Net Primary Production (NPP)/ Gross Primary Production (GPP)) may be due to changes in wood residence time (Biomass/NPP_wood), temperature, or soil nutrient status. We tested these hypotheses in 14, one ha plots across Amazonian and Andean forests where we measured most key components of net primary production (NPP: wood, fine roots, and leaves) and autotrophic respiration (R_a; wood, rhizosphere, and leaf respiration). We found that lower fertility sites were less efficient at producing biomass and had higher rhizosphere respiration, indicating increased carbon allocation to belowground components. We then compared wood respiration to wood growth and rhizosphere respiration to fine root growth and found that forests with residence times <40 yrs had significantly lower maintenance respiration for both wood and fine roots than forests with residence times >40 yrs. A comparison of rhizosphere respiration to fine root growth showed that rhizosphere growth respiration was significantly greater at low fertility sites. Overall, we found that Amazonian forests produce biomass less efficiently in stands with residence times >40 yrs and in stands with lower fertility, but changes to long‐term mean annual temperatures do not impact CUE.     

增温和刈割对高寒草甸土壤呼吸及其组分的影响

评估土壤呼吸及其组分对增温等全球变化的响应对于预测陆地生态系统碳循环至关重要。本研究利用红外线辐射加热器(Infrared heater)装置在青藏高原高寒草甸生态系统设置增温和刈割野外控制实验。通过测定2018年生长季(5—9月)土壤呼吸和异养呼吸,探究增温和刈割对土壤呼吸及其组分的影响。研究结果表明：(1) 单独增温使土壤呼吸显著增加31.65% (P<0.05),异养呼吸显著增加27.12% (P<0.05),土壤自养呼吸没有显著改变(P>0.05);单独刈割对土壤呼吸和自养呼吸没有显著影响(P>0.05),单独刈割刺激异养呼吸增加32.54% (P<0.05);(2) 增温和刈割之间的交互作用对土壤呼吸和异养呼吸没有显著影响(P>0.05),但是对自养呼吸的影响是显著的(P<0.05),土壤呼吸和异养呼吸的季节效应显著(P<0.05);(3)土壤呼吸及其组分与土壤温度均成显著指数关系,与土壤湿度呈显著的正相关关系(P<0.05),处理影响它们的响应敏感性。本研究表明青藏高原东缘高寒草甸土壤碳排放与气候变暖存在正反馈。     

寒温带岛状林沼泽土壤呼吸速率和季节变化

2011年生长季内利用静态箱-气相色谱法,研究了寒温带典型湿地白桦(Betula platyphylla)岛状林沼泽、兴安落叶松(Larix gmelinii)岛状林沼泽土壤呼吸速率的季节动态及其主要环境因子,利用壕沟隔断法对土壤呼吸各组分间的差异进行研究。结果表明:生长季白桦和兴安落叶松岛状林沼泽土壤呼吸速率具有明显的季节性规律,土壤呼吸总速率分别为368.60和312.46 mg m-2h-1,异养呼吸速率分别为300.57和215.70 mg m-2h-1,占土壤呼吸总速率的81.5%和69.0%;自养呼吸速率为68.03和96.76 mg m-2h-1,占土壤呼吸总速率的18.5%和31.0%。不同处理条件下的土壤呼吸在季节变化上表现基本一致,高峰期都发生在夏季;土壤呼吸与温度呈极显著相关性,但与土壤湿度的相关性较差。生长季白桦和兴安落叶松岛状林沼泽土壤呼吸总量分别为12.64和10.61 t/hm2。     

若尔盖高寒草本沼泽土壤呼吸对水位下降的响应

土壤呼吸会影响全球碳循环,而湿地水位与土壤呼吸息息相关。然而,由于原位观测有限,目前尚不清楚高寒沼泽土壤呼吸及其组分如何响应水位下降。在若尔盖高原纳勒乔沼泽建立了水位下降控制实验平台,定位监测土壤呼吸及其组分的变化,并初步探讨土壤呼吸及其组分与生物和非生物因素的潜在联系。结果发现,水位下降对高寒草本沼泽土壤呼吸（Rs）没有显著影响,但自养呼吸（Ra）和异氧呼吸（Rh）对水位下降表现出明显不同反应。其中,自养呼吸速率下降了67.2%,异养呼吸速率上升了67.3%。异养呼吸和自养呼吸在土壤呼吸中的占比发生显著变化,水位下降后,Rh/Rs较对照增加了88%,Ra/Rs减少了61%。水位下降引起的自养呼吸和异养呼吸变化的驱动因素不同,植株高度、地上及地下生物量解释了自养呼吸的变化,土壤温度、C：N则是异氧呼吸变化的关键影响因素。综上,在高寒草本沼泽生态系统中,水位下降对土壤呼吸组分的影响强度及其驱动因素存在明显差异,这需要在陆地表层碳循环模型中加以考虑,以便更好评估高寒草本沼泽碳循环对气候变化的反馈作用。     

Partitioning of belowground C in young sugar maple forest

Background and aims

Trees allocate a high proportion of assimilated carbon belowground, but the partitioning of that C among ecosystem components is poorly understood thereby limiting our ability to predict responses of forest C dynamics to global change drivers.

Methods

We labeled sugar maple saplings in natural forest with a pulse of photosynthetic ¹³C in late summer and traced the pulse over the following 3 years. We quantified the fate of belowground carbon by measuring ¹³C enrichment of roots, rhizosphere soil, soil respiration, soil aggregates and microbial biomass.

Results

The pulse of ¹³C contributed strongly to root and rhizosphere respiration for over a year, and respiration comprised about 75 % of total belowground C allocation (TBCA) in the first year. We estimate that rhizosphere carbon flux (RCF) during the dormant season comprises at least 6 % of TBCA. After 3 years, 3.8 % of the C allocated belowground was recovered in soil organic matter, mostly in water-stable aggregates.

Conclusions

A pulse of carbon allocated belowground in temperate forest supplies root respiration, root growth and RCF throughout the following year and a small proportion becomes stabilized in soil aggregates.     

Thawing permafrost increases old soil and autotrophic respiration in tundra: Partitioning ecosystem respiration using δ13C and ∆14C

Ecosystem respiration (R_eco) is one of the largest terrestrial carbon (C) fluxes. The effect of climate change on R_eco depends on the responses of its autotrophic and heterotrophic components. How autotrophic and heterotrophic respiration sources respond to climate change is especially important in ecosystems underlain by permafrost. Permafrost ecosystems contain vast stores of soil C (1672 Pg) and are located in northern latitudes where climate change is accelerated. Warming will cause a positive feedback to climate change if heterotrophic respiration increases without corresponding increases in primary production. We quantified the response of autotrophic and heterotrophic respiration to permafrost thaw across the 2008 and 2009 growing seasons. We partitioned R_eco using Δ¹⁴C and δ¹³C into four sources–two autotrophic (above – and belowground plant structures) and two heterotrophic (young and old soil). We sampled the Δ¹⁴C and δ¹³C of sources using incubations and the Δ¹⁴C and δ¹³C of R_eco using field measurements. We then used a Bayesian mixing model to solve for the most likely contributions of each source to R_eco. Autotrophic respiration ranged from 40 to 70% of R_eco and was greatest at the height of the growing season. Old soil heterotrophic respiration ranged from 6 to 18% of R_eco and was greatest where permafrost thaw was deepest. Overall, growing season fluxes of autotrophic and old soil heterotrophic respiration increased as permafrost thaw deepened. Areas with greater thaw also had the greatest primary production. Warming in permafrost ecosystems therefore leads to increased plant and old soil respiration that is initially compensated by increased net primary productivity. However, barring large shifts in plant community composition, future increases in old soil respiration will likely outpace productivity, resulting in a positive feedback to climate change.     

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.     

Variations of root and heterotrophic respiration along environmental gradients in China's forests

Aims Root and heterotrophic respiration may respond differently to environmental variability, but little evidence is available from large-scale observations. Here we aimed to examine variations of root and heterotrophic respiration across broad geographic, climatic, soil and biotic gradients.Methods We conducted a synthesis of 59 field measurements on root and heterotrophic respiration across China's forests.Important findings Root and heterotrophic respiration varied differently with forest types, of which evergreen broadleaf forest was significantly different from those in other forest types on heterotrophic respiration but without statistically significant differences on root respiration. The results also indicated that root and heterotrophic respiration exhibited similar trends along gradients of precipitation, soil organic carbon and satellite-indicated vegetation growth. However, they exhibited different relationships with temperature: root respiration exhibited bimodal patterns along the temperature gradient, while heterotrophic respiration increased monotonically with temperature. Moreover, they showed different relationships with MOD17 GPP, with increasing trend observed for root respiration whereas insignificant change for heterotrophic respiration. In addition, root and heterotrophic respiration exhibited different changes along the age sequence, with insignificant change for root respiration and decreasing trend for heterotrophic respiration. Overall, these results suggest that root and heterotrophic respiration may respond differently to environmental variability. Our findings could advance our understanding on the different environmental controls of root and heterotrophic respiration and also improve our ability to predict soil CO₂ flux under a changing environment.