Soil acidification reduces the effects of short‐term nutrient enrichment on plant and soil biota and their interactions in grasslands

Soil nitrogen (N) and phosphorus (P) contents, and soil acidification have greatly increased in grassland ecosystems due to increased industrial and agricultural activities. As major environmental and economic concerns worldwide, nutrient enrichment and soil acidification can lead to substantial changes in the diversity and structure of plant and soil communities. Although the separate effects of N and P enrichment on soil food webs have been assessed across different ecosystems, the combined effects of N and P enrichment on multiple trophic levels in soil food webs have not been studied in semiarid grasslands experiencing soil acidification. Here we conducted a short‐term N and P enrichment experiment in non‐acidified and acidified soil in a semiarid grassland on the Mongolian Plateau. We found that net primary productivity was not affected by N or P enrichment alone in either non‐acidified or acidified soil, but was increased by combined N and P enrichment in both non‐acidified and acidified soil. Nutrient enrichment decreased the biomass of most microbial groups in non‐acidified soil (the decrease tended to be greatest with combined N and P enrichment) but not in acidified soil, and did not affect most soil nematode variables in non‐acidified or acidified soil. Nutrient enrichment also changed plant and microbial community structure in non‐acidified but not in acidified soil, and had no effect on nematode community structure in non‐acidified or acidified soil. These results indicate that the responses to short‐term nutrient enrichment were weaker for higher trophic groups (nematodes) than for lower trophic groups (microorganisms) and primary producers (plants). The findings increase our understanding of the effects of nutrient enrichment on multiple trophic levels of soil food webs, and highlight that soil acidification, as an anthropogenic stressor, reduced the responses of plants and soil food webs to nutrient enrichment and weakened plant–soil interactions.     

Grazing intensity significantly affects belowground carbon and nitrogen cycling in grassland ecosystems: a meta‐analysis

Livestock grazing activities potentially alter ecosystem carbon (C) and nitrogen (N) cycles in grassland ecosystems. Despite the fact that numerous individual studies and a few meta‐analyses had been conducted, how grazing, especially its intensity, affects belowground C and N cycling in grasslands remains unclear. In this study, we performed a comprehensive meta‐analysis of 115 published studies to examine the responses of 19 variables associated with belowground C and N cycling to livestock grazing in global grasslands. Our results showed that, on average, grazing significantly decreased belowground C and N pools in grassland ecosystems, with the largest decreases in microbial biomass C and N (21.62% and 24.40%, respectively). In contrast, belowground fluxes, including soil respiration, soil net N mineralization and soil N nitrification increased by 4.25%, 34.67% and 25.87%, respectively, in grazed grasslands compared to ungrazed ones. More importantly, grazing intensity significantly affected the magnitude (even direction) of changes in the majority of the assessed belowground C and N pools and fluxes, and C : N ratio as well as soil moisture. Specifically,light grazing contributed to soil C and N sequestration whereas moderate and heavy grazing significantly increased C and N losses. In addition, soil depth, livestock type and climatic conditions influenced the responses of selected variables to livestock grazing to some degree. Our findings highlight the importance of the effects of grazing intensity on belowground C and N cycling, which may need to be incorporated into regional and global models for predicting effects of human disturbance on global grasslands and assessing the climate‐biosphere feedbacks.     

食微线虫对植物生长及土壤养分循环的影响

近二十多年来, 土壤动物的生态功能受到广泛重视。越来越多的证据表明, 土壤动物和微生物间的相互作用对土壤生态系统过程和植物生长起着重要的调节作用。本文综述了食细菌线虫和食真菌线虫对土壤微生物、土壤氮矿化和植物生长的影响。大量研究发现, 食细菌线虫和食真菌线虫都有助于土壤氮素等养分矿化, 从而促进植物生长。这种作用主要是线虫通过取食活动加速微生物周转, 并通过代谢分泌和释放微生物所固持的养分而实现的。但这种作用会因不同的线虫、微生物和植物的种类以及土壤基质的C/N营养状况而异, 此外还受线虫的营养类群及其与其他土壤动物之间复杂关系的影响。今后应该加强以下几方面的研究: (1)深入研究线虫、微生物和植物之间相互作用的机制; (2) 增加控制实验系统的复杂性, 研究线虫不同功能群之间及其与其他土壤动物之间的关系; (3)加强长期实验和观察, 在较长的时间尺度上了解线虫的生态功能; (4)加强对不同生态系统的研究, 在更大的空间尺度上综合了解土壤线虫的生态功能; (5)在全球气候变化的背景下了解土壤线虫的响应, 并预测土壤线虫对全球变化的反馈。     

The response of a three trophic level soil food web to the identity and diversity of plant species and functional groups

Despite considerable recent interest in how biodiversity may influence ecosystem properties, the issue of how plant diversity and composition may affect multiple trophic levels in soil food webs remains essentially unexplored. We conducted a glasshouse experiment in which three plant species of each of three functional groups (grasses, N‐fixing legumes and forbs) were grown in monoculture and in mixtures of three species (with the three species being in the same or different functional groups) and all nine species. Plant species identity had important effects on the biomasses or population densities of belowground primary consumers (microbial biomass, herbivorous nematodes) and two groups of secondary consumers (microbe‐feeding nematodes and enchytraeids); the third consumer trophic level (predatory nematodes) was marginally not significantly affected at P=0.05. Plant species also influenced the relative importance of the bacterial‐based and fungal‐based energy channels for both the primary and secondary consumer trophic levels. Within‐group diversity of only the soil microflora and herbivorous nematodes (both representing the basal consumer trophic level) were affected by plant species identity. However, community composition within all trophic groupings considered (herbivorous nematodes, microbes, microbe‐feeding nematodes, predatory nematodes) was strongly influenced by what plant species were present. Despite the strong responses of the soil biota to plant species identity, there were few effects of plant species or functional group richness on any of the belowground response variables measured. Further, net primary productivity (NPP) was unaffected by plant diversity. Since some belowground response variables were correlated with NPP across treatments, it is suggested that belowground responses to plant diversity might become more apparent in situations when NPP itself responds to plant diversity. Our results point to plant species identity as having important multitrophic effects on soil food webs, both at the whole trophic group and within‐group levels of resolution, and suggest that differences in plant traits across species may be important in driving the decomposer subsystem.     

不同强度牦牛放牧对青藏高原高寒草地土壤和植物生物量的影响

放牧作为家畜饲养方式之一,是草地最简单、有效的利用方式,放牧中的家畜对草地生态系统的影响是全球畜牧生态学研究的焦点。过度放牧导致草地退化严重,虽然在青藏高原地区已有较多放牧对草地影响的研究,但探究连续4年放牧对高寒草地生态系统影响的定位实验却鲜见报道。本研究在青藏高原东缘选取典型高寒草地,使用高原特有且分布最广的牦牛作为大型草食放牧家畜,设置了4个牦牛放牧强度（禁牧：无放牧、轻牧：1头/hm²、中牧：2头/hm²和重牧：3头/hm²）以研究其对高寒草地土壤和植物功能的影响。开展4年试验后的结果表明：放牧条件下土壤含水率显著增加;而土壤容重、全磷和有机质含量对放牧强度均无显著性响应;土壤全氮和pH的响应主要在表层0-20 cm,其中全氮为轻牧和重牧处理分别显著高于中牧,中牧处理下的土壤pH为显著高于轻牧;土壤全钾含量在禁牧处理中显著高于放牧处理;而土壤有效氮和速效钾均为中牧处理显著高于禁牧;放牧可以显著降低植物地上生物量。牦牛放牧强度显著影响土壤含水率、有效养分和植物地上生物量,而对其它土壤理化性质影响较弱。本研究结果揭示放牧对高寒草地土壤理化性质和植物地上生物量的影响,为青藏高原高寒草甸生态系统保护、可持续管理和合理放牧率提供理论依据。     

Intensified plant N and C pool with more available nitrogen under experimental warming in an alpine meadow ecosystem

Nitrogen (N) availability is projected to increase in a warming climate. But whether the more available N is immobilized by microbes (thus stimulates soil carbon (C) decomposition), or is absorbed by plants (thus intensifies C uptake) remains unknown in the alpine meadow ecosystem. Infrared heaters were used to simulate climate warming with a paired experimental design. Soil ammonification, nitrification, and net mineralization were obtained by in situ incubation in a permafrost region of the Qinghai‐Tibet Plateau (QTP). Available N significantly increased due to the stimulation of net nitrification and mineralization in 0–30 cm soil layer. Microbes immobilized N in the end of growing season in both warming and control plots. The magnitude of immobilized N was lower in the warming plots. The root N concentration significantly reduced, but root N pool intensified due to the significant increase in root biomass in the warming treatment. Our results suggest that a warming‐induced increase in biomass is the major N sink and will continue to stimulate plant growth until plant N saturation, which could sustain the positive warming effect on ecosystem productivity.     

Heterogeneity of soil and plant N and C associated with individual plants and openings in North American shortgrass steppe

Small-scale spatial heterogeneity of soil organic matter (SOM) associated with patterns of plant cover can strongly influence population and ecosystem dynamics in dry regions but is not well characterized for semiarid grasslands. We evaluated differences in plant and soil N and C between soil from under individual grass plants and from small openings in shortgrass steppe. In samples from 0 to 5 cm depth, root biomass, root N, total and mineralizable soil N, total and respirable organic C, C:N ratio, fraction of organic C respired, and ratio of respiration to N mineralization were significantly greater for soil under plants than soil from openings. These differences, which were consistent for two sites with contrasting soil textures, indicate strong differentiation of surface soil at the scale of individual plants, with relative enrichment of soil under plants in total and active SOM. Between-microsite differences were substantial relative to previously reported differences associated with landscape position and grazing intensity in shortgrass steppe. We conclude that microscale heterogeneity in shortgrass steppe deserves attention in investigation of controls on ecosystem and population processes and when sampling to estimate properties at plot or site scales.     

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.     

The effect of reindeer grazing on decomposition, mineralization and soil biota in a dry oligotrophic Scots pine forest

Reindeer grazing in the Fennoscandian area has a considerable influence on the ground vegetation, and this is likely in turn to have important consequences for the soil biota and decomposition processes. The effects of reindeer grazing on soil biota, decomposition and mineralization processes, and ecosystem properties in a lichen‐dominated forest in Finnish Lapland were studied inside and outside a large long term fenced reindeer exclosure area. Decomposition rates of Vaccinium myrtillus leaves in litter bags were retarded in the grazed area relative to the ungrazed area, as well as in subplots from which lichens had been artificially removed to simulate grazing. The effect of reindeer grazing on soil respiration and microbial C was positive in the lichen and litter layers of the soil profile, but retarded in the humus layer. There was no effect of grazing on gross N mineralization and microbial biomass N in the humus and upper mineral soil layer, but net N mineralization was increased by grazing. In these layers soil respiration was reduced by grazing, indicating that reindeer effects reduce the ratio of C to N mineralized by soil microorganisms. Grazing stimulated populations of all trophic groupings of nematodes in the lichen layer and microbe feeding nematodes in the litter layer, indicating that grazing by reindeer has multitrophic effects on the decomposer food‐web. Grazing decreased lichen and dwarf shrub biomasses and increased the mass of litter present in the litter layer on an areal basis, but did not significantly alter total C storage per unit area in the humus and mineral soil layers. The N concentration of lichens was increased by grazing, but the N concentrations of both living and dead Pinus sylvestris needles and Empetrum hermaphroditum leaves were not affected.
There was some evidence for each of three mechanisms which could account for the grazing effects that we observed in our study. Firstly, reindeer may have changed the composition and quality of litter input by affecting plant species composition and through addition of N from urine and faeces, resulting in a lack of available C relative to N for decomposer organisms. Secondly, the organic matter in the soil may be older in the grazed area, because of reduction of recent production of lichen litter relative to the ungrazed area. The organic matter in the grazed area may have been in a different phase of decomposition from that in the exclosure. Thirdly, the soil microclimate is likely to be affected by reindeer grazing through physical removal of lichen cover on the ground, and this can have a significant influence on soil microbial processes. This is supported by the strong observed effects of experimental removal of lichens on decomposer processes. The impact of reindeer grazing on soil processes may be a result of complex interactions between different mechanisms, and this could help to explain why the below‐ground effects of reindeer grazing have different consequences to those which have been observed in recent investigations on other grazing systems.     

Invasive plants have different effects on trophic structure of green and brown food webs in terrestrial ecosystems: a meta‐analysis

Although invasive plants are a major source of terrestrial ecosystem degradation worldwide, it remains unclear which trophic levels above the base of the food web are most vulnerable to plant invasions. We performed a meta‐analysis of 38 independent studies from 32 papers to examine how invasive plants alter major groupings of primary and secondary consumers in three globally distributed ecosystems: wetlands, woodlands and grasslands. Within each ecosystem we examined if green (grazing) food webs are more sensitive to plant invasions compared to brown (detrital) food webs. Invasive plants have strong negative effects on primary consumers (detritivores, bacterivores, fungivores, and/or herbivores) in woodlands and wetlands, which become less abundant in both green and brown food webs in woodlands and green webs in wetlands. Plant invasions increased abundances of secondary consumers (predators and/or parasitoids) only in woodland brown food webs and green webs in wetlands. Effects of invasive plants on grazing and detrital food webs clearly differed between ecosystems. Overall, invasive plants had the most pronounced effects on the trophic structure of wetlands and woodlands, but caused no detectable changes to grassland trophic structure.     

Herbivore influence on soil microbial biomass and nitrogen mineralization in a northern grassland ecosystem: Yellowstone National Park

Microorganisms are largely responsible for soil nutrient cycling and energy flow in terrestrial ecosystems. Although soil microorganisms are affected by topography and grazing, little is known about how these two variables may interact to influence microbial processes. Even less is known about how these variables influence microorganisms in systems that contain large populations of free-roaming ungulates. In this study, we compared microbial biomass size and activity, as measured by in situ net N mineralization, inside and outside 35- to 40-year exclosures across a topographic gradient in northern Yellowstone National Park. The objective was to determine the relative effect of topography and large grazers on microbial biomass and nitrogen mineralization. Microbial C and N varied by almost an order of magnitude across sites. Topographic depressions that contained high plant biomass and fine-textured soils supported the greatest microbial biomass. We found that plant biomass accurately predicted microbial biomass across our sites suggesting that carbon inputs from plants constrained microbial biomass. Chronic grazing neither depleted soil C nor reduced microbial biomass. We hypothesize that microbial populations in grazed grasslands are sustained mainly by inputs of labile C from dung deposition and increased root turnover or root exudation beneath grazed plants. Mineral N fluxes were affected more by grazing than topography. Net N mineralization rates were highest in grazed grassland and increased from dry, unproductive to mesic, highly productive communities. Overall, our results indicate that topography mainly influences microbial biomass size, while mineral N fluxes (microbial activity) are affected more by grazing in this grassland ecosystem. Received: 4 June 1997 / Accepted: 16 December 1997     

Woody Plant Encroachment by <Emphasis Type="Italic">Juniperus virginiana</Emphasis> in a Mesic Native Grassland Promotes Rapid Carbon and Nitrogen Accrual

The cover and abundance of Juniperus virginiana L. in the U.S. Central Plains are rapidly increasing, largely as a result of changing land-use practices that alter fire regimes in native grassland communities. Little is known about how conversion of native grasslands to Juniperus-dominated forests alters soil nutrient availability and ecosystem storage of carbon (C) and nitrogen (N), although such land-cover changes have important implications for local ecosystem dynamics, as well as regional C and N budgets. Four replicate native grasslands and adjacent areas of recent J. virginiana encroachment were selected to assess potential changes in soil N availability, leaf-level photosynthesis, and major ecosystem C and N pools. Net N mineralization rates were assessed in situ over two years, and changes in labile soil organic pools (potential C and N mineralization rates and microbial biomass C and N) were determined. Photosynthetic nitrogen use efficiencies (PNUE) were used to examine differences in instantaneous leaf-level N use in C uptake. Comparisons of ecosystem C and N stocks revealed significant C and N accrual in both plant biomass and soils in these newly established forests, without changes in labile soil N pools. There were few differences in monthly in situ net N mineralization rates, although cumulative annual net N mineralization was greater in forest soils compared to grasslands. Conversely, potential C mineralization was significantly reduced in forest soils. Encroachment by J. virginiana into grasslands results in rapid accretion of ecosystem C and N in plant and soil pools with little apparent change in N availability. Widespread increases in the cover of woody plants, like J. virginiana, in areas formerly dominated by graminoid species suggest an increasing role of expanding woodlands and forests as regional C sinks in the central U.S.     

Evidence for top predator control of a grazing ecosystem

The importance of top predators in controlling ecological processes in large, intact ecosystems is unclear. In grasslands that support abundant ungulates, top–down control by predators may be particularly important, because of the tight biogeochemical linkages of ungulate prey with plants and soil microbes. Here, I examined the effects of the recent reintroduction of the gray wolf Canis lupus on ecosystem processes in Yellowstone National Park, where herds of grazing ungulates previously have been shown to stimulate several processes, including soil net nitrogen (N) mineralization. Rates of ungulate grazing intensity and soil net N mineralization were compared before and after wolf reintroduction in grasslands ranging five‐fold in aboveground production. Grazing intensity and grassland net N mineralization declined after wolf reintroduction, a likely partial function of fewer ungulates; wolf predation has been one of several factors implicated in causing the decline in Yellowstone ungulates. In addition, the spatial pattern of grazing and net N mineralization changed after reintroduction. A shift in the spatial patterns of grazer‐associated processes is consistent with a growing body of work indicating that wolves have changed habitat use patterns of ungulates in Yellowstone National Park. These findings suggest widespread wolf effects on ungulate prey, plants, and microbial activity that have spatially reorganized grassland energy and nutrient dynamics in Yellowstone Park.     

Uncovering trophic positions and food resources of soil animals using bulk natural stable isotope composition

Despite the major importance of soil biota in nutrient and energy fluxes, interactions in soil food webs are poorly understood. Here we provide an overview of recent advances in uncovering the trophic structure of soil food webs using natural variations in stable isotope ratios. We discuss approaches of application, normalization and interpretation of stable isotope ratios along with methodological pitfalls. Analysis of published data from temperate forest ecosystems is used to outline emerging concepts and perspectives in soil food web research. In contrast to aboveground and aquatic food webs, trophic fractionation at the basal level of detrital food webs is large for carbon and small for nitrogen stable isotopes. Virtually all soil animals are enriched in ¹³C as compared to plant litter. This ‘detrital shift’ likely reflects preferential uptake of ¹³C‐enriched microbial biomass and underlines the importance of microorganisms, in contrast to dead plant material, as a major food resource for the soil animal community. Soil organic matter is enriched in ¹⁵N and ¹³C relative to leaf litter. Decomposers inhabiting mineral soil layers therefore might be enriched in ¹⁵N resulting in overlap in isotope ratios between soil‐dwelling detritivores and litter‐dwelling predators. By contrast, ¹³C content varies little between detritivores in upper litter and in mineral soil, suggesting that they rely on similar basal resources, i.e. little decomposed organic matter. Comparing vertical isotope gradients in animals and in basal resources can be a valuable tool to assess trophic interactions and dynamics of organic matter in soil. As indicated by stable isotope composition, direct feeding on living plant material as well as on mycorrhizal fungi is likely rare among soil invertebrates. Plant carbon is taken up predominantly by saprotrophic microorganisms and channelled to higher trophic levels of the soil food web. However, feeding on photoautotrophic microorganisms and non‐vascular plants may play an important role in fuelling soil food webs. The trophic niche of most high‐rank animal taxa spans at least two trophic levels, implying the use of a wide range of resources. Therefore, to identify trophic species and links in food webs, low‐rank taxonomic identification is required. Despite overlap in feeding strategies, stable isotope composition of the high‐rank taxonomic groups reflects differences in trophic level and in the use of basal resources. Different taxonomic groups of predators and decomposers are likely linked to different pools of organic matter in soil, suggesting different functional roles and indicating that trophic niches in soil animal communities are phylogenetically structured. During last two decades studies using stable isotope analysis have elucidated the trophic structure of soil communities, clarified basal food resources of the soil food web and revealed links between above‐ and belowground ecosystem compartments. Extending the use of stable isotope analysis to a wider range of soil‐dwelling organisms, including microfauna, and a larger array of ecosystems provides the perspective of a comprehensive understanding of the structure and functioning of soil food webs.     

Drought decreases incorporation of recent plant photosynthate into soil food webs regardless of their trophic complexity

Theory suggests that more complex food webs promote stability and can buffer the effects of perturbations, such as drought, on soil organisms and ecosystem functions. Here, we tested experimentally how soil food web trophic complexity modulates the response to drought of soil functions related to carbon cycling and the capture and transfer below‐ground of recent photosynthate by plants. We constructed experimental systems comprising soil communities with one, two or three trophic levels (microorganisms, detritivores and predators) and subjected them to drought. We investigated how food web trophic complexity in interaction with drought influenced litter decomposition, soil CO₂ efflux, mycorrhizal colonization, fungal production, microbial communities and soil fauna biomass. Plants were pulse‐labelled after the drought with ¹³C‐CO₂ to quantify the capture of recent photosynthate and its transfer below‐ground. Overall, our results show that drought and soil food web trophic complexity do not interact to affect soil functions and microbial community composition, but act independently, with an overall stronger effect of drought. After drought, the net uptake of ¹³C by plants was reduced and its retention in plant biomass was greater, leading to a strong decrease in carbon transfer below‐ground. Although food web trophic complexity influenced the biomass of Collembola and fungal hyphal length, ¹³C enrichment and the net transfer of carbon from plant shoots to microbes and soil CO₂ efflux were not affected significantly by varying the number of trophic groups. Our results indicate that drought has a strong effect on above‐ground–below‐ground linkages by reducing the flow of recent photosynthate. Our results emphasize the sensitivity of the critical pathway of recent photosynthate transfer from plants to soil organisms to a drought perturbation, and show that these effects may not be mitigated by the trophic complexity of soil communities, at least at the level manipulated in this experiment.     

土壤动物多样性及其生态功能

土壤无脊椎动物生物量通常小于土壤生物总生物量的10%,但它们种类丰富,取食行为及生活史策略多种多样,且土壤动物之间,土壤动物与微生物之间存在着复杂的相互作用关系。土壤动物的生态功能主要通过取食作用(trophic effect)和非取食作用(non-trophic effect)来实现。原生动物数量大、周转快,故原生动物本身的代谢活动(即取食作用)对碳氮矿化的贡献可以接近甚至超过细菌的贡献;然而大多数中小型土壤动物的本身代谢过程对碳氮矿化的贡献远低于土壤微生物,但它们可以通过取食作用来调节微生物进而影响碳氮的矿化。大型节肢动物中的蜘蛛和地表甲虫等捕食者经常活跃于地表,它们常常会通过级联效应对土壤生态系统产生重要的影响。蚯蚓、白蚁等大型土壤动物除可以通过取食作用以外,还可以通过非取食作用调控土壤微生物,进而显著影响土壤碳氮过程。土壤动物取食行为的多样性和复杂的非营养关系的存在造就了多维度的土壤食物网,给土壤动物的生态功能研究带来了巨大的挑战。介绍了土壤动物的多样性及主要的生态功能,并对研究的热点和前沿问题进行了探讨,以期引起关于土壤动物多样性及其生态功能的深入思考。     

Soil animal responses to moisture availability are largely scale,not ecosystem dependent: insight from a cross‐site study

Climate change will result in reduced soil water availability in much of the world either due to changes in precipitation or increased temperature and evapotranspiration. How communities of mites and nematodes may respond to changes in moisture availability is not well known, yet these organisms play important roles in decomposition and nutrient cycling processes. We determined how communities of these organisms respond to changes in moisture availability and whether common patterns occur along fine‐scale gradients of soil moisture within four individual ecosystem types (mesic, xeric and arid grasslands and a polar desert) located in the western United States and Antarctica, as well as across a cross‐ecosystem moisture gradient (CEMG) of all four ecosystems considered together. An elevation transect of three sampling plots was monitored within each ecosystem and soil samples were collected from these plots and from existing experimental precipitation manipulations within each ecosystem once in fall of 2009 and three times each in 2010 and 2011. Mites and nematodes were sorted to trophic groups and analyzed to determine community responses to changes in soil moisture availability. We found that while both mites and nematodes increased with available soil moisture across the CEMG, within individual ecosystems, increases in soil moisture resulted in decreases to nematode communities at all but the arid grassland ecosystem; mites showed no responses at any ecosystem. In addition, we found changes in proportional abundances of mite and nematode trophic groups as soil moisture increased within individual ecosystems, which may result in shifts within soil food webs with important consequences for ecosystem functioning. We suggest that communities of soil animals at local scales may respond predictably to changes in moisture availability regardless of ecosystem type but that additional factors, such as climate variability, vegetation composition, and soil properties may influence this relationship over larger scales.     

The influence of nematodes on below-ground processes in grassland ecosystems

This review summarises recent information on beneficial roles that soil nematodes play in the cycling of carbon and other plant nutrients in grassland ecosystems. In particular, we focus on the role of the two dominant functional groups of nematodes, namely the microbial- and root-feeders, and how their activities may enhance soil ecosystem-level processes of nutrient cycling and, ultimately, plant productivity in managed and unmanaged grassland ecosystems. We report recent experiments which show that low amounts of root herbivory by nematodes can increase the allocation of photoassimilate carbon to roots, leading to increased root exudation and microbial activity in the rhizosphere. The effects of these interactions on soil nutrient cycling and plant productivity are discussed. Evidence is presented to show that the feeding activities of microbial-feeding nematodes can enhance nutrient mineralization and plant nutrient uptake in grasslands, but that these responses are highly species-specific and appear to be strongly regulated by higher trophic groups of fauna (top-down regulation). We recommend that future studies of the roles of nematodes in grasslands ecosystems should consider these more complex trophic interactions and also the effects of species diversity of nematodes on soil ecosystem-level processes. This revised version was published online in June 2006 with corrections to the Cover Date.     

Linking soil food web structure to above‐ and belowground ecosystem processes: a meta‐analysis

Soil fauna can be an important regulator of community parameters and ecosystem processes, but there have been few quantitative syntheses of the role of soil fauna in terrestrial soil communities and ecosystems. Here, we conducted a meta‐analysis to investigate the impacts of invertebrate soil micro‐ and mesofauna (grazers and predators) on plant productivity and microbial biomass. Overall our results indicate that an increase in the biomass of soil fauna increased aboveground plant productivity across ecosystems by 35% and decreased microbial biomass by 8%. In addition, we found no evidence for trophic cascades in terrestrial soil food webs, but the bacterivorous component of soil fauna influenced plant productivity and microbial biomass more than did the fungivorous component. Furthermore, changes in the biomass of soil fauna differentially affected plant productivity among plant functional groups: a higher biomass of soil fauna increased aboveground productivity by 70% in coniferous systems. However, in ecosystems dominated by legumes, a functional group with lower inorganic nitrogen requirements, there was no response of aboveground productivity to increases in the biomass of soil fauna. In sum, the results of this meta‐analysis indicate that soil fauna help to regulate ecosystem production, especially in nutrient‐limited ecosystems.     

Trickle-down effects of aboveground trophic cascades on the soil food web

Trophic cascades are increasingly being regarded as important features of aboveground and belowground food webs, but the effects of aboveground cascades on soil food webs, and vice versa, remains essentially unexplored. We conducted an experiment consisting of model synthesised communities containing grassland plant and invertebrate species, in which treatments included soil only, soil+plants, soil+plants+aphids, and soil+plants+aphids+predators; predator treatments consisted of the lacewing Micromus tasmaniae and ladybird beetle Coccinella undecimpunctata added either singly or in combination. Addition of Micromus largely reversed the negative effects of aphids on plant biomass, while both of the predator species caused large changes in the relative abundances of dominant plant species. Predators of aphids also affected several components of the belowground subsystem. Micromus had positive indirect effects on the primary consumer of the soil decomposer food web (microflora), probably through promoting greater input of basal resources to the decomposer subsystem. Predator treatments also influenced densities of the tertiary consumers of the soil food web (top predatory nematodes), most likely through inducing changes in plant community composition and therefore the quality of resource input to the soil. The secondary consumers of the soil food web (microbe‐feeding nematodes) were, however, unresponsive. The fact that some trophic levels of the soil food web but not others responded to aboveground manipulations is explicable in terms of top‐down and bottom‐up forces differentially regulating different belowground trophic levels. Addition of aphids also influenced microbial community structure, promoted soil bacteria at the expense of fungi, and enhanced the diversity of herbivorous nematodes; in all cases these effects were at least partially reversed by addition of Micromus. These results in tandem point to upper level consumers in aboveground food webs as a potential driver of the belowground subsystem, and provide evidence that predator‐induced trophic cascades aboveground can have effects that trickle through soil food webs.