Trophic structure of oribatid mite communities from six different oak forests (Quercus robur)

The degree of trophic plasticity in soil animals is intensely debated. We used stable isotope ratios (¹⁵N/¹⁴N, ¹³C/¹²C) of oribatid mite species from six oak (Quercus robur) forests to investigate (1) if trophic niches vary between forests and (2) the range of trophic levels spanned by oribatid mites. Using litter as baseline stable isotope signatures of most oribatid mite species differed between forests. Therefore, the stable isotope signatures were re-calibrated using stable isotope values of Platynothrus peltifer as primary decomposer species occurring in each of the six forests. Re-calibrated values of nine species (Cerachipteria jugata, Damaeus clavipes, Neotrichoppia variabilis, Oppia denticulata, Hermaniella dolosa, Steganacarus magnus, Ceratozetes peritus, Nanhermannia nana, Xenillus tegeocranus, Eremaeus cordiformis) differed significantly between forests indicating trophic plasticity in most of the studied oribatid mite species. Overall, calibrated stable isotope ratios spanned over 8.7 δ units for ¹⁵N and 5.9 δ units for ¹³C indicating that in forest ecosystems oribatid mite species span about three trophic levels.     

Trophic niche differentiation in soil microarthropods (Oribatida, Acari): evidence from stable isotope ratios (N/N)

The large number of animals that coexist in soil without any clear niche differentiation has puzzled biologists for a long time. We investigated stable isotope ratios (¹⁵N/¹⁴N) in a diverse group of soil microarthropods, oribatid mites, to evaluate trophic niche differentiation. The natural variation of the stable isotopes ¹⁵N/¹⁴N was measured in 36 species/taxa from four beech and beech-oak forests. Signatures of δ¹⁵N formed a gradient spanning over 12 δ units suggesting that (a) different species occupy different trophic niches and (b) oribatid mites span three to four trophic levels. This study for the first time documented strong trophic niche differentiation in decomposer microarthropods. The results suggest that trophic niche differentiation within taxonomic groups significantly contributes to the high diversity of soil animal taxa.     

Compartmentalization of the soil animal food web as indicated by dual analysis of stable isotope ratios (N/N and C/C)

The soil animal food web has become a focus of recent ecological research but trophic relationships still remain enigmatic for many taxa. Analysis of stable isotope ratios of N and C provides a powerful tool for disentangling food web structure. In this study, animals, roots, soil and litter material from a temperate deciduous forest were analysed. The combined measurement of δ¹⁵N and δ¹³C provided insights into the compartmentalization of the soil animal food web. Leaf litter feeders were separated from animals relying mainly on recent belowground carbon resources and from animals feeding on older carbon. The trophic pathway of leaf litter-feeding species appears to be a dead end, presumably because leaf litter feeders (mainly diplopods and oribatid mites) are unavailable to predators due to large size and/or strong sclerotization. Endogeic earthworms that rely on older carbon also appear to exist in predator-free space. The data suggest that the largest trophic compartment constitutes of ectomycorrhizal feeders and their predators. Additionally, there is a smaller trophic compartment consisting of predators likely feeding on enchytraeids and potentially nematodes.     

Trophic stability of soil oribatid mites in the face of environmental change

A key issue in ecology is the degree to which trophic structure within communities responds to environmental change. Organisms with generalist diets are more flexible in their feeding habits than are specialists, and may be affected less in a changing environment. Soil fauna fulfill crucial ecosystem functions in terrestrial ecosystems and many are thought to have generalized diets. They may therefore be buffered from negative effects of environmental change. Here, we used ¹⁵N isotope analysis to study trophic differentiation among 91 species of oribatid mites and their responses to chronic atmospheric N deposition. Combining our own measurements with published data, we established that the trophic positions of mite species were remarkably stable within and among forests, as well as between ambient and experimental N deposition. Trophic stability indicates a higher than expected level of feeding specialization, which may foster diversity, but limit the ability to switch food resources in a changing environment.     

Changes in the community composition and trophic structure of microarthropods in sporocarps of the wood decaying fungus Fomitopsis pinicola along an altitudinal gradient

Soil microarthropods colonize a wide range of habitats including microhabitats such as earthworm burrows, ant nests, tree trunks, moss mats and wood decaying fungi. While many of these microhabitats have been investigated intensively, the role of wood decaying fungi as a habitat and food resource for microarthropods found little attention. We investigated the density, community structure, reproductive mode and trophic structure of microarthropods, in particular oribatid mites, in the wood decaying fungus Fomitopsis pinicola (Schwarts: Fr) Karst. along an altitudinal gradient in Germany spanning from 350 m to 1160 m. Microarthropods were extracted from sporocarps, and stable isotope ratios (¹⁵N/¹⁴N; ¹³C/¹²C) of the fungus and the microarthropods were measured. Densities of most microarthropod taxa were highest at lower altitudes and decreased with increasing altitude. Oribatid mites were the dominant animal taxon. Their community structure gradually changed with altitude. Stable isotope ratios indicated that oribatid mite and other arthropod species occupy distinct trophic niches but most do not feed on F. pinicola. Notably, species of the same genus, e.g. Carabodes, occupied different trophic niches. Most oribatid mite species in F. pinicola reproduced sexually which is similar to the bark of trees but in contrast to the soil where most species reproduce via parthenogenesis. The findings indicate that (1) at high altitudes microarthropod density in fungal fruiting bodies is limited by low temperatures reducing animal metabolism and reproduction, and this also affects oribatid mite community structure, (2) despite the uniform habitat trophic niches of oribatid mite species differ and this also applies to morphologically similar species of the same genus, and (3) feeding on F. pinicola or associated resources facilitates the dominance of sexual reproducing species.     

Effects of food quality, starvation and life stage on stable isotope fractionation in Collembola

Naturally occurring stable isotopes of carbon and nitrogen are powerful tools to investigate food webs, where the ratio of ¹⁵N/¹⁴N is used to assign trophic levels and of ¹³C/¹²C to determine the food source. A shift in δ¹⁵N value of 3‰ is generally suggested as mean difference between two trophic levels, whereas the carbon isotope composition of a consumer is assumed to reflect the signal of its diet. This study investigates the effects of food quality, starvation and life stage on the stable isotope fractionation in fungal feeding Collembola. The fractionation of nitrogen was strongly affected by food quality, i.e. the C/N ratio of the fungal diet. Collembola showed enrichment in the heavier isotope with increasing N concentration of the food source. Δ¹⁵N varied between 2.4‰, which assigns a shift in one trophic level, and 6.3‰, suggesting a shift in two trophic levels. Starvation up to 4 weeks resulted in an increase in the total δ¹⁵N value from 2.8‰ to 4.0‰. Different life stages significantly affected the isotope discrimination by Collembola with juveniles showing a stronger enrichment (Δ¹⁵N=4.9‰) compared to adults (Δ¹⁵N=3.5‰). Δ¹³C varied between −2.1‰ and −3.3‰ depending on the food quality, mainly due to compensational feeding on low quality diet. During starvation δ¹³C value decreased by 1.1‰, whereas the life stage of Collembola had no significant effect on isotopic ratios. The results indicate that the food resource and the physiological status of the consumer have important impact on stable isotope discrimination. They may cause differences in fractionation rate comparable to trophic level shifts, a fact to consider when analysing food web structure.     

Feeding guilds in Collembola based on nitrogen stable isotope ratios

In soil a high number of species co-exist without extensive niche differentiation, which was assigned as ‘the enigma of soil animal species diversity’. In particular, the detritivores are regarded as food generalists. We have investigated nitrogen stable isotope ratios (¹⁵N/¹⁴N) of a major decomposer group, the Collembola, to evaluate trophic relationship and determine feeding guilds. Additionally, the δ¹⁵N values of potential food sources such as mosses, lichens and other plant derived material (bark, nuts, leaves) were analysed. The natural variation in nitrogen isotopes was assessed in 20 Collembola taxa from three deciduous forest stands. The δ¹⁵N signature formed a continuum from phycophages/herbivores to primary and secondary decomposers, reflecting a gradual shift from more detrital to more microbial diets. The δ¹⁵N gradient spanned over 9 δ units, which implies a wide range in food sources used. Assuming a shift in ¹⁵N of about 3 ‰ per trophic level, the results indicate a range of three trophic levels. These variations in ¹⁵N/¹⁴N ratios suggest that trophic niches of Collembola species differ and this likely contributes to Collembola species diversity.     

Evaluating N/N and C/C isotope ratio analysis to investigate trophic relationships of elaterid larvae (Coleoptera: Elateridae)

Carbon and nitrogen isotope ratios in consumer tissues can be used to analyse the diet and trophic level of soil animals. However, life history traits may significantly influence stable isotope patterns. We evaluated in a series of experiments how stable isotope ratios of carbon (¹³C/¹²C) and nitrogen (¹⁵N/¹⁴N) at natural abundance can be used to study the diet and trophic position of long-lived macro-invertebrates, elaterid larvae, which are major below-ground herbivores. Small, but significant differences in δ¹³C signatures were found between the larvaes’ anterior and posterior body segments, whereas exuvia reflected the body's overall isotopic composition. The species-specific trophic shift (±SE) in δ¹⁵N for Agriotes obscurus and Agriotes sputator (1.62±0.24‰ and 1.08±0.27‰, respectively) was significantly lower than “mean enrichment estimates” reported in the literature, showing the limited applicability of such generalised estimates in studies of invertebrate trophic ecology. To avoid false-positive assignments to two trophic levels due to variation in δ¹⁵N values, a minimum sample size of three and five individuals for A. obscurus and A. sputator, respectively, was needed to reduce this risk to below α=5%. Keeping elaterid larvae for up to 128 days without food did not affect their isotopic signatures, in contrast to previous studies on starving animals. Switching wireworms to isotopically different diets induced changes in their isotopic signatures within 2 weeks. Changes, however, were significant only when the isotopic difference between diets was large. We conclude that experimental studies evaluating how specific life history traits affect stable isotope signatures in consumers have to precede any interpretation of stable isotope data gathered in the field.     

Dual stable isotope analysis (δC and δN) of soil invertebrates and their food sources

More research is required to validate and refine natural abundance stable isotope ratio techniques as a tool for the investigation of the feeding ecology of soil animals and trophic relations in soil food webs. Isotope ratios of C (δ¹³C) and N (δ¹⁵N) were measured in herbivorous and detritivorous invertebrate groups, namely lumbricid earthworms (7 species), enchytraeid worms (3 species), slugs (3 taxa), and their potential food sources in an arable system. Intrapopulation δ¹⁵N variation in the slug Deroceras reticulatum (n=52) was large (range 4.2‰), possibly reflecting spatial variability in the food sources. Significant correlations between C:N ratios and isotope ratios in earthworms suggest that factors other than feeding may influence isotopic patterns. One enchytraeid species, Enchytraeus buchholzi, was enriched in ¹³C and strongly depleted in ¹⁵N compared to all other groups. Invertebrates formed a continuum when considered in relation to C and N separately, but fell into two distinct groups on the basis of combined C and N isotope ratios. The less enriched group represents herbivorous and litter-feeding species, while the more enriched group represents soil feeders. It is concluded that δ¹³C measurements could provide a means of assigning separate baseline δ¹⁵N values to primary and secondary decomposers, which in turn could improve the inference of higher trophic levels, omnivory and intraguild predation.     

Trophic fractionation (ΔN) in Collembola depends on nutritional status: A laboratory experiment and mini-review

Isotopic studies in temperate forest soils suggested that collembolans occupy a broad range of trophic niches. This notion is mainly based on the wide (up to 8-9‰) range of δ¹⁵N signatures of different species. The interpretation of these data depends strongly on the correct estimation of nitrogen isotope fractionation per trophic level. It is usually assumed that a consumer is on average enriched in ¹⁵N by about 3.4‰ relative to its diet. However, trophic fractionation is not uniform across different animal taxa and different tissues, and varies with a range of factors, including quality of the diet and the nutritional status of animals. We performed a laboratory experiment aiming to compare the trophic fractionation of nitrogen isotopes (Δ¹⁵N) in different Collembola species fed with three fungi of different palatability. Collembolans were fed with a single-species fungal diet for 75 days. There was a positive correlation between the whole body C:N ratio and reproduction rates of collembolans. We therefore used the C:N ratio as a proxy of nutritional status. In all species of collembolans, the trophic fractionation decreased with increased whole body C:N ratio, and this factor explained most of the within-species variation in Δ¹⁵N. The analysis of published data on the trophic fractionation of collembolans in laboratory experiments confirmed this conclusion. However, the mean trophic fractionation in most collembolan species studied in the laboratory was fairly similar. In contrast, field studies have documented a consistent difference in δ¹⁵N among different families of collembolans. In particular, in nearly all published cases Onychiuridae had δ¹⁵N higher than Isotomidae or Entomobryidae. A sharp contrast between laboratory-based estimates of the trophic fractionation (generally uniform in different species and families) and field data (which show a consistent difference in δ¹⁵N among different species and even families of collembolans) confirms that δ¹⁵N values of field-collected animals bear important information on the trophic position occupied by a particular species, and are not heavily affected by the species-specific differences in trophic fractionation. As the Δ¹⁵N in collembolans depends on their nutritional status, we suggest that the C:N values should be reported along with isotopic data to allow cross-study comparisons.     

Molecular detection of nematode predation and scavenging in oribatid mites: Laboratory and field experiments

Recent stable isotope analyses indicate that a number of putative detritivorous soil microarthropods is not typical detritivores but rather live as predators or scavengers. Using molecular gut content analyses the present study investigates if nematodes indeed form part of the diet of oribatid mites. First, in a no-choice laboratory feeding experiment two nematode species (Phasmarhabditis hermaphrodita and Steinernema feltiae) were offered to eight species of oribatid mites and one gamasid mite. Second, after feeding for 4 and 48 h on each nematode species the detection time of prey DNA in the oribatid mite species Steganacarus magnus was investigated. Third, in a field experiment nematode prey (P. hermaphrodita and S. feltiae) in the diet of microarthropods was investigated distinguishing between scavenging and predation. In the no-choice laboratory experiment not only the gamasid mite but also several of the studied oribatid mite species consumed nematodes. After feeding on nematodes for 4 h prey DNA was detectable in S. magnus for only 4 h, but after feeding for 48 h prey DNA was detectable for 128 h, indicating that the duration of feeding on prey is an important determinant for prey DNA detection. The field experiment confirmed that oribatid mite species including Liacarus subterraneus, Platynothrus peltifer and S. magnus intensively prey on nematodes. Interestingly, DNA of dead P. hermaphrodita was detectable to a similar degree as that of living individuals indicating that scavenging is of significant importance in decomposer food webs. Results of our study indicate that predation and scavenging on nematodes by “detritivorous” microarthropods in soil food webs need to be reconsidered.     

Dissipation of bacterially derived C and N through the meso- and macrofauna of a grassland soil

Feeding relationships between organisms may be determined by observations of behaviour in manipulative experiments or, as in more recent times, by the use of stable isotope labelling to trace the passage of ¹³C and ¹⁵N through food webs. Here we introduce living bacteria, labelled with both ¹³C and ¹⁵N into intact soil cores to understand further the movement of bacterially sourced C and N into the meso- and macrofauna of a grassland soil. We found that these groups showed a range of isotope levels which relate to their feeding strategies. Some had no label (e.g. dipterous larvae), whilst others were highly labelled which may indicate a preference for the added bacteria. This latter group included Collembola, generally perceived as being predominantly fungal feeders. This work describes a novel technique which has the potential to provide critical information about the dissipation of bacterially derived C and N through the soil food web.     

A GC/MS method for the assessment of N and C incorporation into soil amino acid enantiomers

Identifying the transformation process of amino acid enantiomers was essential to probe into the fate, turnover and aging of soil nitrogen due to their important roles in the biogeochemical cycling. If this can be achieved by differentiating between the newly biosynthesized and the inherent compounds in soil, then the isotope tracer method can be considered most valid. We thereby developed a gas chromatography/mass spectrometry (GC/MS) method to trace the ¹⁵N or ¹³C isotope incorporation into soil amino acid enantiomers after being incubated with ¹⁵NH₄⁺ or U-¹³C-glucose substrates. The most significant fragments (F) as well as the related minor ions were monitored by the full scan mode and the isotope enrichment in amino acids was estimated by calculating the atom percentage excess (APE). ¹⁵NH₄⁺ incorporation was evaluated according to the relative abundance increase of m/z F+1 to F for neutral and acidic amino acids and F+2 to F (mass 439) for lysine. The assessment of ¹³C enrichment in soil amino acids was more complicated than that of ¹⁵N due to multi-carbon atoms in amino acid molecules. The abundance ratio increment of m/z F+n to F (n is the original skeleton carbon number in each fragment) indicated the direct conversion from the added glucose to amino acids, but the total isotope incorporation from the added ¹³C can only be calculated according to all target isotope fragments, i.e. the abundance ratio increment summation from m/z (Fa+1) through m/z (Fa+T) represented the total incorporation of the added ¹³C (Fa is the fragment containing all original skeleton carbons and T is the carbon number in the amino acid molecule). This method has a great advantage especially for the evaluation of high-abundance isotope enrichment in organic compounds compared with GC/C/IRMS. And in principle, this technique is also valid for amino acids besides enantiomers if stereoisomers are not concerned. Our assessment approach could shine a light on investigating the biochemical mechanism of microbial transformation of N and C in soils of terrestrial ecosystem.     

The feeding ecology of elaterid larvae in central European arable land: New perspectives based on naturally occurring stable isotopes

To understand soil food webs, empirically generated data on the trophic connections and the feeding ecology of the major below-ground animal taxa are needed. Here we used stable isotope analysis to assess the trophic ecology of wireworms, the larvae of click beetles, in Central European arable land. Wireworms are amongst the major soil macroinvertebrates and are of practical importance in arable soils. Besides feeding on crops, they are thought to feed on weeds, soil organic matter (SOM), and even animal prey, but their feeding ecology is poorly studied under natural conditions. Elaterid larvae and their putative feeding substrates—plant roots, SOM, and litter—were sampled at 17 locations in Austria, Germany, and Italy and their isotope ratios of carbon (¹²C/¹³C) and nitrogen (¹⁴N/¹⁵N) measured to determine the wireworms’ trophic level, the importance of SOM and weeds within the diet of Agriotes larvae, as well as the individual diet variation in Agriotes obscurus larvae. δ¹⁵N signatures suggested that Agriotes larvae are predominately herbivorous, whereas the other wireworm species primarily fed on animal prey. In contrast to SOM, weeds were readily eaten by Agriotes larvae: their dietary contribution ranged between 28% and 67% in weedy maize fields. Most A. obscurus larvae fed on a mixed diet of weeds and maize, although ～15% of the larvae fed primarily on one of the two food sources only. δ¹⁵N signatures indicated that ～10% of the “herbivorous” A. obscurus larvae fed primarily on animal prey, revealing high intraspecific trophic plasticity in these soil insects. Wireworm feeding behaviour is apparently complex at the individual level: the population consists of types A and B generalists, a phenomenon which needs further assessment.     

Savanna-derived organic matter remaining in arable soils of the South African Highveld long-term mixed cropping: Evidence from C and N natural abundance

Sustainable agriculture requires the formation of new humus from the crops. We utilized ¹³C and ¹⁵N signatures of soil organic matter to assess how rapidly wheat/maize cropping contributed to the humus formation in coarse-textured savanna soils of the South African Highveld. Composite samples were taken from the top 20 cm of soils (Plinthustalfs) cropped for lengths of time varying from 0 to 98 years, after conversion from native grassland savanna (C4). We performed natural ¹³C and ¹⁵N abundance measurements on bulk and particle-size fractions. The bulk soil δ¹³C values steadily decreased from −14.6 in (C4 dominated) grassland to −16.5‰ after 90 years of arable cropping. This δ¹³C shift was attributable to increasing replacement of savanna-derived C by wheat crop (C3) C which dominated over maize (C4) inputs. After calculating the annual C input from the crop yields and the output from literature data, by using a stepwise C replacement model, we were able to correct the soil δ¹³C data for the irregular maize inputs for a period of about one century. Within 90 years of cropping 41-89% of the remaining soil organic matter was crop-derived in the three studied agroecosystems. The surface soil C stocks after 90 years of the wheat/maize crop rotation could accurately be described with the Rothamsted Carbon Model, but modelled C inputs to the soil were very low. The coarse sand fraction reflected temporal fluctuations in ¹³C of the last C₃ or C₄ cropping and the silt fraction evidenced selective erosion loss of old savanna-derived C. Bulk soil ¹⁵N did not change with increasing cropping length. Decreasing δ¹⁵N values caused by fertilizer N inputs with prolonged arable cropping were only detected for the coarse sand fraction. This indicated that the present N fertilization was not retained in stable soil C pool. Clearly, conventional cropping practices on the South African highlands neither contribute to the preservation of old savanna C and N, nor the effective humus reformation by the crops.     

Tree identity surpasses tree diversity in affecting the community structure of oribatid mites (Oribatida) of deciduous temperate forests

The role of tree diversity and identity as determinants of soil animal community structure is little understood. In a mature deciduous forest dominated by beech we identified clusters of one, two and three tree species of beech, ash and lime allowing to investigate the role of tree species diversity and identity on the density and community structure of oribatid mites. To relate oribatid mite community structure to environmental factors we measured leaf litter input, fine root biomass, mass of organic layers, topsoil pH and C and N content. We expected oribatid mite density to increase with increasing tree diversity, but we expected the effects of tree species identity to override effects of tree diversity. Further, we hypothesized the density of oribatid mites to be reduced by the presence of beech but increased by the presence of lime and ash. As expected tree diversity little affected oribatid mite communities, whereas tree species identity strongly altered density and community structure of oribatid mites. However, in contrast to our expectations the density of oribatid mites was highest in presence of beech indicating that many oribatid mite species benefit from the presence of recalcitrant litter forming thick organic layers. Especially Oppioidea benefited from the presence of beech presumably due to an increased availability of food resources such as fungi and nematodes. Lower density of oribatid mites in monospecific clusters of lime and ash suggests that oribatid mites did not benefit from high quality litter of these species. Notably, large and strongly sclerotized oribatid mite species, such as Steganacarus magnus and Chamobates voigtsi, benefited from the presence of ash and lime. Presumably, these large species better resist harsh microclimatic conditions in shallow organic layers.     

Do oribatid mites live in enemy-free space? Evidence from feeding experiments with the predatory mite Pergamasus septentrionalis

To examine whether their strongly hardened cuticle permits adult oribatid mites (Acari) to live in enemy-free space, we investigated (1) if Pergamasus septentrionalis, a widespread and abundant predatory mesostigmate mite species, is able to feed on oribatid mites, (2) if this predator preferentially feeds on certain oribatid mite species and (3) to what extent oribatid mites are consumed compared to collembolans and juvenile Mesostigmata. Single adult individuals of six different oribatid mite species (Steganacarus magnus; Nothrus silvestris; Damaeus riparius; Liacarus coracinus; Eupelops plicatus; Achipteria coleoptrata), one collembolan species (Folsomia quadrioculata) and juvenile Pergamasus spp. were offered separately to adult P. septentrionalis in a no-choice feeding experiment. The predators quickly and preferentially fed on collembolans and juvenile Pergamasus; three oribatid mite species were occasionally eaten (L. coracinus, N. silvestris, A. coleoptrata); the other oribatid mite species were rejected as food (E. plicatus, S. magnus, H. riparius). When preying on oribatid mites, P. septentrionalis typically first cut off the legs of the mite, then opened the body through the region of the genital plates or the mouthparts. The results suggest that predator pressure on adult oribatid mites in the field is low, since few relevant predators at the study site are more abundant and powerful than P. septentrionalis. Adult oribatid mites therefore likely indeed live in enemy-free space, i.e. are little affected by predators, but that may not apply to soft-bodied immatures. Collembolans were quickly consumed indicating that they comprise a major part of the diet of P. septentrionalis. Strong feeding on juveniles of Pergamasus suggests that P. septentrionalis also functions as an intra-guild predator.     

施氮量对嘎啦幼苗~(15)N、~(13)C分配利用特性影响

【目的】采用15N、13C同位素示踪技术,通过对不同施氮量下嘎啦幼苗生长状况及氮、碳分配、利用特性等的研究,以期为苹果生产合理施肥提供依据。【方法】将2年生盆栽嘎啦幼苗进行低、中、高三个氮水平处理,同时进行15N标记。在新梢旺长初始期、新梢旺长期、新梢缓长期分别进行整株13C标记,72小时后,整株解析为叶、梢、根三部分,进行15N、13C测定。样品全氮用凯氏定氮法测定,15N丰度用ZHT-03质谱计测定。13C丰度用DELTA V Advantage同位素比率质谱仪测定。【结果】1)中、高氮水平的施肥处理可在不同程度上提高整株及叶片干物质量和新梢长度。新梢旺长初始期和新梢缓长期嘎啦幼苗整株干物质量、新梢旺长期叶片干物质分配比率在中、高氮水平处理间差异不显著,中氮水平经济有效。新梢旺长期以后新梢长度以中氮高氮低氮,三者间差异性显著,中氮处理有利于新梢生长。2)在新梢旺长初始期,低氮处理植株叶片15N分配率达50%,比其他处理高出13个百分点左右,表明低氮处理更多的氮被叶片所利用,中氮和高氮处理间差异不显著,说明在本试验施氮条件下中氮供应水平已能满足氮素营养需求。3)新梢旺长期和新梢缓长期幼苗13C固定量均以中氮处理最高,新梢旺长初始期3个处理间根系13C分配率中氮高氮低氮,表明中氮处理有利于碳同化物在嘎啦幼苗中的分配。4)不同施氮量处理的嘎啦幼苗,15N利用率随施氮水平提高而降低,高氮处理对碳同化物分配没有显著贡献。【结论】低、中、高氮不同处理新梢缓长期碳同化物在各器官间的分配比较均衡,氮素水平不能影响碳同化物的分配。盆栽试验表明,中氮水平在保证营养供应的同时,能够促进新梢生长和树势健壮。     

C and N natural abundance of the soil microbial biomass

Stable isotope analysis is a powerful tool in the study of soil organic matter formation. It is often observed that more decomposed soil organic matter is ¹³C, and especially ¹⁵N-enriched relative to fresh litter and recent organic matter. We investigated whether this shift in isotope composition relates to the isotope composition of the microbial biomass, an important source for soil organic matter. We developed a new approach to determine the natural abundance C and N isotope composition of the microbial biomass across a broad range of soil types, vegetation, and climates. We found consistently that the soil microbial biomass was ¹⁵N-enriched relative to the total (3.2 ‰) and extractable N pools (3.7 ‰), and ¹³C-enriched relative to the extractable C pool (2.5 ‰). The microbial biomass was also ¹³C-enriched relative to total C for soils that exhibited a C3-plant signature (1.6 ‰), but ¹³C-depleted for soils with a C4 signature (−1.1 ‰). The latter was probably associated with an increase of annual C3 forbs in C4 grasslands after an extreme drought. These findings are in agreement with the proposed contribution of microbial products to the stabilized soil organic matter and may help explain the shift in isotope composition during soil organic matter formation.     

Roots from beech (Fagus sylvatica L.) and ash (Fraxinus excelsior L.) differentially affect soil microorganisms and carbon dynamics

Knowledge about the influence of living roots on decomposition processes in soil is scarce but is needed to understand carbon dynamics in soil. We investigated the effect of dominant deciduous tree species of the Central European forest vegetation, European beech (Fagus sylvatica L.) and European ash (Fraxinus excelsior L.), on soil biota and carbon dynamics differentiating between root- and leaf litter-mediated effects. The influence of beech and ash seedlings on carbon and nitrogen flow was investigated using leaf litter enriched in ¹³C and ¹⁵N in double split-root rhizotrons planted with beech and ash seedlings as well as a mixture of both tree species and a control without plants. Stable isotope and compound-specific fatty acid analysis (¹³C-PLFA) were used to follow the incorporation of stable isotopes into microorganisms, soil animals and plants. Further, the bacterial community composition was analyzed using pyrosequencing of 16S rRNA gene amplicons. Although beech root biomass was significantly lower than that of ash only beech significantly decreased soil carbon and nitrogen concentrations after 475 days of incubation. In addition, beech significantly decreased microbial carbon use efficiency as indicated by higher specific respiration. Low soil pH probably increased specific respiration of bacteria suggesting that rhizodeposits of beech roots induced increased microbial respiration and therefore carbon loss from soil. Compared to beech δ¹³C and δ¹⁵N signatures of gamasid mites in ash rhizotrons were significantly higher indicating higher amounts of litter-derived carbon and nitrogen to reach higher trophic levels. Similar δ¹³C signatures of bacteria and fine roots indicate that mainly bacteria incorporated root-derived carbon in beech rhizotrons. The results suggest that beech and ash differentially impact soil processes with beech more strongly affecting the belowground system via root exudates and associated changes in rhizosphere microorganisms and carbon dynamics than ash.