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Ticks are obligate haematophagous arthropods that represent a major economic drain upon the world's livestock as well being a significant medical and veterinary risk through the transmission of tick-borne pathogens such as Borrelia burgdorferi, the causative agent of Lyme disease. The tick-host relationship is a function of both ecological and physiological factors. Successful feeding requires the effective acquisition and digestion of a bloodmeal by the tick. Acquisition relies upon the ability of the tick to counteract host immune responses induced by the extended feeding periods of ixodid ticks (up to 2 weeks). The host response to tick infestation and the consequent countermeasures employed by the tick, constitute the tick-host interface. The immune response of hosts to Ixodes ricinus infestations was examined through antigenic profiling. The antigens exposed to the host were shown to vary throughout the feeding period and differed between the different development stages of I. ricinus. It was also shown that different host species infested with I. ricinus recognised different antigens. This was true of both natural and non-natural hosts, and even closely related species. Anti-complement activity was investigated in the salivary glands of Ixodes ticks. This activity was shown to inhibit some host species but not others. The pattern of inhibitory activity varied between the tick species tested in a way that was consistent with known tick host-preferences. The mechanisms of anti-complement activity in I. ricinus salivary glands were explored. The alternative but not the classical pathway of complement was inhibited. Activity was present in unfed ticks and throughout the feeding period. Three targets of the complement system were identified as being modulated by the tick. Digestion of the bloodmeal was explored and a haemolytic activity was associated with the salivary glands of I. ricinus ticks. The activity was demonstrated to be Mg²⁺- dependent. In addition, a subtractive cDNA library enriched for saliva-associated transcripts was successfully produced. Random sampling identified putative differentially expressed genes. The results of this thesis illustrate the complexity of tick-host interactions at the molecular level. It is apparent that the research described poses many more questions than answers.

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Immune responses are presumed to contribute to host fitness, either by fighting off infections or via immunopathology. Research in this thesis sought to relate the magnitude of a putative immune response to infection and host and parasite fitness. The experiments and field studies presented here all focus on the interactions between the freshwater crustacean, Daphnia magna and its sterilizing bacterial endoparasite, Pasteuria ramosa, using the number of circulating haemocytes as a measure of host immune activity. I found substantial genetic variation in Daphnia’s cellular response to P. ramosa, and that Daphnia genotypes that mount the strongest cellular responses are the most likely to get infected and suffer sterilization. Thus, a strong cellular response is associated with low, as opposed to high host fitness potential. There were also some host genotypes that mounted a weaker cellular response and did not go on to suffer infection, and some that lacked a cellular response and also never suffered infection with P. ramosa. These findings led to a heuristic two-stage model for infection, where the parasite has to (1) pass from the host gut to haemolymph and then (2) successfully overcome haemolymph-based immune effectors to reproduce and achieve fitness. I also demonstrate that both the magnitude of host cellular response and likelihood of infection increases with initial parasite dose in susceptible host genotypes, and that host cellular response is associated with likely infection under both host and parasite genetic variation. Parasitised Daphnia also have substantially more circulating haemocytes than their healthy counterparts in both the laboratory and in the wild, where there is substantial genetic and environmental variation. This is one of the very few examples of how an immune response designates low host and high parasite fitness potential in a wild system. Finally, using a mixture of field study and common garden experiment, I demonstrate evolution in parasite infection traits over the course of an epidemic in a wild population, and that this evolution is associated with a decline in host abundance.

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Bibliography: leaves 113-144. Investigates the functions of the maternal, protein secretions of the endoparasitoid wasps Venturia canescens Gravenhorst regarding their role in providing protection against the host's immune system. Also investigates mucinous protein secretions on the egg surface and coagulation reactions and their role in protecting eggs against host cellular attacks.

Most, if not all, animals will at some stage in their lives encounter parasites. Some of the most widespread and abundant parasites belong to the Class Trematoda. Trematodes often have a substantial negative impact on individual intermediate host ecology. In this thesis, I investigate ecological and evolutionary consequences of the interaction between the microphallid trematode, Maritrema novaezealandensis, and its intermediate snail and crustacean hosts. Parasites often show a heterogeneous spatial distribution pattern in natural animal host populations. In this study, factors determining the spatial distribution of larval trematodes in Zeacumantus subcarinatus were investigated at two spatial scales (within and among bays). The distribution of shorebird definitive hosts explained a significant amount of the variation in the distribution of trematodes among bays. However, within a bay, other factors override the effect of bird distribution. The influence of larval trematodes on reproduction, survival and population density of Z. subcarinatus was investigated using laboratory and field studies. In the laboratory, it was found that larval trematodes induce castration and mortality of Z. subcarinatus. The field study revealed that the local prevalence of trematodes had a significant negative effect on population density of Z. subcarinatus. Through castration, trematodes act as strong selective agents on snail host life history. The effect of trematodes on life history characteristics (reproductive effort, juvenile growth, size at maturity and susceptibility to trematode infections) were investigated among natural populations of Z subcarinatus. Reproductive effort was not higher for uninfected females from populations where the risk of becoming infected was high. However, offspring from those populations were significantly larger, and laboratory-reared juveniles grew significantly faster than conspecifics from other populations. In addition, size at maturity was negatively correlated with trematode prevalence across snail populations. Z. subcarinatus thus adapts to a high local risk of trematode infection by reaching maturity early, thereby increasing the chance of reproducing. The influence of M. novaezealandensis on the survial of the amphipod host, Paracalliope novizealandiae was examined using experimental infections and field observations. The experimental infections demonstrated that parasite-induced mortality was intensity-dependent. The number of M. novaezealandensis per amphipod was too low to significantly induce host mortality in the field. However, the transmission strategy of this parasite allows it to affect host populations during weather conditions ideal for trematode transmission. Trematode strategies in the second intermediate host are important to the understanding of host-parasite co-evolution and the evolution of parasite life cycles. In this study, potential density-dependent effects at the metacercarial stage on size and fecundity of in vitro adult M. novaezealandensis was examined in both experimentally infected P. novizealandiae and naturally infected Macrophthalmus hirtipes. For this purpose, a method to excyst and cultivate M. novaezealandensis metacercariae to an egg producing stage, was developed. Naturally infected M. hirtipes also harboured larval stages of three other helminths. Crowding effects in the two crustacean hosts were expressed as a decreased volume and smaller egg production of in vitro adult M. novaezealandensis. In addition, interspecific interactions among parasite species were observed in crab hosts. The work in this thesis provides evidence that M. novaezealandensis significantly and negatively affect intermediate host ecology. The heterogeneous distribution of trematodes causes differential effects among host populations with subsequent effects on the life history of snail hosts. In addition, this study demonstrates that parasites interact within their second intermediate host with possible implications for the way parasites exploit their hosts.