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[Host-parasite interactions in Coccus hesperidum L. (Hom. Coccidae) and its parasite Coccophagus scutellaris Dalman (hum. Aphelinidae). III. Effect of the number of hosts and their sanitary conditions on the oviposition of the parasite].

Intensity of egg laying of Coccophagus scutellaris depends on the number of hosts available. It is found to be more important when the female has several hosts. Furthermore a better distribution of eggs is observed in that case. When parasited and parasiteless hosts are available, the female of Coccophagus deposits more eggs on the latter. This discriminating ability in relation to number and kinds of hosts available is discussed.

Animals↗

Four-year monitoring of parasite communities in gobiid fishes of the southwest Baltic: III. parasite species diversity and applicability of monitoring.

The parasite infrapopulations of five goby species (Pomatoschistus minutus, P. pictus, P.microps, Gobiusculus flavescens and Gobius niger) were investigated during spring, summer and autumn of the years 1997-2000. In total, 34 parasite species were found: 17 Digenea, 6 Nematoda, 5 Cestoda, 3 Acanthocephala, 2 Protozoa, and 1 Monogenea. The dominant parasites were the digeneans Podocotyle atomon and Cryptocotyle concavum, which represent different ecological groups in terms of their modes of transmission, either indirectly by prey or directly by larvae. The relationship between the parasite Cryptocotyle concavum and the host P. microps is a special one which results in a mean intensity of several hundred cysts (max. 1,329) which settle in the kidney. The diversity of the parasite component community was highest in autumn, but low in spring and summer, with the exception of P. microps for which high values were already found in spring when direct parasites were disregarded. These results depend on the respective seasonal variation in species, some of which occur in huge numbers in some hosts. The diversity of the prey parasite assemblage is higher in Pomatoschistus microps and Gobius niger than in the whole parasite spectrum; the other hosts present the opposite trend. A combination of the island theory of biogeography as modified for parasite infection with the theory of screens and filters leads to a model which considers three handicaps or distances for parasite colonisation: genetic, phylogenetic and ecological. Long-term investigations, as performed here over a time-span of 4 years, can detect more than 80% of parasite species in single hosts after 3 years, and in the whole goby guild after 2 years. Long-term investigations can be useful for finding rare parasites, in analysing parasite diversity, and for determining the seasonality of parasites.

Acanthocephala↗

Low survival of parasite chicks may result from their imperfect adaptation to hosts rather than expression of defenses against parasitism.

Host parents exhibit a variety of behaviors toward avian brood parasites, but not all of their actions have necessarily evolved in response to costs imposed by parasites. To investigate whether common waxbills (Estrilda astrild) have evolved defenses specifically against parasitic pin-tailed whydahs (Vidua macroura), I studied the specificity and flexibility of host behaviors toward nestlings at two sites that differed significantly in parasitism rates and intensities. I focused on documenting nestling survival because V. macroura young match the elaborate gape morphology of E. astrild nestlings, a pattern that suggests hosts may possess unique defenses against parasite chicks. Parasite young survived significantly worse than host young in mixed broods. However, this apparent discrimination was not associated with parasitism risk as would be expected if defenses had evolved specifically to counter parasitism. Parasite young may have survived poorly compared to host young because individual chicks were less able to stimulate sufficient care from foster parents or because they were more susceptible to nestling competition, disease, or reduced provisioning by hosts. Mortality may have also been exacerbated by poor timing of parasite egg laying. In nonparasitized and parasitized nests, rates of nestling survival were similar, further suggesting that parenting behaviors that result in chick mortality did not evolve solely in response to parasite young. In addition, orange-breasted waxbills (Amandava subflava) and zebra finches (Taeniopygia guttata), rarely parasitized and nonparasitized relatives of E. astrild, experience similar levels of nestling mortality presumably as a result of phylogenetically widespread parenting strategies. Despite the similarity of parasitic V. macroura nestlings and E. astrild nestlings, I found no evidence that E. astrild parents possess defenses that allow for specific discrimination against parasite chicks during the nestling period. Rather than being subject to host defenses evolved in an arms race, Vidua chicks may simply be imperfectly adapted to life in the nests of their hosts.

Adaptation, Biological↗

The effect of host heterogeneity and parasite intragenomic interactions on parasite population structure.

Understanding the processes that shape the genetic structure of parasite populations and the functional consequences of different parasite genotypes is critical for our ability to predict how an infection can spread through a host population and for the design of effective vaccines to combat infection and disease. Here, we examine how the genetic structure of parasite populations responds to host genetic heterogeneity. We consider the well-characterized molecular specificity of major histocompatibility complex binding of antigenic peptides to derive deterministic and stochastic models. We use these models to ask, firstly, what conditions favour the evolution of generalist parasite genotypes versus specialist parasite genotypes? Secondly, can parasite genotypes coexist in a population? We find that intragenomic interactions between parasite loci encoding antigenic peptides are pivotal in determining the outcome of evolution. Where parasite loci interact synergistically (i.e. the recognition of additional antigenic peptides has a disproportionately large effect on parasite fitness), generalist parasite genotypes are favoured. Where parasite loci act multiplicatively (have independent effects on fitness) or antagonistically (have diminishing effects on parasite fitness), specialist parasite genotypes are favoured. A key finding is that polymorphism is not stable and that, with respect to functionally important antigenic peptides, parasite populations are dominated by a single genotype.

Animals↗

Parasite mediated mortality and host immune response explain age-related differences in blood parasitism in birds.

An important pattern in host-parasite assemblages is a higher intensity of parasites in juveniles than in adults, but the reasons for these differences remain obscure. Three non-mutually exclusive hypotheses have been proposed: (1) heavily parasitized juveniles die before being recruited into the adult population ('selection' hypothesis); (2) the development of an acquired immunity by the host in front of the parasite reduces the intensity of the parasite in adult hosts ('immunity' hypothesis); and (3) differences in behavior makes adults less exposed to the parasite than juveniles ('vector exposure' hypothesis). Having rejected the 'vector exposure' hypothesis in a previous study, here we tested the 'selection' and 'immunity' hypotheses in feral pigeons (Columba livia) infected by the blood parasite Haemoproteus columbae. In agreement with the 'selection' hypothesis, young (but not adult) pigeons that were highly parasitized had a lower probability of surviving until adulthood, independent of their body condition. However, selection was not strong enough to account for the observed differences in parasite intensity between age-classes, and after selection parasite intensity of survivors still remained 85% higher in juveniles than in adults. In contrast, the 'immunity' hypothesis offered a greater explanatory power. The intensity of blood parasites in young pigeons, but not in adults, decreased over time so dramatically that by the time they had become adults their intensities were indistinguishable from that typically seen in adults. Therefore, while selection against highly parasitized juveniles can contribute to some extent to a reduction in parasitism seen in the adult population, age-specific blood parasitism in feral pigeons is best explained as a transitory phase just before the host develops an effective immune response.

Age Factors↗

Parasite faunas of freshwater fish: the relationship between richness and the specificity of parasites.

The relationship between the host specificity of parasites and the richness of the assemblages in which they occur was examined among the parasite faunas of Canadian freshwater fishes. The prediction tested was that rich faunas would consist of both generalist and specialist parasites, whereas poor faunas would include only generalists, a pattern known as nestedness. An index of nestedness was computed for parasite faunas of fish species in 5 large families (Salmonidae, Cyprinidae, Catostomidae, Centrarchidae and Percidae) and compared with the value expected if parasite faunas are random assemblages of parasites. There was no evidence of nested patterns among any of the 5 families of fish hosts. However, since both measures of host range of parasites (i.e. number of known host species) and richness of parasite faunas are affected by how intensely the different parasites and hosts have been studied, tests of nestedness may be flawed. After correcting both variables for study effort, negative correlations were found between the mean host range of parasites and the richness of the faunas to which they belong. In other words, parasites in rich faunas occurred on average in fewer host species, because of the many specialists, than parasites in poor faunas, which are mainly generalists. This relationship was apparent in all fish but the Salmonidae; fish species in this family have been introduced to new areas much more frequently than other fish, and their parasite faunas have thus had a distinct recent history. The general trend observed in non-salmonid fish suggests that parasite colonization and speciation may have been facilitated in some fish species, but not in others.

Animals↗

Biomathematical enzyme kinetics model of prebiotic autocatalytic RNA networks: degenerating parasite-specific hyperparasite catalysts confer parasite resistance and herald the birth of molecular immunity.

Catalysis and specifically autocatalysis are the quintessential building blocks of life. Yet, although autocatalytic networks are necessary, they are not sufficient for the emergence of life-like properties, such as replication and adaptation. The ultimate and potentially fatal threat faced by molecular replicators is parasitism; if the polymerase error rate exceeds a critical threshold, even the fittest molecular species will disappear. Here we have developed an autocatalytic RNA early life mathematical network model based on enzyme kinetics, specifically the steady-state approximation. We confirm previous models showing that these second-order autocatalytic cycles are sustainable, provided there is a sufficient nucleotide pool. However, molecular parasites become untenable unless they sequentially degenerate to hyperparasites (i.e. parasites of parasites). Parasite resistance-a parasite-specific host response decreasing parasite fitness-is acquired gradually, and eventually involves an increased binding affinity of hyperparasites for parasites. Our model is supported at three levels; firstly, ribozyme polymerases display Michaelis-Menten saturation kinetics and comply with the steady-state approximation. Secondly, ribozyme polymerases are capable of sustainable auto-amplification and of surmounting the fatal error threshold. Thirdly, with growing sequence divergence of host and parasite catalysts, the probability of self-binding is expected to increase and the trend towards cross-reactivity to diminish. Our model predicts that primordial host-RNA populations evolved via an arms race towards a host-parasite-hyperparasite catalyst trio that conferred parasite resistance within an RNA replicator niche. While molecular parasites have traditionally been viewed as a nuisance, our model argues for their integration into the host habitat rather than their separation. It adds another mechanism-with biochemical precision-by which parasitism can be tamed and offers an attractive explanation for the universal coexistence of catalyst trios within prokaryotes and the virosphere, heralding the birth of a primitive molecular immunity.

Kinetics↗

To eject or to abandon? Life history traits of hosts and parasites interact to influence the fitness payoffs of alternative anti-parasite strategies.

Hosts either tolerate avian brood parasitism or reject it by ejecting parasitic eggs, as seen in most rejecter hosts of common cuckoos, Cuculus canorus, or by abandoning parasitized clutches, as seen in most rejecter hosts of brown-headed cowbirds, Molothrus ater. What explains consistent variation between alternative rejection behaviours of hosts within the same species and across species when exposed to different types of parasites? Life history theory predicts that when parasites decrease the fitness of host offspring, but not the future reproductive success of host adults, optimal clutch size should decrease. Consistent with this prediction, evolutionarily old cowbird hosts, but not cuckoo hosts, have lower clutch sizes than related rarely- or newly parasitized species. We constructed a mathematical model to calculate the fitness payoffs of egg ejector vs. nest abandoner hosts to determine if various aspects of host life history traits and brood parasites' virulence on adult and young host fitness differentially influence the payoffs of alternative host defences. These calculations showed that in general egg ejection was a superior anti-parasite strategy to nest abandonment. Yet, increasing parasitism rates and increasing fitness values of hosts' eggs in both currently parasitized and future replacement nests led to switch points in fitness payoffs in favour of nest abandonment. Nonetheless, nest abandonment became selectively more favourable only at lower clutch sizes and only when hosts faced parasitism by a cowbird- rather than a cuckoo-type brood parasite. We suggest that, in addition to evolutionary lag and gape-size limitation, our estimated fitness differences based on life history trait variation provide new insights for the consistent differences observed in the anti-parasite rejection strategies between many cuckoo- and cowbird-hosts.

Animals↗

Behavioural defense against parasites: interaction with parasite invasiveness.

Behavioural patterns involved in avoiding, repelling, or removing parasites allow mammalian hosts to defend themselves from an array of parasites that threaten the host's fitness in the natural world. Some examples of behavioural defenses and the presumed target parasites are: grooming to remove ticks, grouping to reduce attack rate of biting flies, fly repelling movements to reduce parasitic flies, and selective grazing to reduce intake of faecal-borne parasites. These behavioural defenses are discussed with regard to effectiveness in controlling the target parasites. Parasites have sometimes evolved behavioural strategies of evading, penetrating or disabling these behavioural defenses. These parasite behavioural strategies, though less studied, are discussed. Also discussed is the possibility that host behavioural patterns that may defend against one parasite may be exploited by a different type of parasite to facilitate its own transmission. The interplay between host defensive strategies, the cost versus the effectiveness of such strategies, and a parasite's evasion or exploitation of such strategies, may be useful in understanding some aspects of host-parasite dynamics in nature.

Animals↗

Density-dependent parasite establishment suggests infection-associated immunosuppression as an important mechanism for parasite density regulation in onchocerciasis.

The modulation of human immune response by filarial parasites has yielded contradictory experimental findings and attracted much controversy. We address the unresolved question of acquisition, establishment and accumulation of Onchocerca volvulus by using a modelling approach that relates computer simulations to cross-sectional data concerning parasite burdens in 913 West African onchocerciasis patients. It is shown that the acquisition of O. volvulus is not constant with host age; instead, the analysis of age profiles of parasite burdens strongly indicate the operation of immunosuppressive processes within the human host, associated with the presence of adult parasites or microfilariae. It is suggested that these processes suppress immunity against incoming infective larvae (L3), which themselves act as an immune modulating component once they have successfully overcome the barrier of concomitant immunity. Suppression of parasite-specific immunity leads to parasite establishment rates which increase along with the parasite burden, but which hardly depend on hyperendemic annual transmission potentials. Children, still immunocompetent due to low parasite burdens, acquire 0.1-0.5 adult female parasites per year, whereas older people, immunosuppressed due to high burdens, acquire 2-4 adult female parasites per year. Differences in parasite establishment between the forest and the savannah strains of O. volvulus are quantified and dynamic aspects of density-dependent parasite establishment discussed.

Adolescent↗

Host-parasite coevolution: comparative evidence for covariation of life history traits in primates and oxyurid parasites.

The environmental factors that drive the evolution of parasite life histories are mostly unknown. Given that hosts provide the principal environmental features parasites have to deal with, and given that these features (such as resource availability and immune responses) are well characterized by the life history of the host, we may expect natural selection to result in covariation between parasite and host life histories. Moreover, some parasites show a high degree of host specificity, and cladistic analyses have shown that host and parasite phylogenies can be highly congruent. These considerations suggest that parasite and host life histories may covary. The central argument in the theory of life history evolution concerns the existence of trade-offs between traits. For parasitic nematodes it has been shown that larger body sizes induce higher fecundity, but this is achieved at the expense of delayed maturity. As high adult mortality would select for reduced age at maturity, the selective benefit of increased fecundity is expressed only if adult mortality is low. Parasite adult mortality may depend on a number of factors, including host longevity. Here we tested the hypothesis concerning the positive covariation between parasite body size (which reflects parasite longevity) and host longevity. To achieve this goal, we used the association between the pinworms (Oxyuridae, Nematoda) and their primate hosts. Oxyurids are highly host specific and are supposed to be involved in a coevolutionary process with their hosts. We found that female parasite body length was positively correlated with host longevity after correcting for phylogeny and host body mass. Conversely, male parasite body length and host longevity were not correlated. These results confirm that host longevity may represent a constraint on the evolution of body size in oxyurids, at least in females. The discrepancy between female and male oxyurids is likely to depend on the particular mode of reproduction of this taxon (haplodiploidy), which should result in weak (or even null) selection pressures to an increase of body size in males.

Animals↗

Host-parasite dynamics and the evolution of host immunity and parasite fecundity strategies.

We explore evolutionarily stable co-evolution of host-macroparasite++ interactions in a discrete-time two-species population dynamics model, in which the dynamics may be stable, cyclic or chaotic. The macroparasites are assumed to harm host individuals through decreased reproductive output. Hosts may develop costly immune responses to defend themselves against parasites. Parasites compete with conspecifics by adjusting their fecundities. Overall, the presence of both parasites and the immune response in hosts produces more stable dynamics and lower host population sizes than that observed in the absence of the parasites. In our evolutionary analyses, we show that maximum parasite fecundity is always an evolutionarily stable strategy (ESS), irrespective of the type of population interaction, and that maximum parasite fecundity generally induces a minimum parasite population size through over-exploitation of the host. Phenotypic polymorphisms with respect to immunity in the host species are common and expected in ESS host strategies: the benefits of immunication depend on the frequency of the immune hosts in the population. In particular, the steady-state proportions of immune hosts depend, in addition to all the parameters of the parasite dynamics only on the cost of immunity and on the virulence of parasites in susceptible hosts. The implicit ecological dynamics of the host-parasite interaction affect the proportion of immune host individuals in the population. Furthermore, when changes in certain population parameters cause the dynamics of the host-parasite interaction to move from stability to cyclicity and then to chaos, the proportion of immune hosts tend to decrease; however, we also detected counter-examples to this result. As a whole, incorporating immunological and genetic aspects, as well as life-history trade-offs, into host-macroparasite dynamics produces a rich extension to the patterns observed in the models of ecological interactions and epidemics, and deserves more attention than is currently the case.

Animals↗

The mortality and fecundity of Haemonchus contortus in parasite-naive and parasite-exposed sheep following single experimental infections.

Parasite-exposed lambs and their parasite-naive controls were experimentally infected once only with 30,000 H. contortus larvae at 3, 9, 12, and 20 weeks following termination of a moderate immunizing infection of 30,000 H. contortus larvae. Previously exposed lambs, challenged at 3 weeks, had a significant reduction in the total H. contortus worm burden as compared to parasite-naive controls. No difference in the total H. contortus worm burden was found between parasite-exposed or parasite-naive lambs challenged at 9 weeks or thereafter. Female worms were found to be significantly smaller in lambs previously exposed to the parasite as compared to those found in parasite-naive lambs. The average parasite fecundity was 4700 eggs per female worm per day. Previous exposure of the lambs to the parasite had no effect on parasite fecundity. Various mathematical models were used to examine parasite fecundity. Parasite fecundity was found to increase in the initial post-challenge period reaching a constant value approximately 58 days after challenge infection. No density-dependent constraints on fecundity were observed.

Animals↗

Discrimination by female mice between the odours of parasitized and non-parasitized males.

The detection and avoidance of parasitized males has been proposed to be a component of female mate choice. We investigated whether or not female laboratory mice, Mus musculus domesticus, could discriminate between parasitized and non-parasitized males on the basis of odour. Female mice were given a choice between the urine and other odorous secretions of either a male mouse sub-clinically infected for five days with the naturally occurring, enteric, single host, protozoan parasite, Eimeria vermiformis, or an uninfected male. Females showed a marked preference for the odours of non-parasitized male mice over those of the parasitized males; as measured by number of investigations, time spent per investigation, and total investigation time of the odours in a choice situation. Female mice also displayed an overwhelming initial, or first choice, preference for the odours of the non-parasitized male mice over those of the parasitized males. These observations show that female mice can distinguish between the odours of parasitized and non-parasitized males, and discriminate against parasitized males on the basis of odour. We suggest that the detection and avoidance of infected males by female mice through odour cues may function to reduce parasite transmission and potentially serve as a component of female mate selection or choice.

Animals↗

Relative number of generations of hosts and parasites does not influence parasite local adaptation in coevolving populations of bacteria and phages.

A potential consequence of host-parasite coevolution in spatially structured populations is parasite local adaptation: local parasites perform better than foreign parasites on their local host populations. It has been suggested that the generally shorter generation times of parasites compared with their hosts contributes to parasites, rather than hosts, being locally adapted. We tested the hypothesis that relative generation times of hosts and parasites affect local adaptation of hosts and parasites, using the bacterium Pseudomonas fluorescens and a lytic phage as host and parasite, respectively. Generation times were not directly manipulated, but instead one of the coevolving partners was regularly removed and replaced with a population from an earlier time point. Thus, one partner underwent more generations than the other. Manipulations were carried out at both early and later periods of coevolutionary interactions. At early stages of coevolution, host and parasites that underwent relatively more generations displayed higher levels of resistance and infectivity, respectively. However, the relative number of generations that bacteria and phages underwent did not change the level of local adaptation relative to control populations. This is likely because generalist hosts and parasites are favoured during early stages of coevolution, preventing local adaptation. By contrast, at later stages manipulations had no effect on either average levels of resistance or infectivity, or alter the level of local adaptation relative to the controls, possibly because traits other than resistance and infectivity were under strong selection. Taken together, these data suggest that the relative generation times of hosts and parasites may not be an important determinant of local adaptation in this system.

Adaptation, Biological↗

Relative infection levels and taxonomic distances among the host species used by a parasite: insights into parasite specialization.

Parasites often exploit more than one host species at any stage in their life-cycle, but the extent to which these host species are used varies greatly. Parasites typically achieve their highest prevalence, intensity and/or abundance in one host species (the principal host), whereas infection levels in auxiliary hosts range from relatively high to very low. The present study examines what influences the distribution of parasite individuals among their different host species, using metazoan parasites that use freshwater fish as their definitive or only host. Specifically, I test the hypothesis that differences in relative infection levels by a parasite among its auxiliary hosts are proportional to the taxonomic distance between the respective auxiliary hosts and the parasite's principal host. Taxonomic distance among hosts is a surrogate measure of their similarity in terms of ecology, physiology and immunology. Using data on 29 parasite species and 6 fish communities, for a total of 47 parasite-locality combinations, it was found that taxonomic distance between the auxiliary hosts and the principal host had no real influence on infection levels in auxiliary hosts, measured as either prevalence, intensity or abundance. The analysis revealed differences in the degree of specialization among major groups of parasites: in terms of abundance or intensity, auxiliary hosts were less important for cestodes than for nematodes and copepods. The lack of an effect of taxonomic distance may indicate that ecological similarity among host species, arising from convergence and not from relatedness, is more important than host phylogeny or taxonomy. Although the results are based on a limited number of parasite taxa, they suggest that parasites may be opportunistic in their colonization of new hosts, and not severely constrained by evolutionary baggage.

Animals↗

Parasite assemblages of crucian carp (Carassius carassius)--is depauperate composition explained by lack of parasite exchange, extreme environmental conditions or host unsuitability?

Recent studies on parasite communities have considered the factors which contribute to variability in parasite infections among populations of one host species. This paper examined parasite assemblages of crucian carp (Carassius carassius) in a natural set-up of two distinctive environments inhabited by the same host species and differing in respect to composition of other fish species: ponds, where extreme conditions prevent other fish species from occurring and lakes, where crucian carp coexist with other fish species. Our aim was to evaluate the significance of parasite exchange, environmental conditions and host suitability for the depauperate parasite assemblages observed in pond crucian carp. As predicted, crucian carp had more diverse parasite species composition in lakes, which supports the hypotheses of increased opportunities for parasite exchange with other fish species and better environmental conditions for the parasites. However, for instance, diplostomids were found only from few fish individuals in very low numbers. Experimental exposure trials with the eye fluke Diplostomum spathaceum in the laboratory indicated that crucian carp were totally resistant to infection, which is exceptional and suggests that the low number of these parasites in crucian carp was due to physiological unsuitability of the host. To obtain a better understanding on the mechanisms underlying the formation of parasite assemblages in these fish, further studies on the relationships between the unique physiology of crucian carp and infection success by other parasite species are needed.

Acanthocephala↗

Interactions between the parasite's previous and current environment mediate the outcome of parasite infection.

The study of parasite virulence has generally focused on the conditions under which virulence is expected to increase or decrease over time and how the interactions between hosts and their environments may mediate the outcome of infection. Recently, parasite traits such as transmission, offspring production, and development have also been shown to be influenced by environmental variation. What is unclear is how variation in the parasite's environment may impact virulence. Recent theory demonstrates that plasticity can promote the evolution of decreased virulence; thus, understanding whether the parasite's environment can mediate virulence can improve predictions regarding the outcome of parasite infection. Here, an obligate mosquito parasite was reared in hosts fed high or low levels of food. Parasite oocysts (offspring) produced in these two host environments were subsequently fed to uninfected hosts. Parasites originating from well-fed hosts were found to be more virulent to these subsequent hosts compared to parasites originating from poorly fed hosts. Additionally, this effect was apparent only when current hosts were food deprived. These results demonstrate that parasite virulence was mediated by a cross-generational effect of the environment and that the overall outcome of infection was modified by variation in both the parasite's and host's environments.

Aedes↗