Large‐scale biogeography and environmental regulation of methanotrophic bacteria across boreal inland waters

Aerobic methanotrophic bacteria (methanotrophs) use methane as a source of carbon and energy, thereby mitigating net methane emissions from natural sources. Methanotrophs represent a widespread and phylogenetically complex guild, yet the biogeography of this functional group and the factors that explain the taxonomic structure of the methanotrophic assemblage are still poorly understood. Here, we used high‐throughput sequencing of the 16S rRNA gene of the bacterial community to study the methanotrophic community composition and the environmental factors that influence their distribution and relative abundance in a wide range of freshwater habitats, including lakes, streams and rivers across the boreal landscape. Within one region, soil and soil water samples were additionally taken from the surrounding watersheds in order to cover the full terrestrial–aquatic continuum. The composition of methanotrophic communities across the boreal landscape showed only a modest degree of regional differentiation but a strong structuring along the hydrologic continuum from soil to lake communities, regardless of regions. This pattern along the hydrologic continuum was mostly explained by a clear niche differentiation between type I and type II methanotrophs along environmental gradients in pH, and methane concentrations. Our results suggest very different roles of type I and type II methanotrophs within inland waters, the latter likely having a terrestrial source and reflecting passive transport and dilution along the aquatic networks, but this is an unresolved issue that requires further investigation.     

Landscape‐scale spatial abundance distributions discriminate core from random components of boreal lake bacterioplankton

Aquatic bacterial communities harbour thousands of coexisting taxa. To meet the challenge of discriminating between a ‘core’ and a sporadically occurring ‘random’ component of these communities, we explored the spatial abundance distribution of individual bacterioplankton taxa across 198 boreal lakes and their associated fluvial networks (188 rivers). We found that all taxa could be grouped into four distinct categories based on model statistical distributions (normal like, bimodal, logistic and lognormal). The distribution patterns across lakes and their associated river networks showed that lake communities are composed of a core of taxa whose distribution appears to be linked to in‐lake environmental sorting (normal‐like and bimodal categories), and a large fraction of mostly rare bacteria (94% of all taxa) whose presence appears to be largely random and linked to downstream transport in aquatic networks (logistic and lognormal categories). These rare taxa are thus likely to reflect species sorting at upstream locations, providing a perspective of the conditions prevailing in entire aquatic networks rather than only in lakes.     

典型草原区不同生境反硝化菌群的空间特征

【背景】锡林河-河滨湿地-阶地草原是蒙古高原典型草原区代表性的水生-湿生-陆生生境,但不同生境中反硝化菌群的空间分布特征尚不明晰。【目的】阐明典型草原区不同生境反硝化菌群的组成、丰度、空间分布特征及异质性成因。【方法】利用16S rRNA基因测序研究锡林河流域水生、湿生、陆生生境6个样带沉积物/土壤细菌群落组成及相对丰度。基于2014年及以前文献报道的反硝化细菌及16S rRNA基因信息构建参比菌库,筛选生境关联的反硝化菌属。通过典范对应分析等探究反硝化菌群空间异质性成因。【结果】参比菌库包含80种反硝化细菌(65个属),6个样带测序获得的469个细菌属中36个为反硝化细菌属。3种生境共存的反硝化细菌有14个属,其中黄杆菌属(1.65%-14.17%)和噬氢菌属(1.56%-1.69%)是水生和湿生生境共有的优势菌,假单胞菌属(1.85%)是低河漫滩样带的优势菌。空间分布特征显示反硝化菌群沿水生-湿生-陆生生境呈现先升后降的分布趋势,在低河漫滩湿地达到最高值。典范对应分析表明:黄杆菌属、噬氢菌属、气单胞菌属、鞘氨醇单胞菌属等与pH值、水分及沙粒含量呈正相关关系,而芽孢杆菌属、链霉菌属、马杜拉放线菌属等与粘粒、粉粒、有机质、总氮含量等呈正相关关系。【结论】典型草原区反硝化菌群组成及丰度具有明显的生境异质性,低河漫滩湿地是反硝化细菌生长繁殖的最佳生境,由颗粒组成、水分含量和pH等环境因子共同驱动。     

The bacterial biogeography of British soils

Despite recognition of the importance of soil bacteria to terrestrial ecosystem functioning there is little consensus on the factors regulating belowground biodiversity. Here we present a multi-scale spatial assessment of soil bacterial community profiles across Great Britain (> 1000 soil cores), and show the first landscape scale map of bacterial distributions across a nation. Bacterial diversity and community dissimilarities, assessed using terminal restriction fragment length polymorphism, were most strongly related to soil pH providing a large-scale confirmation of the role of pH in structuring bacterial taxa. However, while α diversity was positively related to pH, the converse was true for β diversity (between sample variance in α diversity). β diversity was found to be greatest in acidic soils, corresponding with greater environmental heterogeneity. Analyses of clone libraries revealed the pH effects were predominantly manifest at the level of broad bacterial taxonomic groups, with acidic soils being dominated by few taxa (notably the group 1 Acidobacteria and Alphaproteobacteria). We also noted significant correlations between bacterial communities and most other measured environmental variables (soil chemistry, aboveground features and climatic variables), together with significant spatial correlations at close distances. In particular, bacterial and plant communities were closely related signifying no strong evidence that soil bacteria are driven by different ecological processes to those governing higher organisms. We conclude that broad scale surveys are useful in identifying distinct soil biomes comprising reproducible communities of dominant taxa. Together these results provide a baseline ecological framework with which to pursue future research on both soil microbial function, and more explicit biome based assessments of the local ecological drivers of bacterial biodiversity.     

Structure and dynamics of an amphibian metacommunity in two regions

1. The concept of metacommunity is based on the hypothesis that species occurrence depends on species dynamics and interactions on local and regional scales via the movements of individuals between localities. Metacommunity approaches are currently being applied to pond breeding taxa such as amphibians. 2. Given that animal movement is also influenced by the physical quality of the matrix to be crossed to reach a breeding habitat and by the affinity of the species for specific terrestrial habitats, matrix characteristics may enhance or hinder dispersal success. These characteristics would, in turn, affect the composition of larval assemblages at local level and, consequently, determine metacommunity structure and dynamics. 3. Here we compared the structures and dynamics of two metacommunities with the same pool of anurans along similar freshwater gradients in two regions that are well differentiated in terms of their respective terrestrial matrix. 4. Abundance of tadpole species and species assemblage in the two regions were determined principally by local processes (at pond level); however, the structure and dynamics of the communities differed. In one region species abundance was explained in part by landscape factors and consequently showed lower co-occurrence and lower colonization rates (species sorting models) indicating that terrestrial habitat could restrict animal movements, whereas in the other region higher co-occurrence and higher colonization rates (mass effect models) indicated low dispersal limitations.     

The relative importance of spatial and environmental processes in distribution of benthic chironomid larvae within a large and shallow lake

Although chironomids are popular model organisms in ecological research and indicators of bioassessment, the relative role of dispersal and environmental filtering in their community assembly is still poorly known, especially at fine spatial scales. In this study, we applied a metacommunity framework and used various statistical tools to examine the relative role of spatial and local environmental factors in distribution of benthic chironomid taxa and their assemblages in large and shallow Lake Balaton, Hungary. Contrary to present predictions on the metacommunity organisation of aquatic insects with winged terrestrial adults, we found that dispersal limitation can considerably affect distribution of chironomids even at lake scale. However, we also revealed the predominant influence of environmental filtering, and strong taxa–environment relationships were observed especially along sediment type, sediment organic matter content and macrophyte coverage gradients. We account that identified reference conditions and assemblages along with specified optima and tolerances of the abundant taxa can contribute to our understanding of chironomid ecology and be utilised in shallow lake bioassessment. Further, we propose that predictive models of species–environment relationships should better take into account pure spatial structuring of local communities and species-specific variability of spatial processes and environmental control even at small spatial scales.     

Contrasting metacommunity structure and beta diversity in an aquatic‐floodplain system

Habitat connectivity and dispersal interact to structure metacommunities, but few studies have examined these patterns jointly for organisms across the aquatic–terrestrial ecotone. We assessed metacommunity structure and beta diversity patterns of instream benthic invertebrates, riparian carabid beetles (Order: Coleoptera; Family: Carabidae) and riparian spiders (Order: Araneae) at fifteen sites in a river‐floodplain system. Sampling took place over a three‐year period (2010–2012) in the Rhine‐Main‐Observatory LTER site on the Kinzig River, central Germany. This allowed disentangling the combined influence, and temporal variability, of habitat connectivity (i.e. between aquatic and terrestrial) and dispersal ability (i.e. between spiders and beetles, and aerial and aquatic dispersing invertebrates) on the dominant paradigms structuring these metacommunities. We found mostly consistent differences in the manner that metacommunities were structured between groups, with lower levels of variability explained for beetles compared to the other groups. Beetles were consistently structured more by turnover than nestedness components, with greater beta diversity than expected by chance and a minor spatial compared to environmental signal emerging with variance partitioning. Conversely, spiders and benthic invertebrates had lower beta diversity and greater nestedness than null expectation, and a clearer spatial signal controlling metacommunity structure. Our results suggest varying levels of mass effects and species sorting shape river‐floodplain metacommunities, depending on habitat connectivity and dispersal ability. That is, greater connectivity and lower fragmentation along the river compared to the terrestrial zone promoted mass effects, and differences in overall dispersal ability and mode (i.e. active and passive) for instream and riparian communities shifted paradigms between mass effects and species sorting.     

Soil Properties and Spatial Processes Influence Bacterial Metacommunities within a Grassland Restoration Experiment

Metacommunity theory proposes that a collection of local communities are linked by dispersal and the resulting compositions are a product of both niche‐based (species sorting) and spatial processes. Determining which of these factors is most important in different habitats can provide insight into the regulation of community assembly. To date, the metacommunity organization of heterotrophic soil bacteria is largely unknown. Spatial variation of soil bacterial communities could arise from (1) the resource heterogeneity produced by plant communities through root exudation and/or litter inputs; (2) the heterogeneity of soil environmental properties; and (3) pure spatial processes, including dispersal limitation and stochastic assembly. Understanding the relative importance of these factors for soil bacterial community structure and function could increase our ability to restore soil communities. We utilized an ongoing tallgrass prairie restoration experiment in northeastern Kansas to assess if restoring native plant communities produced changes in bacterial communities 6 years after restoration. We further examined the relative importance of the spatial heterogeneity of plant communities, soil properties, and pure spatial effects for bacterial community structure in the old‐field restoration site. We found that soil bacterial communities were not influenced by plant restoration, but rather, by the local heterogeneity of soil environmental properties (16.9% of bacterial community variation) and pure spatial effects (11.1%). This work also stresses the idea that restoring bacterial communities can take many years to accomplish due to the inherent changes that occur to the soil after cultivation and the time it takes for the re‐establishment of soil quality.     

Metacommunity size influences aquatic community composition in a natural mesocosm landscape

Ecosystems are often arranged in naturally patchy landscapes with habitat patches linked by dispersal of species in a metacommunity. The size of a metacommunity, or number of patches, is predicted to influence community dynamics and therefore the structure and function of local communities. However, such predictions have yet to be experimentally tested using full food webs in natural metacommunities. We used the natural mesocosm system of aquatic macroinvertebrates in bromeliad phytotelmata to test the effect of the number of patches in a metacommunity on species richness, abundance, and community composition. We created metacommunities of varying size using fine mesh cages to enclose a gradient from a single bromeliad up to the full forest. We found that species richness, abundance, and biomass increased from enclosed metacommunities to the full forest size and that diversity and evenness also increased in larger enclosures. Community composition was affected by metacommunity size across the full gradient, with a more even detritivore community in larger metacommunities, and taxonomic groups such as mosquitoes going locally extinct in smaller metacommunities. We were able to divide the effects of metacommunity size into aquatic and terrestrial habitat components and found that the importance of each varied by species; those with simple life cycles were only affected by local aquatic habitat whereas insects with complex life cycles were also affected by the amount of terrestrial matrix. This differential survival of obligate and non‐obligate dispersers allowed us to partition the beta‐diversity between metacommunities among functional groups. Our study is one of the first tests of metacommunity size in a natural metacommunity landscape and shows that both diversity and community composition are significantly affected by metacommunity size. Synthesis Natural food webs are sensitive to meta‐community size, i.e. the number of patches connected through dispersal. We provide an empirical test using the aquatic foodweb associated within bromeliads as a model system. When we reduced the number of bromeliad patches connect through dispersal, we found a clear change of the foodweb in terms of population sizes, beta diversity, community composition and predator‐prey ratios. The response of individual taxa was predictable based on species traits including dispersal modes, life cycle, and adult resource requirements. Our study demonstrates that community structure is strongly influenced by the interplay of species traits and landscape properties.     

Regional community assembly drivers and microbial environmental sources shaping bacterioplankton in an alpine lacustrine district (Pyrenees,Spain)

Microbial communities in natural ecosystems are subject to strong ecological rules. The study of local communities along a regional metacommunity can reveal patterns of community assembly, and disentangle the underlying ecological processes. In particular, we seek drivers of community assembly at the regional scale using a large lacustrine dataset (>300 lakes) along the geographical, limnological and physico-chemical gradients in the Pyrenees. By using high throughput amplicon sequencing of the 16S rRNA gene, and inferring environmental sources of bacterial immigrants, we showed that surface aquatic bacterial assemblages were strongly influenced by terrestrial populations from soil, biofilms or sediments, and primarily selected by a pH-alkalinity gradient. Indeed, source proportions explained 27% of the community variation, and chemistry 15% of the total variation, half of it shared with the sources. Major taxonomic groups such as Verrucomicrobia, Actinobacteria and Bacteroidetes showed higher aquatic affinities than Parcubacteria, Gammaproteobacteria, Alphaproteobacteria or Betaproteobacteria, which may be recruited and selected through different hydrographic habitats. A regional fingerprint was observed with lower alpha diversity and higher beta diversity in the central Pyrenees than in both ends. We suggest an ecological succession process, likely influenced by complex interactions of environmental source dispersal and environmental filtering along the mountain range geography.     

Species-sorting may explain an apparent minimal effect of immigration on freshwater bacterial community dynamics

Long distance atmospheric transport of bacterial cells is often implied as a driver of the apparent cosmopolitan distribution of bacterial taxa. Surprisingly, efforts to measure immigration in bacterial communities are rare. An 8-week time series of within-lake bacterial community composition and atmospheric deposition rates and composition were used to estimate the influence of immigration on bacterial community dynamics in two north temperate lakes. Characterization of bacterial community dynamics using automated ribosomal intergenic spacer analysis suggested moderate overlap in composition between the lakes and atmospherically deposited cells. However, taxa that appeared to be delivered by atmospheric deposition had a relatively minor influence on lake bacterial community dynamics. The weak influence of immigrating bacterial taxa suggests that a species-sorting concept best describes aquatic bacterial metacommunity dynamics.     

Metacommunity organisation,spatial extent and dispersal in aquatic systems: patterns,processes and prospects

相似文献   

The role of metacommunity processes in shaping invertebrate rock pool communities along a dispersal gradient

Explaining the variance of local communities in a spatial‐environmental matrix is one of the core interests of ecology today. Recent progress in metacommunity theory has made a substantial contribution to this field, however good empirical data in support of available theories are still relatively scarce. In this study we sampled a cluster of 36 temporary rock pools four times during one season to assess invertebrate metacommunity structure and dynamics and to search for steering processes and variables. Both Mantel tests and redundancy models indicate that local abiotic factors were dominant over spatial factors in explaining community structure and both were acting independently. Spatial variables were only important for passive dispersers and significantly explained 11% of variation in this community component. Pools connected by temporary overflows hosted more similar communities of passive dispersers than unconnected ones while community dissimilarity significantly increased with inter‐pool distance. A negative curvilinear relation was discovered between taxon richness and isolation in passive dispersers, providing some support for existing theoretical models of Mouquet and Loreau. Of different metacommunity perspectives, a combination of species sorting and mass effects best explains the observed patterns. Additionally, priority effects and monopolization may buffer against the homogenising effects of dispersal and contribute to the distinctness of isolated communities. This is one of the first studies to present evidence for spatial patterns in aquatic communities on such a small spatial scale (a rock ledge of ±9000 m²). Bridging the gap between theory and observed patterns in natural systems is one of the main challenges for future metacommunity research. Small aquatic habitats such as pitcher plants and freshwater rock pools may well have an important role to play as model systems to study ecological processes in a natural spatially explicit environment.     

Relationship Between Soil Properties and Patterns of Bacterial β-diversity Across Reclaimed and Natural Boreal Forest Soils

Productivity gradients in the boreal forest are largely determined by regional-scale changes in soil conditions, and bacterial communities are likely to respond to these changes. Few studies, however, have examined how variation in specific edaphic properties influences the composition of soil bacterial communities along environmental gradients. We quantified bacterial compositional diversity patterns in ten boreal forest sites of contrasting fertility. Bulk soil (organic and mineral horizons) was sampled from sites representing two extremes of a natural moisture-nutrient gradient and two distinct disturbance types, one barren and the other vegetation-rich. We constructed 16S rRNA gene clone libraries to characterize the bacterial communities under phylogenetic- and species-based frameworks. Using a nucleotide analog to label DNA-synthesizing bacteria, we also assessed the composition of active taxa in disturbed sites. Most sites were dominated by sequences related to the α-Proteobacteria, followed by acidobacterial and betaproteobacterial sequences. Non-parametric multivariate regression indicated that pH, which was lowest in the natural sites, explained 34% and 16% of the variability in community structure as determined by phylogenetic-based (UniFrac distances) and species-based (Jaccard similarities) metrics, respectively. Soil pH was also a significant predictor of richness (Chao1) and diversity (Shannon) measures. Within the natural edaphic gradient, soil moisture accounted for 32% of the variance in phylogenetic (but not species) community structure. In the boreal system we studied, bacterial β-diversity patterns appear to be largely related to “master” variables (e.g., pH, moisture) rather than to observable attributes (e.g., plant cover) leading to regional-scale fertility gradients.     

Connecting through space and time: catchment-scale distributions of bacteria in soil,stream water and sediment

Terrestrial and aquatic environments are linked through hydrological networks that transport abiotic components from upslope environments into aquatic ecosystems. However, our understanding of how bacteria are transported through these same networks is limited. Here, we applied 16S rRNA gene sequencing to over 500 soil, stream water and stream sediment samples collected within a native forest catchment to determine the extent to which bacterial communities in these habitats are connected. We provide evidence that while the bacterial communities in each habitat were significantly distinct from one another (PERMANOVA pairwise P < 0.001), the bacterial communities in soil and stream samples were weakly connected to each other when stream sediment sample locations were downhill of surface runoff flow paths. This pattern decreased with increasing distance between the soil and sediment samples. The connectivity between soil and stream water samples was less apparent and extremely transient; the greatest similarity between bacterial communities in soil and stream water overall was when comparing stream samples collected 1 week post soil sampling. This study shows how bacterial communities in soil, stream water and stream sediments are connected at small spatial scales and provides rare insights into the temporal dynamics of terrestrial and aquatic bacterial community connectivity.     

集合群落及其在河流鱼类群聚研究中的应用

集合群落(metacommunity)是指多个潜在相互作用的物种通过它们之间的扩散而连接在一起的一组局域群落,目前已成为斑块生境下生物群落结构、格局和动态的重要理论基础之一。斑块动态、物种排序、群体效应和中性模型等4种理论模型,可用于解释不同情形下集合群落内物种的迁移状况,描述集合群落的动态。可采用群落结构或生态学机制等途径,来阐述所研究的群落是属于哪一种特定的集合群落类型。集合群落可用于研究河流鱼类群聚,解释鱼类的群落结构等问题。另外本文还结合我国水域生态环境及水生生物现状,对今后集合群落的研究作了展望。     

Evidence for structuring of bacterial community composition by organic carbon source in temperate lakes

Water entering lakes from the surrounding watershed often delivers large amounts of terrestrial-derived dissolved organic carbon (DOC) that can contribute to aquatic bacterial production. However, research suggests that phytoplankton-derived DOC is more labile than its terrestrial counterpart, owing to microbial processing of terrestrial-derived DOC along its flow path to surface waters. The ratio of water colour (absorbance at 440 nm) to chlorophyll a has been suggested as a simple measure of the relative contribution of terrestrial and aquatic primary production to aquatic secondary production. To explore the correlation between primary DOC source and the occurrence of bacterial taxonomic groups, we conducted a survey of bacterial 16S rRNA gene composition in 15 lakes positioned along a water colour : chlorophyll a gradient. Our goal was to identify bacterial taxa occurrence patterns along the colour : chlorophyll a gradient that may indicate a competitive advantage for bacterial taxa using terrestrial or aquatic carbon. We observed a large number of bacterial taxa occurrence patterns suggestive of carbon substrate niche partitioning, especially when relatively highly resolved taxonomic groups were considered. Our survey supports the hypothesis that bacterial taxa partition along a carbon substrate source gradient and highlights carbon source–bacterial interactions that should be explored further.     

Catchment-scale biogeography of riverine bacterioplankton

Lotic ecosystems such as rivers and streams are unique in that they represent a continuum of both space and time during the transition from headwaters to the river mouth. As microbes have very different controls over their ecology, distribution and dispersion compared with macrobiota, we wished to explore biogeographical patterns within a river catchment and uncover the major drivers structuring bacterioplankton communities. Water samples collected across the River Thames Basin, UK, covering the transition from headwater tributaries to the lower reaches of the main river channel were characterised using 16S rRNA gene pyrosequencing. This approach revealed an ecological succession in the bacterial community composition along the river continuum, moving from a community dominated by Bacteroidetes in the headwaters to Actinobacteria-dominated downstream. Location of the sampling point in the river network (measured as the cumulative water channel distance upstream) was found to be the most predictive spatial feature; inferring that ecological processes pertaining to temporal community succession are of prime importance in driving the assemblages of riverine bacterioplankton communities. A decrease in bacterial activity rates and an increase in the abundance of low nucleic acid bacteria relative to high nucleic acid bacteria were found to correspond with these downstream changes in community structure, suggesting corresponding functional changes. Our findings show that bacterial communities across the Thames basin exhibit an ecological succession along the river continuum, and that this is primarily driven by water residence time rather than the physico-chemical status of the river.     

Fate of glacier surface snow-originating bacteria in the glacier-fed hydrologic continuums

Glaciers represent important biomes of Earth and are recognized as key species pools for downstream aquatic environments. Worldwide, rapidly receding glaciers are driving shifts in hydrology, species distributions and threatening microbial diversity in glacier-fed aquatic ecosystems. However, the impact of glacier surface snow-originating taxa on the microbial diversity in downstream aquatic environments has been little explored. To elucidate the contribution of glacier surface snow-originating taxa to bacterial diversity in downstream aquatic environments, we collected samples from glacier surface snows, downstream streams and lakes along three glacier-fed hydrologic continuums on the Tibetan Plateau. Our results showed that glacier stream acts as recipients and vectors of bacteria originating from the glacier environments. The contributions of glacier surface snow-originating taxa to downstream bacterial communities decrease from the streams to lakes, which was consistently observed in three geographically separated glacier-fed ecosystems. Our results also revealed that some rare snow-originating bacteria can thrive along the hydrologic continuums and become dominant in downstream habitats. Finally, our results indicated that the dispersal patterns of bacterial communities are largely determined by mass effects and increasingly subjected to local sorting of species along the glacier-fed hydrologic continuums. Collectively, this study provides insights into the fate of bacterial assemblages in glacier surface snow following snow melt and how bacterial communities in aquatic environments are affected by the influx of glacier snow-originating bacteria.     

植物群落和土壤理化性质对碧塔海湿地土壤细菌群落的影响

土壤微生物是湿地生态系统中土壤-植物系统生源要素迁移转化的引擎。探究湿地生态系统地上植物群落、土壤理化性质和空间结构与土壤细菌间的相互关系是维护湿地生态系统健康和稳定的关键。本研究运用双向指示种分析法(TWINSPAN)对碧塔海湿地采集的35个样方中的植物群落进行分类,并采用高通量测序技术对样方的表层土壤细菌进行测序,分析植物群落、土壤理化性质和空间结构与细菌群落间的关系。结果表明: 双向指示种分析将样方的植物群落划分为3种群落类型,相同类型的植物群落在外貌和结构上相似,说明基于双向指示种分析的数量分类方法在高原湿地生态系统植物群落分类中有较好的适用性;细菌相对丰度统计结果表明,酸杆菌门(21.0%)、绿弯菌门(15.5%)、变形菌门(15.3%)和拟杆菌门 (10.1%)是碧塔海湿地总丰度高于10%的门类,相似性分析(ANOSIM)显示,不同植物群落类型对应的土壤细菌群落存在显著差异,说明植物群落对土壤细菌组成有一定的影响;典范对应分析(CCA)结果显示,植物群落的多样性、含水率、pH、铁和空间结构是影响土壤细菌群落的重要因素。方差分解结果显示,细菌群落既受单一环境变量的影响,也受环境变量间复合作用的影响。综上,地上植物群落、土壤理化环境和空间结构共同塑造细菌群落,地上植物群落-细菌-土壤理化性质是不可分割的整体。