积雪和冻结土壤系统中的微生物碳排放和碳氮循环的季节性特征

土壤微生物作为生态系统中重要的分解者,在对动植物残体以及土壤有机质降解的过程中,一方面释放CO2到大气中,是土壤碳排放的重要组成部分;另一方面,在分解的过程中,形成了可供给植物利用的无机养分.由于温度对代谢活动的直接影响,过去对微生物代谢的研究主要集中在生长季,通常假设冬季土壤微生物的活力可以忽略.陆地表面近60％的区域经历着季节性积雪覆盖和季节性土壤冻结的影响.近年来的研究表明,由于积雪的覆盖,形成很好的绝缘层,雪被下土壤中微生物仍然具有显著的活性,对土壤碳排放和植物的养分吸收具有重要的贡献.本文就积雪和冻结土壤系统中的微生物碳排放和碳氮循环的季节性特征进行了全面的分析,综述了国内外冬季雪下碳氮循环的研究现状,提出了目前研究中存在的问题和未来的研究方向,强调了开展温带冬季雪下土壤微生物碳氮循环研究的必要性和重要性.     

Biotic degradation at night,abiotic degradation at day: positive feedbacks on litter decomposition in drylands

The arid and semi‐arid drylands of the world are increasingly recognized for their role in the terrestrial net carbon dioxide (CO₂) uptake, which depends largely on plant litter decomposition and the subsequent release of CO₂ back to the atmosphere. Observed decomposition rates in drylands are higher than predictions by biogeochemical models, which are traditionally based on microbial (biotic) degradation enabled by precipitation as the main mechanism of litter decomposition. Consequently, recent research in drylands has focused on abiotic mechanisms, mainly photochemical and thermal degradation, but they only partly explain litter decomposition under dry conditions, suggesting the operation of an additional mechanism. Here we show that in the absence of precipitation, absorption of dew and water vapor by litter in the field enables microbial degradation at night. By experimentally manipulating solar irradiance and nighttime air humidity, we estimated that most of the litter CO₂ efflux and decay occurring in the dry season was due to nighttime microbial degradation, with considerable additional contributions from photochemical and thermal degradation during the daytime. In a complementary study, at three sites across the Mediterranean Basin, litter CO₂ efflux was largely explained by litter moisture driving microbial degradation and ultraviolet radiation driving photodegradation. We further observed mutual enhancement of microbial activity and photodegradation at a daily scale. Identifying the interplay of decay mechanisms enhances our understanding of carbon turnover in drylands, which should improve the predictions of the long‐term trend of global carbon sequestration.     

Carbon uptake in Eurasian boreal forests dominates the high-latitude net ecosystem carbon budget

Arctic-boreal landscapes are experiencing profound warming, along with changes in ecosystem moisture status and disturbance from fire. This region is of global importance in terms of carbon feedbacks to climate, yet the sign (sink or source) and magnitude of the Arctic-boreal carbon budget within recent years remains highly uncertain. Here, we provide new estimates of recent (2003–2015) vegetation gross primary productivity (GPP), ecosystem respiration (R_eco), net ecosystem CO₂ exchange (NEE; R_eco − GPP), and terrestrial methane (CH₄) emissions for the Arctic-boreal zone using a satellite data-driven process-model for northern ecosystems (TCFM-Arctic), calibrated and evaluated using measurements from >60 tower eddy covariance (EC) sites. We used TCFM-Arctic to obtain daily 1-km² flux estimates and annual carbon budgets for the pan-Arctic-boreal region. Across the domain, the model indicated an overall average NEE sink of −850 Tg CO₂-C year⁻¹. Eurasian boreal zones, especially those in Siberia, contributed to a majority of the net sink. In contrast, the tundra biome was relatively carbon neutral (ranging from small sink to source). Regional CH₄ emissions from tundra and boreal wetlands (not accounting for aquatic CH₄) were estimated at 35 Tg CH₄-C year⁻¹. Accounting for additional emissions from open water aquatic bodies and from fire, using available estimates from the literature, reduced the total regional NEE sink by 21% and shifted many far northern tundra landscapes, and some boreal forests, to a net carbon source. This assessment, based on in situ observations and models, improves our understanding of the high-latitude carbon status and also indicates a continued need for integrated site-to-regional assessments to monitor the vulnerability of these ecosystems to climate change.     

大气二氧化碳浓度升高对植物根际微生物及菌根共生体的影响

综述了大气CO2浓度升高条件下,植物根际土壤环境、根际土壤微生物和植物菌根形成的变化趋势等方面的研究进展,CO2浓度升高,运转到根系的碳水化合物增加,根际环境、根际微生物活性、微生物群落结构以及菌根共生体的形成发生变化．提出在CO2浓度升高条件下,根际微生物和菌根真菌群落的变化对植物群落和陆地生态系统碳动态的调节是今后的研究趋向。     

Structure of microbial communities in <Emphasis Type="Italic">Sphagnum</Emphasis> peatlands and effect of atmospheric carbon dioxide enrichment

Little is known about the structure of microbial communities in Sphagnum peatlands, and the potential effects of the increasing atmospheric CO2 concentration on these communities are not known. We analyzed the structure of microbial communities in five Sphagnum-dominated peatlands across Europe and their response to CO2 enrichment using miniFACE systems. After three growing seasons, Sphagnum samples were analyzed for heterotrophic bacteria, cyanobacteria, microalgae, heterotrophic flagellates, ciliates, testate amoebae, fungi, nematodes, and rotifers. Heterotrophic organisms dominated the microbial communities and together represented 78% to 97% of the total microbial biomass. Testate amoebae dominated the protozoan biomass. A canonical correspondence analysis revealed a significant correlation between the microbial community data and four environmental variables (Na+, DOC, water table depth, and DIN), reflecting continentality, hydrology, and nitrogen deposition gradients. Carbon dioxide enrichment modified the structure of microbial communities, but total microbial biomass was unaffected. The biomass of heterotrophic bacteria increased by 48%, and the biomass of testate amoebae decreased by 13%. These results contrast with the absence of overall effect on methane production or on the vegetation, but are in line with an increased below-ground vascular plant biomass at the same sites. We interpret the increase in bacterial biomass as a response to a CO2-induced enhancement of Sphagnum exudation. The causes for the decrease of testate amoebae are unclear but could indicate a top-down rather than a bottom-up control on their density.     

Environmental effects on CO2 efflux from riparian tundra in the northern foothills of the Brooks Range,Alaska, USA

Summary Carbon dioxide efflux and soil microenvironmental factors were measured diurnally in Carex aquatilus-and Eriophorum angustifolium-dominated riparian tundra communities to determine the relative importance of soil environmental factors controlling ecosystem carbon dioxide exchange with the atmosphere. Measurements were made weekly between 18 June and 24 July 1990. Diurnal patterns in carbon dioxide efflux were best explained by changes in soil temperature, while seasonal changes in efflux were correlated with changes in depth to water table, depth to frozen soil and soil moisture. Carbon dioxide efflux rates were lowest early in the growing season when high water tables and low soil temperatures limited microbial and root activity. Individual rainfall events that raised the water table were found to strongly reduce carbon dioxide efflux. As the growing season progressed, rainfall was low and depth to water table and soil temperatures increased. In response, carbon dioxide efflux increased strongly, attaining rates late in the season of approximately 10 g CO₂ m^–2 day^–1. These rates are as high as maxima recorded for other arctic sites. A mathematical model is developed which demonstrates that soil temperature and depth to water table may be used as efficient predictors of ecosystem CO₂ efflux in this habitat. In parallel with the field measurements of CO₂ efflux, microbial respiration was studied in the laboratory as a function of temperature and water content. Estimates of microbial respiration per square meter under field conditions were made by adjusting for potential respiring soil volume as water table changed and using measured soil temperatures. The results indicate that the effect of these factors on microbial respiration may explain a large part of the diurnal and seasonal variation observed in CO₂ efflux. As in coastal tundra sites, environmental changes that alter water table depth in riparian tundra communities will have large effects on ecosystem CO₂ efflux and carbon balance.     

温度、pH对非光合微生物菌群固碳效率的影响及其成因分析

CO2是主要的温室气体,也是地球上最丰富的碳源和可再生资源,本实验通过研究温度和pH对非光合固碳微生物固碳效率的影响,以期获得非光合固碳微生物适宜的固碳条件,并通过16SrDNA序列分析,研究了不同条件对固碳微生物群落结构的影响及其与固碳效率的相关性。结果表明,好氧／厌氧固碳微生物菌群生长的最适pH均在7左右。好氧固碳微生物菌群可在较大的温度范围（10℃-40℃）内良好生长,但其固碳速率在10℃左右时最高,之后微生物固碳速率随温度上升而下降,在40℃时,微生物固碳速率开始回升。厌氧时微生物的固碳效率随温度的变化情况与好氧时相似,但总体效果要比其在好氧时低20％～25％。由DGGE图谱及戴斯系数分析可得培养温度和pH对群落结构有显著影响,不同pH和温度条件下优势菌差异显著,这可能是固碳效率变化的重要原因之一。     

Effect of Elevated CO2 and Drought on Soil Microbial Communities Associated with Andropogon gerardii

Our understanding of the effects of elevated atmospheric CO2, singly and In combination with other environmental changes,on plant-soil interactions is incomplete. Elevated CO2 effects on C4 plants, though smaller than on C3 species, are mediated mostly via decreased stomatal conductance and thus water loss. Therefore, we characterized the interactive effect of elevated CO2 and drought on soil microbial communities associated with a dominant C4 prairie grass, Andropogon gerardii Vitman. Elevated CO2 and drought both affected resources available to the soil microbial community. For example, elevated CO2 increased the soil C:N ratio and water content during drought, whereas drought alone decreased both. Drought significantly decreased soil microbial biomass. In contrast, elevated COz increased biomass while ameliorating biomass decreases that were induced under drought. Total and active direct bacterial counts and carbon substrate use (overall use and number of used sources) increased significantly under elevated CO2. Denaturing gradient gel electrophoresis analysis revealed that drought and elevated CO2, singly and combined, did not affect the soil bacteria community structure.We conclude that elevated CO2 alone increased bacterial abundance and microbial activity and carbon use, probably in response to increased root exudation. Elevated CO2 also limited drought-related impacts on microbial activity and biomass,which likely resulted from decreased plant water use under elevated CO2. These are among the first results showing that elevated CO2 and drought work in opposition to modulate plant-associated soil-bacteria responses,which should then Influence soil resources and plant and ecosystem function.     

Carbon dioxide supersaturation promotes primary production in lakes

A majority of the world's lakes are supersaturated with respect to carbon dioxide (CO(2) ). By experimental manipulation of the CO(2) concentration in supersaturated boreal lakes, we demonstrate that phytoplankton primary production was up to 10 times higher in supersaturated lake water in comparison with water with CO(2) at equilibrium concentrations and that CO(2) , together with nutrients, explained most of the variation in pelagic primary production and phytoplankton biomass over a wide variety of unproductive lakes. These results suggest that phytoplankton can be co-limited by CO(2) and nutrients in unproductive lakes. As import of terrestrial organic carbon and its subsequent microbial mineralisation in lakes is a driving force of CO(2) -supersaturation our results suggest that lake productivity and carbon cycling may respond to variations in terrestrial organic carbon export, (e.g. caused by land use or climate change) in ways not described before.     

Carbon and nitrogen allocation in Lolium perenne in response to elevated atmospheric CO2 with emphasis on soil carbon dynamics

The effect of elevated CO2 on the carbon and nitrogen distribution within perennial ryegrass (L. perenne L.) and its influence on belowground processes were investigated. Plants were homogeneously 14C-labelled in two ESPAS growth chambers in a continuous 14C-CO2 atmosphere of 350 and 700 L L-1 CO2 and at two soil nitrogen regimes, in order to follow the carbon flow through all plant and soil compartments.After 79 days, elevated CO2 increased the total carbon uptake by 41 and 21% at low (LN) and high nitrogen (HN) fertilisation, respectively. Shoot growth remained unaffected, whereas CO2 enrichment stimulated root growth by 46% and the root/soil respiration by 111%, irrespective of the nitrogen concentration. The total 14C-soil content increased by 101 and 28% at LN and HN, respectively. The decomposition of the native soil organic matter was not affected either by CO2 or by the nitrogen treatment.Elevated CO2 did not change the total nitrogen uptake of the plant either at LN or at HN. Both at LN and HN elevated CO2 significantly increased the total amount of nitrogen taken up by the roots and decreased the absolute and relative amounts translocated to the shoots.The amount of soil nitrogen immobilised by micro-organisms and the size of the soil microbial biomass were not affected by elevated CO2, whereas both were significantly increased at the higher soil N content.Most striking was the 88% increase in net carbon input into the soil expressed as: 14C-roots plus total 14C-soil content minus the 12C-carbon released by decomposition of native soil organic matter. The net carbon input into the soil at ambient CO2 corresponded with 841 and 1662 kg ha-1 at LN and HN, respectively. Elevated CO2 increased these amounts with an extra carbon input of 950 and 1056 kg ha-1. Combined with a reduced decomposition rate of plant material grown at elevated CO2 this will probably lead to carbon storage in grassland soils resulting in a negative feed back on the increasing CO2 concentration of the atmosphere.     

Radioisotopic tracing of carbon monoxide conversion by anaerobic thermophilic prokaryotes

The rate of CO conversion by a pure culture of a thermophilic CO-oxidizing, H₂-producing bacterium Carboxydocella sp. strain 1503 was determined by the radioisotopic method. The overall daily uptake of ¹⁴CO by the bacterium was estimated at 38–56 μmol CO per 1 ml of the culture. A radioisotopic method was developed to separate and quantitatively determine the products of anaerobic CO conversion by microbial communities in hot springs. The new method was first tested on the microbial community from a sample obtained from a hot spring in Kamchatka. The potential rate of CO conversion by the anaerobic microbial community was found to be 40.75 nmol CO/cm³ sediment per day. 85% of the utilized ¹⁴CO was oxidized to carbon dioxide; 14.5% was incorporated into dissolved organic matter, including 0.2% that went into volatile fatty acids; 0.5% was used for cell biomass production; and only just over 0.001% was converted to methane.     

Elevated carbon dioxide accelerates the spatial turnover of soil microbial communities

Although elevated CO₂ (eCO₂) significantly affects the α‐diversity, composition, function, interaction and dynamics of soil microbial communities at the local scale, little is known about eCO₂ impacts on the geographic distribution of micro‐organisms regionally or globally. Here, we examined the β‐diversity of 110 soil microbial communities across six free air CO₂ enrichment (FACE) experimental sites using a high‐throughput functional gene array. The β‐diversity of soil microbial communities was significantly (P < 0.05) correlated with geographic distance under both CO₂ conditions, but declined significantly (P < 0.05) faster at eCO₂ with a slope of ?0.0250 than at ambient CO₂ (aCO₂) with a slope of ?0.0231 although it varied within each individual site, indicating that the spatial turnover rate of soil microbial communities was accelerated under eCO₂ at a larger geographic scale (e.g. regionally). Both distance and soil properties significantly (P < 0.05) contributed to the observed microbial β‐diversity. This study provides new hypotheses for further understanding their assembly mechanisms that may be especially important as global CO₂ continues to increase.     

外源碳输入对陆地生态系统碳循环关键过程的影响及其微生物学驱动机制

外源碳输入对土壤碳源可利用性的改变不仅直接影响着微生物参与陆地生态系统的碳循环过程,而且也制约着微生物对其它营养元素的需求。在大气氮沉降持续增加的全球变化背景下,部分地区已出现生态系统氮养分条件的显著变化甚至土壤中活性氮素的过量积累,进而带来微生物对碳源需求的增加。通过人为调控碳源的可利用性,改善微生物的碳限制状况,将对科学的增加陆地生态系统固碳能力具有极为重大的意义。综述了国内外有关外源碳输入对土壤碳排放、凋落物分解以及土壤碳库影响及其主要的微生物作用机制的相关研究结果,以期能够为未来氮沉降持续增加情景下,如何科学有效地提高生态系统的碳汇潜力提供一定的参考。     

Microbial functional diversity covaries with permafrost thaw‐induced environmental heterogeneity in tundra soil

Permafrost soil in high latitude tundra is one of the largest terrestrial carbon (C) stocks and is highly sensitive to climate warming. Understanding microbial responses to warming‐induced environmental changes is critical to evaluating their influences on soil biogeochemical cycles. In this study, a functional gene array (i.e., geochip 4.2) was used to analyze the functional capacities of soil microbial communities collected from a naturally degrading permafrost region in Central Alaska. Varied thaw history was reported to be the main driver of soil and plant differences across a gradient of minimally, moderately, and extensively thawed sites. Compared with the minimally thawed site, the number of detected functional gene probes across the 15–65 cm depth profile at the moderately and extensively thawed sites decreased by 25% and 5%, while the community functional gene β‐diversity increased by 34% and 45%, respectively, revealing decreased functional gene richness but increased community heterogeneity along the thaw progression. Particularly, the moderately thawed site contained microbial communities with the highest abundances of many genes involved in prokaryotic C degradation, ammonification, and nitrification processes, but lower abundances of fungal C decomposition and anaerobic‐related genes. Significant correlations were observed between functional gene abundance and vascular plant primary productivity, suggesting that plant growth and species composition could be co‐evolving traits together with microbial community composition. Altogether, this study reveals the complex responses of microbial functional potentials to thaw‐related soil and plant changes and provides information on potential microbially mediated biogeochemical cycles in tundra ecosystems.     

Changes in biocrust cover drive carbon cycle responses to climate change in drylands

Dryland ecosystems account for ca. 27% of global soil organic carbon (C) reserves, yet it is largely unknown how climate change will impact C cycling and storage in these areas. In drylands, soil C concentrates at the surface, making it particularly sensitive to the activity of organisms inhabiting the soil uppermost levels, such as communities dominated by lichens, mosses, bacteria and fungi (biocrusts). We conducted a full factorial warming and rainfall exclusion experiment at two semiarid sites in Spain to show how an average increase of air temperature of 2–3 °C promoted a drastic reduction in biocrust cover (ca. 44% in 4 years). Warming significantly increased soil CO₂ efflux, and reduced soil net CO₂ uptake, in biocrust‐dominated microsites. Losses of biocrust cover with warming through time were paralleled by increases in recalcitrant C sources, such as aromatic compounds, and in the abundance of fungi relative to bacteria. The dramatic reduction in biocrust cover with warming will lessen the capacity of drylands to sequester atmospheric CO₂. This decrease may act synergistically with other warming‐induced effects, such as the increase in soil CO₂ efflux and the changes in microbial communities to alter C cycling in drylands, and to reduce soil C stocks in the mid to long term.     

海拔梯度变化对中亚热带黄山松土壤微生物生物量和群落结构的影响

全球变暖对陆地生态系统造成一系列生态问题,使这些问题将随着全球平均气温的升高而进一步加剧。海拔梯度变化是研究气候变暖对陆地生态系统影响的一种重要手段。目前为止利用海拔梯度对微生物影响的研究尚未定论,其主要原因是忽略了植被类型的影响。因此,以中亚热带戴云山的3个海拔(1300、1450、1600 m)的黄山松(Pinus taiwanensis)林为研究对象,探究沿海拔梯度的变化,森林土壤微生物生物量和微生物群落结构的响应变化。结果表明:土壤碳氮磷养分(SOC、TN、TP)、微生物生物量氮(MBN)、微生物生物量磷(MBP)和丛枝菌根真菌(AMF)、革兰氏阴性菌(GN)、真菌(Fungi)、总磷脂脂肪酸(T_(PLFA)),细菌∶真菌(F∶B)均随海拔升高显著下降,而革兰氏阳性菌∶革兰氏阴性菌(GP∶GN)随海拔升高呈相反的趋势。冗余分析(RDA)表明,温度(T)和可溶性有机氮(DON)是影响微生物群落结构的最重要的环境因子。研究表明:与1600 m海拔相比,1300 m海拔温度较高,土壤有机质矿化作用较强,土壤速效养分及微生物生物量随之增加,从而提高(Fungi)、细菌(Bacteria)等。因此,未来气候变暖将通过改变土壤碳氮磷养分来影响本区域微生物群落组成结构。这对进一步深入了解气候变化对山地生态系统土壤养分循环过程具有重要意义。     

Enhancing soil carbon storage for carbon remediation: potential contributions and constraints by microbes

Terrestrial carbon sequestration represents an important option for partially mitigating anthropogenic CO(2) emissions. Evidence suggests that terrestrial ecosystems can be managed for carbon sequestration, but it is not certain to what extent the microbes within them can be manipulated. Challenges include identifying which specific microbes and mechanisms contribute to sequestered carbon; understanding how microbial communities respond over large spatial and long temporal scales to crucial environmental variables; and developing management strategies suitable for large spatial and long temporal scales. The growing recognition that microbes produce proteins that limit organic matter degradation suggests targets for basic research. Directly manipulating microbes to sequester CO(2) through other processes such as mineral formation offers intriguing alternatives that merit further attention, but at present the prospects for practical implementation appear remote.     

土壤呼吸对秸秆与秸秆生物炭还田的响应及其微生物机制

土壤呼吸释放CO_2是温室气体排放的重要途径之一,减少土地利用中温室气体排放、增强土壤碳汇聚能力对于减缓全球温室效应具有重要意义。生物炭具有改善土壤理化性质、增加作物产量、调节土壤微生物性质等特性。本研究采用室外盆栽的方式,以地肤草为目标植物,研究了芦苇、水稻、互花米草三种农林秸秆及其秸秆生物炭还田对土壤的改良效应,以及对土壤呼吸的影响及其微生物机制。结果表明:秸秆及其秸秆生物炭均可改善土壤肥力,促进植物生长,且生物炭改良效果略好于秸秆直接还田。但秸秆生物炭还田的土壤呼吸显著低于秸秆直接还田,其中芦苇生物炭最低。秸秆直接还田可促进土壤β-糖苷酶、脱氢酶和活性微生物量等微生物活性指标,从而促进土壤呼吸,增加土壤CO_2的释放,而生物炭还田对土壤微生物活性无显著的促进作用,反而有一定的抑制作用,这可能是由于生物炭中易降解有机物含量很低,可降解性较低的缘故。     

Microbial mediation of biogeochemical cycles revealed by simulation of global changes with soil transplant and cropping

Despite microbes'' key roles in driving biogeochemical cycles, the mechanism of microbe-mediated feedbacks to global changes remains elusive. Recently, soil transplant has been successfully established as a proxy to simulate climate changes, as the current trend of global warming coherently causes range shifts toward higher latitudes. Four years after southward soil transplant over large transects in China, we found that microbial functional diversity was increased, in addition to concurrent changes in microbial biomass, soil nutrient content and functional processes involved in the nitrogen cycle. However, soil transplant effects could be overridden by maize cropping, which was attributed to a negative interaction. Strikingly, abundances of nitrogen and carbon cycle genes were increased by these field experiments simulating global change, coinciding with higher soil nitrification potential and carbon dioxide (CO₂) efflux. Further investigation revealed strong correlations between carbon cycle genes and CO₂ efflux in bare soil but not cropped soil, and between nitrogen cycle genes and nitrification. These findings suggest that changes of soil carbon and nitrogen cycles by soil transplant and cropping were predictable by measuring microbial functional potentials, contributing to a better mechanistic understanding of these soil functional processes and suggesting a potential to incorporate microbial communities in greenhouse gas emission modeling.     

施氮对不同有机碳水平桉树林土壤微生物群落碳代谢的影响

土壤微生物群落碳代谢功能既受土壤氮素水平的影响,也与土壤有机碳水平密切相关,但二者如何共同影响土壤微生物群落碳代谢功能的研究尚不多见。以我国南方广泛种植的桉树林为对象,采用野外控制实验比较研究了4种施氮处理(对照:0kg/hm2,低氮:84.2 kg/hm2,中氮:166.8 kg/hm2,高氮:333.7 kg/hm2)对有机碳水平差异显著的两桉树林样地土壤微生物群落碳代谢功能的影响,结果表明:(1)两种有机碳水平桉树林土壤微生物群落碳代谢强度和代谢碳源丰富度显著不同,高有机碳水平桉树林土壤微生物群落碳代谢强度和代谢碳源丰富度显著高于低有机碳水平桉树林(P0.01);(2)施氮显著改变了桉树林土壤微生物群落的碳代谢强度和代谢碳源丰富度(P0.05),随着施氮水平的升高,土壤微生物群落碳代谢强度和代谢碳源丰富度均呈现先增加后降低的变化规律,但是高、低有机碳水平桉树林土壤微生物群落碳代谢强度和代谢碳源丰富度对施氮梯度的响应各不相同,高、低有机碳水平桉树林的土壤微生物群落碳代谢指标分别在中氮、低氮处理中达到最高值;(3)施氮影响土壤微生物群落代谢的碳源类型主要是碳水化合物类、氨基酸类和羧酸类,土壤微生物生物量是影响土壤微生物碳代谢强度和代谢碳源丰富度的重要因素。由此可知,施氮对土壤微生物碳代谢功能影响,也与土壤本底中有机碳水平的调节有关,所以在研究土壤微生物群落对施氮等条件的响应时,不能忽略土壤中有机碳水平。