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.     

Global change belowground: impacts of elevated CO2, nitrogen,and summer drought on soil food webs and biodiversity

The world's ecosystems are subjected to various anthropogenic global change agents, such as enrichment of atmospheric CO₂ concentrations, nitrogen (N) deposition, and changes in precipitation regimes. Despite the increasing appreciation that the consequences of impending global change can be better understood if varying agents are studied in concert, there is a paucity of multi‐factor long‐term studies, particularly on belowground processes. Herein, we address this gap by examining the responses of soil food webs and biodiversity to enrichment of CO₂, elevated N, and summer drought in a long‐term grassland study at Cedar Creek, Minnesota, USA (BioCON experiment). We use structural equation modeling (SEM), various abiotic and biotic explanatory variables, and data on soil microorganisms, protozoa, nematodes, and soil microarthropods to identify the impacts of multiple global change effects on drivers belowground. We found that long‐term (13‐year) changes in CO₂ and N availability resulted in modest alterations of soil biotic food webs and biodiversity via several mechanisms, encompassing soil water availability, plant productivity, and – most importantly – changes in rhizodeposition. Four years of manipulation of summer drought exerted surprisingly minor effects, only detrimentally affecting belowground herbivores and ciliate protists at elevated N. Elevated CO₂ increased microbial biomass and the density of ciliates, microarthropod detritivores, and gamasid mites, most likely by fueling soil food webs with labile C. Moreover, beneficial bottom‐up effects of elevated CO₂ compensated for detrimental elevated N effects on soil microarthropod taxa richness. In contrast, nematode taxa richness was lowest at elevated CO₂ and elevated N. Thus, enrichment of atmospheric CO₂ concentrations and N deposition may result in taxonomically and functionally altered, potentially simplified, soil communities. Detrimental effects of N deposition on soil biodiversity underscore recent reports on plant community simplification. This is of particular concern, as soils house a considerable fraction of global biodiversity and ecosystem functions.     

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.     

Nitrogen Addition Regulates Soil Nematode Community Composition through Ammonium Suppression

Nitrogen (N) enrichment resulting from anthropogenic activities has greatly changed the composition and functioning of soil communities. Nematodes are one of the most abundant and diverse groups of soil organisms, and they occupy key trophic positions in the soil detritus food web. Nematodes have therefore been proposed as useful indicators for shifts in soil ecosystem functioning under N enrichment. Here, we monitored temporal dynamics of the soil nematode community using a multi-level N addition experiment in an Inner Mongolia grassland. Measurements were made three years after the start of the experiment. We used structural equation modeling (SEM) to explore the mechanisms regulating nematode responses to N enrichment. Across the N enrichment gradient, significant reductions in total nematode abundance, diversity (H' and taxonomic richness), maturity index (MI), and the abundance of root herbivores, fungivores and omnivores-predators were found in August. Root herbivores recovered in September, contributing to the temporal variation of total nematode abundance across the N gradient. Bacterivores showed a hump-shaped relationship with N addition rate, both in August and September. Ammonium concentration was negatively correlated with the abundance of total and herbivorous nematodes in August, but not in September. Ammonium suppression explained 61% of the variation in nematode richness and 43% of the variation in nematode trophic group composition. Ammonium toxicity may occur when herbivorous nematodes feed on root fluid, providing a possible explanation for the negative relationship between herbivorous nematodes and ammonium concentration in August. We found a significantly positive relationship between fungivores and fungal phospholipid fatty acids (PLFA), suggesting bottom-up control of fungivores. No such relationship was found between bacterivorous nematodes and bacterial PLFA. Our findings contribute to the understanding of effects of N enrichment in semiarid grassland on soil nematode trophic groups, and the cascading effects in the detrital soil food web.     

Consumer-resource stoichiometry in detritus-based streams

Stoichiometric relationships between consumers and resources in detritus‐based ecosystems have received little attention, despite the importance of detritus in most food webs. We analysed carbon (C), nitrogen (N), and phosphorus (P) content of invertebrate consumers, and basal food resources in two forested headwater streams (one reference and the other nutrient‐enriched). We found large elemental imbalances between consumers and food resources compared with living plant‐based systems, particularly in regard to P content, which were reduced with enrichment. Enrichment significantly increased nutrient content of food resources (consistent with uptake of N and P by detritus‐associated microbes). P content of some invertebrates also increased in the enriched vs. reference stream, suggesting deviation from strict homeostasis. Nutrient content varied significantly among invertebrate functional feeding groups, orders and, to some extent, size classes. Future application of stoichiometric theory to detritus‐based systems should consider the potential for relatively large consumer‐resource elemental imbalances and P storage by insect consumers.     

Convergent responses of nitrogen and phosphorus resorption to nitrogen inputs in a semiarid grassland

Human activities have significantly altered nitrogen (N) availability in most terrestrial ecosystems, with consequences for community composition and ecosystem functioning. Although studies of how changes in N availability affect biodiversity and community composition are relatively common, much less remains known about the effects of N inputs on the coupled biogeochemical cycling of N and phosphorus (P), and still fewer data exist regarding how increased N inputs affect the internal cycling of these two elements in plants. Nutrient resorption is an important driver of plant nutrient economies and of the quality of litter plants produce. Accordingly, resorption patterns have marked ecological implications for plant population and community fitness, as well as for ecosystem nutrient cycling. In a semiarid grassland in northern China, we studied the effects of a wide range of N inputs on foliar nutrient resorption of two dominant grasses, Leymus chinensis and Stipa grandis. After 4 years of treatments, N and P availability in soil and N and P concentrations in green and senesced grass leaves increased with increasing rates of N addition. Foliar N and P resorption significantly decreased along the N addition gradient, implying a resorption‐mediated, positive plant–soil feedback induced by N inputs. Furthermore, N : P resorption ratios were negatively correlated with the rates of N addition, indicating the sensitivity of plant N and P stoichiometry to N inputs. Taken together, the results demonstrate that N additions accelerate ecosystem uptake and turnover of both N and P in the temperate steppe and that N and P cycles are coupled in dynamic ways. The convergence of N and P resorption in response to N inputs emphasizes the importance of nutrient resorption as a pathway by which plants and ecosystems adjust in the face of increasing N availability.     

Interplay between the paradox of enrichment and nutrient cycling in food webs

Nutrient cycling is fundamental to ecosystem functioning. Despite recent major advances in the understanding of complex food web dynamics, food web models have so far generally ignored nutrient cycling. However, nutrient cycling is expected to strongly impact food web stability and functioning. To make up for this gap, we built an allometric and size structured food web model including nutrient cycling. By releasing mineral nutrients, recycling increases the availability of limiting resources for primary producers and links each trophic level to the bottom of food webs. We found that nutrient cycling can provide a significant part of the total nutrient supply of the food web, leading to a strong enrichment effect that promotes species persistence in nutrient poor ecosystems but leads to a paradox of enrichment at high nutrient inputs. The presence of recycling loops linking each trophic level to the basal resources weakly affects species biomass temporal variability in the food web. Recycling loops tend to slightly dampen the destabilising effect of nutrient enrichment on consumer temporal variability while they have opposite effects for primary producers. By considering nutrient cycling, this new model improves our understanding of the response of food webs to nutrient availability and opens perspectives to better link studies on food web dynamics and ecosystem functioning.     

松嫩盐碱草地几种植物的土壤营养位分析

针对松嫩盐碱草地上的几种主要植物,以植物地上生物量和总和优势度（SDR）为土壤营养位效能指标,分别从多项土壤因子综合、可溶盐含量、总碱度、Na^ 含量、有机质含量、全氮含量角度进行了土壤营养位分析。讨论了松嫩盐碱草地植物种群分布格局与土壤营养的关系,指出松嫩盐碱草地植物种群分布格局的形成是可溶盐含量、总碱度、Na^ 含量、有机质含量、全氮含量5项主要土壤因子综合作用的结果,对于各植物种群土壤营养位的分化结果而言,没有一项土壤因子是起决定性作用的。     

Aboveground mammal and invertebrate exclusions cause consistent changes in soil food webs of two subalpine grassland types,but mechanisms are system‐specific

Ungulates, smaller mammals, and invertebrates can each affect soil biota through their influence on vegetation and soil characteristics. However, direct and indirect effects of the aboveground biota on soil food webs remain to be unraveled. We assessed effects of progressively excluding aboveground large‐, medium‐ and small‐sized mammals as well as invertebrates on soil nematode diversity and feeding type abundances in two subalpine grassland types: short‐ and tall‐grass vegetation. We explored pathways that link exclusions of aboveground biota to nematode feeding type abundances via changes in plants, soil environment, soil microbial biomass, and soil nutrients. In both vegetation types, exclusions caused a similar shift toward higher abundance of all nematode feeding types, except plant feeders, lower Shannon diversity, and lower evenness. These effects were strongest when small mammals, or both small mammals and invertebrates were excluded in addition to excluding larger mammals. Exclusions resulted in a changed abiotic soil environment that only affected nematodes in the short‐grass vegetation. In each vegetation type, exclusion effects on nematode abundances were mediated by different drivers related to plant quantity and quality. In the short‐grass vegetation, not all exclusion effects on omni–carnivorous nematodes were mediated by the abundance of lower trophic level nematodes, suggesting that omni–carnivores also depended on other prey than nematodes. We conclude that small aboveground herbivores have major impacts on the soil food web of subalpine short‐ and tall‐grass ecosystems. Excluding aboveground animals caused similar shifts in soil nematode assemblages in both subalpine vegetation types, however, mechanisms turned out to be system‐specific.     

氮磷输入对过度放牧退化草原土壤线虫群落的影响

线虫是陆地生态系统中数量最多的一类无脊椎后生动物,在土壤碎屑食物网中占据多个营养级,并在能量流动和养分循环中发挥重要的生态功能.土壤线虫的群落结构和多样性水平可以反映生态系统功能的变化.多年来,过度放牧等不合理的开发利用导致我国大面积的草原处在不同程度的退化之中,外源性养分输入是维持其养分平衡进而恢复其生态和生产功能的重要措施.本研究以内蒙古锡林郭勒退化草原为对象,就氮磷输入对土壤线虫丰度和群落结构的单独效应和交互效应进行了研究.本研究获得线虫38属,其中垫咽属、滑刃属、索努斯属和盾垫属是该退化草原线虫群落优势属.氮输入降低了土壤线虫总丰度、捕食杂食线虫丰度以及植物寄生线虫丰度,而磷输入提高了土壤线虫总丰度、食真菌线虫丰度、捕食杂食类线虫丰度以及植物寄生线虫丰度,氮输入抑制了磷输入对线虫总丰度、捕食杂食线虫和植物寄生线虫丰度的促进作用.氮磷输入对线虫多样性无影响,可能与不同养分输入下稳定的植物群落多样性有关.氮输入显著提升了退化草原线虫成熟度指数,降低了植物寄生线虫成熟度指数(PPI),并且能缓解磷输入对PPI和瓦斯乐斯卡指数的负面作用,表明氮输入提高退化草地土壤健康程度,促进线虫群落和食物网的稳定性.上述研究结果有助于从土壤生物学角度理解氮磷输入对退化草地恢复的影响机理.     

Soil acidity, ecological stoichiometry and allometric scaling in grassland food webs

The factors regulating the structure of food webs are a central focus of community and ecosystem ecology, as trophic interactions among species have important impacts on nutrient storage and cycling in many ecosystems. For soil invertebrates in grassland ecosystems in the Netherlands, the site-specific slopes of the faunal biomass to organism body mass relationships reflected basic biochemical and biogeochemical processes associated with soil acidity and soil C : N : P stoichiometry. That is, the higher the phosphorus availability in the soil, the higher, on average, the slope of the faunal biomass size spectrum (i.e., the higher the biomass of large-bodied invertebrates relative to the biomass of small invertebrates). While other factors may also be involved, these results are consistent with the growth rate hypothesis from biological stoichiometry that relates phosphorus demands to ribosomal RNA and protein production. Thus our data represent the first time that ecosystem phosphorus availability has been associated with allometry in soil food webs (supporting information available online). Our results have broad implications, as soil invertebrates of different size have different effects on soil processes.     

Divergence of feeding channels within the soil food web determined by ecosystem type

Understanding trophic linkages within the soil food web (SFW) is hampered by its opacity, diversity, and limited niche adaptation. We need to expand our insight between the feeding guilds of fauna and not just count biodiversity. The soil fauna drive nutrient cycling and play a pivotal, but little understood role within both the carbon (C) and nitrogen (N) cycles that may be ecosystem dependent. Here, we define the structure of the SFW in two habitats (grassland and woodland) on the same soil type and test the hypothesis that land management would alter the SFW in these habitats. To do this, we census the community structure and use stable isotope analysis to establish the pathway of C and N through each trophic level within the ecosystems. Stable isotope ratios of C and N from all invertebrates were used as a proxy for trophic niche, and community‐wide metrics were obtained. Our empirically derived C/N ratios differed from those previously reported, diverging from model predictions of global C and N cycling, which was unexpected. An assessment of the relative response of the different functional groups to the change from agricultural grassland to woodland was performed. This showed that abundance of herbivores, microbivores, and micropredators were stimulated, while omnivores and macropredators were inhibited in the grassland. Differences between stable isotope ratios and community‐wide metrics, highlighted habitats with similar taxa had different SFWs, using different basal resources, either driven by root or litter derived resources. Overall, we conclude that plant type can act as a top‐down driver of community functioning and that differing land management can impact on the whole SFW.     

Asymmetric effects of grazing intensity on macroelements and microelements in grassland soil and plants in Inner Mongolia Grazing alters nutrient dynamics of grasslands

Grazing is a traditional grassland management technique and greatly alters ecosystem nutrient cycling. The effects of grazing intensity on the nutrient dynamics of soil and plants in grassland ecosystems remain uncertain, especially among microelements. A 2‐year field grazing experiment was conducted in a typical grassland with four grazing intensities (ungrazed control, light, moderate, and heavy grazing) in Inner Mongolia, China. Nutrient concentration was assessed in soil and three dominant plant species (Stipa krylovii, Leymus chinensis, and Cleistogenes squarrosa). Assessed quantities included four macroelements (carbon (C), nitrogen (N), phosphorus (P), and magnesium (Mg)) and four microelements (copper (Cu), iron (Fe), manganese (Mn), and zinc (Zn)). Soil total C, total N, total P, available N, and available P concentrations significantly increased with grazing intensity but soil Mg, Cu, Fe, Mn, Zn concentrations had no significant response. Plant C concentration decreased but plant N, P, Mg, Cu, Fe, Mn, and Zn concentrations significantly increased with grazing intensity. In soil, macroelement dynamics (i.e., C, N, and P) exhibited higher sensitivity with grazing intensity, conversely in plants, microelements were more sensitive. This result indicates macroelements and microelements in soil and plants had asymmetric responses with grazing intensity. The slopes of nutrient linear regression in C. squarrosa were higher than that of S. krylovii and L. chinensis, indicating that C. squarrosa had higher nutrient acquisition capacity and responded more rapidly to heavy grazing. These findings indicate that short‐term heavy grazing accelerates nutrient cycling of the soil–plant system in grassland ecosystems, elucidate the multiple nutrient dynamics of soil and plants with grazing intensity, and emphasize the important function of microelements in plant adaptation in grazing management.     

Trophic level and basal resource use of soil animals are hardly affected by local plant associations in abandoned arable land

Plants provide resources and shape the habitat of soil organisms thereby affecting the composition and functioning of soil communities. Effects of plants on soil communities are largely taxon‐dependent, but how different functional groups of herbaceous plants affect trophic niches of individual animal species in soil needs further investigation. Here, we studied the use of basal resources and trophic levels of dominating soil meso‐ and macrofauna using stable isotope ratios of carbon and nitrogen in arable fallow systems 3 and 14–16 years after abandonment. Animals were sampled from the rhizosphere of three plant species of different functional groups: a legume (Medicaco sativa), a nonlegume herb (Taraxacum officinale), and a grass (Bromus sterilis). We found virtually no consistent effects of plant identity on stable isotope composition of soil animals and on thirteen isotopic metrics that reflect general food‐web structure. However, in old fallows, the carbon isotope composition of some predatory macrofauna taxa had shifted closer to that of co‐occurring plants, which was particularly evident for Lasius, an aphid‐associated ant genus. Trophic levels and trophic‐chain lengths in food webs were similar across plant species and fallow ages. Overall, the results suggest that variations in local plant diversity of grassland communities may little affect the basal resources and the trophic level of prey consumed by individual species of meso‐ and macrofauna belowground. By contrast, successional changes in grassland communities are associated with shifts in the trophic niches of certain species, reflecting establishment of trophic interactions with time, which shapes the functioning and stability of soil food webs.     

Changes in plant species richness induce functional shifts in soil nematode communities in experimental grassland

Background

Changes in plant diversity may induce distinct changes in soil food web structure and accompanying soil feedbacks to plants. However, knowledge of the long-term consequences of plant community simplification for soil animal food webs and functioning is scarce. Nematodes, the most abundant and diverse soil Metazoa, represent the complexity of soil food webs as they comprise all major trophic groups and allow calculation of a number of functional indices.

Methodology/Principal Findings

We studied the functional composition of nematode communities three and five years after establishment of a grassland plant diversity experiment (Jena Experiment). In response to plant community simplification common nematode species disappeared and pronounced functional shifts in community structure occurred. The relevance of the fungal energy channel was higher in spring 2007 than in autumn 2005, particularly in species-rich plant assemblages. This resulted in a significant positive relationship between plant species richness and the ratio of fungal-to-bacterial feeders. Moreover, the density of predators increased significantly with plant diversity after five years, pointing to increased soil food web complexity in species-rich plant assemblages. Remarkably, in complex plant communities the nematode community shifted in favour of microbivores and predators, thereby reducing the relative abundance of plant feeders after five years.

Conclusions/Significance

The results suggest that species-poor plant assemblages may suffer from nematode communities detrimental to plants, whereas species-rich plant assemblages support a higher proportion of microbivorous nematodes stimulating nutrient cycling and hence plant performance; i.e. effects of nematodes on plants may switch from negative to positive. Overall, food web complexity is likely to decrease in response to plant community simplification and results of this study suggest that this results mainly from the loss of common species which likely alter plant – nematode interactions.     

Multi‐nutrient vs. nitrogen‐only effects on carbon sequestration in grassland soils

Human activities have greatly increased the availability of biologically active forms of nutrients [e.g., nitrogen (N), phosphorous (P), potassium (K), magnesium (Mg)] in many soil ecosystems worldwide. Multi‐nutrient fertilization strongly increases plant productivity but may also alter the storage of carbon (C) in soil, which represents the largest terrestrial pool of organic C. Despite this issue is important from a global change perspective, key questions remain on how the single addition of N or the combination of N with other nutrients might affect C sequestration in human‐managed soils. Here, we use a 19‐year old nutrient addition experiment on a permanent grassland to test for nutrient‐induced effects on soil C sequestration. We show that combined NPKMg additions to permanent grassland have ‘constrained’ soil C sequestration to levels similar to unfertilized plots whereas the single addition of N significantly enhanced soil C stocks (N‐only fertilized soils store, on average, 11 t C ha^?1 more than unfertilized soils). These results were consistent across grazing and liming treatments suggesting that whilst multi‐nutrient additions increase plant productivity, soil C sequestration is increased by N‐only additions. The positive N‐only effect on soil C content was not related to changes in plant species diversity or to the functional composition of the plant community. N‐only fertilized grasslands show, however, increases in total root mass and the accumulation of organic matter detritus in topsoils. Finally, soils receiving any N addition (N only or N in combination with other nutrients) were associated with high N losses. Overall, our results demonstrate that nutrient fertilization remains an important global change driver of ecosystem functioning, which can strongly affect the long‐term sustainability of grassland soil ecosystems (e.g., soils ability to deliver multiple ecosystem services).     

Differential responses of size‐based functional groups to bottom–up and top–down perturbations in pelagic food webs: a meta‐analysis

We performed a meta‐analysis of 31 lake mesocosm experiments to investigate differences in the responses of pelagic food chains and food webs to nutrient enrichment and fish presence. Trophic levels were divided into size‐based functional groups (phytoplankton into highly edible and poorly edible algae, and zooplankton into small herbivores, large herbivores and omnivorous zooplankton) in the food webs. Our meta‐analysis shows that 1) nutrient enrichment has a positive effect on phytoplankton and zooplankton, while fish presence has a positive effect on phytoplankton and a negative effect on zooplankton in the food chains; 2) nutrient enrichment has a positive effect on highly edible algae and small herbivores, but no effect on poorly edible algae, large herbivores and omnivorous zooplankton in the food webs. Planktivorous fish have a positive effect on highly edible algae and small herbivores, a negative effect on large herbivores and omnivorous zooplankton, and no effect on poorly edible algae. Our meta‐analysis confirms that nutrient enrichment and planktivorous fish affect functional groups differentially within trophic levels, revealing important changes in the functioning of food webs. The analysis of fish effects shows the well‐described trophic cascade in the food chain and reveals two trophic cascades in the food web: one transmitted by large herbivores that benefit highly edible phytoplankton, and one transmitted by omnivorous zooplankton that benefit small herbivores. Comparison between the responses of food webs and simple food chains also shows consistent biomass compensation between functional groups within trophic levels.     

Most soil trophic guilds increase plant growth: a meta‐analytical review

Trophic cascades are important drivers of plant and animal abundances in aquatic and aboveground systems, but in soils trophic cascades have been thought to be of limited importance due to omnivory and other factors. Here we use a meta‐analysis of 215 studies with 1526 experiments that measured plant growth responses to additions or removals of soil organisms to test how different soil trophic levels affect plant growth. Consistent with the trophic cascade hypothesis, we found that herbivores and plant pathogens (henceforth pests) decreased plant growth and that predators of pests increased plant growth. The magnitude of this trophic cascade was similar to that reported for aboveground systems. In contrast, we did not find evidence for trophic cascades in decomposer‐ and symbiont‐based (henceforth mutualist) food chains. In these food chains, mutualists increased plant growth and predators of mutualists also increased plant growth, presumably by increasing nutrient cycling rates. Therefore, mutualists, predators of mutualists and predators of pests all increased plant growth. Further, experiments that added multiple organisms from different trophic levels also increased plant growth. As a result, across the dataset, soil organisms increased plant growth 29% and non‐pest soil organisms increased plant growth 46%. Omnivory has traditionally been thought to confound soil trophic dynamics, but here we suggest that omnivory allows for a simplified perspective of soil food webs – one in which most soil organisms increase plant growth by preying on pests or increasing nutrient cycling rates. An implication of this perspective is that processes that decrease soil organism abundance (e.g. soil tillage) are likely to decrease aboveground productivity. Synthesis Soil foodwebs have resisted generalizations due to their diversity and interconnectedness. Here we use results from a meta‐analysis to inform a simplified perspective of soil foodwebs: one in which most soil trophic guilds increase plant growth. Our review also includes the first widespread support for the presence of trophic cascades in soils.     

大气氮沉降影响草地植物物种多样性机制研究综述

大气氮沉降对草地生态系统结构和功能的影响已成为全球变化生物学研究重点。大气氮沉降导致草地群落物种多样性降低已成为全球普遍现象,但其生物学机制还不清楚,因此有必要系统梳理大气氮沉降对全球不同草地生态系统的研究结果,以便在氮沉降背景下为我国草地生态系统的研究和管理制定科学决策。系统综述了氮沉降降低草地群落物种多样性的可能机制,主要包括资源竞争排斥、群落更新限制、土壤酸化及其离子毒害、养分失衡、氮素本身的毒害、次生胁迫。氮沉降导致草地物种多样性降低是多种机制综合作用的结果,每种机制在不同时空具有不同的相对贡献。同时,与欧洲酸性土壤草地和美国高草草原相比,我国草地土壤类型和植被属性具有明显差异。因此,应根据我国草地生态系统的特征、不同植物功能利用养分策略,从土壤养分变化、根系养分吸收转运、叶片生理过程等方面的整合研究思路,探讨氮沉降影响我国草地群落物种多样性的生物学机制,为我国草地生态系统的科学管理提供理论依据。     

Nitrogen deposition weakens plant–microbe interactions in grassland ecosystems

Soil carbon (C) and nitrogen (N) stoichiometry is a main driver of ecosystem functioning. Global N enrichment has greatly changed soil C : N ratios, but how altered resource stoichiometry influences the complexity of direct and indirect interactions among plants, soils, and microbial communities has rarely been explored. Here, we investigated the responses of the plant‐soil‐microbe system to multi‐level N additions and the role of dissolved organic carbon (DOC) and inorganic N stoichiometry in regulating microbial biomass in semiarid grassland in northern China. We documented a significant positive correlation between DOC and inorganic N across the N addition gradient, which contradicts the negative nonlinear correlation between nitrate accrual and DOC availability commonly observed in natural ecosystems. Using hierarchical structural equation modeling, we found that soil acidification resulting from N addition, rather than changes in the plant community, was most closely related to shifts in soil microbial community composition and decline of microbial respiration. These findings indicate a down‐regulating effect of high N availability on plant–microbe interactions. That is, with the limiting factor for microbial biomass shifting from resource stoichiometry to soil acidity, N enrichment weakens the bottom‐up control of soil microorganisms by plant‐derived C sources. These results highlight the importance of integratively studying the plant‐soil‐microbe system in improving our understanding of ecosystem functioning under conditions of global N enrichment.