[Study under certain African conditions of the antiparasitic action of thiabendazole on various helminths of domestic animals. I. Helminths of the zebu].
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Helminth identification and egg counts were performed in 316 fecal samples during 4 years in 199 golden lion tamarins (GLTs), Leontopithecus rosalia, from two Brazilian conservation units. Tamarin sex and age, area of occurrence and helminth co-infection were tested as potential factors that could affect helminth prevalence and egg shedding in host individuals. Three nematodes species were found at low prevalences but not in both conservation units: Ancylostomatidae (12%), Ascarididae (1%) and Tripanoxyuris minutus, an Oxyuridae (4%). Three other species had high prevalences and were found in both conservation units: one acanthocephalan, Oncicola sp. (30%), and two nematodes, Spiruridae (24%), and Trichostrongylidae (31%). These three latter helminths had distinct prevalences between the conservation units, probably due to differences in availability of helminth infective stages in each area. Prevalences were greater in females for all helminth species; this was especially the case for Oncicola sp. Sex and age differences in helminth prevalence may be associated with changes in sexual steroid levels that accompany age and reproductive status. Frequency of simultaneous infections by the two helminths considered most pathogenic (Oncicola and Trichostrongylidae) were statistically lower than expected; this may be related to: (i) higher tamarin death rate caused by the association; (ii) differences in exposure to helminth infective stages in the various areas of occurrence; (iii) and competition between these helminth species. All helminth species followed a negative binomial distribution, with stronger clumping occurring in Trichostrongylidae and in female GLTs. Distinct transmission strategies of Oncicola sp. and Spiruridae, in contrast with Trichostrongylidae, may partially explain the different clumping levels of these helminths in GLTs. Mean fecal egg counts of all helminths were not different between GLT sexes, ages or areas of occurrence. The three most common helminth species may be a threat to isolated or dense GLT populations.
Parasitic infections are a major theme in the "hygiene hypothesis", as allergies and autoimmune diseases are less prevalent in countries with higher burdens of helminths and other parasitic organisms. Helminths"-the grouping of multicellular worm parasites including nematodes, cestodes and trematodes-tend to establish long-lived, chronic infections indicating successful down-modulation of the host immune system. In this review, we describe the intricate immunology of host-helminth interactions and how parasites manipulate immune responses to enhance their survival. In so doing, they often minimise immunopathology and, it is suggested, reduce host susceptibility to, and severity of allergic and autoimmune diseases. Studies on helminth-infected communities and individuals support the hypothesis that an immuno-regulatory network promoted by parasites extends its influence to limiting allergies. Experimental models are now probing more deeply into the area of immune modulation by helminths, and we discuss the likely mechanisms by which helminths could be establishing a strongly regulatory environment. Understanding and harnessing the modulatory capacity of helminths may uncover novel therapeutic interventions, mimicking and exploiting their evolution for our benefit. Parasitic infections are a major theme in the "hygiene hypothesis", as allergies and autoimmune diseases are less prevalent in countries with higher burdens of helminths and other parasitic organisms. Helminths"-the grouping of multicellular worm parasites including nematodes, cestodes and trematodes-tend to establish long-lived, chronic infections indicating successful down-modulation of the host immune system. In this review, we describe the intricate immunology of host-helminth interactions and how parasites manipulate immune responses to enhance their survival. In so doing, they often minimise immunopathology and, it is suggested, reduce host susceptibility to, and severity of allergic and autoimmune diseases. Studies on helminth-infected communities and individuals support the hypothesis that an immuno-regulatory network promoted by parasites extends its influence to limiting allergies. Experimental models are now probing more deeply into the area of immune modulation by helminths, and we discuss the likely mechanisms by which helminths could be establishing a strongly regulatory environment. Understanding and harnessing the modulatory capacity of helminths may uncover novel therapeutic interventions, mimicking and exploiting their evolution for our benefit.
This paper analyses the association between infection with helminth parasites, the elevated production of IgE and the expression of allergies. Interpretations of this interaction have taken place in a scientific environment whose most secure element is the immunochemistry of allergic reactions resulting in a substantial body of literature that has sought a biological role for allergic reactivity in protective immunity directed against helminth parasites. While the association between helminth infections and elevated levels of IgE, mast cells and eosinophils is well established, a functional role for allergic reactions in protection against helminths has eluded experimental proof. Instead of this hypothesis, it is proposed that allergic reactivity is rarely present in helminth-infected individuals because allergic reactions do not function to regulate helminth infections. Data from many sources are used to establish that the 'normal' state of all mammals is to be infected with helminth parasites from shortly after birth until well into adulthood. Only in the last 100 years or so have people living in areas of high development with sophisticated water and sewage systems been able to escape helminth infection. Allergies are as conspicuously present in these human populations as they are absent in populations that are still regularly exposed to helminths. Furthermore, in populations with endemic helminthoses there is little overt expression of allergic pathology that could be connected to the acquisition or elimination of helminth parasites. Based on these observations, it is suggested that endemic helminthoses activate the Th2 system, particularly at mucosal surfaces, to provide a different level of immunological homeostasis than currently occurs in developed societies. Under these conditions, mast cells, eosinophils and IgE rarely participate in reactions that we would recognize as 'allergic', although their participation in the control of helminth infections is still envisaged. Allergic reactions are considered to be a purely pathologic consequence of the disruption of this homeostatic mechanism and are not protective at all for the individual expressing them. This interpretation is derived from the immunobiology of the host-parasite interaction rather than the biology of allergies and should lead to new concepts regarding both allergic disease and the role of helminth infections in human and animal populations.
Knowledge about the source, metabolism and functions of cobalamins and folates in helminths remains fragmentary. It is likely that all helminths, whether free-living or parasitic, cannot synthesize cobalamins and folates de novo. Folates, but not cobalamins, appear to be ubiquitous in helminths. Of the parasitic helminths that take up free cobalamins in vitro, all but one species showed no uptake of cobalamin bound to transport proteins, although the latter type of cobalamin by far predominates in vivo. Certain free-living and parasitic helminths in vitro and in vivo took up a variety of folyl and antifolyl monoglutamates, but it is not known whether helminths can take up any folyl polyglutamates. Helminths that have any folate-dependent metabolism appear able to produce polyglutamylated forms of the required tetrahydrofolate coenzymes. Helminths that possess a functional cobalamin-dependent pathway from succinyl CoA to propionyl CoA appear able to form the required adenosylcobalamin coenzyme. Only free-living helminths may possess a cobalamin (and 5-methyltetrahydrofolate)-dependent pathway from homocysteine to methionine. It is likely that all helminths possess the 5,10-methylenetetrahydrofolate-dependent pathway from deoxyuridylate to thymidylate. All helminths appear able to salvage purine bases and nucleosides, but 10-formyltetrahydrofolate-dependent de novo purine ribonucleotide synthesis has been demonstrated unequivocally only in nematodes. The primitive parasitic groups of helminths exhibit cobalamin metabolism, whereas the more highly evolved ones seem to have lost the mechanisms for uptake and the associated biochemical pathways utilizing cobalamins.
Pathogens frequently use vectors to facilitate transmission between hosts and, for vertebrate hosts, the vectors are typically ectoparasitic arthropods. However, other parasites that are intimately associated with their hosts may also be ideal candidate vectors; namely the parasitic helminths. Here, we present empirical evidence that helminth vectoring of pathogens occurs in a range of vertebrate systems by a variety of helminth taxa. Using a novel theoretical framework we explore the dynamics of helminth vectoring and determine which host-helminth-pathogen characteristics may favour the evolution of helminth vectoring. We use two theoretical models: the first is a population dynamic model amalgamated from standard macro- and microparasite models, which serves as a framework for investigation of within-host interactions between co-infecting pathogens and helminths. The second is an evolutionary model, which we use to predict the ecological conditions under which we would expect helminth vectoring to evolve. We show that, like arthropod vectors, helminth vectors increase pathogen fitness. However, unlike arthropod vectors, helminth vectoring increases the pathogenic impact on the host and may allow the evolution of high pathogen virulence. We show that concomitant infection of a host with a helminth and pathogen are not necessarily independent of one another, due to helminth vectoring of microparasites, with profound consequences for pathogen persistence and the impact of disease on the host population.
The helminth community of the wood mouse, Apodemus sylvaticus, in the Sierra Espuna was characterized after a complete analysis of its helminth community component and infracommunity structure relative to host age, sex and year of capture. The helminth community comprised 13 species: one trematode, four cestodes and eight nematodes. The cestode Pseudocatenotaenia matovi and the nematode Syphacia frederici were the most prevalent and abundant helminth species, respectively. Sixty four percent of mice analysed presented helminths with a direct cycle and 42% presented helminths with an indirect cycle. The helminth community presents a low diversity with infracommunities usually made up of only one or two helminth species. Host age and year of capture seem to play a major role in determining species richness and helminth diversity, but not in determining the abundance of helminths. Host sex does not seem to affect the infection rate nor the diversity. Further studies on more samples of wood mice and other small mammal species in this regional park are needed to explore any possible interactions between helminth communities in the host populations.
It has been proposed that helminth infection may exacerbate HIV progression by promoting activation of 'type 2' immune responses. To examine this hypothesis, we investigated helminth infection in a cohort of HIV-1-seropositive adults in Entebbe, Uganda, during November 1999 to January 2000. Individuals with helminths were treated. At enroLlment, after 5 weeks and after 4 months, CD4+ and CD8+ T cell counts and viral load were measured. Cytokine responses (interferon [IFN]-gamma, interleukin [IL]-2, IL-4 and IL-5) to Schistosoma mansoni adult worm antigen (SWA), Mycobacterium tuberculosis culture filtrate proteins (CFPs) and phytohaemagglutinin (PHA) were measured in a whole blood assay. At baseline, CD4+ T cell counts and CD4+: CD8+ ratios were higher in individuals with helminths than in those without (median CD4+ T cell counts 467/microL and 268/microL, respectively, P = 0.005). Viral load was lower among those with helminths but this was not statistically significant. During follow-up, CD4+ T cell counts and cytokine responses to PHA fell among individuals without helminths. Among those treated for helminths, CD4+ counts remained stable. Viral loads showed a transient increase at 5 weeks, which was more marked among those treated for helminths, but the levels at 4 months were similar to baseline in both groups. Among those with schistosomiasis, IFN-gamma and IL-2 responses to CFP, and IL-2 and IL-4 responses to PHA declined but there was a sustained increase in cytokine responses to SWA following treatment. These data do not support the hypothesis that helminth infection exacerbates HIV infection. The possibility that chronic helminth infection may suppress HIV replication and that effects on HIV replication may vary during helminth infection and treatment should be considered.
BACKGROUND: Infection with intestinal helminths may stimulate dysfunctional immune responses in human immunodeficiency virus (HIV)-infected persons. Studies have yielded conflicting results regarding the impact of antihelminthic treatment on plasma concentrations of HIV-1 RNA.Methods. We conducted a prospective study of 54 HIV-1- and helminth-coinfected and 57 HIV-1-infected, helminth-uninfected asymptomatic adults living in Lusaka, Zambia, to assess the impact of antihelminthic treatment on plasma concentrations of HIV-1 RNA. RESULTS: Median baseline viral load was 0.33 log(10) copies/mL lower in the helminth-infected group than in the uninfected group. Mean viral load between pretreatment and posttreatment visits increased in the helminth-infected (mean, 4.23 vs. 4.29 log(10) copies/mL; P=.6) and helminth-uninfected (mean, 4.39 vs. 4.52 log(10) copies/mL; P=.2) groups. Helminth-infected participants with high pretreatment viral loads had a mean 0.25-log(10) copies/mL decrease after treatment (P=.3), and helminth-uninfected participants had a mean 0.02-log(10) copies/mL decrease (P=.8). CONCLUSIONS: We did not find an overall association between treatment of intestinal helminth infections and reduction in viral load in coinfected adults. Future studies may need to focus on adults with intense helminth infections who live in rural areas or on adults or children who harbor higher helminth burdens and plasma concentrations of HIV-1 RNA.
A total of 239 green frogs Rana clamitans, collected between June 3 and August 27, 1998 from 6 locations in southwestern Michigan, was examined for helminths. Of the 26 helminth taxa found, the larval cestode Mesocestoides sp. had the highest mean intensity, followed by the larval trematode Fibricola sp. Of the helminths that mature in frogs, Haematoloechus varioplexus had the highest prevalence and Gorgodera amplicava had the highest mean intensity. Frogs from 118th Pond had the highest species richness (20), mean helminth species richness (5.2), and mean helminth abundance (153.7). Frogs from Constantine East had the highest mean helminth species diversity (0.8778) and evenness (0.6033), followed by frogs from 118th Pond. In all comparisons of mean helminth community species richness, abundance, diversity, and evenness, adult frogs had significantly higher or higher values than did juveniles at each location. Jaccard's coefficients of similarity for the helminth communities for location pairs ranged from 0.545 to 0.823. Nine and 2 core helminth taxa occurred at the local and regional levels, respectively. The differences in several helminth community measures in green frogs among locations stress the importance of local ecological conditions on helminth community structure.
One-hundred twelve amphibians, including 51 blue-spotted salamanders, Ambystoma laterale, 30 eastern American toads, Bufo americanus americanus, and 31 northern leopard frogs, Rana pipiens, were collected during April-October 1996 from Waukesha County, Wisconsin and examined for helminth parasites. The helminth compound community of this amphibian assemblage consisted of at least 10 species: 9 in American toads, 8 in leopard frogs, and 3 in blue-spotted salamanders. American toads shared 7 species with leopard frogs, and 2 species occurred in all 3 host species. Although there was a high degree of helminth species overlap among these sympatric amphibians, statistically significant differences were found among host species and percent of indirect or direct-life cycle parasites of amphibian species individual component communities (chi2 = 1,015, P < 0.001). American toads had a higher relative abundance of nematodes, 59%, than larval cestodes, 31%, and larval and adult trematodes, 10%, whereas leopard frogs had a higher relative abundance of larval cestodes, 71.3%, and larval and adult trematodes, 25.3%, than nematodes 3.4%. This is related to ecological differences in habitat and dietary preferences between these 2 anuran species. Helminth communities of blue-spotted salamanders were depauperate and were dominated by larval trematodes, 94%, and few nematodes, 6%. Low helminth species richness in this host species is related to this salamander's relatively small host body size, smaller gape size, lower vagility, and more fossorial habitat preference than the other 2 anuran species. Adult leopard frogs and toads had significantly higher mean helminth species richness than metamorphs, but there was no significant difference in mean helminth species richness among adult and metamorph blue-spotted salamanders. Considering adult helminths, the low species richness and low vagility of caudatans as compared with anurans suggest that local factors may be more important in structuring caudatan helminth communities of salamanders than of anuran hosts. Helminth species infecting salamanders may be more clumped in their geographic distribution as compared with anurans, and the role of other hosts and their parasites at the compound community level may be important in structuring helminth communities of salamanders.
The helminths of the alimentary tract of wood mice, Apodemus sylvaticus, were studied at two sites over a 33-month period. Nine helminth species were recovered regularly. All but one was absent for at least 1 month. Monthly samples from these helminth communities were more similar to samples taken in the following month than to samples taken at progressively greater intervals up to a year. Helminth communities at both sites, however, had cyclical elements. Comparison of mean similarity indices for helminth faunas from 33 monthly samples and those from data sets generated by four null models suggests that observed values did not differ from a model where relative abundance of each species was determined randomly with specific maxima of abundance and species absences based on observed data. Loss of helminth species may decrease measurements of community stability based on relative abundance while persistence of abundant species increases stability. Helminth faunas in samples of A. sylvaticus from six localities taken at the same time of year at 5-year intervals indicated that some changed radically while others remained virtually unchanged. The considerable variation in helminth communities from different localities was not related to proximity or gross habitat characteristics. Nematospiroides dubius, Corrigia vitta and Capillaria murissylvatici were important in discriminating between the parasite faunas at different sites. Abundance of Syphacia stroma varied considerably between spatial surveys reflecting differences in host population dynamics in the 2 years. The present report and data from elsewhere in Ireland suggest that species composition of the helminths associated with A. sylvaticus may be stable over a wider geographical scale. There were neither strong nor consistent positive or negative interactions between pairs of helminth species. It is concluded that the stability characteristics of this parasite community, in terms of species composition and relative abundance, are the product of the population biology of independent parasite species rather than interspecific interactions. Variation in the role of competition in parasite communities is discussed.
Helminth parasites are the cause of very significant morbidity, mortality and economic losses in man and domestic animals. Most parasitic helminths infect their hosts via the oral route, and live either at the mucosal surface of the gastro-intestinal tract (GIT), or cross this mucosal barrier on their way to predilection sites. Many helminths live at mucosal surfaces, typically the gut or respiratory tract, and some cross these barriers, either temporarily, spending a period of time in the mucosa before returning to the mucosal surface, or to access other tissues and sites in the host. Typically, helminths induce strongly polarised Th2 responses, which are often effective in mediating protective immunity against those parasites living at mucosal surfaces, but less so in protecting against tissue-dwelling parasites. Induction of strongly-polarised Th2 responses may impair the ability of parasites hosts to eliminate other pathogens. Control of helminth infections relies largely on chemotherapy, together with management and environmental measures designed to keep hosts away from infective stages. Drug resistance has become a significant problem in some helminth populations, and this has promoted interest in the development of immunoprophylactic strategies. However, despite intensive research efforts, helminth vaccines have not become part of regular control strategies. In addition to the considerable technical difficulties posed in the production of vaccines against these complex organisms, further difficulties in securing acceptance for anti-helminth vaccine by regulatory authorities and by users, will be encountered. Such vaccines need not result in sterile immunity, as is required of anti-bacterial and anti-viral vaccines. Recent evidence indicates that while helminths are responsible for disease, immunopathology and impairment of immunity to other pathogens, a complete absence of helminth infection during early life may be a predisposing factor for the development of auto-immune pathology.
Anthelmintic treatment of sick preschool-age children at health facilities is a potentially effective strategy for intestinal helminth control in this age-group. We conducted a study from July 1998 to February 1999 in western Kenya to determine whether the Integrated Management of Childhood Illness (IMCI) guidelines' clinical assessment can be used to identify helminth-infected children, and to evaluate the nutritional benefit of treating sick children without pallor with an anthelmintic (mebendazole is already part of IMCI treatment for sick children aged 2-4 years with palmar pallor in areas where hookworm and Trichuris trichiura infections are endemic). Sick children aged 2-4 years seen at 3 rural health facilities were clinically evaluated and tested for haemoglobin concentration, malaria parasites, and intestinal helminths. Children without pallor were randomly assigned to receive a single dose of 500 mg of mebendazole or a placebo and re-examined 6 months later. Among the 574 children enrolled, 11% had one or more intestinal helminths. Most infections were of light intensity. Selected clinical signs and symptoms available from the IMCI assessment, including palmar pallor and low weight-for-age, were not associated with helminth infection. Six months after enrollment, no differences in growth of children without pallor were observed between the mebendazole (n = 166) and placebo (n = 181) groups. However, there was a significantly greater mean increase in weight, height, and weight-for-age Z score among the helminth-infected children in the mebendazole group (n = 22) as compared with helminth-infected children in the placebo group (n = 20). We conclude that even lightly infected preschool-age children without palmar pallor benefit from anthelmintic treatment; however, in this study setting of low helminth prevalence and intensity, helminth-infected children could not be identified using the IMCI guidelines. Cost-effectiveness studies are needed to help define helminth prevalence thresholds for routine anthelmintic treatment of sick preschool-age children seen at first-level health facilities.
Human T cell lymphotropic virus type 1 (HTLV-1) infection is associated with an exacerbated type 1 immune response and secretion of high levels of proinflammatory cytokines. In contrast, helminthic infection induces a type 2 immune response. In the present study, the cytokine profile in HTLV-1 carriers coinfected with helminths (Strongyloides stercoralis and/or Schistosoma mansoni) was compared with that in HTLV-1 carriers not coinfected with helminths. Levels of interferon (IFN)- gamma were higher in HTLV-1 carriers not coinfected with helminths than in HTLV-1 carriers coinfected with helminths (P<.05). The overall frequency of IFN- gamma -expressing CD8+ and CD4+ cells was decreased in HTLV-1 carriers coinfected with helminths (P<.05). The percentage of interleukin (IL)-5- and IL-10-expressing T cells in HTLV-1 carriers coinfected with helminths was higher than that in HTLV-1 carriers not coinfected with helminths (P<.05). Moreover, we found that the prevalence of helminthic infection was 7-fold higher in HTLV-1 carriers than in patients with HTLV-1-associated myelopathy/tropical spastic paraparesis (P<.05). These data show that helminthic infection decreases activation of type 1 cells, which may influence the clinical outcome of HTLV-1 infection.
The detection and identification of intestinal helminths were studied retrospectively by comparing the Formalin concentration technique with the trichrome-stained smear technique. A total of 3,997 stool samples from 1,570 patients were examined by both methods. Of the 3,997 samples, 31% (1,239 of 3,997) contained helminths or protozoans or both. A total of 11% (434 of 3,997) of the samples representing 14% (221 of 1,570) of the patients were positive for one or more helminth species. A total of 570 separate identifications of helminth ova/larvae were made. Among the helminth ova/larvae identified. 14.6% (83 of 570) were detected only in the trichrome-stained smear, representing 6.3% (14 of 221) of the patients. From these data, it can be concluded that unless a diligent search of the stained smear for helminths is made, a significant number of helminth infections may be missed. It is, therefore, recommended that stained stool smears be used to aid the detection of helminth ova/larvae in conjunction with Formalin concentration. The appearance of the most common helminth ova/larvae in trichrome-stained smears is described, along with specific characteristics that may be used for their identification. In addition, the advantages and disadvantages of the trichrome staining technique for helminth identification are discussed.
From the above discussion it is quite obvious that the bioenergetics in helminths are different in many ways from those found in higher organisms. All adult helminths appear to be able to consume oxygen when it is available but none of them can use it to drive the pathways of complete substrate degradation, like typical aerobic organisms, as a major strategy for energy generation. These properties hold also true for those worms residing in a highly aerobic environment, such as the blood stream or the muscle and lung tissues. Although in a number of recent studies oxygen was found to play apparently a greater role in the bioenergetics of adult helminths than originally thought, energy-generating mechanisms in adult worms seem to place greater emphasis on fermentations and anaerobic electron transport processes. These exhibit relatively low energy conservation efficiencies and result in the formation of a variety of organic end products, most of which must be excreted. The obvious correlation between the type of bioenergetic strategy operative in a particular helminth species and its environmental conditions is not well understood. The increased capacity to generate chemical energy and key metabolites of helminths possessing multiple fermentations and anaerobic respirations may give the organism greater versatility and metabolic flexibility to respond to the environmental changes observed in its corresponding habitat. Other helminths, such as schistosomes and filariids, which have continuous access to a fairly constant nutrient supply, were found to depend primarily on the more inefficiently functioning and primitive strategy of glycolysis for energy production. The reason for the occurrence in helminths of limited oxidative capacities is not completely clear. It may be assumed that the variety of alternative anaerobic pathways have evolved in response to the lack of a circulatory system and/or to the specific, often peculiar, environmental conditions prevailing in most parasitic habitats. An alternative idea put forward by Barrett [8] is that helminth metabolism represents a form of biochemical economy. Most endoparasites have an abundant supply of food and swim as if in a land of Cockain, obviously without any need to extract a maximum amount of chemical energy from the nutrients they take up. On the other hand, the fact that free-living and other larval or juvenile stages of helminths often have a typical aerobic bioenergetic pattern is a clear indication that the DNA of these organisms carries the genetic message for all the enzymes involved in complete substrate degradation.(ABSTRACT TRUNCATED AT 400 WORDS)
Larval helminths of different species that share the same intermediate host and are transmitted by predation to the same definitive host may cooperate in their attempts to manipulate the behaviour of the intermediate host, while at the same time having conflicts of interests over the use of host resources. A few studies have indicated that intermediate hosts harbouring larval helminths have altered concentrations of neurotransmitters in their nervous system, and thus measuring levels of neurotransmitters in host brains could serve to assess the respective and combined effect of different helminth species on host behaviour. Here, we investigate potential cooperation and conflict among three helminths in two species of crab intermediate hosts. The acanthocephalan Profilicollis spp., the trematode Maritrema sp. and an acuariid nematode, all use Macrophthalmus hirtipes (Ocypodidae) as intermediate host, whereas Profilicollis and Maritrema also use Hemigrapsus crenulatus (Grapsidae). All three helminths mature inside gulls or other shore birds. There was a significant decrease in the mean volume of Profilicollis cystacanths as the intensity of infection by this parasite increased in H. crenulatus, the only host in which this was investigated; however, there was no measurable effect of other helminth species on the size of acanthocephalans, suggesting no interspecific conflict over resource use within crabs. There was, in contrast, evidence of a positive interspecific association between the two most common helminth species: numbers of Profilicollis and Maritrema were positively correlated among crabs, independently of crab size, in M. hirtipes but not H. crenulatus. More importantly, we found that the total number of larval helminths per crab correlated significantly, and negatively, with concentrations of serotonin in crab brains, again only in M. hirtipes; numbers of each parasite species separately did not covary in either crab species with serotonin or dopamine, the other neurotransmitter investigated in this study. The relationship with serotonin appears due mainly to numbers of Profilicollis and Maritrema and not to nematodes. This is the first demonstration of a potentially synergistic manipulation of host behaviour by different helminth species, one that appears host-specific; our results also point toward the neurobiological mechanism underlying this phenomenon.