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PubMed1998

The identification, cloning and characterization of earthworm metallothionein.

Combining standard gel chromatographic techniques and novel molecular methodologies (Directed Differential Display and quantitative PCR), it has been possible to isolate and sequence two isoforms of the first true earthworm metallothionein. Both proteins are characteristically high in cysteine residues and possess no significant aromatic residues. Metal responsiveness was confirmed by determining metallothionein specific expression profiles in earthworms exposed to soils of differing heavy metal concentrations. Analysis of the derived amino acid sequence of isoform 2 identified two putative N-glycosylation signal sequences, suggesting that the two isoforms may have different subcellular distributions and functions. Possible implications for intracellular metal trafficking are discussed.

Amino Acid Sequence↗
PubMed2002

Crystal structure of earthworm fibrinolytic enzyme component a: revealing the structural determinants of its dual fibrinolytic activity.

Earthworm fibrinolytic enzyme component A (EFEa) from Eisenia fetida is a strong fibrinolytic enzyme that not only directly degrades fibrin, but also activates plasminogen. Proteolytic assays further revealed that it cleaved behind various P1 residue types. The crystal structure of EFEa was determined using the MIR method and refined to 2.3A resolution. The enzyme, showing the overall polypeptide fold of chymotrypsin-like serine proteases, possesses essential S1 specificity determinants characteristic of elastase. However, the beta strand at the west rim of the S1 specificity pocket is significantly elongated by a unique four-residue insertion (Ser-Ser-Gly-Leu) after Val217, which not only provides additional substrate hydrogen binding sites for distal P residues, but also causes extension of the S1 pocket at the south rim. The S2 subsite of the enzyme was partially occluded by the bulky side-chain of residue Tyr99. Structure-based inhibitor modeling demonstrated that EFEa's S1 specificity pocket was preferable for elastase-specific small hydrophobic P1 residues, while its accommodation of long and/or bulky P1 residues was also feasible if enhanced binding of the substrate and induced fit of the S1 pocket were achieved. EFEa is thereby endowed with relatively broad substrate specificity, including the dual fibrinolysis. The presence of Tyr99 at the S2 subsite indicates a preference for P2-Gly, while an induced fit of Tyr99 was also suggested for accommodation of bigger P2 residues. This structure is the first reported for an earthworm fibrinolytic enzyme component and serine protease originating from annelid worms.

Amino Acid Sequence↗
PubMed1984

Assembly of erythrocruorin from the earthworm Octolasium complanatum.

The spontaneous assembly of the earthworm erythrocruorin molecule (60 S) from its 1/12 subunits (10 S) obtained by alkaline dissociation is a long debated problem, since the 60 S to 10 S dissociation step has been regarded as essentially irreversible or as only partially reversible when freshly dissociated solutions are used. Erythrocruorin from the earthworm Octolasium complanatum has been reassembled from its 10 S subunits. "Age" of the subunits, pH, and divalent cation concentration are the factors that influence the assembly reaction. Of primary importance is the age of the subunits, i.e. their exposure time to the alkaline dissociating pH. Parallel sedimentation velocity and sodium dodecyl sulfate/polyacrylamide gel electrophoresis experiments on the dissociated and reassembled solutions indicate that two processes take place at alkaline pH values: disulfide exchange and limited proteolysis. These processes, whose relative importance differs in the various preparations, might be responsible for the loss of reassociating capacity of the 10 S subunits. With freshly dissociated subunits, reassembly up to 80% may be achieved at pH 6.2 to 6.5 in the absence of divalent cations; the presence of 25 to 50 mM-Ca2+ renders the reaction essentially pH-independent in the range 6.2 to 8. The effect of Ca2+ is discussed in the light of the presence of structure-stabilizing binding sites for divalent cations at the 10 S intersubunit's contact regions.

Animals↗
PubMed2002

Acute toxicity of chlorophenols to earthworms using a simple paper contact method and comparison with toxicities to fresh water organisms.

An acute toxicity test of chlorophenols on earthworms (Eisenia fetida) was performed using a simple paper contact method proposed by OECD testing guideline no. 207, that were applied as an earthworm toxicity test. The median lethal concentration, EC50, had significant correlation with logP(ow) (1-octanol/water partition coefficient) of the chemicals. The toxicity of chlorophenols on E. fetida was compared with toxicities for other species: an algae (Selenostrum capricornutum), a crustacean (Daphnia magna), and a fish (Oryzias latipes). It was found that the toxicity of chlorophenols was almost same for E. fetida and for fresh water organisms. These results suggest the possibility of drawing correlations between the effects of pollutants on living things in different environments, fresh water and soil.

Animals↗
PubMed2002

Chemical assays of availability to earthworms of polycyclic aromatic hydrocarbons in soil.

The bioavailable concentration of an organic pollutant is less than the concentration determined by vigorous extraction of soil. Because bioavailability varies with the particular compound, soil, and aging time, an assay for bioavailability is needed. Three methods were tested: extraction with a 25% aqueous solution of tetrahydrofuran (THF), 95% ethanol, and C18 membranes. Evaluations were conducted with a mixture of four polycyclic aromatic hydrocarbons (PAHs) added to five dissimilar soils and with pyrene as sole PAH added to six soils, and the availability of aged and freshly added compounds was determined. Assimilation by earthworms (Eisenia fetida) was used to assess bioavailability. For extraction with THF and earthworm uptake, the correlation coefficients of determination (R2) for anthracene, chrysene, pyrene, and benzo(a)pyrene added as a mixture exceeded 0.85. The R2 values for assays with the C18 membrane were 0.77 or higher. The values for pyrene added alone were 0.710 and 0.823, respectively. R2 values for assays with ethanol often exceeded 0.87, but lower values were sometimes obtained. We suggest that such solvent or solid-phase extractions may be useful in assessing PAH bioavailability.

Animals↗
PubMed2002

Effect of alpha2M on earthworm fibrinolytic enzyme III-1 from Lumbricus rubellus.

Though it is known that human alpha2-macroglobulin (alpha2M) inhibits most proteases, the effect of alpha2M has not been investigated on earthworm fibrinolytic enzymes (EFEs) from Lumbricus rubellus, which could be transported from intestine epithelium into blood as an intact molecule (Fan et al., Biochim. Biophys. Acta 1526 (2001) 286). The activity of earthworm fibrinolytic III-1 (EFE-III-1) decreased to 65% when incubated with alpha2M, while it decreased to 30% in plasma under the same conditions. The first order rate of the inactivation of EFE-III-1 with alpha2M was similar to that of fast phase with plasma, indicating that alpha2M may be the inhibitor initially binding to the enzyme in blood. SDS-PAGE showed that incubation of EFE-III-1 with alpha2M a released fragment ( approximately 90 kDa), followed by formation of a high molecular weight complex (approximately 700 kDa). There was a linear relationship between the apparent inhibition rate constant (k1) and [alpha2M], by double reciprocal plot. It was suggested, as described by Tsou (Acta Biochem. Biophys. Sinica 5 (1965) 398) and Tian (Biochem. J. 21 (1982) 1028), that the mechanism of alpha2M/EFE-III-1 interaction could be coincided with a complexing irreversible inhibition. Experiments in both the inactivation and the intrinsic fluorescence showed that alpha2M bound to the enzyme mole by mole equivalently. The intrinsic fluorescence of alpha2M was enhanced with an observable blue shift in emission maxima, suggesting that alpha2M was one of the important inhibitors to EFEs when it absorbed into blood.

Animals↗
PubMed1982

Agglutinins in the earthworm Lumbricus terrestris: naturally occurring and induced.

A naturally occurring hemagglutinin against rabbit and rat erythrocytes is contained in the coelomic fluid of the earthworm Lumbricus terrestris. The hemagglutinin reacts with some chicken and human erythrocytes, but not others, and does not react with the erythrocytes of seven other vertebrates. Hemagglutinins appear in increased amounts within 24 hr after injection of rat, rabbit, horse and sheep erythrocytes, and some chicken and human erythrocytes, and are the highest (approximately four- to sevenfold) with rabbit erythrocytes. The response is brief, and increased or more rapid responses do not occur after multiple injections. Cross reactivity and absorption data indicate a close or possibly identical relationship between agglutinins induced against different erythrocyte types. Effects of heating, enzyme and chemical treatment on induced anti-rabbit erythrocyte agglutinins indicate at least two and perhaps three different agglutinins. Two of the agglutinins are protein, one trypsin-sensitive and the other trypsin-resistant. Agglutinin activity is reduced in the absence of divalent cations. Sensitivity to heat varies with the type of agglutinin. The naturally occurring agglutinin is a protein, trypsin-resistant and unaffected by heating at 100 degrees C for 30 min. These hemagglutinins constitute one of the earthworm's humoral factors that may participate in immune responses.

Absorption↗
PubMed2003

Toxic effects of chlorpyrifos on morphology and acetylcholinesterase activity in the earthworm, Eisenia foetida.

The acute toxicity of chlorpyrifos (O, O-diethyl-O-(3,5,6-trichloro-2-pyridyl) phosphorothioate) was determined in the earthworm, Eisenia foetida. A 48-h contact test as described by the Organization for Economic Cooperation and Development (OECD) guideline 207 was carried out. The LC(50) of chlorpyrifos was 0.063 microg/cm(2). Inhibition of acetylcholinesterase (AChE: EC 3.1.1.7) activity indicated by in vitro neurotoxic potentiality revealed competitive inhibition and altered K(m) values widely in a dose-dependent manner. The K(i) value of chlorpyrifos was 4.20 x 10(-6)M. AChE activity of LC(50)-exposed worms was 62%, 79%, 85%, and 91% inhibited at 12, 24, 36, and 48h, respectively. Scanning electron microscopic studies revealed the morphological abnormalities in the worms. The present study demonstrates a dose- and time-dependent exposure of chlorpyrifos through skin results, morphological abnormalities, and inhibition of AChE in the earthworm, E. foetida.

Acetylcholinesterase↗
PubMed1999

Identification of a coelomic mitogenic factor in Eisenia foetida earthworm.

Coelomic fluid of earthworms Eisenia foetida (Oligochaeta, Annelida) exerts a mitogenic activity on murine splenocytes. Total coelomic fluid was subjected to size-exclusion chromatography and a semi-purified mitogenic fraction (fraction 5) was isolated and further characterized. Both coelomic fluid and the semi-purified fraction 5 block concanavalin A (ConA)-induced spleen cell proliferation but exert a synergistic effect on LPS-triggered spleen cell proliferation. Using a polyclonal antiserum neutralizing the mitogenic activity of the semi-purified fraction 5, a 60-kDa component was identified and named CMF (coelomic mitogenic factor). CMF was found to bind ConA which could account for its ability to inhibit ConA-induced spleen cell proliferation. CMF is present in the coelomic fluid as a trimer of a 20-kDa protein. N-terminal amino acid sequence of monomeric CMF reveals partial sequence homology with phospholipase A2 (PLA2). Moreover, CMF-enriched coelomic fluid fraction 5 exerts phospholipase activity comparable with that of bovine pancreatic PLA2. Our results suggest that coelomic fluid of E. foetida contains a ubiquitous PLA2-like enzyme which might be involved in immune reactions in earthworms such as anti-bacterial mechanisms.

Amino Acid Sequence↗
PubMed2001

Serotonin-induced nitric oxide production in the ventral nerve cord of the earthworm, Eisenia fetida.

Effect of serotonin on nitric oxide (NO) production in the ventral nerve cord (VNC) of the earthworm Eisenia fetida was investigated by a bio-imaging and an electrochemical technique. In the bio-imaging, the spatial pattern of NO production in VNC was visualized using an NO-specific fluorescent dye, diaminofluorescein-2 diacethyl (DAF-2 DA). Application of serotonin (100 microM) increased NO production in VNC by about 65% (P<0.05), compared with basal NO production. The increase was mainly from the nitergic neurons in the ventral side of VNC. In the electrochemical technique, real-time basal and serotonin-induced NO production was estimated with an NO-specific electrode. On the ventral surface of VNC, the estimated basal NO production was stable at 200+/-52 nM, and was transiently augmented to 840+/-193 nM by the addition of 10 microM serotonin. In conclusion, the estimated basal NO production in the earthworm VNC is relatively high compared with other nervous systems earlier reported, and transiently augmented by serotonin. Our results suggest that NO signaling in VNC is involved in neuromodulation by serotonin.

Animals↗
PubMed1984

Mitogenic effect of earthworm (Lumbricus terrestris) coelomic fluid on mouse and human lymphocytes.

We have cultured mouse and human lymphocytes with earthworm (Lumbricus terrestris) coelomic fluid and measured their mitogenic responses. Normal fluid was collected from untreated worms, while induced fluid was harvested from worms injected 24 h earlier with rabbit erythrocytes. At low protein concentrations in the coelomic fluid, human and mouse lymphocytes were significantly activated, as measured by incorporation of 3HTdr. The activation index for induced fluid was approximately 2.5 times that for uninduced fluid. Separation of mouse lymphocytes into B- and T-cell populations revealed that primarily T cells were activated. Normal and induced coelomic fluid contains 12 and 28 electrophoretic bands and agglutinin titers of 32 and 1024, respectively. Addition of agglutinin inhibitors or absorption of agglutinins from coelomic fluid did not alter levels of mitogenic activity, thus the relation between earthworm agglutinins and mitogens is problematic. Techniques designed to separate and purify the agglutinins are in progress to elucidate this point.

Agglutinins↗
PubMed2000

Activity and localisation of the lysosomal marker enzymes acid phosphatase, N-acetyl-beta-D-glucosaminidase, and beta-galactosidase in the earthworms Eisenia fetida and E. veneta.

The present study describes the activity and localisation of three putative lysosomal marker enzymes, acid phosphatase (AP), N-acetyl-beta-D-glucosaminidase (beta-NAG), and beta-galactosidase (beta-Gal), in whole individuals and in distinct parts of the earthworms, Eisenia veneta and Eisenia fetida. Activities of AP and beta-NAG were high in the two species with most of the activity located to the anterior and mid-parts of the worms. The activity of beta-Gal was low in all body regions. We found interspecies difference in the AP activity as E. veneta had significantly higher activity of AP than E. fetida in posterior and mid-parts, as well as in whole individuals. Of the three enzymes tested, AP was the only enzyme located to lysosomes, yielding high latency all over the worms with especially high latency in the coelomic fluids and posterior regions. The lysosomal APs in E. veneta and E. fetida may be utilised as a new biomarker for xenobiotic-induced lysosomal membrane damage in earthworms.

Acetylglucosaminidase↗
PubMed2001

Effects of acclimation temperature on enzymatic capacities and mitochondrial membranes from the body wall of the earthworm Lumbricus terrestris.

Many ectotherms respond to low temperature by adjusting capacities of enzymes from energy metabolism, restructuring membrane phospholipids and modulating membrane fluidity. Although much is known about the temperature biology of earthworms, it is not known to what extent earthworms employ compensatory changes in enzymatic capacities and membrane physical properties after exposure to low temperature. We examined activities of enzymes from glycolysis and central oxidative pathways as well as fluidity and phospholipid fatty acid composition of mitochondrial membranes prepared from the body wall of the temperate oligochaete Lumbricus terrestris after a one month acclimation to 5 degrees and 15 degrees C. No compensation occurs in central pathways of oxidative metabolism since activities of cytochrome-c oxidase and citrate synthase, when measured at a common temperature, are similar for 5 degrees C and 15 degrees C-acclimated animals. In contrast, activity of pyruvate kinase is elevated 1.3-fold after acclimation to 5 degrees C. Mitochondrial membranes display inverse compensation with respect to temperature (membranes from 5 degrees C animals are more ordered than membranes from 15 degrees C animals). Our results, in combination with earlier reports, indicate that routine metabolism in L. terrestris may be maintained at reduced temperatures with little or no change in enzymatic capacities and inverse compensation of mitochondrial membranes.

Adaptation, Physiological↗
PubMed2001

Toxicity of coelomic fluid of the earthworm Eisenia foetida to vertebrates but not invertebrates: probable role of sphingomyelin.

The coelomic fluid (CF) of the earthworm Eisenia foetida exhibits a wide variety of biological activities. We found that the CF was not toxic to 42 species, belonging to seven invertebrate phyla, almost all in aquatic adults and larvae exposed to CF. Eleven teleostean species tested died in 0.2-1% CF mostly between 10 and 120 min and the effects were dose-dependent. Tadpoles of the toad Bufo japonicus formosus died in 0.4-2% CF between 80 and 225 min depending upon size, with larger tadpoles surviving longer. Before dying, all experimental tadpoles developed curled and shrunken tails. The Okinawa tree lizard, soft-shelled turtle, Japanese quail, mouse and rat all died after i.v. injection of CF (above 20 microl/kg). Thus, CF was not toxic to invertebrates, but toxic to vertebrates. After heating, CF lost its toxicity to fish, tadpoles and mice. Both CF and lysenin incubated with sphingomyelin-liposomes (SM-liposomes) were no longer toxic, suggesting the involvement of SM in the toxicity. Lysenin, which is a constituent of CF and known to bind specifically to sphingomyelin, exhibited toxicity similar to that of CF. Thus, lysenin in CF is probably responsible for the toxic effects of CF by binding to SM in vertebrate tissues. The bodies of invertebrates might contain little or no SM, while those of vertebrates do contain SM. The coelomic fluid of the earthworm Pheretima communissima has no toxicity to mouse.

Animals↗
PubMed2005

Parasitism and growth in the earthworm Lumbricus terrestris: fitness costs of the gregarine parasite Monocystis sp.

Parasites inflict fitness costs on their hosts, but often the exact reduction in fitness is not well understood. We investigated the influence of infection by the gregarine genus Monocystis sp. on growth and female investment (cocoon production) of its earthworm host, Lumbricus terrestris. Earthworms (n = 81) were observed in a laboratory setting for 8 months, after which parasite load was determined. The results revealed a significant negative relationship between parasite load and growth, yet no association to cocoon production was found. Although the exact nature, strength, and evolutionary consequence of reduced growth are still unclear, the results are the first indication for a clear, albeit weak effect of Monocystis on host fitness.

Animals↗
PubMed2001

Novel model describing trace metal concentrations in the earthworm, Eisenia andrei.

We developed a novel model describing Eisenia andrei body concentrations for Cd, Cu, Pb, and Zn as a function of pH, metals, and soluble organic carbon (SOC) in soil extracts for potential use in predicting values in contaminated field sites. Data from 17 moderately contaminated Dutch field soils in which earthworms were cultured were used in model development. Model parameters quantify biological phenomena important for metal bioavailability, and soil variables quantify relevant soil chemistry characteristics. Earthworm body concentration (EBC) was modeled so that soil metal soluble at bulk soil pH was considered available for dermal exposure, and gut exposure was due to soil metal in solution near neutral regulated gut pH. The efficiency parameter values indicated that metals are biologically regulated in the following order (most to least): Zn - Cu > Pb > Cd. The values determined for the exposure-route constant indicate that Cd, Cu, and Pb EBCs are almost exclusively (>96%) due to dermal exposure and that only 18% of Zn EBC was due to gut exposure. The minimum healthful EBCs determined were Zn > Cu > Pb > Cd, and the values for Pb and Cd were near zero. The Cu model was normalized by soluble organic carbon to be meaningful. The model was most accurate in describing Zn behavior.

Animals↗
PubMed2004

Cadmium detoxification in earthworms: from genes to cells.

Cadmium pollution has clear ecotoxicological consequences as it is readily bio-available and has a reported tendency to bio-accumulate in soil biota. Understanding the mechanisms of cadmium trafficking pathways within sentinel terrestrial invertebrates, such as the earthworm, is therefore considered to be of importance. Using X-ray microanalysis, quantitative polymerase chain reaction, and immunohistochemical techniques, we were able to demonstrate that the earthworm has the intrinsic capacity to efficiently sequester and compartmentalize cadmium via a metallothionein-mediated trafficking pathway. There is evidence that wMT-2, rather than wMT-1, is the major isoform implicated in the detoxification of cadmium and the identification of three independent wMT-2 loci (totalling over 25 kb of genomic sequence) has revealed a complex genomic organization. Complementary in silico analysis of over 6500 expressed sequence tags has identified a third metallothionein isoform, wMT-3, found to be highly enriched in embryonic tissue. In summary, this paper provides a detailed dissection of the genetic, molecular, and cellular basis of a sophisticated pathway that facilitates the uptake, accumulation, transport, and excretion of cadmium.

Amino Acid Sequence↗
PubMed2005

Effect of sewage-sludge application on concentrations of higher-brominated diphenyl ethers in soils and earthworms.

Polybrominated diphenyl ethers (PBDEs), including octa-decaBDEs, were found in soil and earthworm samples collected in 2000 from three research stations (reference plots and sewage-sludge-amended plots) and two farms (reference and amended/flooded soils) in Sweden. Sewage-sludge amendment at the research stations increased concentrations of all BDE congeners 2- to 13-fold, with the highest increases for BDE-209. Concentrations 100-to 1000-fold higher were seen in contaminated soils at both farms. BDE-209 was the predominant congener in all soils. sigmaPBDE concentrations in worms ranged from 3.1 to 38 000 ng/g lipid weight and were correlated to soil concentrations, including the octa-decaBDEs. Biota-soil accumulation factors declined in the following order: TeBDE > PeBDE > HxBDE > OcBDE > NoBDE > DeBDE, and ranged from 0.3 to 2 for the octa-decaBDEs. Thus, higher-brominated PBDEs, including BDE-209, are bioavailable from soils and accumulate in earthworms, presenting an exposure pathway into the terrestrial food web. High levels found at one farm 20 years after the last use of PBDEs indicate high persistence of PBDEs in soils, including BDE-209. No evidence of photolytic debromination of BDE-209 in soils was seen.

Animals↗