Community food web, decomposition and nitrogen mineralisation in a stratified Scots pine forest soil

A soil community food web model was used to improve the understanding of what factors govern the mineralisation of nutrients and carbon and the decay of dead organic matter. The model derives the rates of C and N mineralisation by organisms by splitting their uptake rate of food resources into a rate at which faeces or prey remains are added to detritus, a rate at which elements are incorporated into biomass, and a rate at which elements are released by organisms as inorganic compounds. The functioning of soil organisms in the mineralisation of C and N was modelled in the soil horizon of a Scots pine forest. The organic horizon was divided into three distinct layers, representing successive stages of decay, i.e. litter, fragmented litter, and humus. Each of the layers had a different, quantitative, biota composition. For each layer the annual C and N mineralisation rates were simulated and compared to observed C and N mineralisation rates from organic matter in stratified litterbags. Simulated C and N mineralisation was relatively close to measured losses of C and N, but the fit was not perfect. Discrepancies between the observed and predicted mineralisation rates are discussed in terms of variation in model parameter values of those organisms that showed the highest contribution to mineralisation rates. The measured, and by the model predicted, significant decrease in mineralisation rates down the profile was not explained by the biomass of the primary decomposers and only partly by the total food web biomass. Modelling results indicated that indirect effects of soil fauna, due to trophic interactions with their resources, are an important explanatory factor. In addition, the analyses suggest that community food web structure is an important factor in the regulation of nutrient mineralisation. The model provided the means to evaluate the contribution of functionally defined groups of organisms, structured in a detrital food web, to losses of C and N from successive decay stages.     

C and N mineralisation in the decomposer food webs of a European forest transect

Belowground processes are essential for the overall carbon and nitrogen fluxes in forests. Neither the functioning of the soil food web mediating these fluxes, nor its modulation by environmental factors is sufficiently understood. In this study the belowground carbon and nitrogen mineralisation of four European coniferous forest sites (northern Sweden to north‐east France) with different climate and N depositional inputs was analysed by investigating the soil food webs using field observations and modelling. The soil fauna directly contributed 7–13% to C mineralisation, among which the testate amoebae (Protozoa) made the largest contribution. Microbial grazing was suggested to have an important indirect effect by stimulating bacterial turnover. Due to relatively high C:N ratios of their substrate, bacteria immobilized N, while the fauna i.e. testate amoebae, nematodes, microarthropods and enchytraeids, counteracted this N immobilisation. Despite similar food web biomass, the sites differed with respect to food web structure and C and N flows. Model calculations suggested a significant influence of food web structure on soil ecosystem processes in addition to environmental factors and resource quality. Mineralisation rates were lowest at the low N input boreal site with a food web dominated by fungal pathways. Further south, as N availability increased, bacterial pathways became more important and the cycling of C and N was faster. The bioavailability of degradable C sources is suggested to be a limiting factor for microbial activity and overall mineralisation rates. In this respect, above‐ and belowground interactions e.g. transfers of labile C sources from the vegetation to the decomposer system deserve further attention. Our study revealed the combined effects of climate and nutrient inputs to ecosystems and the subsequent changes in the structure and functioning of the systems. If decomposition, and therefore carbon loss, is stimulated as a consequence of structural and/or nutritional changes, resulting for example from continuous industrial N emission, the storage capacity of forest ecosystems could be altered.     

Influence of decomposer food web structure and nitrogen availability on plant growth

We studied the sensitivity of soil microbial communities and ecosystem processes to variation in the vertical and horizontal structure of decomposer food web under nitrogen poor and N-enriched conditions. Microcosms with humus and litter layer of boreal forest floor, birch seedlings infected with mycorrhizal fungi, and decomposer food webs with differing trophic group and species composition of soil fauna were constructed. During the second growing period for the birch, we irrigated half of the microcosms with urea solution, and the other half with de-ionised water to create two levels of N concentration in the substrate. During the experiment night time respirations of the microcosms were measured, and the water leached through the microcosms was analysed for concentration of mineral N, and nematode numbers. The microcosms were destructively sampled after 37 weeks for plant biomass and N uptake, structure of soil animal and microbial community (indicated by PLFA profiles), and physical and chemical properties of the humus and litter materials. Predatory mites and nematodes had a negative influence on the biomass of their microbivorous and microbi-detritivorous prey, and microbi-detritivores affected the biomass and community structure of microbes (indicated by PLFA-analysis). Moreover, predatory mites and nematodes increased microbial biomass and changed the microbial community structure. The decomposer food web structure affected also N uptake and growth of plants. Microbi-detritivorous fauna had a positive effect, whereas predators of microbial and detritus feeding fauna exerted a negative influence on plant N uptake and biomass production. The impact of a trophic group on the microbes and plant was also strongly dependent on species composition within the group. Nitrogen addition magnified the influence of food web structure on microbial biomass and plant N uptake. We suggest that addition of urea-N to the soil modified the animal-microbe interaction by increasing microbial growth and altering community structure of microbes. The presence of microbi-detritivores and predators reduced loss of carbon from the microcosms, and the food web structure influenced also water holding capacity of the materials. The changes in plant growth, nutrient cycling, size of N and C pools, and in the physical properties of the soil emphasize the importance and diversity of indirect consequences of decomposer food web structure. This revised version was published online in June 2006 with corrections to the Cover Date.     

Using Stable Isotopes to Differentiate Trophic Feeding Channels within Soil Food Webs

The soil is probably the most diverse habitat there is, with organisms ranging in sizes from less than 1 μm to several metres in length. However, it is increasingly evident that we know little about the interactions occurring between these organisms, the functions that they perform as individual species, or together within their different feeding guilds. These interactions between groups of organisms and physical and chemical processes shape the soil as a habitat and influence the nature of the soil food web with consequences for the above‐ground vegetation and food web. Protists are known as one of the most abundant groups of bacterivores within the soil; however, they are also consumers of a number of other food sources. Even though they are responsible for a large proportion of the mineralisation of bacterial biomass and have a large impact on the C and N cycles within the soil they are regularly overlooked when investigating the complete soil food web. Recently, stable isotopes have been used to determine trophic interactions and here we describe how this technique has been used to highlight linkages between protists and the soil food web.     

Nitrogen Addition and Warming Independently Influence the Belowground Micro-Food Web in a Temperate Steppe

Climate warming and atmospheric nitrogen (N) deposition are known to influence ecosystem structure and functioning. However, our understanding of the interactive effect of these global changes on ecosystem functioning is relatively limited, especially when it concerns the responses of soils and soil organisms. We conducted a field experiment to study the interactive effects of warming and N addition on soil food web. The experiment was established in 2006 in a temperate steppe in northern China. After three to four years (2009–2010), we found that N addition positively affected microbial biomass and negatively influenced trophic group and ecological indices of soil nematodes. However, the warming effects were less obvious, only fungal PLFA showed a decreasing trend under warming. Interestingly, the influence of N addition did not depend on warming.Structural equation modeling analysis suggested that the direct pathway between N addition and soil food web components were more important than the indirect connections through alterations in soil abiotic characters or plant growth. Nitrogen enrichment also affected the soil nematode community indirectly through changes in soil pH and PLFA. We conclude that experimental warming influenced soil food web components of the temperate steppe less than N addition, and there was little influence of warming on N addition effects under these experimental conditions.     

Legacy effects of drought on plant growth and the soil food web

Soils deliver important ecosystem services, such as nutrient provision for plants and the storage of carbon (C) and nitrogen (N), which are greatly impacted by drought. Both plants and soil biota affect soil C and N availability, which might in turn affect their response to drought, offering the potential to feed back on each other’s performance. In a greenhouse experiment, we compared legacy effects of repeated drought on plant growth and the soil food web in two contrasting land-use systems: extensively managed grassland, rich in C and with a fungal-based food web, and intensively managed wheat lower in C and with a bacterial-based food web. Moreover, we assessed the effect of plant presence on the recovery of the soil food web after drought. Drought legacy effects increased plant growth in both systems, and a plant strongly reduced N leaching. Fungi, bacteria, and their predators were more resilient after drought in the grassland soil than in the wheat soil. The presence of a plant strongly affected the composition of the soil food web, and alleviated the effects of drought for most trophic groups, regardless of the system. This effect was stronger for the bottom trophic levels, whose resilience was positively correlated to soil available C. Our results show that plant belowground inputs have the potential to affect the recovery of belowground communities after drought, with implications for the functions they perform, such as C and N cycling.     

On the Importance of Planktonic Protozoans in the Eutrophication Process of the Baltic Sea

Analysing the results of various authors recent studies in the pelagic region of the Baltic revealed that protozoan biomass is in the same range or even higher than metazooplankton biomass. The dominant groups of planktonic protozoans are heterotrophic pico- and nanoflagellates (various taxonomic groups), large heterotrophic flagellates (mainly dinoflagellates) and ciliates. Regularly the spring bloom of phytoplankton is accompanied by a maximum of protozoan biomass which declines in early summer as a result of intensive grazing pressure by metazooplankton and changing food conditions. The analysis of results from different stations indicated that biomasses of protozoans increase with an increasing degree of eutrophication. Several trophic levels within the microbial web should be added to the traditional view on the pelagic food web of the Baltic. Our knowledge regarding the quantitative aspect of the microbial matter flux of the Baltic is very limited up to now and complex ecological (and taxonomical) studies using standardized methods including all protozoan components are necessary. Protozoans (various trophic groups and levels), besides bacteria, should be viewed as the metabolically most active heterotrophic component in the pelagic region of the Baltic, their activity should increase with an increasing degree of eutrophication.     

土壤微食物网结构与生态功能

土壤微食物网是碎屑食物网中与土壤生态过程密切相关的一部分,通过取食资源基质直接或间接地参与养分循环过程,影响陆地生态系统功能.本文从土壤微食物网的组成、结构和生态功能等方面综述了近年来土壤微食物网的研究进展.通过对土壤微食物网的能量通道及营养级联效应的介绍,阐述了土壤微食物网在碳(C)、氮(N)转化、凋落物分解和植物生长等方面的重要作用.针对目前的研究现状,提出未来土壤生态学研究应与高通量测序及稳定同位素技术相结合;通过构建模型进一步加强对土壤食物网结构和功能的研究,从而深入揭示地下生态过程及其对地上植物生长的反馈作用机理.     

Grazing simplifies soil micro‐food webs and decouples their relationships with ecosystem functions in grasslands

Livestock grazing often alters aboveground and belowground communities of grasslands and their mediated carbon (C) and nitrogen (N) cycling processes at the local scale. Yet, few have examined whether grazing‐induced changes in soil food webs and their ecosystem functions can be extrapolated to a regional scale. We investigated how large herbivore grazing affects soil micro‐food webs (microbes and nematodes) and ecosystem functions (soil C and N mineralization), using paired grazed and ungrazed plots at 10 locations across the Mongolian Plateau. Our results showed that grazing not only affected plant variables (e.g., biomass and C and N concentrations), but also altered soil substrates (e.g., C and N contents) and soil environment (e.g., soil pH and bulk density). Grazing had strong bottom‐up effects on soil micro‐food webs, leading to more pronounced decreases at higher trophic levels (nematodes) than at lower trophic levels (microbes). Structural equation modeling showed that changes in plant biomass and soil environment dominated grazing effects on microbes, while nematodes were mainly influenced by changes in plant biomass and soil C and N contents; the grazing effects, however, differed greatly among functional groups in the soil micro‐food webs. Grazing reduced soil C and N mineralization rates via changes in plant biomass, soil C and N contents, and soil environment across grasslands on the Mongolian Plateau. Spearman's rank correlation analysis also showed that grazing reduced the correlations between functional groups in soil micro‐food webs and then weakened the correlation between soil micro‐food webs and soil C and N mineralization. These results suggest that changes in soil micro‐food webs resulting from livestock grazing are poor predictors of soil C and N processes at regional scale, and that the relationships between soil food webs and ecosystem functions depend on spatial scales and land‐use changes.     

A Vigorous Specialized Microbial Food Web in the Suboxic Waters of a Shallow Subtropical Coastal Lagoon

To examine the extent of the microbial food web in suboxic waters of a shallow subtropical coastal lagoon, the density and biomass of bacteria and protozooplankton were quantified under different dissolved oxygen (DO) levels. In addition, bottom waters of a stratified site were compared with bottom waters of a homogeneous site under periods of high and low biological oxygen production/consumption in the lagoon. At the stratified site, microbial biomass decreased with oxygen decline, from oxia to suboxia, with a recovery of the initial total biomass after a 20-day period of persistent suboxia. A peak in density and biomass of purple sulfur bacteria (PSB) (90 μg C L(-1)) occurred in the suboxic waters 20 days prior to the peak in biomass of ciliates >50 μm (Loxophyllum sp. of 150 μm) (160 μg C L(-1)), demonstrating a top down biomass control. Ciliates >50 μm were positively correlated with PSB and bacteriochlorophyll a (photosynthetic pigment of PSB). Total protozoan biomass reached 430 μg C L(-1) in the suboxic waters of the stratified site, with ciliates >50 μm accounting for 90% of the total ciliate biomass and of 55 % of biomass of protozoa. At the homogeneous site, total protozoan biomass was only 66 μg C L(-1), where flagellates and ciliates <25 μm were the dominant microorganisms. Therefore, as light is available for primary producers in the bottom waters of shallow stratified coastal lagoons or estuaries, one can expect that high primary production of PSB may favor a specialized microbial food web composed by larger microorganisms, accessible to zooplankton that tolerate low DO levels.     

线虫区系分析指示土壤食物网结构和功能研究进展

土壤食物网结构复杂,功能众多,直接测定土壤食物网各功能群生物量并结合数学模型来推断土壤食物网结构和功能,工作量大且分析过程繁琐。线虫生态学的发展为土壤食物网的研究开辟了一条新的思路,即利用线虫区系分析来定性推断食物网的结构和功能。线虫作为土壤中数量最丰富的后生动物,占据着土壤食物网的中心位置,其物种多样性、食性多样性、生活史策略多样性、功能团多样性奠定了其作为土壤食物网结构和功能指示生物的生态学基础。线虫区系分析根据发展历史可以分为个体分类、生活史策略分类、功能团分类和代谢足迹分类四个时期,其中后两个时期主要用于推断土壤食物网结构和功能。基于功能团的线虫区系分析将线虫的食性和生活史策略结合起来,发展出一系列指数来判断土壤食物网的连通性、食物网链长度、外界养分投入情况、分解途径及对外界干扰的响应等。基于代谢足迹的线虫区系分析在功能团分析基础上,加入线虫能流分析,从而定性反映了土壤食物网功能的大小。两者在指示土壤食物网自下而上调节及对植物线虫控制等方面起着重要的作用。     

Nematode Indicators of Organic Enrichment

The organisms of the soil food web, dependent on resources from plants or on amendment from other sources, respond characteristically to enrichment of their environment by organic matter. Primary consumers of the incoming substrate, including bacteria, fungi, plant-feeding nematodes, annelids, and some microarthropods, are entry-level indicators of enrichment. However, the quantification of abundance and biomass of this diverse group, as an indicator of resource status, requires a plethora of extraction and assessment techniques. Soluble organic compounds are absorbed by bacteria and fungi, while fungi also degrade more recalcitrant sources. These organisms are potential indicators of the nature of incoming substrate, but current methods of biomass determination do not reliably indicate their community composition. Guilds of nematodes that feed on bacteria (e.g., Rhabditidae, Panagrolaimidae) and fungi (e.g., Aphelenchidae, Aphelenchoididae) are responsive to changes in abundance of their food. Through direct herbivory, plant-feeding nematodes (e.g., many species of Tylenchina) also contribute to food web resources. Thus, analysis of the nematode community of a single sample provides indication of carbon flow through an important herbivore channel and through channels mediated by bacteria and fungi. Some nematode guilds are more responsive than others to resource enrichment. Generally, those bacterivores with short lifecycles and high reproductive potential (e.g., Rhabditidae) most closely mirror the bloom of bacteria or respond most rapidly to active plant growth. The feeding habits of some groups remain unclear. For example, nematodes of the Tylenchidae may constitute 30% or more of the individuals in a soil sample; further study is necessary to determine which resource channels they portray and the appropriate level of taxonomic resolution for this group. A graphic representation of the relative biomass of bacterivorous, fungivorous, and herbivorous nematodes provides a useful tool for assessing the importance of the bacterial, fungal, and plant resource channels in an extant food web.     

Soil Food Web Changes during Spontaneous Succession at Post Mining Sites: A Possible Ecosystem Engineering Effect on Food Web Organization?

Parameters characterizing the structure of the decomposer food web, biomass of the soil microflora (bacteria and fungi) and soil micro-, meso- and macrofauna were studied at 14 non-reclaimed 1– 41-year-old post-mining sites near the town of Sokolov (Czech Republic). These observations on the decomposer food webs were compared with knowledge of vegetation and soil microstructure development from previous studies. The amount of carbon entering the food web increased with succession age in a similar way as the total amount of C in food web biomass and the number of functional groups in the food web. Connectance did not show any significant changes with succession age, however. In early stages of the succession, the bacterial channel dominated the food web. Later on, in shrub-dominated stands, the fungal channel took over. Even later, in the forest stage, the bacterial channel prevailed again. The best predictor of fungal bacterial ratio is thickness of fermentation layer. We argue that these changes correspond with changes in topsoil microstructure driven by a combination of plant organic matter input and engineering effects of earthworms. In early stages, soil is alkaline, and a discontinuous litter layer on the soil surface promotes bacterial biomass growth, so the bacterial food web channel can dominate. Litter accumulation on the soil surface supports the development of the fungal channel. In older stages, earthworms arrive, mix litter into the mineral soil and form an organo-mineral topsoil, which is beneficial for bacteria and enhances the bacterial food web channel.     

The role of prey nutritional status in governing protozoan nitrogen regeneration efficiency

Laboratory experiments were conducted to study nitrogen (N) regeneration by the heterotrophic marine dinoflagellate Oxyrrhis marina when ingesting phytoplankton prey of two different species and of two alternative carbon:nitrogen (C:N) ratios. Experiments were conducted in the presence of L-methionine sulfoximine (MSX) which acts as a glutamine synthetase inhibitor. Utilisation by phytoplankton of N regenerated by protozoans and other organisms drives secondary production in marine food webs. However, the rapid utilisation of this N by phytoplankton has previously hampered accurate assessment of the efficiency of protozoan N regeneration. This phenomenon is particularly problematic when the phytoplankton are nutrient stressed and most likely to rapidly utilise N. The use of MSX prevented significant utilisation by phytoplankton of protozoan regenerated N. Hence, by removing the normal pathway of N cycling, we were able to determine the N regeneration efficiency (NRE) of the protozoan. The results suggested that predator NRE could be explained in terms of the relative CN stoichiometry of prey and predator. Using a mathematical model we demonstrated that changing the method used to simulate the NRE of the protozoan trophic level has the potential to markedly modify the predicted dynamics of the simulated microbial food web.     

Effects of microbivore species composition and basal resource enrichment on trophic-level biomasses in an experimental microbial-based soil food web

Previous theoretical and empirical evidence suggests that species composition within trophic levels may profoundly affect the response of trophic-level biomasses to enhanced basal resources. To test whether species composition of microbivorous nematodes has such an effect in microbial-based soil food webs, I created three microcosm food webs, consisting of bacteria, fungi, bacterial-feeding nematodes (Acrobeloides tricornus, Caenorhabditis elegans), fungal-feeding nematodes (Aphelenchus avenae, Aphelenchoides sp.) and a predatory nematode (Prionchulus punctatus). The food webs differed in species composition at the second trophic level: food web A included A. tricornus and Aph. avenae, food web B included C. elegans and Aphelenchoides sp., and food web AB included all four species. I increased basal resources by adding glucose to half of the replicates of each food web, and sampled microcosms destructively four times during a 22-week experiment to estimate the biomass of organisms at each trophic level. Microbivore species composition significantly affected bacterivore and fungivore biomass but not bacterial, fungal or predator biomass. Greatest bacterivore and fungivore biomass was found in food web A, intermediate biomass in food web AB, and smallest biomass in food web B. Basal resource addition increased the biomass of microbes and microbivores but did not affect predator biomass. Importantly, microbivore species composition did not significantly modify the effect of additional resources on trophic-level biomasses. The presence of a competitor reduced the biomass of A. tricornus and Aph. avenae, in that the biomass of these species was less in food web AB than in food web A, whereas the biomass of C. elegans and Aphelenchoides sp. was not affected by their potential competitors. The biomass of Aph. avenae increased with additional resources in the absence of the competitor only, while the biomass of A. tricornus and Aphelenchoides sp. increased also in the presence of their competitors. The results imply that microbivore species composition may determine the second-level biomass in simple microbe-nematode food webs, but may not significantly affect biomass at other levels or modify the response of trophic-level biomasses to enhanced basal resources. The study also shows that even if the role of predation in a food web is diminished, the positive response of organisms to increased resource availability may still be hindered by competition. Received: 22 June 1998 / Accepted: 28 August 1998     

Contribution of nematodes to the structure and function of the soil food web

As carbon and energy flow through the soil food web they are depleted by the metabolic and production functions of organisms. To be sustained, a "long" food web, with a large biomass at higher trophic levels, must receive a high rate of rhizodeposition or detrital subsidy, or be top-populated by organisms of slow growth and long life cycle. Disturbed soil food webs tend to be bottom heavy and recalcitrant to restoration due to the slow growth of upper predator populations, physical and chemical constraints of the soil matrix, biological imbalances, and the relatively low mobility and invasion potential of soil organisms. The functional roles of nematodes, determined by their metabolic and behavioral activities, may be categorized as ecosystem services, disservices or effect-neutral. Among the disservices attributable to nematodes are overgrazing, which diminishes services of prey organisms, and plant-damaging herbivory, which reduces carbon fixation and availability to other organisms in the food web. Unfortunately, management to ameliorate potential disservices of certain nematodes results in unintended but long-lasting diminution of the services of others. Beneficial roles of nematodes may be enhanced by environmental stewardship that fosters greater biodiversity and, consequently, complementarity and continuity of their services.     

Effects of food‐web structure on periphyton stoichiometry in eutrophic lakes: a mesocosm study

1. Aquatic herbivores typically have much higher concentrations of nutrients (e.g. N and P) in their tissues than there is in the food they eat. These stoichiometric differences can cause herbivores to be limited by the elemental quality of their food, which could affect, in turn, the structure of consumer communities and even alter key ecosystem processes. 2. In streams and in the littoral zone of shallow lakes, periphyton is an important food resource for benthic animals. Studying the elemental composition of periphyton may help us to understand food‐web structure, and any reciprocal effect of this structure on periphyton stoichiometry. 3. To understand how alterations in the food‐web structure affect the elemental composition of periphyton in a eutrophic lake, we carried out a long‐term experiment (14 months) in large‐scale mesocosms (40 m³), in which we manipulated food‐web structure, and which were dominated either by planktivorous fish (Rutilus rutilus) or herbivorous invertebrates (without fish). Periphyton was sampled monthly at three depths (0.5, 1.5 and 2.5 m) to determine its biomass and elemental composition (C/N/P ratio). Food‐web structure, physical and chemical parameters were monitored throughout the experiment. 4. Fish had indirect positive effect on periphyton biomass, leading to twofold higher levels than in herbivore‐dominated mesocosms. This result was probably due to control of benthic consumers by fish, suggesting a strong top–down control on periphyton by their consumers in fishless enclosures. 5. The elemental ratios C/P and C/N were lower in deep water in both treatments, mainly mediated by light availability, in accordance with the light/nutrient ratio hypothesis. These ratios were also lower in fishless treatments, probably due to increases in inorganic nutrient availability and grazing pressure in herbivore‐dominated systems. During winter, periphyton elemental composition was similar in both treatments, and was unrelated to inorganic nutrient availability. 6. These results indicate that any alteration of food‐web structure in lakes, such as in biomanipulation experiments, is likely to modify both the biomass and elemental quality of periphyton. Resultant effects on the consumers of periphyton and macrophytes could play a key role in the success of biomanipulations and should be taken into account in further studies.     

Establishment of trophic continuum in the food web of the Yellow Sea and East China Sea ecosystem: Insight from carbon and nitrogen stable isotopes

The determination of trophic level for the biology in a marine ecosystem is very important as alteration of its structure and function may be reflected in the tro-phic level of component species. A change in trophic level indicates variation in an organism’s feeding bi-ology or in the pathway of energy flow from primary producers to the consumer. The gut content analysis is a traditional method for studying trophodynamics of food web in marine ecosystems. Species composition and amounts in al…     

基于土壤食物网的生态系统复杂性-稳定性关系研究进展

复杂性-稳定性关系是生态学核心问题之一。作为模式食物网,土壤食物网在探索生态系统复杂性-稳定性关系中起了极大的作用。总结了以Moore、de Ruiter、Neutel等为代表的理论生态学家以土壤食物网为工具研究生态系统复杂性-稳定性关系的方法、结论及不足之处,并展望了未来的发展方向。Moore、de Ruiter、Neutel等将土壤食物网功能群生物量数据、土壤食物网Lotka-Volterra模型和面向过程模型三者结合起来,描述相互作用强度大小格局、分室、能流组织形式等复杂性特征;将土壤食物网Lotka-Volterra模型与群落矩阵结合起来分析局域稳定性,进而探讨生态系统复杂性-稳定性关系的一般规律。在此基础上,Moore、de Ruiter、Neutel等证明了与随机食物网相比,真实食物网的相互作用强度格局、分室等复杂性特征提高了生态系统稳定性,生产力与稳定性共同决定了食物链的长度,并指出建立在平衡态基础上的静态土壤食物网模型在探索生态系统复杂性-稳定性关系方面具有较大的不足,动态土壤食物网是未来以土壤食物网为工具研究生态系统复杂性-稳定性关系的发展方向。     

Food web complexity affects stoichiometric and trophic interactions

The stoichiometry of trophic interactions has mainly been studied in simple consumer–prey systems, whereas natural systems often harbour complex food webs with abundant indirect effects. We manipulated the complexity of trophic interactions by using simple laboratory food webs and complex field food webs in enclosures in Lake Erken. In the simple food web, one producer assemblage (periphyton) and its consumers (benthic snails) were amended by perch, which was externally fed by fish food. In the complex food web, two producer assemblages (periphyton and phytoplankton), their consumers (benthic invertebrates and zooplankton) and perch feeding on zooplankton were included. In the simple food web perch affected the stoichiometry of periphyton and increased periphyton biomass and the concentration of dissolved inorganic nitrogen. Grazers reduced periphyton biomass but increased its nutrient content. In the complex food web, in contrast to the simple food web, perch affected periphyton biomass negatively but increased phytoplankton abundance. Perch had no influence on benthic invertebrate density, zooplankton biomass or periphyton stoichiometry. Benthic grazers reduced periphyton biomass and nutrient content. The difference between the simple and the complex food web was presumably due to the increase of pelagic cyanobacteria ( Gloeotrichia sp.) with fish presence in the complex food web, thus fish had indirect negative effects on periphyton biomass through nutrient competition and shading by cyanobacteria. We conclude that the higher food web complexity through the presence of pelagic primary producers (in this case Gloeotrichia sp.) influences the direction and strength of trophic and stoichiometric interactions.