1. The most striking successes with the method of biological control by the introduction of the parasitic and predaceous enemies of foreign pests, appear to have been obtained mainly in island areas; some doubt exists concerning the utility of the method in continental areas.

2. Three main objections may be raised to the use of the method of parasitic introduction in continental areas. The first is based on the similarity between the faunas and floras of such continental areas as Europe and North America, the second on the polyphagous habit of many parasites, the third on the behaviour of hyperparasites in relation to introduced primaries.

3. The studies made on the natural control of indigenous species in continental areas show that control is effected by complexes of many factors differing both quantitatively and qualitatively in the different parts of the habitat.

4. This fact appears at first sight to indicate the inutility of many of the factors involved, as, for example, the parasitic enemies.

5. The answer to this difficulty is to be found principally in the fact that the quantitative and qualitative fluctuations in the composition of the controlling complexes are much less important than they appear.

6. The effect of the disappearance of controlling factors of the two main types from controlling complexes of various kinds, is studied in detail in order to make this clear.

7. The action of many factors is thus shown to be much less important than it appears at first sight. Nevertheless, this action is real and to a certain degree indispensable in a given environment. The disappearance of a factor from a system in equilibrium will thus permit the increase and spread of the host.

8. The effects which follow the introduction of foreign pests in continental areas constitute a clear demonstration of their escape from control.

9. The evidence indicates that the factor whose absence has permitted increase and spread is the parasitic factor.

10. Equilibrium can be re-established in several ways. Mechanical methods must not be neglected and will normally interfere very little with the biological method; but it is to the latter method that we must turn for a hope of permanent and inexpensive control.

11. The practical results obtained by parasite introduction in continental areas tend in general to support the ideas defended in this paper, and justify the introduction of the native parasites and predators of foreign pests even in the case of exchanges between continental areas.

A cationic Schistosoma mansoni cercarial antigen was shown to be a serine protease as it was capable of hydrolysing N-acetyl-dl-phenylalanine β-naphthyl ester (NAPBNE) after precipitation by immunoelectrophoresis, and this reaction was modulated by the serine protease inhibitors phenylmethanesulfonyl fluoride (PMSF) and diisopropylfluorophosphate (DFP). The antigen in the immunoprecipitin arcs could also be radio-isotope labelled with tritiated DFP. The peptidolytic enzyme identified in immunoelectrophoresis with polyspecific sera and radio-isotope labelled with tritiated DFP had a relative molecular size of approximately 27 kDa in sodium dodecyl sulphate polyacrylamide gel electrophoresis (SDS–PAGE), and evidence obtained after partial purification, SDS–PAGE and immunoblotting supported this size estimate for the enzyme. A rabbit antiserum raised against the peptidolytic antigen reacted against a doublet of antigens at 27/28 kDa in immunoelectrophoresis arcs and against an antigen of 60 kDa in Western immunoblots of crude cercarial homogenate. However, the latter serum precipitated the cationic antigen in immunoelectrophoresed cercarial homogenates only after pre-incubation of the homogenates with PMSF. Fractions containing the partially purified protease also degraded radio-isotope labelled human IgG. The reactivity of a range of polyspecific and monospecific rabbit antisera in Western blots with larval extracts indicated that antibody responses against the 27/28 kDa doublet may be modulated. When immunized with material which contained the 27 kDa enzyme as a major constituent, and which was secreted by S. mansoni cercariae during transformation, only 5 of 16 mice produced antibody to this antigen that was detectable in Western blots. The 5 antibody ‘responder’ mice were significantly (P<0·001) protected against challenge with a percutaneous infection of S. mansoni cercariae compared with a group of mice also immunized with CTF, but which had not produced antibodies against the 27/28 kDa doublet. The results indicate that the 27 kDa serine protease of S. mansoni larvae is a target that is sensitive to immunological attack.

A protein phosphatase exclusively dependent upon micromolar amounts of Ca2+ and calmodulin has been identified and partially purified from Leishmania spp. Complete obliteration of its activity is observed in the presence of calmodulin antagonists such as trifluoperazine, fluphenazine and calmidazolium. Relative insensitivity to okadaic acid and lack of activation in the absence of Ca2+ and calmodulin distinguishes this enzyme from PP1, PP2A and PP2C-type protein phosphatases. Cross-reactivity of the enzyme was observed with antibodies that recognize both the A and B chains of calcineurin, a PP2B type Ca2+ and calmodulin-dependent phosphatase from brain. FK506, an immunosuppresive drug that inhibits the enzyme from other sources inhibited the enzyme only in the presence of exogenous FK binding protein, whereas Cyclosporin A inhibited the enzyme in crude preparations. Taken together these results reveal the presence of a Ca2+ and calmodulin-dependent phosphatase from Leishmania. This is the first report of the presence of a PP2B-type protein phosphatase from a pathogenic protozoa.

The phylum of Apicomplexa comprises parasitic protozoa that share distinctive features such as the apical complex, the apicoplast, specialized cytoskeletal components and secretory organelles. Other unique cytoplasmic inclusions sharing similar features have been described in some representatives of Apicomplexa, although under different denominations. These are the crystalloid body, present for example in Cryptosporidium, Plasmodium and Cystoisospora; the refractile body in Eimeria and Lankesterella; and virus-like particles, also present in Eimeria and Cryptosporidium. Yet, the specific role of these cytoplasmic inclusions in the cell cycle of these protozoa is still unknown. Here, we discuss their morphology, possible inter-relatedness and speculate upon their function to bring these organelles back to the attention of the scientific community and promote new interest towards original research on these elusive structures.

While parasitism is recognized as the most common mode of existence on the planet, and hosts from virtually all ecosystems have been studied, very little is known about the parasites found in deep-sea hydrothermal vent ecosystems and even less is known about their ecology, evolution, and effects on their hosts. The purpose of this work is to offer a comprehensive review of our state of knowledge about parasitism in the deep-sea vents and to pose and address specific questions for future studies. Because the deep-sea environment itself may influence the number and types of parasites found in the vents, non-vent (below 1000 m) and vent deep-sea data were used in a comparative analysis to account for this factor as a potential major determinant of the parasite fauna in the vents. Based upon analysis of these data, it is highly likely that the reason why so few parasites are currently known from deep-sea vents, even given the low diversity of hosts in this ecosystem, is simply that their inconspicuous nature has caused them to be overlooked by vent biologists.

The direct effects of phytohormones (auxin and kinetin) and root diffusates on the surface lipophilicity of the plant parasitic nematodes Globodera rostochiensis and Meloidogyne incognita were investigated. The fluorescent lipid probe AF18 (5-N-(octodecanoyl) aminofluorescein) was used to detect surface changes. Root diffusates increased AF18 uptake by G. rostochiensis while it had no effect on M. incognita. Kinetin and auxin decreased AF18 uptake in G. rostochiensis, while they had the opposite effect on M. incognita. Auxin/kinetin ratio was also found to be important in triggering the surface changes, especially at high concentrations. Whether plant nematodes have auxin and/or kinetin binding proteins is discussed as well as the mechanism behind the surface lipophilicity changes due to root diffusates and phytohormones.

Previous laboratory studies have indicated that tenebrionid beetles infected with Hymenolepis diminuta display behavioural alterations. These are assumed to increase the likelihood of this intermediate host being predated by the Rattus sp. definitive host, and hence provide support for the Manipulation Hypothesis. We tested this hypothesis by both assessing beetle behaviour and predation rates in semi-naturalistic environments. Behavioural assays showed that infected beetles were more often exposed (not concealed under boxes) than uninfected beetles. However, there were no differences in predation rates between infected and uninfected beetles. We discuss this in terms of the historical and evolutionary contexts under which such altered behaviours could evolve and their implications for the Manipulation Hypothesis.

Adult Haemonchus contortus obtained from freshly killed sheep were incubated in a medium similar to Tyrode's saline containing known amounts of D-[U-14C]glucose. The worms appeared to remain in a healthy condition throughout the incubations which lasted for up to 6 h. All the radioactivity was recovered either within the worms or in the incubation vessel in the form of CO2, excretory products and unmetabolized glucose. An oxygen electrode in the incubation liquid showed that, because of their rapid oxygen consumption, the worms were under anaerobic conditions during all or nearly all of the incubation period, despite the presence of oxygen in the gas phase of the vessel. In terms both of quantity and radioactivity the main excretory products in solution were propan-1-ol, propionate and acetate. Smaller amounts of ethanol, lactate and succinate were excreted. Much radioactivity was associated with the expired CO2. At the end of the incubations glucose other than the initial radioactive glucose was detected indicating that the absorption of glucose by H. contortus is complex and needs further investigation. The present experiments suggest that the rate of absorption depends on the worm's need for nutrients. The results are compared with those found for other parasitic nematodes, particularly Ascaris lumbricoides, Heterakis gallinae and Trichuris vulpis. The importance of CO2 fixation in the utilization of energy from glucose by H. contortus is discussed.

Blastocrithidia familiaris were found to be parasitic in the midgut, ileum and rectum of Lygaeus pandurus. The host—parasite relationship is described. Attachment of parasites in the midgut and ileum occurs by interdigitation of expanded flagella over and between the microvilli. No attachment to microvilli was observed where extracellular membranes form well-organized layers which lie parallel to the gut wall and completely separate the microvilli border from the lumen. The extracellular membranes originate from delamination of the outer unit membrane of microvilli which consists of a double plasma membrane. The function of the extracellular membrane layers and their possible role as a peritrophic membrane in preventing parasite attachment is discussed in relation to previous studies on midgut cells in Hemiptera with a similar apical coat on midgut microvilli. In the rectum, parasites attach to the cuticle of the gland cells and not to the rest of the rectal wall. Attachment to the cuticle occurs by the formation of hemidesmosomes. A comparison of the relationship of B. familiaris and its host to previous ultrastructural studies of associations between kinetoplastid flagellates and their respective hosts is discussed.

Individual susceptibility to malaria infection, disease and death is influenced by host genotype, parasite virulence and a number of environmental factors including malaria-specific immunity. Immune responses are themselves determined by a combination of host genes and environmental effects. The extent to which host genotype limits the spectrum of possible immune responses may influence the outcome of infection and has consequences for vaccine design. Associations have been observed between human major histocompatibility complex (MHC) genotype and susceptibility to severe malaria, but no similar associations have been observed for mild malarial disease or for specific antibody responses to defined malaria antigens. Epidemiological studies have shown that, in practice, neither T helper cell nor antibody responses to malaria parasites are limited by host MHC genotype, but have revealed that genes lying outside the MHC may influence T cell proliferative responses. These genes have yet to be identified, but possible candidates include T cell receptor (TcR) genes, and genes involved in TcR gene rearrangements. More importantly, perhaps, longitudinal epidemiological studies have shown that the anti-malarial antibody repertoire is selective and becomes fixed in malaria-immune individuals, but is independent of host genotype. These findings suggest that the antibody repertoire may be determined, at least in part, by stochastic events. The first of these is the generation of the T and B cell repertoire, which results from random gene recombinations and somatic mutation and is thus partially independent of germline genes. Secondly, of the profusion of immunogenic peptides which are processed and presented by antigen presenting cells, a few will, by chance, interact with T and B cell surface antigen receptors of particularly high affinity. These T and B cell clones will be selected, will expand and may come to dominate the immune response, preventing the recognition of variant epitopes presented by subsequent infections - a process known as original antigenic sin or clonal imprinting. The immune response of an individual thus reflects the balance between genetic and stochastic effects. This may have important consequences for subunit vaccine development.

A longitudinal investigation on natural populations of Ascaris in humans and pigs and an investigation of soil contamination with Ascaris eggs were carried out from June 1993 to June 1994 in 2 villages, Manhu area, Xinjian County, Jiangxi Province, China. Results from these studies indicate that although human ascariasis is endemic there is significant fluctuation in both prevalence and the mean number of eggs/g faeces (epg) of the communities. Fluctuation of age-stratified prevalence and mean epg was detected in children but not in most adult groups. Most cases of human ascariasis were judged to involve low intensities of infection and a typical overdispersion distribution pattern was observed through the year. It was estimated that during the year, nearly half of the eggs discharged in the environment came from infections in children aged between 2 and 15 years which accounted for about 30% of the total population. Soil in and around houses and in vegetable gardens was found to be contaminated by Ascaris eggs and this situation remained relatively stable throughout the year. Monthly developmental rate of Ascaris eggs in soil was detected and the results suggest that the fluctuation in prevalence observed during the year should be directly attributed to the effect of seasonality of egg development. Features of Ascaris infection in pigs were found to be similar to those in humans except for a lower mean intensity of infection. The possibility of cross-infection of Ascaris between human and pig hosts is discussed.

The transmission of Ascaris suum was studied in outdoor reared pigs. From May to June 2001, 6 farrowing paddocks were naturally contaminated with A. suum using experimentally infected seeder pigs. Early July, 1 sow farrowed on each paddock. One piglet per litter was slaughtered every second week starting at week 3 post-partum (p.p.) for registration of liver white spots and recovery of A. suum from the lungs and the small intestine. The last pigs were slaughtered at week 19 p.p. Faeces was examined for parasite eggs and blood was analysed for A. suum-specific antibodies. Weaning took place at week 7 p.p. by removing the sow. Paddock infection levels were estimated by regular examination of soil samples and in late June and late November using parasite naïve tracer pigs. Paddock contamination was high but eggs developed slowly resulting in a low initial transmission to the experimental pigs. By week 5 p.p. transmission had increased and the numbers of infective eggs in the soil increased during the study. The results indicate a continuous uptake of infective eggs, but visceral larval migration was reduced with time, probably due to the development of a pre-hepatic barrier. Nevertheless, a rather large population of adult worms remained in the pigs throughout the study, and it may primarily have been eggs ingested in the early infection phase that gave rise to the patent infections. It is suggested that neonatal exposure may result in increased persistence and size of adult worm burden and that the higher ‘life-time worm burden’ may be of significant economic importance.

Few studies have focussed on the health and immunity of triploid Atlantic salmon and therefore much is still unknown about their response to commercially significant pathogens. This is important if triploid stocks are to be considered for full-scale commercial production. This study aimed to investigate and compare the response of triploid and diploid Atlantic salmon to an experimental challenge with Neoparamoeba perurans, causative agent of amoebic gill disease (AGD). This disease is economically significant for the aquaculture industry. The results indicated that ploidy had no significant effect on gross gill score or gill filaments affected, while infection and time had significant effects. Ploidy, infection and time did not affect complement or anti-protease activities. Ploidy had a significant effect on lysozyme activity at 21 days post-infection (while infection and time did not), although activity was within the ranges previously recorded for salmonids. Stock did not significantly affect any of the parameters measured. Based on the study results, it can be suggested that ploidy does not affect the manifestation or severity of AGD pathology or the serum innate immune response. Additionally, the serum immune response of diploid and triploid Atlantic salmon may not be significantly affected by amoebic gill disease.

We investigated the timing of seasonal activity of the seabird tick Ixodes uriae by examining engorgement rates of off-host ticks and the prevalence of infested common guillemots (Uria aalge) and kittiwakes (Rissa tridactyla) on the Isle of May, Scotland. More than 99% of I. uriae fed during the period late April to early August which coincided with the seabird breeding season. No ticks fed in the autumn and winter, and none over-wintered in the engorged state. Peak tick feeding activity was during the hosts' incubation period in May and early June, when over 70% of birds were parasitized. Few ticks were active when guillemot chicks were being brooded, or during the early part of kittiwake chick-rearing. Adult female tick feeding peaked in the first week of June, while nymphal and larval activity peaked in the third week of May. Adult female ticks fed later than nymphs on guillemots and larvae on both guillemots and kittiwakes; however, there was considerable temporal overlap in the feeding periods of the 3 stages on both hosts. All larvae and nymphs fed in May and early June, but up to 18% of adult females remained unengorged at this time, and probably deferred feeding by 1 year. A small proportion of individuals which fed as larvae in May appeared to feed again as nymphs in July of the same season, thereby shortening their life-cycle by 1 year. The duration of the life-cycle of I. uriae on the Isle of May is typically 3 years but varies from 2 to 4 years. Ticks feeding on guillemots did so significantly earlier than those feeding on kittiwakes, possibly because guillemots start breeding earlier in the year. Breeding of both guillemots and kittiwakes was later in 1994 than in 1993. This delayed the activity of all tick stages on kittiwakes and adult female ticks on guillemots, but did not affect the timing of activity of nymphs or larvae feeding on guillemots.

An in vitro investigation has been made of the relative roles of the gut and tegument in the absorption of the neutral amino acids L-alanine and L-leucine by a marine fish-gill parasite, Diclidophora merlangi. The use of ligatures to preclude oral ingestion of trace-labelled medium has proved inadequate, invariably damaging the tegument, as revealed by stereoscan electron microscopy, and resulting in artifactual levels of absorption. Three alternative procedures have given consistently reliable data on the route of entry of low molecular weight substrates. (1) Ultrastructural examination of worms previously incubated in electron-dense cationic tracers has shown that, in vitro, there is no oral intake of sea water. (2) The suspending of worms in trace-labelled medium with the mouth out of the medium and comparing amino acid uptake with that of worms totally immersed in medium has revealed no statistically significant difference in the absorption levels. (3) Application of section (freeze-dried) auto-radiography to detect diffusible isotope has demonstrated directly transtegumental absorption of a neutral amino acid. It is concluded from these experiments that Diclidophora has a tegumental transport system for absorbing certain neutral amino acids, and whilst, clearly, the worm is sanguinivorous and digests blood in a well-developed gut, it may also be capable of supplementing this diet with low molecular weight organic nutrient absorbed directly from sea water via the tegument.

Mean villus height, crypt depth and the number of 5-HT-positive enterochromaffin (EC) cells have been examined in two regions of the small intestine (20–30° and 60–70° distance from the pylorus) of male, 6 to 8-week-old, C57 mice following a 5-cysticercoid infection of the rat tapeworm, Hymenolepis diminuta. Test mice and sham-infected controls were autopsied 0, 4, 8, 10, 14 and 28 days post-primary infection (p-1°-i) and 2, 4, 5, 7 and 14 days post-secondary infection (p-2°-i), administered 28 days p-1°-i. Morphometric analysis revealed a statistically significant increase in crypt depth in the 60–70°o intestine region in infected mice during both primary and secondary infections; no significant deviation from the control was observed for villus height in infected mice. Statistical analysis showed a significant increase in the number of 5-HT-positive EC cells in infected mice. This response occurred in the lower portion of the intestine on days 10-p-1°-i and 5-p-2°-i, and was not due to increased mucosal surface area in this region. Results are discussed with reference to murine cestode rejection and the possible involvement therein of the neuroendocrine system.

125I-Schistosoma mansoni schistosomulum surface antigens were immunoprecipitated with human antibodies from individual Egyptian patients diagnosed as being either acutely or chronically infected with S. mansoni. Both sets of patients were found to have IgG antibodies in their sera capable of immunoprecipitating the major Mr > 200, 38 and 32K antigens. However, the immunoprecipitation of the Mr 200K antigen was found to constitute a significantly greater proportion of the total precipitate achieved with acute sera than with chronic sera. The Mr 38 and 32K antigens were more variably precipitated by the acute sera than the chronic sera but the proportion of the total precipitation that these two antigens constituted was not found to be significantly different between the two sets of sera. Immunoprecipitation with pooled antibodies absorbed with egg and adult worm homogenates which had been treated to remove either carbohydrate or polypeptide epitopes demonstrated that the Mr > 200K antigen was the principal target of egg-cross-reactive anti-carbohydrate antibody amongst the antigens detected. The Mr 38 and 32K antigens were found to be precipitated by antibodies to protease-sensitive and periodate-insensitive polypeptide epitopes. These results are consistent with egg-cross-reactive anti-carbohydrate IgG antibody making a greater contribution to schistosomulum surface recognition in acute infection than in chronic infection. Indeed the presence of a higher level of egg-cross-reactive and anti-carbohydrate antibody directed against schistosomulum surface epitopes in an acute serum pool than in a chronic serum pool was confirmed by measurement of antibody binding to whole schistosomula.

The frequency distribution of parasites in hosts commonly follows a negative binomial or similar distribution. Under certain conditions the magnitude of parasite-associated host mortality can be estimated by comparing the tail of the observed distribution to that of the distribution predicted from the first few points of the data. For the technique to work the following assumptions need to be met: mortality in lightly infected fish must be rare; infection and consequent mortality occur only in fish younger than those sampled; and the frequency distribution of the parasite at the time of infection should conform to a known probability distribution. The method was applied to frequency distributions of blastocysts of Callitetra rhynchus gracilis in 898 Saurida tumbil (Bloch) and 5013 S. undosquamis (Richardson). Parasite-associated mortality in S. tumbil was calculated to be at least 11 % in males and 2% in females. For S. undosquamis, estimated mortality was about 5% in males and 3% in females. The numbers of parasites estimated to produce a 0·5 probability of death, the parasitological equivalent of an LD50 were 3·4 and 5·7 for S. tumbil males and females, and 18 and 3 for S. undosquamis males and females respectively.

Virulence is part of the proximate base to the understanding of host–parasite interactions, and therefore it is crucial to interpret parasite-induced damage as an outcome of co-evolution between parasite and host. We experimentally investigated the impact of 2 congeneric ticks with contrasting transmission modes and host specificity, on the health of songbird nestlings (Parus major). The nidicolous Ixodes arboricola lives in tree holes where it obtains blood from adult birds and their nestlings, hence is prone to vertical transmission. All developmental stages of the field-dwelling Ixodes ricinus feed on different host taxa, hence are prone to horizontal transmission. Within each nest we infested 3 nestlings with varying loads of I. ricinus and 3 with Ixodes arboricola. Ixodes arboricola had no negative effects on nestling health (haematocrit, inflammation, body size, body condition). Body size was positively associated with the number of feeding I. arboricola adults. In contrast, I. ricinus infestations by nymphs and adult ticks led to haematocrit reductions. Furthermore, I. arboricola was a more prudent feeder, with longer feeding durations for smaller bloodmeals. Although the negative effects on nestling health were limited, our results are in line with current theories, stating that virulence increases with horizontal transmission, reduced dependence on individual hosts and reduced relatedness among co-exploiting ticks.

We describe the infectivity, virulence, cultivating conditions, and phylogenetic positions of naturally occurring oomycete parasites of Daphnia, invertebrates which play a major role in aquatic food webs. Daphnia pulex individuals were found dead and covered by oomycete mycelia when exposed to pond sediments. We were able to extract 4 oomycete isolates from dead Daphnia and successfully cultivate them. Using the ITS and LSU rDNA sequences, we further showed these isolates to be distinct species. The isolates were experimentally demonstrated to be parasitic and not saprobic. After exposure to the parasites, Daphnia mortality was much higher than that reported for Daphnia infected with other known parasite species. Therefore, it is likely that oomycete parasites are important selective pressures in natural Daphnia populations. Moreover, their close phylogenetic relationship to parasites of fish and algae suggests that the stability of aquatic food webs (i.e. fish–Daphnia–algae) might be influenced by the shared parasite communities.