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The role of parasitic diseases as causes of mortality in small ruminants in a high-potential farming area in central Kenya.

A 15-year retrospective study was performed to determine the role of parasitic diseases in causing mortalities in small ruminants. In total, 115 (32 %) sheep were diagnosed as having been killed by parasitic diseases out of 366 that died as a result of disease. The major cause of mortality was helminthosis (63 % of all parasitic cases). Most of the helminthosis cases were attributed to haemonchosis (40% of parasitic cases). Heartwater was the second most important parasitic disease (27% of all parasitic cases). Ninety-five (26%) goats were diagnosed to have been killed by parasitic diseases out of 365 cases presented at the post mortem facility. Helminthosis was the most frequent cause of mortality (55% of the total parasitic diseases). Twenty-six goats were killed by haemonchosis (27% of all parasitic diseases). Heartwater was the second most important parasitic disease, accounting for about 20% of all parasitic diseases. These findings indicate that viable helminth and tick control strategies should be devised in order to reduce mortality caused by helminthosis and heartwater and thereby achieve improved productivity.

Animals↗

Community ecology of the metazoan parasites of the grey triggerfish, Balistes capriscus Gmelin, 1789 and queen triggerfish B. vetula Linnaeus,1758 (Osteichthyes: Balistidae) from the State of Rio de Janeiro, Brazil.

Sixty-six specimens of grey triggerfish Balistes capriscus Gmelin, 1788 and thirty specimens of queen triggerfish B. vetula Linnaeus, 1758 (Osteichthyes: Balistidae) collected from the coastal zone of the State of Rio de Janeiro (21-23 degrees S, 41-45 degrees W), Brazil, from April 2001 to May 2003, were necropsied to study their metazoan parasites. All fish were parasitized by at least one parasite species. Twenty-seven parasites species were collected: 22 in B. capriscus and 15 in B. vetula. Ten parasite species were common to the two host species. Balistes capriscus and B. vetula were a new host record for 16 and seven parasite species, respectively. Hypocreadium biminensis and Taeniacanthus balistae were recorded for the first time in Brazil. The copepod T. balistae and the nematode Contracaecum sp. were the dominant species with highest parasitic prevalence and abundance in the parasite community of B. capriscus and B. vetula, respectively. The metazoan parasites of B. capriscus and B. vetula showed typical aggregated pattern of distribution. The infracommunities of adult endoparasites showed highest values of mean abundance and parasite species richness. The parasite species richness, the total number of specimens and the mean Berger-Parker's index of the infracommunities of B. capriscus and B. vetula showed significant differences.

Animals↗

Antimalarial action of hydrophilic drugs: involvement of aqueous access routes to intracellular parasites.

The antimalarial action and intracellular distribution of the hydrophilic agents phloridzin (PHL) (a bioflavonoid glycoside) and desferrioxamine (DFO) (an iron chelator) were studied in cultures of Plasmodium falciparum-infected human erythrocytes. When added to cultures, these agents arrested parasite growth with IC50 values of 12 microM (PHL) and 22 microM (DFO). At 37 degrees, PHL (40 microM) was virtually impermeant to uninfected cells but permeated with a mean t1/2 of 1.5 hr in trophozoites (30% accessible cell volume) and 8 hr in rings (10% of accessible cell volume). PHL, in analogy with DFO, was demonstrably permeant to infected cells harboring mature forms of the parasites. Permeation was restricted to only a fraction of the infected cell volume. PHL elicited inhibition of nucleic acid synthesis within 1 hr of exposure of trophozoites to PHL (40 microM) and in > 8 hr of exposure of rings. Red cell containers into which millimolar concentrations of PHL or DFO were encapsulated demonstrably supported parasite invasion and subsequent parasite growth and maturation (48-hr incubation). Under culture conditions, uninfected or parasite-infected red cell containers that were loaded with either agent retained the drugs for at least 42 hr at hundred-micromolar concentrations. The agent present in the cells was fully active after release from cells and administration to test cultures of parasites. PHL added to parasite cultures was active at micromolar concentrations, but when present intracellularly it was virtually inactive even at millimolar concentrations. The data presented are consistent with direct access of hydrophilic agents from medium to parasite, a process referred to as fenestration. Permeation into parasites might constitute the rate-limiting step in drug uptake and drug-mediated arrest of parasite growth by PHL and DFO. The putative role of the parasitophorous duct in providing aqueous access routes from medium to parasites is discussed.

Animals↗

Interactions between malaria and mosquitoes: the role of apoptosis in parasite establishment and vector response to infection.

Malaria parasites of the genus Plasmodium are transmitted from host to host by mosquitoes. Sexual reproduction occurs in the blood meal and the resultant motile zygote, the ookinete, migrates through the midgut epithelium and transforms to an oocyst under the basal lamina. After sporogony, sporozoites are released into the mosquito haemocoel and invade the salivary gland before injection when next the mosquito feeds on a host. Interactions between parasite and vector occur at all stages of the establishment and development of the parasite and some of these result in the death of parasite and host cells by apoptosis. Infection-induced programmed cell death occurs in patches of follicular epithelial cells in the ovary, resulting in follicle resorption and thus a reduction in egg production. We argue that fecundity reduction will result in a change in resource partitioning that may benefit the parasite. Apoptosis also occurs in cells of the midgut epithelium that have been invaded by the parasite and are subsequently expelled into the midgut. In addition, the parasite itself dies by a process of programmed cell death (PCD) in the lumen of the midgut before invasion has occurred. Caspase-like activity has been detected in the cytoplasm of the ookinetes, despite the absence of genes homologous to caspases in the genome of this, or any, unicellular eukaryote. The putative involvement of other cysteine proteases in ancient apoptotic pathways is discussed. Potential signal pathways for induction of apoptosis in the host and parasite are reviewed and we consider the evidence that nitric oxide may play a role in this induction. Finally, we consider the hypothesis that death of some parasites in the midgut will limit infection and thus prevent vector death before the parasites have developed into mature sporozoites.

Animals↗

Associations between fish reproductive cycle and the dynamics of metazoan parasite infection.

The parasite fauna of the stone loach (Barbatula barbatula) was investigated monthly from May to December 2001. Four parasite species were recorded: Gyrodactylus spp. (Monogenea), Tylodelphys clavata (Digenea), Proteocephalus sagittus (Cestoda), and Raphidascaris acus (Nematoda). The changes in the seasonal dynamics of parasite infection were investigated throughout the 8 months. The potential effect of the reproductive investment of the host, measured by gonad mass and gonado-somatic index (GSI), on the parasite infection was tested against the prediction that, during periods of high reproductive investment (beginning of the breeding period or forming gonads after breeding), the fish are more susceptible to parasite infection. Differences between parasite loads between genders were also hypothesized. Seasonal differences in infection were observed for all parasite species studied. The values of GSI showed a pattern of energy accumulation in the pre-reproductive period and at the beginning of breeding, a decrease during breeding, and an increase in the post-breeding period. A similar pattern was observed for parasite abundance, a strong or weak increase in spring and/or autumn and a decrease during summer (July and August). Positive correlations between the abundance of Gyrodactylus spp. and R. acus and both gonad mass and GSI were found in females after eliminating the effect of fish weight. Our results suggest that stone loach females are more susceptible to parasite infection in periods of higher reproductive investment. The main factor determining the infection of T. clavata was fish size. The abundance of P. sagittus was positively correlated with GSI in the total fish sample with no detectable effect of sex or fish weight. The increase in cestode infection in spring supports the hypothesis that the parasite life cycle could by synchronized with the beginning of host reproduction, probably induced by increasing fish hormone levels in the spring.

Animals↗

Altered parasite assemblages in raccoons in response to manipulated resource availability.

The role that host aggregation plays in structuring parasite assemblages was examined by experimentally increasing the contact rates of raccoons, Procyon lotor. Two populations of raccoons in southern New York were monitored for 2 years to determine baseline levels of host interaction and to identify the parasite assemblage. In the third year of the study, one population was provisioned with the addition of clumped food resources, while the other was provisioned with equal quantities of dispersed food resources. Remote photography showed that raccoons aggregated at clumped resources but not at dispersed resources, and therefore contact rates between individuals were higher in the site with clumped resources. There were no differences in parasitism between the sites prior to resource augmentation. Among ectoparasites, there were no significant changes in the prevalence or abundance of any species in response to the perturbation. In contrast, across the endoparasite assemblage within and across hosts, the prevalence of infection increased as a result of increased host contact. Strong increases in the prevalence of a few directly transmitted species and slight increases among most species lead to increased evenness in parasite prevalence, suggesting that parasites in this system are transmission limited. In addition, the number of parasite species per host (the parasite infracommunity) was higher in the clumped-resource population. These endoparasite results suggest that intraspecific variation in the species richness of parasite communities of individual hosts, and the prevalence of parasitic species in host populations as assessed across entire parasitic assemblages, is robustly influenced by intraspecific variation in the degree of host contact. Further, these results suggest that anthropogenic changes which alter resource availability may have important consequences for disease transmission in wildlife.

Analysis of Variance↗

Vaccination against animal parasites.

A decade of molecular parasitology is beginning to bear fruit, with the appearance of several new, highly effective, practical vaccines against parasitic diseases. Recombinant antigen vaccines have been developed against cestode, nematode, trematode, protozoan and arthropod parasites. Greatest progress has been made with veterinary vaccines, where the ability to test numerous vaccine formulations in challenge trials has allowed more rapid identification of host-protective antigens than is possible with many medically important parasites. Several quite different approaches to vaccine development have been successful. The traditional approach using live, attenuated parasites continues to provide effective vaccines against several protozoan and nematode parasites. Recombinant DNA technology, monoclonal antibody technology, protein chemistry and immunochemistry have played critical roles in the outstanding success which has been achieved over the last 5 years in the development of defined-antigen vaccines. Two approaches have been successful in research towards defined antigen vaccines against parasites: (1) the 'natural antigen' approach where immune responses are stimulated to parasite molecules which are normally antigenic, and possibly host-protective, in infected hosts; (2) the 'naive antigen' approach where parasite molecules which are not antigenic, or of very low antigenicity, in infected hosts are used to raise immune responses capable of killing the parasite. This review examines the successful approaches taken towards the development of effective anti-parasite vaccines and the vaccines which have been produced to date.

Animals↗

No evidence for specificity between host and parasite genotypes in experimental Strongyloides ratti (Nematoda) infections.

A key requirement for several theories involving the evolution of sex and sexual selection is a specificity between host and parasite genotypes, i.e. the resistance of particular host genotypes to particular parasite genotypes and the infectivity of particular parasite genotypes for particular host genotypes. Determining the scope and nature of any such specificity is also of applied relevance, since any specificity for different parasite genotypes to infect particular host genotypes may affect the level of protection afforded by vaccination, the efficacy of selective breeding of livestock for parasite resistance and the long-term evolution of parasite populations in response to these control measures. Whereas we have some evidence for the role of specificity between host and pathogen genotypes in viral and bacterial infections, its role in macroparasitic infections is seldom considered. The first empirical test of this specificity for a vertebrate-nematode system is provided here using clonal lines of parasite and inbred and congenic strains of rat that differ either across the genome or only at the major histocompatibility complex. Although significant differences between the resistance of host genotypes to infection and between the fitness of different parasite genotypes are found, there is no evidence for an interaction between host and parasite genotypes. It is concluded that a specificity between host and parasite genotypes is unlikely in this system.

Animals↗

The host-parasite relationship in bovine neosporosis.

Infection with the protozoan parasite Neospora caninum is thought to be a major cause of reproductive failure in cattle worldwide. Cattle infected with the parasite are three to seven times more likely to abort compared to uninfected cattle. The parasite may be transmitted to cattle through the ingestion of oocysts that are shed in the faeces of acutely infected dogs (definitive host of N. caninum) or by congenital infection from mother to foetus via the placenta. Interestingly, transplacental transmission can occur over consecutive pregnancies and congenitally infected heifers can transmit the parasite to their own offspring. This repeated vertical transmission observed in naturally infected cattle suggests that cattle do not easily develop effective immunity to the parasite, presenting a significant challenge to the development of a control strategy based on vaccination. Neosporosis is a disease of pregnancy and studying the bovine maternal and foetal immune responses during pregnancy will help us to understand the change in the balance between the parasite and the host that may result in disease of the foetus. Studies in non-pregnant cattle and in murine models of infection have shown the importance of T-helper 1-type immune responses involving pro-inflammatory cytokines, such as IFNgamma and IL-12, in limiting intracellular multiplication of the parasite. During pregnancy, changes occur in the immune system allowing the mother to accept the foetal allograft. Research in other species has stressed the crucial role of T-helper 2-type cytokines at the materno-foetal interface in maintaining the pregnancy and regulating the potentially damaging effect of Th-1 responses. Studies in cattle have shown that cell proliferation and IFNgamma responses may be significantly down-regulated around mid-gestation. This may mean that cattle are less able to cope with N. caninum infection at this time and are more likely to transmit the parasite to the foetus. Another important factor is the gestational age and hence immuno-competence of the foetus at the time of infection. Early in gestation, N. caninum infection of the placenta and subsequently the foetus usually proves fatal, whereas infection occurring in mid to late pregnancy may result in the birth of a congenitally infected but otherwise healthy calf. Studies of foetal immune responses have shown that at 14 weeks of gestation, lymphocytes only respond to mitogen, while by 24 weeks (mid-gestation), they respond to antigen by proliferating and releasing IFNgamma. Clearly, there are several factors influencing the outcome of N. caninum infection in pregnancy: the timing, quantity and duration of parasitaemia, the effectiveness of the maternal immune response and the ability of the foetus to mount an immune response against the parasite. The challenge is to design a vaccine that will prevent foetal infection by N. caninum. This is likely to involve a fine balancing act with the immune system that will allow intervention in a manner that will tip the host-parasite balance in favour of the host without compromising the pregnancy.

Animals↗

Parasite diversity/host age and size relationship in three coral-reef fishes from French Polynesia.

The parasite communities of three coral-reef fish species (Stegastes nigricans, Dascyllus aruanus and Cephalopholis argus) were on Tiahura reef, French Polynesia. The age and growth of each fish was analysed by otolith increment counts and a significant correlation between these variables was found. Stegastes nigricans was parasitised by six adult parasite species, D. aruanus by two adult parasite species and C. argus by five adult parasite species. The most common parasite species were found in all fish size classes. Ectoparasites showed a positive relationship between their abundance and host body length for all three reef fish species. A positive relationship was found only between host size and parasite abundance for common endoparasite species. Parasite species richness, Brillouin's diversity index, and host size and age were positively related. Finally, we discuss the influence of different biological (host diet, host immune response, parasite life-cycle) and ecological factors on parasite community structure in these three reef fishes. Host diet quality seems to be one of the major factors affecting the endoparasite community structure in these reef fishes. Ectoparasite communities seem to be influenced more by biological factors such as, for example, host immunity for the caligid larvae or parasite life-cycle for the gnathiid praniza larvae. In addition, the effect of ecological factors such as cleaning symbiosis on these ectoparasites cannot be dismissed.

Aging↗

Introduced species and their missing parasites.

Damage caused by introduced species results from the high population densities and large body sizes that they attain in their new location. Escape from the effects of natural enemies is a frequent explanation given for the success of introduced species. Because some parasites can reduce host density and decrease body size, an invader that leaves parasites behind and encounters few new parasites can experience a demographic release and become a pest. To test whether introduced species are less parasitized, we have compared the parasites of exotic species in their native and introduced ranges, using 26 host species of molluscs, crustaceans, fishes, birds, mammals, amphibians and reptiles. Here we report that the number of parasite species found in native populations is twice that found in exotic populations. In addition, introduced populations are less heavily parasitized (in terms of percentage infected) than are native populations. Reduced parasitization of introduced species has several causes, including reduced probability of the introduction of parasites with exotic species (or early extinction after host establishment), absence of other required hosts in the new location, and the host-specific limitations of native parasites adapting to new hosts.

Adaptation, Physiological↗

Antigenic variation and the within-host dynamics of parasites.

Many parasites exhibit antigenic variation within their hosts. We use mathematical models to investigate the dynamical interaction between an antigenically varying parasite and the host's immune system. The models incorporate antigenic variation in the parasite population and the generation of immune responses directed against (i) antigens specific to individual parasite variants and (ii) antigens common to all the parasite variants. Analysis of the models allows us to evaluate the relative importance of variant-specific and cross-reactive immune responses in controlling the parasite. Early in the course of infection within the host, when parasite diversity is below a defined threshold value (the value is determined by the biological properties of the parasite and of the host's immune response), the variant-specific immune responses are predominant. Later, when the parasite diversity is high, the cross-reactive immune response is largely responsible for controlling the parasitemia. It is argued that increasing antigenic diversity leads to a switch from variant-specific to cross-reactive immune responses. These simple models mimic various features of observed infections recorded in the experimental literature, including an initial peak in parasitemia, a long and variable duration of infection with fluctuating parasitemia that ends with either the clearance of the parasite or persistent infection.

Animals↗

Potential interactions between metazoan parasites of the Mayan catfish Ariopsis assimilis and chemical pollution in Chetumal Bay, Mexico.

The effect of pollutants on the intensity of infection of metazoan parasites in the Mayan catfish, Ariopsis assimilis was investigated. Data were collected on pollutants and metazoan parasites from 76 catfish from five localities in Chetumal Bay in October, 1996. Nineteen pollutants (pesticides, polychlorinated biphenyls (PCBs) and polycyclic aromatic hydrocarbons (PAHs)) were found in the catfish livers. Heavy metal content was not determined. Nineteen metazoan parasite species were recovered. After controlling for fish length and sampling station, there was a significant negative linear relationship between the intensity of the larval digenean Mesostephanus appendiculatoides and 1,1,1,-trichloro-2,2-bis (4-chlorophenyl) ethane (DDT) concentrations. This negative relationship may be explained either by the effect of the pesticide on the mortality of (i) free-living larval forms, (ii) metacercariae in the fish, (iii) infected fish or (iv) intermediate host snails. There were significant differences between fish parasitized and not parasitized with M. appendiculatoides with respect to their DDT concentrations. There were also significant differences between the variances of the mean Clark's coefficient of condition values between catfish parasitized and not parasitized by M. appendiculatoides, with the variance of non-parasitized catfish being significantly larger. The results provided statistical evidence that DDT has a detrimental effect on M. appendiculatoides infection intensity. Furthermore, the significantly larger variance value of Clark's coefficient for non-parasitized fish suggested that DDT affects both the parasite and general host condition.

Animals↗

Parasites and the evolutionary diversification of primate clades.

Coevolutionary interactions such as those between hosts and parasites have been regarded as an underlying cause of evolutionary diversification, but evidence from natural populations is limited. Among primates and other mammalian groups, measures of host diversification rates vary widely among lineages, but comparative studies have not yet identified a reliable explanation for this variation. In this study, we used a comprehensive data set of disease-causing organisms from free-living primates to illustrate how phylogenetic comparative methods can be used to examine mammalian lineage diversity in relation to parasite species richness. Our results provide evidence that the phylogenetic diversity of primate clades is correlated positively with the number of parasite species harbored by each host and that this pattern is largely independent of other host traits that have been shown to influence diversification rates and parasite species richness in primates. We investigated two possible mechanisms that could explain this association, namely that parasites themselves drive host evolutionary diversification through processes linked with sexual selection and that host shifts or host sharing increases parasite species richness among diverse primate clades. Neither parasite species richness nor host diversification is related to measures of sexual selection in primates. Further, we found only partial evidence that more rapidly diversifying host lineages produced increased opportunities for host sharing or host shifting by parasites through mechanisms involving species' geographic range overlap. Thus, our analyses provide evidence for an important link between the evolutionary diversification of primates and the richness of their parasite communities, but other mechanisms, particularly those related to reciprocal selection or coextinction of hosts and parasites, require further investigation.

Animals↗

Genomic contingence beneath ecological convergence: the tempo and mode of gene loss in parasitic bilaterians.

Parasitism has independently evolved hundreds of times among metazoans. Nonetheless, parasites have explored only a limited range of ecologies, and they display frequent convergence in morphological, behavioral, and life-history traits. Although gene loss in particular parasitic species has been documented, it is not known if gene loss converges along the same lines as these other traits. To test for convergent gene loss, we characterized the housekeeping, regulatory, and DNA-repair complements of 48 bilaterian species, including 20 parasites belonging to 6 different bilaterian phyla. We found that different parasitic strategies do not display characteristic tempos or modes of gene loss. Further, the accelerated rates of gene loss seen in some parasites were almost always shared with their free-living relatives, indicating that the increased rate of loss preceded the rise of parasitism. Therefore, the convergent ecological strategies and adaptations that have arisen in distantly related parasitic lineages overlay contingent gene losses, which largely reflect their phylogenetic history. These results have important implications for how ecologists and evolutionary biologists should model the acquisition of parasitism, especially regarding the long-held assumption that reversion from a parasitic to a free-living state is impossible.

Animals↗

Parasite-mediated predation between native and invasive amphipods.

Parasites can structure biological communities directly through population regulation and indirectly by processes such as apparent competition. However, the role of parasites in the process of biological invasion is less well understood and mechanisms of parasite mediation of predation among hosts are unclear. Mutual predation between native and invading species is an important factor in determining the outcome of invasions in freshwater amphipod communities. Here, we show that parasites mediate mutual intraguild predation among native and invading species and may thereby facilitate the invasion process. We find that the native amphipod Gammarus duebeni celticus is host to a microsporidian parasite, Pleistophora sp. (new species), with a frequency of infection of 0-90%. However, the parasite does not infect three invading species, G. tigrinus, G. pulex and Crangonyx pseudogracilis. In field and laboratory manipulations, we show that the parasite exhibits cryptic virulence: the parasite does not affect host fitness in single-species populations, but virulence becomes apparent when the native and invading species interact. That is, infection has no direct effect on G. d. celticus survivorship, size or fecundity; however, in mixed-species experiments, parasitized natives show a reduced capacity to prey on the smaller invading species and are more likely to be preyed upon by the largest invading species. Thus, by altering dominance relationships and hierarchies of mutual predation, parasitism strongly influences, and has the potential to change, the outcome of biological invasions.

Amphipoda↗

Immunological investments reflect parasite abundance in island populations of Darwin's finches.

The evolution of parasite resistance can be influenced by the abundance of parasites in the environment. However, it is yet unresolved whether vertebrates change their investment in immune function in response to variation in parasite abundance. Here, we compare parasite abundance in four populations of small ground finches (Geospiza fuliginosa) in the Galapagos archipelago. We predicted that populations exposed to high parasite loads should invest more in immune defence, or alternatively use a different immunological defence strategy. We found that parasite prevalence and/or infection intensity increased with island size. As predicted, birds on large islands had increased concentrations of natural antibodies and mounted a strong specific antibody response faster than birds on smaller islands. By contrast, the magnitude of cell-mediated immune responses decreased with increasing parasite pressure, i.e. on larger islands. The data support the hypothesis that investments into the immune defence are influenced by parasite-mediated selection. Our results are consistent with the hypothesis that different immunological defence strategies are optimal in parasite-rich and parasite-poor environments.

Analysis of Variance↗

Parasite transmission modes and the evolution of virulence.

A mathematical model is presented that explores the relationship between transmission patterns and the evolution of virulence for horizontally transmitted parasites when only a single parasite strain can infect each host. The model is constructed by decomposing parasite transmission into two processes, the rate of contact between hosts and the probability of transmission per contact. These transmission rate components, as well as the total parasite mortality rate, are allowed to vary over the course of an infection. A general evolutionarily stable condition is presented that partitions the effects of virulence on parasite fitness into three components: fecundity benefits, mortality costs, and morbidity costs. This extension of previous theory allows us to explore the evolutionary consequences of a variety of transmission patterns. I then focus attention on a special case in which the parasite density remains approximately constant during an infection, and I demonstrate two important ways in which transmission modes can affect virulence evolution: by imposing different morbidity costs on the parasite and by altering the scheduling of parasite reproduction during an infection. Both are illustrated with examples, including one that examines the hypothesis that vector-borne parasites should be more virulent than non-vector-borne parasites (Ewald 1994). The validity of this hypothesis depends upon the way in which these two effects interact, and it need not hold in general.

Animals↗