To understand the relationship between mode of transmission and virulence, we investigated 2 microsporidian parasites in Daphnia magna laboratory populations. Pleistophora intestinalis is only transmitted horizontally, while Tuzetia sp. is transmitted vertically with high efficiency from mothers to parthenogenetic male and female offspring. We were not able to transmit Tuzetia horizontally in the laboratory. Tuzetia reduces host life-time reproductive success and host survival to a much greater extent than does P. intestinalis. Tuzetia-infected hosts were rapidly outcompeted by uninfected hosts. We suspect that Tuzetia infections may persist in natural populations by an as yet undiscovered horizontal transmission. It is possible that an alternate host species may be involved. We present a mathematical model to analyse the conditions for the persistence of a parasite with perfect vertical and an additional degree of horizontal transmission. We show that horizontal and vertical transmission contribute additively to a parasite's ability to invade and persist. Since the fitness contribution of horizontal transmission increases with population size, only very low transmission probabilities per host-to-host contact are necessary for the persistence of parasites occurring in large populations such as those commonly found for Daphnia. The detection of such low rates of horizontal transmission appears unlikely under laboratory conditions since the necessary number of host-to-host (or host-to-spore) contacts is not feasible. We review mechanisms that can maintain vertically transmitted parasites in nature.

Vertically transmitted parasites are transmitted from generation to generation of hosts usually via the host's gametes. Owing to gamete size dimorphism, the major transmission route is transovarial and selection (on the parasite) favours strategies which increase the relative frequency of the transmitting (female) host sex. These strategies impose unusual selection pressures on the host, and coevolution between hosts and vertically transmitted parasites has been implicated in speciation, in the evolution of symbiosis, and in the evolution of novel systems of host reproduction and sex determination. We review the evolutionary implications of vertically transmitted parasites in arthropods before focusing on strategies of transmission of a parasitic sex ratio distorter in Gammarus duebeni. The efficiency of parasite transmission to new hosts is a key factor underlying the relationship between vertically transmitted parasites and their hosts. Vertically transmitted parasites must overcome 2 bottlenecks in order to ensure successful infection of future host generations: first, transmission from adult to gamete; and secondly, transmission to the germ-line of the infected host. We investigate these 2 processes with regard to transovarial transmission by a microsporidian parasite in Gammarus duebeni. Parasite transmission from adult to eggs is highly efficient, with 96% of eggs of infected mothers inheriting the infection, whereas transmission to germ-line within infected embryos is relatively inefficient (72%). We measure parasite distribution between cells of developing embryos, and use these distributions to infer possible mechanisms of parasite transmission to germ-line. Parasite distribution within the embryo is dependent on host cell lineage, and is not consistent with unbiased segregation between daughter cells. These results indicate that parasites segregate together at host cell division, and may reflect a strategy of differential segregation to the host germ-line. We consider alternative parasite strategies at the cell-level in terms of their evolutionary implications.

This is an investigation of the impact of salinity on transovarial transmission and burden of a microsporidian sex ratio distorter in the inter-tidal crustacean Gammarus duebeni. Exposure of parasitized mothers to increased salinity during the gonotrophic cycle caused an increase in parasite burden in the follicle cells and a decrease in burden in the oocytes. It appears that salinity impedes parasite transmission from the follicle cells to the oocytes during host oogenesis. A lower proportion of the young were infected in broods from elevated salinity and, in infected offspring, parasite burden was lower than in control embryos. Parasite replication occurred during embryogenesis. However, the pattern of parasite growth did not differ between salinities, indicating that differences in parasite burden could be attributed to a reduction in the initial parasite burden transmitted to the gamete, rather than to a reduction in parasite replication during host embryogenesis. We discuss our findings with respect to parasite/host dynamics and the ecology of the host.

The cellular distribution of a vertically transmitted, feminizing microsporidian was followed in its host Gammarus duebeni. In adult females the parasite was restricted to gonadal tissue, in particular primary and secondary follicle cells. Spores were diplokaryotic with a thin spore wall and a short polar filament, characteristics typical of ‘early’ spores involved in autoinfection. The diplokaryotic life-cycle, absence of spore groupings and of a pansporoblast membrane typify the genus Nosema. However, the unusual globular polaroplast of the spore and restriction of this stage to host ovarian tissue have not previously been described in Nosema. Sporogony occurred only in follicle cells adjacent to developing oocytes and was in synchrony with the process of vitellogenesis. Oocytes were infected after formation of intracellular connections with follicle cells but harboured only vegetative stages of the parasite. Parasites were associated with the perinuclear cytoplasm and, in developing embryos, segregated to daughter cells along the axis of the spindle. In juvenile animals there was no evidence of pathology linked with feminization and the parasite was found at low density in cells under the cuticle. The parasite is highly adapted to transovarial transmission with an efficient mechanism of oocyte infection and no evidence of pathology.

This is a report of the prevalence, transmission and intensity of infection of a microsporidian sex ratio distorter in natural populations of its crustacean host Gammarus duebeni. Prevalence in the adult host population reflects differences in the intensity of infection in transovarially infected embryos and in adult gonadal tissue. The efficiency of transovarial parasite transmission to young also differs between populations, but this alone is insufficient to explain observed patterns of prevalence. Infection intensity may be important in determining future infection of target tissue in the adult and subsequent transmission to future host generations. We consider patterns of parasite infection in terms of selection on transmission and virulence.

Laboratory-reared tsetse, Glossina pallidipes were inoculated with the tsetse virus by microinjection into the haemocoel, and feeding through micropipettes. The inoculated tsetse were reared on rabbits for 45 days, and feeding, flight and mating activities recorded. The tsetse were dissected and the conditions of the salivary glands and gonads noted. In males, tsetse were examined under a compound microscope and the presence of spermatozoa noted. F1 pupae were allowed to emerge, dissected, and the salivary glands examined for hypertrophy, and the gonads for sterility.

There was no reduction in the activity of inoculated tsetse. Infection level was 23.5% in the treated adults. All infected ♂♂ were sterile while ♀♀ were fertile. There was no significant difference in the maternal age at larviposition, F1 pupal weight, or incubation period of F1 pupae, between treated and untreated tsetse.

A high proportion of F1 adults (65%) were infected, with category four hypertrophied salivary glands. All males with enlarged glands were sterile. The evidence obtained shows that the tsetse virus may be used in biological control of G. pallidipes.