Model of apparent and real priming effects: Linking microbial activity with soil organic matter decomposition

The most frequently used models simulating soil organic matter (SOM) dynamics are based on first-order kinetics. These models fail to describe and predict such interactions as priming effects (PEs), which are short-term changes in SOM decomposition induced by easily available C or N sources. We hypothesized that if decomposition rate depends not only on size of the SOM pool, but also on microbial biomass and its activity, then PE can be simulated. A simple model that included these interactions and that consisted of three C pools - SOM, microbial biomass, and easily available C - was developed. The model was parameterized and evaluated using results of ¹²C-CO₂ and ¹⁴C-CO₂ efflux after adding ¹⁴C-labeled glucose to a loamy Haplic Luvisol. Experimentally measured PE, i.e., changes in SOM decomposition induced by glucose, was compared with simulated PE. The best agreement between measured and simulated CO₂ efflux was achieved by considering both the total amount of microbial biomass and its activity. Because it separately described microbial turnover and SOM decomposition, the model successfully simulated apparent and real PE.The proposed PE model was compared with three alternative approaches with similar complexity but lacking interactions between the pools and neglecting the activity of microbial biomass. The comparison showed that proposed new model best described typical PE dynamics in which the first peak of apparent PE lasted for 1 day and the subsequent real PE gradually increased during 60 days. This sequential decomposition scheme of the new model, with immediate microbial consumption only of soluble substrate, was superior to the parallel decomposition scheme with simultaneous microbial consumption of two substrates with different decomposability. Incorporating microbial activity function in the model improved the fit of simulation results with experimental data, by providing the flexibility necessary to properly describe PE dynamics. We conclude that microbial biomass should be considered in models of C and N dynamics in soil not only as a pool but also as an active driver of C and N turnover.     

Three-source-partitioning of microbial biomass and of CO2 efflux from soil to evaluate mechanisms of priming effects

We propose and successfully applied a new approach for 3-source-partitioning based on a combination of ¹⁴C labeling with ¹³C natural abundance. By adding ¹⁴C-labeled glucose to soil after C₃ - C₄ vegetation change, we partitioned three C sources in three compartments, namely CO₂, microbial biomass and dissolved organic C (DOC). This enabled us to estimate mechanisms and sources of priming effects (PE).Glucose application at low and high rate (GL: 100 and GH: 1000 μg C g⁻¹, respectively) caused positive PE both short-term (during 1-3 days) and long-term (3-55 days). Despite a 10-fold difference in the amount of substrate added, the PE observed was larger by a factor of only 1.6 at the high versus low rate of glucose. The real and apparent priming effects were distinguished by partitioning of microbial C for glucose-C and SOM-derived C. As the amount of primed CO₂ respired during short-term PE was 40% lower than microbial C, and the contribution of soil C in microbial biomass did not increase, we concluded that such short-term PE was apparent and was mainly caused by accelerated microbial turnover (at GL) and by pool substitution (at GH). Both the amount of primed CO₂-C, which was 1.3-2.1 times larger than microbial C, and the increased contribution of soil C in microbial biomass allowed us to consider the long-term PE as being real. The sole source of real PE (GL treatment) was the “recent” soil organic matter, which is younger than 12-year-old C. The real PE-induced by a glucose amount exceeding microbial biomass (GH) was due to the almost equal contribution of ‘recent’ (<12 years) and ‘old’ (>12 years) C. Thus, the decomposition of old recalcitrant SOM was induced only by an amount of primer exceeding microbial C. We conclude that combining ¹⁴C labeling with ¹³C natural abundance helped disentangle three C sources in CO₂, microbial biomass and DOC and evaluate mechanisms and sources of PE.     

Priming effects: Interactions between living and dead organic matter

In this re-evaluation of our 10-year old paper on priming effects, I have considered the latest studies and tried to identify the most important needs for future research. Recent publications have shown that the increase or decrease in soil organic matter mineralization (measured as changes of CO₂ efflux and N mineralization) actually results from interactions between living (microbial biomass) and dead organic matter. The priming effect (PE) is not an artifact of incubation studies, as sometimes supposed, but is a natural process sequence in the rhizosphere and detritusphere that is induced by pulses or continuous inputs of fresh organics. The intensity of turnover processes in such hotspots is at least one order of magnitude higher than in the bulk soil. Various prerequisites for high-quality, informative PE studies are outlined: calculating the budget of labeled and total C; investigating the dynamics of released CO₂ and its sources; linking C and N dynamics with microbial biomass changes and enzyme activities; evaluating apparent and real PEs; and assessing PE sources as related to soil organic matter stabilization mechanisms. Different approaches for identifying priming, based on the assessment of more than two C sources in CO₂ and microbial biomass, are proposed and methodological and statistical uncertainties in PE estimation and approaches to eliminating them are discussed. Future studies should evaluate directions and magnitude of PEs according to expected climate and land-use changes and the increased rhizodeposition under elevated CO₂ as well as clarifying the ecological significance of PEs in natural and agricultural ecosystems. The conclusion is that PEs - the interactions between living and dead organic matter - should be incorporated in models of C and N dynamics, and that microbial biomass should regarded not only as a C pool but also as an active driver of C and N turnover.     

Experimental warming effects on the microbial community of a temperate mountain forest soil

Soil microbial communities mediate the decomposition of soil organic matter (SOM). The amount of carbon (C) that is respired leaves the soil as CO₂ (soil respiration) and causes one of the greatest fluxes in the global carbon cycle. How soil microbial communities will respond to global warming, however, is not well understood. To elucidate the effect of warming on the microbial community we analyzed soil from the soil warming experiment Achenkirch, Austria. Soil of a mature spruce forest was warmed by 4 °C during snow-free seasons since 2004. Repeated soil sampling from control and warmed plots took place from 2008 until 2010. We monitored microbial biomass C and nitrogen (N). Microbial community composition was assessed by phospholipid fatty acid analysis (PLFA) and by quantitative real time polymerase chain reaction (qPCR) of ribosomal RNA genes. Microbial metabolic activity was estimated by soil respiration to biomass ratios and RNA to DNA ratios. Soil warming did not affect microbial biomass, nor did warming affect the abundances of most microbial groups. Warming significantly enhanced microbial metabolic activity in terms of soil respiration per amount of microbial biomass C. Microbial stress biomarkers were elevated in warmed plots. In summary, the 4 °C increase in soil temperature during the snow-free season had no influence on microbial community composition and biomass but strongly increased microbial metabolic activity and hence reduced carbon use efficiency.     

Soil microbial biomass C:N:P stoichiometry and microbial use of organic phosphorus

Microbial mineralization and immobilization of nutrients strongly influence soil fertility. We studied microbial biomass stoichiometry, microbial community composition, and microbial use of carbon (C) and phosphorus (P) derived from glucose-6-phosphate in the A and B horizons of two temperate Cambisols with contrasting P availability. In a first incubation experiment, C, nitrogen (N) and P were added to the soils in a full factorial design. Microbial biomass C, N and P concentrations were analyzed by the fumigation-extraction method and microbial community composition was analyzed by a community fingerprinting method (automated ribosomal intergenic spacer analysis, ARISA). In a second experiment, we compared microbial use of C and P from glucose-6-phosphate by adding ¹⁴C or ³³P labeled glucose-6-phosphate to soil. In the first incubation experiment, the microbial biomass increased up to 30-fold due to addition of C, indicating that microbial growth was mainly C limited. Microbial biomass C:N:P stoichiometry changed more strongly due to element addition in the P-poor soils, than in the P-rich soils. The microbial community composition analysis showed that element additions led to stronger changes in the microbial community in the P-poor than in the P-rich soils. Therefore, the changed microbial biomass stoichiometry in the P-poor soils was likely caused by a shift in the microbial community composition. The total recovery of ¹⁴C derived from glucose-6-phosphate in the soil microbial biomass and in the respired CO₂ ranged between 28.2 and 37.1% 66 h after addition of the tracer, while the recovery of ³³P in the soil microbial biomass was 1.4–6.1%. This indicates that even in the P-poor soils microorganisms mineralized organic P and took up more C than P from the organic compound. Thus, microbial mineralization of organic P was driven by microbial need for C rather than for P. In conclusion, our experiments showed that (i) the microbial biomass stoichiometry in the P-poor soils was more susceptible to additions of C, N and P than in the P-rich soils and that (ii) even in the P-poor soils, microorganisms were C-limited and the mineralization of organic P was mainly driven by microbial C demand.     

基于meta分析的放牧压力对内蒙古高原草地生态系统的影响

放牧是最主要的草地利用模式,直接或间接地影响草地物质循环和能量流动,放牧强度对草地的健康状况和演替方向起决定作用。本文基于40篇内蒙古草原放牧相关文献数据,通过meta分析探讨温带草原对放牧强度的响应特征。结果表明,与未放牧草地相比,轻度放牧草地对群落植物地上、地下生物量和土壤全氮和全磷含量无显著影响,而土壤有机碳、微生物生物量碳、细菌和真菌数量分别显著上升3.60%、7.80%、11.40%和10.83%（P<0.05）;中度放牧下群落植物地下生物量和土壤微生物数量无显著变化,而地上生物量和土壤有机碳、全氮、全磷和微生物生物量氮含量分别显著降低21.62%、4.44%、2.15%、8.35%和6.76%（P<0.05）;重度放牧下群落植物地上和地下生物量,土壤有机碳、全氮、全磷、微生物生物量碳含量,细菌和放线菌数量分别显著下降39.72%、16.30%、7.62%、6.46%、8.03%、8.76%、12.92%和18.27%（P<0.05）。以上结果表明轻度放牧有利于土壤肥力和草地生产力的保持和提升,而当放牧干扰超出一定的限度时,草地各项功能均显著下降而发生退化。本研究可为内蒙古温带草原的合理利用和适应性管理提供理论基础。     

Labile,recalcitrant, microbial carbon and nitrogen and the microbial community composition at two Abies faxoniana forest elevations under elevated temperatures

We investigated the interactions of altitude and artificial warming on the soil microbial community structure in a subalpine Abies faxoniana forest in southwestern China after four years of warming. Open top chambers (OTCs) at two elevations (3000 m and 3500 m) were established, and their soil microbial characteristics, organic carbon (C) and nitrogen (N) were measured. The microbial community structure was quantified by phospholipid fatty acid (PLFA) analysis. A two-step sulfuric acid hydrolysis was used to quantify the labile and recalcitrant C fractions in the soil organic matter. The results showed that bacterial PLFAs and gram-negative bacterial PLFAs increased and the fungal PLFAs and the fungi/bacteria ratio decreased with warming at the high altitude. By contrast, the warming effects on those parameters at low altitude were small. The higher proportion of labile easily decomposable soil C may explain the different responses of the microbial community composition at the two altitudes. An RDA analysis confirmed that the variations in the soil community structure were significantly associated with soil organic matter properties such as the sizes of the soil labile N pool (LP-N), the recalcitrant N pool (RP-N), and the labile C pool as well as dissolved organic C (DOC) and dissolved organic N concentrations (DON). Our results also showed that labile C and N pools increased with the altitude, but the microbial biomass C as measured with chloroform fumigation techniques decreased. Warming increased only the recalcitrant C pools at the high altitude. Given the longer mean residence time for recalcitrant C and the much greater size of this soil organic carbon pool, the results indicated that a rise in temperature in our case increased soil C pools at higher altitudes, at least during the early stages of experimental soil warming. Warming could also cause changes in the composition of the microbial community and enzyme activities, consequently leading to functional changes in soil ecosystem processes at the high altitude.     

废弃农耕地自然演替过程中微生物群落与土壤特性间的关系

The changes of microbial biomass carbon (MBC) and nitrogen (MBN) and microbial community in the topsoil of the abandoned agricultural land on the semi-arid Loess Plateau in China during the natural succession were evaluated to understand the relationship between microbial community and soil properties. MBC and MBN were measured using fumigation extraction, and microbial community was analyzed by the method of fatty acid methyl ester (FAME). The contents of organic C, total N, MBC, MBN, total FAME, fungal FAME, bacterial FAME and Gram-negative bacterial FAME at the natural succession sites were higher than those of the agricultural land, but lower than those of the natural vegetation sites. The MBC, MBN and total FAME were closely correlated with organic C and total N. Furthermore, organic C and total N were found to be positively correlated with fungal FAME, bacterial FAME, fungal/bacterial and Gram-negative bacterial FAME. Natural succession would be useful for improving soil microbial properties and might be an important alternative for sustaining soil quality on the semi-arid Loess Plateau in China.     

Soil organic matter quality as influenced by tillage,lime, and phosphorus

In Eastern Canada, cereal yields are often restricted by soil acidity and low fertility. Continuous cereal production can also lead to soil structural degradation. The addition of lime and fertilizers and the adoption of conversation tillage practices are proposed solutions which may have a positive impact on soil quality. The objective of the present work was to assess the impact of 3 years of different tillage practices and P additions, and of a single lime addition on organic C and total N, microbial biomass C, and on N mineralization at the surface layer (0–7.5 cm) of a Courval sandy clay loam (Humic Gleysol). The easily mineralizable N, total amount of N mineralized in 22.1 weeks, the rate of N mineralization, and microbial biomass C were significantly greater in the minimum tillage than in the moldboard plow treatment. Chisel plow treatment showed intermediate values. The ratios of potentially mineralizable N and of easily mineralizable to total soil N were also significantly larger under minimum tillage and chisel plowing than under moldboard plowing. The lime and P treatments had no significant effect on the measured soil quality parameters. The total amount of N mineralized per unit of biomass C decreased as the tillage intensity increased, suggesting a decrease in the efficiency of the biomass in transforming organic N into potentially plant-available forms and thus a loss in soil organic matter quality. The results of this study indicate that conservation tillage practices such as rototilling and chisel plowing are efficient ways of maintaining soil organic matter quality when old pastures are brought back into cultivation.     

土壤微生物量碳氮作为土壤肥力指标的探讨

通过对不同肥力水平黑土、棕壤、黄棕壤、红壤土壤微生物量碳氮的比较,研究和探讨土壤微生物量碳氮与土壤肥力的关系。结果表明:有机肥或无机肥施入土壤均能增加土壤微生物量C、N的含量,且有机肥与无机肥配合有显著的正交互作用。土壤微生物量C、N从北到南的地带性变化规律不强。BC/BN及TC/TN从北到南逐渐减小,BC/TC及BN/TN从北到南逐渐增加。相关分析结果表明:土壤微生物量C、N与全C、全N均呈极显著相关,可以作为指示土壤肥力的重要指标。     

CN-SIM—a model for the turnover of soil organic matter. I. Long-term carbon and radiocarbon development

A novel computer model is presented which describes the flow of C and N in the soil. It employs a structure with conceptual compartments. Organic matter is represented by seven different compartments, two for added matter, two for soil microbial biomass, one for microbial residues, one for native (‘humified’) organic matter, and one for inert organic matter. The latter pool represents both truly inert matter, and matter with negligible turnover in a time-span of decades to a century. This paper describes the parameterisation and performance of this model on selected long-term field carbon and radiocarbon data from United Kingdom, Sweden and Denmark. Previously unpublished radiocarbon data series from Denmark are included. Statistical methods were employed to estimate parameters, and obtain proximate confidence intervals for these parameters. Simulations in good agreement with measured values could be achieved, using the same set of parameters on all sites. It was demonstrated that the inert pool might constitute any amount between approx. 10 and 50% of total soil C, so that modelling cannot be used as a tool to obtain narrow estimates for this pool.     

Carbon and nitrogen mineralization in acidic, limed and calcareous agricultural soils: Apparent and actual effects

Soil pH and calcium carbonate contents are often hypothesized to be important factors controlling organic matter turnover in agricultural soils. The aim of this study was to differentiate the effects of soil pH from those related to carbonate equilibrium on C and N dynamics. The relative contributions of organic and inorganic carbon in the CO₂ produced during laboratory incubations were assessed. Five agricultural soils were compared: calcareous (74% CaCO₃), loess (0.2% CaCO₃) and an acidic soil which had received different rates of lime 20 years ago (0, 18 or 50 t ha⁻¹). Soil aggregates were incubated with or without rape residues under aerobic conditions for 91 days at 15 °C. The C and N mineralized, soil pH, O₂ consumption and respiratory quotient (RQ=ΔCO₂/ΔO₂) were monitored, as well as the δ¹³C composition of the evolved CO₂ to determine its origin (mineral or organic). Results showed that in non-amended soils, the cumulative CO₂ produced was significantly greater in the limed soil with a pH>7 than in the same soil with less or no lime added, whereas there was no difference in N mineralization or in O₂ consumption kinetics. We found an exponential relationship between RQ values and soil pH, suggesting an excess production of CO₂ in alkaline soils. This CO₂ excess was not related to changes in substrate utilization by the microbial biomass but rather to carbonates equilibrium. The δ¹³C signatures confirmed that the CO₂ produced in soils with pH>7 originated from both organic and mineral sources. The contribution of soil carbonates to CO₂ production led to an overestimation of organic C mineralization (up to 35%), the extent of which depended on the nature of soil carbonates but not on the amount. The actual C mineralization (derived from organic C) was similar in limed and unlimed soil. The amount of C mineralized in the residue-amended soils was ten times greater than in the basal soil, thus masking the soil carbonate contribution. Residue decomposition resulted in a significant increase in soil pH in all soils. This increase is attributed to the alkalinity and/or decarboxylation of organic anions in the plant residue and/or to the immobilization of nitrate by the microbial biomass and the corresponding release of hydroxyl ions. A theoretical composition (C, O, H, N) of residue and soil organic matter is proposed to explain the RQ measured. It emphasizes the need to take microbial biomass metabolism, O₂ consumption due to nitrification and carbon assimilation yield into account when interpreting RQ data.     

Nutrient limitations to soil microbial biomass and activity in loblolly pine forests

We performed an assay of nutrient limitations to soil microbial biomass in forest floor material and intact cores of mineral soil collected from three North Carolina loblolly pine (Pinus taeda) forests. We added solutions containing C, N or P alone and in all possible combinations, and we measured the effects of these treatments on microbial biomass and on microbial respiration, which served as a proxy for microbial activity, during a 7-day laboratory incubation at 22 °C. The C solution used was intended to simulate the initial products of fine root decay. Additions of C dramatically increased respiration in both mineral soil and forest floor material, and C addition increased microbial biomass C in the mineral soil. Additions of N increased respiration in forest floor material and increased microbial biomass N in the mineral soil. Addition of P caused a small increase in forest floor respiration, but had no effect on microbial biomass.     

Humification-mineralization pyrolytic indices and carbon fractions of soil under organic and conventional management in central Italy

The aim of this study was to evaluate chemical changes in soil organic matter (SOM) in organically and conventionally managed fields, using pyrolytic indices and the extraction of different carbon fractions. Pyrolysis-gas chromatography (Py-GC) was used to study structural changes in SOM, whereas the different soil extractions gave a fractionation of C forms. Organic management increased both humic and labile C forms (microbial biomass C and water soluble organic C). A significant positive relationship was found between the living SOM fraction, expressed as microbial biomass/total organic C ratio (MBC/TOC) and humification rate. A negative relationship was found between the pyrrole to phenol ratio (O/Y) and total extractable C (TEC).An opposite trend has been observed for the second pyrolytic index (N/O), which represents the mineralization of fresh organic matter. Mineralization was higher in organically managed soil, probably because of consistent input of fresh material to the organic field. Carbon fraction pools and pyrolytic indices provided complementary indications of SOM quality under organic and conventional management.     

Microbial community composition and substrate use in a highly weathered soil as affected by crop rotation and P fertilization

A better understanding of soil microbial processes is required to improve the synchrony between nutrient release from plant residues and crop demand. Phospholipid fatty acid analysis was used to investigate the effect of two crop rotations (continuous maize and maize-crotalaria rotation) and P fertilization (0 and 50 kg P ha⁻¹ yr⁻¹, applied as triple superphosphate) on microbial community composition in a highly weathered soil from western Kenya. Microbial substrate use in soils from the field experiment was compared in incubation experiments. Higher levels of soil organic matter and microbial biomass in the maize-crotalaria rotation were connected with higher total amounts of phospholipid fatty acids and an increase in the relative abundances of indicators for fungi and gram-negative bacteria. P fertilization changed the community profile only within the continuous maize treatment. The decomposition of glucose, cellulose and three plant residues (all added at 2.5 g C kg⁻¹ soil) proceeded faster in soil from the maize-crotalaria rotation, but differences were mostly transient. Microbial P and N uptake within one week increased with the water-soluble carbon content of added plant residues. More P and N were taken up by the greater microbial biomass in soil from the maize-crotalaria rotation than from continuous maize. Re-mineralization of nutrients during the decline of the microbial biomass increased also with the initial biological activity of the soil, but occurred only for a high quality plant residue within the half year incubation period. Compared to the effect of crop rotation, P fertilization had a minor effect on microbial community composition and substrate use.     

Is there a linear relationship between priming effect intensity and the amount of organic matter input?

Inputs of fresh organic matter (FOM) are known to affect the rate of soil organic matter (SOM) mineralization. SOM mineralization can be accelerated or decelerated by FOM inputs. This phenomenon, known as the Priming effect (PE), may largely influence the carbon (C) storage capacity of soils. However, the link between PE intensity and FOM inputs is not clearly understood. Indeed, almost all the studies about PE used only one FOM amount which is generally largely below the amount of FOM observed in field conditions. In our study, we incubated soil amended with three levels of ¹³C-labeled straw as FOM and a control without FOM amendment for 80 days. The three levels used were in the same range as the natural FOM inputs observed on our sampling site. Various levels of mineral nitrogen were added within each level of straw supply so that the final input C:N ratios ranged among 44, 30 and 20. CO₂ and δ¹³C-CO₂ were measured during the experiment allowing us to distinguish the FOM respired CO₂ from the SOM respired CO₂. We observed that PE intensity did not increase linearly with increasing FOM additions. Moreover, decreasing the input C:N ratios did not systematically affect PE intensity probably because of shifts in the microbial characteristics such as their C:N ratio or their assimilation yields. These results suggest that PE is a saturating function of FOM inputs that is only weakly influenced by initial N availability. Our results may be explained (i) by the existence of a limited SOM pool subject to PE (ii) or by the occurrence of two simultaneous and antagonistic mechanisms: an increase of the total active microbial biomass accelerating SOM mineralization (i.e. a positive PE) and a preferential substrate utilization of FOM over SOM decreasing SOM mineralization (i.e. a negative PE). Finally, irrespective of the mechanisms implied, our results suggest that the importance of positive PE relatively to the amount of FOM may decrease when FOM inputs increase, which is favorable to carbon sequestration in soils. Indeed, in the case of the lower amount of FOM, the PE corresponded to 6.25% of the total amount of CO₂ mineralized at the end of the experiment while, for the higher amount of FOM, the PE corresponded to 5% of the total amount of CO₂ mineralized at the end of the experiment.     

Mineralization of soil organic matter initiated by the application of an available substrate to the profiles of surface and buried podzolic soils

The priming effect (PE) initiated by the application of ¹⁴C-glucose was studied for copiotrophic microbial communities of organic horizons and for oligotrophic microbial communities of mineral soil horizons, as well as for mineral horizons of buried soils depleted in the input of fresh organic matter. The intensity of the PE depended on the reserves of C_org, the initial amount of the microbial biomass, and the enzymatic activity, which decreased from the organic to the mineral soil horizons. The ratio of the PE to the applied carbon was two times higher in the mineral horizons as compared with the organic horizons. This is explained by the predominance of K-strategists capable of decomposing difficultly available organic compounds in the mineral horizons, so that the turnover of the microbial biomass in the mineral horizons was more active than that in the organic horizons. The predominance of K-strategists was confirmed by the close correlation between the PE and the activity of the cellobiohydrolase enzyme decomposing cellulose (R = 0.96). In general, the absolute value of the PE was controlled by the soil organic matter content, whereas the specific PE was controlled by the functional features of the microorganisms. It was shown that the functional features of the soil microorganisms remained unchanged under the conditions of their preservation in the buried soil.     

Organic matter, microbial biomass and enzyme activity of soils under different crop rotations in the tropics

Soil organic matter level, soil microbial biomass C, ninhydrin-N, C mineralization, and dehydrogenase and alkaline phosphatase activity were studied in soils under different crop rotations for 6 years. Inclusion of a green manure crop of Sesbania aculeata in the rotation improved soil organic matter status and led to an increase in soil microbial biomass, soil enzyme activity and soil respiratory activity. Microbial biomass C increased from 192 mg kg^–1 soil in a pearl millet-wheat-fallow rotation to 256 mg kg^–1 soil in a pearl millet-wheat-green manure rotation. Inclusion of an oilseed crop such as sunflower or mustard led to a decrease in soil microbial biomass, C mineralization and soil enzyme activity. There was a good correlation between microbial biomass C, ninhydrin-N and dehydrogenase activity. The alkaline phosphatase activity of the soil under different crop rotations was little affected. The results indicate the green manuring improved the organic matter status of the soil and soil microbial activity vital for the nutrient turnover and long-term productivity of the soil. Received: 7 January 1996     

Microbial phospholipid biomarkers and stable isotope methods help reveal soil functions

This review targets microbial phospholipid biomarkers, their isotope analysis and their ability to reveal soil functions. The amount and composition of phospholipid fatty acids (PLFAs) measured in environmental samples strongly depend on the methodology. To achieve comparable results the extraction, separation and methylation method must be kept constant. PLFAs patterns are sensitive to microbial community shifts even though the taxonomic resolution of PLFAs is low. The possibility to easily link lipid biomarkers with stable isotope techniques is identified as a major advantage when addressing soil functions. Measurement of PLFA isotopic ratios is sensitive and enables detecting isotopic fractionation. The difference between the carbon isotopic ratio of single PLFAs and their substrate (Δ¹³C) can vary between −6 and +11‰. This difference derives from the fractionation during biosynthesis and from substrate inhomogeneity. Consequently, natural abundance studies are restricted to quantifying substrate uptake of the total microbial biomass. In contrast, artificial labelling enables quantifying carbon uptake into single PLFAs, but labelling success depends on homogeneous and undisturbed label application. Current developments in microbial ecology (e.g. ¹³C and ¹⁵N proteomics) and isotope techniques (online monitoring of CO₂ isotope ratios) will likely improve soil functional interpretations in the future. ¹³C PLFA analysis will continue to contribute because it is affordable, sensitive and allows frequent sampling combined with the use of small amounts of ¹³C label.     

Surface and subsurface microbial biomass, community structure and metabolic activity as a function of soil depth and season

Microbial biomass, size and community structure along with an estimate of microbial activity and soil chemical parameters were determined at three depths in two soils (e.g. sandy loam Ultic Hapludalf and silt loam Mollic Hapludalf) replicated three times under one winter and summer season. Microbial biomass and community structure were estimated from phospholipid-PO₄ content and fatty acid methyl ester (FAME) measurements. Microbial activity and assimilative capacity were estimated using a ³H-acetate incorporation into phospholipids and by incubating the soil samples at the average winter and summer temperatures, 3 and 20 °C, respectively. We found that the size of the microbial biomass in both the surface and the subsurface soils was not significantly affected by the seasonal variation but activity increased by as much as 83% at the summer temperatures in the surface soil. We demonstrated using FAME analysis that for both soils seasonal changes in the subsurface microbial community occurred. These findings suggest that winter conditions will shift the population activity level in both the surface and subsurface systems and the biochemical structure of the community in the subsurface. In all cases, the inorganic chemical properties of the soil, as a function of season, remained constant. The greatly increased activity of microbial population at the higher temperature will favor the capacity of the system to utilize nutrients or organic materials that may enter soil. During low temperature seasons the capacity of either surface or subsurface soils to assimilate materials is generally diminished but the reduction reflects changes in metabolism and not a reduced biomass size.