Precipitation modifies the effects of warming and nitrogen addition on soil microbial communities in northern Chinese grasslands

Terrestrial ecosystems experience simultaneous shifts in multiple drivers of global change, which can interactively affect various resources. The concept that different resources co-limit plant productivity has been well studied. However, co-limitation of soil microbial communities by multiple resources has not been as thoroughly investigated. Specifically, it is not clearly understood how microbial communities respond to shifts in multiple interacting resources such as water, temperature, and nitrogen (N), in the context of global change. To test the effects of these various resources on soil microorganisms, we established a field experiment with temperature and N manipulation in three grasslands of northern China, where there is a decrease in precipitation from east to west across the region. We found that microbial responses to temperature depended upon seasonal water regimes in these temperate steppes. When there was sufficient water present, warming had positive effects on soil microorganisms, suggesting an interaction between water and increases in temperature enhanced local microbial communities. When drought or alternating wet–dry stress occurred, warming had detrimental effects on soil microbial communities. Our results also provide clear evidence for serial co-limitation of microorganisms by water and N at the functional group and community levels, where water is a primary limiting factor and N addition positively affects soil microorganisms only when water is sufficient. We predict that future microbial responses to changes in temperature and N availability could be seasonal or exist only in non-drought years, and will strongly rely on future precipitation regimes.     

Plant species richness does not attenuate responses of soil microbial and nematode communities to a flood event

Human activities are causing climatic changes and alter the composition and biodiversity of ecosystems. Climate change has been and will be increasing the frequency and severity of extreme climate events and natural disasters like floods in many ecosystems. Therefore, it is important to investigate the effects of disturbances on ecosystems and identify potential stabilizing features of ecological communities. In this study, soil microbial and nematode communities were investigated in a grassland biodiversity experiment after a natural flood to investigate if plant diversity is able to attenuate or reinforce the magnitude of effects of the disturbance on soil food webs. In addition to community analyses of soil microorganisms and nematodes, the stability indices proportional resilience, proportional recovery, and proportional resistance were calculated. Generally, soil microbial biomass decreased significantly due to the flood with the strongest reduction in gram-negative bacteria, while gram-positive bacteria were less affected by flooding. Fungal biomass increased significantly three months after the flood compared to few days before the flood, reflecting elevated availability of dead plant biomass in response to the flood. Similar to the soil microbial community, nematode community structure changed considerably due to the flood by favoring colonizers (in the broadest sense r-strategists; c–p 1, 2 nematodes), particularly so at high plant diversity. None of the soil microbial community stability indices and few of the nematode stability indices were significantly affected by plant diversity, indicating limited potential of plant diversity to buffer soil food webs against flooding disturbance. However, plant diversity destabilized colonizer populations, while persister populations (in the broadest sense K-strategists; c–p 4 nematodes) were stabilized, suggesting that plant diversity can stabilize and destabilize populations depending on the ecology of the focal taxa. The present study shows that changes in plant diversity and subsequent alterations in resource availability may significantly modify the compositional shifts of soil food webs in response to disturbances.     

Early-successional vegetation changes after roadside prairie restoration modify processes related with soil functioning by changing microbial functional diversity

Because of their rapidly changing vegetation dynamics and harsh environmental conditions, roadside prairies in semi-arid regions represent an exceptional study system in which to investigate the effects of plant-soil interactions on ecosystem functioning. We conducted a two-year field experiment on two roadside embankments in semi-arid central Spain differing in construction age to answer the following questions: (i) do commonly used restoration treatments (hydroseeding, fertilization and irrigation) affect soil microbial functional diversity and processes related to soil functioning (basal respiration, total N and P and in situ N availability rate)? (ii) what portion of plant effects on processes related to soil functioning is mediated indirectly by microbial functional diversity? Except for a small and negative irrigation effect on the microbial functional diversity in the three-year old site, the restoration treatments employed did not affect this variable. Fertilization increased plant diversity, an effect likely mediated by the enhanced soil nutrient availability with this treatment at early stages of secondary succession. In contrast, hydroseeding did not affect processes related to soil functioning. The total effect of the plant community on these processes was higher than that of the microbial functional diversity alone, suggesting that the studied slopes are to the greater extent regulated by plants. However, soil microbes are a key proximate influence in the system, as the indirect effects of plant community on soil functioning processes mediated by soil microbes represented 37-41% of the total plant effects observed. Our results indicate that the restoration of recently built slopes can potentially be improved with treatments that promote plant compositional shifts, such as fertilization, or alter soil function, such as the enhancement of soil microbial functional diversity. They also highlight that plant-soil interactions are an important process that can be manipulated for restoration purposes in early-successional stages, especially in nutrient-poor semi-arid ecosystems.     

Enzyme activities as a component of soil biodiversity: A review

Soil enzyme activities are the direct expression of the soil community to metabolic requirements and available nutrients. While the diversity of soil organisms is important, the capacity of soil microbial communities to maintain functional diversity of those critical soil processes through disturbance, stress or succession could ultimately be more important to ecosystem productivity and stability than taxonomic diversity. This review examines selected papers containing soil enzyme data that could be used to distinguish enzyme sources and substrate specificity, at scales within and between major nutrient cycles. Developing approaches to assess soil enzyme functional diversity will increase our understanding of the linkages between resource availability, microbial community structure and function, and ecosystem processes.     

Resource dynamics in an early-successional plant community are influenced by insect exclusion

The exclusion of insects from terrestrial ecosystems may change productivity, diversity and composition of plant communities and thereby nutrient dynamics. In an early-successional plant community we reduced densities of above- and below-ground insects in a factorial design using insecticides. Beside measuring vegetation dynamics we investigated the effects of insect exclusion on above- and below-ground plant biomass, below-ground C and N storage by plants, litter quality, decomposition rate, soil water content, soil C:N ratio, nutrient availability and soil microbial activity and biomass.The application of soil insecticide had only minor effects on above- and below-ground biomass of the plant community but increased carbon content in root biomass and total carbon and nitrogen storage in roots. In one of the three investigated plant species (Cirsium arvense), application of soil insecticide decreased nitrogen concentration of leaves (−12%). Since C. arvense responded positively to soil insecticide application, this effect may be due to drought stress caused by root herbivory. Decomposition rate was slightly increased by the application of above-ground insecticide, possibly due to an impact on epigeic predators. The application of soil insecticide caused a slightly increased availability of soil water and an increased availability of mineralised nitrogen (+30%) in the second season. We explain these effects by phenological differences between the plant communities, which developed on the experimental plots. Microbial biomass and activity were not influenced by insecticide application, but were correlated to above-ground plant biomass of the previous year. Overall, we conclude that the particular traits of the involved plant species, e.g. their phenology, are the key to understand the resource dynamics in the soil.     

Changes of the functional diversity of soil microbial community during the colonization of abandoned grassland by a forest

The impact of secondary succession of grassland communities towards a Norway spruce forest on soil microbial community was studied on a belt transect established in the Pol’ana Mts., Central Europe. Data on understory vegetation, light availability, soil properties and microbial activity were collected on 147 plots distributed over regular grid. Moreover, distributions of functional groups of microorganisms were assessed using BIOLOG analysis on a subset of 27 plots. Mantel partial correlations between microbial community indicators and environmental variables showed that microbial activity generally decreased with increasing tree density and size, whereas it increased with increasing radiation at the soil surface, soil temperature, and cover and diversity of understory vegetation. Functional richness and diversity of microorganisms were positively correlated with solar radiation, but also with plant species richness and diversity. Abundance of several functional groups correlated closely with succession-related variables. Redundance analysis of microbial data provided slightly different outcomes. Forward selection yielded only two environmental variables significantly influencing the composition of the microbial community: tree influence potential and organic carbon content. Abundances of several functional microbial groups correlated with tree influence, documenting that microbial community changes are at least partially driven by the colonization of grassland by trees. Nevertheless, the relative importance of abiotic environment change and plant community succession on microbial community dynamics remains unresolved.     

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.     

Land-use intensification and agroforestry in the Kenyan highland: Impacts on soil microbial community composition and functional capacity

This study investigates microbial communities in soil from sites under different land use in Kenya. We sampled natural forest, forest plantations, agricultural fields of agroforestry farms, agricultural fields with traditional farming and eroded soil on the slopes of Mount Elgon, Kenya. We hypothesised that microbial decomposition capacity, biomass and diversity (1) decreases with intensified cultivation; and (2) can be restored by soil and land management in agroforestry. Functional capacity of soil microbial communities was estimated by degradation of 31 substrates on Biolog EcoPlates™. Microbial community composition and biomass were characterised by phospholipid fatty acid (PLFA) and microbial C and N analyses. All 31 substrates were metabolised in all studied soil types, i.e. functional diversity did not differ. However, both the substrate utilisation rates and the microbial biomass decreased with intensification of land use, and the biomass was positively correlated with organic matter content. Multivariate analysis of PLFA and Biolog EcoPlate™ data showed clear differences between land uses, also indicated by different relative abundance of PLFA markers for certain microorganism groups. In conclusion, our results show that vegetation and land use control the substrate utilisation capacity and microbial community composition and that functional capacity of depleted soils can be restored by active soil management, e.g. forest plantation. However, although 20–30 years of agroforestry farming practises did result in improved soil microbiological and chemical conditions of agricultural soil as compared to traditional agricultural fields, the change was not statistically significant.     

Effect of cinnamic acid on soil microbial characteristics in the cucumber rhizosphere

In order to elucidate the effect of allelochemicals on soil microbial characteristics in the cucumber rhizosphere, the soil microbial biomass and respiration, community functional diversity and RAPD marker diversity as affected by exogenous cinnamic acid were studied. Exogenous cinnamic acid increased soil microbial respiration and the metabolic quotient, but decreased soil microbial biomass-C. Soil microbial community functional diversity and genetic diversity (as indicated by RAPD markers) were also significantly altered by exogenous cinnamic acid. These results suggest that allelochemicals can change soil microbial genetic diversity, biological activity and microbial metabolic activity, which alter soil microbial ecology and accordingly affect the growth of cucumber with accumulation in the soil of allelochemicals.     

Temporal variability in plant and soil nitrogen pools in a high-Arctic ecosystem

This study determined temporal variability in N pools, both aboveground and belowground, across two contrasting plant communities in high-Arctic Spitsbergen, Svalbard (78°N). We measured N pools in plant material, soil microbial biomass and soil organic matter in moist (Alopecurus borealis dominated) and dry (Dryas octopetala dominated) meadow communities at four times during the growing season. We found that plant, microbial and dissolved inorganic and organic N pools were subject to significant, but surprisingly low, temporal variation that was controlled primarily by changes in temperature and moisture availability over the short growing season. This temporal variability is much less than that experienced in other seasonally cold ecosystems such as alpine tundra where strong seasonal partitioning of N occurs between plant and soil microbial pools. While only a small proportion of the total ecosystem N, the microbial biomass represented the single largest of the dynamic N pools in both moist and dry meadow communities (3.4% and 4.6% of the total ecosystem N pool, respectively). This points to the importance of soil microbial community dynamics for N cycling in high-Arctic ecosystems. Microbial N was strongly and positively related to soil temperature in the dry meadow, but this relationship did not hold true in the wet meadow where other factors such as wetter soil conditions might constrain biological activity. Vascular live belowground plant parts represented the single largest plant N pool in both dry and moist meadow, constituting an average of 1.6% of the total N pool in both systems; this value did not vary across the growing season or between plant communities. Overall, our data illustrate a surprisingly low growing season variability in labile N pools in high-Arctic ecosystems, which we propose is controlled primarily by temperature and moisture.     

Direct and indirect effects of tree diversity drive soil microbial diversity in temperate deciduous forest

We investigated the link between aboveground and belowground diversity in temperate deciduous forest ecosystems. To this end, we determined the effects of the tree species composition on the biomass and composition of the soil microbial community using phospholipid fatty acid (PLFA) profiles in the Hainich National Park, a deciduous mixed forest on loess over limestone in Central-Germany. We investigated the effects of the leaf litter composition on the microbial community, hypothesizing that distinctive leaf litter compositions increase signature PLFAs. In addition, we studied the impact of clay content, pH and nutrient status of the soil on the microbial community in different surface soil layers. Consequently, soil was sampled from depths of 0-5 cm, 5-10 cm and 10-20 cm. Plots with highest leaf litter diversity had the largest total amounts of fatty acids, but only PLFA 16:1ω5, which is a common marker for arbuscular mycorrhizal fungi, was significantly increased. In the uppermost soil layer, the pH explained most of the variance in microbial composition. In the deeper surface soil layers, nutrients such as carbon, nitrogen and phosphorus determined the microbial abundances and composition. Our results suggest that the soil microbial community is mainly indirectly influenced by aboveground diversity. Changes in soil pH or the soil nutrient status that are driven by specific plant traits like leave litter quality drive these indirect changes. Specific direct interactions are most reasonable for mycorrhizal fungi.     

Mechanisms of real and apparent priming effects and their dependence on soil microbial biomass and community structure: critical review

The number of studies on priming effects (PE) in soil has strongly increased during the last years. The information regarding real versus apparent PE as well as their mechanisms remains controversial. Based on a meta-analysis of studies published since 1980, we evaluated the intensity, direction, and the reality of PE in dependence on the amount and quality of added primers, the microbial biomass and community structure, enzyme activities, soil pH, and aggregate size. The meta-analysis allowed revealing quantitative relationships between the amounts of added substrates as related to microbial biomass C and induced PE. Additions of easily available organic C up to 15% of microbial biomass C induce a linear increase of extra CO₂. When the added amount of easily available organic C is higher than 50% of the microbial biomass C, an exponential decrease of the PE or even a switch to negative values is often observed. A new approach based on the assessment of changes in the production of extracellular enzymes is suggested to distinguish real and apparent PE. To distinguish real and apparent PE, we discuss approaches based on the C budget. The importance of fungi for long-term changes of SOM decomposition is underlined. Priming effects can be linked with microbial community structure only considering changes in functional diversity. We conclude that the PE involves not only one mechanism but a succession of processes partly connected with succession of microbial community and functions. An overview of the dynamics and intensity of these processes as related to microbial biomass changes and C and N availability is presented.     

The influence of different types of urban land use on soil microbial biomass and functional diversity in Beijing,China

Soil microbes in urban ecosystems are affected by a variety of abiotic and biotic factors resulting from changes in land use. However, the influence of different types of land use on soil microbial properties and soil quality in urban areas remains largely unknown. Here, by comparing five types of land use: natural forest, park, agriculture, street green and roadside trees, we examined the effects of different land uses on soil microbial biomass and microbial functional diversity in Beijing, China. We found that soil properties varied with land uses in urban environments. Compared to natural forest, soil nutrients under the other four types of urban land use were markedly depleted, and accumulation of Cu, Zn, Pb and Cd was apparent. Importantly, under these four types of land use, there was less microbial biomass, but it had greater functional diversity, particularly in the roadside‐tree soils. Furthermore, there were significant correlations between the microbial characteristics and physicochemical properties, such as organic matter, total nitrogen and total phosphorus (P < 0.05), suggesting that lack of nutrients was the major reason for the decrease in microbial biomass. In addition, the larger C/N ratio, Ni concentration and pool of organic matter together with a higher pH contributed to the increase in microbial functional diversity in urban soils. We concluded that different land uses have indirect effects on soil microbial biomass and microbial community functional diversity through their influence on soil physicochemical properties, especially nutrient availability and heavy metal content.     

中国北方温带草原土壤微生物对气温变暖与添加氮素的响应特征

The responses of soil microbes to global warming and nitrogen enrichment can profoundly affect terrestrial ecosystem functions and the ecosystem feedbacks to climate change. However, the interactive effect of warming and nitrogen enrichment on soil microbial community is unclear. In this study, individual and interactive effects of experimental warming and nitrogen addition on the soil microbial community were investigated in a long-term field experiment in a temperate steppe of northern China. The field experiment started in 2006 and soils were sampled in 2010 and analyzed for phospholipid fatty acids to characterize the soil microbial communities. Some soil chemical properties were also determined. Five-year experimental warming significantly increased soil total microbial biomass and the proportion of Gram-negative bacteria in the soils. Long-term nitrogen addition decreased soil microbial biomass at the 0-10 cm soil depth and the relative abundance of arbuscular mycorrhizal fungi in the soils. Little interactive effect on soil microbes was detected when experimental warming and nitrogen addition were combined. Soil microbial biomass positively correlated with soil total C and N, but basically did not relate to the soil C/N ratio and pH. Our results suggest that future global warming or nitrogen enrichment may significantly change the soil microbial communities in the temperate steppes in northern China.     

长期施用化肥对塿土微生物多样性的影响

【目的】土壤微生物在土壤有机质分解、营养循环、植物生长等方面都发挥着重要作用,研究长期施用化肥对其产生的影响可为农田合理施用化肥、培肥土壤和高产高效可持续性农业生产提供理论依据。【方法】以陕西杨凌"国家黄土肥力与肥料效益监测基地"的长期定位试验为基础,利用BIOLOG分析并结合常规分析研究了6种长期不同化肥施用方式不施肥(CK)、单施氮肥(N)、氮钾配施(NK)、磷钾配施(PK)、氮磷配施(NP)和氮磷钾配施(NPK)对土土壤微生物量和微生物功能多样性的影响。【结果】与不施肥CK相比,长期单施氮肥(N)的SMBC、SMBN没有显著变化,但明显降低了土壤微生物商和土壤微生物对氮素的利用;NP和NPK配施能够显著增加土SMBC和SMBN含量,明显提高了土壤微生物商,使土壤微生物种群结构发生了明显变化但土壤微生物对氮素的利用没有显著提高;长期偏施肥处理(NK、PK)的SMBC、SMBN和微生物商虽轻微增加,但土壤微生物种群结构没有显著改变。BIOLOG分析结果显示施磷处理(PK、NP和NPK)对微生物代谢活性的促进作用较大且在培养初期代谢活性较不施磷处理(CK、N和NK)增加显著。长期单施氮肥(N)对于提高土壤微生物多样性没有显著作用而其他化肥施用处理可以提高土土壤微生物群落的碳源利用能力、物种的丰富度和优势度,其中NP和NPK处理配施效果最好。化肥施用对土土壤微生物群落的均匀度没有显著影响。主成分分析的结果表明不同处理的土壤微生物对碳源利用表现出显著差异,氮磷养分的差异是产生分异的主要原因。【结论】土区小麦玉米轮作下,平衡施肥(NP或NPK配施)对于改善农田土壤微生物特性具有良好作用。     

Structural and functional diversity of soil bacterial and fungal communities following woody plant encroachment in the southern Great Plains

In the southern Great Plains (USA), encroachment of grassland ecosystems by Prosopis glandulosa (honey mesquite) is widespread. Mesquite encroachment alters net primary productivity, enhances stores of C and N in plants and soil, and leads to increased levels of soil microbial biomass and activity. While mesquite’s impact on the biogeochemistry of the region is well established, it effects on soil microbial diversity and function are unknown. In this study, soils associated with four plant types (C₃ perennial grasses, C₄ midgrasses, C₄ shortgrasses, and mesquite) from a mesquite-encroached mixed grass prairie were surveyed to in an attempt to characterize the structure, diversity, and functional capacity of their soil microbial communities. rRNA gene cloning and sequencing were used in conjunction with the GeoChip functional gene array to evaluate these potential differences. Mesquite soil supported increased bacterial and fungal diversity and harbored a distinct fungal community relative to other plant types. Despite differences in composition and diversity, few significant differences were detected with respect to the potential functional capacity of the soil microbial communities. These results may suggest that a high level of functional redundancy exists within the bacterial portion of the soil communities; however, given the bias of the GeoChip toward bacterial functional genes, potential functional differences among soil fungi could not be addressed. The results of this study illustrate the linkages shared between above- and belowground communities and demonstrate that soil microbial communities, and in particular soil fungi, may be altered by the process of woody plant encroachment.     

Effects of harvester ant (<Emphasis Type="Italic">Messor</Emphasis> spp.) activity on soil properties and microbial communities in a Negev Desert ecosystem

Harvester ants (Messor spp.) function as an essential link between aboveground resources and below-ground biota such as the microbial community. We examined changes in soil microbial biomass and functional diversity resulting from harvester ant (Messor spp.) activity in the Negev Desert, Israel. Abiotic and biotic soil parameters were recorded during two seasons—wet and dry—also representing food availability periods for the ants (low and high seed availability, respectively). Soil samples were collected monthly from the 0- to 10- and 10- to 20-cm soil layers: (1) near the nest entrance, (2) under chaff piles, and (3) at a 2-m radius from the nest entrance (control). Harvester ant activity increased the percentage of organic matter, total soluble nitrogen, and microbial activity in nest-modified soils in comparison to the control soils. Higher CO₂ evolution was recorded in the low-seed season in ant nest soils than in the control soils. During the high-seed season, higher carbon dioxide evolution was recorded only at the nest entrance locations. There were no differences in microbial biomass between the low- and high-seed seasons, but highest microbial biomass was found under chaff in low-seed season and in nest soils in high-seed season. Microbial functional diversity was higher in nest-modified soils than in the control soils. This study suggests that the effect of harvester ant nests on soil fertility is due to increased microbial biomass and microbial activity in ant nest-modified soils.     

Rapid effects of plant species diversity and identity on soil microbial communities in experimental grassland ecosystems

Changes in plant community structure, including the loss of plant diversity may affect soil microbial communities. To test this hypothesis, plant diversity and composition were experimentally varied in grassland plots cultivated with monocultures or mixtures of 2, 3 or 4 species. We tested the effects of monocultures versus mixtures and of plant species composition on culturable soil bacterial activity, number of substrates used and catabolic diversity, microbial biomass N, microbial respiration, and root biomass. These properties were all measured 10 months after seeding the experiment. Soil bacterial activity, number of substrates used and catabolic diversity were measured in the different plant communities using BIOLOG GN and GP microplates, which are redox-based tests measuring capacity of soil culturable bacteria to use a variety of organic substrates. Microbial biomass N, microbial respiration, and root biomass were insensitive to plant diversity. Culturable soil microbial activity, substrates used and diversity declined with declining plant diversity. Their activity, number of substrates used and diversity were significantly higher in plots with 3 and 4 plant species than in monocultures and in plots with 2 species. There was also an effect of plant species composition. Culturable soil microbial activity and diversity was higher in the four-species plant community than in any of the plant monocultures suggesting that the effect of plant diversity could not be explained by the presence of a particular plant species. Our results showed that changes in plant diversity and composition in grassland ecosystems lead to a rapid response of bacterial activity and diversity.     

Effect of vegetation manipulation of abandoned arable land on soil microbial properties

 The effect of vegetation composition on various soil microbial properties in abandoned arable land was investigated 2 years after agricultural practice had terminated. Microbial numbers and processes were determined in five replicate plots of each of the following treatments: continued agricultural practice (monoculture of buckwheat in 1997), natural colonization by the pioneer community (arable weeds), and manipulated colonization from low (four species, three functional groups: grasses, forbs and legumes) or high diversity (15 species, three functional groups) seed mixtures from plant species that are characteristic of abandoned fields in later successional stages. The results indicated that differences in above-ground plant biomass, plant species composition and plant species diversity had no significant effect on soil microbial processes (net N mineralization, short-term nitrification, respiration and Arg ammonification), microbial biomass C and N (fumigation-incubation) or colony-forming units of the major microbial groups. Hence, there were no indications that soil microbial processes responded differently within 2 years of colonization of abandoned arable land by later successional plants as compared to that by plants from the natural pioneer weed community. Therefore, it seems that during the first few years after arable field abandonment, plants are more dependent on the prevailing soil microbiological conditions than vice versa. Received: 8 April 1999     

Influence of disturbance and nitrogen addition on plant and soil animal diversity in grassland

Two key determinants of biological diversity that have been examined in aboveground and aquatic systems are productivity, or resource supply, and physical disturbance. In this study, we examined how these factors interact under field conditions to determine belowground diversity using microarthropods (mites and Collembola) as our test community. To do this, we established a field manipulation experiment consisting of crossed, continuous gradients of nitrogenous (N) fertilizer addition (up to 240 kg N ha^?1) and disturbance (imitated trampling by cattle) to produce a gradient of soil nutrient availability and disturbance. Due to the relatively short-term nature of our study (i.e. 2 years), we only detected minimal changes in plant diversity due to the experimental manipulations; in the longer term we would expect to detect changes in plant diversity that could potentially impact on soil fauna. However, disturbance reduced, and additions of N increased, aboveground biomass, reflecting the potential effects of these manipulations on resource availability for soil fauna. We found that disturbance strongly reduced the abundance, diversity, and species richness of oribatid mites and Collembola, but had little effect on predatory mites (Mesostigmata). In contrast, N addition, and therefore resource availability, had little effect on microarthropod community structure, but did increase mesostigmatan mite richness and collembolan abundance at high levels of disturbance. Oribatid community structure was mostly influenced by disturbance, whereas collembolan and mesostigmatan diversity were responsive to N addition, suggesting bottom-up control. That maximal species richness of microarthropod groups overall occurred in undisturbed plots, suggests that the microarthropod community was negatively affected by disturbance. We found no change in microarthropod species richness with high N additions, where plant productivity was greatest, indicating that soil biotic communities are unlikely to be strongly regulated by competition. We conclude that the diversity of soil animals is best explained as a combination of their many varied life history tactics, phenology and the heterogeneity of soils that enable so many species to co-exist.