Experimental Evidence that Fungi are Dominant Microbes in Carbon Content and Growth Response to Added Soluble Organic Carbon in Moss‐rich Tundra Soil

Global warming significantly affects Arctic tundra, including permafrost thaw and soluble C release that may differentially affect tundra microbial growth. Using laboratory experiments, we report some of the first evidence for the effects of soluble glucose‐C enrichment on tundra soil prokaryotes (bacteria and archaea) and fungi, with comparisons to microbial eukaryotes. Fungal increase in C‐biomass was equivalent to 10% (w/w) of the added glucose‐C, and for prokaryote biomass 2% (w/w), the latter comparable to prior published results. The C‐gain after 14 d was 1.3 mg/g soil for fungi, and ~200 μg/g for prokaryotes.     

不同密度巨桉人工林土壤有机碳及微生物量碳氮特征

通过原位试验,对华西雨屏区不同密度[833株·hm-2(A1)、1 333株·hm-2(A2)、2 222株·hm-2(A3)]巨桉(Eucalyptus grandis)人工林土壤有机碳和微生物生物量碳氮动态特征进行了研究。结果显示:(1)各密度巨桉林分土壤有机碳含量在夏季和秋季较高,春季和冬季较低,但季节变化相对平稳,而它们土壤有机碳含量年平均值分别为22.54g·kg-1(A1)、19.76g·kg-1(A2)、16.84g·kg-1(A3),且密度间差异达到显著水平。(2)各密度林分土壤微生物生物量碳氮呈现出与土壤有机碳相似的规律性季节变化,随着林分密度增加,林下土壤微生物生物量碳氮含量减小。(3)土壤微生物熵一年内的波动较小,分别处于2.30%~2.44%(A1)、2.14%~2.39%(A2)、2.47%~2.69%(A3)之间。(4)各密度林分土壤微生物生物量碳氮含量与其立地土壤有机碳、水解氮、有效磷、速效钾均存在显著相关关系。研究表明,华西雨屏区不同密度巨桉人工林土壤有机碳含量、微生物生物量碳氮含量、微生物熵季节变化相对平稳,但受到人工林密度的显著影响,并随巨桉密度增加土壤有机碳含量、微生物生物量碳氮含量有降低的趋势。     

Rapid turnover of DOC in temperate forests accounts for increased CO2 production at elevated temperatures

The evidence for the contribution of soil warming to changes in atmospheric CO₂ concentrations and carbon stocks of temperate forest ecosystems is equivocal. Here, we use data from a beech/oak forest on concentrations and stable isotope ratios of dissolved organic carbon (DOC), phosphate buffer-extractable organic carbon, soil organic carbon (SOC), respiration and microbial gross assimilation of N to show that respired soil carbon originated from DOC. However, the respiration was not dependent on the DOC concentration but exceeded the daily DOC pool three to four times, suggesting that DOC was turned over several times per day. A mass flow model helped to calculate that a maximum of 40% of the daily DOC production was derived from SOC and to demonstrate that degradation of SOC is limiting respiration of DOC. The carbon flow model on SOC, DOC, microbial C mobilization/immobilization and respiration is linked by temperature-dependent microbial and enzyme activity to global warming effects of CO₂ emitted to the atmosphere.     

荒漠草原3种典型群落类型的土壤微生物量碳氮研究

采用氯仿熏蒸-浸提法,以宁夏盐池荒漠草原3种典型群落(柠条、沙蒿、短花针茅)类型为研究对象,分析了不同生境(冠下、丛间)和不同土层间(0~5、5~10、10~15cm)土壤理化性质及微生物量——微生物量碳(MBC)和微生物量氮(MBN)的变化特征。结果表明:(1)3种群落土壤微生物量变化差异较大,柠条、沙蒿和短花针茅群落土壤MBC含量分别为77.00~393.18、17.27~221.71和81.05~173.37mg/kg,MBN含量分别为7.59~64.81、1.43~13.95和4.25~22.13mg/kg,MBC和MBN含量均表现为:冠下丛间,且随土层深度的增加而降低,有明显的"沃岛效应"。(2)群落类型对土壤微生物量碳氮含量的变化有显著影响,3种典型群落类型下土壤微生物量熵(qMB)、碳氮比(C/N)、微生物量碳氮比(MBC/MBN)分别在0.76~4.10、15.02~52.50、5.34~23.07范围内变化,其比值在不同生境和不同土层深度的分布特征有明显差异。(3)3种典型群落类型的土壤MBC与SOC、MBN、qMB具有显著相关关系,土壤C/N与MBC/MBN呈显著正相关关系,表明土壤MBC、MBN具有一定的生物学指示特性,可以作为评价土壤质量的生物学指标。     

Tree Species Effects on Soil Organic Matter Dynamics: The Role of Soil Cation Composition

Abstract We studied the influence of tree species on soil carbon and nitrogen (N) dynamics in a common garden of replicated monocultures of fourteen angiosperm and gymnosperm, broadleaf and needleleaf species in southwestern Poland. We hypothesized that species would influence soil organic matter (SOM) decomposition primarily via effects on biogeochemical recalcitrance, with species having tissues with high lignin concentrations retarding rates of decomposition in the O and A horizons. Additionally, because prior work demonstrated substantial divergence in foliar and soil base cation concentrations and soil pH among species, we hypothesized that species would influence chemical stabilization of SOM via cation bridging to mineral surfaces in the A-horizon. Our hypotheses were only partially supported: SOM decomposition and microbial biomass were unrelated to plant tissue lignin concentrations, but in the mineral horizon, were significantly negatively related to the percentage of the cation exchange complex (CEC) occupied by polyvalent acidic (hydrolyzing) cations (Al and Fe), likely because these cations stabilize SOM via cation bridging and flocculation and/or because of inhibitory effects of Al or low pH on decomposers. Percent CEC occupied by exchangeable Al and Fe was in turn related to both soil clay content (a parent material characteristic) and root Ca concentrations (a species characteristic). In contrast, species influenced soil N dynamics largely via variation in tissue N concentration. In both laboratory and in situ assays, species having high-N roots exhibited faster rates of net N mineralization and nitrification. Nitrification:mineralization ratios were greater, though, under species with high exchangeable soil Ca²⁺. Our results indicate that tree species contribute to variation in SOM dynamics, even in the mineral soil horizons. To our knowledge the influence of tree species on SOM decomposition via cation biogeochemistry has not been demonstrated previously, but could be important in other poorly buffered systems dominated by tree species that differ in cation nutrition or that are influenced by acidic deposition.     

荒漠草原3种典型群落类型下土壤粒径分布与微生物量碳氮的关系

为了探讨不同群落类型的土壤粒径分布（PSD）与土壤微生物生物量（MB）的关系,以宁夏盐池荒漠草原3种典型群落类型（冰草、沙蒿、短花针茅）为研究对象,测定了不同群落2种生境（冠下、丛间）0~5、5~10和10~15 cm表土层土壤PSD和微生物量碳（MBC）、微生物量氮（MBN）含量变化,并分析了土壤颗粒组成中砂粒、粉粒和黏粒体积分数变化与土壤有机碳（SOC）、全氮（TN）和MBC、MBN间的关系。结果表明：（1）不同群落类型土壤粒径都呈“倒V”型分布趋势,但土壤退化最严重的沙蒿群落中100~500 μm粒径颗粒含量相对较多,与其他两种群落形成显著差异。（2）不同群落类型SOC、MBC、MBN含量均随土层深度的增加而降低,同土层SOC、MBN含量均表现为冠下明显大于丛间,表现出“肥岛效应”,且0~5 cm土层差异显著（P＜0.05）。（3）对土壤粒径组成与土壤SOC、MBC、MBN间相关性研究表明,在土壤SOC、MBC和MBN含量较高的冰草、短花针茅群落类型中,0.01~2、2~50和50~100 μm土壤粒径的颗粒含量也高,SOC、MBC和MBN含量与＜100 μm的粉粒含量呈正相关关系;在沙蒿群落类型中粒径为100~250和250~500 μm的土粒含量增高,导致其SOC、MBC和MBN含量较低,表明不同群落类型对土壤理化结构产生影响的同时,对微生物生物量也有显著的影响。     

黄土旱塬区冬小麦不同施肥处理的土壤呼吸及土壤碳动态

依据黄土旱塬区黑垆土上中国科学院长武站长期定位试验 (始于1984年),于2008年3月到6月,测定了冬小麦连作系统中返青期、拔节期、抽穗期、灌浆期和收获期土壤呼吸日变化、生育期变化以及土壤可溶性有机碳(Dissolved organic C, DOC)和微生物量碳(Soil microbial biomass C, MBC),研究了施肥措施对土壤呼吸、DOC和MBC的影响以及土壤呼吸与碳组分之间的关系.研究涉及6个处理:休闲地(F)、不施肥(CK)、有机肥(M)、氮肥(N)、氮磷肥(NP)和氮磷有机肥(NPM).结果表明,冬小麦连作系统中土壤呼吸的日变化格局呈单峰曲线,最高值出现在12:00左右(拔节期)和14:30左右(成熟期),最小值出现在0:00～3:00之间或6:00左右;冬小麦土壤呼吸速率拔节期最高,其次是灌浆后期,抽穗期最低;不同施肥条件下,各生育期土壤呼吸速率大小顺序:NPM＞M＞NP＞N＞CK＞F.土壤水分亏缺是导致抽穗期和灌浆期土壤呼吸速率降低的重要原因.各施肥处理DOC含量高低顺序为灌浆期＞抽穗期＞成熟期＞返青期＞拔节期;除M,NPM处理MBC含量拔节期＞灌浆期外,各施肥处理MBC含量高低顺序为成熟期＞抽穗期＞灌浆期＞拔节期＞返青期.同一处理不同生育期土壤呼吸速率与DOC,MBC的相关性较低,但同生育期不同施肥处理土壤呼吸与土壤有机碳组分间存在显著的相关性.以F处理土壤呼吸为基础,估算CK、N和NP处理生育期根系对土壤呼吸的平均贡献率依次为36%、45%和54%.     

Evaluation of Microbial Biomass and Activity in Different Soils Exposed to Increasing Level of Arsenic Pollution: A Laboratory Study

Soil contamination with arsenic is a widespread phenomenon in many parts of the world and it is well known that it may affect the soil microbial community. A laboratory experiment was performed to evaluate the acute toxicity effect of different As concentrations (0, 0.1, 1.0, 10, 100, and 200 mg/kg) on soil biological parameters like microbial biomass carbon (MBC), active microbial biomass carbon (AMBC), basal soil respiration (BSR), fluorescein diacetate hydrolase activity (FDA), and dehydrogenase activity (DHA) under three different soil types (Inceptisol, Vertisol, and Entisol). Soil enzyme activities, microbial biomass, and respiration were significantly decreased by arsenic exposure. The soil metabolic quotient increased with increasing As concentration, the maximum being observed for Entisol followed by Vertisol and Inceptisol. Except for soil enzyme activities, the majority of which started to decrease after the addition of 10 mg/kg arsenic, all other parameters decreased at arsenic dose ≥0.1 mg/kg. The decrease of AMBC/MBC ratio showed that acute arsenic stress affected the active microorganisms more than the dormant component of total microbial biomass. Among the three soils studied, Vertisol was comparatively more tolerant to As exposure in terms of soil enzymes and biomass, followed by Inceptisol and Entisol. MBC has evolved as a distinct parameter in cluster analysis, while some others, such as DHA and AMBC, and FDA and BSR, appeared essentially redundant.     

荒漠草原典型植物群落土壤活性有机碳组分特征及其与酶活性的关系

以宁夏荒漠草原典型植物柠条(Caragana korshinskii)、沙蒿(Artemisia ordosica)、短花针茅(Stipa breviflora)和蒙古冰草(Agropyron mongolicum)群落为研究对象,分析不同植物群落不同土层深度(0~5、5~10和10~15cm)土壤活性有机碳组分土壤微生物量碳(MBC)、可溶性有机碳(DOC)和易氧化有机碳(EOC)特征及其与土壤酶(蔗糖酶、脲酶、碱性磷酸酶和过氧化氢酶)活性之间的关系。结果表明:(1)4种典型植物群落土壤SOC、MBC、EOC含量均随土层深度的增加而减少,且表层(0~5cm)土壤显著高于亚表层(5~10cm)和深层(10~15cm)土壤(P0.05),而土壤DOC含量随土层深度的增加呈先增加后减少的趋势。在同一土层深度,灌木(柠条和沙蒿)群落土壤活性有机碳组分含量高于禾本科植物(短花针茅和蒙古冰草)。(2)4种典型植物群落土壤酶(蔗糖酶、脲酶、磷酸酶和过氧化氢酶)活性整体上随土层深度的增加而降低,局部土层深度表现出波动性;同一土层不同植被群落土壤酶活性未表现出一定的变化规律。(3)4种典型群落土壤活性有机碳各组分除DOC外,其余均与SOC呈显著正相关关系,与土壤酶活性、微生物量熵以及有机碳活度具有一定的相关关系,表明土壤活性有机碳不仅依赖于总有机碳,也与土壤酶活性密切相关。     

Soil [N] modulates soil C cycling in CO2‐fumigated tree stands: a meta‐analysis

Under elevated atmospheric CO₂ concentrations, soil carbon (C) inputs are typically enhanced, suggesting larger soil C sequestration potential. However, soil C losses also increase and progressive nitrogen (N) limitation to plant growth may reduce the CO₂ effect on soil C inputs with time. We compiled a data set from 131 manipulation experiments, and used meta‐analysis to test the hypotheses that: (1) elevated atmospheric CO₂ stimulates soil C inputs more than C losses, resulting in increasing soil C stocks; and (2) that these responses are modulated by N. Our results confirm that elevated CO₂ induces a C allocation shift towards below‐ground biomass compartments. However, the increased soil C inputs were offset by increased heterotrophic respiration (Rh), such that soil C content was not affected by elevated CO₂. Soil N concentration strongly interacted with CO₂ fumigation: the effect of elevated CO₂ on fine root biomass and –production and on microbial activity increased with increasing soil N concentration, while the effect on soil C content decreased with increasing soil N concentration. These results suggest that both plant growth and microbial activity responses to elevated CO₂ are modulated by N availability, and that it is essential to account for soil N concentration in C cycling analyses.     

The Fate and Retention of Organic and Inorganic <Superscript>15</Superscript>N-Nitrogen in an Old-Growth Forest Soil in Western Oregon

Forests in the American Pacific Northwest receive very little nitrogen (N) through atmospheric deposition; therefore, they can provide insights into how the N cycle functioned in other regions before heavy atmospheric deposition of inorganic N began. Our objectives were to determine (a) if the fate of organic N differed from the fate of inorganic N, (b) the effect that polyphenols have on the fate of organic N, and (c) the effect of season of addition on the fate of N inputs. We traced N added to in situ soil cores as ammonium, organic N, tannin-complexed organic N, and the N₂-fixing lichen Lobaria oregana. Total ¹⁵N recovery was between 74% and 109% for all N additions. Total ¹⁵N recovery did not vary significantly from the first sampling date to the last date. The litter/organic horizon, as a bulk pool, was the largest N retention pool for all forms of N addition. Within the litter/organic horizon, the chloroform-extractable microbial biomass initially accounted for nearly all of the added N from the ammonium additions. On a different time scale, microbial biomass also played a noteworthy role in the retention of N from organic N, tannin-complexed organic N, and Lobaria. Complexing organic matter with tannin appeared to slow N cycling, but it did not significantly change the ultimate distribution of added organic N. Season of N addition had little effect on the retention of added N; however, where differences did occur, spring additions had lower recoveries than autumn additions.     

小兴安岭原始阔叶红松林和枫桦次生林土壤有机碳库比较研究

土壤有机碳含量是全球生态系统碳储量变化的重要指标之一,本研究以空间替代时间序列的方法,分别选取了小兴安岭地区原始阔叶红松林和枫桦次生林并测定土壤有机碳库、土壤全氮含量、土壤微生物量碳及土壤相关理化性质,结果表明,土壤有机碳含量(SOC)、土壤全氮含量(TN)、土壤微生物量碳(MBC)、土壤含水率等指标随着土壤层的深度增加而逐渐减少最后趋于稳定,而土壤容重随着土壤层的加深而增大。在原始林中0~10和10~20cm层的SOC、TN含量差异不显著,而次生林则差异显著。原始阔叶红松林和枫桦次生林的土壤有机碳密度(SOCD)分别为21.46和21.3 kg·m^-2,差异不显著。原始林和次生林的平均有机碳含量分别为35.79,28.6 g·kg^-1,土壤全氮含量分别为2.86,1.83 g·kg^-1,枫桦次生林MBC与SOC的线性相关性高于原始林。结果表明原始林土壤肥力高于次生林,在今后次生林的管理中应适当混栽针叶树种,原始林中应适当间伐使地下碳储量增加。     

Bacterial and Heterotrophic Nanoflagellate Densities and C‐biomass Estimates Along an Alaskan Tundra Transect with Prediction of Respiratory CO2 Efflux

Although tundra terrestrial ecology is significantly affected by global warming, we know relatively little about how eukaryotic microbial communities respond and how much microbial respiratory CO₂ may be released due to available organic nutrient sources in the permafrost melt. Prior research has shown a strong positive correlation between bacteria and fungi in some Arctic locales; this research focused on the relationships of terrestrial bacteria and heterotrophic nanoflagellates. The densities and estimated C‐biomass of bacteria and heterotrophic nanoflagellates (a major occurring group of protozoa) were assessed in 14 samples obtained along a 10 km transect in northwest AK during the summer of 2012. Two samples were taken, one at the top and one near the base of seven hummocks along the transect. Densities (no./g soil) of bacteria varied from 2.7–16 × 10⁹, and nanoflagellates 0.7–7.9 × 10⁷. C‐biomass (μg/g soil) of bacteria varied from 358 to 2,114, and nanoflagellates 12–37. Additionally, the rate of respiration was analyzed in the laboratory for each soil sample. A linear relationship between soil respiration and bacterial densities was obtained (20 °C): R_s = 12.32 + 14.07 B_d (p ? 0.01); where R_s is soil respiration (nmol/min/g soil) and B_d = bacterial density (no. × 10⁹/g soil).     

Effects of soil moisture on the temperature sensitivity of heterotrophic respiration vary seasonally in an old‐field climate change experiment

Microbial decomposition of soil organic matter produces a major flux of CO₂ from terrestrial ecosystems and can act as a feedback to climate change. Although climate‐carbon models suggest that warming will accelerate the release of CO₂ from soils, the magnitude of this feedback is uncertain, mostly due to uncertainty in the temperature sensitivity of soil organic matter decomposition. We examined how warming and altered precipitation affected the rate and temperature sensitivity of heterotrophic respiration (R_h) at the Boston‐Area Climate Experiment, in Massachusetts, USA. We measured R_h inside deep collars that excluded plant roots and litter inputs. In this mesic ecosystem, R_h responded strongly to precipitation. Drought reduced R_h, both annually and during the growing season. Warming increased R_h only in early spring. During the summer, when R_h was highest, we found evidence of threshold, hysteretic responses to soil moisture: R_h decreased sharply when volumetric soil moisture dropped below ~15% or exceeded ~26%, but R_h increased more gradually when soil moisture rose from the lower threshold. The effect of climate treatments on the temperature sensitivity of R_h depended on the season. Apparent Q₁₀ decreased with high warming (~3.5 °C) in spring and fall. Presumably due to limiting soil moisture, warming and precipitation treatments did not affect apparent Q₁₀ in summer. Drought decreased apparent Q₁₀ in fall compared to ambient and wet precipitation treatments. To our knowledge, this is the first field study to examine the response of R_h and its temperature sensitivity to the combined effects of warming and altered precipitation. Our results highlight the complex responses of R_h to soil moisture, and to our knowledge identify for the first time the seasonal variation in the temperature sensitivity of microbial respiration in the field. We emphasize the importance of adequately simulating responses such as these when modeling trajectories of soil carbon stocks under climate change scenarios.     

三江平原4种典型湿地土壤碳氮分布差异和微生物特征

选择三江平原洪河湿地保护区4种典型湿地类型：小叶章+沼柳湿地、小叶章湿地、毛苔草湿地和芦苇湿地作为研究对象,分析了不同湿地土壤有机碳(SOC)、全氮(TN)含量和微生物活性指标(土壤蔗糖酶、纤维素酶、过氧化氢酶、微生物生物量碳MBC、微生物生物量氮MBN、微生物呼吸速率MBR、微生物商q_MB和代谢商q CO₂)的变化及其相互关系.结果表明： SOC和TN含量均随土层深度增加而减少,不同湿地类型之间具有极显著性差异(P<0.01).各湿地土壤酶活性(除过氧化氢酶)、MBC、MBN含量和MBR均以表层(0～10 cm)最高,并随着土层深度的增加而降低.在0～30 cm土层内,小叶章+沼柳湿地和小叶章湿地SOC、TN含量、土壤酶活性、MBC、MBN含量、MBR、q_MB和qCO₂均高于淹水区的毛苔草湿地和芦苇湿地.统计分析表明,SOC、TN与微生物活性指标(qCO₂除外)均存在极显著正相关关系(P<0.01).表明研究区土壤微生物特征对SOC、TN的变化具有重要的影响和指示作用.     

Short‐term soil carbon sink potential of oil palm plantations

Oil palm plantations cover ≈14.6 million ha worldwide and the total area under cultivation is expected to increase during the 21st century . Indonesia and Malaysia together account for 87% of global palm oil production and the combined harvested area in these countries has expanded by 6.5 million ha since 1990. Despite this, soil C cycling in oil palm systems is not well quantified but such information is needed for C budget inventories. We quantified soil C storage (root biomass, soil organic matter (SOM) and microbial biomass) and losses [potential soil respiration (R_s) and soil surface CO₂ flux (F_s)] in mineral soils from an oil palm plantation chronosequence (11–34 years since planting) in Selangor, Malaysia. There were no significant effects of plantation age on SOM, microbial biomass, R_s or F_s, implying soil C was in dynamic equilibrium over the chronosequence. However, there was a significant increase in root biomass with plantation age, indicating a short‐term C sink. Across the chronosequence, R_s was driven by soil moisture, soil particle size, root biomass and soil microbial biomass N but not microbial biomass C. This suggests that the nutrient status of the microbial community may be of equal or greater importance for soil CO₂ losses than substrate availability and also raises particular concerns regarding the addition of nitrogenous fertilizer, i.e. increased yields will be associated with increased soil CO₂ emissions. To fully assess the impact of oil palm plantations on soil C storage, initial soil C losses following land conversion (e.g. from native forest or other previous plantations) must be accounted for. If initial soil C losses are large, our data show that there is no accumulation of stable C in the soil as the plantation matures and hence the conversion to oil palm would probably represent a net loss of soil C.     

锡林河流域两类植物群落土壤呼吸特征的比较

采用碱液吸收法对锡林河流域一个半干旱典型草原群落的土壤呼吸进行了5个月的野外测定,并对其与气候因子和生物量之间的关系进行了分析.另选择了锡林河岸边的一个沼泽化草甸群落作为对比来研究土壤湿度和植被类型对土壤呼吸的影响.主要结果包括:1)两个群落土壤呼吸的季节动态基本一致,均出现了两个峰值,其中草原群落和草甸群落土壤呼吸速率的变化范围分别为312.8～1 738.9mgC@m-2@d-1和354.6～2 235.6 mg C@m -2@d-1.草甸群落的土壤呼吸速率明显高于草原群落,它们的日平均土壤呼吸速率分别为1 349.6 mg C@m-2@d-1和785.9mg C@m-2@d1;2)在草原群落中,土壤呼吸速率与土壤湿度的相关性比其与温度的关系更加显著,而在草甸群落正好相反,反映出这两种气候因子在不同生境中起着不同的作用.根据土壤呼吸与气温之间的同归关系外推出2001年生长季草原群落和草甸群落的土壤呼吸量分别为142.4 g C/m2和236.1 g C/m2;3)在草甸群落中,地上总生物量与土壤呼吸速率之间没有显著的相关关系,而地上部活体生物量与土壤呼吸速率之间则存有很显著的幂函数关系.在草原群落中,土壤呼吸速率与地上活体生物量或地上总生物量的相关性均很弱.     

贵州山区土壤中微生物担是能源物质碳流动的源与汇

在传统的农业生态系统的研究中 ,主要精力放在营养物 (如Ｎ)上 ,认为它们是限制生产力的因素 ;而往往忽略了土壤中碳的重要性 ,认为收获不受Ｃ限制的影响。然而 ,碳循环中的有机碳的分解作用部分控制着出现在地表下和显露在地表上的农业过程[4]。土壤中所储存的有机质 ,其数量既反映土壤从植物残留物的输入所获得的有机质与微生物群落的能量和营养需求之间的平衡 ,又反映植物对营养物的需求与有机质分解作用之间的平衡。因此 ,土壤中碳的平衡能反映出有机质中能量物质的储存[5]。大部分由光合作用形成的碳 ,是通过地表下的生态系统来流动的[…