Influence of elevated CO2 and nitrogen nutrition on rice plant growth,soil microbial biomass,dissolved organic carbon and dissolved CH4

Rice (Oryza sativa) was grown in six sunlit, semi-closed growth chambers for two seasons at 350 L L^–1 (ambient) and 650 L L^–1 (elevated) CO₂ and different levels of nitrogen (N) supplement. The objective of this research was to study the influence of CO₂ enrichment and N nutrition on rice plant growth, soil microbial biomass, dissolved organic carbon (DOC) and dissolved CH₄. Elevated CO₂ concentration ([CO₂]) demonstrated a wide range of enhancement to both above- and below-ground plant biomass, in particular to stems and roots (for roots when N was not limiting) in the mid-season (80 days after transplanting) and stems/ears at the final harvest, depending on season and the level of N supplement. Elevated [CO₂] significantly increased microbial biomass carbon in the surface 5 cm soil when N (90 kg ha^–1) was in sufficient supply. Low N supplement (30 kg ha^–1) limited the enhancement of root growth by elevated [CO₂], leading consequently to diminished response of soil microbial biomass carbon to CO₂ enrichment. The concentration of dissolved CH₄ (as well as soil DOC, but to a lesser degree) was observed to be positively related to elevated [CO₂], especially at high rate of N application (120 kg ha^–1) or at 10 cm depth (versus 5 cm depth) in the later half of the growing season (at 80 kg N ha^–1). Root senescence in the late season complicated the assessment of the effect of elevated [CO₂] on root growth and soil organic carbon turnover and thus caution should be taken when interpreting respective high CO₂ results.     

Hydrophilicity effect on CO2/CH4 separation using carbon nanotube membranes: insights from molecular simulation

Abstract

Molecular simulation methods were applied to study the effect of hydrophilicity on CO₂/CH₄ separation using carbon nanotube (CNT) membranes. CNTs with a diameter of ~1 nm were functionalised by varying amounts of carbonyl groups, in order to achieve various hydrophilicity. The presence of –CO groups inside the CNT allow a significant gain in the diffusion selectivity of CO₂, while in contrast the adsorption selectivity is hardly changed. The corresponding permeation selectivity increases as the hydrophilicity of the CNT-based membrane increases. However, the permeability of CO₂ decreases due to a combination of the intermolecular interactions between the gas and functional groups and the steric effects of the added functional groups. Considering both the permeation selectivity and permeability, it was found that the maximum separation performance is achieved in a certain hydrophilic CNT membrane. Moreover, the separation performance of hydrophilic CNTs for CO₂/CH₄ mixtures breaks the Robeson upper bound.     

Responses of soil microbiota of a late successional alpine grassland to long term CO2 enrichment

We investigated microbial responses in a late successional sedge-dominated alpine grassland to four seasons of CO₂ enrichment. Part of the plots received fertilizer equivalent to 4.5g N m⁻² a⁻¹. Soil basal respiration (R _mic ), the metabolic quotient for CO₂ (qCO₂=R _mic /C _mic ), microbial C and N (C _mic and N _mic ) as well as total soil organic C and N showed no response to CO₂ enrichment alone. However, when the CO₂ treatment was combined with fertilizer addition R _mic and qCO₂ were statistically significantly higher under elevated CO₂ than under ambient conditions (+57% and +71%, respectively). Fertilizer addition increased microbial N pools by 17%, but this was not influenced by elevated CO₂. Microbial C was neither affected by elevated CO₂ nor fertilizer. The lack of a CO₂-effect in unfertilized plots was suprising in the light of our evidence (based on C balance) that enhanced soil C inputs must have occurred under elevated CO₂ regardless of fertilizer treatment. Based on these data and other published work we suggest that microbial responses to elevated CO₂ in such stable, late-successional ecosystems are limited by the availability of mineral nutrients and that results obtained with fertile or heavily disturbed substrates are unsuitable to predict future microbial responses to elevated CO₂ in natural systems. However, when nutrient limitation is removed (e.g. by wet nitrogen deposition) microbes make use of the additional carbon introduced into the soil system. We believe that the response of natural ecosystems to elevated CO₂ must be studied in situ in natural, undisturbed systems.     

Do Root Exudates Enhance Peat Decomposition?

Despite the importance of understanding controls on microbial carbon (C) mineralization in peat soils, the role of vascular plant root exudates is still unclear. To determine whether root exudates could stimulate enhanced decomposition of peat, we utilized an in-vitro method involving the addition of a solution similar to root exudates (6 glucose-C: 2 citrate-C: 2 amino acid-C, at 3 addition levels) to peat, incubating the mixture and measuring CO₂ produced over 20 d and microbial biomass and dissolved organic carbon (DOC) at the end of the incubation. We defined priming as inorganic C (IC) production (CO₂ + calculated dissolved inorganic C) during the incubation being greater than that attributed to the control peat plus the added C. An addition level of 0.083 mg C g^?1 dry peat, estimated to represent root exudation over one 12-h sunny day in a bog, caused an enhancement in IC production that exceeded that produced in the controls and the amount of added C after 8 d, with rates levelling to control values after 15 d. At the end of the incubation nearly 3 times the amount added C had been mineralized, relative to the control, however this represented only 4% of total microbial respiration in the controls. Although the priming effect pattern appeared to be real throughout repeated measurements in our experiments, the statistical probabilities were not always large due to high variability in background CO₂ production levels. Given the observed long lag-times and overall small magnitude and large variability in observed effects, we conclude that although priming of decomposition appears to occur in peatlands, it likely has only a minor overall impact on net C loss to the atmosphere.     

Effect of carbon dioxide concentration on microbial respiration in soil

In order to assess the validity of conventional methods for measuring CO₂ flux from soil, the relationship between soil microbial respiration and ambient CO₂ concentration was studied using an open-flow infra-red gas analyser (IRGA) method. Andosol from an upland field in central Japan was used as a soil sample. Soil microbial respiration activity was depressed with the increase of CO₂ concentration in ventilated air from 0 to 1000 ppmv. At 1000 ppmv, the respiration rate was less than half of that at 0 ppmv. Thus, it is likely that soil respiration rate is overestimated by the alkali absorption method, because CO₂ concentration in the absorption chamber is much lower than the normal level. Metabolic responses to CO₂ concentration were different among groups of soil microorganisms. The bacteria actinomycetes group cultivated on agar medium showed a more sensitive response to the CO₂ concentration than the filamentous fungi group.     

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     

Contribution of root respiration to soil respiration in a C3/C4 mixed grassland

The spatial and temporal variations of soil respiration were studied from May 2004 to June 2005 in a C₃/C₄ mixed grassland of Japan. The linear regression relationship between soil respiration and root biomass was used to determine the contribution of root respiration to soil respiration. The highest soil respiration rate of 11-54 Μmol m^-2 s^-1 was found in August 2004 and the lowest soil respiration rate of 4.99 Μmol m^-2 s^-1 was found in April 2005. Within-site variation was smaller than seasonal change in soil respiration. Root biomass varied from 0.71 kg m^-2 in August 2004 to 102 in May 2005. Within-site variation in root biomass was larger than seasonal variation. Root respiration rate was highest in August 2004 (5.7 Μmol m^-2 s^-1) and lowest in October 2004 (1.7 Μmol m^-2 s^-1). Microbial respiration rate was highest in August 2004 (5.8 Μmol m^-2 s^-1) and lowest in April 2005 (2.59 Μmol m^-2 s^-1). We estimated that the contribution of root respiration to soil respiration ranged from 31% in October to 51% in August of 2004, and from 45% to 49% from April to June 2005.     

Methanogenesis and Methanogen Diversity in Three Peatland Types of the Discontinuous Permafrost Zone,Boreal Western Continental Canada

Because recent patterns of permafrost collapse in boreal peatlands appear to enhance emissions of CH ₄ to the atmosphere, we examined methanogenesis and methanogen diversity in peat soil from peatlands with and without permafrost in two peatland complexes situated in continental western Canada. Peat soil from the active layer of permafrost bogs had very low rates of CH ₄ production (ca. 10 nmol g ^?1 day ^?1 ), and we were unable to PCR-amplify 16s rRNA gene sequences using Archaea-specific primers in four peat samples. Surface peat soil from continental bogs with no permafrost supported moderate rates of CH ₄ production (20–600 nmol g ^?1 day ^?1 ), with maximum rates in soil located close to the mean water table level. Additions of ethanol stimulated CH ₄ production rates, suggesting metabolic substrate limitations. Peat from internal lawns, which have experienced surface permafrost degradation in the past 150 years, had very rapid rates of CH₄ production (up to 800 nmol g ^?1 day ^?1 ) occurring within the soil profile. Concomitant rates of anaerobic CO ₂ production were greater in continental bogs (ca. 6 μmol g ^?1 day ^?1 ) than in internal lawns (ca. 4 μ mol g ^?1 day ^?1 ) or in permafrost bogs (2.8 μ mol g ^?1 day ^?1 ). Analysis of the 16s rRNA gene for Archaea in the continental bog indicated mostly sequences associate with Methanobacteriales and RC-I with a Methanosarcinaceae sequence in the deepest peat soil. In the internal lawn, Methanosarcinaceae were most common in peat soil with a Methanosaetaceae sequence in the deepest peat soil. This study showed that patterns of discontinuous permafrost and ongoing permafrost degradation in boreal regions create patchy soil environments for methanogens and rates of methanogenesis.     

Effects of soil water content on soil respiration in forests and cattle pastures of eastern Amazonia

The effect of soil water content on efflux of CO₂ from soils has been described by linear, logarithmic, quadratic, and parabolic functions of soil water expressed as matric potential, gravimetric and volumetric water content, water holding capacity, water-filled pore space, precipitation indices, and depth to water table. The effects of temperature and water content are often statistically confounded. The objectives of this study are: (1) to analyze seasonal variation in soil water content and soil respiration in the eastern Amazon Basin where seasonal temperature variation is minor; and (2) to examine differences in soil CO₂ emissions among primary forests, secondary forests, active cattle pastures, and degraded cattle pastures. Rates of soil respiration decreased from wet to dry seasons in all land uses. Grasses in the active cattle pasture were productive in the wet season and senescent in the dry season, resulting in the largest seasonal amplitude of CO₂ emissions, whereas deep-rooted forests maintained substantial soil respiration during the dry season. Annual emissions were 2.0, 1.8, 1.5, and 1.0 kg C m^-2 yr^-1 for primary forest, secondary forest, active pasture, and degraded pasture, respectively. Emissions of CO₂ were correlated with the logarithm of matric potential and with the cube of volumetric water content, which are mechanistically appropriate functions for relating soil respiration at below-optimal water contents. The parameterization of these empirical functions was not consistent with those for a temperate forest. Relating rates of soil respiration to water and temperature measurements made at some arbitrarily chosen depth of the surface horizons is simplistic. Further progress in defining temperature and moisture functions may require measurements of temperature, water content and CO₂ production for each soil horizon.     

Elevated CO2 and temperature effects on soil carbon and nitrogen cycling in ryegrass/white clover turves of an Endoaquept soil

Effects of elevated CO₂ (700 L L^–1) and a control (350 L L^–1 CO₂) on the productivity of a 3-year-old ryegrass/white clover pasture, and on soil biochemical properties, were investigated with turves of a Typic Endoaquept soil in growth chambers. Temperature treatments corresponding to average winter, spring, and summer conditions in the field were applied consecutively to all of the turves. An additional treatment, at 700 L L^–1 CO₂ and a temperature 6°C higher throughout than in the other treatments, was included.Under the same temperature conditions, overall herbage yields in the 700 L L^–1 CO₂ treatment were ca. 7% greater than in the control at the end of the summer period. Root mass (to ca 25 cm depth) in the 700 L L^–1 CO₂ treatment was then about 50% greater than in the control, but in the 700 L L^–1 CO₂+6°C treatment it was 6% lower than in the control. Based on decomposition results, herbage from the 700 L L^–1+6°C treatment probably contained the highest proportion of readily decomposable components.Elevated CO₂ had no consistent effect on soil total C and N, microbial C and N, or extractable C concentrations in any of the treatments. Under the same temperature conditions, it did, however, enhance soil respiration (CO₂-C production) and invertase activity. The effects of elevated CO₂ on rates of net N mineralization were less distinct, and the apparent availability of N for the sward was not affected. Under elevated CO₂, soil in the higher-temperature treatment had a higher microbial C:N ratio; it also had a greater potential to degrade plant materials.Data interpretation was complicated by soil spatial variability and the moderately high background levels of organic matter and biochemical properties that are typical of New Zealand pasture soils. More rapid cycling of C under CO₂ enrichment is, nevertheless, indicated. Futher long-term experiments are required to determine the overall effect of elevated CO₂ on the soil C balance.     

Elevated CO2 effects on carbon and nitrogen cycling in grass/clover turves of a Psammaquent soil

Effects of elevated CO₂ (525 and 700 L L^–1), and a control (350 L L^–1 CO₂), on biochemical properties of a Mollic Psammaquent soil in a well-established pasture of C3 and C4 grasses and clover were investigated with continuously moist turves in growth chambers over four consecutive seasonal temperature regimes from spring to winter inclusive. After a further spring period, half of the turves under 350 and 700 L L^–1 were subjected to summer drying and were then re-wetted before a further autumn period; the remaining turves were kept continuously moist throughout these additional three consecutive seasons. The continuously moist turves were then pulse-labelled with ¹⁴C-CO₂ to follow C pathways in the plant/soil system during 35 days.Growth rates of herbage during the first four seasons averaged 4.6 g m^–2 day^–1 under 700 L L^–1 CO₂ and were about 10% higher than under the other two treatments. Below-ground net productivity at the end of these seasons averaged 465, 800 and 824 g m^–2 in the control, 525 and 700 L L^–1 treatments, respectively.in continuously moist soil, elevated CO₂ had no overall effects on total, extractable or microbial C and N, or invertase activity, but resulted in increased CO₂-C production from soil, and from added herbage during the initial stages of decomposition over 21 days; rates of root decomposition were unaffected. CO₂ produced h^–1 mg^–1 microbial C was about 10% higher in the 700 L L^–1 CO₂ treatment than in the other two treatments. Elevated CO₂ had no clearly defined effects on N availability, or on the net N mineralization of added herbage.In the labelling experiment, relatively more ¹⁴C in the plant/soil system occurred below ground under elevated CO₂, with enhanced turnover of ¹⁴C also being suggested.Drying increased levels of extractable C and organic-N, but decreased mineral-N concentrations; it had no effect on microbial C, but resulted in lowered microbial N in the control only. In soil that had been previously summer-dried, CO₂ production was again higher, but net N mineralization was lower, under elevated CO₂ than in the control after autumn pasture growth.Over the trial period of 422 days, elevated CO₂ generally appears to have had a greater effect on soil C turnover than on soil C pools in this pasture ecosystem.     

The C4 pathway: an efficient CO2 pump

The C₄ pathway is a complex combination of both biochemical and morphological specialisation, which provides an elevation of the CO₂ concentration at the site of Rubisco. We review the key parameters necessary to make the C₄ pathway function efficiently, focussing on the diffusion of CO₂ out of the bundle sheath compartment. Measurements of cell wall thickness show that the thickness of bundle sheath cell walls in C₄ species is similar to cell wall thickness of C₃ mesophyll cells. Furthermore, NAD-ME type C₄ species, which do not have suberin in their bundle sheath cell walls, do not appear to compensate for this with thicker bundle sheath cell walls. Uncertainties in the CO₂ diffusion properties of membranes, such as the plasmalemma, choroplast and mitochondrial membranes make it difficult to estimate bundle sheath diffusion resistance from anatomical measurements, but the cytosol itself may account for more than half of the final calculated resistance value for CO₂ leakage. We conclude that the location of the site of decarboxylation, its distance from the mesophyll interface and the physical arrangement of chloroplasts and mitochondria in the bundle sheath cell are as important to the efficiency of the process as the properties of the bundle sheath cell wall. Using a mathemathical model of C₄ photosynthesis, we also examine the relationship between bundle sheath resistance to CO₂ diffusion and the biochemical capacity of the C₄ photosynthetic pathway and conclude that bundle sheath resistance to CO₂ diffusion must vary with biochemical capacity if the efficiency of the C₄ pump is to be maintained. Finally, we construct a mathematical model of single cell C₄ photosynthesis in a C₃ mesophyll cell and examine the theoretical efficiency of such a C₄ photosynthetic CO₂ pump. This revised version was published online in August 2006 with corrections to the Cover Date.     

The effect of CO2 enrichment on leaf photosynthetic rates and instantaneous water use efficiency of Andropogon gerardii in the tallgrass prairie

Open-top chambers were used to study the effects of CO₂ enrichment on leaf-level photosynthetic rates of the C₄ grass Andropogon gerardii in the native tallgrass prairie ecosystem near Manhattan, Kansas. Measurements were made during a year with abundant rainfall (1993) and a year with below-normal rainfall (1994). Treatments included: No chamber, ambient CO₂ (A); chamber with ambient CO₂ (CA); and chamber with twice-ambient CO₂ (CE). Measurements of photosynthesis were made at 2-hour intervals, or at midday, on cloudless days throughout the growing season using an open-flow gas-exchange system. No significant differences in midday rates of photosynthesis or in daily carbon accumulation as a result of CO₂ enrichment were found in the year with abundant precipitation. In the dry year, midday rates of photosynthesis were significantly higher in the CE treatment than in the CA or A treatments throughout the season. Estimates of daily carbon accumulation also indicated that CO₂ enrichment allowed plants to maximize carbon acquisition on a diurnal basis. The increased carbon accumulation was accounted for by greater rates of photosynthesis in the CE plots during midday. During the wet year, CO₂ enrichment decreased stomatal conductance, which allowed plants to decrease transpiration while still photosynthesizing at rates similar to plants in ambient conditions. During the dry year, CO₂ enrichment allowed plants to maintain photosynthetic rates even though stomatal conductance and transpiration had been reduced in all treatments due to stress. Estimates of instantaneous water-use efficiency were reduced under CO₂ enrichment for both years. This revised version was published online in June 2006 with corrections to the Cover Date.     

Novel technique for component monitoring of CO2 exchange in plants

A novel technique designed for component monitoring of CO₂ exchange in plants is described. The system is based on application of self-clamping leaf chambers connected to an open gas-exchange measuring system and on automatic recording of CO₂ concentration. This technique was implemented in a commercially available instrument, PTM-48A Photosynthesis Monitor, which provides for long-term measurements of gas exchange and for discrimination of its separate components. Furthermore, many other plant functions can be monitored during plant growth and development under laboratory, greenhouse, and field conditions.     

Magnitude and variations of groundwater seepage along a Florida marine shoreline

Direct groundwater inputs are receiving increasingattention as a potential source of nutrients and otherdissolved constituents to the coastal ocean. Seepageinto St. George Sound, Florida was measuredextensively from 1992 to 1994 using seepage meters. Spatial and temporal variations were documented alonga 7-km stretch of coastline and up to 1 km from shore. Measurements were made at 3 transects perpendicular toshore and 1 transect parallel to shore. The generalresults indicated that seepage decreased with distancefrom shore (2 of 3 transects), and substantialtemporal and spatial variability was observed inseepage flow from nearshore sediments. In addition,trends in mean monthly integrated seepage rates weresimilar to precipitation patterns measured at a nearbycoastal weather station. Based on these measurements, weestimate that the magnitude of groundwater seepage intothe study area is substantial, representing from 0.23 to4.4 m3 sec-1of flow through the sediments, approximately equivalentto a first magnitude spring. Although it is unknown howrepresentative this region is with respect to globalgroundwater discharge, it demonstrates thatgroundwater flow can be as important as riverine andspring discharge in some cases. Our subsurfacedischarge rates suggest groundwater is an importanthydrologic source term for this region and may beimportant to the coastal biogeochemistry as well.     

岩溶区不同土地利用方式下土壤CO2排放模拟研究

土地利用变化作为全球气候变化研究的重要内容之一,对土壤CO₂的排放具有重要影响。岩溶区石漠化治理过程中植被恢复伴随着土地利用方式的转变,其对土壤CO₂排放的影响有待进一步研究。基于控制性实验,以土壤、岩溶含水介质初始条件相同,仅土地利用方式不同的贵州普定沙湾模拟试验场为研究对象,通过1年的土壤CO₂浓度和通量数据,研究岩溶区不同土地利用方式下土壤CO₂的排放规律及其影响因素。结果表明：(1)土壤CO₂的浓度和通量具有明显的季节变化规律,不同季节下的土壤CO₂通量呈现昼夜变化规律,温度和降雨影响着土壤CO₂的排放,前者可促进排放量,后者可抑制排放量,且不同土地利用方式受影响的程度不同;(2)耕作活动也会影响土壤CO₂的排放,耕作使得土壤变得松散,加上岩溶区下伏基岩的溶蚀作用,增加了土壤CO₂向含水层的扩散,导致春季耕地表现为负通量;(3)不同土地利用方式下土壤CO₂的年排...     

土地利用对崇明岛围垦区土壤有机碳库和土壤呼吸的影响

土地利用方式是影响农业土壤碳固持和温室气体减排的关键因子之一,而准确地评价土地利用变化的影响往往因土壤本底的不均一和土地利用历史多变而复杂化。为此,在崇明东滩湿地围垦区选取了本底均匀、利用历史简单的几种土地利用类型(水-旱轮作农田、人工林、鱼塘撂荒地),研究其土壤有机碳库和土壤呼吸的变化及其与土壤环境间的关系,以期评价其各自的固碳和温室气体减排潜力。农田土壤的表层(20cm)有机碳和微生物生物量碳含量最高,分别为12.62g/kg和225.34mg/kg,包括苗圃栾树林、水杉林带以及桔园在内的人工林地次之,鱼塘撂荒地最低;但撂荒地深层土壤(40—100cm)的有机碳含量高于其它类型,反映了围垦前湿地土壤有机碳累积的残留影响。土壤呼吸强度的顺序则为鱼塘撂荒地农田桔园苗圃栾树林水杉林带。农耕地在前作小麦收割种植水稻后,土壤CO2通量显著下降,不及旱作时的10%。除农田和撂荒地以外,土壤表层5 cm深处温度可以很好地解释土壤呼吸速率的变化,但在高温高湿季节呼吸速率较为离散。研究表明:在有机质含量较低的土壤中,水-旱轮作可增加土壤有机碳的储量;受人类活动干扰较小的林地土壤,有机碳含量反而有可能低于农田土壤。在中国南方湿润亚热带地区,水旱轮作可较好地协调农业土壤的碳固持和释放过程的矛盾,可能具有相当大的农业减排潜力。     

喀斯特峰丛洼地土壤剖面微生物特性对植被和坡位的响应

选取广西环江县喀斯特峰丛洼地:草丛(T)、灌丛(S)、原生林(PF)(中坡位)不同植被类型,原生林上、中、下不同坡位,按土壤发生层采集淋溶层(A层,0-10 cm)、过渡层(AB层,20-30 cm,草丛和灌丛;30-50 cm,原生林)、淀积层(B层,70-100cm)样品,研究土壤微生物量碳、氮(Soil microbial biomass carbon (SMBC)、soil microbial biomass nitrogen (SMBN))、微生物碳熵、氮熵(ratio of SMBC to soil organic carbon (qMBC)、ratio of SMBN to soil total nitrogen (qMBN))、土壤基础呼吸(soil basic respiration (SBR))以及代谢熵(microbial metabolic quotient(qCO2))的剖面分异特征及其影响因素.结果表明,植被、土层深度显著影响土壤微生物量及基础呼吸,随植被恢复,SMBC、SMBN、SBR由草丛、灌丛、原生林依次上升,并随土壤发生层位的加深逐渐减少,qCO2在3种植被类型间差异显著:T＞PF＞S;原生林A层SMBC,SMBN在各坡位间均显著高于AB层、B层,SBR在A层由下坡位至上坡位递减,而在AB和B层的上、下坡位间无显著差异,qCO2坡位间无显著差异(P＞0.05);SMBC与SMBN之间存在显著正相关(r=0.825,P＜0.01,n=45),且SMBC、SMBN、SBR分别与有机碳、全氮、碱解氮均呈显著正相关.因此,随植被恢复,土壤质量明显改善,且坡位对A层土壤的影响较AB层和B层更显著,对于维持土壤微生物调节的土壤养分循环功能,调控土壤氮素营养与土壤有机质同等重要,这为合理制订喀斯特生态恢复措施提供了理论依据.