不同蚯蚓采样方法对比研究

蚯蚓种类组成和丰度变化是反映农业管理措施对土壤质量影响的重要指标。为研究利用芥末悬液等方法进行蚯蚓样品采集的有效性和准确性,寻求合理有效的蚯蚓种群特征调查方法,在我国东北玉米连作黑土上分别运用手拣法、福尔马林溶液驱赶法、芥末+水悬液及芥末+乙酸悬液驱赶法,以及驱虫剂与手拣法相结合的方法进行蚯蚓样品采集,将上述7种方法测得的蚯蚓丰度、生物量及年龄组成信息进行了系统分析。结果表明,福尔马林溶液驱赶法不结合手拣法使用时测得的蚯蚓丰度及生物量远远低于其他方法,尤其幼年蚯蚓比例偏低,并不能反映真实的蚯蚓种群特征。使用芥末悬液做驱虫剂有效性明显提高,尤其芥末+乙酸悬液,测得的蚯蚓丰度比手拣法高22.3%。福尔马林溶液结合手拣法后有效性大大提高,测得的蚯蚓丰度和生物量分别是其单独使用时的19.1和9.3倍,但相对其他驱虫剂方法仍高估了成年蚯蚓的比例;芥末+水悬液和芥末+乙酸悬液方法结合手拣法后收集到蚯蚓的数量分别提高了67.8%和89.1%,平均个体重量分别是原来的1.8和1.3倍,说明芥末悬液不会杀死小个体的幼年蚯蚓,但少部分大个体蚯蚓及洞穴不与地表相接的蚯蚓可能并不能被其驱赶至地表,结合手拣法后可以弥补这个缺陷。芥末+乙酸悬液结合手拣法收集到的成年及幼年蚯蚓数量和生物量均显著高于其他方法(P<0.05),既能有效地驱赶小个体和幼年蚯蚓,又可以收集到洞穴不与地表相接的土栖型蚯蚓,因此可以作为真实有效地反映蚯蚓种群特征的采样方法之一。     

Autofluorescence in eleocytes of some earthworm species

Immunocompetent cells of earthworms, coelomocytes, comprise adherent amoebocytes and granular eleocytes (chloragocytes). Both cell populations can be expelled via dorsal pores of adult earthworms by exposure to an electric current (4.5 V) for 1 min. Analysis by phase contrast/fluorescence microscopy and flow cytometry demonstrated that eleocyte population of several species exhibits a strong autofluorescence. A high percentage (11-35%) of autofluorescent eleocytes was recorded in Allolobophora chlorotica, Dendrodrilus rubidus, Eisenia fetida, and Octolasion sp. (O. cyaneum, O. tyrtaeum tyrtaeum and O. tyrtaeum lacteum). In contrast, autofluorescent coelomocytes were exceptionally scarce (less than 1%) in representative Aporrectodea sp. (A. caliginosa and A. longa) and Lumbricus sp. (L. castaneus, L. festivus, L. rubellus, L. terrestris). Thus, this paper for the first time describes profound intrinsic fluorescence of eleocytes in some--but not all--earthworm species. The function (if any) and inter-species differences of the autofluorescent coelomocytes still remain elusive.     

Tissue distribution and characterization of cholinesterase activity in six earthworm species

To validate cholinesterase activity as a biomarker of pesticide exposure, we characterized the tissue distribution (whole body, nervous tissue and crop/gizzard), activity at two seasons of cholinesterase in six different species of earthworms collected in an unpolluted field: Lumbricus terrestris, Lumbricus castaneus, Aporrectodea nocturna, Aporrectodea caliginosa, Allolobophora chlorotica and Aporrectodea rosea. The major part of total cholinesterase activity was found in the nervous tissue while activity in crop/gizzard was weak. The level of the total cholinesterase activity was stable for each species considered throughout the year (spring and autumn). Lumbricus species exhibited three-fold higher specific activity than the others (0.086+/-0.015 U mg(-1) and 0.235+/-0.036 U mg(-1) for Allolobophora or Aporrectodea, and Lumbricus species respectively). This stability of the base level makes cholinesterase activity a useful biomarker for monitoring effects of pesticide under natural conditions. Cholinesterase activity was characterized using different substrates and inhibitors. It seems likely that the cholinesterases are acetylcholinesterases in most species investigated as they preferentially hydrolyzed acetylthiocholine and were inhibited by eserine, but not by tetraisopropyl pyrophoramide (iso-OMPA). Characterization of cholinesterase from Allolobophora chlorotica is uncertain and it cannot be classified as a true AChE.     

STUDIES ON THE RELATIONSHIPS BETWEEN EARTHWORMS AND SOIL FERTILITY: I. BIOLOGICAL STUDIES IN THE FIELD

Changes in the seasonal activity of several species of earthworms have been followed in a permanent pasture field for 18 months, 1945–6. The two soil conditions which chiefly determine activity are temperature and moisture. Other factors are the occurrence of an obligatory diapause in the two species Allolobophora nocturna and A. longa , and changes in population in A. chlorotica and Lumbricus terrestris.
Soil temperature and soil moisture also determined the weight of soil thrown up in the form of wormcasts during autumn, winter and spring. It is suggested that only two species, A. nocturna and A. longa , are responsible for wormcasts and that the other four common species present play little or no part in this activity.
At Rothamsted it was found that the previous agricultural history of the field is an important factor in determining the fauna. Old permanent pasture is characterized by a high percentage of A. nocturna and a rather lower percentage of A. caliginosa. Ploughing old permanent pasture and reseeding to grass after 1 or 2 years arable reduces the proportions of A. nocturna and A. caliginosa and increases that of Eisenia rosea. Arable fields have A. chlorotica as the dominant species and pasture fields 2–7 years grass after many years of arable fanning still show a high percentage of A. chlorotica and a low proportion of A. nocturna.
A survey of the Carse of Stirling, Scotland, showed that soil type is also an important factor in determining the earthworm fauna. A. caliginosa was the dominant species on the three soil types studied, but the subdominant species varied. On clay soil, A. longa was subdominant; on loam, A. longa and L. rubellus ; on sandy soil, A. longa, L. rubellus and A. chlorotica. The dominant species of pasture land at Rothamsted, A. nocturna , was not found at Stirling.     

Seasonal variation in the abundance, biomass and biodiversity of earthworms in soils contaminated with metal emissions from a primary smelting works

1. Earthworms (Annelida: Oligochaeta) were sampled on four occasions (spring, summer, autumn and winter) at 14 sites along two transects from a primary lead/ zinc/cadmium smelting works at Avonmouth, UK.
2. Total abundance and biomass of earthworms decreased with proximity to the smelter. No worms were collected from the two sites closest to the factory (<0·6 km) and catches were significantly lower than controls at a further five sites (<3 km).
3. Seasonal composition of sampled communities differed only for summer with lower numbers of individuals and species collected at all sites. Reduced catches in the summer sample is a response to drought.
4. Species richness was lowest at sites close to the factory. For example, worms such as Aporrectodea caliginosa (Savigny) and Allolobophora chlorotica (Savigny) that were dominant at relatively clean sites further from the smelter are absent from the most contaminated soils.
5. Reduced species richness resulted in lower Shannon–Weiner diversity and higher Berger–Parker dominance. Multivariate cluster analysis for spring, summer and winter indicated that sites could be split into three groups based upon relative species composition. In autumn, two clusters were identified.
6. The absence of sensitive species from sites close to the smelting works supports the inclusion of earthworms as a key group in a terrestrial prediction and classification scheme for quantifying the effects of pollutants on soil biodiversity. However, sampling should be carried out in spring or autumn to obtain an accurate picture of community structure.     

Glutathione S-transferases in earthworms (Lumbricidae).

Glutathione S-transferase activity (EC 2.5.1.18) was demonstrated in six species of earthworms of the family Lumbricidae: Eisenia foetida, Lumbricus terrestris, Lumbricus rebellus, Allolobophora longa, Allolobophora caliginosa and Allolobophora chlorotica. Considerable activity was obtained with 1-chlorl-2,4-dinitrobenzene and low activity with 3,4-dichloro-1-nitrobenzene, but no enzymic reaction was detectable with sulphobromophthalein 1,2-epoxy-3-(p-nitrophenoxy)propane of trans-4-phenylbut-3-en-2-one as substrates. Enzyme prepartations from L. rubellus and A. longa were the most active, whereas A. chlorotica gave the lowest activity. The ratio of the activities obtained with 1-chloro-2,4-dinitrobenzene and 3,4-cichloro-1-nitrobenzene was very different in the various species, but no phylogenetic pattern was evident. Isoelectric focusing gave rise to various activity peaks as measured with 1-chloro-2,4-dinitrobenzene as a substrate, and the activity profiles of the species examined appeared to follow a taxonomic pattern. The activity of Allolobophora had the highest peak in the alkaline region, whereas that of Lumbricus had the highest peak in the acid region. Eisenia showed a very complex activity profile, with the highest peak ne pH 7. As determined by an enzymic assay, all the species contained glutathione, on an average about 0.5 mumol/g wet wt. Conjugation with glutathione catalysed by glutathione S-transferases may consequently be an important detoxification mechanism in earthworms.     

Studies on Artioposthia triangulata (Dendy) (Tricladida: Terricola), a predator of earthworms

Observations had linked the disappearance of earthworms from a grass field to the presence of Artioposthia triangulata. An experiment demonstrated that this land planarian could severely and quickly reduce numbers of four earthworm species. Two different approaches to sampling A. triangulata were investigated on a grassland site. The first used four different trap types (wood, ceramic tile, 'corriboard' plastic and 5 mm polystyrene beneath a ceramic tile) measuring 15 cm x 15 cm which were compared over a period of 18 wk. The second used one, or two, applications of dilute formalin solution to quadrats followed by counting the residual planarians by hand-sorting the soil beneath the quadrat to a depth of 30 cm. The polystyrene traps were the most effective for detecting the presence of A. triangulata. No planarians were ever found by hand-sorting after two formalin applications and it was concluded that formalin sampling provided a good estimate of the population density. Counts under traps were not related to absolute estimates of population densities so polystyrene type traps should only be used for detection of the planarians. A multiple regression relationship of A. triangulata weight over time in 5% formalin is presented and allows the original liveweight to be calculated from the weight in preservative. Planarian and earthworm population densities in eight fields were sampled and the planarians were found to be randomly distributed. Counts of planarians in 0.25 m² quadrats were not related to numbers of earthworms but overall field population densities were. It is concluded that A. triangulata is a severe threat to the earthworm populations of Northern Ireland.     

Opening a can of worms: unprecedented sympatric cryptic diversity within British lumbricid earthworms

Earthworms play a major role in many aspects of soil fertility, food web ecology and ecosystem functioning, and hence are frequently the subjects of, for example, ecological and toxicological research. Our aim was to examine the genetic structure of common earthworm species, to identify cryptic lineages or species that may be distinct ecotypes or biotypes (and hence confound current research based upon morphotypes) and to try to explain the massive cryptic diversity that eventually emerged. We demonstrated that species such as Allolobophora chlorotica, Aporrectodea longa, Aporrectodea rosea and Lumbricus rubellus all comprise highly divergent lineages with species-level divergence at the mitochondrial cytochrome oxidase I (COI) gene. In Allo. chlorotica alone, we found 55 haplotypes for COI, with 35 of these being found in pink and 20 in green morph worms. There were no cases of the two colour morphs sharing COI haplotypes. Phylogenetic analyses of mitochondrial COI and 16S genes showed the presence of five highly divergent lineages, suggesting the presence of multiple cryptic species within Allo. chlorotica. There was no clear geographical pattern to lineage distribution and many populations were polymorphic for both mitochondrial DNA lineage and colour morph. Amplified fragment length polymorphism results, based on two primer combinations, were broadly congruent with mitochondrial DNA results with one significant exception. Despite showing over 14% divergence at COI, amplified fragment length polymorphism markers showed that the two green morph lineages may be interbreeding and therefore represent a single taxon. The cryptic diversity revealed by these results has profound consequences for all areas of earthworm research.     

Effects of Soil Nutrient Heterogeneity and Earthworms on Aboveground Biomass of Experimental Plant Communities

Soil nutrients are commonly heterogeneously distributed and earthworms are one of the most common soil organisms. While effects of both soil nutrient heterogeneity and earthworms have been well studied, their interactive effect on plant community productivity has rarely been tested. In a greenhouse experiment, we constructed experimental plant communities by sowing seed mixtures of four grasses, two legumes and two forbs in either a heterogeneous soil consisting of low and high nutrient soil patches or a homogeneous soil where the low and high nutrient soil patches were evenly mixed. The earthworm Eisenia fetida was either added to these soils or not. Aboveground biomass of the whole communities, grasses and legumes did not differ between the homogeneous and heterogeneous soils or between the soils with and without earthworms. However, soil nutrient heterogeneity reduced aboveground biomass of forbs, and such an effect did not interact with earthworms. In response to soil heterogeneity and earthworms, biomass ratio of the three functional groups showed similar patterns as that of their biomass. At the patch level, aboveground biomass of the whole community, grasses and legumes were greater in the high than in the low nutrient soil patches within the heterogeneous soil. A similar pattern was found for the forbs, but this was only true in the absence of earthworms. Our results suggest that soil nutrient heterogeneity and earthworms may not influence aboveground biomass of plant communities, despite the fact that they may modify the growth of certain plant functional groups within the community.     

Biogeographical zonation of the western Iberian peninsula on the basis of the distribution of earthworm species

We used TWINSPAN and CANOCO to investigate the biogeography of the western Iberian Peninsula on the basis of the distribution of earthworms among ninety areas of Galicia, Portugal, Asturias, León, Zamora, Salamanca, Cáceres, Badajoz and Huelva. This part of Iberia is clearly divisible into a Eurosiberian Region in the north and west and a Mediterranean Region in the south and east. For earthworm faunas, however, the limit between these regions is considerably further south than is conventionally accepted on the basis of Rivas Martinez's (1987) vegetation-type classification. Species character- istic of the Eurosiberian Region include Dendrobaena madeirensis, D. octaedra, Allolobophora caliginosa and various species of the genus Lumbricus . Within this region the earthworm fauna of Asturias is clearly distinct from that of Galicia and northern Portugal. Species characteristic of the Mediterranean Region include Allolobophora caliginosa , A. chlorotica and A. rosea , but inventories from this region are most easily identified by the absence or scarcity of those species characteristic of the Eurosiberian Region. Many areas show intermediate characteristics between the two major regions, and in some cases one region 'intrudes' into the other.     

Invasion of exotic earthworms into ecosystems inhabited by native earthworms

The most conspicuous biological invasions in terrestrial ecosystems have been by exotic plants, insects and vertebrates. Invasions by exotic earthworms, although not as well studied, may be increasing with global commerce in agriculture, waste management and bioremediation. A number of cases has documented where invasive earthworms have caused significant changes in soil profiles, nutrient and organic matter dynamics, other soil organisms or plant communities. Most of these cases are in areas that have been disturbed (e.g., agricultural systems) or were previously devoid of earthworms (e.g., north of Pleistocene glacial margins). It is not clear that such effects are common in ecosystems inhabited by native earthworms, especially where soils are undisturbed. We explore the idea that indigenous earthworm fauna and/or characteristics of their native habitats may resist invasion by exotic earthworms and thereby reduce the impact of exotic species on soil processes. We review data and case studies from temperate and tropical regions to test this idea. Specifically, we address the following questions: Is disturbance a prerequisite to invasion by exotic earthworms? What are the mechanisms by which exotic earthworms may succeed or fail to invade habitats occupied by native earthworms? Potential mechanisms could include (1) intensity of propagule pressure (how frequently and at what densities have exotic species been introduced and has there been adequate time for proliferation?); (2) degree of habitat matching (once introduced, are exotic species faced with unsuitable habitat conditions, unavailable resources, or unsuited feeding strategies?); and (3) degree of biotic resistance (after introduction into an otherwise suitable habitat, are exotic species exposed to biological barriers such as predation or parasitism, “unfamiliar” microflora, or competition by resident native species?). Once established, do exotic species co-exist with native species, or are the natives eventually excluded? Do exotic species impact soil processes differently in the presence or absence of native species? We conclude that (1) exotic earthworms do invade ecosystems inhabited by indigenous earthworms, even in the absence of obvious disturbance; (2) competitive exclusion of native earthworms by exotic earthworms is not easily demonstrated and, in fact, co-existence of native and exotic species appears to be common, even if transient; and (3) resistance to exotic earthworm invasions, if it occurs, may be more a function of physical and chemical characteristics of a habitat than of biological interactions between native and exotic earthworms.     

Acidovorax-like symbionts in the nephridia of earthworms

Dense accumulations of bacteria in the excretory organs, nephridia, were first described more than 75 years ago in members of the annelid family Lumbricidae (earthworms). These nephridial symbionts were assumed to play a role in the degradation of proteins in the excretory fluid for nitrogen recycling. In the present study, the phylogenetic affiliation of the nephridial bacteria of the earthworms Lumbricus terrestris, Aporrectodea tuberculata, Octolasion lacteum and Eisenia foetida was resolved. The 16S rRNA gene sequences of the symbionts formed a monophyletic cluster within the genus Acidovorax. Similarity between symbiont sequences from different host species was 95.5-97.6%, whereas similarity was> 99% between symbiont sequences from individuals of the same species. Densely packed bacteria were detected in the ampulla of the nephridia by fluorescence in situ hybridization (FISH) using Acidovorax-specific oligonucleotide probes. No other bacterial cells could be found by FISH, although a few sequences other than Acidovorax had been found by PCR and cloning. These results suggest that the Acidovorax-earthworm symbiosis is a stable, host-specific association that has evolved from a common bacterial ancestor. Given the close phylogenetic relationship of the symbionts to proteolytic, free-living Acidovorax species, they may indeed play a role in protein degradation during nitrogen excretion by earthworms.     

Local-scale correlates of native and non-native earthworm distributions in juniper-encroached tallgrass prairie

European and Asian earthworms have invaded much of North America with profound impacts to soils, plant communities, and animal populations. However, few studies have assessed local-scale correlates of earthworm distributions, and most invasive earthworm research has occurred in northern forests. Additionally, despite several studies showing facilitative relationships between invasive earthworms and invasive plants, no research has assessed a potential facilitative interaction between earthworms and woody plants encroaching into prairies. We conducted the first assessment of factors influencing local-scale distributions of native and non-native earthworms for the U.S. Great Plains in a tallgrass prairie-woodland mosaic experiencing eastern redcedar (Juniperus virginiana) encroachment. We documented both native and non-native earthworms, including non-native species from Eurasia (Aporrectodea spp.) and South America (Family Ocnerodrilidae). Native and non-native earthworm distributions were strongly correlated, yet local-scale predictors of distribution also differed between the groups. Native earthworms were more likely to occur near roads and in areas with moist soils. Contrary to expectation, we found no evidence that non-native earthworms occurred more frequently in areas with eastern redcedar-encroachment; instead, non-native earthworms were most likely to occur in tallgrass prairie. Our results suggest that, within prairies and woodlands of the Great Plains, native and non-native earthworms occur most frequently near roadways and in locations with moist soil. Because the few approaches for controlling invasive earthworms are only likely to be feasible on a small scale, findings from such local-scale studies are important for directing management to reduce earthworm impacts on biodiversity and ecosystem services.     

Effects of nickel, zinc, and lead-contaminated soil on burrowing rate and coelomocytes of the earthworm, Allolobophora chlorotica

We have shown previously that stubby worms Allolobophora chlorotica are sensitive to environmental stress, including metal-polluted soil. In order to discern the mechanisms of this sensitivity, adult (clitellate) Al. chlorotica were exposed in the laboratory to soil samples soaked with water (control) or Ni (1 and 2 mg/kg), Zn (1.25 and 2.5 g/kg) or Pb (5 and 10 g/kg) chlorides. Worms avoided contact with metal contaminants by prolonging burrowing time in metal-soaked samples, especially in the case of lead. Higher concentrations of the investigated metals were lethal for worms. During a 3 week exposure to lower metal concentrations, nickel and lead readily accumulated in the bodies of worms while zinc was efficiently regulated. However, body weights and numbers of non-invasively retrieved free coelomocytes (consisting of amoebocytes and riboflavin-loaded eleocytes) were significantly lower only in zinc-exposed worms. We assume that zinc regulation in worm bodies is more energy-demanding than nickel or lead bioaccumulation, thus this might be responsible for inhibition of the body gain and diminution of immunocompetent cells in zinc-exposed earthworms. Alternatively, missing free coelomocytes may actually be involved in Zn trafficking and removal through nephridia and/or in the formation of multicellular brown bodies, since metal can unbalance host/bacteria relationships.     

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。     

Denitrifying Bacteria in the Earthworm Gastrointestinal Tract and In Vivo Emission of Nitrous Oxide (N(inf2)O) by Earthworms

Earthworms (Lumbricus rubellus and Octolasium lacteum) and gut homogenates did not produce CH(inf4), and methanogens were not readily culturable from gut material. In contrast, the numbers of culturable denitrifiers averaged 7 x 10(sup7) and 9 x 10(sup6) per g (dry weight) of gut material for L. rubellus and O. lacteum, respectively; these values were 256- and 35-fold larger than the numbers of culturable denitrifiers in the soil from which the earthworms were obtained. Anaerobically incubated earthworm gut homogenates supplemented with nitrate produced N(inf2)O at rates exceeding that of soil homogenates. Furthermore, living earthworms emitted N(inf2)O under aerobic conditions, and N(inf2)O emission was stimulated by acetylene. For earthworms collected from a mildly acidic (pH 6) beech forest soil, the rates of N(inf2)O emission for earthworms and soil averaged 884 and 2 pmol per h per g (fresh weight), respectively. In contrast, for earthworms collected from a more acidic (pH 4.6) oak-beech forest soil, N(inf2)O emission by earthworms and soil averaged 145 and 45 pmol per h per g (fresh weight), respectively. Based on the extrapolation of this data, earthworms accounted for an estimated 16 and 0.25% of the total N(inf2)O produced at the stand level of these beech and oak-beech forest soils, respectively.     

长期不同施肥下太湖地区黄泥土蚯蚓的多样性、蛋白质含量与氨基酸组成的变化

水稻土在不同施肥管理下的土壤质量及其生态系统功能的变化是当前土壤学和农业生态学的关注领域。对太湖地区一个15a的长期不同施肥处理下稻田进行了蚯蚓群落的调查采样,并测定了蚯蚓蛋白质含量和氨基酸组成。供试水稻土中共检出7种蚯蚓。长期不同施肥措施影响了蚯蚓的群落结构,单施化肥下农田蚯蚓的种类和数量明显减少,多样性指数和丰富度明显降低,且蚯蚓总氨基酸的含量和大部分种类氨基酸含量降低,同时蚯蚓中分子量小于25kd的蛋白质含量降低,而分子量33kd附近的蛋白质含量明显增加;相反,长期化肥配合秸杆还田和配施猪粪趋向于提高蚯蚓蛋白质含量或氨基酸含量。看来．农田中蚯蚓不但在种群变化上,而且在体内生命活性物质组成上均响应稻田不同施肥措施下的土壤环境变化。     

Changes in diversity, protein content, and amino acid composition of earthworms from a paddy soil under different long-term fertilizations in the Tai Lake Region, China

Influence of the agricultural management practices on soil quality and the ecosystem functioning has been an increasing concern in soil science and ecology with sustainable agriculture. This study deals with the changes of soil earthworm communityfrom a paddy soil under different long-term fertilizations. The soil earthworms were collected and counted from different fertilizer treated plots in the field after the rape harvest in May 2004, and their taxonomic groups were determined under a binocular stereoscope at the laboratory. The body of the earthworm (Metaphire californica) was crushed by a cell crusher to collect protein, and the protein molecules with different sizes were analyzed by electrophoresis. Furthermore, the Metaphire californica collected was hydrolyzed and the aliquots were subject to an amino acid auto-analyzer. The results showed that totally seven species of earthworms were recognized in the paddy field with the number varying with different fertilization treatments. The structure of earthworm communities was dramatically affected by the fertilization practice. Under chemical fertilization only, both the number of earthworm species and the quantity of individuals were significantly smaller than those under other treatments, or even than those under no fertilization. Furthermore, there was an obvious decrease in the total amino acid and the contents of most individual amino acids of Metaphire californica under chemical fertilization only, compared with those under the combined fertilization of chemical and organic fertilizers. Although chemical fertilizers in combination with rice straw return increased earthworm amino acid content, long-term pig manure application tended to increase earthworm protein content. As a molecular footprint, long-term chemical fertilization caused a reduction in the content of protein with MW less than 25 kd, but a significant increase in that of protein with molecule size around 33 kd. Our study demonstrated that different fertilizations affected not only earthworm population but also diversity and richness in the paddy soil after 16 years of treatment, and that long-term chemical fertilization may impact the soil animal community and, thus, influence the paddy ecosystem functioning for yield stability. This study implicated that not only the community structure but also the amino acid metabolism for life functioning of earthworms in cropland soils may pose significant responses to the agricultural management practices.     

Earthworm numbers,biomass and respiratory metabolism in a beech woodland-Wytham Woods,Oxford

Summary The mean annual population density of earthworms was found to be 164.6 m^-2 during a period of detailed study between October 1971 and September 1972. In a year of less detailed study between November 1972 and October 1973 the population density was 117.5 m^-2 (139.8 m^-2 when the type of extraction method was allowed for). Mean biomass densities in the two years of investigation were 41.0 g preserved wet wt m^-2 (1971–1972) and 38.6 (possibly 39.2) g preserved wet wt m^-2 (1972–1973).Comparison of the Brogden's Belt population and biomass densities with those reported from other woodlands indicates that soil type is more important than leaf litter type in determining the numerical abundance of earthworms. Population densities are lower in beechwoods on mor soils, mor soils also support significantly fewer species. As with numbers, mean biomass density in beechwoods on mor soils was significantly lower than that occurring in beechwoods on mull soils; the latter, in turn, being lower than those found in mixed deciduous woods on mull soils. Unlike population density biomass density is influenced by both soil and litter type, this is discussed by reference to mean body weights and food quality as reflected by tannin, nitrogen and carbohydrate content.The annual respiratory metabolism of the Brogden's Belt earthworms was calculated to be between 10.7 and 13.41 O₂ m^-2 a^-1, which is equivalent to between 4.1 and 5.1% of the total soil metabolism. A production/biomass ratio of 0.49–0.58 was estimated, as was a net population efficiency of 22%.     

Nonlinearity of effects of invasive ecosystem engineers on abiotic soil properties and soil biota

Invasions of non‐indigenous species into natural communities are currently rated as one of the most important threats to biodiversity. Particularly exotic ecosystem engineers such as earthworms potentially have profound impacts on community assembly and functioning. We investigated the impact of invasion by the lumbricid earthworms into an aspen forest of the Canadian Rocky Mountains on soil organic matter, microorganisms and microarthropod communities. Building on the results of previous studies in this forest, we expected positive effects of Lumbricus terrestris middens and negative effects of Octolasion tyrtaeum on soil biota (increase and decrease in soil nutrient concentrations, microbial parameters and soil microarthropod density and diversity, respectively). Further, we expected that earthworm effects change with time. Combined results of previous and the present study suggest a wavelike colonization pattern for Dendrobaena octaedra and O. tyrtaeum and that indeed the impact of earthworms on soil biota changed with time, likely due to changes in earthworm density. Unexpectedly, L. terrestris middens neither affected soil abiotic nor soil biotic properties. By contrast and in contrast to our hypothesis, carbon and nitrogen concentration and C‐to‐N ratio in deeper soil layers increased in presence of O. tyrtaeum, thereby likely enhancing nutrient availability for soil microorganisms and microarthropods. Even though the density of this endogeic species was rather low, presence of O. tyrtaeum resulted in increased densities of a number of microarthropod taxa and increased microarthropod diversity. The results suggest that at low density, invasive ecosystem engineers, such as O. tyrtaeum, cause disturbances of intermediate strength thereby beneficially affecting soil microorganisms and most microarthropods. This contrasts earlier effects during the wavelike invasion of O. tyrtaeum into the aspen forest when densities of O. tyrtaeum were high resulting in generally detrimental effects on soil biota. The results emphasize the nonlinearity of earthworm effects on abiotic and biotic soil properties and call for further long‐term investigations.