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PubMed2005

Identification of membrane-bound and secreted proteins from Echinococcus granulosus by signal sequence trap.

The signal sequence trap technique was applied to identify genes coding for secreted and membrane bound proteins from Echinococcus granulosus, the etiologic agent of cystic hydatid disease. An E. granulosus protoscolex cDNA library was constructed in the AP-PST vector such that randomly primed cDNAs were fused with a placental alkaline phosphatase reporter gene lacking its endogenous signal peptide. E. granulosus cDNAs encoding a functional signal peptide were selected by their ability to rescue secretion of alkaline phosphatase by COS-7 cells that had been transfected with the cDNA library. Eighteen positive clones were identified and sequenced. Their deduced amino acid sequences showed significant similarity with amino acid transporters, Krebs cycle intermediates transporters, presenilins and vacuolar protein sorter proteins. Other cDNAs encoded secreted proteins without homologues. Three sequences were transcribed antisense to E. granulosus expressed sequence tags. All the mRNAs were expressed in protoscoleces and adult worms, but some of them were not found in oncospheres. The putative E. granulosus secreted and membrane bound proteins identified are likely to play important roles in the metabolism, development and survival in the host and represent potential targets for diagnosis, drugs and vaccines against E. granulosus.

Alkaline Phosphatase↗
PubMed2006

Complexity and function of cytokine responses in experimental infection by Echinococcus granulosus.

Cytokines are important in the regulation of the immune system and are secreted by a variety of cells in response to self and non-self stimuli. Communication within cells, in the same or distant anatomical sites, occurs via cytokines which determine the quality and intensity of inflammatory and adaptive immune responses. Infection by helminths is characterized by a dominant secretion of type-2 cytokines; IL-4, IL-5, IL-10 (among others), which down-regulates the induction and functions of type-1 cytokines. The molecular mechanisms involved in the polarization of type-2 responses and their biological significance in helminthic infections are unknown, and probably depends on each host-parasite system. Understanding these issues may contribute to immune therapy against parasitic infections. Here we summarize our data obtained in Echinococcus granulosus experimental infection regarding type-2 cytokine induction and its putative role in the host-parasite interaction. Results suggest that induction of cytokine responses at different stages of infection is complex and depends on several parameters. In addition, they support the hypothesis that early IL-10, secreted by B cells in response to non-proteic antigens, may favour parasite survival and the establishment of a polarized type-2 cytokine response.

Animals↗
PubMed2005

Patent and pre-patent detection of Echinococcus granulosus genotypes in the definitive host.

The detection of Echinococcus granulosus in dogs is important for epidemiological surveillance and evaluation of cystic hydatic disease control programs. We report the efficacy of two PCR-based methods to detect patent and pre-patent infection in dogs experimentally infected with E. granulosus. The detection is based on amplification of a fragment of a mitochondrial gene (Mit-PCR) and a DNA repetitive element (Rep-PCR) of E. granulosus. We tested the ability of both methods to detect several genotypes of the parasite. Both PCR methods could detect E. granulosus in pre-patent and patent periods, even when microscopical observation of eggs resulted negative in fecal samples. The Mit-PCR produced the same amplification pattern for all the parasite genotypes tested while the amplification patterns with the Rep-PCR differed among groups of strains. Fecal samples collected from dogs of an endemic area were diagnosed with more sensitivity than arecoline hydrobromide purgation. These molecular methods could be applied in the confirmation of coproantigen-positive fecal samples and to verify the success of control programs.

Animals↗
PubMed2004

Purification, cloning, and expression of the mitochondrial malate dehydrogenase (mMDH) from protoscolices of Echinococcus granulosus.

Protoscolices of the parasitic helminth Echinococcus granulosus contain two malate dehydrogenases (EC 1.1.1.37), one cytosolic and one mitochondrial. The latter has been separated from the other isoform and purified to protein homogeneity. Sequencing of tryptic peptides by Edman degradation allowed the design of oligonucleotide primers for PCR, leading to the cloning and sequencing of a full length cDNA. The encoding gene is present as a single copy per haploid genome and codes for a protein with high sequence identity (56-58%) with the similar enzymes from mammals, Caenorhabditis elegans and yeast. Active recombinant mitochondrial malate dehydrogenase was expressed in Escherichia coli, as protein fusions with glutathione S-transferase or a poly-His tail. The purified recombinant enzymes had a kinetic behaviour similar to that of the native enzyme, being inhibited by excess of the substrate oxaloacetate and unaffected by excess L-malate. The results indicate that E. granulosus contains two typical eukaryotic malate dehydrogenases, with relative levels quite different from those present in mammalian tissues like heart, in good agreement with the predominantly fermentative metabolism of the protoscolices.

Amino Acid Sequence↗
PubMed2004

Effect of treatment with free and liposomized albendazole on selected immunological parameters and cyst growth in mice infected with Echinococcus multilocularis.

Selected immunological parameters in healthy mice and mice infected with Echinococcus multilocularis and the effect of free and liposomized albendazole (lip.ABZ) upon these parameters in relation to the reduction of parasite growth were investigated over 26 weeks. Proliferative response of splenic T and B lymphocytes, number of CD4+ and CD8+ spleen T cell subpopulations, serum concentration of IFN-gamma and IL-5, and generation of superoxide anion (O2-) by peritoneal macrophages were the chosen parameters. Both drug forms were given to mice at a dose of 10 mg kg(-1) twice a week from week 4 to week 10 post infection (p.i.) (6 weeks in total). The reduction of cyst growth after treatment with ABZ and lip.ABZ was similar up to week 4 after last dose, but the parasitostatic effect of lip.ABZ lasted 4 weeks longer than the effect of free drug. After administration of both drug forms, the proliferative responses of T and B cells were restored, and also the number of CD4+ and CD8+ increased markedly. In lip.ABZ-treated mice, stimulation of mentioned lymphocyte parameters, except that of CD8+ numbers, persisted for longer period than after ABZ therapy, where values peaked at week 12 p.i., then declined more rapidly. A very strong stimulatory effect was seen on B lymphocytes during the period of lip.ABZ administration, although interestingly, numbers of CD8+ cells were higher in free ABZ-treated group. Low concentrations of IFN-gamma (Th1 response) were present in infected, untreated mouse serum. Only moderate IFN-gamma elevation was observed after treatment with free ABZ. A profound increase of its concentration was seen shortly after administration of lip.ABZ, and persisted until the experiment ended. In infected untreated mice, concentration of IL-5 (Th2 response) was highest on week 2 p.i. Significantly more IL-5 was recorded in serum of mice treated with free ABZ treatment than with lip.ABZ from week 12 to 18 p.i. (weeks 2-8 after the last dose). After the initial increase of superoxide anions (weeks 4-11 p.i.), generation of O2- by peritoneal macrophages was gradually inhibited by E. multilcoularis infection. In general, treatment abolished this suppression and macrophages from lip.ABZ-treated mice produced elevated amounts of O2- over a longer period than macrophages from ABZ-treated mice. Our data indicate that anthelmintic potency of ABZ could be increased after incorporation into liposomes, not only because of improved pharmacokinetics and consequent bioavailability, but also because of significant stimulation of Th1-type cytokine IFN-gamma response and effector macrophage functions.

Albendazole↗
PubMed2005

Molecular survival strategies of Echinococcus multilocularis in the murine host.

Larval infection with Echinococcus multilocularis starts with the intrahepatic postoncospheral development of a metacestode that-at its mature stage-consists of an inner germinal and an outer laminated layer (GL & LL). In certain cases, an appropriate host immune response may inhibit parasite proliferation. Several lines of evidence obtained in vivo and in vitro indicate the important bio-protective role of the LL. For instance, the LL has been proposed to protect the GL from nitric oxide produced by periparasitic macrophages and dendritic cells, and also to prevent immune recognition by surrounding T cells. On the other hand, the high periparasitic NO production by peritoneal exsudate cells contributes to periparasitic immunosuppression, explaining why iNOS deficienct mice exhibit a significantly lower susceptibility towards experimental infection. The intense periparasitic granulomatous infiltration indicates a strong host-parasite interaction, and the involvement of cellular immunity in control of the metacestode growth kinetics is strongly suggested by experiments carried out in T cell deficient mouse strains. Carbohydrate components of the LL, such as Em2(G11) and Em492, as well as other parasite metabolites yield immunomodulatory effects that allow the parasite to survive in the host. I.e., the IgG response to the Em2(G11)-antigen takes place independently of alpha-beta+CD4+T cells, and in the absence of interactions between CD40 and CD40 ligand. Such parasite molecules also interfere with antigen presentation and cell activation, leading to a mixed Th1/Th2-type response at the later stage of infection. Furthermore, Em492 and other (not yet published) purified parasite metabolites suppress ConA and antigen-stimulated splenocyte proliferation. Infected mouse macrophages (AE-MØ) as antigen presenting cells (APC) exhibited a reduced ability to present a conventional antigen (chicken ovalbumin, C-Ova) to specific responder lymph node T cells when compared to normal MØ. As AE-MØ fully maintain their capacity to appropriately process antigens, a failure in T cell receptor occupancy by antigen-Ia complex or/and altered co-stimulatory signals can be excluded. Studying the status of accessory molecules implicated in T cell stimulation by MØ, it could be shown that B7-1 (CD80) and B7-2 (CD86) remained unchanged, whereas CD40 was down-regulated and CD54 (=ICAM-1) slightly up-regulated. FACS analysis of peritoneal cells revealed a decrease in the percentage of CD4+ and CD8+T cells in AE-infected mice. Taken together the obstructed presenting-activity of AE-MØ appeared to trigger an unresponsiveness of T cells leading to the suppression of their clonal expansion during the chronic phase of AE infection. Interesting information on the parasite survival strategy and potential can be obtained upon in vitro and in vivo treatment. Hence, we provided very innovative results by showing that nitazoxanide, and now also, respectively, new modified compounds may represent a useful alternative to albendazole. In the context of chemotherapeutical repression of parasite growth, we searched also for parasite molecules, whose expression levels correlate with the viability and growth activity of E. multilocularis metacestode. Expression levels of 14-3-3 and II/3-10, relatively quantified by realtime reverse transcription-PCR using a housekeeping gene beta-actin, were studied in permissive nu/nu and in low-permissive wild type BALB/c mice. At 2 months p.i., the transcription level of 14-3-3 was significantly higher in parasites actively proliferating in nu/nu mice compared to parasites moderately growing in wild type mice. Immunoblotting experiments confirmed at the protein level that 14-3-3 was over-expressed in parasites derived from nu/nu mice at 2 months p.i. In vitro-treatment of E. multilocularis with an anti-echinococcal drug nitazoxanide for a period of 8 days resulted in a significant decrease of both 14-3-3 and II/3-10 transcription levels, which correlated with the kinetics of a housekeeping gene, beta-actin. This indicates that 14-3-3-exhibits a good potential as a molecular marker to assess viability and growth activity of the parasite.

14-3-3 Proteins↗
PubMed2005

Survival strategy of Echinococcus multilocularis in the human host.

As exemplified by "aborted" calcified liver lesions commonly found in patients from endemic areas, Echinococcus multilocularis metacestodes develop only in a minority of individuals exposed to infection with the papasite. Clinical research has disclosed some aspects of the survival strategy of E. multilocularis in human hosts. Clinical observations in liver transplantation and AIDS suggest that suppression of cellular/Th1-related immunity increases disease severity. Most of the studies have stressed a role for CD8+ T cells and for Interleukin-10 in the development of tolerance. A spontaneous secretion of IL-10 by the PBMC seems to be the immunological hallmark of patients with progressive forms of alveolar echinococcosis (AE). IL-10-induced inhibition of effector macrophages, but also of antigen-presenting dendritic cells, may be operating and allowing parasite growth and survival. The genetic correlates of susceptibility to infection with E. multilocularis are clearer in humans than in the mouse model. A significant link between MHC polymorphism and clinical presentation of AE has been shown, and the spontaneous secretion of IL-10 in patients with a progressive AE is higher in patients with the HLA DR3+, DQ2+ haplotype. Clustering of cases in certain families, in communities otherwise exposed to similar risk factors, also points to immuno-genetic predisposition factors that may allow the larva to escape host immunity more easily. The first stage of larval development may be crucial in producing "danger signals" stimulating the initial production of cytokines. Therapeutic use of Interferon alpha is an attempt to foil the survival strategy of E. multilocularis.

Animals↗
PubMed2005

Identification and intra-specific variability analysis of secreted and membrane-bound proteins from Echinococcus granulosus.

Echinococcus granulosus, the etiological agent of cystic hydatid disease, exists as a series of strains or genotypes, differing in biological features. Many of the secreted and membrane-bound proteins (S/M) from helminth parasites are involved in the host-parasite interplay and constitute potential targets for diagnosis, anti-parasitic drugs and vaccines. A number of E. granulosus S/M proteins were identified using the signal sequence trap technique. Six out of seven cDNA fragments of these newly identified proteins showed nucleotide and amino acid sequence variation. Inter-strain variation was reported for other characterized S/M proteins as the vaccine target EG95 and the major hydatid cyst fluid antigen, Antigen B (AgB). AgB is highly polymorphic, 101 different sequences related to AgB were reported so far and were grouped in 5 genes (EgB1-EgB5) and one pseudogene (EgB2p) exclusive of G5, G6/G7 genotypes. The significance of AgB polymorphism and possible consequences in diagnostic performance are discussed. The diagnostic value of the new protein variants detected in E. granulosus strains could be determined through standardized inter-laboratory studies as the recently done by the South American Network for Hydatid Serology.

Amino Acid Sequence↗
PubMed2005

Distribution and ecology of Echinococcus multilocularis in Central Asia.

Much of Central Asia is characterized by an arid landscape. Foci of Echinococcus multilocularis have a very patchy distribution. This is due to a significant deficiency of moisture in the superficial layers of the ground and is related to the types of animal colonies. In the plain, the abundance of endemic foci increases in a direction from the sandy deserts to steppe. In mountains the highest levels of infection of animals are observed in the eastern part of region. In arid regions, E. multilocularis is typically located in humid mesophilous biotopes. Here there is sufficient moisture in the spring to promote survival of eggs and there is a close interrelation between the definitive and intermediate hosts.

Animals↗
PubMed2005

Transmission ecology of Echinococcus multilocularis: what are the ranges of parasite stability among various host communities in China?

A striking feature of the transmission ecology of Echinococcus multilocularis in China is the diversity of hosts that contribute to the parasite cycle. Considering the population dynamics of key reservoir intermediate hosts and the ratio of their preferred habitat in a landscape (ROMPA) is essential to understanding transmission, but the numerous communities in which the parasite cycles and the extent of those communities is currently far from being fully documented. On the Tibetan plateau grassland management influences intermediate host species populations and the sheer size of the area could be a major contributing factor to sustaining transmission in the region. In Southern Gansu and Southern Ningxia the processes of deforestation have temporarily provided optimal habitat for key reservoir intermediate hosts of E. multilocularis, resulting in high transmission and human disease, however currently the parasite may be extinct locally. Faced with this pattern of potentially transient transmission in a diversity of communities the question of the dispersal potential of the parasite arises. The reforestation program currently active across much of Western China has the potential to give rise to a massive increase in habitat favourable to suitable intermediate hosts and emergence or re-emergence of the zoonosis alveolar echinococcosis in many areas. This potential epidemic could be mitigated by both natural and human induced parasite dispersal mechanisms including fox migration and the sale of infected dogs originating from stable endemic foci on the Tibetan plateau. However, currently the degree of genetic exchange between discrete transmission foci is unknown and it is expected that genetic techniques could provide crucial information regarding this important question.

Animals↗
PubMed2005

Mathematical modeling of Echinococcus multilocularis transmission.

A mathematical model for the transmission cycle of Echinococcus multilocularis would be useful for estimating its prevalence, and the model simulation can be instrumental in designing various control strategies. This review focuses on the epidemiological factors in the E. multilocularis transmission cycle and the recent advances of mathematical models for E. multilocularis transmission.

Animals↗
PubMed2005

Towards the control of Echinococcus multilocularis in the definitive host in Japan.

Echinococcus multilocularis is distributed all over Hokkaido, the northern island of Japan. The prevalence in foxes has been around 40% in the last decade. Three trials of anthelmintic bait distribution have been conducted in Hokkaido to reduce the prevalence in foxes. In those trials, bait distribution was done along roads in the study area using cars and/or around fox breeding dens by hand. Changes in the prevalence in foxes were evaluated either by necropsy of captured foxes or by coproantigen and egg detection of faeces collected in field. All of the trials showed bait distribution was effective for the reduction of the prevalence in foxes; however, it was also suggested that a frequent and continuous baiting program is necessary for effective and stable control of the prevalence in foxes. As observed in some cities in Europe, urban foxes infected with the parasite were also recognized in Sapporo. A survey of pet dogs showed that 0.4% of surveyed dogs were determined infected. In addition, a dog which was transported from Hokkaido to the main island of Japan was found excreting E. multilocularis eggs. The results raised the public recognition of canine infections, which in turn lead to the modification of a Japanese law for infectious diseases and to the enforcement of a national reporting system of dogs infected with E. multilocularis by veterinarians.

Animals↗
PubMed2005

Spatial modelling and ecology of Echinococcus multilocularis transmission in China.

Recent research in central China has suggested that the most likely transmission mechanism for Echinococcus multilocularis to humans is via domestic dogs which are allowed to roam freely and hunt (infected) small mammals within areas close to villages or in areas of tented pasture. This assertion has led to the hypothesis that there is a landscape control on transmission risk since the proximity of suitable habitat for susceptible small mammals appears to be the key. We have tested this hypothesis in a number of endemic areas in China, notably south Gansu Province and the Tibetan region of western Sichuan Province. The fundamental landscape control is its effect at a regional scale on small mammal species assemblages (susceptible species are not ubiquitous) and, at a local scale, the spatial distributions of small mammal populations. To date the research has examined relationships between landscape composition and patterns of human infection, landscape and small mammal distributions and recently the relationships between landscape and dog infection rates. The key tool to characterize landscape is satellite remote sensing and these data are used as inputs to drive spatial models of transmission risk. This paper reviews the progress that has been made so far in spatial modeling of the ecology of E. multilocularis with particular reference to China, outlines current research issues, and describes a framework for building a spatial-temporal model of transmission ecology.

Animals↗
PubMed2006

Spatial spreading of Echinococcus multilocularis in Red foxes (Vulpes vulpes) across nation borders in Western Europe.

The occurrence of the fox tapeworm Echinococcus multilocularis in Red foxes was studied in Belgium and a neighbouring region in The Netherlands. A total number of 1202 foxes were analysed (1018 in Belgium and 184 in The Netherlands) of which 179 were infected with E. multilocularis (164 in Belgium and 15 in The Netherlands). Further, the spatial distribution of infection among sampled foxes was analysed with an ellipsoidal gradient, demonstrating a decreasing prevalence in northwestern direction. Using this gradient, we showed that the spatial patterns of infection in Belgium and the neighbouring region in The Netherlands correspond, indicating a continuous distribution of E. multilocularis across the nation borders. Part of the Belgian data allowed investigating temporal changes in the spatial distribution of E. multilocularis. This revealed a northwestern spread of E. multilocularis.

Animals↗
PubMed2004

Wilderness in the city: the urbanization of Echinococcus multilocularis.

A distinct increase in fox populations, particularly in urban areas, has been observed in Europe. This is of particular concern in endemic regions of the small fox tapeworm Echinococcus multilocularis, the aetiological agent of human alveolar echinococcosis. Novel tools have facilitated the investigation of the ecology of urban foxes and have demonstrated the urban wildlife cycle of E. multilocularis. Such studies are essential for estimating the risk of transmission to humans and to determine the basics for the development of control strategies.

Animals↗
PubMed2003

Prevalence of Echinococcus multilocularis in the red fox (Vulpes vulpes) in southern Belgium.

Between June 1998 and February 2002, 709 red foxes killed in Wallonia (south of Belgium) were available for parasitological examination of the gut. The identification of Echinococcus multilocularis was based on morphological data. E. multilocularis adults were observed in 20.2% of the animals. The analysis of data revealed marked differences between the geological areas of Wallonia; the highest prevalence (33%) was found in the Ardenne and the lowest (0%) on the Plateau de Herve. Host gender and the collection season had no effect on the prevalence. However, the latter was significantly higher in juveniles (<8 months of age). The geographical distribution of E. multilocularis in Belgium is much wider than originally thought.

Age Factors↗
PubMed2004

Local and systemic immune responses to Echinococcus granulosus in experimentally infected dogs.

Local and systemic immune responses were studied in six dogs experimentally infected with the dog/sheep tapeworm Echinococcus granulosus. All dogs developed similar IgG antibody response to parasite antigens. In contrast, IgE and IgA responses differed widely. No relationship between IgA responses and parasite burden at the end of the infection were observed. Further, clear differences in the anti-parasite IgA response in serum as compared with specific IgA forming cells in mesenteric lymph nodes were observed within the same dog. An inverse association of anti-parasite IgE and parasite load seemed to be present, with the strongest IgE response in the one dog that had no worms in the intestine at the end of the experiment. No differences were observed in the numbers of intestinal mast cells and goblet cells among all infected dogs. However, the dog with no detectable parasite load had a marked reduction of detected mast cells in the submuscular and muscular layer of the mucosa. Our data give new insight into the immune response of dogs during E. granulosus infection and provide information that may be useful for the rational design of vaccines for the control of hydatid disease.

Animals↗
PubMed2004

Base line prevalence and spatial distribution of Echinococcus multilocularis in a newly recognized endemic area in the Netherlands.

The base line prevalence of Echinococcus multilocularis in foxes was determined in the eastern part of the province of Groningen in the Netherlands adjacent to the German border. This region has been identified in a previous study in 1998 as one of the westernmost border areas of E. multilocularis. Base line prevalence data are important for a better insight in the possible spread of the parasite and its changes in time. As fox feces containing E. multilocularis eggs are an important source for human exposure this base line prevalence is also an indicator for the potential risk for public health. The base line prevalence was estimated at 9.4% (95% CI: 5.2-16.5%). These results confirm previous findings of E. multilocularis in the same region. The spatial distribution of the infected foxes has been analyzed as a spatial gradient using a logistic model. The prevalence appeared to change strongest in east-western direction and was highest near the German border, adjacent to a German endemic area. These results suggest that the border areas in the Netherlands are the most margin of E. multilocularis territory.

Animals↗