Reaction of microorganisms to the digestive fluid of earthworms

The reaction of soil bacteria and fungi to the digestive fluid of the earthworm Aporrectodea caliginosa was studied. The fluid was obtained by centrifugation of the native enzymes of the digestive tract. The inhibition of growth of certain bacteria, spores, and fungal hyphae under the effect of extracts from the anterior and middle sections of the digestive tract of A. caliginosa was discovered for the first time. In bacteria, microcolony formation was inhibited as early as 20–30 s after the application of the gut extracts, which may indicate the nonenzymatic nature of the effect. The digestive fluid exhibited the same microbicidal activity whether the earthworms were feeding on soil or sterile sand. This indicates that the microbicidal agents are formed within the earthworm’s body, rather than by soil microorganisms. The effect of the digestive fluid from the anterior and middle divisions is selective in relation to different microorganisms. Of 42 strains of soil bacteria, seven were susceptible to the microbicidal action of the fluid (Alcaligenes faecalis 345-1, Microbacterium sp. 423-1, Arthrobacter sp. 430-1, Bacillus megaterium 401-1, B. megaterium 413-1, Kluyvera ascorbata 301-1, Pseudomonas reactans 387-2). The remaining bacteria did not die in the digestive fluid. Of 13 micromycetes, the digestive fluid inhibited spore germination in Aspergillus terreus and Paecilomyces lilacinus and the growth of hyphae in Trichoderma harzianum and Penicillium decumbens. The digestive fluid stimulated spore germination in Alternaria alternata and the growth of hyphae in Penicillium chrysogenum. The reaction of the remaining micromycetes was neutral. The gut fluid from the posterior division of the abdominal tract did not possess microbicidal activity. No relation was found between the reaction of microorganisms to the effects of the digestive fluid and the taxonomic position of the microorganisms. The effects revealed are similar to those shown earlier for millipedes and wood lice in the following parameters: quick action of the digestive fluid on microorganisms, and the selectivity of the action on microorganisms revealed at the strain level. The selective effect of the digestive gut fluid of the earthworms on soil microorganisms is important for animal feeding, maintaining the homeostasis of the gut microbial community, and the formation of microbial communities in soils.     

Specific features of nitrogen transformation in the gut and coprolites of earthworms

It has been found that nitrogenase activity in the guts and coprolites of three earthworm species (Lumbricus terrestris, Aporrectodea rosea, and Aporrectodea caliginosa) is one to three orders of magnitude higher than that in the control soil. In A. caliginosa earthworms, the actual nitrogenase activity remains high upon keeping them in soils of different types. However, it reaches a peak in the gut lumen and decreases to a minimum on its wall, which is evidence for a major role of transitional microflora in intestinal nitrogen fixation. Nitrogen fixation activity in freshly excreted coprolites decreases to one-half or one-third of that in the gut and then increases again, reaching a peak on days 3–5 of exposure in the soil. According to the results of multisubstrate testing, the functional diversity of nitrogen-fixing soil microorganisms increases in the course of passage through the earthworm gut. Thus, the microbial community in coprolites retains its functional potential and, within a few days, shows the second peak of activity in the soil. Due to a short-term increase in the rate of nitrogen fixation, coprolites contain a pool of bound amino acids, which become involved in the formation of new humus substances.     

Earthworm effects on plant growth do not necessarily decrease with soil fertility

Earthworms are known to generally increase plant growth. However, because plant-earthworm interactions are potentially mediated by soil characteristics the response of plants to earthworms should depend on the soil type. In a greenhouse microcosm experiment, the responsiveness of plants (Veronica persica, Trifolium dubium and Poa annua) to two earthworm species (in combination or not) belonging to different functional groups (Aporrectodea. caliginosa an endogeic species, Lumbricus terrestris an anecic species) was measured in term of biomass accumulation. This responsiveness was compared in two soils (nutrient rich and nutrient poor) and two mineral fertilization treatments (with and without). The main significant effects on plant growth were due to the anecic earthworm species. L. terrestris increased the shoot biomass and the total biomass of T. dubium only in the rich soil. It increased also the total biomass of P. annua without mineral fertilization but had the opposite effect with fertilization. Mineral fertilization, in the presence of L. terrestris, also reduced the total biomass of V. persica. L. terrestris did not only affect plant growth. In P. annua and V. persica A. caliginosa and L. terrestris also affected the shoot/root ratio and this effect depended on soil type. Finally, few significant interactions were found between the anecic and the endogeic earthworms and these interactions did not depend on the soil type. A general idea would be that earthworms mostly increase plant growth through the enhancement of mineralization and that earthworm effects should decrease in nutrient-rich soils or with mineral fertilization. However, our results show that this view does not hold and that other mechanisms are influential.     

Root Foraging Influences Plant Growth Responses to Earthworm Foraging

Interactions among the foraging behaviours of co-occurring animal species can impact population and community dynamics; the consequences of interactions between plant and animal foraging behaviours have received less attention. In North American forests, invasions by European earthworms have led to substantial changes in plant community composition. Changes in leaf litter have been identified as a critical indirect mechanism driving earthworm impacts on plants. However, there has been limited examination of the direct effects of earthworm burrowing on plant growth. Here we show a novel second pathway exists, whereby earthworms (Lumbricus terrestris L.) impact plant root foraging. In a mini-rhizotron experiment, roots occurred more frequently in burrows and soil cracks than in the soil matrix. The roots of Achillea millefolium L. preferentially occupied earthworm burrows, where nutrient availability was presumably higher than in cracks due to earthworm excreta. In contrast, the roots of Campanula rotundifolia L. were less likely to occur in burrows. This shift in root behaviour was associated with a 30% decline in the overall biomass of C. rotundifolia when earthworms were present. Our results indicate earthworm impacts on plant foraging can occur indirectly via physical and chemical changes to the soil and directly via root consumption or abrasion and thus may be one factor influencing plant growth and community change following earthworm invasion. More generally, this work demonstrates the potential for interactions to occur between the foraging behaviours of plants and soil animals and emphasizes the importance of integrating behavioural understanding in foraging studies involving plants.     

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     

A comparative study of gut microbiota profiles of earthworms fed in three different substrates

The gut microbiome of earthworms has a complex interdependence with the host. When the soil minerals pass through earthworm’s gut, they may affect the gut microbiota. To gain insight into the response of gut microbiota to the passed minerals, we fed earthworm (Eisenia fetida) on nutrient-poor soil and ore powder, and used high throughput sequencing to characterize the earthworm intestinal microbial community to find evidence for a core bacterial community of the E. fetida. The results showed that earthworms’ gut maintained a core microbiome that appeared in all samples. These core microbiota may play a significant role in a species’ environmental interactions. The composition of intestinal microbiomes varied with substrates. The earthworm guts from two nutrient-poor substrates had similar microbial communities and they were different from nutrient-rich substrate. Proteobacteria and Bacteroidetes were more abundant in the gut of earthworms kept on a nutrient-poor substrate such as ore powder or mineral soil than in the gut of earthworms kept in organic-rich compost soil; some of these microorganisms may help earthworms survive in nutrient-poor substrates.     

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.     

蚯蚓在植物修复芘污染土壤中的作用

采用盆栽试验法,研究了蚯蚓(Pheretima hupeiensis)在植物修复芘污染土壤中的作用。结果显示,试验浓度(20.24-321.42 mg/kg) 范围内,蚯蚓活动促进了芘污染土壤中修复植物黑麦草(Lolium multiforum)黑麦草的生长,其根冠比明显增大。添加蚯蚓72 d后,种植黑麦草的土壤中芘的去除率高达60.01%-86.26%,其平均去除率(74.66%)比无蚯蚓活动的土壤-植物系统(64.55%)提高10.11%,比无植物对照组(18.24%)提高56.42%。各种生物、非生物修复因子中,植物-微生物交互作用对芘去除的平均贡献率(51.75%)最为突出,比无蚯蚓活动时(44.94%)提高6.81%。说明蚯蚓活动可强化土壤-植物系统对土壤芘污染的修复作用。     

The role of substrate feeding earthworms (Lumbricidae) for bioturbation in a beechwood soil

Summary Burrow formation (burrow length and faeces production) of the substrate feeding earthworm Aporrectodea caliginosa (Savigny) in relation to three temperatures (5, 10 and 15°C) and soil moisture contents (48, 60 and 73% water of dry wt) was determined. Soil moisture content affected the burrowing activity of A. caliginosa only below a distinct threshold (60% water of dry wt). At sufficient moisture an, increase in temperature by 5° C approximately doubled the amount of egesta produced. Another substrate feeding earthworm (Octolasion lacteum (Örley)) showed a similar pattern of burrowing activity.The data for the dependence of the faeces production of A. caliginosa on soil temperature and moisture content were combined with field data. The amount of faeces produced by the population of this earthworm species in a beechwood on limestone was calculated to be about 4.23 kg ha^-1 a^-1. The amount of egesta produced by all substrate feeding species is assumed to exceed 6 kg ha^-1 a^-1, which is equivalent to a soil layer of approximately 9 mm. This group of earthworms is therefore considered to be of greater importance for bioturbation and the formation of mull than previously assumed.     

The Effect of Biochar and Its Interaction with the Earthworm Pontoscolex corethrurus on Soil Microbial Community Structure in Tropical Soils

Biochar effects on soil microbial abundance and community structure are keys for understanding the biogeochemical cycling of nutrients and organic matter turnover, but are poorly understood, in particular in tropical areas. We conducted a greenhouse experiment in which we added biochars produced from four different feedstocks [sewage sludge (B1), deinking sewage sludge (B2), Miscanthus (B3) and pine wood (B4)] at a rate of 3% (w/w) to two tropical soils (an Acrisol and a Ferralsol) planted with proso millet (Panicum milliaceum L.). The interactive effect of the addition of earthworms was also addressed. For this purpose we utilized soil samples from pots with or without the earthworm Pontoscolex corethrurus, which is a ubiquitous earthworm in tropical soils. Phospholipid fatty acid (PLFA) measurements showed that biochar type, soil type and the presence of earthworms significantly affected soil microbial community size and structure. In general, biochar addition affected fungal but not bacterial populations. Overall, biochars rich in ash (B1 and B2) resulted in a marked increase in the fungi to bacteria ratio, while this ratio was unaltered after addition of biochars with a high fixed carbon content (B3 and B4). Our study remarked the contrasting effect that both, biochar prepared from different materials and macrofauna, can have on soil microbial community. Such changes might end up with ecosystem-level effects.     

Role of earthworm-straw interactions on phytoremediation of Cu contaminated soil by ryegrass

It is well known that the earthworm's activities can increase the availability of soil nutrients, improve soil structure, and enhance the biomass of plants in uncontaminated soil. Recently, many researchers found that some metal-tolerant earthworms can survive and even change the fractional distribution of heavy metals in contaminated soil. Furthermore, it has been revealed that earthworms are able to increase metal availability, and therefore, accumulate more metals in plants through their burrowing and casting activity. It is clear that the influence of soil animals is an important factor for phyto-remedation that must be taken into account. ~In this article, the authors studied some effects of addition of earthworms (Metaphire guillelmi), corn straw, and in combinations of earthworms and corn straw on the growth and Cu uptake by ryegrass in Cu contaminated pot soils. The experiment consisted of four levels of Cu addition (0, 100, 200, 400 mg·kg?1) and four treatments. The treatments were 1. control (CK); 2.straw mulching only (M); 3. earthworm additions to soil only (E); and 4.straw mulching plus earthworm additions (ME). Each treatment had three replicates. 10 seeds of ryegrass (Lolium multiflorum) were sowed in each pot and harvested after 30 days. After 30 days of incubation, all earthworms were found to be alive and the pot soils were burrowed through by earthworms. Results showed that the biomass of earthworm declined with the increase of the dosage of Cu additions. The biomass of earthworm increased significantly in treatment 4 (ME) as compared with treatment 3 (E). Not only the earthworms could get more food from the straw, but also could counteract some negative effects of Cu on the earthworms. The rates of straw decomposition in ME treatment increased by about 58.11% ?77.32%. The earthworm activities increased root biomass of ryegrass significantly, but did not show the effect on plant root growth. On the contrary, straw enhanced roots biomass significantly instead of shoots biomass. It was also found that the concentration of Cu in the plant shoot and the plant root, as well as plant Cu uptake were enhanced by earthworm's activities and straw mulching. The increased amount by straw mulching was lower than that of earthworms (E). The treatment of the earthworm–straw mulching combinations enhanced plant Cu concentration, and the amount increased by it was lower than that of the earthworm treatment (E) but higher than that of straw mulching treatment (M). The accumulation factors of copper in the shoots of ryegrass were increased by 31.22% ?121.07%, 2.12% ?61.28% and 25.56% ?132.64%, respectively, in treatment 3(E), 2(M), and 4(ME), respectively. In conclusion, the earthworm activities, straw-mulching and their interactions may have potential roles in elevating phyto-extraction efficiency in low to medium level Cu contaminated soil.     

Protozoa,Nematoda and Lumbricidae in the rhizosphere of Hordelymus europeaus (Poaceae): faunal interactions,response of microorganisms and effects on plant growth

Interactions among protozoa (mixed cultures of ciliates, flagellates and naked amoebae), bacteria-feeding nematodes (Pellioditis pellio Schneider) and the endogeic earthworm species Aporrectodea caliginosa (Savigny) were investigated in experimental chambers with soil from a beechwood (Fagus sylvatica L.) on limestone. Experimental chambers were planted with the grass Hordelymus europeaus L. (Poaceae) and three compartments separated by 45-m mesh were established: rhizosphere, intermediate and non-rhizosphere. The experiment lasted for 16 weeks and the following parameters were measured at the end of the experiment: shoot and root mass of H. europaeus, carbon and nitrogen content in shoots and roots, density of ciliates, amoebae, flagellates and nematodes, microbial biomass (SIR), basal respiration, streptomycin sensitive respiration, ammonium and nitrate contents, phosphate content of soil compartments. In addition, leaching of nutrients (nitrogen and phosphorus) and leachate pH were measured at regular intervals in leachate obtained from suction cups in the experimental chambers. Protozoa stimulated the recovery of nitrifying bacteria following defaunation (by chloroform fumigation) and increased nitrogen losses as nitrate in leachate. In contrast, protozoa and nematodes reduced leaching of phosphate, an effect ascribed to stimulation of microbial growth early in the experiment. Earthworms strongly increased the amount of extractable mineral nitrogen whereas it was strongly reduced by protozoa and nematodes. Both protozoa and nematodes reduced the stimulatory effect of earthworms on nitrogen mineralization. Microbial biomass, basal respiration, and numbers of protozoa and nematodes increased in the vicinity of the root. Protozoa generally caused a decrease in microbial biomass whereas nematodes and earthworms reduced microbial biomass only in the absence of protozoa. None of the animals studied significantly affected basal respiration, but specific respiration of microorganisms (O₂ consumption per unit biomass) was generally higher in animal treatments. The stimulatory effect of nematodes and earthworms, however, occurred only in the absence of protozoa. The sensitivity of respiration to streptomycin suggested that protozoa selectively grazed on bacterial biomass but the bacterial/fungal ratio appeared to be unaffected by grazing of P. pellio. Earthworms reduced root biomass of H. europaeus, although shoot biomass remained unaffected, and concentrations of nitrogen in shoots and particularly in roots were strongly increased by earthworms. Both nematodes and protozoa increased plant biomass, particularly that of roots. This increase in plant biomass was accompanied by a marked decrease in nitrogen concentrations in roots and to a lesser extent in shoots. Generally, the effects of protozoa on plant growth considerably exceeded those of nematodes. It is concluded that nematodes and protozoa stimulated plant growth by non-nutritional effects, whereas the effects of earthworms were caused by an increase in nutrient supply to H. europaeus.     

Distribution, prevalence and intensity of earthworm populations in arable land and grassland in Scotland

The earthworms of arable and pasture fields from 100 randomly chosen arable farms were identified and counted and their relationship with soil factors examined. Thirteen species of earthworm were recorded including Lumbricus friendi from Scotland for the first time. The most prevalent and numerous species were Aporrectodea longa, Aporrectodea caliginosa and Lumbricus terrestris. All earthworm species had cosmopolitan distributions apart from Aporrectodea nocturna and Lumbricus festivus which were confined to southern/central Scotland. Four soil characteristics (% moisture, % sand, % organic matter and pH) shared no relationship with species recorded but tillage may have had a detrimental effect on species composition and size of population. It is suggested that the magnitudes of the earthworm populations found, particularly in permanent pasture, were important in maintaining soil structure and fertility, and the spread of the New Zealand flatworm (Artioposthia triangulata), an obligate predator of earthworms, could have a considerable impact on agricultural productivity and wildlife in Scotland.     

Influence of Different Ecological Groups of Earthworms on the Intensity of Nitrogen Fixation

The results of studying nitrogen fixation in organic substrates processed by different ecological groups of earthworms suggest that the earthworms Aporrectodea caliginosa actively stimulate nonsymbiotic nitrogen fixation. In this respect, they exceed the manure worms Eizenia foetida tens of times due to the formation in the organic substrate of conditions favorable for the nitrogen fixing bacteria, and namely: low content of nitrogen easily accessible for microorganisms, changes in the structure of the microbial community of the substrate in favor of non-spore forms of bacteria, and suppression of the growth of saprophyte bacilli, the main competitors of nitrogen fixing bacteria for carbon nourishment sources.     

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.     

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。     

Culturable microorganisms from the earthworm digestive tract

The cultured aerobic copiotrophic bacteria and fungi from food-free digestive tracts of Aporrectodea caliginosa, Lumbricus terrestris, and Eisenia fetida earthworms, soil (compost), and fresh earthworm excrements were investigated. The microorganisms were isolated on nutrient media and identified by sequencing the fragments of bacterial 16S rRNA and fungal 28S rRNA (D1/D2 domain) gene sequences with subsequent phylogenetic analysis. Bacteria isolated from the digestive tracts of earthworms belonged to the families Aeromonadaceae, Comamonadaceae, Enterobacteriaceae, Flavobacteriaceae, Moraxellaceae, Pseudomonadaceae, and Sphingobacteriaceae (Bacteroidetes), as well as Actinobacteria. For five strains, namely Ochrobactrum sp. 341-2 (α-Proteobacteria), Massilia sp. 557-1 (β-Proteobacteria), Sphingobacterium sp. 611-2 (Bacteroidetes), Leifsonia sp. 555-1, and a bacterium from the family Microbacteriaceae, isolate 521-1 (Actinobacteria), the similarity to known 16S rRNA sequences was 93–97%; they therefore, probably belong to new species and genera. Bacterial groups isolated from the digestive tracts of earthworms were significantly different from those isolated from soil and excrements. Some bacterial taxa occurred in different sections of A. caliginosa intestine and in intestines of different earthworm species; however, the overall composition of bacterial communities in these objects is different. Existence of bacterial groupings symbiotically associated with intestines is proposed. Among the fungi, Bjerkandera adusta and Syspastospora parasitica were isolated from the cleaned digestive tracts as light-colored, sterile mycelium, as well as Geotrichum candidum, Acremonium murorum (A. murorum var. felina), Alternaria alternata, Aspergillus candidus, A. versicolor, Cladosporium cladosporioides, Rhizomucor racemosus, Mucor hiemalis, Fusarium (F. oxysporum, Fusarium sp.), and Penicillium spp. These fungi survive for a long time in the earthworm’s digestive environment. Investigation of the functional characteristics and role in the host organism is required to confirm the symbiotic status of the microorganisms associated with the earthworm digestive tract.     

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.     

Effects of an Ecosystem Engineer on Belowground Movement of Microarthropods

Ecosystem engineers affect other species by changing physical environments. Such changes may influence movement of organisms, particularly belowground where soil permeability can restrict dispersal. We investigated whether earthworms, iconic ecosystem engineers, influence microarthropod movement. Our experiment tested whether movement is affected by tunnels (i.e., burrows), earthworm excreta (mucus, castings), or earthworms themselves. Earthworm burrows form tunnel networks that may facilitate movement. This effect may be enhanced by excreta, which could provide resources for microarthropods moving along the network. Earthworms may also promote movement via phoresy. Conversely, negative effects could occur if earthworms alter predator-prey relationships or change competitive interactions between microarthropods. We used microcosms consisting of a box connecting a “source” container in which microarthropods were present and a “destination” container filled with autoclaved soil. Treatments were set up within the boxes, which also contained autoclaved soil, as follows: 1) control with no burrows; 2) artificial burrows with no excreta; 3) abandoned burrows with excreta but no earthworms; and 4) earthworms (Lumbricus rubellus) present in burrows. Half of the replicates were sampled once after eight days, while the other half were sampled repeatedly to examine movement over time. Rather than performing classical pairwise comparisons to test our hypotheses, we used AIC_c to assess support for three competing models (presence of tunnels, excreta, and earthworms). More individuals of Collembola, Mesostigmata, and all microarthropods together dispersed when tunnels were present. Models that included excreta and earthworms were less well supported. Total numbers of dispersing Oribatida and Prostigmata+Astigmata were not well explained by any models tested. Further research is needed to examine the impact of soil structure and ecosystem engineering on movement belowground, as the substantial increase in movement of some microarthropods when corridors were present suggests these factors can strongly affect colonization and community assembly.