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PubMed1997

The leech excitatory peptide, a member of the GGNG peptide family: isolation and comparison with the earthworm GGNG peptides.

A member of the GGNG peptide family was isolated from Hirudo nipponia (leech). GGNG peptides had only been isolated previously from earthworms. The C-terminus structure of the leech peptide, LEP (leech excitatory peptide), was -Gly-Gly-Asn-amide, while that of the earthworm peptides, EEP (earthworm excitatory peptide), was -Gly-Gly-Asn-Gly. LEP exerted 1000-fold more potent activities on leech gut than did EEP-2. On the other hand, EEP-2 was 1000-fold more potent than LEP on the crop-gizzard of the earthworm. Analog peptides of LEP and EEP-2 were synthesized, and the myoactive potency of each analog on the leech and earthworm tissues was compared.

Amino Acid Sequence↗

PubMed2002

Method for determining toxicologically relevant cadmium residues in the earthworm Eisenia fetida.

We investigated a method to isolate toxicologically relevant Cd in earthworms (Eisenia fetida) exposed in a 14-d Cd bioaccumulation study. A procedure involving acid insoluble ash (AIA) content was combined with homogenization and centrifugation techniques to divide total earthworm Cd burdens into supernatant (metallothionein-bound), pellet (toxicologically active), and soil-associated Cd fractions. Whereas the supernatant fraction of the earthworm digests increased linearly throughout the exposure period (from approximately 0 to 3.59 mmol/kg), the pellet fraction reached a steady-state concentration (95% CI) of 1.2 (0.9-1.4) mmol/kg, suggesting the ability of the metallothionein detoxification system to sequester incoming Cd. The AIA method was useful for correcting earthworm Cd concentrations for ingested soil-associated Cd and observing soil ingestion, which was suppressed in Cd-spiked artificial soil (1.06 (0.57-1.55)%) compared to controls (17.25 (14.36-20.15)%). These methods may be useful in investigating soil ingestion and Cd uptake and detoxification in earthworms.

Animals↗

PubMed1984

Characterization of vomeronasally-mediated response-eliciting components of earthworm wash-II.

Garter snakes reliably spend more time and tongue flick more frequently at a dish containing surface washings of earthworm (EW) than at a control dish containing water or sodium chloride. In a series of experiments EW was examined to further characterize and purify the snake-attracting component(s). Earthworm wash boiled for up to two hours did not lose its snake-attracting activity, but EW boiled for four hours was not discriminated from a control substance. The active components of EW were completely precipitated by 90 and 100% saturated ammonium sulfate solutions. The molecular weight of the active components, as determined by dialysis, is larger than 67,000 daltons. Earthworm wash was separated into two peaks on either G-75 or AcA 44. The larger peak (F2) contains snake-attracting material of molecular weight greater than 67K. The smaller peak material (F4) is inactive as a snake-attractant and contains material of approximate molecular weight of 3K. Biological activity of EW covaried with Lowry (protein) assay and Dubois (carbohydrate) assay values and 280 nm absorbance. The response eliciting components of EW are not extractable from earthworms at temperatures below 60 degrees C, but can be extracted at 60 degrees C (for one minute) from the worms or the soil that forms the bedding for the earthworms.

Ammonium Sulfate↗

PubMed2003

Comparative efficacy of three epigeic earthworms under different deciduous forest litters decomposition.

An experiment was conducted during 1998-1999, in a deciduous forest located in the semi-arid tropics of central India, to evaluate the suitability of different forest litters as food material for the tropical epigeic earthworms i.e. Eisenia fetida (Savigny), Perionyx excavatus (Perrier) and Dicogaster bolaui (michaelsen). The aim was to examine the influence of these earthworms on the decomposition processes of three types of forest litters i.e. Tectona grandis (teak), Madhuca indica (mahua) and Butea monosperma (palas), on the maintenance of quality in a vermicomposting system, and to assess the effect of applications of in situ prepared vermicomposts on the growth of forest trees. The results indicated that T. grandis litter was the most suitable food material for the earthworms possibly because it contained high reserves of mineral nutrients. Comparisons of the survival and reproduction rates of the three epigeic earthworm species indicated that a higher reproduction rate was maintained for E. fetida compared to P. excavatus and D. bolaui in the decomposition of these forest litters. The rates of growth and population increases of E. fetida approximately doubled after 12 weeks of litter decomposition. The litter decomposition process was associated strongly with the quality of the materials and their chemical composition. Irrespective of earthworm inoculations, the levels of available nutrient such as NH(4)-N, NO(3)-N, available P and K increased significantly (p</=0.05) in the order T. grandis litter compost>M. indica litter compost>B. monosperma litter compost. The mature decomposed litter had lower C/N ratios (11.3-24.8:1), water-soluble carbon (0.30-0.58%), water-soluble carbohydrates (0.35-0.71%) and larger cation exchange capacity/total organic carbon ratios than the values in the parent forest litter. The lignin content increased with maturation with a concomitant decrease in cellulose resulting in higher lignin/cellulose ratios. Application of all three vermicomposts to forest trees significantly improved their heights and diameters over those of control trees, although the increases were lower than those resulting from the chemical fertilizer applications. However, soil biological activities i.e. soil respiration, soil microbial biomass carbon and dehydrogenase activity were greater by application of vermicomposts over that after application of inorganic fertilizer in a new plantation of T. grandis.

Animals↗

PubMed2004

Do earthworms mobilize fixed zinc from ingested soil?

A wide range of organisms inhabit the soil and has to deal with soil-bound metals. The bioavailable fraction of metals may be estimated explicitly using the isotopic dilution technique. In the present paper, we evaluated the isotopic exchange technique for assessing the bioavailability of soil Zn (using 65Zn) to earthworms. To validate the technique, the worms were first exposed to various 65Zn levels, and errors due to soil entrained in the gut were evaluated. This exposure indicated no effect of gamma-radiation on growth (wet weight gain) of the organisms and that depuration of the earthworms minimized errors in labile pools determined by isotopic dilution. Our study further showed that the earthworms accessed 55-65% of the total Zn in the soil. The labile pool for the earthworms Eisenia andrei was similar to that for the plant Lactuca sativa, indicating that earthworms and plants to a large extent access the same fraction of soil Zn. Hence, the isotopic dilution technique has the potential to assess biologically available pool of Zn in soils. As lettuce is not known to significantly mobilize nonlabile metals in soil, this study indicates that Zn uptake by E. andrei is predominantly via the exchangeable pools (possibly the soil pore water) rather than dissolution of Zn held within soil particles or within soil organic matter or other food sources.

Animals↗

PubMed2005

Ligand arsenic complexation and immunoperoxidase detection of metallothionein in the earthworm Lumbricus rubellus inhabiting arsenic-rich soil.

Although earthworms have been found to inhabit arsenic-rich soils in the U.K., the mode of arsenic detoxification is currently unknown. Biochemical analyses and subcellular localization studies have indicated that As3+-thiol complexes may be involved; however, it is not known whether arsenic is capable of inducing the expression of metallothionein (MT) in earthworms. The specific aims of this paper were (a) to detect and gain an atomic characterization of ligand complexing by X-ray absorption spectrometry (XAS), and (b) to employ a polyclonal antibody raised against an earthworm MT isoform (w-MT2) to detect and localize the metalloprotein by immunoperoxidase histochemistry in the tissues of earthworms sampled from arsenic-rich soil. Data suggested that the proportion of arsenate to sulfur-bound species varies within specific earthworm tissues. Although some arsenic appeared to be in the form of arsenobetaine, the arsenic within the chlorogogenous tissue was predominantly coordinated with S in the form of -SH groups. This suggests the presence of an As::MT complex. Indeed, MT was detectable with a distinctly localized tissue and cellular distribution. While MT was not detectable in the surface epithelium or in the body wall musculature, immunoperoxidase histochemistry identified the presence of MT in chloragocytes around blood vessels, within the typhlosolar fold, and in the peri-intestinal region. Focal immunostaining was also detectable in a cohort of cells in the intestinal wall. The results of this study support the hypothesis that arsenic induces MT expression and is sequestered by the metalloprotein in certain target cells and tissues.

Animals↗

PubMed2002

Arsenic-speciation in arsenate-resistant and non-resistant populations of the earthworm, Lumbricus rubellus.

Arsenic speciation was determined in Lumbricus rubellus Hoffmeister from arsenic-contaminated mine spoil sites and an uncontaminated site using HPLC-MS, HPLC-ICP-MS and XAS. It was previously demonstrated that L. rubellus from mine soils were more arsenate resistant than from the uncontaminated site and we wished to investigate if arsenic speciation had a role in this resistance. Earthworms from contaminated sites had considerably higher arsenic body burdens (maximum 1,358 mg As kg-1) compared to the uncontaminated site (maximum 13 mg As kg-1). The only organo-arsenic species found in methanol/water extracts for all earthworm populations was arsenobetaine, quantified using both HPLC-MS and HPLC-ICP-MS. Arsenobetaine concentrations were high in L. rubellus from the uncontaminated site when concentrations were expressed as a percentage of the total arsenic burden (23% mean), but earthworms from the contaminated sites with relatively low arsenic burdens also had these high levels of arsenobetaine (17% mean). As arsenic body burden increased, the percentage of arsenobetaine present decreased in a dose dependent manner, although its absolute concentration rose with increasing arsenic burden. The origin of this arsenobetaine is discussed. XAS analysis of arsenic mine L. rubellus showed that arsenic was primarily present as As(III) co-ordinated with sulfur (30% approx.), with some As(v) with oxygen (5%). Spectra for As(III) complexed with glutathione gave a very good fit to the spectra obtained for the earthworms, suggesting a role for sulfur co-ordination in arsenic metabolism at higher earthworm arsenic burdens. It is also possible that the disintegration of As(III)-S complexes may have taken place due to (a) processing of the sample, (b) storage of the extract or (c) HPLC anion exchange. HPLC-ICP-MS analysis of methanol extracts showed the presence of arsenite and arsenate, suggesting that these sulfur complexes disintegrate on extraction. The role of arsenic speciation in the resistance of L. rubellus to arsenate is considered.

Adaptation, Physiological↗

PubMed2002

Metabolism of 4-fluoroaniline and 4-fluorobiphenyl in the earthworm Eisenia veneta characterized by high-resolution NMR spectroscopy with directly coupled HPLC-NMR and HPLC-MS.

1. Little is known about metabolism of xenobiotics by earthworms, despite their importance in soil ecotoxicity testing. Normal earthworms and earthworms treated with antibiotics to ensure inhibition of gut microflora were exposed to two model xenobiotic compounds, 4-fluoroaniline and 4-fluorobiphenyl, to determine which metabolites were produced, and whether the pattern of metabolism was affected by the presence of microbial transformation ability. 2. (19)F-NMR spectroscopy detected the number and relative proportions of metabolites and directly coupled HPLC-(1)H-NMR spectroscopy and HPLC-MS then identified the metabolites. 3. Despite uptake, no metabolism of 4-fluorobiphenyl was observed at any stage, which appears to be a consequence of the lack of oxidative Phase I metabolic activity of the earthworms towards this substrate. In contrast, 4-fluoroaniline exhibited dose-dependent metabolism. At high doses (leading to mortality within 24 h) one predominant metabolite was observed, which was identified as the N-beta-glucoside conjugate. At lower dose levels, the predominant metabolite was the gamma-glutamyl conjugate, although the glucoside and another as yet unidentified metabolite were also detected. 4. The inhibition of gut microflora did not have any influence on metabolism. The study represents the first evidence for glucoside and glutamyl conjugation as a pathway for xenobiotic metabolism in earthworms.

Aniline Compounds↗

PubMed1999

Feasibility of vermicomposting dairy biosolids using a modified system to avoid earthworm mortality.

A laboratory study was conducted to examine the feasibility of vermicomposting dairy biosolids (dairy sludge), either alone or with either of the bulking agents-cereal straw or wood shavings, using the epigeic earthworm-Eisinea andrei. Earthworms added directly to these three substrates died within 48 hours. A system was developed to overcome the toxic effect of unprocessed dairy biosolids. The substrates were placed over a layer of vermicomposted sheep manure into which the earthworms were inoculated. Within two weeks, all earthworms were within the upper layer of substrate. Compared to sheep manure which is a favourable substrate for vermicomposting, the three substrates containing dairy biosolids were more effective in supporting earthworm growth and reproduction. The final products obtained after 63 days of vermicomposting had 39-53% less organic carbon than the initial substrates. Organic fractionation indicated that vermicomposting increased the stability of the materials to biological decomposition. The vermicomposts obtained from the three substrates with dairy biosolids had low heavy metal contents and electrical conductivities, and did not inhibit plant growth when compared with a commercial vermicompost in a bioassay.

Animals↗

PubMed2001

Evaluating environmental hazards of land applying composted diazinon using earthworm bioassays.

Environmental hazards resulting from land application of composted pesticide residue have not been rigorously evaluated. This study was conducted to examine the toxicity of a composted pesticide residue using earthworms (Eisenia foetida Savigny) as a microinvertebrate model in a soil bioassay system. Diazinon, which was used in these experiments as a test pesticide, was removed from simulated rinsate (wastewater) by sorption onto peat moss. Following the rinsate clean-up phase, diazinon-laden peat moss was placed into bioreactors and composted for either 30 or 60 days. Earthworms were then exposed to soil amended with the composted material. Mortality and symptomatic effects characteristic of acetylcholinesterase inhibition, including weight loss, reduced burying ability and curling, occurred in earthworms exposed to soil amended with either uncomposted or 30-day composted diazinon, but not in those exposed to soil amended with 60-day composted diazinon. The amount of solvent-extractable diazinon from compost was not directly related to acute earthworm toxicity based on the selected criteria. These results indicated a reduction in diazinon bioavailability during latter 30 d of composting that did not correspond to a reduction in solvent-extractable diazinon concentrations. Measuring symptomatic effects of xenobiotics as described in this study may increase the sensitivity and diagnostic ability of earthworm bioassays.

Acetylcholinesterase↗

PubMed2006

Influence of ultraviolet radiation on selected physiological responses of earthworms.

The purpose of this study was to investigate the adverse effects of ultraviolet (UV) radiation on earthworms. Earthworms that crawl out of the soil may die within a few hours after sunrise. This study shows that UV exposure can be lethal. In general, UV-B had a stronger damaging effect than UV-A. Different species of earthworms had different tolerances to UV exposure. In this study, Pontoscolex corethrurus showed the highest tolerance of the three tested species to UV radiation, while Amynthas gracilis was the most sensitive. UV radiation induced both acute and chronic responses. The acute response, which occurred immediately on or after UV exposure, was characterized by the appearance of abnormally strong muscle contractions, including S-shaped movements and jumping behavior, possibly caused by bad coordination between the circular and longitudinal muscles. The chronic response included damage to the skin and muscle cells, which resulted in a high mortality rate. Oxygen consumption by A. gracilis was significantly decreased after exposure to UV-A or UV-B. Since the circulation in earthworms is mediated by muscle contraction and the skin is the main organ of respiration, it is reasonable to expect that abnormal muscle contraction and a damaged epithelium could cause suffocation. Because of their sensitive responses, we propose that some earthworms, such as A. gracilis, could serve as a new model for studying UV-induced photodamage.

Animals↗

PubMed1998

Ontogenetic scaling of hydrostatic skeletons: geometric, static stress and dynamic stress scaling of the earthworm lumbricus terrestris

Soft-bodied organisms with hydrostatic skeletons range enormously in body size, both during the growth of individuals and in the comparison of species. Therefore, body size is an important consideration in an examination of the mechanical function of hydrostatic skeletons. The scaling of hydrostatic skeletons cannot be inferred from existing studies of the lever-like skeletons of vertebrates and arthropods because the two skeleton types function by different mechanisms. Hydrostats are constructed of an extensible body wall in tension surrounding a fluid or deformable tissue under compression. It is the pressurized internal fluid (rather than the rigid levers of vertebrates and arthropods) that enables the maintenance of posture, antagonism of muscles and transfer of muscle forces to the environment. The objectives of the present study were (1) to define the geometric, static stress and dynamic stress similarity scaling hypotheses for hydrostatic skeletons on the basis of their generalized form and function, and (2) to apply these similarity hypotheses in a study of the ontogenetic scaling of earthworms, Lumbricus terrestris, to determine which parameters of skeletal function are conserved or changed as a function of body mass during growth (from 0.01 to 8 g). Morphometric measurements on anesthetized earthworms revealed that the earthworms grew isometrically; the external proportions and number of segments were constant as a function of body size. Calculations of static stresses (forces per cross-sectional area in the body wall) during rest and dynamic stresses during peristaltic crawling (calculated from measurements of internal pressure and body wall geometry) revealed that the earthworms also maintained static and dynamic stress similarity, despite a slight increase in body wall thickness in segment 50 (but not in segment 15). In summary, the hydrostatic skeletons of earthworms differ fundamentally from the rigid, lever-like skeletons of their terrestrial counterparts in their ability to grow isometrically while maintaining similarity in both static and dynamic stresses.

Journal Article↗

PubMed1999

Kinematic scaling of locomotion by hydrostatic animals: ontogeny of peristaltic crawling by the earthworm lumbricus terrestris

This study examined the relationship between ontogenetic increase in body size and the kinematics of peristaltic locomotion by the earthworm Lumbricus terrestris, a soft-bodied organism supported by a hydrostatic skeleton. Whereas the motions of most vertebrates and arthropods are based primarily on the changes in the joint angles between rigid body segments, the motions of soft-bodied organisms with hydrostatic skeletons are based primarily on the changes in dimensions of the deformable body segments themselves. The overall kinematics of peristaltic crawling and the dynamic shape changes of individual earthworm segments were measured for individuals ranging in body mass (mb) by almost three orders of magnitude (0.012-8.5 g). Preferred crawling speed varied both within and among individuals: earthworms crawled faster primarily by taking longer strides, but also by taking more strides per unit time and by decreasing duty factor. On average, larger worms crawled at a greater absolute speed than smaller worms (U p2finity mb0.33) and did so by taking slightly longer strides (l p2finity mb0.41, where l is stride length) than expected by geometric similarity, using slightly lower stride frequencies (f p2finity mb-0.07) and the same duty factor (df p2finity mb-0.03). Circumferential and longitudinal body wall strains were generally independent of body mass, while strain rates changed little as a function of body mass. Given the extent of kinematic variation within and among earthworms, the crawling of earthworms of different sizes can be considered to show kinematic similarity when the kinematic variables are normalized by body length. Since the motions of peristaltic organisms are based primarily on changes in the dimensions of the deformable body wall, the scaling of the material properties of the body wall is probably an especially important determinant of the scaling of the kinematics of locomotion.

Journal Article↗

PubMed2001

Stable strontium accumulation by earthworms: a paradigm for radiostrontium interactions with its cationic analogue, calcium.

The accumulation of stable strontium and its chemical analogue calcium by four species of earthworm, representing three distinct ecophysiological groups inhabiting celestite (SrSO4)-rich natural soils, was investigated. An increase in soil strontium concentration over a four-orders-of-magnitude range was accompanied by an increase in earthworm tissue strontium concentration. In contrast, no relationship between soil and tissue calcium concentrations was apparent, indicating that this essential cation is strongly regulated within earthworm tissues but that nonessential strontium burdens are not physiologically regulated. Concentration factors for strontium ranged from 0.1 to 1.42, with an inverse relationship between this parameter and soil strontium concentration. The use of concentration factors and discrimination coefficients lead to the conclusion that earthworms discriminate against strontium accumulation at increased soil calcium concentrations, such that strontium would be predicted to be more efficiently accumulated by worms inhabiting acidic, calcium-poor soils. Although few consistent trends in strontium and calcium concentration were apparent between earthworm species, the species with the highest calcium turnover (i.e., the litter-inhabiting Lumbricus rubellus) typically accumulated higher concentrations of strontium than representatives of contrasting ecophysiological classes. These findings are considered in the context of the distribution, retention, and cycling of radiostrontium in soils.

Animals↗

PubMed2000

Absence of obvious short-term impact of the nematode-trapping fungus Duddingtonia flagrans on survival and growth of the earthworm Aporrectodea longa.

The nematode-trapping fungus Duddingtonia flagrans may be used in biological control of parasitic nematode larvae in faeces of domestic host animals after feeding the hosts with fungal chlamydospores. In this experiment a possible undesirable fungal impact on earthworms, of the species Aporrectodea longa, was investigated. As earthworms eat animal faeces, D. flagrans may come into contact with earthworms both in their alimentary tract and on their body surface. However during the experimental period of 20 days, when earthworms were living in soil and eating cattle faeces that were heavily infested with viable chlamydospores of D. flagrans there were no indications of internal or external mycosis among the earthworms.

Animals↗

PubMed2002

Effects of earthworm (Eisenia fetida) and wheat (Triticum aestivum) straw additions on selected properties of petroleum-contaminated soils.

Current bioremediation techniques for petroleum-contaminated soils are designed to remove contaminants as quickly and efficiently as possible, but not necessarily with postremediation soil biological quality as a primary objective. To test a simple postbioremediation technique, we added earthworms (Eisenia fetida) or wheat (Triticum aestivum) straw to petroleum land-farm soil and measured biological quality of the soil as responses in plant growth, soil respiration, and oil and grease (O&G) and total petroleum hydrocarbon (TPH) concentrations. Results indicated that plant growth was greater in earthworm-treated land-farm soil. Furthermore, addition of wheat straw resulted in greater total respiration in all soils tested (land-farm soil, noncontaminated reference soil, and a 1:1 mixture of land-farm and reference soils). We observed a 30% increase in soil respiration in straw-amended oily soil, whereas respiration increased by 246% in straw-amended reference soil. Much of the difference between oily and reference soils was attributable to higher basal respiration rates of nonamended oily soil compared to nonamended reference soil. Addition of earthworms resulted in greater total respiration of all soil and straw treatments except two (the land-farm and the 1:1 mixture soil treatments without straw). Straw and earthworm treatments did not affect O&G or TPH concentrations. Nevertheless, our findings that earthworm additions improved plant growth and that straw additions enhanced microbial activity in land-farm soil suggest that these treatments may be compatible with plant-based remediation techniques currently under evaluation in field trials, and could reduce the time required to restore soil ecosystem function.

Animals↗

PubMed2002

Fate and sublethal effects of isoproturon on mature earthworm (Lumbricus terrestris L.).

This study was conducted to investigate the effects of isoproturon in mature earthworm (Lumbricus terrestris L.) under laboratory condition. Earthworms were exposed to soils contaminated with different concentrations for various duration. Residues were monitored in soil and earthworms after 7, 15, 30, 45, and 60 days of exposure to different isoproturon concentrations. Acute toxicity of isoproturon was determined together with growth rate and total soluble protein content of worms. These parameters were related to isoproturon concentration in soil and earthworms. No lethal effect of isoproturon was observed even at the highest concentration tested (1.4 g/kg soil) after 60 days after treatment. Residues of isoproturon have caused a significant reduction of the growth rate. Additionally a reduction of total soluble protein was observed in all treated worms. Decrease of isoproturon concentration in soil was slow and depended on herbicide initial concentration. In the worms, it increased during the first 15 days and decreased thereafter. This study is suggesting the use of the growth rate of earthworms as biomarker of exposure to isoproturon.

Animals↗

PubMed2003

[Single and binary-combined toxicity of methamidophos, acetochlor and Cu on earthworm Eisenia foetida].

A population of earthworm Eisenia foetida was exposed to single and binary-combined contamination of phaeozem by methamidophos, acetochlor and Cu. The result showed that one of three test chemicals had its toxicity on the earthworm population, and the single toxic sequence of the chemicals was acetochlor > methamidophos > Cu. The values of their LD50 were 0.307, 0.708 and 118.70 mg.kg-1, respectively. The difference was depended on the biological mechanisms of the earthworm population. Acetochlor and Cu in soil could be absorbed by the earthworm population through penetrating through the skin of an earthworm. The result also showed that Cu could swell the toxicity of methamidophos, whether it was in low or high concentration by the binary-combined toxic effect test. Cu in low concentration could decrease the toxicity of acetochlor, but in high concentration, Cu could increase the acetochlor toxicity in soil. Therefore, these three pollutants were dangerous to the ecological security of soil ecosystem and soil-health quality. Furthermore, when the chemicals in same soil environment act one another, they could boost up the potential danger of soil ecosystem contaminated by the three pollutants. The joint toxic effects of various chemicals were in relation to their different concentration combinations in soil.

Animals↗