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PubMed2000

Purification and partial characterisation of tentatively classified acid phosphatase from the earthworm Eisenia veneta.

In order to use leakage of lysosomal acid phosphatase (AP) as a biomarker of stress to earthworms, more information about AP's in earthworms are needed. This paper describes the details about tentatively classified APs in the earthworm Eisenia veneta. Two isoenzymes (enzyme I and II) of acid phosphatase (AP) and one alkaline phosphatase (enzyme III) from the earthworm E. veneta were separated by gel filtration. All three enzymes were further purified and concentrated on a Con A Sepharose 4B column. Enzyme I was inhibited by tartrate, showed an optimal pH range between 4.0 and 5.0 and was assumed to be of lysosomal origin. Enzyme II was the major enzyme showing the highest activity of the three enzymes. It was expected to be a lysosomal AP under physiological conditions. Enzyme II had a molecular mass 113 kDa and was composed of apparently identical polypeptide chains of 36 kDa each. This enzyme was inhibited by tartrate, showed an optimal pH in the range 6.0-7.5 and was slowly degraded at temperatures above 40 degrees C. Enzyme III is not inhibited by tartrate and has a pH-optimum > 9. The subcellular location under physiological conditions was assumed to be the cytosol.

Acid Phosphatase↗

PubMed1998

Cu-Cd interactions in earthworms maintained in laboratory microcosms: the examination of a putative copper paradox.

Earthworms (Lumbricus rubellus) from Ecton (predominantly Cu-contaminated), Shipham (Cd-contaminated) and Dinas Powys (uncontaminated, reference) were maintained in the laboratory on soil from the sampling sites. Two principle exposure protocols were used: (1) a 4-week 'no pre-exposure experiment', where batches of earthworms were maintained on soils from each habitat and (2) a 'pre-exposure experiment' where uncontaminated control worms were maintained on Shipham soil for 4 weeks (the pre-exposure period), and then transferred to Ecton soil for a further 4 weeks, and vice versa. In summary, there were two primary findings. Firstly, exposure of worms to Cd-rich soil (Shipham) before exposure to Cu-rich soil (Ecton) increased the Cu burden and secondly, biochemical analysis of metal binding proteins in analogous earthworm populations indicated that Cu and Cd, but not Zn, are bound to a metallothionein homologue. These observations are consistent with the hypothesis that Cu does not induce thionein synthesis efficiently in earthworm tissues, but may bind avidly to Cd-induced thionein by substituting for Cd in the molecule because Cu-thionein is a more stable complex.

Animals↗

PubMed2001

Water-soluble luminal contents of the gut of the earthworm Lumbricus terrestris L. and their physiological significance.

In the post-gizzard gut of the earthworm Lumbricus terrestris, distinguishing the functions of the luminal epithelium from those of the chloragogenous tissue has been hindered by the close apposition of these two tissues. Moreover, both tissues may have different functions from the anterior to the posterior of the animal. We analyzed the gut luminal contents of L. terrestris so as to gain a better understanding of the function of the luminal epithelium. The intestine was divided into four regions from anterior to posterior, and the water-soluble portion of the luminal contents of these four regions was analyzed for protease and amylase activity, calcium and ammonium ions, and protein. The same four regions of the gut wall were analyzed for glutamate dehydrogenase (GDH) and serine dehydratase (SDH) to determine their location with reference to the site of ammonia production. We observed high levels of proteases, amylase, protein and calcium ions in the gut luminal contents of the first two regions, and a significant decline of all four variables in region III. Conversely, ammonia was low in the gut contents of regions I and II but rose sharply in region III, which was also the region to which the tissue enzymes GDH and SDH were localized. The ammonia content of earthworm casts was observed to be much higher than that of the surrounding soil. These data are presented as partial evidence for the proposal that the excretory ammonia produced by feeding earthworms is a product of the luminal epithelium of region III of the gut. It is also proposed that ammonia and calcium may function as ion-exchangers in the absorptive function of the earthworm gut.

Animals↗

PubMed1992

Ceramide glycanase from the earthworm, Lumbricus terrestris.

Ceramide glycanase (CGase) is an enzyme that cleaves the linkage between the sugar chain and the ceramide. To make this enzyme readily available, we have developed a simple method for preparing it from the earthworm, Lumbricus terrestris. The method involves Bio-Gel A-0.5m, octyl-Sepharose and p-aminophenylthiogalactoside-agarose column chromatography. By gel filtration, the molecular mass of earthworm CGase was found to be 43.7 kDa. With ganglioside GM1 as substrate, the optimal pH of this enzyme was found to be between pH 3.5 and 4.0. Earthworm CGase hydrolyses glycolipids only in the presence of a detergent. Among various bile salts tested, sodium cholate was found to be the most effective in stimulating the hydrolysis of GM1 by this enzyme. Earthworm CGase released intact glycan chains from various glycosphingolipids in which the glycan chain is linked to the ceramide through a beta-glucosyl linkage. It also detached glycan chains from lactosyldialkylglycerol and alkyl-beta-lactosides.

Animals↗

PubMed2003

Effects of transgenic Bt corn litter on the earthworm Lumbricus terrestris.

A 200-day study was carried out to investigate the impact of transgenic Bacillus thuringiensis (Bt) corn on immature and adult Lumbricus terrestris in the field and in the laboratory. Another objective of this study was to develop test methods that could be used for standard testing of the impact of transgenic plants on different earthworm species in the field and in the laboratory. For this purpose two different experiments were involved, a laboratory experiment with adult L. terrestris and a field experiment with immature L. terrestris. No lethal effects of transgenic Bt corn on immature and adult earthworms were observed. Immature L. terrestris in the field had a very similar growth pattern when fed either (Bt+) or (Bt-) corn litter. No significant differences in relative weights of (Bt+) and (Bt-) corn-fed adult L. terrestris were observed during the first 160 days of the laboratory trial, but after 200 days adult L. terrestris had a significant weight loss of 18% of their initial weight when fed (Bt+) corn litter compared to a weight gain of 4% of the initial weight of (Bt-) corn-fed earthworms. Further studies are necessary to see whether or not this difference in relative weight was due to the Bt toxin or other factors discussed in the study. Degradation of Cry1Ab toxin in corn residues was significantly slower in the field than at 10 degrees C in the laboratory. Enzyme-linked immunosorbent assay results indicated that earthworms in both experiments were exposed to the Bt toxin throughout the whole experimental time.

Animals↗

PubMed2003

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.

Animals↗

PubMed2002

[Study of the effect of earthworm Lumbricus terrestris on the speciation of heavy metals in soils].

Evaluation of the effect of earthworm Lumbricus terrestris on the speciation of copper and cadmium was carried out on two types of soils with a high metallic contamination due to municipal wastes spreading. The concentrations of total dissolved metals were higher in the soil containing earthworms. This increase was larger for the soil submitted to disturbance by earthworms for a long time (3 months). The main chemical species in the lixiviates of all type of soils including controls, were labile forms of cadmium with 52 to 87% and stable forms of copper which represents 67 to 95% of total concentration of dissolved metal. In the solid phase, there was a slight transfer of cadmium and copper from the oxidizable fraction into the exchangeable and acid soluble fractions. This suggests that soil disturbance by earthworms increases the mobility of these metals particularly cadmium. On the contrary copper appears in lixiviates as non labile organic complexes. Analysis of the whole results showed differences between soils as regards the mobility of the metals studied, which reflected the role of the mains physico-chemical characteristics (pH, C.E.C. and total calcareous content).

Animals↗

PubMed2005

Bioavailability of pentachlorophenol to earthworms (Eisenia fetida) in artificially contaminated soils.

The bioaccumulation of pentachlorophenol in earthworms (Eisenia fetida) was studied for two artificially contaminated soils (S1 and S2). The uptake kinetics of pentachlorophenol (PCP) in earthworms increased quickly within the initial 10 days. This was followed by a nearly steady state for the next 20 days that fit with the equilibrium partitioning model. The correlation coefficients were 0.812 and 0.715 for S1 and S2, respectively. The average biota-to-soil accumulation factor of PCP in S1 was 0.51 +/- 0.09, whereas that of S2 was 0.79 +/- 0.12. There was a significant correlation between log Csoil and log Cworm, demonstrating the validity of the equilibrium partitioning model. The bioavailability of PCP was assessed by chemical extraction methods. The results demonstrated a close correlation between extractable amounts of PCP freshly added in soils and those in earthworms. With increasing residence time of PCP in soil, there was a progressively smaller amount of PCP assimilated by the earthworms. In contrast, the amount extracted by Soxhlet extraction did not show a similar decline. However, the extractable amount of PCP by methanol and methanol-water (1:1) significantly decreased over 440 days. Compared with the methanol-water (1:1) extraction method, the methanol extraction method was preferred to the prediction of the bioavailability of PCP in aged soils.

Animals↗

PubMedDate not supplied

Evaluation of tissue and cellular biomarkers to assess 2,4,6-trinitrotoluene (TNT) exposure in earthworms: effects-based assessment in laboratory studies using Eisenia andrei.

The lysosomal neutral red retention time (NRRT) assay, a biomarker for lysosomal membrane stability, and the total immune activity (TIA) assay, a measure of non-specific immune system activity, were used in laboratory studies to assess the toxic effects of 2,4,6-trinitrotoluene (TNT) on earthworms (Eisenia andrei) in vivo. The results were compared with the concentration of TNT and its metabolites in earthworm tissue, as well as standard sublethal toxicity endpoints including growth (i.e. weight change) and reproduction effects from previously published studies. Filter paper experiments indicated a significant decrease in NRRT at >or=1.8 micro g TNT cm(-2), whereas sublethal (weight loss) and lethal effects to earthworms were detected at >or=3.5 and 7.1 micro g TNT cm(-2), respectively. Experiments in artificial soil showed that NRRT effects could be detected at lower TNT concentrations (>or=55 mg TNT kg(-1) soil dry weight) compared with other sublethal endpoints (effects on growth and reproduction). The TIA biomarker did not significantly respond to TNT. Copper (as CuSO4, filter paper contact tests) and 2-chloroacetamide (soil tests), which were used as reference toxicants, also decreased the NRRT. The use of the NRRT assay linked with tissue concentrations of TNT metabolites in earthworms was identified as a potentially appropriate biomarker approach for TNT exposure assessment under laboratory conditions and a novel tool for effects-based risk assessment.

Animals↗

PubMed1963

SOLUBLE EARTHWORM CUTICLE COLLAGEN: A POSSIBLE DIMER OF TROPOCOLLAGEN.

When soluble earthworm cuticle collagen molecules are subjected to the shearing forces of a flow birefringence instrument, they are broken into particles approximately half the original size. The broken particles resemble vertebrate tropocollagen molecules in their hydrodynamic properties, in levorotatory powers, and in their appearance in the electron microscope. Most significantly, the broken earthworm particles form ordered aggregates similar to the segmented-long-spacing aggregations formed by vertebrate tropocollagen. These phenomena are explained by the suggestion that earthworm collagen molecules are dimers of tropocollagen-like particles. On this basis, an explanation is presented for the lack of striations in the gross collagen fibrils of earthworm cuticle.

Animals↗

PubMed2004

Molecular cloning and characterization of cDNA encoding fibrinolytic enzyme-3 from earthworm Eisenia foetida.

The earthworm fibrinolytic enzyme-3 (EFE-3, GenBank accession No: AY438622), from the earthworm Eisenia foetida, is a component of earthworm fibrinolytic enzymes. In this study, cDNA encoding the EFE-3 was cloned by RT-PCR. The cDNA contained an open reading frame of 741 nucleotides, which encoded a deduced protein of 247 amino acid residues, including signal sequences. EFE-3 showed a high degree of homology to earthworm (Lumbricus rebullus) proteases F-III-1, F-III-2, and bovine trypsin. The recombinant EFE-3 was expressed in E. coli as inclusion bodies, and the gene encoding the native form of EFE-3 was expressed in COS-7 cells in the medium. Both the refolding product of inclusion bodies and the secreted protease could dissolve the artificial fibrin plate.

Amino Acid Sequence↗

PubMed1994

The complete sequence of a 40-kDa actin-modulating protein from the earthworm Lumbricus terrestris.

The complete primary structure of a 40-kDa actin-modulating protein from the earthworm Lumbricus terrestris is presented. A muscle-specific cDNA library of the earthworm was constructed and screened with a specific DNA probe obtained by polymerase chain reaction considering information from peptide sequencing. A full-length clone with a coding region of 1098 bp was isolated. The deduced polypeptide sequence of 366 amino acids (41457 Da) reveals the segmental structure typical of both the 40-kDa and 80-kDa actin-modulating proteins. Prominent similarities to the 80-kDa protein gelsolin especially exist with respect to the first segment and to the C-terminal segment. The comparatively high nucleation efficiency of the earthworm actin modulator is probably determined by its third segment which seems to enable the earthworm actin modulator to bind a second G-actin molecule more tightly than other previously described 40-kDa modulators.

Actins↗

PubMed1995

Comparative assessment of the aerobic and anaerobic microfloras of earthworm guts and forest soils.

Aerobic and anaerobic microbial potentials of guts from earthworms (Lumbricus rubellus Hoffmeister and Octolasium lacteum (Oerl.)) collected from a beech forest were evaluated. On the basis of enumeration studies, microbes capable of growth under both aerobic and anaerobic conditions were more numerous in the earthworm intestine than in the beech forest soil from which the worms were obtained. The intestine of worms displayed nearly equivalent aerobic and anaerobic microbial growth potentials; in comparison, soils displayed greater aerobic than anaerobic microbial growth potentials. Hence, the ratio of microbes capable of growth under obligately anaerobic conditions to those capable of growth under aerobic conditions was higher with the worm intestine than with the soil. Process level studies corroborated these population differentials: (i) under anaerobic conditions, worm gut homogenates consumed glucose, cellobiose, or ferulate more readily than did soil homogenates; and (ii) under aerobic conditions, worm gut homogenates consumed cellobiose or oxygen more readily than did soil homogenates. Collectively, these results reinforce the general concept that the earthworm gut is not microbiologically equivalent to soil and also suggest that the earthworm gut might constitute a microhabitat enriched in microbes capable of anaerobic growth and activity.

Journal Article↗

PubMed1999

Earthworm egg capsules as vectors for the environmental introduction of biodegradative bacteria.

Earthworm egg capsules (cocoons) may acquire bacteria from the environment in which they are produced. We found that Ralstonia eutropha (pJP4) can be recovered from Eisenia fetida cocoons formed in soil inoculated with this bacterium. Plasmid pJP4 contains the genes necessary for 2,4-dichlorophenoxyacetic acid (2,4-D) and 2, 4-dichlorophenol (2,4-DCP) degradation. In this study we determined that the presence of R. eutropha (pJP4) within the developing earthworm cocoon can influence the degradation and toxicity of 2,4-D and 2,4-DCP, respectively. The addition of cocoons containing R. eutropha (pJP4) at either low or high densities (10(2) or 10(5) CFU per cocoon, respectively) initiated degradation of 2,4-D in nonsterile soil microcosms. Loss of 2,4-D was observed within the first week of incubation, and respiking the soil with 2,4-D showed depletion within 24 h. Microbial analysis of the soil revealed the presence of approximately 10(4) CFU R. eutropha (pJP4) g-1 of soil. The toxicity of 2,4-DCP to developing earthworms was tested by using cocoons with or without R. eutropha (pJP4). Results showed that cocoons containing R. eutropha (pJP4) were able to tolerate higher levels of 2,4-DCP. Our results indicate that the biodegradation of 2, 4-DCP by R. eutropha (pJP4) within the cocoons may be the mechanism contributing to toxicity reduction. These results suggest that the microbiota may influence the survival of developing earthworms exposed to toxic chemicals. In addition, cocoons can be used as inoculants for the introduction into the environment of beneficial bacteria, such as strains with biodegradative capabilities.

2,4-Dichlorophenoxyacetic Acid↗

PubMed2002

Molecular and culture-based analyses of prokaryotic communities from an agricultural soil and the burrows and casts of the earthworm Lumbricus rubellus.

The microbial populations in no-till agricultural soil and casts of the earthworm Lumbricus rubellus were examined by culturing and molecular methods. Clone libraries of the 16S rRNA genes were prepared from DNA isolated directly from the soil and earthworm casts. Although no single phylum dominated the soil library of 95 clones, the largest numbers of clones were from Acidobacteria (14%), Cytophagales (13%), Chloroflexi (8%), and gamma-Proteobacteria (8%). While the cast clone library of 102 clones was similar to the soil library, the abundances of several taxa were different. Representatives of the Pseudomonas genus as well as the Actinobacteria and Firmicutes increased in number, and one group of unclassified organisms found in the soil library was absent in the cast library. Likewise, soil and cast archaeal 16S rRNA gene libraries were similar, although the abundances of some groups were different. Two hundred and thirty aerobic bacteria were also isolated on general heterotrophic media from casts, burrows, and soil. The cast isolates were both phenotypically and genotypically different from the soil isolates. The cast isolates were more likely to reduce nitrate, grow on acetate and Casamino Acids, and utilize fewer sugars than the soil isolates. On the basis of their ribotypes, the cast isolates were dominated by Aeromonas spp. (28%), which were not found in the soil isolates, and other gamma-Proteobacteria (49%). In contrast, the soil isolates were mostly Actinobacteria (53%), Firmicutes (16%), and gamma-Proteobacteria (19%). Isolates obtained from the sides of earthworm burrows were not different from the soil isolates. Diversity indices for the collections of isolates as well as rRNA gene libraries indicated that the species richness and evenness were decreased in the casts from their levels in the soil. These results were consistent with a model where a large portion of the microbial population in soil passes through the gastrointestinal tract of the earthworm unchanged while representatives of some phyla increase in abundance.

Agriculture↗

PubMed2000

Ontogenetic scaling of burrowing forces in the earthworm Lumbricus terrestris.

In hydrostatic skeletons, it is the internal fluid under pressure surrounded by a body wall in tension (rather than a rigid lever) that enables the stiffening of the organism, the antagonism of muscles and the transmission of force from the muscles to the environment. This study examined the ontogenetic effects of body size on force production by an organism supported with a hydrostatic skeleton. The earthworm Lumbricus terrestris burrows by forcefully enlarging crevices in the soil. I built a force-measuring apparatus that measured the radial forces as earthworms of different sizes crawled through and enlarged pre-formed soil burrows. I also built an apparatus that measured the radial and axial forces as earthworms of different sizes attempted to elongate a dead-end burrow. Earthworms ranging in body mass m(b) from hatchlings (0.012 g) to adults (8.9 g) exerted maximum forces (F, in N) during active radial expansion of their burrows (F=0.32 m(b)(0.43)) and comparable forces during axial elongation of the burrow (F=0.26 m(b)(0.47)). Both these forces were almost an order of magnitude greater than the radial anchoring forces during normal peristalsis within burrows (F=0.04 m(b)(0.45)). All radial and axial forces scaled as body mass raised to the 2/5 power rather than to the 2/3 power expected by geometric similarity, indicating that large worms exert greater forces than small worms on an absolute scale, but the difference was less than predicted by scaling considerations. When forces were normalized by body weight, hatchlings could push 500 times their own body weight, while large adults could push only 10 times their own body weight.

Animals↗

PubMed2006

Acephaline gregarine parasites (Monocystis sp.) are not transmitted sexually among their lumbricid earthworm hosts.

The precise transmission mode(s) of acephaline gregarines in their earthworm hosts has long been questioned, yet a rigorous experimental evaluation of sexual transmission is currently lacking. That Monocystis sp., a common gregarine parasite of the earthworm Lumbricus terrestris, infects the sexual organs of its host is suggestive of sexual transmission. Considering the divergent evolutionary consequences of various modes of transmission, excluding or proving sexual transmission in this host-parasite system is critical to fully understanding it. We cultured uninfected earthworms from cocoons and subsequently mated them to either an infected or uninfected partner (from the wild). We then compared these individuals with an orally infected group, which were infected using a newly developed gavage (oral injection) method. Our data have unambiguously established that (1) horizontal sexual transmission does not play a significant role in the transmission of Monocystis sp., and (2) oral transmission through the soil is likely the principal mode of transmission between earthworms. This finding is important to models of mate-choice because infection avoidance does not appear to drive mating decisions. Finally, we further report a successful and relatively simple method to obtain infection-free individuals, which can subsequently be infected via oral gavage and used in empirical studies.

Animals↗

PubMed2004

Acute and chronic toxicity of the new explosive CL-20 to the earthworm (Eisenia andrei) exposed to amended natural soils.

Monocyclic nitramine explosives such as 1,3,5-trinitro-1,3,5-triazacyclohexane (RDX) and octahydro-1,3,5,7-tetranitro-1,3,5,7-tetrazocine (HMX) are toxic to a number of ecological receptors, including earthworms. The polycyclic nitramine CL-20 (2,4,6,8,10,12-hexanitro-2,4,6,8,10,12-hexaazaisowurtzitane) is a powerful explosive that may replace RDX and HMX, but its toxicity is not known. In the present study, the lethal and sublethal toxicities of CL-20 to the earthworm (Eisenia andrei) are evaluated. Two natural soils, a natural sandy forest soil (designated RacFor2002) taken in the Montreal area (QC, Canada; 20% organic carbon, pH 7.2) and a Sassafras sandy loam soil (SSL) taken on the property of U.S. Army Aberdeen Proving Ground (Edgewood, MD, USA; 0.33% organic carbon, pH 5.1), were used. Results showed that CL-20 was not lethal at concentrations of 125 mg/kg or less in the RacFor2002 soil but was lethal at concentrations of 90.7 mg/kg or greater in the SSL soil. Effects on the reproduction parameters such as a decrease in the number of juveniles after 56 d of exposure were observed at the initial CL-20 concentration of 1.6 mg/kg or greater in the RacFor2002 soil, compared to 0.2 mg/kg or greater in the SSL soil. Moreover, low concentrations of CL-20 in SSL soil (approximately 0.1 mg/kg; nominal concentration) were found to reduce the fertility of earthworms. Taken together, the present results show that CL-20 is a reproductive toxicant to the earthworm, with lethal effects at higher concentrations. Its toxicity can be decreased in soils favoring CL-20 adsorption (high organic carbon content).

Adsorption↗