The behaviour of cercariae was recorded on video at 10, 20 and 30 °C and a combination of population and individual behavioural patterns studied. Initially cercarial populations were extremely active, individuals alternating between swimming and sinking behaviour. As they aged the percentage sinking and resting increased resulting in the establishment of a vertical distribution in the water column. An increased environmental temperature reduced the active life-span of the parasites. Over time cercarial swimming rates declined. Swimming rates increased with environmental temperature and as a result of a shadow stimulus. As cercariae aged, the duration of swimming events decreased. Post-stimulation swimming periods were significantly longer than pre-stimulation periods. Pre- and post-stimulation swimming distances also declined with age. Post-stimulation swimming distances were significantly longer than pre-stimulation distances. The sinking rates of cercariae decreased with age while sinking periods and distances increased. Data for proportional time spent swimming and the ratio of sinking to swimming events indicated that as cercariae aged, less time was spent swimming. The frequency of swimming/sinking events declined with age. The implications of this behaviour for dispersal and transmission are discussed.

This review considers three case studies based on macroparasites of anurans: (a) natural infections in the permanently-aquatic Xenopus laevis which represent the worm burdens acquired, and the implications for pathology, when hosts are exposed to continuous, year-round, transmission; (b) the desert toad, Scaphiopus couchii, which experiences invasion very briefly each year and provides a simplified system involving only a single significant infection (Pseudodiplorchis americanus); (c) the mesic Bufo bufo which has been the subject of experimental laboratory studies designed to measure the effects of Rhabdias bufonis infection on host growth, physical performance and survival. Experimental manipulation of both Scaphiopus and Bufo provide quantitative data on disease effects of macroparasites, including precise measurements of parasite-induced host mortality. Field data for Xenopus and Scaphiopus show that, despite high initial worm burdens from efficient transmission, infection levels at parasite maturity are modulated below those leading to significant disease. Experimental data for Scaphiopus and Bufo have documented the time-course and magnitude of this decline in intensities, and there is circumstantial evidence for Scaphiopus that this regulation is host-mediated. Immunological studies on Xenopus show that disease effects of the pathogenic Pseudocapillaroides xenopodis are exacerbated in thymectomised hosts and reversed by implantation of thymuses from MHC-compatible donors. Thus, whilst factorial experiments can demonstrate the potential of helminths to cause significant disease and mortality in anuran host-macroparasite interactions, powerful post-invasion regulation of worm burdens appears to exert a strong control of parasite-induced disease in natural host populations.

A haemoprotein released in vitro by adult Fasciola hepatica was purified by gel filtration chromatography on Sephacryl S-200 and ion-exchange chromatography on DEAE-Sepharose. The molecule, with an apparent molecular weight of > 200 kDa, contains a haem group and has absorption spectra characteristics similar to haemoglobins. N-terminal amino acid sequence analysis revealed no similarity between the F. hepatica haemoglobin and other vertebrate or invertebrate haemoglobins. Antibodies to the haemoglobin molecule can be detected in the sera of F. hepatica-infected bovines as early as 1 week after infection.

Much of what is currently known of the host-parasite interaction between mice and the parasitic nematode Trichuris muris has come from experiments using a single parasite isolate (E/N). This isolate has been compared with 2 others which, on morphological criteria, belong to the same species. In 3 inbred strains of mouse that show distinct, genetically determined response phenotypes, there was a consistent pattern in terms of parasite survival time regardless of host strain, E/K worms being expelled early, E/N expelled later and S worms very late or not at all. High-responder CBA mice expelled E/K and E/N worms earlier than low-responder C57 B1/10 mice. B10.BR mice were permissive to S isolate infection, mounted a very late response to E/N worms but expelled E/K worms effectively by day 25. The differential response of mice to these isolates provides an experimental system for identifying the basis of variation in this host-parasite relationship.

Using a simple infection model the infection interaction of Steinernema feltiae (Site 76 strain) Filipjev and Galleria mellonella L. was investigated under laboratory conditions. The model analyses showed that the infection interaction was consistent with a proportion of the S. feltiae infective stages being infectious and, alone amongst the infective-stage population, being capable of infection. The investigation also showed that the infection behaviour of the infectious stages could be reduced to a slightly overdispersed probability of transmission. The importance of these findings with respect to the quantitative description and explanation of infection are discussed.

African trypanosomes are parasitic protozoa causing sleeping sickness in humans and related diseases in domestic animals against which no entirely satisfactory forms of chemotherapy are yet available. It was previously shown that related species of trypanosomes, as well as procyclic (insect) forms of Trypanosoma brucei are extremely sensitive to the action of phenothiazine neuroleptic drugs in vitro. In this work, we have carried out a more detailed investigation of the effects of thioridazine, one of the most potent neuroleptic phenothiazine drugs known, on the morphology of the infective bloodstream forms of T. brucei, with particular reference to the parasite's prominent pellicular membrane complex. Our data show that this drug induces rapid changes in cell shape that appear to involve some reorganization of the microtubular membrane skeleton, but does not affect the structural integrity of the microtubular complex. Another early consequence of drug action involved damage to nuclear and cytoplasmic membranes and the appearance of tubular arrays of coated membrane within the flagellar pocket. It was also revealed that the drug induces a rapid release of the variant-specific glycoprotein (VSG) which makes up the surface coat protecting bloodstream forms of the parasite against the host immune system. Our evidence suggests that this release of VSG involves cleavage of the protein's glycosyl-phosphatidylinositol (GPI) membrane anchor by endogenous GPI-specific phospholipase C, probably as a consequence of minor damage to the parasite plasma membrane induced by the drug.

To test the hypothesis that the relatively high developmental temperature threshold of the parasitic nematode Elaphostrongylus rangiferi in the intermediate snail host is an adaptation to minimize larval mortality during winter, an experiment was set up in which snails of the species Arianta arbustorum were experimentally infected with the parasite. The snails were divided into 3 groups known to contain 1st, 2nd or 3rd-stage larvae, and incubated at 3 °C for an experimental period of 18 weeks. First-stage larvae showed a significantly higher survival rate within snails than 2nd or 3rd-stage larvae. We also found that snails carrying 1st-stage larvae survived better than snails with other larval stages. It is concluded that if the nematode has started development before the hibernation, this has a real and significant effect on the risk of dying. The high developmental threshold is therefore likely to be an adaptation to reduce the chance of hibernating as developing larvae during long periods of low temperatures.

Migration by nematode larvae through the tissues of their mammalian hosts can cause considerable pathology, and yet the evolutionary factors responsible for this migratory behaviour are poorly understood. The behaviour is particularly paradoxical in genera such as Ascaris and Strongylus in which larvae undergo extensive migrations which begin and end in the same location. The orthodox explanation for this apparently pointless behaviour is that a tissue phase is a developmental requirement following the evolutionary loss of skin penetration or intermediate hosts. Yet tissue migration is not always necessary for development, and navigation and survival in an array of different habitats must require costly biochemical and morphological adaptations. Migrating larvae also risk becoming lost or killed by the host. Natural selection should therefore remove such behaviour unless there are compensating benefits. Here we propose that migration is a selectively advantageous life-history strategy. We show that taxa exploiting tissue habitats during development are, on average, bigger than their closest relatives that develop wholly in the gastrointestinal tract. Time to reproduction is the same, indicating that worms with a tissue phase during development grow faster. This previously unsuspected association between juvenile habitat and size is independent of any effects of adult habitat, life-cycle, or host size, generation time or diet. Because fecundity is intimately linked with size in nematodes, this provides an explanation for the maintenance of tissue migration by natural selection, analogous to the pre-spawning migrations of salmon.

Third-stage (dauer) juveniles of the entomopathogenic nematode Steinernema feltiae from 3 laboratory populations were tested for infectivity to larvae of the sciarid fly Lycoriellia solani. Bioassay experiments were done to assess the effects of nematode application density, exposure time and host density on the proportion of individuals that established infection. When dauer stages were presented to a host singly, establishment rates of 28, 26 and 18% were obtained for the 3 populations, but the presence of conspecifics consistently increased the rate of establishment to 42, 33 and 21% respectively, at 20 nematodes per host. A modified binomial model was developed to show that nematode establishment was not related to application density in a linear fashion, as initial infection facilitated secondary colonization. The data suggest that 3 subpopulations of nematodes may be distinguished by their infection behaviour: a first group of individuals with the behavioural propensity to initiate infection in unparasitized insects, a second that only invaded infected hosts, and a third group of non-invaders that failed to establish in L. solani.

Naturally infected Biomphalaria glabrata snails were collected at two sites near Belo Horizonte, Brazil, and Schistosoma mansoni cercariae isolated from single snails were used to infect individual mice. Genetic comparison of single worm DNA was accomplished by hybridization of Southern blots to a polymorphic repetitive DNA element. Genetic profiles of parasite individuals revealed a diverse array of parasite genotypes in naturally infected intermediate hosts. The observed distribution of schistosome genotypes among intermediate hosts indicates that over half of the infected snails harbour multiple miracidia. Snails were more likely to carry multiple infections than expected by chance. This degree of overdispersion combined with high levels of genetic variability facilitates multi-genotype transmission and helps maintain parasite genetic diversity.

The natural history of fig-pollinating wasps and their associated species-specific nematodes allows the measurement of many parameters which are relevant to testing hypotheses concerning host-parasite ecology and evolution. Within fig wasps species, it is possible to estimate lifetime reproductive success of foundress wasps as a function of presence or absence of nematode parasitism (virulence). Across species, there is a wide range of host population structures which, in turn, results in a range of opportunities for either horizontal or vertical nematode transmission. Therefore, estimates of virulence can be related to opportunities for transmission across a group of closely related hosts and their parasites. Further, the dynamics of the nematode infections over ecological and short-term evolutionary timescales can be monitored, giving added insight into the interpretation of the virulence estimates. Moreover, several scales of longer term evolutionary relationships are either known directly from fossil evidence or can be inferred from molecular data, providing deeper temporal context for the observed patterns. This combination of attributes permits detailed testing of hypotheses concerning the factors that potentially influence the evolution of virulence in host-parasite systems, and further, population and simulation models of the system that incorporate the parameter estimates can clarify the interpretation of how those factors act. There is little evidence suggesting that intermediate and long-term evolutionary relationships explain current levels of virulence. That is, it appears that virulence can change rapidly relative to speciation events, and that the nematodes do not tend to become ‘benign over time’. Instead, it appears that host population structure can influence the evolution of parasite virulence by affecting the relative opportunities for horizontal to vertical transmission, which, in turn, influences the relative costs and benefits of virulence to the nematodes. At one level, increased opportunities for horizontal transmission decouple the reproductive interests of the individual nematodes from those of the individual hosts that they are directly parasitizing, thereby reducing the cost of virulence to individual nematodes. At another level, increased opportunities of horizontal transmission also increases the relative frequency of hosts infected by multiple strains of nematodes. This promotes the evolution of more virulent forms by increasing the relative importance of within-host competition among nematode strains, thereby favouring strains that ‘eat more host sooner’. An interesting property of the fig-nematode systems is that the proportion of infected hosts does not change dramatically through time. This finding implies that there can be considerable negative effects on survival of infected hosts in addition to the previously documented reductions in fecundity of infected foundresses, because the latter are insufficient to account for the observed stability of wasp infection rates.

As a major morphological feature in establishing the form of the nervous system, it is recognized that neurons are initially overproduced, then naturally occurring cell death reduces the neuron number to the functional requirement. However, the mechanisms controlling the selective elimination of certain neurons during a general phase of cell death are not fully understood. One event that seems to be pivotal is the establishment of neural connections, the degree of which may be influential regarding the fate of specific neurons. However, little quantitative evidence is available to either support or refute this theory. In this current study, a Stereological measurement of gap junction per neuron was carried out within the invertebrate model system of the tapeworm metacestode Mesocestoides corti, which has previously been shown to overproduce neurons during the asexual reproduction stage of its life-cycle. Novel Stereological estimation methods with ‘ vertical sections’ indicated that prior to asexual division the cerebral ganglion possessed approximately 268 neurons, each with a gap junction surface area of 250 μm2. As division progressed, the neuron number increased to approximately 700, while the total surface area of gap junction remained statistically unchanged. As a result the surface area of gap junction per neuron decreased to 106 μm2, less than half that in the undividing stage. These results provide the first non-biased quantitative data regarding changes in the mean surface area of gap junction per neuron in a developing cerebral ganglion.

The status of bluegill (Lepomis macrochirus) and green sunfish (Lepomis cyanellus) as homologous hosts for the acanthocephalan Neoechinorhynchus cylindratus was experimentally determined. It was found that the adult parasite did not establish in bluegills, but that these fish could serve as paratenic host. In contrast, complete growth and development to the adult stage occurred in the green sunfish. When green sunfish were intubated with 10 cystacanths/fish, the parasite exhibited a clear preference for the anterior half of the intestine; when 50 cystacanths/fish were intubated, the parasites showed a preference for the posterior half of the intestine. With repeated exposure of cystacanths, the parasites were distributed throughout the intestine. The extent of histopathology induced by N. cylindratus was related to the numbers of parasites present. In light infections (10 cystacanths), the parasite penetrated deeply into the intestinal wall and connective tissue developed around the proboscis. In infections with 50 cystacanths, the proboscis penetration was shallow and little if any connective tissue accumulated. There was also an indication that in crowded areas, the parasites appeared to change their sites of attachment frequently. In both heavy and repeated infections, the parasites evoked significant goblet cell hyperplasia and substantial quantities of mucus covered the intestinal wall. It is suggested that the sticky covering and the presumed presence of antibodies in the mucus combined to create a protective barrier thereby reducing the numbers of parasites that could attach and become established.

Trematode cercariae typically become unable to successfully infect a host many hours before they die. We examined the hypothesis that both time to 50% mortality and time to loss of infective capacity are controlled to the same degree by rates of energy expenditure, by determining the relative effects of temperature on both parameters. Infective capacity was assessed by exposing Echinostoma trivolvis cercariae of different ages to a suitable second intermediate host (the gastropod Biomphalaria glabrata) and counting 1–2 days later the number of metacercarial cysts formed. Temperature had a remarkably similar effect on time to 50% mortality and loss of infective capacity, supporting the hypothesis that both absolute and functional cercarial life-spans are limited by the rates at which energy stores are utilized.

We present a discrete time model for age-related changes in the mean and variance of the number of helminth parasites per host. We assess the degree of aggregation as the negative binomial parameter, k, and use the model to examine the influence of various factors on changes in aggregation with host age: discrete versus continuous infection; the degree of predisposition to infection; infection rate; parasite survival rate; and the variance in exposure to infective stages. The model can produce both increases and decreases in k with host age. However, with parameter values typical of many human helminth infections, a monotonic increase in k (decrease in aggregation) with age is predicted. With an age-dependent infection rate, convex relationships between k and age are possible. These predictions are consistent with data from field studies, but differ from those of previous models which have suggested that k is independent of host age in the absence of density dependence in parasite population dynamics. Differences between the models, and some difficulties in the interpretation of field data, are discussed.

This paper demonstrates that the establishment and growth of the filarial nematode parasite, Litomosoides carinii, is reduced in pyridoxine-deficient cotton rats. Young cotton rats were assigned to one of three dietary groups: vitamin B6-deficient cotton rats (B6 –AL) were fed a pyridoxine-free diet ad libitum; pair-fed controls (B6 + PF) were fed the same amount of pyridoxine-free diet as animals in the deficient group and given daily oral supplements of 100 μg pyridoxine; and pyridoxine-sufficient controls (B6 + AL) were fed the pyridoxine-free diet ad libitum and supplemented daily with 100 μg pyridoxine. Half of each group was infected with 50 L3 of L. carinii by subcutaneous injection 8 weeks after the start of the experimental feeding period. B6-deficient cotton rats ate less (P < 0·001) and gained less weight (P < 0·001) than B6-supplemented controls. The levels of microfilaraemia in deficient animals, measured weekly throughout the experiment by taking blood smears, was significantly lower than in supplemented animals (P < 0·001). The deficient rats became latent for L. carinii at 20 weeks post-infection, whereas there was patent microfilaraemia in rats in the other dietary groups until the end of the experiment. Smaller (P < 0·001) and fewer (P < 0·05) adult worms were recovered from the pleural and abdominal cavities of deficient animals than from either pair-fed or sufficient controls at autopsy 28 weeks post-infection.

Peptides belonging to the FMRFamide family are widely distributed amongst invertebrates. We report here on the isolation of the FMRFamide neuropeptide AF2 (Lys-His-Glu-Tyr-Leu-Arg-Phe-NH2) from the parasitic nematode Haemonchus contortus. Immunocytochemical techniques showed that FMRFamide-like material was distributed in several regions of these organisms including nerve cords and cell bodies of the central nervous system. AF2 was isolated using a method that employed 6 steps of reverse-phase HPLC. The concentration of AF2 in this organism was approximately 30 pmol/g of nematode.

The prediction that richness and diversity of helminth communities would be greater in tropical fish was tested by analysing total and intestinal helminth communities in samples of Anguilla reinhardtii taken from 10 localities in tropical and semi-tropical freshwaters in Queensland, Australia. Comparison of findings with those from A. anguilla in the UK revealed that regional species richness was higher in Queensland and that helminth communities harboured a suite of common species, virtually all eel specialists, that were responsible for the observed higher levels of similarity between component communities. All measures of community richness and diversity adopted indicated that the poorest helminth communities in A. reinhardtii were comparable with the richest ones reported from the northern temperate A. anguilla. The richest communities in A. reinhardtii were more diverse than those reported from any other species of fish, whether marine or freshwater, to date, and were comparable with the communities found in some species of aquatic birds. Both tropical conditions and age and endemism of the host, i.e. the time hypothesis, were considered as explanations for this richness but neither could be favoured. A cautious approach to generalizations based on data from temperate regions only is advocated, and the need for more studies on tropical parasite communities is strongly emphasized.