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Solute flux into parasitic plants.

Parasitic plants form intimate contacts with host tissue in order to gain access to host solutes. There are a variety of cell types within the host which parasitic plants could access to extract solutes. Depending on the degree to which the parasite has embraced the parasitic lifestyle, the extent of solute flux and the pathways used to transfer solutes from host to parasite will vary. To date, a variety of experimental approaches argue for diversity in the mechanisms and the routes by which parasites accumulate host solutes. Contact between host and parasite ranges from direct lumen-to-lumen links between host and parasite xylem and continuity between the sieve elements of host and parasite, to the involvement of transfer cells between host and parasite. Progress has been slow since Solms-Laubach distinguished types of parasitic plants that fed from host phloem or xylem in 1867, but advances in clearly delineating the pathways that link host and parasite should now be possible using fluorescent proteins expressed and restricted to particular cell types of the host. This will initially necessitate using Arabidopsis, but should allow the types of connection, i.e. symplasmic or apoplasmic, to be determined and then the identification of parasite transporters responsible for solute flux.

Adaptation, Physiological↗

[Role of host behavior in the life cycle of parasites].

Parasite is an organism which, at least in a part of its ontogeny uses another living organism as a proper environment for its life. In the "parasite-host" relationships, formed by both these components, the parasite itself bears the burden of formation and maintenance of these relationships in a balance. Three factors play the main role in this process: (1) physiological adaptations of the parasite, enabling survival in the host body and resistance against defence reactions of the host; (2) morphological adaptations, leading to changes in the body structure of the parasite, dependent on location in the host body; and (3) behavioural adaptations assuring contacts of parasites with their hosts. In the process of evolution most groups of parasites evolved a complicated life cycle, with change of host and outer environment, succeeding in maturation of the parasite and production of the offspring. To pursuit this aim the parasite takes advantage from the behaviour of its potential host, its food preferences (e.g. by inclusion into a food chain), periodic or circadian migrations, and generally from its mode of life. The parasite modifies behaviour of its hosts, sometimes to a high degree, especially the behaviour of intermediate hosts, making them more conspicuously displayed for predators, the most often their final hosts. The parasite itself changes also its behaviour to be more attractive for a potential host or to enhance the possibility of finding a proper host. Finally, in a host population the parasite bears upon the position of particular host individuals by degradation of dominants and shifting them from reproduction. This phenomenon may be considered as a self-defence of the host population against reproduction of ill individuals, weaken by a burden of parasites.

Animals↗

Community ecology of the metazoan parasites of grey mullets, Mugil platanus (Osteichthyes: Mugilidae) from the littoral of the state of Rio de Janeiro, Brazil.

One hundred and fifty specimens of grey mullets, Mugil platanus Günther, collected from the littoral of the State of Rio de Janeiro, Brazil, between June 1984 to August 1988, were necropsied to study their infracommunities of metazoan parasites. Twenty-five species of metazoan parasites were collected. One hundred and forty-nine (99.3%) fishes were parasitized by one or more metazoan species. According to their importance values, 13 species were considered dominant, 7 species codominant and 2 species subordinate (digeneans were the majority of the total number of parasite specimens collected, with 63.7%). The presence of larval stages of digeneans and cestodes suggest that M. platanus is at an intermediate level in the marine trophic web. The Simpson index for all parasite species was low (0.059) indicating lack of dominance by a particular species in the parasite community. The parasites of M. platanus showed a typical overdispersed pattern of distribution. The capture localities of the hosts were consolidated into 3 groups and its parasite communities showed high qualitative and quantitative similarity, with high values for the Sorenson similarity coefficient and Percent similarity coefficient. Six species had a positive correlation between the host's total body length and parasite intensity, while three species had a negative correlation. Four species had a positive correlation between the host's total body length and parasite prevalence and one species had negative correlation. Five parasite species had differences in prevalence and intensity in relation to sex of host, but these differences were confounded with the total length of male and female fishes. The mean diversity in the infracommunities of M. platanus was H' = 0.936 +/- 0.477, without correlation with the host's total length and without significant difference between male and female fishes. In the ectoparasites, the infracommunities showed overall positive association, 6 pairs of copepod species had a significant positive association. One pair of species showed a negative covariation between parasite intensities. Infracommunities of endoparasites also showed overall positive association, and four species pairs of digeneans had a significant positive association. Only one had a negative association. The specialist components with low prevalence values predominated in the parasite community, which, based on these factors, was considered closer to the isolationist type.

Animals↗

[The fate of parasites of animal origin transmitted to humans].

The fate of a parasite transmitted from an animal to man depends on the ability of the contaminating agent to reach a place where it can thrive, to find necessary nutrients, and to resist host defense mechanisms. The purpose of this study was to evaluate the incidence of transmission of parasites from animals to man and to determine to what extent transmission is followed by development. Stenoxenic parasites whose life cycle requires transmission from animals to man obviously develop in man and then return to animals. These parasites cause holozoonoses of the cyclozoonosis type. Some euryxenic parasites can develop as well in man as in animals. These parasites can cause holozoonoses of the amphixenoses type. Other presumably euryxenic parasites can be transmitted from animals to man but not vice versa. These parasites are hemizoonoses agents. Non-transmission back from man to animals can be observed under several circumstances: incomplete development in man with failure to reach the stage at which transmission back to animals is possible; full development but with immaturity or sterility of the elements of dissemination necessary for transmission back to animals; full development but no way of evacuating elements of dissemination; full development and evacuation but with failure of elements of dissemination to survive. In these four cases man constitutes a dead-end for the parasite. A fifth possibility is that the parasite reaches full development but transmission back to animals cannot occur because man is not preyed upon by a carnivorous animal. In this case parasites are potential agents of holozoonoses and man is a cul-de-sac for the involved parasites.

Angiostrongylus↗

Egg-laying behaviour by shiny cowbirds parasitizing brown-and-yellow marshbirds.

We studied the laying behaviour of shiny cowbirds, Molothrus bonariensis, parasitizing brown-and-yellow marshbirds, Pseudoleistes virescens. Shiny cowbirds lay two egg morphs, spotted and white immaculate. Brown-and-yellow marshbirds eject the white egg morph but accept the spotted morph. The incidence of parasitism in this host was 66.5%, and half of the parasitized nests had more than one shiny cowbird egg. There was a positive relationship between the number of parasitic events and the availability of nests in the laying stage, but parasitic events reached a plateau when the availability of nests was high. The distribution of parasitic eggs per nest was more clumped than expected by chance. Shiny cowbird females synchronized parasitism with host laying in 80% of the cases. They seldom parasitized nests before the host started laying or after the nest had been deserted or predated. The majority of multiply parasitized nests were parasitized by more than one female. Females that lay white eggs did not avoid parasitizing brown-and-yellow marshbird nests. Egg pecking by cowbird females resulted, on average, in one egg lost per parasitic event and the probability of being broken was greater for host eggs when host and parasitic eggs where both in the nest. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Heme biosynthesis by the malarial parasite. Import of delta-aminolevulinate dehydrase from the host red cell.

The mouse and human malarial parasites, Plasmodium berghei and Plasmodium falciparum, respectively, synthesize heme de novo following the standard pathway observed in animals despite the availability of large amounts of heme, derived from red cell hemoglobin, which is stored as hemozoin pigment. The enzymes, delta-aminolevulinate dehydrase (ALAD), coproporphyrinogen oxidase, and ferrochelatase are present at strikingly high levels in the P. berghei infected mouse red cell in vivo. The isolated parasite has low levels of ALAD and the data clearly indicate it to be of red cell origin. The purified enzyme preparations from the uninfected red cell and the parasite are identical in kinetic properties, subunit molecular weight, cross-reaction with antibodies to the human enzyme, and N-terminal amino acid sequence. Immunogold electron microscopy of the infected culture indicates that the enzyme is present inside the parasite and, therefore, is not a contaminant. The parasite derives functional ALAD from the host and the enzyme binds specifically to isolated parasite membrane in vitro, suggestive of the involvement of a receptor in its translocation into the parasite. While, ALAD, coproporphyrinogen oxidase, and ferrochelatase from the parasite and the uninfected red cell supernatant have identical subunit molecular weights on SDS-polyacrylamide gel electrophoresis and show immunological cross-reaction with antibodies to the human enzymes, as revealed by Western analysis, the first enzyme of the pathway, namely, delta-aminolevulinate synthase (ALAS) in the parasite, unlike that of the red cell host, does not cross-react with antibodies to the human enzyme. However, ALAS enzyme activity in the parasite is higher than that of the infected red cell supernatant. We therefore conclude that the parasite, while making its own ALAS, imports ALAD and perhaps most of the other enzymes of the pathway from the host to synthesize heme de novo, and this would enable it to segregate this heme from the heme derived from red cell hemoglobin degradation. ALAS of the parasite and the receptor(s) involved in the translocation of the host enzymes into the parasite would be unique drug targets.

Animals↗

Acute Plasmodium chabaudi chabaudi malaria infection induces antibodies which bind to the surfaces of parasitized erythrocytes and promote their phagocytosis by macrophages in vitro.

CBA/Ca mice infected with 5 x 10(4) Plasmodium chabaudi chabaudi AS-parasitized erythrocytes experience acute but self-limiting infections of relatively short duration. Parasitemia peaks ( approximately 40% infected erythrocytes) on day 10 or 11 and is then partially resolved over the ensuing 5 to 6 days, a period referred to as crisis. How humoral and cellular immune mechanisms contribute to parasite killing and/or clearance during crisis is controversial. Humoral immunity might be parasite variant, line, or species specific, while cellular immune responses would be relatively less specific. For P. c. chabaudi AS, parasite clearance is largely species and line specific during this time, which suggests a primary role for antibody activity. Accordingly, acute-phase plasma (APP; taken from P. c. chabaudi AS-infected mice at day 11 or 12 postinfection) was examined for the presence of parasite-specific antibody activity by enzyme-linked immunosorbent assay. Antibody binding to the surface of intact, live parasitized erythrocytes, particularly those containing mature (trophozoite and schizont) parasites, was demonstrated by immunofluorescence in APP and the immunoglobulin G (IgG)-containing fraction thereof. Unfractionated APP (from P. c. chabaudi AS-infected mice), as well as its IgG fraction, specifically mediated the opsonization and internalization of P. c. chabaudi AS-parasitized erythrocytes by macrophages in vitro. APP from another parasite line (P. c. chabaudi CB) did not mediate the same effect against P. c. chabaudi AS-parasitized erythrocytes. These results, which may represent one mechanism of parasite removal during crisis, are discussed in relation to the parasite variant, line, and species specificity of parasite clearance during this time.

Acute Disease↗

Surface properties of extracellular malaria parasites: morphological and cytochemical study.

Morphological and cytochemical surface characteristics of isolated malaria parasites (Plasmodium berghei) and host erythrocytes were compared by electron microscopy by using thin section and carbon replica techniques. Erythrocytes were uniform in shape and had fine, granular surfaces. In contrast, free parasites exhibited a variety of sizes, shapes, and surface textures. Fine surface stippling was a common topographical feature of isolated parasites. Small, infective forms often had patterned surfaces resulting from the protuberance of an underlying thick intermediate layer. Results of cytochemical analysis using a sialophilic colloidal iron stain indicated that the malaria parasite's surface lacked exposed sialic acid groups which would normally give rise to a net negative surface charge common to erythrocytes. Biochemical assay demonstrated that malaria parasites contained about one-half the amount of sialic acid per unit weight as did control red cell extracts. Similarly, external acidic mucopolysaccharide coats of free parasites, as revealed by ruthenium red staining were extremely thin as compared with the thick glycocalyx layer of red cells. Lipid plaques at the surface of parasites and red cells were localized by lipophilic iron colloid staining. Although the gross patchwork distribution of plaques was somewhat similar for the two cell types, the parasites were stained more intensely and had a closer-knit patchwork pattern than those exhibited by the erythrocytes. Such findings indicate that there are slight differences in the arrangement of phospholipids at the surfaces of limiting membranes of host cells and parasites. The significance of the above cytochemical surface properties of the malaria parasite (which are seemingly akin to those of intracellular organelles is discussed in relation to certain host-parasite interactions, such as parasite adhesion to target cells and enhanced clearance of extracellular parasites.

Cell Membrane↗

Altered dietary nutrient intake maintains metabolic homeostasis in parasitized larvae of the insect Manduca sexta L.

Manduca sexta larvae exhibited altered food selection over a 2- or 3-day feeding period when parasitized by Cotesia congregata, and offered a choice of two chemically defined diets, one containing casein without sucrose and a second with sucrose but no casein. While normal larvae consumed the diets in a ratio of approximately 2:1 protein:carbohydrate (w/w), parasitized insects consumed a ratio of approximately 1:1. The altered nutrient ratio consumed by parasitized insects was principally due to a decrease in consumption of the protein diet, and was only partially explained by their lower growth. Conditioning larvae for 1 day to either one of the choice diets had little effect on subsequent dietary intake over a 2-day feeding period. Conditioned larvae, regardless of parasitism, initially fed on the opposite diet immediately after conditioning. Although this suggests that the altered nutrient intake displayed by parasitized insects was not due to any failure in their capacity for dietary selection, these results do not definitively demonstrate an altered nutrient intake target by parasitized larvae. Rather, parasitism may compromise dietary selection, resulting in random feeding. When parasitized larvae were maintained on several isocaloric diets with a varying ratio of casein and sucrose, those larvae feeding on the diet with a ratio of 1:1 of these nutrients supported the largest parasite population. Previous investigation of larvae maintained on a single artificial diet established that parasitized insects display an aberrant induction of gluconeogenesis, so that haemolymph trehalose is maintained at a level equivalent to that of normal insects. In contrast, the present results demonstrated that parasitized larvae offered a choice of diets, and feeding at the altered nutrient ratio above, maintain haemolymph sugar but have the same level of gluconeogenesis as normal larvae given the same dietary choice. These investigations suggest that altered food selection by parasitized M. sexta larvae maintains metabolic homeostasis and, moreover, may be adaptive for C. congregata, potentially maximizing the number of parasites developing in a single host larva.

Animal Nutritional Physiological Phenomena↗

Parasite enzymes as potential targets for antiparasitic chemotherapy.

I have thus far listed a total of 10 potential targets for antiparasitic chemotherapeutic consideration. This is by no means a completed list. Many more will be added to it with time and with more future findings. Among these 10 targets (summarized in Table I), however, one may gain some insight and see a few interesting general trends: (1) Nucleic acid metabolism and carbohydrate-energy metabolism in protozoan parasites appear to be targets for fruitful chemotherapeutic attacks. Their being useful targets results generally from the deficient metabolism in the protozoan parasites. Thus, the main vulnerability among the protozoan parasites is closely associated with their parasitic nature. (2) Microtubules and nervous systems appear to be the main chemotherapeutic targets in helminths. They differ from those in the host not because of their parasitic nature but, more likely, because of the evolutionary distance separating the mammalian hosts and the primitive metazoa. Thus, free-living nematodes, such as Caenorhabditis elegans, have their microtubules just as susceptible to the benzimidazole anthelmintics as those from the parasitic worms. The motoneuronal map of C. elegans is identical with that of Ascaris lumbricoides. Both worms are similarly immobilized by levamisole, piperazine, avermectins, etc. The dual insecticidal and antiexoparasite activities found in the avermectins and milbemycins may also suggest that the free-living insects and the ticks and lice may have the same GABA nervous system. This main discrepancy between protozoan parasites and metazoan parasites may be partly attributable to the higher mutation rates and higher frequencies of genetic recombination among the protozoa, evidenced by the higher rates of development of drug resistance among them. The fast adaptation to a new environment may be essential for survival, but it would also lead to metabolic deficiencies after the protozoa lived in a luxurious environment for a while. This revelation may suggest that future chemotherapeutic studies on parasitic helminths can utilize free-living helminths as models to eliminate many unnecessary technical difficulties. Also, there perhaps could be a further classification among the parasites to term the protozoa "true parasites" and the helminth "pseudo-parasites" from the viewpoint of chemotherapy.

Animals↗

The contagion indicator hypothesis for parasite-mediated sexual selection.

Hamilton and Zuk [Hamilton, W. D. & Zuk, M. (1982) Science 218, 384-387] proposed that females choosing mates based on the degree of expression of male characters obtain heritable parasite resistance for their offspring. Alternatively, the "contagion indicator" hypothesis posits that females choose mates based on the degree of expression of male characters because the latter indicate a male's degree of infestation of parasites and thus the risk that choosing females and their offspring will acquire these parasites. I examined whether parasite transmittability affects the probability that parasite intensity and male mating success are negatively correlated in intraspecific studies of parasite-mediated sexual selection. When females risk infection of themselves or their future offspring as a result of mating with a parasitized male, negative relationships between parasite intensity and male mating success are significantly more likely to occur than when females do not risk such infection. The direct benefit to females of avoiding parasitic infection is proposed to lead to the linkage between variable secondary sexual characters and the intensity of transmittable parasites. The direct benefits of avoiding associatively transmittable parasites should be considered in future studies of parasite-mediated sexual selection.

Adaptation, Biological↗

Increased host cell-Trypanosoma cruzi interaction following phospholipase D treatment of the parasite surface.

We examined the effect of phospholipase D (PLD) treatment on the ability of Trypanosoma cruzi to interact with phagocytic and nonphagocytic host cells. The presence of PLD during the incubation of parasites with mouse peritoneal macrophages caused significant increases in both the number of parasites per 100 macrophages and the percentage of macrophages associated with parasites. Parasites pretreated with PLD, washed, and then incubated with untreated macrophages showed a marked increase in parasite-host cell association. In contrast, when only the macrophages were pretreated with PLD, there was no significant change in the association. Parasites required 45 min of PLD treatment before a significant enhancement in parasite-host cell association was observed. The action of PLD could be blocked by the presence of a competitive substrate, phosphatidylethanolamine, during enzyme treatment. The enhancing effect of PLD treatment of the parasites was relatively long lasting since it was still seen 3 h after the enzyme had been removed. The enhancing effect of PLD probably reflected an increased capacity of T. cruzi to associate with host cells rather than increased phagocytosis of PLD-altered parasites by macrophages since similar results were obtained when rat heart myoblasts, which are not phagocytic, were used as host cells. Neither the presence of phospholipids or PLD phospholipid cleavage products during the incubation of T. cruzi with macrophages had any effect on parasite-host cell association. These results show that PLD-mediated alterations to parasite phospholipids increase parasite-host cell association, and suggest that these phospholipids play a role in the initial stages of host cell infection by T. cruzi.

Animals↗

Parasitism, host immune function, and sexual selection.

Parasite-mediated sexual selection may arise as a consequence of 1) females avoiding mates with directly transmitted parasites, 2) females choosing less-parasitized males that provide parental care of superior quality, or 3) females choosing males with few parasites in order to obtain genes for parasite resistance in their offspring. Studies of specific host-parasite systems and comparative analyses have revealed both supportive and conflicting evidence for these hypotheses. A meta-analysis of the available evidence revealed a negative relationship between parasite load and the expression of male secondary sexual characters. Experimental studies yielded more strongly negative relationships than observations did, and the relationships were more strongly negative for ectoparasites than for endoparasites. There was no significant difference in the magnitude of the negative effect for species with and without male parental care, or between behavioral and morphological secondary sexual characters. There was a significant difference between studies based on host immune function and those based on parasite loads, with stronger effects for measures of immune function, suggesting that the many negative results from previous analyses of parasite-mediated sexual selection may be explained because relatively benign parasites were studied. The multivariate analyses demonstrating strong effect sizes of immune function in relation to the expression of secondary sexual characters, and for species with male parental care as compared to those without, suggest that parasite resistance may be a general determinant of parasite-mediated sexual selection.

Animals↗

Water flows in the parasitic association Rhinanthus minor/Hordeum vulgare.

Using the facultative root hemiparasite Rhinanthus minor and its host Hordeum vulgare several aspects of water relations have been measured in this parasitic association. Extraction of xylem sap by the parasite from the host's roots is facilitated by con siderably higher transpiration per leaf area in the parasite than in the host and by the fact that stomata of attached Rhinanthus were open all day and night despite extremely high ABA concentrations in the leaves. By comparison, another root hemiparasite, Melampyrum arvense, parasitizing various grasses in the field, showed normal diurnal stomatal behaviour. The abnormal behaviour of Rhinanthus stomata was not due to anatomical reasons as closure could be induced by applying high external ABA concentrations. Remarkable differences have been detected between the hydraulic conductance of barley seminal roots showing relatively low values and that of Rhinanthus seminal roots showing very high values. The latter could be related to the observed high ABA concentrations in these roots. Whole plant water uptake, transpirational losses, growth-dependent deposition, and the flows of water within the plants have been measured in singly growing Rhinanthus and Hordeum plants and in the parasitic association between the two. Water uptake, deposition and transpiration in Rhinanthus were dramatically increased after attachment to the barley host; most of the water used by the parasite was extracted as xylem sap from the host, thereby scavenging 20% of the total water taken up by the host's roots. This water uptake by the parasitized host, however, due to a parasite-induced reduction in the host's growth, was decreased by 22% as compared to non-parasitized barley. The overall changes in growth-related water deposition in the host and parasite pointed to decreased shoot growth and relatively favoured root growth in the host and to strongly favoured shoot growth in the parasite. These changes in the host became more severe, when more than one Rhinanthus was parasitizing one barley plant.

Abscisic Acid↗

Prevalence of parasitic infections among Thai patients at the King Chulalongkorn Memorial Hospital, Bangkok, Thailand.

Parasitic diseases are still considered to be a major public health problem. Most patients with parasitic infections are asymptomatic and therefore remain undetected. Asymptomatic parasitic infections are usually discovered by routine parasite examination. To determine the result of parasite examination at the Parasitology Unit, Out Patient Department, King Chulalongkorn Memorial Hospital, Bangkok, Thailand, the authors collected the data of individuals examined for parasite infections from June to December 1997. A total of 6,231 Thais provided the data for analysis. Evidence of parasitic infections was found in 557 (8.94%) cases. The disease was most prevalent in males (57.3%), and in the age group >15-30 years old (11.13%). The population from the Northeast of Thailand was found to harbor parasites with the highest prevalence rate (17.03%), while it was 11.90 per cent in the northern group. The parasitic prevalence rates in the West, East, South and Central regions were 10.60 per cent, 8.90 per cent, 7.74 per cent, and 4.92 per cent, respectively. The parasite most commonly identified was Strongyloides stercoralis (33.39%), while giardiasis was the most common protozoan infection (14.36%). The highest infection rates of S. stercoralis, hookworms, Opisthorchis viverrini, and Gnathostoma spinigerum were found in northeasterners. People from the North of Thailand were infected mostly with G. lamblia. People of working-age from northeastern as well as northern regions harbored pathogenic parasites with high prevalence rates. To prevent parasitic infections, health education for these high risk groups should be provided.

Adolescent↗

Unique characteristics of local responses in host resistance to mucosal parasitic infections.

Because of the tremendous impact that parasitic infections have on the health and productivity of humans and domestic animals, considerable research effort has been focused upon understanding the mechanisms of host-parasite coexistence, host resistance and immunopathology. Studies have employed a range of approaches including: kinetic analysis of parasite establishment, development, fecundity and survival in naive and previously-infected hosts; correlation between parasite survival and histopathologic responses at the site of infection; vaccination with attenuated parasites or their products; cellular and serum transfer of immunity to naive or immunocompromised hosts; pharmacologic manipulation of potential mediators of host defense using agonistic and antagonistic drugs. However, it is becoming increasingly clear that to understand the mechanisms associated with host resistance and parasite survival, one must define the characteristics of the local microenvironment at the host-parasite interface. One of the approaches by which such studies can be made involves the isolation and characterization of cells derived from the local infection site. This manuscript reviews some of these studies on local aspects of mucosal immune responses in parasitic infections. Examples that will be discussed include IgA antibody, intraepithelial leukocytes from the intestine, intestinal mast cell populations, macrophages derived from bronchoalveolar lavage, and local immunoregulatory responses during respiratory and intestinal parasitic infection. These studies have established unequivocally that local responses to mucosal parasitic infection can only be appropriately investigated using cells derived from the specific microenvironment. This conclusion should encourage others to further study these local responses and to be innovative in investigating unexplored aspects of the host-parasite interface.

Animals↗

Parasites and supernormal manipulation.

Social parasites may exploit their hosts by mimicking other organisms that the hosts normally benefit from investing in or responding to in some other way. Some parasites exaggerate key characters of the organisms they mimic, possibly in order to increase the response from the hosts. The huge gape and extreme begging intensity of the parasitic common cuckoo chick (Cuculus canorus) may be an example. In this paper, the evolutionary stability of manipulating hosts through exaggerated signals is analysed using game theory. Our model indicates that a parasite's signal intensity must be below a certain threshold in order to ensure acceptance and that this threshold depends directly on the rate of parasitism. The only evolutionarily stable strategy (ESS) combination is when hosts accept all signallers and parasites signal at their optimal signal intensity, which must be below the threshold. Supernormal manipulation by parasites is only evolutionarily stable under sufficiently low rates of parasitism. If the conditions for the ESS combination are not satisfied, rejector hosts can invade using signal intensity as a cue for identifying parasites. These qualitative predictions are discussed with respect to empirical evidence from parasitic mimicry systems that have been suggested to involve supernormal signalling, including evicting avian brood parasites and insect-mimicking Ophrys orchids.

Adaptation, Physiological↗

The effect of parasites on wildlife.

Populations of animals which live in the wild are regulated by many biotic and abiotic factors. Parasites are one of the biotic factors. Parasites may influence their hosts in different ways. They may cause the death of the host due to a direct lethal effect or an indirect effect. Direct lethal effects may occur if killing is a part of the life cycle of the parasite or if hosts and parasites have not developed an equilibrium. The introduction of hosts or parasites into a new environment with suitable hosts or parasites is an example. Death by parasitism may also be caused by a combination of the emaciating effects of parasites combined with factors such as bad weather conditions, environmental pollution or human handling. Parasites may also influence the behaviour of their hosts. If the hosts are intermediate hosts in the life cycle of the parasites, the alterations in behaviour may make them an easier prey for their predators, the final hosts. Parasites may also influence the reproductive success of the hosts. In this respect the relationship between the red grouse (Lagopus lagopus scoticus) and the caecal nematode Trichostrongylus tenuis has been well worked out.

Animals↗