Epigeic earthworms exert a bottleneck effect on microbial communities through gut associated processes

Background

Earthworms play a critical role in organic matter decomposition because of the interactions they establish with microorganisms. The ingestion, digestion, assimilation of organic material in the gut and then casting is the first step in earthworm-microorganism interactions. The current knowledge of these direct effects is still limited for epigeic earthworm species, mainly those living in man-made environments. Here we tested whether and to what extent the earthworm Eisenia andrei is capable of altering the microbiological properties of fresh organic matter through gut associated processes; and if these direct effects are related to the earthworm diet.

Methodology

To address these questions we determined the microbial community structure (phospholipid fatty acid profiles) and microbial activity (fluorescein diacetate hydrolysis) in the earthworm casts derived from three types of animal manure (cow, horse and pig manure), which differed in microbial composition.

Principal Findings

The passage of the organic material through the gut of E. andrei reduced the total microbial biomass irrespective of the type of manure, and resulted in a decrease in bacterial biomass in all the manures; whilst leaving the fungi unaffected in the egested materials. However, unlike the microbial biomass, no such reduction was detected in the total microbial activity of cast samples derived from the pig manure. Moreover, no differences were found between cast samples derived from the different types of manure with regards to microbial community structure, which provides strong evidence for a bottleneck effect of worm digestion on microbial populations of the original material consumed.

Conclusions/Significance

Our data reveal that earthworm gut is a major shaper of microbial communities, thereby favouring the existence of a reduced but more active microbial population in the egested materials, which is of great importance to understand how biotic interactions within the decomposer food web influence on nutrient cycling.     

Species-specific effects of epigeic earthworms on microbial community structure during first stages of decomposition of organic matter

Background

Epigeic earthworms are key organisms in organic matter decomposition because of the interactions they establish with microorganisms. The earthworm species and the quality and/or substrate availability are expected to be major factors influencing the outcome of these interactions. Here we tested whether and to what extent the epigeic earthworms Eisenia andrei, Eisenia fetida and Perionyx excavatus, widely used in vermicomposting, are capable of altering the microbiological properties of fresh organic matter in the short-term. We also questioned if the earthworm-induced modifications to the microbial communities are dependent on the type of substrate ingested.

Methodology/Principal Findings

To address these questions we determined the microbial community structure (phospholipid fatty acid profiles) and microbial activity (basal respiration and microbial growth rates) of three types of animal manure (cow, horse and rabbit) that differed in microbial composition, after being processed by each species of earthworm for one month. No differences were found between earthworm-worked samples with regards to microbial community structure, irrespective of type of manure, which suggests the existence of a bottleneck effect of worm digestion on microbial populations of the original material consumed. Moreover, in mesocosms containing cow manure the presence of E. andrei resulted not only in a decrease in bacterial and fungal biomass, but also in a reduced bacterial growth rate and total microbial activity, while no such reduction was found with E. fetida and P. excavatus.

Conclusions/Significance

Our results point to the species of earthworm with its associated gut microbiota as a strong determinant of the process shaping the structure of microbial communities in the short-term. This must nonetheless be weighed against the fact that further knowledge is necessary to evaluate whether the changes in the composition of microbiota in response to the earthworm species is accompanied by a change in the microbial community diversity and/or function.     

Animal ecosystem engineers modulate the diversity-invasibility relationship

Background

Invasions of natural communities by non-indigenous species are currently rated as one of the most important global-scale threats to biodiversity. Biodiversity itself is known to reduce invasions and increase stability. Disturbances by ecosystem engineers affect the distribution, establishment, and abundance of species but this has been ignored in studies on diversity-invasibility relationships.

Methodology/Principal Findings

We determined natural plant invasion into 46 plots varying in the number of plant species (1, 4, and 16) and plant functional groups (1, 2, 3, and 4) for three years beginning two years after the establishment of the Jena Experiment. We sampled subplots where earthworms were artificially added and others where earthworm abundance was reduced. We also performed a seed-dummy experiment to investigate the role of earthworms as secondary seed dispersers along a plant diversity gradient. Horizontal dispersal and burial of seed dummies were significantly reduced in subplots where earthworms were reduced in abundance. Seed dispersal by earthworms decreased with increasing plant species richness and presence of grasses but increased in presence of small herbs. These results suggest that dense vegetation inhibits the surface activity of earthworms. Further, there was a positive relationship between the number of earthworms and the number and diversity of invasive plants. Hence, earthworms decreased the stability of grassland communities against plant invasion.

Conclusions/Significance

Invasibility decreased and stability increased with increasing plant diversity and, most remarkably, earthworms modulated the diversity-invasibility relationship. While the impacts of earthworms were unimportant in low diverse (low earthworm densities) and high diverse (high floral structural complexity) plant communities, earthworms decreased the stability of intermediate diverse plant communities against plant invasion. Overall, the results document that fundamental processes in plant communities like plant seed burial and invader establishment are modulated by soil fauna calling for closer cooperation between soil animal and plant ecologists.     

蚯蚓对土壤微生物残留物的影响研究评述与展望

蚯蚓如何影响土壤有机碳的固持是土壤生态学的关键科学问题之一。蚯蚓能同时促进土壤有机碳分解和稳定,这种两面作用带来的不确定性被研究者称为"蚯蚓困境"。研究证据和新兴的"土壤微生物碳泵"概念模型表明土壤微生物残留物是土壤有机质的主要贡献者。为系统了解蚯蚓对土壤微生物残留物的影响与可能的机制,研究分析和总结了已有的国内外蚯蚓与微生物残留物（氨基糖）的相关研究成果,表明：（1）过往的研究忽略了蚯蚓对微生物残留物的影响,导致这一方向的研究严重滞后;（2）蚯蚓对土壤微生物残留物影响的方向和大小仍有很大的不确定性,可供量化分析其驱动机制的研究还很缺乏。研究尝试将蚯蚓整合到"土壤微生物碳泵"概念框架中,分析蚯蚓影响土壤微生物残留物3个方面的可能机制,即：（1）改变土壤微生物量、群落结构,（2）改变微生物生理特性,（3）改变土壤团聚体结构等,影响土壤有机碳的积累。同时,本文提出了未来相关研究的6个重点方向,包括：（1）蚯蚓对微生物的选择性取食,（2）肠道介导的微生物"涨落"现象,（3）蚯蚓对矿质结合有机物的"破坏"与"重组",（4）蚯蚓引起的"激发"和"续埋"效应,（5）多生态型相互作用,（6）全球变化背景下的蚯蚓生态学等,以期为进一步揭示蚯蚓-微生物相互作用影响土壤有机碳累积与稳定性的机制提供参考。     

The effect of earthworms on the physiological state of the microbial community at vermicomposting

The effect of earthworms on the microbial community of composts and vermicomposts was assayed by the following parameters: mineralization activity, the levels of physiologically active and growing microbial biomass, the requirement for growth factors, and the spectrum of assimilation of organic substrates by the microbial community. The substrate affinities of microbial enzyme systems in vermicompost were found to be lower than in compost without earthworms, which is evidence of a higher amount of r-strategists in the microbial community of vermicomposts. Physiologically active biomass of microorganisms is higher in peat-based vermicompost than in compost. The microorganisms of vermicomposts and composts experience deficiency in growth factors to a lesser extent than the microorganisms in soil. The presence of earthworms influences the physiological diversity: the Shannon index increases or decreases depending on the type of composted substrate and incubation time. The growth rate of microorganisms increases on various test substrates in the presence of worms.     

Carbon availability controls the growth of detritivores (Lumbricidae) and their effect on nitrogen mineralization

Activity of soil decomposer microorganisms is generally limited by carbon availability, but factors controlling saprophagous soil animals remain largely unknown. In contrast to microorganisms, animals are unable to exploit mineral nutrient pools. Therefore, it has been suggested that soil animals, and earthworms in particular, are limited by the availability of nitrogen. In contrast to this view, a strong increase in density and biomass of endogeic earthworms in response to labile organic carbon addition has been documented in field experiments. The hypothesis that the growth of endogeic earthworms is primarily limited by carbon availability was tested in a laboratory experiment lasting for 10 weeks. In addition, it was investigated whether the effects of earthworms on microbial activity and nutrient mineralization depend on the availability of carbon resources. We manipulated food availability to the endogeic earthworm species Octolasion tyrtaeum by using two soils with different organic matter content, providing access to different amounts of soil, and adding labile organic carbon (glucose) enriched in ¹³C.Glucose addition strongly increased the growth of O. tyrtaeum. From 8 to 17% of the total C in earthworm tissue was assimilated from the glucose added. Soil microbial biomass was not strongly affected by the addition of glucose, though basal respiration was significantly increased and up to 50% of the carbon added as glucose was incorporated into soil organic matter. The impact of earthworms on the mineralization and leaching of nitrogen depended on C availability. As expected, in C-limited soil, the presence of earthworms strongly increased nitrogen leaching. However, when C availability was increased by the addition of glucose, this pattern was reversed, i.e. the presence of O. tyrtaeum decreased nitrogen leaching and its availability to soil microflora. We conclude that irrespective of the total carbon content of soils, O. tyrtaeum was primarily limited by carbon, and that increased carbon availability allowed earthworms to be more effective in mobilizing N. The presence of earthworms increases C limitation of soil microorganisms, due to increased availability of N and P in earthworm casts or a direct depletion of easily available carbon resources by earthworms.     

<Emphasis Type="Italic">Eisenia fetida</Emphasis> (Oligochaeta: Lumbricidae) Modifies the Structure and Physiological Capabilities of Microbial Communities Improving Carbon Mineralization During Vermicomposting of Pig Manure

Although microorganisms are largely responsible for organic matter decomposition, earthworms may also affect the rates of decomposition directly by feeding on and digesting organic matter and microorganisms, or indirectly affect them through their interactions with the microorganisms, basically involving stimulation or depression of the microbial populations. We tested the general hypothesis that microbial populations, and especially fungi, are enhanced by earthworm activity, and also whether earthworms are able to modify the biodiversity of microbial populations, and its relation to the function of the system. In addition, we examined the metabolic quotient and the effect of labile organic C to assess the relationships between earthworm and microbes. We found that decomposition of pig manure has two stages characterized by the presence or absence of earthworms. Thus, the presence of earthworms was related with increases in overall microbial biomass and activity, which decreased when earthworms left the substrate; the same pattern was observed for fungi. Furthermore, earthworms modified the physiological profiles of microbial communities of pig manure, increasing the diversity of substrates utilized. In addition, earthworms promoted a more efficient use of energy of microbial communities, as the metabolic quotient showed. The rate of carbon loss was almost twice where earthworms were present, revealing faster decomposition. Our data match with the recent findings that to maintain essential processes the functional properties of present species are at least as important as the number of species per se. This is in accordance with the “insurance hypothesis,” which states that a large number of species is probably essential for maintaining stable processes in changing environments, as the presence of earthworms would have promoted in pig manure.     

Degradation of potassium rock by earthworms and responses of bacterial communities in its gut and surrounding substrates after being fed with mineral

Background

Earthworms are an ecosystem''s engineers, contributing to a wide range of nutrient cycling and geochemical processes in the ecosystem. Their activities can increase rates of silicate mineral weathering. Their intestinal microbes usually are thought to be one of the key drivers of mineral degradation mediated by earthworms,but the diversities of the intestinal microorganisms which were relevant with mineral weathering are unclear.

Methodology/Principal Findings

In this report, we show earthworms'' effect on silicate mineral weathering and the responses of bacterial communities in their gut and surrounding substrates after being fed with potassium-bearing rock powder (PBRP). Determination of water-soluble and HNO₃-extractable elements indicated some elements such as Al, Fe and Ca were significantly released from mineral upon the digestion of earthworms. The microbial communities in earthworms'' gut and the surrounding substrates were investigated by amplified ribosomal DNA restriction analysis (ARDRA) and the results showed a higher bacterial diversity in the guts of the earthworms fed with PBRP and the PBRP after being fed to earthworms. UPGMA dendrogram with unweighted UniFrac analysis, considering only taxa that are present, revealed that earthworms'' gut and their surrounding substrate shared similar microbiota. UPGMA dendrogram with weighted UniFrac, considering the relative abundance of microbial lineages, showed the two samples from surrounding substrate and the two samples from earthworms'' gut had similarity in microbial community, respectively.

Conclusions/Significance

Our results indicated earthworms can accelerate degradation of silicate mineral. Earthworms play an important role in ecosystem processe since they not only have some positive effects on soil structure, but also promote nutrient cycling of ecosystem by enhancing the weathering of minerals.     

Experimental Evaluation of Herbivory on Live Plant Seedlings by the Earthworm Lumbricus terrestris L. in the Presence and Absence of Soil Surface Litter

Background

Recent studies suggested that the earthworm Lumbricus terrestris might act as a seedling predator by ingesting emerging seedlings, and individuals were observed damaging fresh leaves of various plant species in the field. To evaluate the significance of herbivore behavior of L. terrestris for plant and earthworm performance we exposed 23- to 33-days-old seedlings of six plant species to earthworms in two microcosm experiments. Plants belonged to the three functional groups grasses, non-leguminous herbs, and legumes. Leaf damage, leaf mortality, the number of leaves as well as mortality and growth of seedlings were followed over a period of up to 26 days. In a subset of replicates 0.1 g of soil surface litter of each of the six plant species was provided and consumption was estimated regularly to determine potential feeding preferences of earthworms.

Results

There was no difference in seedling growth, the number of live seedlings and dead leaves between treatments with or without worms. Fresh leaves were damaged eight times during the experiment, most likely by L. terrestris, with two direct observations of earthworms tearing off leaf parts. Another nine leaves were partly pulled into earthworm burrows. Lumbricus terrestris preferred to consume legume litter over litter of the other plant functional groups. Earthworms that consumed litter lost less weight than individuals that were provided with soil and live plants only, indicating that live plants are not a suitable substitute for litter in earthworm nutrition.

Conclusion

Our results demonstrate that L. terrestris damages live plants; however, this behavior occurs only rarely. Pulling live plants into earthworm burrows might induce microbial decomposition of leaves to make them suitable for later consumption. Herbivory on plants beyond the initial seedling stage may only play a minor role in earthworm nutrition and has limited potential to influence plant growth.     

Transformation of buffalo manure by composting or vermicomposting to rehabilitate degraded tropical soils

The addition of composted buffalo manure may lead to qualitative and quantitative improvement of the organic matter content of degraded tropical agricultural soils in Northern Vietnam. The objectives of this study were to follow the biochemical changes occurring during composting of buffalo manure with and without earthworms during 3 months and to study the effect of the end products (compost and vermicompost) on soil biochemical parameters and plant growth after two months of incubation under controlled conditions in an open pot experiment. Our conceptual approach included characterisation of organic matter of the two composts before and after addition to soil by elemental, isotopic analysis and analytical pyrolysis and comparison with conventional fertilisation. We also analysed for lignin content and composition.Our results showed that composting in the presence of earthworms led to stronger transformation of buffalo manure than regular composting. Vermicompost was enriched in N-containing compounds and depleted in polysaccharides. It further contained stronger modified lignin compared to regular compost. In the bulk soil, the amendment of compost and vermicompost led to significant modification of the soil organic matter after 2 months of exposure to natural weather conditions. The lignin component of SOM was unaffected whatever the origin of the organic amendment. Compost and vermicompost amendments both enhanced aggregation and increased the amount of organic matter in water stable aggregates. However, vermicompost is preferable to compost due to its beneficial effect on plant growth, while having similar positive effects on quantity and quality of SOM.     

Fruit and Soil Quality of Organic and Conventional Strawberry Agroecosystems

Background

Sale of organic foods is one of the fastest growing market segments within the global food industry. People often buy organic food because they believe organic farms produce more nutritious and better tasting food from healthier soils. Here we tested if there are significant differences in fruit and soil quality from 13 pairs of commercial organic and conventional strawberry agroecosystems in California.

Methodology/Principal Findings

At multiple sampling times for two years, we evaluated three varieties of strawberries for mineral elements, shelf life, phytochemical composition, and organoleptic properties. We also analyzed traditional soil properties and soil DNA using microarray technology. We found that the organic farms had strawberries with longer shelf life, greater dry matter, and higher antioxidant activity and concentrations of ascorbic acid and phenolic compounds, but lower concentrations of phosphorus and potassium. In one variety, sensory panels judged organic strawberries to be sweeter and have better flavor, overall acceptance, and appearance than their conventional counterparts. We also found the organically farmed soils to have more total carbon and nitrogen, greater microbial biomass and activity, and higher concentrations of micronutrients. Organically farmed soils also exhibited greater numbers of endemic genes and greater functional gene abundance and diversity for several biogeochemical processes, such as nitrogen fixation and pesticide degradation.

Conclusions/Significance

Our findings show that the organic strawberry farms produced higher quality fruit and that their higher quality soils may have greater microbial functional capability and resilience to stress. These findings justify additional investigations aimed at detecting and quantifying such effects and their interactions.     

Earthworm invasions of ecosystems devoid of earthworms: effects on soil microbes

Recent studies document North American earthworm invasions and their profound effects on the structure of the soil profile, which is the habitat for soil microorganisms (mainly fungi and bacteria). Dramatic alterations made to these layers during earthworm invasion significantly change microbial community structure and therefore microbial activities such as C transformations. Understanding the impacts of earthworm invasion on the microbes themselves will give insight into earthworm effects on microbial activities. Bacterial and actinomycete communities in earthworm guts and casts have not been studied in environments recently invaded by earthworms. Earthworm invasion tended to decrease fungal species density and fungal species diversity and richness. The presence of earthworms decreased zygomycete species abundance probably due to disruption of fungal hyphae. Physical disruption of hyphae may also explain decreased mycorrhizal colonization rates, decreased mycorrhizal abundance and altered mycorrhizal morphology in the presence of earthworms. Mixing of organic layers into mineral soil during earthworm invasion tended to decrease microbial biomass in forest floor materials while increasing it in mineral soil. In newly invaded forest soils, microbial respiration and the metabolic quotient tended to decline. In forests where either the microbial community has had time to adapt to earthworm activities, or where the destruction of the forest floor is complete, as in invasions by the Asian Amynthas hawayanus, the presence of earthworms tends to increase the metabolic quotient indicating a shift to a smaller, more active microbial community.     

Soil microbial properties and plant growth responses to carbon and water addition in a temperate steppe: the importance of nutrient availability

Background

Global climatic change is generally expected to stimulate net primary production, and consequently increase soil carbon (C) input. The enhanced C input together with potentially increased precipitation may affect soil microbial processes and plant growth.

Methodology/Principal Findings

To examine the effects of C and water additions on soil microbial properties and plant growth, we conducted an experiment lasting two years in a temperate steppe of northeastern China. We found that soil C and water additions significantly affected microbial properties and stimulated plant growth. Carbon addition significantly increased soil microbial biomass and activity but had a limited effect on microbial community structure. Water addition significantly increased soil microbial activity in the first year but the response to water decreased in the second year. The water-induced changes of microbial activity could be ascribed to decreased soil nitrogen (N) availability and to the shift in soil microbial community structure. However, no water effect on soil microbial activity was visible under C addition during the two years, likely because C addition alleviated nutrient limitation of soil microbes. In addition, C and water additions interacted to affect plant functional group composition. Water addition significantly increased the ratio of grass to forb biomass in C addition plots but showed only minor effects under ambient C levels. Our results suggest that soil microbial activity and plant growth are limited by nutrient (C and N) and water availability, and highlight the importance of nutrient availability in modulating the responses of soil microbes and plants to potentially increased precipitation in the temperate steppe.

Conclusions/Significance

Increased soil C input and precipitation would show significant effects on soil microbial properties and plant growth in the temperate steppe. These findings will improve our understanding of the responses of soil microbes and plants to the indirect and direct climate change effects.     

A Feeding Induced Switch from a Variable to a Homogenous State of the Earthworm Gut Microbiota within a Host Population

Background

The distribution pattern of the earthworm gut microbiota at the host population level is of fundamental importance to understand host-microbiota interactions. Our current understanding of these interactions is very limited. Since feeding represents a main perturbation of the gut microbiota, we determined the effect of a single dose of feed on the microbiota associated with an earthworm population in a simulated microenvironment.

Methodology

Earthworms were sampled 0, 1 and 7 days after feeding. We determined the overall composition of the earthworm-associated microbiota by 16S rRNA gene cloning and sequencing. Based on the 16S rRNA gene data we constructed quantitative PCR''s (Q-PCR) for the seven most dominating bacterial groups.

Principal Findings

Q-PCR revealed low density and highly variable microbiota among the earthworms before feeding, while a high-density homologous microbiota resulted from feeding. We found that the microbiota 1 day after feeding was more equal to the microbiota after 7 days than before feeding. Furthermore, we found that the gut microbiota was very distinct from that of the bedding and the feed.

Significance

The homogenous population response represents fundamental new knowledge about earthworm gut associated bacteria.     

Deposit-Feeding Sea Cucumbers Enhance Mineralization and Nutrient Cycling in Organically-Enriched Coastal Sediments

Background

Bioturbators affect multiple biogeochemical interactions and have been suggested as suitable candidates to mitigate organic matter loading in marine sediments. However, predicting the effects of bioturbators at an ecosystem level can be difficult due to their complex positive and negative interactions with the microbial community.

Methodology/Principal Findings

We quantified the effects of deposit-feeding sea cucumbers on benthic algal biomass (microphytobenthos, MPB), bacterial abundance, and the sediment–seawater exchange of dissolved oxygen and nutrients. The sea cucumbers increased the efflux of inorganic nitrogen (ammonium, NH₄ ⁺) from organically enriched sediments, which stimulated algal productivity. Grazing by the sea cucumbers on MPB (evidenced by pheopigments), however, caused a net negative effect on primary producer biomass and total oxygen production. Further, there was an increased abundance of bacteria in sediment with sea cucumbers, suggesting facilitation. The sea cucumbers increased the ratio of oxygen consumption to production in surface sediment by shifting the microbial balance from producers to decomposers. This shift explains the increased efflux of inorganic nitrogen and concordant reduction in organic matter content in sediment with bioturbators.

Conclusions/Significance

Our study demonstrates the functional role and potential of sea cucumbers to ameliorate some of the adverse effects of organic matter enrichment in coastal ecosystems.     

Estimating the power of indirect comparisons: a simulation study

Background

Indirect comparisons are becoming increasingly popular for evaluating medical treatments that have not been compared head-to-head in randomized clinical trials (RCTs). While indirect methods have grown in popularity and acceptance, little is known about the fragility of confidence interval estimations and hypothesis testing relying on this method.

Methods

We present the findings of a simulation study that examined the fragility of indirect confidence interval estimation and hypothesis testing relying on the adjusted indirect method.

Findings

Our results suggest that, for the settings considered in this study, indirect confidence interval estimation suffers from under-coverage while indirect hypothesis testing suffers from low power in the presence of moderate to large between-study heterogeneity. In addition, the risk of overestimation is large when the indirect comparison of interest relies on just one trial for one of the two direct comparisons.

Interpretation

Indirect comparisons typically suffer from low power. The risk of imprecision is increased when comparisons are unbalanced.     

Horizontal gene transfer in an acid mine drainage microbial community

Background

Horizontal gene transfer (HGT) has been widely identified in complete prokaryotic genomes. However, the roles of HGT among members of a microbial community and in evolution remain largely unknown. With the emergence of metagenomics, it is nontrivial to investigate such horizontal flow of genetic materials among members in a microbial community from the natural environment. Because of the lack of suitable methods for metagenomics gene transfer detection, microorganisms from a low-complexity community acid mine drainage (AMD) with near-complete genomes were used to detect possible gene transfer events and suggest the biological significance.

Results

Using the annotation of coding regions by the current tools, a phylogenetic approach, and an approximately unbiased test, we found that HGTs in AMD organisms are not rare, and we predicted 119 putative transferred genes. Among them, 14 HGT events were determined to be transfer events among the AMD members. Further analysis of the 14 transferred genes revealed that the HGT events affected the functional evolution of archaea or bacteria in AMD, and it probably shaped the community structure, such as the dominance of G-plasma in archaea in AMD through HGT.

Conclusions

Our study provides a novel insight into HGT events among microorganisms in natural communities. The interconnectedness between HGT and community evolution is essential to understand microbial community formation and development.

Electronic supplementary material

The online version of this article (doi:10.1186/s12864-015-1720-0) contains supplementary material, which is available to authorized users.     

The influence of invasive earthworms on indigenous fauna in ecosystems previously uninhabited by earthworms

Recent studies on earthworm invasion of North American soils report dramatic changes in soil structure, nutrient dynamics and plant communities in ecosystems historically free of earthworms. However, the direct and indirect impacts of earthworm invasions on animals have been largely ignored. This paper summarizes the current knowledge on the impact of earthworm invasion on other soil fauna, vertebrates as well as invertebrates.Earthworm invasions can have positive effects on the abundance of other soil invertebrates, but such effects are often small, transient, and restricted to habitats with harsh climates or a long history of earthworm co-occurrence with other soil invertebrates. Middens and burrows can increase soil heterogeneity and create microhabitats with a larger pore size, high microbial biomass, and microclimates that are attractive to micro- and mesofauna. Under harsh climatic conditions, the aggregates formed by earthworms may increase the stability of soil microclimates. Positive effects can also be seen when comminution and mucus secretion increase the palatability of unpalatable organic material for microorganisms which are the main food of most micro- and mesofaunal groups. For larger invertebrates or small vertebrates, invasive earthworms may become important prey, with the potential to increase resource availability. In the longer-term, the activity of invading earthworms can have a strong negative impact on indigenous faunal groups across multiple trophic levels. Evidence from field and laboratory studies indicates that the restructuring of soil layers, particularly the loss of organic horizons, physical disturbance to the soil, alteration of understory vegetation, and direct competition for food resources, lead directly and indirectly to significant declines in the abundance of soil micro- and mesofauna. Though studies of invasive earthworm impacts on the abundance of larger invertebrates or vertebrates are generally lacking, recent evidence suggests that reduced abundance of small soil fauna and alteration of soil microclimates may be contributing to declines in vertebrate fauna such as terrestrial salamanders. Preliminary evidence also suggests the potential for earthworm invasions to interact with other factors such as soil pollution, to negatively affect vertebrate populations.     

蚯蚓与微生物的相互作用

蚯蚓从成体到卵内均有微生物,微生物来源于蚯蚓生活的环境,在消化消化道时,随食物进入体内的真菌营养体及大部分细菌被杀死,只有真菌的孢子和部分细菌仍保持生活力,生长缓慢的细菌通过蚯蚓消化道后群体下降;而生长快的细菌,由于在消化道内迅速繁殖,在蚯蚓排泄物中的群体数量甚至会超过进入蚯蚓体内时的数量,蚯蚓能促进土表有益和致病微生物在土壤内从向传播,但也能减轻由病原真菌引起的病害。真菌是蚯蚓食物的一部分,消化     

两种代表性蚯蚓对设施菜地土壤微生物群落结构及理化性质的影响

不同生态型蚯蚓的取食偏好和生境有所差异,因此蚯蚓的生态型差异可能关乎其对土壤性质的不同影响;有关不同生态型蚯蚓对土壤性质尤其是微生物学性质影响的研究有助于了解蚯蚓生态功能的作用机制。在野外调控试验的第4年采集土壤,研究了牛粪混施和表施处理下内层种威廉腔环蚓(Metaphire guillelmi)和表层种赤子爱胜蚓(Eisenia foetida)对设施菜地土壤微生物群落结构和主要理化性质的影响。结果表明,土壤微生物群落结构同时受到蚯蚓种类和牛粪施用方式的影响。牛粪表施时,两种蚯蚓均显著降低了菌根真菌、真菌生物量和原生动物生物量(P0.05);牛粪混施时,不同蚯蚓的影响有所差异,威廉腔环蚓明显增加了菌根真菌、真菌生物量和放线菌生物量,而赤子爱胜蚓的作用不明显。此外,两种蚯蚓均提高了土壤孔隙度、团聚体稳定性和土壤p H、矿质氮以及微生物生物量碳氮水平,但提高幅度取决于蚯蚓种类和牛粪施用方式。冗余分析表明蚯蚓影响下土壤微生物群落结构的变化与团聚体稳定性、p H、速效磷、矿质氮呈正相关,而与土壤容重呈负相关。