Different metazoan parasites, different transcriptomic responses, with new insights on parasitic castration by digenetic trematodes in the schistosome vector snail Biomphalaria glabrata.

BACKGROUND: Gastropods of the genus Biomphalaria (Family Planorbidae) are exploited as vectors by Schistosoma mansoni, the most common causative agent of human intestinal schistosomiasis. Using improved genomic resources, overviews of how Biomphalaria responds to S. mansoni and other metazoan parasites can provide unique insights into the reproductive, immune, and other systems of invertebrate hosts, and their responses to parasite challenges. RESULTS: Using Illumina-based RNA-Seq, we compared the responses of iM line B. glabrata at 2, 8, and 40 days post-infection (dpi) to single infections with S. mansoni, Echinostoma paraensei (both digenetic trematodes) or Daubaylia potomaca (a nematode parasite of planorbid snails). Responses were compared to unexposed time-matched control snails. We observed: (1) each parasite provoked a distinctive response with a predominance of down-regulated snail genes at all time points following exposure to either trematode, and of up-regulated genes at 8 and especially 40dpi following nematode exposure; (2) At 2 and 8dpi with either trematode, several snail genes associated with gametogenesis (particularly spermatogenesis) were down-regulated. Regarding the phenomenon of trematode-mediated parasitic castration in molluscs, we define for the first time a complement of host genes that are targeted, as early as 2dpi when trematode larvae are still small; (3) Differential gene expression of snails with trematode infection at 40dpi, when snails were shedding cercariae, was unexpectedly modest and revealed down-regulation of genes involved in the production of egg mass proteins and peptide processing; and (4) surprisingly, D. potomaca provoked up-regulation at 40dpi of many of the reproduction-related snail genes noted to be down-regulated at 2 and 8dpi following trematode infection. Happening at a time when B. glabrata began to succumb to D. potomaca, we hypothesize this response represents an unexpected form of fecundity compensation. We also document expression patterns for other Biomphalaria gene families, including fibrinogen domain-containing proteins (FReDs), C-type lectins, G-protein coupled receptors, biomphalysins, and protease and protease inhibitors. CONCLUSIONS: Our study is relevant in identifying several genes involved in reproduction that are targeted by parasites in the vector snail B. glabrata and that might be amenable to manipulation to minimize their ability to serve as vectors of schistosomes.

Brobdingnagians and Goliaths: two forms of gigantism in fish.

Two forms of gigantism are differentiated in fish, Brobdingnagian and Goliathan gigantism, the former applying to populations whose individuals are all larger than is typical for the taxon, the latter to single individuals within a population. While Brobdingnagian gigantism is largely explained by various ecological and evolutionary rules, Goliathan gigantism is not. A mechanistic hypothesis is proposed which explains Goliathan gigantism in terms of the reduction of oxygen requirements of individual fish via moving to cooler temperatures and/or acquiring larger, more energy-dense prey, which enable them to get bigger, and, in the process, sometimes generate bimodal size distributions that may qualify as gradual forms between Goliathan and Brobdingnagian gigantism. This mechanism, which relies on the manner in which their gill surface area grows, is more likely to operate in fish that can get big in the first place than in fish that remain small.

Invasive slipper limpets Crepidula fornicata are hosts for sterilizing digenean parasites.

Invasion and spread of alien species can drive ecosystem changes, such as, the dynamics of infectious diseases. The non-native, marine gastropod Crepidula fornicata has become established across European coastlines over the last century, but there remains little insight into its disease carrying capacity and potential role as a source/sink of parasites. To address this knowledge gap, we surveyed limpets from two sites in South Wales, UK for signatures of disease/pathology using polymerase chain reaction-based methods (haemolymph) and histology (solid tissue). We encountered trematode-like parasites in ~1% individuals (5 out of 462). Three limpets displayed gross damage in the gonad, i.e. castration, and encysted metacercariae were found in the muscle of two other individuals. On the basis of 28S rDNA and internal transcribed spacer 2 genomic targets, we identified the gonad-infecting trematodes as members of the family Microphallidae ­ putative novel species related to the genus Longiductotrema. Earlier reports suggest that C. fornicata is not a host for trematode parasites in either its native or alien range but may act as a sink due to its filter feeding lifestyle. We provide clear evidence that C. fornicata is parasitized by at least one trematode species at two sites in Wales, UK, and likely act as a spillback or accidental host among native littorinids.

Heterorhabditis bacteriophora (Rhabditida: Heterorhabditidae), isolate HP88, induces reproductive and physiological alterations in Biomphalaria glabrata (Gastropoda: Planorbidae): an alternative for biological control of schistosomiasis.

Heterorhabditis bacteriophora is an entomopathogenic nematode (EPN) that is mutually associated with Photorhabdus luminescens, utilized globally for biological control of numerous organisms. Freshwater snails of the species Biomphalaria glabrata have been incriminated as the main intermediate hosts of Schistosoma mansoni in Brazil, but virtually nothing is known about the susceptibility of these gastropod to EPNs. Information in this respect is relevant for control of these intermediate hosts, and thus of the helminthiases they transmit. This paper for the first time reports the susceptibility of B. glabrata to infective juveniles of H. bacteriophora (isolate HP88) under laboratory conditions. For that purpose, six groups were formed: three Control groups (not exposed) and three Treated groups, in which the snails were exposed to 300 juveniles infecting the nematode over three weeks. The entire experiment was conducted in triplicate, using a total of 270 snails. Significant physiological alterations in B. glabrata were observed in response to the infection by H. bacteriophora HP88, characterized by decreased levels of hemolymphatic glucose as well as reduced contents of glycogen stored in the host's digestive gland. In parallel, the hemolymphatic activity of lactate dehydrogenase increased in the infected snails, indicating that the infection induces breakdown of carbohydrate homeostasis in B. glabrata. Additionally, all the reproductive parameters analyzed were reduced as a consequence of the infection. The results indicate the occurrence of the phenomenon of parasitic castration in the B. glabrata/H. bacteriophora HP88 interface, probably due to the depletion of galactogen in the parasitized organism. Although the infection did not cause lethality in the population of infected snails, H. bacteriophora HP88 compromised the reproductive performance of B. glabrata, suggesting its applicability in programs for biological control of this planorbid.

Parasitism-Induced Intersexuality in a Sexually Dimorphic Varunid Crab, Ptychognathus ishii (Decapoda: Varunidae).

Although many animals that perform sexual reproduction exhibit sexual dimorphism, individuals with intersex traits between the traits of males and females appear in some species, depending on environmental factors. Ptychognathus ishii, a varunid crab, exhibits distinctive sexual dimorphism in the morphology of its abdomen, chelipeds and setal tufts on the chelipeds. In this study, however, we report for the first time that intersex individuals with intermediate characters between those of males and females were occasionally found in wild populations. Morphological features of intersex individuals are described. Their taxonomic positions are identified based on DNA sequences of part of the mitochondrial cytochrome c oxidase I (COI) gene. It was shown that the intersexuality was induced by entoniscid parasites, because all intersex individuals were parasitized by entoniscid isopods, identified as Entionella sp. The apparent correlation between parasitism and morphological anomalies suggests that the parasitic isopods affect physiological conditions, leading to the feminization of male hosts.

Tetracotyle wayanadensis (Trematoda: Digenea) metacercaria - A potential parasitic castrator of the freshwater fish Aplocheilus lineatus (Valenciennes, 1846): A histopathological and temporal variation study in the South Western Ghats, India.

Parasitic castration is a phenomenon where the parasite blocks or reduces the host reproductive output. This study explored the impact of Tetracotyle wayanadensis infection on the reproductive performance of the freshwater fish Aplocheilus lineatus. Heavily infected A. lineatus showed slow and sluggish movements with reduced food intake and feeding activities. Histopathological observations of the ovary revealed severe necrosis and degeneration of previtellogenic and vitellogenic oocytes and follicular atresia. The oogonial nests were absent; chromatin nucleolar oocytes irregularly shaped and vacuolated. Perinucleolar oocytes have a vacuolated appearance with the hypertrophied highly columnar follicular cells invading them. Vitellogenic oocytes had a shrunken appearance with folded zona radiata; yolk globules broken down into smaller granules together with vacuolar degeneration of the ooplasm. The hypertrophied highly columnar follicular cells invaded the early vitellogenic oocytes to phagocytize the degenerating material. Zona radiata was found broken and liquefaction of yolk globules was evident with advancement in follicular atresia. Seasonal variation of parasitic infection has also been studied. Metacercaria infecting the vital organs showed seasonal variation with the highest values for prevalence and abundance during monsoon and mean intensity during winter. The lowest values for these descriptors were documented during the post-monsoon period.

Prevalence of a nematode castrator of the carrot weevil and impact on fecundity and survival.

Bradynema listronoti is a parasitic nematode described from infected specimens of the carrot weevil Listronotus oregonensis. Prevalence of infection by B. listronoti under field conditions was followed over a period of 16 years in an untreated carrot field. Susceptibility of different carrot weevil life stages was evaluated as well as the impact of infection on fecundity and mortality. Gene expression in infected and uninfected carrot weevils was also compared to evaluate the impact of the parasite on the host transcriptome. Prevalence of B. listronoti in carrot weevil populations was sustained over the years ranging from 20 to 63%. All the weevil stages exposed to B. listronoti inoculum were susceptible to infection, larvae being more vulnerable (59 ± 8% infected) compared with pupae (4 ± 3% infected) and adults (7 ± 3% infected). The fecundity of infected female weevils was greatly reduced (60-fold) due to an inhibition of the maturation of the reproductive system. Transcriptomic analyses revealed that this parasitic castration may have been triggered by the inhibition of reproductive hormone production. The B. listronoti-L. oregonensis interaction represents a case of parasitic castration with a unique potential for biological control of an important pest of carrots.

Effects of experimental Angiostrongylus cantonensis infection on the reproductive biology of Biomphalaria straminea and Biomphalaria tenagophila.

Eosinophilic meningoencephalitis is an endemic zoonosis in Southeast Asia and the Pacific Islands, but in recent years, new cases have been reported in various countries outside these regions, including Brazil, where it is considered an emerging disease. In this study, the effect of infection by the nematode Angiostrongylus cantonensis, one of the main etiologic agent of this disease, on the reproductive biology of the planorbid snails Biomphalaria straminea and B. tenagophila was investigated during the pre-patent period. Alterations in the reproductive biology of B. straminea and B. tenagophila were analyzed in laboratory-reared specimens infected by A. cantonensis during 21days; the number of eggs, number of egg masses, number of eggs/mass, number of eggs/snail, viable eggs/snail, survival and galactogen content in the albumen gland were measured. The results indicated the occurrence of initial compensation in reproductive effort in both snail species, but at different moments in the pre-patent period. More specifically, a reduction of 46.53% in the eggs/egg mass ratio in infected B. straminea was observed, a reflection of a 50% decline in the concentration of galactogen contained in the albumen gland. Changes in this parameter were also noted in B. tenagophila, but only at the end of the study period, with a reduction of 15.49%. Histological analyses indicate that changes observed can be explained by the tissue damages caused by the migration and development of the larvae. These results shed more light on the host-parasite relationship and indicate the importance of studying reproductive aspects for efforts to control infected snails. Considering that terrestrial snails can also transmit eosinophilic meningitis (in addition to aquatic mollusks), the data obtained expand knowledge of this host-parasite relationship and provide support for programs to control this zoonosis.

Subtle effect of Xenos vesparum (Xenidae, Strepsiptera) on the reproductive apparatus of its male host: Parasite or parasitoid?

Parasitic castration is an adaptive strategy where parasites usurp the hosts' reproductive physiology to complete their life cycle. The alterations in the host traits vary in their magnitude, from subtle changes in the host morpho-physiology and behaviour to the production of complex aberrant phenotypes, which often depend on the host gender. The strepsipteran macroparasite Xenos vesparum induces dramatic behavioural and physiological changes in its female host, the paper wasp Polistes dominula, while its effect on the male phenotype is largely unknown. In this study we investigated how a single X. vesparum parasite influences the functional morphology of P. dominula male reproductive apparatus. We performed morphometry and ultrastructure characterization of corpora allata, testes, seminal vesicles and accessory glands in parasitized and unparasitized males, and also in young and old males to control for the effect of age on the natural deterioration of these organs. Our results show that age significantly affects the development of male reproductive apparatus. A low parasite load - one parasite per host is the common prevalence in the field - has only a marginal impact on the reproductive morphology of P. dominula males, affecting quantitatively but not qualitatively the protein content of male accessory glands. Thus, in male P. dominula wasps, X. vesparum appears to behave as a true "parasite", in clear opposition to the role of "parasitoid" that it takes in female hosts where castration causes the reproductive death.

Molluscicidal potential of Heterorhabditis baujardi (Rhabditida: Heterorhabditidae), strain LPP7, on Lymnaea columella (Gastropoda: Pulmonata): An alternative for biological control of fasciolosis.

This study elucidated for the first time, under laboratory conditions, the susceptibility of Lymnaea columella to infective juveniles of Heterorhabditis baujardi LPP7. Exposure to the nematodes induced an average mortality rate of 66.66% in the population of L. columella, with the highest values attained from the second week after exposure onward. In addition, all the reproductive parameters analyzed (total number of eggs, number of egg masses, number of eggs laid/snail, embryo hatching rate and content of galactogen stored in the albumen gland) changed as a result of the infection. The results indicate the occurrence of the phenomenon of parasitic castration in L. columella infected by H. baujardi LPP7, probably through depletion of energy reserves such as galactogen, necessary to meet the intense metabolic demands of the nematode's larval stages. Finally, histopathological analysis demonstrated an intense process of cell disorganization, characterized by the occurrence of granulomatous inflammatory reactions in tissues of exposed snails, induced by the spoliative action of the bacteria/nematode. The results suggest the use of H. baujardi LPP7 as an alternative for biological control of the population of this intermediate host, and thus of the diseases in whose epidemiological chain it participates, especially fasciolosis, in line with the recommendations of the World Health Organization (WHO).

Morphological allometry and intersexuality in horsehair-worm-infected mantids, Hierodula formosana (Mantodea: Mantidae).

Parasitic castration is a strategy used by parasites to minimize damage to the host by consuming its reproductive system, which results in the morphological alteration of the host. We determined that the forewing shape and density of the antennal sensilla of field-collected adult male mantids (Hierodula formosana), infected by horsehair worms (Chordodes formosanus) was partially feminized (intersexuality), and both male and female mantids infected by horsehair worms exhibited allometric changes in their wings and walking legs. In addition, the testes of most infected male adults disappeared or reduced in size, whereas the number of ovarioles in infected female adults was unaffected. The infection mainly influenced the structures related to host reproduction and locomotion, suggesting unbalanced energy exploitation and the reduction of parasitic virulence. In addition, the intersexuality of infected male adults indicated that sexual differentiation in insects, which researchers have considered to be an autonomous process, was influenced by the infection. The similarity of the antennae of infected male adults with those of last-instar female nymphs suggested that parasitic juvenilization may cause such feminization, but the mechanism of parasitic influence on insect sex characteristics should be studied further.

Host age modulates parasite infectivity, virulence and reproduction.

Host age is one of the most striking differences among hosts within most populations, but there is very little data on how age-dependent effects impact ecological and evolutionary dynamics of both the host and the parasite. Here, we examined the influence of host age (juveniles, young and old adults) at parasite exposure on host susceptibility, fecundity and survival as well as parasite transmission, using two clones of the water flea Daphnia magna and two clones of its bacterial parasite Pasteuria ramosa. Younger D. magna were more susceptible to infection than older ones, regardless of host or parasite clone. Also, younger-infected D. magna became castrated faster than older hosts, but host and parasite clone effects contributed to this trait as well. Furthermore, the early-infected D. magna produced considerably more parasite transmission stages than late-infected ones, while host age at exposure did not affect virulence as it is defined in models (host mortality). When virulence is defined more broadly as the negative effects of infection on host fitness, by integrating the parasitic effects on host fecundity and mortality, then host age at exposure seems to slide along a negative relationship between host and parasite fitness. Thus, the virulence-transmission trade-off differs strongly among age classes, which in turn affects predictions of optimal virulence. Age-dependent effects on host susceptibility, virulence and parasite transmission could pose an important challenge for experimental and theoretical studies of infectious disease dynamics and disease ecology. Our results present a call for a more explicit stage-structured theory for disease, which will incorporate age-dependent epidemiological parameters.

Pistil Smut Infection Increases Ovary Production, Seed Yield Components, and Pseudosexual Reproductive Allocation in Buffalograss.

Sex expression of dioecious buffalograss [Bouteloua dactyloides Columbus (syn. Buchloë dactyloides (Nutt.) Engelm.)] is known to be environmentally stable with approximate 1:1, male to female, sex ratios. Here we show that infection by the pistil smut fungus [Salmacisia buchloëana Huff & Chandra (syn. Tilletia buchloëana Kellerman and Swingle)] shifts sex ratios of buffalograss to be nearly 100% phenotypically hermaphroditic. In addition, pistil smut infection decreased vegetative reproductive allocation, increased most seed yield components, and increased pseudosexual reproductive allocation in both sex forms compared to uninfected clones. In female sex forms, pistil smut infection resulted in a 26 fold increase in ovary production and a 35 fold increase in potential harvest index. In male sex forms, pistil smut infection resulted in 2.37 fold increase in floret number and over 95% of these florets contained a well-developed pistil. Although all ovaries of infected plants are filled with fungal teliospores and hence reproductively sterile, an average male-female pair of infected plants exhibited an 87 fold increase in potential harvest index compared to their uninfected clones. Acquiring an ability to mimic the effects of pistil smut infection would enhance our understanding of the flowering process in grasses and our efforts to increase seed yield of buffalograss and perhaps other grasses.

Mercury concentration in the freshwater bonefish Cyphocharax gilbert (Curimatidae) and its parasite the crustacean Riggia paranensis (Cymothoidae)

Fish parasites can accumulate heavy metals reaching higher concentrations than the host and may affect the host's bioaccumulation. The present study compared total mercury concentration in the liver and muscle of Cyphocharax gilbert and in the parasite Riggia paranensis sampled in the middle Itabapoana River, Brazil, also considering the reproductive stages of both fish and parasite. Mean concentrations of mercury in muscle of fish varied from 20.8 ng.g-1 in mature females to 38.1 ng.g-1 in post-spawning females. The mean concentrations in fish liver varied from 60.9 ng/g in post-spawning females to 110.4 ng.g-1 in infested males. The mean concentration of mercury in parasites varied from 26.2 ng.g-1 in specimens carrying early embryo to 39.5 ng.g-1 in specimens with eggs. Positive and significant associations (P<0.05) were found between the total mercury concentrations in parasites and muscle of host (both females and males), and between concentrations in parasites and in the liver of male hosts. These results suggest that R. paranensis can be used to indicate mercury levels in edible parts of C. gilbert.

Parasitos de peixes podem acumular metais pesados em concentrações acima dos hospedeiros, podendo afetar a bioacumulação nestes. O presente estudo comparou a concentração total do mercúrio no fígado e no músculo de Cyphocharax gilbert e no parasito Riggia paranensis, coletados no trecho médio do rio Itabapoana, Brasil, considerando o estágio reprodutivo tanto dos peixes como dos parasitos. A concentração média do mercúrio nos músculos dos peixes variou entre 20,8 ng.g-1 nas fêmeas maturas e 38,1 ng.g-1 nas fêmeas desovadas. As concentrações médias nos fígados dos peixes variaram entre 60,9ng/g em fêmeas desovadas e 110,4 ng.g-1 em machos parasitados. Entre os parasitos, as concentrações médias de mercúrio variaram entre 26,2 ng.g-1 em espécimes com embriões iniciando o desenvolvimento to 39,5 ng.g-1 em espécimes com ovos . Associações positivas e significativas (P<0.05) foram encontradas entre as concentrações totais de mercúrio em parasitos e músculos dos hospedeiros (tanto em macho como em fêmeas) e entre as concentrações nos parasitos e os fígados dos hospedeiros machos. Estes resultados sugerem que o parasito R. paranensis pode ser utilizado como bioindicador dos níveis de mercúrio nas partes comestíveis de C. gilbert.

Infestation pattern and parasitic castration of the crustacean Riggia paranensis (Crustacea: Cymothoidea) on the fresh water fish Cyphocharax gilbert (Teleostei: Curimatidae)

Cyphocharax gilbert infested by Riggia paranensis shows parasitic castration. The prevalence of parasitism in C. gilbert varied among different environments, being higher in the middle rio Itabapoana. Fish were collected monthly using two cast nets (thrown 30 times during the day) and gillnets kept in the river during 12 hour, from sunset to sunrise, between September 1997 and August 2000. Infestation pattern was investigated on 1358 specimens. Most of them were infested (57.9 percent), with one or two parasites; the majority (62.9 percent) was collected during the rainy season (spring-summer). The parasite did not show preference for sex or size of hosts. A total of 91.5 percent of the 511 examined parasites had a body size that represented 10.1 percent to 20 percent of host standard length. The reproductive condition of 311 specimens of R. paranensis was analyzed checking the presence of oocytes in the ovarian and eggs or embryos in the marsupium. Nearly 73 percent of them were at reproductive phase, and had a body size that represented 5.1 percent to 20 percent of host standard length. The size of the immature parasites varied from 0.1 percent to 5 percent of the host size. The results suggest that R. paranensis may adopt a fast growth rate strategy and increase the investment in reproduction when they occupy most of the host's pericardial space.

Cyphocharax gilbert infestado por Riggia paranensis apresenta castração parasitária. A prevalência do parasito varia entre diferentes ambientes, sendo maior no trecho médio do rio Itabapoana. Os peixes foram coletados mensalmente neste rio usando duas tarrafas (lançadas por 30 vezes durante o dia) e redes de espera por 12 horas, do crepúsculo ao amanhecer, entre setembro de 1997 e agosto de 2000. O padrão de infestação foi investigado em 1358 peixes. Peixes infestados apresentaram um ou dois parasitos, representando 57,9 por cento da população e a maioria deles (62,9 por cento) foi coletada no período de maior precipitação (primavera-verão). O parasito não apresentou preferência por sexo ou tamanho do hospedeiro. No total, 91,5 por cento de 511 parasitos examinados apresentaram um tamanho corpóreo que representou de 10,1 por cento a 20 por cento do comprimento padrão do hospedeiro. A condição reprodutiva em 311 espécimes R. paranensis foi analisada através da presença de oócitos no ovário e ovos ou embriões no marsúpio, sendo que cerca de 73 por cento estavam em fase de reprodução, apresentando um tamanho corporal que variou entre 5,1 por cento a 20 por cento do comprimento padrão do hospedeiro. O tamanho dos parasitos imaturos variou de 0,1 por cento a 5 por cento do tamanho do hospedeiro. Os resultados sugerem que R. paranensis pode adotar uma estratégia de rápido crescimento e deve aumentar o investimento em reprodução quando ocupa grande parte do espaço disponível na cavidade pericardial do hospedeiro.

Effect of starvation on parasite-induced mortality in a freshwater snail (Potamopyrgus antipodarum).

The level of host exploitation is expected, under theory, to be selected to maximise (subject to constraints) the lifetime reproductive success of the parasite. Here we studied the effect of two castrating trematode species on their intermediate snail host, Potamopyrgus antipodarum. One of the trematode species, Microphallus sp., encysts in the snail host and the encysted larvae "hatch" following ingestion of infected snails by birds. The other species, Notocotylus gippyensis, by contrast, releases swimming larvae; ingestion of the snail host is not required for, and does not aid, transmission to the final host. We isolated field-collected snails for 3 months in the laboratory, and followed the survival of infected and uninfected snails under two conditions: not fed and fed ad libitum. Mortality of the infected hosts was higher than mortality of the uninfected ones, but the response to starvation treatment was parasite species specific. N. gippyensis induced significantly higher mortality in starved snails than did Microphallus. Based on these results, we suggest that host exploitation by different species of trematodes may depend on the type of transmission. Encysting in the snail host may select for a reduced rate of host exploitation so as to increase the probability of transmission to the final host.

Parasite-induced parthenogenesis in a freshwater snail: stable, persistent patterns of parasitism.

The role of parasites in the evolution of host reproductive modes has gained renewed interest in evolutionary ecology. It was previously argued that obligate parthenogenesis (all-female reproduction) arose in a freshwater snail, Campeloma decisum, as a consequence of severe sperm limitation caused by an unencysted trematode, Leucochloridiomorpha constantiae. In the present study, certain conditions are examined for parasitic castration to account for the maintenance of parthenogenesis: the spatial patterns of the prevalence and intensity of infection on a broad geographical scale and its relationship to host genotype; the recovery from infection after isolation from sources of infection; age-related patterns of infections; and the effects of L. constantiae on snail fecundity.In contrast to the common pattern of the aggregated distribution of parasites within host populations, many snail populations with high prevalence and intensity of infection have non-aggregated parasite distributions. Clonal genotype of the host explained little of the variation in intensity and prevalence of infection by the parasite. Female snails maintained similar prevalence and intensity of infection after isolation, and individuals accumulated parasites throughout their lifespan, both of which suggest there is no effective immune response to infection by L. constantiae. Snail fecundity is not significantly influenced by the intensity of infection. These results suggest that L. constantiae may have represented a strong selective force against males during the initial introduction of this parasite into sexual snail populations because of the persistent nature of infection.

Parasitic castration: host species preferences, size-selectivity and spatial heterogeneity.

This study investigates host-parasite population dynamics in a marine intertidal community of three barnacle host species (Balanus glandula, Chthamalus fissus andC. dalli). Our paper addresses the following questions: (1) Does prevalence (percentage parasitism) differ among the three host species? (2) What are the spatial and temporal population dynamics within the community? and (3) Does the parasite exhibit size-selective behaviour in any of the three host species? Significant differences in prevalence were found among the three host species; the parasitic castrator (Hemioniscus balani) most heavily infected the least abundant host. Parasitism occurred throughout the year and also showed significant spatial variation.H. balani showed size-selective parasitism inC. fissus, but not inC. dalli. Consequently, the population effects of parasitic castration inC. fissus depend both upon the host population size structure and the intensity of the parasite's size-selectivity.