Effect of Collembola on mineralization of litter and soil organic matter

Although soil Collembola are known to contribute to soil carbon (C) cycling, their contribution to the mineralization of C sources that differ in bioavailability, such as soil organic C (SOC) and leaf litter, is unknown. Stable C isotopes are often used to quantify the effects of both soil C and litter C on C mineralization. Here, ¹³C-labeled litter was used to investigate the effects of Collembola (Folsomia candida) on the mineralization of both SOC and litter C in laboratory microcosms. The three microcosm treatments were soil alone (S); soil treated with δ¹³C-labeled litter (SL); and soil treated with δ¹³C-labeled litter and Collembola (SLC). The presence of Collembola did not significantly affect soil microbial biomass or litter mass loss and only had a small effect on CO₂ release during the first week of the experiment, when most of the CO₂ was derived from litter rather than from SOC. Later, during the experiment (days 21 and 63), when litter-derived labile C had been depleted and when numbers of Collembola had greatly increased, Collembola substantially increased the emission of SOC-derived CO₂. These results suggest that the effect of Collembola on soil organic C mineralization is negatively related to C availability.     

Carbon fluxes in soil food webs of increasing complexity revealed by C labelling and C natural abundance

Soil food webs are mainly based on three primary carbon (C) sources: root exudates, litter, and recalcitrant soil organic matter (SOM). These C sources vary in their availability and accessibility to soil organisms, which could lead to different pathways in soil food webs. The presence of three C isotopes (¹²C, ¹³C and ¹⁴C) offers an unique opportunity to investigate all three C sources simultaneously. In a microcosm experiment we studied the effect of food web complexity on the utilization of the three carbon sources. We choose an incomplete three factorial design with (i) living plants, (ii) litter and (iii) food web complexity. The most complex food web consisted of autochthonous microorganisms, nematodes, collembola, predatory mites, endogeic and anecic earthworms. We traced C from all three sources in soil, in CO₂ efflux and in individual organism groups by using maize grown on soil developed under C₃ vegetation and application of ¹⁴C labelled ryegrass shoots as a litter layer. The presence of living plants had a much greater effect on C pathways than food web complexity. Litter decomposition, measured as ¹⁴CO₂ efflux, was decreased in the presence of living plants from 71% to 33%. However, living plants increased the incorporation of litter C into microbial biomass and arrested carbon in the litter layer and in the upper soil layer. The only significant effect of food web complexity was on the litter C distribution in the soil layers. In treatments with fungivorous microarthropods (Collembola) the incorporation of litter carbon into mineral soil was reduced. Root exudates as C source were passed through rhizosphere microorganisms to the predator level (at least to the third trophic level). We conclude that living plants strongly affected C flows, directly by being a source of additional C, and indirectly by modifying the existing C flows within the food web including CO₂ efflux from the soil and litter decomposition.     

Collembola switch diet in presence of plant roots thereby functioning as herbivores

Collembola are abundant and ubiquitous soil decomposers, being particularly active in the rhizosphere of plants where they are assumed to be attracted by high microbial activity and biomass. While feeding on root associated microorganisms or organic matter they may also ingest plant roots, e.g. particularly root hairs and fine roots. Employing stable isotope analysis we investigated Collembola (Protaphorura fimata Gisin) feeding preferences and types of ingested resources. We offered Collembola two resources with distinct isotope signatures: a C4 plant (Zea mays L.) planted in soil mixed with ¹⁵N labelled litter of Lolium perenne L. (C3 plant). We hypothesised that Collembola obtain their nutrients (C and N) from different resources, with their carbon being mainly derived from resources that are closely associated to the plant root, e.g. root exudates, causing enrichment in ¹³C in Collembola tissue, while the incorporated nitrogen originating from litter resources. In contrast to our hypothesis, stable isotope analysis suggests that in absence of plant roots Collembola derived both the incorporated C and N predominantly from litter whereas in presence of plant roots they switched diet and obtained both C and N almost exclusively from plant roots.The results indicate that Collembola in the rhizosphere of plants, being assumed to be mainly decomposers, in fact predominately live on plant resources, presumably fine roots or root hairs, i.e. are herbivorous rather than detritivorous or fungivorous. These findings have major implications on the view how plants respond to decomposers in the rhizosphere.     

Impact of litter quality on mineralization processes in managed and abandoned pasture soils in Southern Ecuador

Tropical regions are currently undergoing remarkable rates of land use change accompanied by altered litter inputs to soil. In vast areas of Southern Ecuador forests are clear cut and converted for use as cattle pastures. Frequently these pasture sites are invaded by bracken fern, when bracken becomes dominant pasture productivity decreases and the sites are abandoned. In the present study implications of invasive bracken on soil biogeochemical properties were investigated. Soil samples (0-5 cm) were taken from an active pasture with Setaria sphacelata as predominant grass and from an abandoned pasture overgrown by bracken. Grass (C₄ plant) and bracken (C₃ plant) litter, differing in C:N ratio (33 and 77, respectively) and lignin content (Klason-lignin: 18% and 45%, respectively), were incubated in soils of their corresponding sites and vice versa for 28 days at 22 °C. Unamended microcosms containing only the respective soil or litter were taken as controls. During incubation the amount of CO₂ and its δ¹³C-signature were determined at different time intervals. Additionally, the soil microbial community structure (PLFA-analysis) as well as the concentrations of KCl-extractable C and N were monitored. The comparison between the control soils of active and abandoned pasture sites showed that the massive displacement of Setaria-grass by bracken after pasture abandonment was characterized by decreased pH values accompanied by decreased amounts of readily available organic carbon and nitrogen, a lower microbial biomass and decreased activity as well as a higher relative abundance of actinomycetes. The δ¹³C-signature of CO₂ indicated a preferential mineralization of grass-derived organic carbon in pasture control soils. In soils amended with grass litter the mineralization of soil organic matter was retarded (negative priming effect) and also a preferential utilization of easily available organic substances derived from the grass litter was evident. Compared to the other treatments, the pasture soil amended with grass litter showed an opposite shift in the microbial community structure towards a lower relative abundance of fungi. After addition of bracken litter to the abandoned pasture soil a positive priming effect seemed to be supported by an N limitation at the end of incubation. This was accompanied by an increase in the ratio of Gram-positive to Gram-negative bacterial PLFA marker. The differences in litter quality between grass and bracken are important triggers of changes in soil biogeochemical and soil microbial properties after land use conversion.     

Collembolan trophic preferences determined using fatty acid distributions and compound-specific stable carbon isotope values

The trophic preferences of soil invertebrates such as Collembola are often determined by the analysis of gut contents, or through visual observations of the location of individuals. As an alternative approach, two species of Collembola, Folsomia candida and Proisotoma minuta, were offered a choice of the soil fungus Cladosporium cladosporioides or the bacterial feeding nematode Panagrellus redivivus; each exhibited distinct fatty acid profiles and stable carbon isotopic compositions. Over 21 days, the fatty acids i15:0, i17:0, 18:1(n-7) and 18:2(n-6) all increased in abundance in both collembolan species consistent with direct routing from the nematode dietary choice which contained a high concentration of these components. Collembolan fatty acid δ¹³C values increased by between 5.7 and 21.6‰ over 21 days reflecting those of the nematode diet. Therefore, both fatty acid profiles and δ¹³C values were consistent with a strong feeding preference of F. candida and P. minuta for the nematodes over the offered fungi. In fact, neither collembolan species consumed any detectable amount of C. cladosporioides. Comparison of the δ¹³C values of the 16:0 and 18:0 fatty acids (which are biosynthesised by the Collembola as well as directly incorporated from the diet) and the 16:1(n-7) and 18:2(n-6) components (which are not biosynthesised by the Collembola) demonstrated that the input of distinct pools of C can lead to large shifts in δ¹³C values between diet and consumer. The fatty acids that were not biosynthesised by Collembola better reflected the δ¹³C values of the diet helping to differentiate between biosynthesised and directly incorporated compounds; an important prerequisite in the interpretation of compound-specific δ¹³C values in trophic behaviour tests. The combination of fatty acid distributions and δ¹³C values is a significant improvement on traditional methods of examining feeding preferences, since it determines directly the assimilated dietary carbon rather than relying on indirect observations, such as the proximity of individuals to a defined food source.     

The effect of diet on isotopic turnover in Collembola examined using the stable carbon isotopic compositions of lipids

Combined compound-specific stable carbon isotopic methods and fatty acid abundance determinations have been used to examine feeding preferences and C allocation in organisms where direct observation of feeding is difficult. In order to examine the effect of differing diets on the δ¹³C values of fatty acids and sterols of Collembola, the diets of two collembolan species, Folsomia candida and Proisotoma minuta, were switched from a yeast diet to one of four isotopically distinct diets, and the δ¹³C values of the lipids monitored over the next 39 d. Cholesterol remained the only sterol detected in both collembolan species, despite the diets containing widely differing sterol compositions. The δ¹³C values of collembolan lipids recorded after long term feeding were often different to those of the same components in the diet, indicating that fractionation or partitioning occurs during digestion, assimilation and biosynthesis within the Collembola, thereby shifting consumer lipid δ¹³C values away from those of the corresponding dietary components. The rates of change of δ¹³C values differed among compounds, with half-lives ranging between 29 min and 14 d. Some of these differences appear to be related to the abundance of dietary components, such that fatty acids present in high abundance in the diet (e.g. 18:2(n−6)) were rapidly assimilated in high proportions into collembolan lipids, leading to a rapid change in δ¹³C values. Similarly, isotopic turnover in the 16:1(n−7) fatty acid, present in the newly presented diets in only low abundances, was significantly correlated to the rate of removal of this component from the consumer fatty acid pool. The rates of change of δ¹³C values in P. minuta lipids did not vary significantly with diet, whilst the rates of change of δ¹³C values of lipids in F. candida were affected by the diets the Collembola consumed. Results of an experiment providing F. candida and P. minuta with two diets of different quality demonstrated that F. candida responded to the high quality diet with increased growth and fecundity, whilst P. minuta responded with increased fecundity only. Thus, the abilities of the two species to respond to diets of varying quality, amongst other factors, is concluded to lead to differences in the rates of change of δ¹³C values reflecting differences in lipid turnover.     

Consumption of mycorrhizal and saprophytic fungi by Collembola in grassland soils

Although soil-dwelling Collembola can influence plant growth and nutrient cycling, their specific role in soil food webs is poorly understood. Soil-free microcosm studies suggest that Collembola are primarily fungivores where they feed preferentially on saprophytic fungi (SF) over other fungal types. We directly assessed collembolan consumption of arbuscular mycorrhizal fungi (AMF) and SF using plant-soil mesocosms and natural abundance stable carbon isotope techniques. Mycorrhizal Andropogon gerardii (C₄ grass) seedlings were placed in pots containing Collembola and soil from a C₃ plant dominated site, while mycorrhizal Pascopyrum smithii (C₃ grass) seedlings were placed in pots with Collembola and soil collected at a C₄ plant dominated site. After 6 weeks, collembolans assimilated carbon derived from C₃ and C₄ sources in both A. gerardii and P. smithii treatments. Comparing Collembola isotope values in AMF vs. AMF-suppressed treatments, our data show that both AMF and SF were consumed in these experimental soil environments.     

Formation and use of microbial residues after adding sugarcane sucrose to a heated soil devoid of soil organic matter

A 67-day incubation experiment was carried out with a soil initially devoid of any organic matter due to heating, which was amended with sugarcane sucrose (C₄-sucrose with a δ¹³C value of ?10.5‰), inorganic N and an inoculum for recolonisation and subsequently at day 33 with C₃-cellulose (δ¹³C value of ?23.4‰). In this soil, all organic matter is in the microbial biomass or in freshly formed residues, which makes it possible to analyse more clearly the role of microbial residues for decomposition of N-poor substrates. The average δ¹³C value over the whole incubation period was ?10.7‰ in soil total C in the treatments without C₃-cellulose addition. In the CO₂ evolved, the δ¹³C values decreased from ?13.4‰ to ?15.4‰ during incubation. In the microbial biomass, the δ¹³C values increased from ?11.5‰ to ?10.1‰ at days 33 and 38. At day 67, 36% of the C₄-sucrose was left in the treatment without a second amendment. The addition of C₃-cellulose resulted in a further 7% decrease, but 4% of the C₃-cellulose was lost during the second incubation period. Total microbial biomass C declined from 200 μg g^?1 soil at day 5 to 70 μg g^?1 soil at day 67. Fungal ergosterol increased to 1.5 μg g^?1 soil at day 12 and declined more or less linearly to 0.4 μg g^?1 soil at day 67. Bacterial muramic acid declined from a maximum of 35 μg g^?1 soil at day 5 to a constant level of around 16 μg g^?1 soil. Glucosamine showed a peak value at day 12. Galactosamine remained constant throughout the incubation. The fungal C/bacterial C ratio increased more or less linearly from 0.38 at day 5 to 1.1 at day 67 indicating a shift in the microbial community from bacteria to fungi during the incubation. The addition of C₃-cellulose led to a small increase in C₃-derived microbial biomass C, but to a strong increase in C₄-derived microbial biomass C. At days 45 and 67, the addition of N-free C₃-cellulose significantly decreased the C/N ratio of the microbial residues, suggesting that this fraction did not serve as an N-source, but as an energy source.     

Atmospheric CO2 enrichment induces life strategy- and species-specific responses of collembolans in the rhizosphere of sugar beet and winter wheat

We studied atmospheric CO₂ enrichment effects on life form types, species composition, dominance structure and individual density of collembolans under cultivation of sugar beet and winter wheat. The study was part of a long-term CO₂ enrichment field experiment (FACE: Free Air CO₂ Enrichment) at the Federal Agricultural Research Centre (FAL) in Braunschweig (Germany), using isotopically labelled CO₂. The stable C-isotopic signature (δ¹³C) of collembolan species, plant material, and soil indicated CO₂ impacts on C translocation. The δ¹³C values of both crops significantly increased from above-ground to below-ground plant parts and significantly decreased under FACE conditions. The δ¹³C values of collembolan species differed significantly depending on CO₂ treatment and crop and showed a distinct tendency depending on plant growth stage. The extent, to which δ¹³C values of collembolans decreased under FACE conditions, was species- and life strategy-dependent. The stable C-isotopic signatures of euedaphic and hemiedaphic species were similar in the control, but, depending on crop, differently affected by atmospheric CO₂ enrichment. Under winter wheat cultivation, hemiedaphic species showed more negative δ¹³C values than euedaphic ones under FACE conditions. CO₂ enrichment effects on occurrence, density and dominance distribution of the collembolan species differed strongly between crops and their developmental stages, which reveal crop-specific below-ground effects due to different food qualities in the rhizosphere. CO₂ impacts were stronger under sugar beet compared to winter wheat cultivation. Independent of crop, CO₂ enrichment enhanced the diversity of collembolans before harvest and increased the proportion of hemiedaphic in relation to euedaphic species in a community. Our results on collembolan communities imply CO₂-induced changes in the root-derived carbon resources used by the soil food web. The present study reveals atmospheric CO₂ enrichment impacts to specifically affect collembolan species according to their food preferences.     

Three-source partitioning of CO2 efflux from soil planted with maize by C natural abundance fails due to inactive microbial biomass

A theoretical approach to the partitioning of carbon dioxide (CO₂) efflux from soil with a C₃ vegetation history planted with maize (Zea mays), a C₄ plant, into three sources, root respiration (RR), rhizomicrobial respiration (RMR), and microbial soil organic matter (SOM) decomposition (SOMD), was examined. The δ¹³C values of SOM, roots, microbial biomass, and total CO₂ efflux were measured during a 40-day growing period. A three-source isotopic mass balance based on the measured δ¹³C values and on assumptions made in other studies showed that RR, RMR, and SOMD amounted to 91%, 4%, and 5%, respectively. Two assumptions were thoroughly examined in a sensitivity analysis: the absence of ¹³C fractionation and the conformity of δ¹³C of microbial CO₂ and that of microbial biomass. This approach strongly overestimated RR and underestimated RMR and microbial SOMD. CO₂ efflux from unplanted soil was enriched in ¹³C by 2.0‰ compared to microbial biomass. The consideration of this ¹³C fractionation in the mass balance equation changed the proportions of RR and RMR by only 4% and did not affect SOMD. A calculated δ¹³C value of microbial CO₂ by a mass balance equation including active and inactive parts of microbial biomass was used to adjust a hypothetical below-ground CO₂ partitioning to the measured and literature data. The active microbial biomass in the rhizosphere amounted to 37% to achieve an appropriate ratio between RR and RMR compared to measured data. Therefore, the three-source partitioning approach failed due to a low active portion of microbial biomass, which is the main microbial CO₂ source controlling the δ¹³C value of total microbial biomass. Since fumigation-extraction reflects total microbial biomass, its δ¹³C value was unsuitable to predict δ¹³C of released microbial CO₂ after a C₃-C₄ vegetation change. The second adjustment to the CO₂ partitioning results in the literature showed that at least 71% of the active microbial biomass utilizing maize rhizodeposits would be necessary to achieve that proportion between RR and RMR observed by other approaches based on ¹⁴C labelling. The method for partitioning total below-ground CO₂ efflux into three sources using a natural ¹³C labelling technique failed due to the small proportion of active microbial biomass in the rhizosphere. This small active fraction led to a discrepancy between δ¹³C values of microbial biomass and of microbially respired CO₂.     

Carbon input belowground is the major C flux contributing to leaf litter mass loss: Evidences from a C labelled-leaf litter experiment

Partitioning of the quantities of C lost by leaf litter through decomposition into (i) CO₂ efflux to the atmosphere and (ii) C input to soil organic matter (SOM) is essential in order to develop a deeper understanding of the litter-soil biogeochemical continuum. However, this is a challenging task due to the occurrence of many different processes contributing to litter biomass loss. With the aim of quantifying different fluxes of C lost by leaf litter decomposition, a field experiment was performed at a short rotation coppice poplar plantation in central Italy. Populus nigra leaf litter, enriched in ¹³C (δ¹³C ∼ +160‰) was placed within collars to decompose in direct contact with the soil (δ¹³C ∼ −26‰) for 11 months. CO₂ efflux from within the collars and its isotopic composition were determined at monthly intervals. After 11 months, remaining litter and soil profiles (0-20 cm) were sampled and analysed for their total C and ¹³C content. Gas chromatography (GC), GC-mass spectrometry (MS) and GC-combustion-isotope ratio (GC/C/IRMS) were used to analyse phospholipid fatty acids (PLFA) extracted from soil samples to identify the groups of soil micro-organisms that had incorporated litter-derived C and to determine the quantity of C incorporated by the soil microbial biomass (SMB). By the end of the experiment, the litter had lost about 80% of its original weight. The fraction of litter C lost as an input into the soil (67 ± 12% of the total C loss) was found to be twice as much as the fraction released as CO₂ to the atmosphere (30 ± 3%), thus demonstrating the importance of quantifying litter-derived C input to soils, in litter decomposition studies. The mean δ¹³C values of PLFAs in soil (δ¹³C = −12.5‰) showed sustained incorporation of litter-derived C after one year (7.8 ± 1.6% of total PLFA-C). Thus, through the application of stable ¹³C isotope analyses, we have quantified two major C fluxes contributing to litter decomposition, at macroscopic and microscopic levels.     

Isotopic discrimination during long-term decomposition in an arid land ecosystem

Discrimination in carbon and nitrogen isotopes of decomposing plant litter in the northern Chihuahuan Desert was determined for a 5-year period. Factors influencing isotopic change were assessed from inter-species comparisons of litter chemistry, mass loss patterns, and isotope values of associated soil. Average δ¹⁵N_litter values of buried roots increased 1.2 and 2.6‰ for Big Blue Stem (Schizachyrium gerardi, grass) and Varital (Drypetes glauca, hardwood) during the study, respectively. Small but inconsistent variations were observed for Slash Pine (Pinus elliotii, conifer) roots. Average δ¹⁵N values of wooden dowels from Ramin (Gonystlylus bancanus, hardwood) increased ca. 2.0‰ during years 1–4, and then decreased slightly during year 5. Changes in δ¹⁵N_litter were independent of N content, and may reflect microbial fractionation or preferential retention of ¹⁵N enriched substrates. Surprisingly, there was no clear relationship between litter N dynamics and C/N ratios. There were no discernable changes in δ¹³C_litter values for Gonystlylus bancanus and Pinus elliotii. Average δ¹³C_litter values for Schizachyrium gerardi decreased ∼2.0‰ during years 0–2 and then increased slightly. In contrast, average δ¹³C_litter values for Drypetes glauca increased ∼0.5‰ from years 0–1 then remained relatively constant until decreasing slightly in year 5. δ¹³C_litter discrimination may have been masked by interfering δ¹³C fractionations or feedbacks between decomposers and litter chemistry. Our data indicate that isotopic discrimination is characteristic of early litter decay stages. These results may highlight aspects of isotope discrimination and nutrient cycling unique to arid land environments. Additional studies will be needed to confirm this.     

C signature of CO2 evolved from incubated maize residues and maize-derived sheep faeces

Analyses of the spatial and temporal variations in the natural abundance of ¹³C are frequently employed to study transformations of plant residues and soil organic matter turnover on sites where long continued vegetation with the C₃-type photosynthesis pathway has been replaced with a C₄-type vegetation (or vice versa). One controversial issue associated with such analyses is the significance of isotopic fractionation during the microbial turnovers of C in complex substrates. To evaluate this issue, C₃-soil and quartz sand were amended with maize residues and with faeces from sheep feed exclusively on maize silage. The samples were incubated at 15 °C for 117 days (maize residues) or 224 days (sheep faeces). CO₂ evolved during incubation was trapped in NaOH and analysed for C isotopic contents. At the end of incubation, 63 and 50% of the maize C was evolved as CO₂ in the soil and sand, respectively, while 32% of the faeces C incubated with soil and with sand was recovered as CO₂. Maize and faeces showed a similar decomposition pattern but maize decomposed twice as fast as faeces. The δ¹³C of faeces was 0.3‰ lower than that of the maize residue (δ¹³C −13.4‰), while the δ¹³C of the C₃-soil used for incubation was −31.6‰. The δ¹³C value of the CO₂ recovered from unamended C₃-soil was similar or slightly lower (up to −1.5‰) than that of the C₃-soil itself except for an initial flush of ¹³C enriched CO₂. The δ¹³C values of the CO₂ from sand-based incubations typically ranged −15‰ to −17‰, i.e. around −3‰ lower than the δ¹³C measured for maize and faeces. Our study clearly demonstrates that the decomposition of complex substrates is associated with isotopic fractionation, causing evolved CO₂ to be depleted in ¹³C relative to substrates. Consequently the microbial products retained in the soil must be enriched in ¹³C.     

Palatability trials on hardwood leaf litter grown under elevated CO2: a stable carbon isotope study

The palatability to isopods and microbes of a broad range of hardwood leaf litter, derived from three field CO₂-enrichment experiments in the USA, was investigated, using δ¹³C, to trace the C flow from litter to isopods and to CO₂ respired by microbial decomposition. Leaf litter grown under elevated CO₂ had δ¹³C values ranging from −39 to −45‰, which were significantly different from ambient litter δ¹³C values of around −30‰. Litter palatability to isopods of the Porcellio sp. was tested by incubating ambient- and elevated-CO₂ litter, and a mixture of the two, in the presence of isopods for 14 days, under environmentally controlled conditions; δ¹³C was measured on litter and isopods' body before and after incubation. In an additional experiment, litter was incubated in the absence of fauna for 30 days, and on five occasions the δ¹³C of the CO₂ respired from litter was measured. The ¹³C label was clearly carried from the litter source to the isopods' bodies, and their faeces. For microbial-respired CO₂, δ¹³C was significantly higher than that of the litter source, suggesting preferential degradation of substrates enriched in ¹³C as compared to those in the overall litter. With the exception of Quercus myrtifolia leaf litter, elevated CO₂ did not affect the palatability to isopods nor the microbial degradation of any of the litters, possibly as a result of unaltered litter N concentration. However, significant differences in litter palatability and decay rates were observed among the different species. With this study, the use of isotopically labelled litter material was confirmed as a key methodology that can significantly contribute to the advancement of the understanding of litter decomposition and of the quantification of C fluxes in the process.     

Collembolans feeding on soil affect carbon and nitrogen mineralization by their influence on microbial and nematode activities

We manipulated Collembola Folsomia candida Willem density and observed the density effect on carbon and nitrogen mineralization and on nematodes in microcosms filled with mineral soil. Collembolan densities were 0 (control), 25 (low), 100 (medium), and 400 (high) individuals per microcosm. The Collembola enhanced soil respiration and nitrogen mineralization rate in a density-dependent manner (P < 0.05). The correlation between collembolan density and the metabolic quotient of microbes, qCO₂, was weakly positive (r = 0.44, P < 0.05). Collembola did not affect microbial biomass. These results suggested that enhanced carbon and nitrogen mineralization was an indirect effect of Collembola mediated by increased microbial activity. Collembola changed the C_nema/C_mic ratio, but only when present at the low density. Thus, Collembola had both positive and negative effects on the nematode population. The positive impact probably depends on the enhancement of microbial activity due to Collembola grazing behavior, while the negative effect appears to result from predation of nematodes.     

Tracking Collembola feeding strategies by the natural C signal of fatty acids in an arable soil with different fertilizer regimes

Compound specific stable isotope analysis (¹³C/¹²C ratio of fatty acids) was used to assess the allocation of plant carbon in soil microbiota, and to identify the trophic links to microbial grazers in an arable field with long-term mineral and organic fertilizer amendments. The feeding strategy of two dominant Collembola species, epedaphic Isotoma viridis and euedaphic Willemia anophthalma was determined. The investigation was conducted following a shift to amaranth, a C₄ plant, after 27 years of continuous C₃ crop rotation. The influence of new C₄ plant carbon was observed in microbial phospholipids (PLFAs) with higher δ¹³C recorded in C₄ amaranth than in C₃ clover soils. The strongest enrichment occurred in the fungal PLFA 18:2ω6,9c and bacterial PLFA 18:1ω9t with 11.2‰ and 6.6‰, respectively. However, other bacterial PLFAs showed no isotopic change, suggesting that the microbial community simultaneously utilized “new” and “old” plant carbon. The δ¹³C of Collembola fatty acids displayed species specific lipid pattern, which was affected by crop type, but not fertilizer amendments. Isotopic separation of Collembola lipids from amaranth and clover plots was more distinct in I. viridis than W. anophthalma. With up to 18‰, the enrichment in Collembola lipids was stronger than in microbial PLFAs, pointing to a distinct incorporation of carbon resources originating from the actual plant residues. The δ¹³C pattern in I. viridis indicated trophic links with bacteria, saprotrophic fungi and plant tissues, while saprotrophic fungi and plant tissues were accountable for the patterns observed in W. anophthalma.     

Effects of collembolan grazing on nutrient release and respiration of a leaf litter inhabiting fungus

An attempt was made to assess the effect of grazing by Onychiurus subtenuis Folsom on nutrient release from and respiration of a sterile dark fungus using simple microcosms held under controlled laboratory conditions.The results showed that the fungus was very efficient in taking up all available soluble NO₃^? and PO₄^2? and that collembolan grazing had no effects in releasing these nutrients during the 10 day experiment. Significant increase in respiration of litter colonized by the fungus was observed following collembolan grazing, but this increase was attributed to the activity of bacteria and fungi tracked into the leaf litter systems by the Collembola.     

Growth of the collembolan Folsomia candida Willem in soil supplemented with glucose

Freely available glucose improves the conditions for soil microorganisms which are utilized as food by Collembola. We examined the effects of glucose application on collembolan (Folsomia candida Willem) growth and on several biotic and abiotic soil parameters (microbial biomass, soil respiration, qCO₂, dissolved organic carbon, inorganic nitrogen, and Olson-P) in an artificial system without predatory pressure on Collembola. Glucose addition increased soil respiration and qCO₂, and decreased nutrient levels in the soil. Collembolan growth increased with increasing glucose doses. We conclude that the availability of carbon substrates can sustain collembolan growth via an improvement of microbial growth conditions.     

Acquisition of microbial communities and enhanced availability of soil nutrients by the isopod Porcellio scaber (Latr.) (Isopoda: Oniscidea)

 In microcosm experiments Porcellio scaber increased litter disappearance of oak and alder litter. Alder litter disappeared at more than twice the rate of oak litter. Soil texture did not influence the disappearance of oak litter; however, disappearance of alder litter was enhanced on silt rather than on sand. P. scaber enhanced microbial communities (i.e. microbial respiration and microbial biomass) on both silt and sand when feeding on either alder or oak. Overall, microbial respiration increased 10-fold when isopods fed on oak litter on sand and 20-fold when isopods fed on alder litter on sand. On silt, the initially high microbial respiration remained constant when isopods fed on oak and doubled when they fed on alder litter. In all treatments without P. scaber there was a decrease in microbial respiration over the 12-week experimental period. The availability of macronutrients (C_org, N_tot, P, K, Mg, Ca) in the topsoil was increased when P. scaber fed on alder litter but less pronounced when P. scaber fed on oak litter. Using sand as a substrate, there was an apparent increase only for C_org, Mg and Ca; on silt, increases in C_org, N_tot and P were measured. Under field conditions the contribution of P. scaber to nutrient fluxes will be higher on sand than on silt. Received: 1 July 1999     

Microbial use of lignite compared to recent plant litter as substrates in reclaimed coal mine soils

In the Lusatian mining district, rehabilitated mine soils contain substantial amounts of lignite in addition to recent carbon derived from plant litter. The aim of this study was to examine the importance of the two organic matter types as substrates for soil microbial biomass in mine soils containing organic matter with a contrasting degree of humification. Samples were taken from the lignite-containing overburden material, from a mine soil under 14-year-old black pine (Pinus nigra) and from a mine soil under 37-year-old red oak (Quercus rubra). Overburden material was ameliorated with alkaline ash and incubated in an identical manner as the 14-year-old and 37-year-old mine soils for 16 months. Carbon mineralisation was monitored throughout. After 0, 3, 6, 12 and 16 months, samples were removed and analysed for chemical parameters and for microbial biomass. In addition, ¹⁴C activity measurements in bulk soil and microbial biomass were used to estimate their lignite content.Despite the high content of organic carbon in lignite-rich overburden material, low contents of microbial biomass were recorded. Ash-amelioration led to high pH values in the overburden material, resulting in high concentrations of dissolved organic carbon most likely derived from lignite. Development of the microbial community was subsequently stimulated by presence of an easily available carbon source. In older mine soils, larger amounts of microbial biomass are most likely related to the presence of recent organic matter. Radiocarbon analysis of the microbial biomass extracted from the 14-year-old mine soil indicated higher lignite carbon contribution than recorded for microbial biomass of the 37-year-old mine soil. The highest concentration of lignite C present in microbial biomass as indicated by the C_mic/C_org ratio was, however, observed in the ameliorated overburden material. Therefore, we conclude that the importance of lignite as a carbon source for micro-organisms decreases when recent organic matter is present in the older stages of mine soil development.