Populational variability of Proteocephalus longicollis (Cestoda: Proteocephalidae) in the vendace Coregonus albula from lakes of Karelia

Populational variability of Proteocephalus longicollis from the vendace inhabiting various types of Karelian water bodies was studied. High phenotypic diversity of the helminth and polymorphism in the cestode's major functional complexes - attachment, trophic and reproduction - are established. High degree of phenotype pool realization was determined for various P. longicollis populations. Traits belonging to different functional complexes were shown to have different variability patterns. The highest stability was shown for the attachment organs of P. longicollis, which had similar values of qualitative indices and nature of variation. The traits of the trophic and reproduction complexes are more variable. The conclusion was made that the high intrapopulation diversity of P. longicollis is a response to high ecological variability of the vendace, and the distinctions in the phenotype variability of the helminth reflect the differences between population development patterns of the parasite (cestode P. longicollis) and its host (vendace Coregonus albula) in lakes of Karelia.     

Common sculpin Cottus gobio as a natural paratenic host of Proteocephalus longicollis (Cestoda: Proteocephalidae), a parasite of salmonids, in Europe.

Common sculpins Cottus gobio L. (Pisces: Cottidae), from the Mlynsky Brook near Ceské Zleby in the Sumava National Park, southwestern Bohemia, Czech Republic, were found to harbour in their intestines juvenile cestodes Proteocephalus longicollis (Zeder, 1800), a common parasite of holarctic salmonids, with a prevalence of 60% and intensity of 1 to 11 (mean 5) parasites per fish; undoubtedly, these prey fish serve as paratenic hosts. In this locality, the definitive host of P. longicollis is the brown trout Salmo trutta m. fario L., large specimens of which apparently acquire infection of this parasite by feeding on infected sculpins. C. gobio is the first known natural paratenic host of P. longicollis in Europe.     

Mediterranean Diplodus annularis (Teleostei: Sparidae) and its brain parasite: unforeseen outcome

Patterns of parasite load and aggregation of the bird trematode Cardiocephaloides longicollis in its main intermediate host in the Mediterranean, the annular sea bream, Diplodus annularis, were studied in a large sample collected off Valencia (Spain) and are discussed within the context of the parasite induced host mortality hypothesis. The metacercariae were located within large composite cysts of host origin in the ventricles of the optic lobes of the cerebrum. A weak immunological response was detected in older fish, which was significantly associated with the total parasite load. Although the mean abundance of C. longicollis showed a tendency to increase with host size, the infection levels were generally homogeneous with a noticeable plateauing in the intermediate size classes. The distribution of the metacercariae was aggregated and agreed with the negative binomial distribution. There was a marked decline in parasite aggregation in the largest size-class, suggesting parasite-induced host mortality in the oldest fish possibly due to predation by large non-host fish predators. On the other hand, levelling off of abundance and decrease in heterogeneity of parasite distribution within the intermediate age cohort could indicate that these sizes are being rapidly and/or constantly removed from the host population due to by-catch fishing. The overall high infection levels and the continuous recruitment across age cohorts provides evidence that an enhanced parasite transmission is taking place in the Gulf of Valencia due to increased spatial overlap of the hosts involved in the life cycle. We suggest a human-induced facilitation of the digenean life cycle due to the fact that gulls in the area feed extensively on discards, thus indicating the possibility of an unforeseen effect of fishing practices in a marine littoral system.     

Ultrastructural Characterisation and Molecular Taxonomic Identification of Nosema granulosis n. sp., a Transovarially Transmitted Feminising (TTF) Microsporidium

A novel microsporidian parasite is described, which infects the crustacean host Gammarus duebeni. The parasite was transovarially transmitted and feminised host offspring. The life cycle was monomorphic with three stages. Meronts were found in host embryos, juveniles, and in the gonadal tissue of adults. Sporoblasts and spores were restricted to the gonad. Sporogony was disporoblastic giving rise to paired sporoblasts, which then differentiated to form spores. Spores were not found in regular groupings and there was no interfacial envelope. Spores were approximately 3.78 x 1.22 microns and had a thin exospore wall, a short polar filament, and an unusual granular polaroplast. All life cycle stages were diplokaryotic. A region from the parasite small subunit ribosomal RNA gene was amplified and sequenced. Phylogenetic analysis based on these data places the parasite within the genus Nosema. We have named the species Nosema granulosis based on the structure of the polaroplast.     

Ecological parasitology of polymorphic Arctic charr, Salvelinus alpinus (L.), in Thingvallavatn, Iceland

The helminth endoparasite fauna in four Arctic charr morphs, Salvelinus alpinus (L.), small benthivorous (SB), large benthivorous (LB), planktivorous (PL) and piscivorous (PI) charr, from Thingvallavatn, Iceland consisted of: Crepidostomum farionis (Trematoda: Allocreadiidae); Diplosttomum sp. (Trematoda: Diplostomatidae); Eubothrium salvelini; Diphyllobothrium dendriticum; D. ditremum (Cestoda: Pseudophyllidae); Proteocephalus longicollis (Cestoda: Proteocepha-lidae): and Philonema oncorhynchi (Nematoda: Filariidae). The morphs exhibited distinctive patterns in prevalences and parasite burdens (mean intensity and mean relative density of parasites). SB charr had high prevalence and parasite burden of the eye fluke Diplostomum sp. and none to very light infections of the other parasite species. LB charr had relatively high prevalence and parasite burden of the intestinal fluke C. farionis , whereas infections of the remaining parasite species were light to moderate. PL and PI charr had high prevalences and worm burdens of Diphyllobothrium spp. and P. longicollis . PL charr differed from PI charr in higher worm burden off P. longicollis and lighter burden of £. salvelini . Prevalences of P. oncorhynchi were high in PL and PI charr. Association of parasite intensities and age and length offish were investigated. The different infection patterns among the morphs agree well with their partitioning in food and habitat utilization, and confirm that there is a high degree of ecological segregation between the morphs. The results demonstrate the importance of ecological factors influencing transmission efficiency of parasites to the fish host.     

Comparative genetic diversity of parasites and their hosts: population structure of an urban cockroach and its haplo-diploid parasite (oxyuroid nematode)

Few studies have investigated the genetic structure of both host and parasite populations at a level of populations and at a level of individuals. We investigated the genetic structure of the urban cockroach Blattella germanica and its oxyuroid parasite Blatticola blattae. Random amplified polymorphic DNA (RAPD) markers were used to quantify genetic diversity between and within four populations (from two cities in France) of the host and its parasite. Diversity based on phenotypic frequencies was calculated for each RAPD marker using Shannon-Wiener's index. We used multivariate analyses to test the significance of genetic differentiation between host and parasite populations. Analysis of molecular variance was also used. Both methods gave similar results. Diversity between pairs of individuals was estimated by Nei & Li's index. Genetic diversity was higher within host or parasite populations (80% and 82%, respectively, of explained diversity) than between host or parasite populations (20% and 18%, respectively, explained diversity). The genetic distances between pairs of parasite populations (or individuals) were not correlated with the genetic distances between the corresponding pairs of host populations (or individuals).     

Relationships among members of the genus Myxobolus (Myxozoa: Bilvalvidae) based on small subunit ribosomal DNA sequences

Sequences representing approximately 1,700 base pairs of the 18S rRNA gene from 10 different species in the genus Myxobolus were found to group them into 3 clusters that showed little correlation with spore morphology and size or host specificity, criteria currently used for both higher and lower taxonomic placements in the Myxozoa. Of the phenotypic criteria examined, tissue tropism was most correlated with the rRNA groupings observed. Spores of similar size and shape (Myxobolus cerebralis vs. Myxobolus squamalis) were distantly related in some instances, whereas spores with divergent morphology and size were sometimes found to be closely related (M. cerebralis and Myxobolus insidiosus). These initial investigations into the phylogenetic relationships of putative members of the genus Myxobolus clearly indicate the potential limitations of groupings based on size and morphological properties of the spores and host species infected. We propose that 18S rRNA gene sequences, combined with information on tissue tropism, host species infected, and developmental cycles in the fish and alternate host (when and if known) be given greater consideration in taxonomic placements of myxosporeans.     

Host use dynamics in a heterogeneous fitness landscape generates oscillations in host range and diversification

Colonization of novel hosts is thought to play an important role in parasite diversification, yet little consensus has been achieved about the macroevolutionary consequences of changes in host use. Here, we offer a mechanistic basis for the origins of parasite diversity by simulating lineages evolved in silico. We describe an individual‐based model in which (i) parasites undergo sexual reproduction limited by genetic proximity, (ii) hosts are uniformly distributed along a one‐dimensional resource gradient, and (iii) host use is determined by the interaction between the phenotype of the parasite and a heterogeneous fitness landscape. We found two main effects of host use on the evolution of a parasite lineage. First, the colonization of a novel host allowed parasites to explore new areas of the resource space, increasing phenotypic and genotypic variation. Second, hosts produced heterogeneity in the parasite fitness landscape, which led to reproductive isolation and therefore, speciation. As a validation of the model, we analyzed empirical data from Nymphalidae butterflies and their host plants. We then assessed the number of hosts used by parasite lineages and the diversity of resources they encompass. In both simulated and empirical systems, host diversity emerged as the main predictor of parasite species richness.     

Parasite fauna of the European river lamprey Lampetra fluviatilis (L.) from Lake Onega

Data on the parasite fauna of the adult European river lamprey Lampetra fluviatilis (L.) from Lake Onega are reported. Ten parasite species are found, including trematodes Diplostomum petromyzifluviatilis and D. spathaceum (metacercariae), cestode Proteocephalus longicollis, nematodes Cucullanus truttae and Raphidascaris acus, acanthocephalan Echinorhynchus salmonis, ectoparasitic infusoria Chilodonella hexastica, Trichodina tenuidens, and Trichodinella epizootica, and fungus Saprolegnia parasitica. Three species are found to be dominate, namely D. petromyzifluviatilis, Cucullanus truttae, and P. longicollis. Comparative analysis of the parasite faunas of the lampreys from other basins is carried out. Some similarity in the parasite faunas of lampreys and salmonids is discovered.     

Parasite-host specificity: experimental studies on the basis of parasite adaptation

Specificity in parasitic interactions can be defined by host genotypes that are resistant to only a subset of parasite genotypes and parasite genotypes that are infective on a subset of host genotypes. It is not always clear if specificity is determined by the genotypes of the interactors, or if phenotypic plasticity (sometimes called acclimation) plays a larger role. Coevolutionary outcomes critically depend on the pervasiveness of genetic interactions. We studied specificity using the bacterial parasite Pasteuria ramosa and its crustacean host Daphnia magna. First, we tested for short-term adaptation of P. ramosa lines that had been rapidly shifted among different host genotypes. Adaptation at this time-scale would demonstrate the contribution of phenotypic plasticity to specificity. We found that infectivity was stable across lines irrespective of recent passage history, indicating that in the short term infection outcomes are fixed by genetic backgrounds. Second, we studied longer-term evolution with two host clones and two parasite lines. In this experiment, P. ramosa lines had the possibility to evolve adaptations to the host genotype (clone) in which they were serially passaged, which allowed us to test for a genetic component to specificity. Substantial differences arose in the two passaged lines: one parasite line gained infectivity on the host clone it was grown on, but it lost infectivity on the other host genotype (this line evolved specificity), while the other parasite line evolved higher infectivity on both host clones. We crossed the two host genotypes used in the serial passage experiment and found evidence that the number of host genes that underlies resistance variation is small. In sum, our results show that P. ramosa specificity is a stably inherited trait, it can evolve rapidly, and it is controlled by few genes in the host. These findings are consistent with the idea of a rapid, ongoing arms race between the bacterium and its host.     

Avian malaria infection intensity influences mosquito feeding patterns

Pathogen-induced host phenotypic changes are widespread phenomena that can dramatically influence host–vector interactions. Enhanced vector attraction to infected hosts has been reported in a variety of host–pathogen systems, and has given rise to the parasite manipulation hypothesis whereby pathogens may adaptively modify host phenotypes to increase transmission from host to host. However, host phenotypic changes do not always favour the transmission of pathogens, as random host choice, reduced host attractiveness and even host avoidance after infection have also been reported. Thus, the effects of hosts’ parasitic infections on vector feeding behaviour and on the likelihood of parasite transmission remain unclear. Here, we experimentally tested how host infection status and infection intensity with avian Plasmodium affect mosquito feeding patterns in house sparrows (Passer domesticus). In separate experiments, mosquitoes were allowed to bite pairs containing (i) one infected and one uninfected bird and (ii) two infected birds, one of which treated with the antimalarial drug, primaquine. We found that mosquitoes fed randomly when exposed to both infected and uninfected birds. However, when mosquitoes were exposed only to infected individuals, they preferred to bite the non-treated birds. These results suggest that the malarial parasite load rather than the infection itself plays a key role in mosquito attraction. Our findings partially support the parasite manipulation hypothesis, which probably operates via a reduction in defensive behaviour, and highlights the importance of considering parasite load in studies on host–vector–pathogen interactions.     

Host availability and the evolution of parasite life-history strategies

Parasites exploit an inherently patchy resource, their hosts, which are discrete entities that may only be available for infection within a relatively short time window. However, there has been little consideration of how heterogeneities in host availability may affect the phenotypic or genotypic composition of parasite populations or how parasites may evolve to cope with them. Here we conduct a selection experiment involving an entomopathogenic nematode (Steinernema feltiae) and show for the first time that the infection rate of a parasite can evolve rapidly to maximize the chances of infecting within an environment characterized by the rate of host availability. Furthermore, we show that the parasite's infection rate trades off with other fitness traits, such as fecundity and survival. Crucially, the outcome of competition between strains with different infection strategies depends on the rate of host availability; frequently available hosts favor "fast" infecting nematodes, whereas infrequently available hosts favor "slow" infecting nematodes. A simple evolutionarily stable strategy (ESS) analysis based on classic epidemiological models fails to capture this behavior, predicting instead that the fastest infecting phenotype should always dominate. However, a novel model incorporating more realistic, discrete bouts of host availability shows that strain coexistence is highly likely. Our results demonstrate that heterogeneities in host availability play a key role in the evolution of parasite life-history traits and in the maintenance of phenotypic variability. Parasite life-history strategies are likely to evolve rapidly in response to changes in host availability induced by disease management programs or by natural dynamics in host abundance. Incorporating parasite evolution in response to host availability would therefore enhance the predictive ability of current epidemiological models of infectious disease.     

Experimental evolution with a multicellular host causes diversification within and between microbial parasite populations—Differences in emerging phenotypes of two different parasite strains

Host‐parasite coevolution is predicted to have complex evolutionary consequences, potentially leading to the emergence of genetic and phenotypic diversity for both antagonists. However, little is known about variation in phenotypic responses to coevolution between different parasite strains exposed to the same experimental conditions. We infected Caenorhabditis elegans with one of two strains of Bacillus thuringiensis and either allowed the host and the parasite to experimentally coevolve (coevolution treatment) or allowed only the parasite to adapt to the host (one‐sided parasite adaptation). By isolating single parasite clones from evolved populations, we found phenotypic diversification of the ancestral strain into distinct clones, which varied in virulence toward ancestral hosts and competitive ability against other parasite genotypes. Parasite phenotypes differed remarkably not only between the two strains, but also between and within different replicate populations, indicating diversification of the clonal population caused by selection. This study highlights that the evolutionary selection pressure mediated by a multicellular host causes phenotypic diversification, but not necessarily with the same phenotypic outcome for different parasite strains.     

Description of embryonic development and ultrastructure in miracidia of Cardiocephaloides longicollis (Digenea,Strigeidae) in relation to active host finding strategy in a marine environment

The functional ultrastructure and embryonic development of miracidia in naturally released eggs of the trematode Cardiocephaloides longicollis were studied using light and transmission electron microscopy. This species has operculated eggs and embryogenesis occurs in the marine environment before an actively infecting ciliated miracidium hatches. Six different developmental stages were identified. The lack of pores in the eggshell indicates its impermeability and the miracidium's dependency on glycogen nutritive reserves, contained in numerous vitellocytes in early embryos. As the development advances, these merge into larger vitelline vacuoles that encircle the miracidium and may aid its hatching. Tissue and primary organ differentiation were observed in advanced stages, i.e., terebratorium, glands, cerebral ganglion, peripheral sensory endings, and eyespots. The anterior part of the body contains a single apical and paired lateral glands, as well as two types of sensory endings, which permit location, adhesion, and penetration of the host. No previous studies describe the embryonic development and ultrastructure of miracidia in strigeids, however, some of the structural features shared with other, well described species with unknown life cycles are emphasised. This study highlights that ultrastructural data have to be interpreted in relation to parasite biology to understand the structural requirements of specific parasite strategies.     

Phylogenetic analysis of European species of Proteocephalus (Cestoda: Proteocephalidea): compatibility of molecular and morphological data, and parasite-host coevolution.

The phylogeny of European species of the tapeworm genus Proteocephalus was studied, based on partial 18S rDNA and morphological data. The group was found to be monophyletic. The analysis showed a low informative value of available morphological characters in comparison with molecular data. The morphological matrix resulted in a poorly resolved tree which is, however, compatible with the topology (Proteocephalus osculatus (Proteocephalus torulosus (Proteocephalus macrocephalus, Proteocephalus filicollis) (Proteocephalus tetrastomus, Proteocephalus percae, Proteocephalus longicollis))) based on the 18S rDNA data. A comparison performed by the program TreeMap showed a lack of significant congruency between parasite and host phylogenies. Therefore, the distribution of species in their hosts appears to be independent of the phylogeny and it is likely to be a result of host-switching, rather than co-speciation events.     

Evolution of phenotypic fluctuation under host-parasite interactions

Robustness and plasticity are essential features that allow biological systems to cope with complex and variable environments. In a constant environment, robustness, i.e., insensitivity of phenotypes, is expected to increase, whereas plasticity, i.e., the changeability of phenotypes, tends to diminish. Under a variable environment, existence of plasticity will be relevant. The robustness and plasticity, on the other hand, are related to phenotypic variances. As phenotypic variances decrease with the increase in robustness to perturbations, they are expected to decrease through the evolution. However, in nature, phenotypic fluctuation is preserved to a certain degree. One possible cause for this is environmental variation, where one of the most important “environmental” factors will be inter-species interactions. As a first step toward investigating phenotypic fluctuation in response to an inter-species interaction, we present the study of a simple two-species system that comprises hosts and parasites. Hosts are expected to evolve to achieve a phenotype that optimizes fitness. Then, the robustness of the corresponding phenotype will be increased by reducing phenotypic fluctuations. Conversely, plasticity tends to evolve to avoid certain phenotypes that are attacked by parasites. By using a dynamic model of gene expression for the host, we investigate the evolution of the genotype-phenotype map and of phenotypic variances. If the host–parasite interaction is weak, the fittest phenotype of the host evolves to reduce phenotypic variances. In contrast, if there exists a sufficient degree of interaction, the phenotypic variances of hosts increase to escape parasite attacks. For the latter case, we found two strategies: if the noise in the stochastic gene expression is below a certain threshold, the phenotypic variance increases via genetic diversification, whereas above this threshold, it is increased mediated by noise-induced phenotypic fluctuation. We examine how the increase in the phenotypic variances caused by parasite interactions influences the growth rate of a single host, and observed a trade-off between the two. Our results help elucidate the roles played by noise and genetic mutations in the evolution of phenotypic fluctuation and robustness in response to host–parasite interactions.     

Successional features of the dynamics of abundance and population structure of the cestode Proteocephalus longicollis (Zeder, 1800) (Cestoda: Proteocephalidae)

Dynamics of abundance and changes of populations structure were investigated in a host-specific parasite of salmonids, cestode Proteocephalus longicollis from the vendace Coregonus albula L. of the Syamozero Lake during the period of more than forty years. A relative stability of the values of infestation indices independently of significant changes in the abundance and population structure of the vendace is established. Changes in the distribution of abundance and population structure of P. longicollis are shown to be connected with the change of the water body's status resulting from eutrophication. These changes can be considered as reliable indices of the succession in freshwater bodies.     

Genotypic and phenotypic variation in transmission traits of a complex life cycle parasite

Characterizing genetic variation in parasite transmission traits and its contribution to parasite vigor is essential for understanding the evolution of parasite life‐history traits. We measured genetic variation in output, activity, survival, and infection success of clonal transmission stages (cercaria larvae) of a complex life cycle parasite (Diplostomum pseudospathaceum). We further tested if variation in host nutritional stage had an effect on these traits by keeping hosts on limited or ad libitum diet. The traits we measured were highly variable among parasite genotypes indicating significant genetic variation in these life‐history traits. Traits were also phenotypically variable, for example, there was significant variation in the measured traits over time within each genotype. However, host nutritional stage had no effect on the parasite traits suggesting that a short‐term reduction in host resources was not limiting the cercarial output or performance. Overall, these results suggest significant interclonal and phenotypic variation in parasite transmission traits that are not affected by host nutritional status.     

THE IMPACT OF SPATIAL SCALE AND HABITAT CONFIGURATION ON PATTERNS OF TRAIT VARIATION AND LOCAL ADAPTATION IN A WILD PLANT PARASITE

Theory indicates that spatial scale and habitat configuration are fundamental for coevolutionary dynamics and how diversity is maintained in host–pathogen interactions. Yet, we lack empirical data to translate the theory to natural host–parasite systems. In this study, we conduct a multiscale cross‐inoculation study using the specialist wild plant pathogen Podosphaera plantaginis on its host plant Plantago lanceolata. We apply the same sampling scheme to a region with highly fragmented (Åland) and continuous (Saaremaa) host populations. Although theory predicts higher parasite virulence in continuous regions, we did not detect differences in traits conferring virulence among the regions. Patterns of adaptation were highly scale dependent. We detected parasite maladaptation among regions, and among populations separated by intermediate distances (6.0–40.0 km) within the fragmented region. In contrast, parasite performance did not vary significantly according to host origin in the continuous landscape. For both regions, differentiation among populations was much larger for genetic variation than for phenotypic variation, indicating balancing selection maintaining phenotypic variation within populations. Our findings illustrate the critical role of spatial scale and habitat configuration in driving host–parasite coevolution. The absence of more aggressive strains in the continuous landscape, in contrast to theoretical predictions, has major implications for long‐term decision making in conservation, agriculture, and public health.     

Morphological variation in the cosmopolitan fish parasite Neobenedenia girellae (Capsalidae: Monogenea)

Intra-species morphological variation presents a considerable problem for species identification and can result in taxonomic confusion. This is particularly pertinent for species of Neobenedenia which are harmful agents in captive fish populations and have historically been identified almost entirely based on morphological characters. This study aimed to understand how the morphology of Neobenedenia girellae varies with host fish species and the environment. Standard morphological features of genetically indistinct parasites from various host fish species were measured under controlled temperatures and salinities. An initial field-based investigation found that parasite morphology significantly differed between genetically indistinct parasites infecting various host fish species. The majority of the morphological variation observed (60%) was attributed to features that assist in parasite attachment to the host (i.e. the posterior and anterior attachment organs and their accessory hooks) which are important characters in monogenean taxonomy. We then experimentally examined the effects of the interaction between host fish species and environmental factors (temperature and salinity) on the morphology of isogenic parasites derived from a single, isolated hermaphroditic N. girellae infecting barramundi, Lates calcarifer. Experimental infection of L. calcarifer and cobia, Rachycentron canadum, under controlled laboratory conditions did not confer host-mediated phenotypic plasticity in N. girellae, suggesting that measured morphological differences could be adaptive and only occur over multiple parasite generations. Subsequent experimental infection of a single host species, L. calcarifer, at various temperatures (22, 30 and 32?°C) and salinities (35 and 40‰) showed that in the cooler environments (22?°C) N. girellae body proportions were significantly smaller compared with warmer temperatures (30 and 32?°C; P?<?0.0001), whereas salinity had no effect. This is evidence that temperature can drive phenotypic plasticity in key taxonomic characters of N. girellae under certain environmental conditions.