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PubMed1997

Structural characterization of the N-glycans from Echinococcus granulosus hydatid cyst membrane and protoscoleces.

Infection by the tapeworm Echinococcus granulosus in the intermediate host results in the development of a hydatid cyst which contains the protoscoleces within a fluid-filled cavity enclosed by the bilayered cyst membrane. N-glycans were enzymatically released from crude extracts of homogenates of hydatid cyst membranes and protoscoleces and their structures were defined by high sensitivity fast atom bombardment mass spectrometry in conjunction with sequential exoglycosidase digestions. The major N-glycans from the cyst membrane were found to be non-charged structures having complex-type antennae and core fucosylation. The antennae are either truncated at the first N-acetylglucosamine or are extended with beta-galactose to form N-acetyllactosamine (lacNAc). A significant proportion of the lacNAc backbones are capped by alpha-galactose. The resulting Gal alpha-Gal beta-terminal structures may account for the earlier observation that antibodies against the blood group P1 epitope recognise components of hydatid cyst extracts. The complex-type N-glycans identified in the protoscoleces extracts were the same as the neutral structures found in the cyst membrane but a small proportion of high mannose structures and truncated di- and trimannosyl core structures were also identified. Sialylated N-glycans were identified as minor constituents of the cyst membrane preparation but were not observed in protoscoleces extracts. Whether the sialylated glycans are host derived or endogenously synthesized by the parasite remains to be established. This is the first reported structural analysis of N-glycans from cestodes and provides new insights into protein glycosylation in helminths.

Animals↗
PubMed1997

Cloning and expression of a cDNA encoding a nonintegrin laminin-binding protein from Echinococcus granulosus with localization of the laminin-binding domain.

We have isolated a cDNA from the hydatid tapeworm, Echinococcus granulosus, encoding a protein that binds laminin. This is the first report of a helminth parasite laminin-binding protein and the first description of a cDNA encoding a laminin-binding protein from a parasite. The cDNA clone (egmo3) was isolated from an E. granulosus protoscolex cDNA expression library, and identified on the basis of sequence homology to the nonintegrin mammalian metastasis-associated 67-kDa laminin receptor (67-LR). The amino acid sequence predicted from the cDNA sequence is 268 residues long with a calculated molecular mass of 29.9 kDa. Southern blot analysis suggested that many copies of the gene may occur in the E. granulosus genome. A Northern blot revealed that the gene is expressed as a single transcript of approximately 1 kb consistent with the size of the cDNA insert. Antibodies raised to the purified protein interacted with a 30 kDa protein in whole E. granulosus protoscoleces. A Western blot of the purified and refolded recombinant protein specifically bound 125I-labelled laminin, as did a synthetic peptide derived from the inferred amino acid sequence of egmo3 which is similar in homology to peptide G, the active ligand-binding site of 67-LR. We also isolated the 3' end of the cDNA encoding the homologous protein from the closely related species, E. multilocularis. The polypeptide encoded by egmo3 also shares substantial identity with the acidic class of ribosomal proteins which are involved in protein synthesis. As such, the egmo3 protein may be multifunctional in E. granulosus, acting as a laminin-binding molecule but also playing a role in cell division and growth.

Amino Acid Sequence↗
PubMed1998

Stage-specific expression of the 14-3-3 gene in Echinococcus multilocularis.

A cDNA expression library representing the metacestode developmental stage of the tapeworm Echinococcus multilocularis was immunoscreened with monospecific antibodies affinity purified following differential immunoblot analysis. Using this procedure, a metacestode-specific clone was isolated representing a 14-3-3 gene of the parasite, which is present as a single copy in the parasite genome. The identity of this clone was demonstrated by cross-reactivity of the recombinant E. multilocularis 14-3-3 protein with antibodies raised against a heterologous 14-3-3 protein from Saccharomyces cerevisiae. In addition, expression of the E. multilocularis 14-3-3 gene in the mutant S. cerevisiae strain, DS9-22, resulted in complementation of the phenotypic deficiency of this strain, thus demonstrating the functionality of the respective gene product. By reverse transcription-polymerase chain reaction (RT-PCR) we showed that the E. multilocularis 14-3-3 protein is about 10-fold overexpressed in the metacestode stage compared with the expression level in the adult stage. Immunolocalization of the 14-3-3 protein in E. multilocularis metacestodes revealed its predominant presence in the germinal layer of the parasite. The results of this study, taken together with the current knowledge on the 14-3-3 protein family, suggest that this parasite molecule may contribute to the promotion of the progressive, potentially unlimited growth behaviour of the E. multilocularis metacestode within the host tissue.

14-3-3 Proteins↗
PubMed1999

A new potent antigen from Echinococcus granulosus associated with muscles and tegument.

An immunoscreening of a cDNA library derived from the adult stage of the parasitic platyhelminth Echinococcus granulosus has been carried out with sera from infected dogs. The EgA31 clone encodes a fibrous protein which shares some sequence elements with paramyosins. The corresponding gene is present as a single copy in the genome. As revealed by an antibody obtained against a fusion protein produced in bacteria, the polypeptide has a molecular weight of 66 kDa. This polypeptide is present at all developmental stages studied and is a potent antigen during an infection by the adult stage in the dog and during cyst growth in human patients. By immunohistology, it was shown that it is present in the tegument and subtegumental parenchyma of the adult with a main location in the region of the suckers where it rapidly accumulates at the time of the head evagination. It is also present in the germinal layer of the cyst and on the protoscolex.

Amino Acid Sequence↗
PubMed2000

Molecular characterization of P-29, a metacestode-specific component of Echinococcus granulosus which is immunologically related to, but distinct from, antigen 5.

In this work the characterization of P-29, a novel 29 kDa antigen from Echinococcus granulosus is reported. E. granulosus was identified while looking for parasite antigens distinct from those present in hydatid cyst fluid. A monoclonal antibody (mAb 47H.PS) prepared against protoscolex components revealed that P-29 is localized to the tegument and rostellum of protoscoleces, and to the germinal layer of the cyst, but it is absent in hydatid cyst fluid or adult worm extracts. Several internal fragments of P-29 showed sequence identity to the amino acid sequence encoded by Eg6, a partial gene sequence reported to code for an epitope of antigen 5 (Ag5), one of the major diagnostic antigens of the parasite. We confirmed that Eg6 encodes a sub-fragment of P-29 by mapping the epitope of mAb 47H.PS, and isolating the full length P-29 cDNA. Since Eg6 had been, postulated to encode a fragment of Ag5, we specifically studied the relationship of P-29 and Ag5 by: (i) examining the cross-reactivity displayed by different mAbs; (ii) comparison of their peptide finger prints; and (iii) a comparative study of their diagnostic value. Our results prove unequivocally that P-29 and Ag5 are immunologically related, but different proteins, raising several questions on the current knowledge of Ag5.

Amino Acid Sequence↗
PubMed2001

Choropleth mapping of regional count data of Echinococcus multilocularis among red foxes in Lower Saxony, Germany.

In this paper, choropleth maps display the geographical distribution of Echinococcus multilocularis in red foxes in Lower Saxony. Areas of high prevalence of the infection in foxes and with high fox population density pose high risk for the human population for alveolar echinococcosis. Spatial statistical methods were used to analyse regional count data obtained from 5365 hunted or found-dead foxes. Spatial smoothing (of raw estimates of regional prevalences based on count data before mapping) was used because raw estimates can give an erratic impression of the spatial pattern of the infection. For smoothing, empirical Bayesian methods are used as an explorative spatial epidemiological tool. The resulting map showed the geographical variation of the estimated prevalences around a median at 9% and indicated the presence of spatial-trend effects. Based on this finding, conditional autoregressive spatial modelling for Freeman-Tukey transformed data was used as an inferential spatial epidemiological tool. There were significant additive linear and quadratic spatial-trend effects with elevated prevalences in the north and south and extreme values (prevalences>38%) for the south of Lower Saxony.

Animals↗
PubMed2003

Controlling Echinococcus multilocularis-ecological implications of field trials.

Two field trials to reduce the prevalence of Echinococcus multilocularis in foxes have been conducted in recent years. Although both trials reduced prevalence considerably, they failed to eradicate the parasite in the study region. Following the control trial in northern Germany, prevalence recovered unexpectedly and rapidly, reaching pre-control levels five quarters (15 months) after the end of control. To understand the internal dynamics of the parasite-host system's reaction to control, we developed a spatially explicit simulation model, Echi. The simulation model incorporates the information available concerning fox tapeworm population dynamics. Using epidemiological parameters to adjust pre-control prevalence, the model predicts the temporal evolution of the prevalence of E. multilocularis in controlled foxes without departing from the range of uncertainty of the field data. However, the model does not predict the rapid pre-control recovery observed in the field trial. The deviation of the model's prediction from field data indicates the involvement of processes not yet taken into account. We modified the model step by step to mimic processes with the potential to cause the rapid post-control recovery of the prevalence of E. multilocularis in foxes. Neither the longevity of tapeworm eggs nor the migratory behaviour of foxes showed any influence on the post-control reaction of the parasite-host system. However, landscape structures leading to a heterogeneous distribution of infected foxes have the potential to alter the system's reaction to control. If infected foxes are concentrated in multiple clusters in the landscape, the model prediction tallied with the range of uncertainty of the field data. Such spatial distribution of infected foxes may be caused by differential abiotic conditions influencing the survival of tapeworm eggs. The model was found to comply best with field data if the foxes acquire partial immunity by being exposed to the fox tapeworm. Both hypotheses explaining the rapid post-control recovery of the prevalence of E. multilocularis observed in the fox population were supported by field data. Both hypotheses have far-reaching consequences for future control trials. The spatial aggregation of infected foxes would enable control efforts to be concentrated on these highly infected areas. However, the acquisition of immunity acts as a buffer to control, necessitating intensified control measures.

Animals↗
PubMed1985

High oral doses of mebendazole interfere with growth of larval Echinococcus multilocularis lesions.

The natural development of the larval stage of Echinococcus multilocularis in man has been studied in 7 patients after presumed radical operation 2 8/12-11 2/12 years prior to detection of a relapse. The volumes of the recurring lesions were assessed by CT-scanning, and assuming linear growth a median increase of 14.8 ml/year (range 3.8-220 ml/year) was calculated. In 6 patients treated for a median duration of 4 5/12 years with high oral doses of mebendazole a median growth rate of -3.0 ml/year (range-470- + 10.2 ml/year) was found, which differed significantly from the natural growth rate (P less than 0.01). Although the patients improved clinically, there was evidence of persistent infection. These data are the first controlled evidence that high oral doses of mebendazole may be parasitostatic in alveolar echinococcosis in man. Although not curative, this pharmacological effect appears to be clinically beneficial.

Adult↗
PubMed2000

How Echinococcus granulosus deals with complement.

Here, Ana Mar a Ferreira and colleagues discuss the interplay between the larval stages of Echinococcus granulosus and an important effector arm of immunity: the host complement system. During early infection, the parasite activates complement, and hence complement-dependent inflammatory responses. However, on differentiation into the hydatid cyst, the parasite exposes to the host a structure - the cyst wall - that does not activate complement strongly. Mechanisms inhibiting complement activation on the cyst wall have been elucidated, contributing to the understanding of how this large, persistent, tissue-dwelling pathogen controls the inflammatory response.

Animals↗
PubMed2000

Salmonella typhimurium as a basis for a live oral Echinococcus granulosus vaccine.

A live attenuated Salmonella typhimurium vaccine candidate, LVR01, was constructed by introducing a null deletion into the aroC gene of the parental canine S. typhimurium isolate, P228067. LVR01 was used to orally deliver to the canine immune system a fatty acid binding protein (FABP) from Echinococcus granulosus (EgDf1), as a fusion protein with fragment C (TetC) of tetanus toxin. Immunization studies demonstrated that live LVR01 is well tolerated by orally vaccinated dogs. There was no detectable shedding of the vaccine strain in the faeces 2 days after immunization. Humoral antibody responses were observed against Salmonella, TetC and EgDf1. Cellular responses were consistently detected against Salmonella and TetC. A cellular response against EgDf1 was also seen in a proportion of the LVR01 vaccinated dogs. We propose S. typhimurium LVR01 as a carrier for recombinant antigens and a vector for the construction of multivalent oral vaccines for dogs.

Administration, Oral↗
PubMed2003

The Echinococcus multilocularis 14-3-3 protein protects mice against primary but not secondary alveolar echinococcosis.

Alveolar echinococcosis (AE), caused by the larval stage (metacestode) of the tapeworm Echinococcus multilocularis, exhibits very similar disease characteristics in humans and rodents. Recently, it has been shown that an over-expression of the parasite 14-3-3 protein could be associated to the proliferative growth of the E. multilocularis metacestode. We now demonstrate the expression of this protein at the E. multilocularis oncospheral stage as well. A recombinant E. multilocularis 14-3-3 protein (E14t) was used to vaccinate mice against either primary or secondary experimental E. multilocularis infection in BALB/c mice. Conversely to non-vaccinated but control infected mice, which developed a very weak anti-E14t response during infection, the response elicited in the E14t-vaccinated and subsequently infected animals exhibited a strong reactivity against the parasite 14-3-3 protein. Major differences became apparent between secondarily and primarily infected animals: whereas no protection against secondary infection was achieved by vaccination, vaccinated animals were protected by 97% against challenge primary infection with 2000 E. multilocularis eggs. Consequently, the parasite 14-3-3 molecule appears crucially involved in the early stage of the host-parasite interplay and exhibits potential to be used as target molecule for the development of protective tools against AE.

14-3-3 Proteins↗
PubMed1999

Synthetic peptides induce antibody against a host-protective antigen of Echinococcus granulosus.

The immunogenicity of four synthetic peptides was investigated in sheep. The sequences of the peptides (6, 12/13, 21/22 and 24) were derived from linear, antibody-binding epitopes of the EG95 recombinant protein, a host-protective antigen of the parasite Echinococcus granulosus. Sheep were immunised with either free peptide or peptide conjugated to diphtheria toxoid. All sheep responded to both conjugated and unconjugated forms of the peptides. For two of the four peptides (6 and 21/22), the amount of antibody elicited was significantly greater for the conjugated form of the peptides than for the corresponding unconjugated forms. For the other two peptides (12/13 and 24), peak antibody levels to both forms of the peptide were equivalent. Maximal antibody titres against peptides 6, 12/13 and 21/22 were established after only one immunisation and were not boosted by a second dose. Antisera to all four peptides reacted with the recombinant antigen, and three of the four peptides generated antibodies, which bound to the native parasite oncosphere antigen. Antisera raised against the peptides were unable to kill the parasite in in vitro culture, although each of the peptides could be used to affinity purify lethal antibody from antisera raised against the recombinant protein. These results indicate that peptides 6, 12/13, 21/22 and 24 of the EG95 recombinant vaccine are immunogenic and suggest that they are associated with host-protective epitopes.

Animals↗
PubMed2001

Mitosis and differentiation in T-cells under cytotoxic action of Echinococcus granulosus hydatid fluid.

In the T-cell line, D10, thymidine uptake was used to measure the proportion of cells in S-phase, and the MTT assay to measure the number of viable cells. The effect of Echinococcus granulosus hydatid fluid (HF) on the lymphocytes was assayed in 3-day cultures of the T-cell line, D10, in increasing concentrations of HF. Apparent cytotoxic effects of HF were recorded as a log-linear decline in S-phase activity, which was reduced by the presence of IL-1, IL-2, or a combination of the two. In the presence of IL-2, however, mitogenic treatment with concanavalin A increased the cytotoxic effect in 3-day cultures, while in day-2 cultures, HF itself showed mitogenic effect. HF-induced decline in S-phase activity was not matched by a parallel decline in viable cells, suggesting that the apparent cytotoxicity of HF could result from cell-cycle arrest. Depending on its origin, HF enhanced membrane expression of CD25 and CD38 on human peripheral blood lymphoblasts, and diminished that of CD28. Taken together, these changes suggest that HF can induce T-cell mitosis and reduce co-stimulation with subsequent T-cell anergy or apoptosis.

Animals↗
PubMed2002

Control of Echinococcus granulosus in Uruguay: evaluation of different treatment intervals for dogs.

This study attempted control of transmission of Echinococcus granulosus from dogs to sheep in different areas in the Department of Florida, Uruguay, by treating dogs with praziquantel at intervals of 6, 12 and 16 weeks. The 6-week interval was based on the prepatent period of infection with E. granulosus, the 12- and 16-week intervals were based on the rate of reinfection with tapeworms in dogs in the area. Dogs had become reinfected with E. granulosus between 2 and 4 months after treatment, whereas they became reinfected with the Taenia spp. tapeworms within 2 months of treatment. One year after the start of treatments sentinel lambs were born and grazed the farms in the three treatment areas. Approximately, 15 months later when the sentinel lambs were killed and examined for parasites the six weekly treatments had stopped the transmission of E. granulosus to the sentinel lambs. Treatment of dogs at 12- and 16-week intervals failed to stop transmission of E. granulosus but both the numbers of farms and the numbers of sheep infected with E. granulosus were lower where dogs received 12 weekly treatments compared with dogs receiving 16 weekly treatments and a fourth area where dogs had received no treatments (chi(2)P=0.002). Lambs continued to become infected with the Taenia spp. tapeworms in all the areas. Control was complicated by large changes in the dog population. From a starting population of 1164 dogs in the three treated areas, 832 new dogs, most of these adult hunting dogs, entered the population and 793 dogs were lost from the population.

Animals↗
PubMed2002

Screening of dogs for Echinococcus granulosus coproantigen in a low endemic situation in Cyprus.

In the framework of an echinococcosis surveillance and control programme in Cyprus, a commercial enzyme-linked immunosorbent assay (ELISA) (CHEKIT ECHINOTEST) designed for the detection of Echinococcus granulosus and E. multilocularis coproantigens was used in 1997-2000 for the investigation of large numbers of dogs. Most of the animals originated from areas where approximately 0.2% of the dogs had been found to be infected with E. granulosus in previous (1993-1996) arecoline surveys. The sensitivity of the coproantigen test was 83%, as determined in 35 dogs naturally infected with this cestode species. The specificity was 98% in 97 randomly selected dogs from Cyprus, but it was reduced to 80% in a group of 50 dogs, infected with Taenia spp. A total of 6551 dogs (mainly of rural origin) was examined, including three large groups (N: 2928, 1761 and 1800) from the Government Control Area (GCA) in southern Cyprus and a small group (N: 62) from the Non-Government Control Area (NonGCA) in the northern part of the island. Among the dogs from the GCA, 184 (2.8%) tested positive for coproantigen; coproantigen prevalences were 2.6, 4.9 and 1.1% in these three groups, and of 62 dogs from the NonGCA 8.1% were positive. The calculated true prevalences of E. granulosus in the dog population of the GCA ranged between 0.0 and 3.58%. The predictive values of the test, based on a 0.2% prevalence, was >99.9% for negative results, but very low (7%) for coproantigen-positive results. However, the relatively small number of coproantigen-positive dogs can be treated with praziquantel or the results can further be confirmed by arecoline purging.

Animals↗
PubMed2002

Evaluation of three enzyme-linked immunosorbent assays (ELISAs) for the detection of serum antibodies in sheep infected with Echinococcus granulosus.

The aim of this study was to develop an immunological method for the identification of sheep infected with Echinococcus granulosus which would allow the monitoring of animals imported into countries free from hydatidosis and as an aid to countries where control schemes for the disease are in operation. Three enzyme-linked immunosorbent assays (ELISAs) were developed and validated, using as antigen either a purified 8 kDa hydatid cyst fluid protein (8kDaELISA), a recombinant EG95 oncosphere protein (OncELISA) or a crude protoscolex preparation (ProtELISA). Sera used for the assay validations were obtained from 249 sheep infected either naturally or experimentally with E. granulosus and from 1012 non-infected sheep. The highest diagnostic sensitivity was obtained using the ProtELISA at 62.7 and 51.4%, depending on the cut-off. Assay sensitivities were lower for the 8kDaELISA and the OncELISA. Diagnostic specificities were high, ranging from 95.8 to 99.5%, depending on the ELISA type and cut-off level chosen. A few sera from 39 sheep infected with T. hydatigena and from 19 sheep infected with T. ovis were recorded as positive. Western immunoblot analysis revealed that the dominant antigenic components in the crude protoscolex antigen preparation were macromolecules of about 70-150 kDa, most likely representing polysaccharides. This study demonstrated that the ProtELISA was the most effective immunological method of those assessed for detection of infection with E. granulosus in sheep. Because of its limited diagnostic sensitivity of about 50-60%, it should be useful for the detection of the presence of infected sheep on a flock basis and cannot be used for reliable identification of individual animals infected with E. granulosus.

Animals↗
PubMed2003

Modelling the transmission dynamics of Echinococcus granulosus in sheep and cattle in Kazakhstan.

Cystic echinococcosis, caused by Echinococcus granulosus, is an emerging disease in many parts of the world and, in particular, in eastern Europe and the former Soviet Union. This paper examines the abundance and prevalence of infection of E. granulosus in cattle and sheep in Kazakhstan. Observed data are fitted to a mathematical model in order to determine if the parasite population is partly regulated by intermediate host immunity and to define parameters in the model. Such data would be useful to develop simulation models for the control of this disease. Maximum likelihood techniques were used to define the parameters and their confidence limits in the model and the negative binomial distribution was used to define the error variance in the observed data. The results indicated that there are significant variations in the infection pressure to sheep depending on their location. In particular sheep from Almaty Oblast and from central and northern Kazakhstan appeared to have a greater exposure than sheep from Jambyl or South Kazakhstan Oblasts. The infection pressure to cattle was somewhat lower in comparison. In common with other similar studies, there was no evidence of parasite-induced immunity in sheep or cattle in Kazakhstan due to natural infection. The highest abundance and prevalence were seen in the oldest age classes of animals.

Age Factors↗
PubMed2003

Echinococcus multilocularis (Cestoda, Taeniidae) in Red foxes (Vulpes vulpes) in northern Belgium.

The first record of the tapeworm Echinococcus multilocularis (Cestoda, Taeniidae) in Red foxes (Vulpes vulpes) in northern Belgium is described. Between 1996 and 1999, 237 dead foxes were examined for the presence of this tapeworm using the intestinal scraping technique. Four foxes (1.7%) were found to be infected with E. multilocularis and showed medium to very high parasitic burdens. Three infected foxes originated from the south of the study area and the fourth animal came from the north of the study area near the border with The Netherlands. These findings are discussed in relation to the high endemicity of E. multilocularis in southern Belgium and to the increased distribution of the Red fox (V. vulpes) in northern Belgium during the last two decades.

Animals↗