There are two ways in which trematodes can be used in the control of other trematodes which transmit diseases of medical or veterinary importance. Either they can interfere with the reproductive capacity of the host species of snails, or they can exert an antagonistic effect against the larval stages of the target trematode species inside the snail. Often the two effects act together.

There are six essential criteria for the selection of a suitable species of snail for control: (1) complete and permanent sterilizing effect; (2) clear dominance over target trematodes; (3) strong infectivity to molluscs of all ages; (4) high egg productivity in the definitive host; (5) lack of pathogenicity towards man and domestic animals; (6) ease and low cost of maintenance of the life-cycle.

Various field trials are discussed and it is considered that trematode species producing rediae are usually dominant over those producing only sporocysts (such as the schistosomes). Most species used in control trials to date have belonged to the family Echinostomatidae as they have rediae, and eggs that can be produced in large numbers in a laboratory host such as the rat.

The mathematical basis and feasibility of control schemes are discussed and it is concluded that much more information is necessary before their potential can be evaluated.

The ‘Muguga cocktail’ live vaccine comprises three Theileria parva stocks (Muguga, Kiambu 5 and the buffalo-derived Serengeti-transformed) and has been used extensively in Eastern, Central and Southern Africa with an infection and treatment protocol to protect cattle against East Coast fever. We report the characterization of the three component vaccine stocks using a panel of polymorphic micro-satellite and mini-satellite markers and the development of a stock-derived PCR method that distinguishes two of the vaccine stocks. These markers, with the use of a recently developed Reverse Line Blot assay, have enabled us to address four important questions in relation to vaccination. First, how closely related are the vaccine stocks, secondly do all three stocks persist post-vaccination and induce a carrier state, thirdly is there evidence for the transmission of the vaccine stocks and fourthly does vaccination prevent infection with local genotypes? The results show that Muguga and Serengeti-transformed stocks are highly related but very distinct from Kiambu 5 that persists in vaccinated cattle establishing a carrier state. No evidence was obtained for the transmission of vaccine stocks to co-grazed animals, although these animals were infected with up to 8 different T. parva genotypes showing there was a significant level of tick challenge. Some of the vaccinated animals become infected with a subset of local genotypes providing evidence for limited vaccine ‘breakthrough’.

The cuticle of adults of Nippostrongylus brasiliensis has been described using histological, histochemical and ultrastructural techniques.

The cuticle has the following layers: an outer triple-layered membrane; a single cortical layer; a fluid-filled layer which is traversed by numerous collagen fibrils; struts which support the fourteen longitudinal ridges of the cuticle and which are suspended by collagen fibrils in the fluid-filled layer; two fibre layers, each layer apparently containing three layers of fibres; and a basement lamella.

The fluid-filled layer contains haemoglobin and esterase.

The muscles of the body wall are attached to either the basement lamella or to the fibre layers of the cuticle.

The mitochondria of the hypodermis are of normal appearance.

The longitudinal ridges of the cuticle appear to abrade the microvilli of the intestinal cells of the host.

Possible functions of the cuticle are discussed.

I wish to thank Dr P. Tate, in whose department this work was done, for helpful suggestions and criticism at all stages of this work, and Mr A. Page for technical assistance. I also wish to thank Professor Boyd for permission to use the electron microscope in the Department of Anatomy.

Infection with Schistosoma haematobium was studied in a rural community of approximately 500 persons in eastern Zimbabwe. The overall prevalence of infection, as determined by urine egg counts, was 40·1%, and of heavy infections (≥ 50 eggs/10 ml urine) was 11·0%. The prevalence of both heavy and all infections was highest in the 8 to 10-year-old age class. During 1987–88 data were obtained from 102 individuals on intensity of reinfection 14 weeks after treatment with praziquantel, the efficacy of treatment having been determined after 4 weeks. The water contact made by these individuals during 2-week periods immediately following treatment was recorded. The relative abundance of patent infected intermediate host snails, Bulinus globosus, was also monitored. An index of exposure was developed which weighted each water contact by its duration, the type of activity, the time of day, and the abundance of infected snails at the site used. The relationships between rates of reinfection, rates of exposure, and age were examined. Although only 13 individuals showed positive rates of reinfection, there were statistically significant and independent effects of both exposure and age on reinfection rate. Quantitative estimates of reinfection rates suggested that individuals aged 12 years or less acquired substantially more infection (measured as egg output) than individuals more than 12 years old.

Biodiversity loss may increase the risk of infectious disease in a phenomenon known as the dilution effect. Circumstances that increase the likelihood of disease dilution are: (i) when hosts vary in their competence, and (ii) when communities disassemble predictably, such that the least competent hosts are the most likely to go extinct. Despite the central role of competence in diversity–disease theory, we lack a clear understanding of the factors underlying competence, as well as the drivers and extent of its variation. Our perspective piece encourages a mechanistic understanding of competence and a deeper consideration of its role in diversity–disease relationships. We outline current evidence, emerging questions and future directions regarding the basis of competence, its definition and measurement, the roots of its variation and its role in the community ecology of infectious disease.

In the present study, we show for the first time that lipophosphoglycan (LPG) stimulated cytokine production by human peripheral blood mononuclear cells is also mediated via Toll-like receptor (TLR2). In addition, in order to verify if TLR2 is involved in recognition of the purified PGs, neutralizing mAbs against TLR2 and TLR4 were used to treat the cells before being stimulated with PGs. We found strong Th1-promoting cytokines induced by sLPG but not by mLPG which was blocked by presence of anti-TLR2 mAb. This finding reveals a mechanism by which the first encounter and recognition of L. major promastigotes by mLPG after interaction with TLR2 provides a cytokine milieu for consequent Th2 differentiation. Moreover, having shown the strong induction of Th1-promoting cytokines and low production of IL-10 in response to sLPG might have vaccine implication since it is recognized by TLR2 providing signals to professional antigen presenting cells that reside in the skin to promote effective T cell responses against Leishmania infection. In addition, it was shown that purified mLPG and sLPG activate reactive oxygen species (ROS) production which is also blocked by anti-TLR2 but not by anti-TLR4. However, no inhibition was seen in PPG-induced cytokine and ROS production in the presence of anti-TLR2 and anti-TLR4, implying involvement of other receptors.

A previous paper reported that the intake of Onchocerca volvulus microfilariae (mff) by different species of Simulium is essentially proportional to the parasite load in the skin of infected carriers. This paper examines the fate of the ingested mff in susceptible vectors to assess the relationship between parasite intake and infective larval output in blackfly species with and without well-developed cibarial armatures. Analysis is based on data from 3 onchocerciasis endemic areas: Guatemala (S. ochraceum s.l.), West Africa (S. damnosum s.l./S. sirbanum) and the Amazonian focus between South Venezuela and Northern Brazil (S. guianense and S. oyapockense s.l.). The data, which include published and unedited information collected in the field, record experimental studies of parasite uptake by wild flies maintained in captivity until the completion of the extrinsic incubation period. The relationship between L3 output (measured as the mean number of successful larvae/fly or, as the proportion of flies with infective larvae) and average microfilarial intake, was strongly non-linear. This non-linearity was best represented by a sigmoid function in case of armed simuliids (S. ochraceum s.l., S. oyapockense s.l.), or by a hyperbolic expression in that of unarmed flies (S. damnosum s.l., S. guianense). These results are compatible, respectively, with the patterns of ‘initial facilitation’ and ‘limitation’ described in culicid vectors of lymphatic filariases. A maximum mean number of 1–3 L3/fly was observed in all 4 vectors. It is concluded that O. volvulus larval development to the infective stage is regulated by density-dependent mechanisms acting at the early phase of microfilarial migration out of the blackfly's bloodmeal. Damage by the bucco-pharyngeal armature may also be density dependent. A hypothesis, based on this density dependence is forwarded to explain initial facilitation, so far only recorded in vectors with well-developed cibarial teeth. Our results provide quantitative support for the conjecture that chemotherapy alone is likely to have a greater impact on reducing onchocerciasis transmission in endemic areas where the main vector has a toothed fore-gut than in foci where the vectors have unarmed cibaria.

Plasmodium falciparum gametocytes grown in vitro were fed through membrane feeders to laboratory-reared Anopheles stephensi mosquitoes. Intact midguts, including entire bloodmeal contents, were removed between 24 and 48 h post-bloodfeeding. Giemsa-stained histological sections were prepared from the midguts and examined by light microscopy. Contrary to previous reports, ookinetes were clearly visible within midgut epithelial cells, demonstrating intracellular migration across the midgut wall. Ookinetes entered epithelial cells through the lateral apical membrane at sites where 3 adjacent cells converged. There was no evidence for the existence of a morphologically distinct group of epithelial cells preferentially invaded by ookinetes. However, ookinete penetration was associated with significant morphological changes to invaded cells, including differential staining, condensation and fragmentation of the nucleus, vacuolization, loss of microvilli and various degrees of extrusion into the midgut lumen. Epithelial cells completely separated from the midgut wall were found within the midgut lumen. These cells were associated with invading parasites suggesting that ookinete penetration resulted in complete ejection of invaded cells from the midgut wall. Small clusters of morphologically altered midgut cells and invading parasites spanning the membranes of adjacent abnormal epithelial cells were observed, consistent with intracellular movement of ookinetes between neighbouring midgut cells. Extruded epithelial cells were also observed rarely in uninfected midguts. Epithelial cell extrusion, therefore, may be a general mechanism of tissue repair through which damaged cells are removed from the midgut wall rather than a parasite-specific response. These observations demonstrate that human malaria parasite infection of mosquitoes is consistent with, and provides further support for, the Time Bomb model of ookinete invasion of the mosquito midgut epithelium previously proposed for rodent malaria parasites.

Infections with irradiated Schistosoma mansoni were established by intramuscular (i.m.) injection of mechanically transformed schistosomula. A dose of 2·3 krad. allowed persistence of a small proportion of worms to adulthood, and of these survivors the majority of the female worms were sexually sterile. However, a small proportion of 2·3 krad. irradiated females and a larger proportion of similarly irradiated males were capable of pairing successfully with non-irradiated partners. Radiation in the range 2·3 to 10 krad. resulted in slightly reduced peak recoveries from the lungs while 20 krad. resulted in a much reduced and 40 krad. a virtual absence of survival to the lung stage. Increasing doses of radiation in the range 2·3 to 10 krad. resulted in successively fewer parasites reaching the liver. Thus, the major sites of the radiation-induced mortality appeared to be as follows: 2·3 krad., mainly in the liver; 4 krad., in the lungs and liver; 10 krad., mainly in the lungs; 20 krad., at the injection site and in the lungs and 40 krad., mainly at the injection site. The infections studied here showed reduced survival following exposure to high doses of radiation compared with the infections, established as percutaneously applied cercariae, which have been reported by other workers. Possible reasons for the disparity are discussed.

Radio-immunoassay of plasma gastrin showed that hypergastrinaemia developed in sheep during experimental infections with Ostertagia circumcincta. Elevations of plasma gastrin occurred within 8 days of the first dose of infective larvae, with the most marked increase being after 11–20 days when adult worms would be expected to be present in the abomasum. Increases of plasma gastrin levels from 69·0 ± 7·6, 28·7 ± 5·3, 48·7 ± 5·9 and 60·6 ± 9·4 pg/ml before infection to maxima of 650, 230, 900 and 750 pg/ml respectively were recorded in 4 sheep infected for the first time. In 2 others which had been previously infected and then treated with anthelmintic, plasma gastrin rose from 16·0 ± 4·0 and 377 ± 87 pg/ml to maxima of 260 and 900 pg/ml at 24 and 29 days after re-infection, respectively. The cause of hypergastrinaemia has not been established. The elevation of abomasal pH which occurs in ostertagiasis may result in increased gastrin levels but is thought not to be the only cause since plasma gastrin increased before the abomasal pH rose and developed in sheep infected for a second time although their abomasal contents remained at pH 4·0 or lower. The presence of the parasite is critical for development of hypergastrinaemia as shown by return of the gastrin level to normal following therapy with an anthelmintic. It was shown that the parasites do not depend on the hypergastrinaemia since O. circumcincta became established in antrectomized sheep in which hypergastrinaemia did not develop.

Genetic studies have demonstrated profound differences between the ‘porcine’ genotype of Cryptosporidium parvum, versus ‘human’ and ‘bovine’ genotypes. The study analysed infectivity and pathogenicity of the ‘porcine’ genotype (CPP-13 isolate) of C. parvum, and compared the results with published data on the ‘bovine’ genotype (CPB-0 isolate). This was investigated in calves and piglets from commercial herds. Piglets were mildly affected by the CPP-13 isolate, contrary to piglets infected with the CPB-0 isolate, which caused diarrhoea of a mean duration of 3·5 days. CPP-13 produced no or very mild clinical signs in piglets despite the excretion of high numbers of oocysts. Concomitant infection with rotavirus, however, caused a dramatic aggravation of the clinical signs, and 5 of 6 experimentally infected piglets died. CPP-13 appeared to be adapted to porcine hosts as illustrated by the lack of infectivity to 1 experimentally inoculated calf, and the absence of clinical signs, the long pre-patent period (15 days), and the excretion of very low numbers of oocysts following experimental infection of another calf. Thus, in accordance with other molecular studies, our results support the genetic evidence for the existence of a new species of Cryptosporidium adapted to pigs.

Trypanosoma cruzi, the etiological agent of Chagas disease, is ingested by triatomines during their bloodmeal on an infected mammal. Aiming to investigate the development and differentiation of T. cruzi inside the intestinal tract of Rhodnius prolixus at the beginning of infection we fed insects with cultured epimastigotes and blood trypomastigotes from infected mice to determine the amount of recovered parasites after ingestion. Approximately 20% of the ingested parasites was found in the insect anterior midgut (AM) 3 h after feeding. Interestingly, a significant reduction (80%) in the numbers of trypomastigotes was observed after 24 h of infection suggesting that parasites were killed in the AM. Moreover, few parasites were found in that intestinal portion after 96 h of infection. The evaluation of the numbers of parasites in the posterior midgut (PM) at the same periods showed a reduced parasite load, indicating that parasites were not moving from the AM. Additionally, incubation of blood trypomastigotes with extracts from R. prolixus AMs revealed that components of this tissue could induce significant death of T. cruzi. Finally, we observed that differentiation from trypomastigotes to epimastigotes is not completed in the AM; instead we suggest that trypomastigotes change to intermediary forms before their migration to the PM, where differentiation to epimastigotes takes place. The present work clarifies controversial points concerning T. cruzi development in insect vector, showing that parasite suffers a drastic decrease in population size before epimastigonesis accomplishment in PM.

Dung beetles are detrivorous insects that feed on and reproduce in the fecal material of vertebrates. This dependency on vertebrate feces implies frequent contact between dung beetles and parasitic helminths with a fecal component to their life-cycle. Interactions between dung beetles and helminths carry both positive and negative consequences for successful parasite transmission, however to date there has been no systematic review of dung beetle-helminth interactions, their epidemiological importance, or their underlying mechanisms. Here we review the observational evidence of beetle biodiversity–helminth transmission relationships, propose five mechanisms by which dung beetles influence helminth survival and transmission, and highlight areas for future research. Efforts to understand how anthropogenic impacts on biodiversity may influence parasite transmission must include the development of detailed, mechanistic understanding of the multiple interactions between free-living and parasitic species within ecological communities. The dung beetle–helminth system may be a promising future model system with which to understand these complex relationships.

The freeze fracture technique has been used to quantify changes in the integral components of the double outer membrane of Schistosoma mansoni during the 6-week period of development within the mouse. The intramembraneous particle (IMP) density on the P1 face begins to rise Within 6 h of host penetration, reaches a maximum at day 4 and then falls rapidly after day 9, so that it is at a low level between 3 and 6 weeks. The E1 face IMP density follows the same course as that of the P1 face except that maximum particle density is recorded on day 1 and the counts begin to fall on day 5. The IMP density on the P1 face remains at a consistently low level throughout development. The E2 face IMP density rises gradually to a peak at day 4, when the parasites have migrated to the lungs, and remains thereafter at a similar level, so that by 6 weeks the E2 face has a higher IMP density than the other three fracture faces. The E2 face IMP show a marked increase in size on day 4. Morphological studies indicate that a different type of inclusion body makes a transient appearance in the tegument of the lung worms, and immunocytochemical techniques show the lung worms to be nonimmunogenic. It is suggested, therefore, that the E2 face IMP may represent complexes of parasite antigens and acquired host antigens. The tegumental membranes of cultured specimens have also been examined by freeze fracturing and the IMP densities compared with those obtained from in vivo parasites; the cultured schistosomula have a lower E2 face particle density than the in vivo specimens.

Salmo salar and Salmo trutta co-exist in coastal river systems in Europe and produce hybrids with little loss of viability or growth. This report describes the susceptibility of pure full-sibs of S. salar and S. trutta and their reciprocal half-sib hybrids to their respective gyrodactylids, Gyrodactylus salaris and Gyrodactylus derjavini. The pure-bred salmon and trout, and half-sib hybrids, were produced using eggs and sperm from wild anadromous S. salar (River Alta stock, North Norway) and wild anadromous S. trutta (River Fossbekk stock, Southwest Norway). Infections were initiated by exposing experimental fishes (0+) to S. salar naturally infected with G. salaris (River Lierelva strain) or S. trutta naturally infected with G. derjavini (River Sandvikselva strain). Fishes were then kept individually isolated under standardized conditions at 12 °C. Pure-bred S. salar were susceptible but frequently mounted a response to G. salaris without eliminating the infection, whereas pure-bred S. trutta were innately resistant to this species. Pure-bred S. trutta ranged from innately resistant to susceptible to G. derjavini but later most of the susceptible trout mounted a host response to G. derjavini. Pure-bred S. salar were also susceptible to this species, although parasite population growth rates were reduced and a host response frequently appeared eliminating G. derjavini. The abundance of both gyrodactylids was lower on the hybrids than on their respective pure-bred natural hosts, and a parental sire- and dam-influence on the resistance of hybrids was observed. When the sire was S. salar, the susceptibility of hybrids to G. salaris was similar to that of pure S. trutta; when the dam was S. salar both innately resistant, intermediately susceptible and responding individuals were present. In the case of G. derjavini, when the sire was S. trutta, infections on hybrids were similar to those on pure S. salar; when the dam was S. trutta, an increased level of susceptibility was observed. The present results provide evidence that: (1) Norwegian salmon stocks are variable in their susceptibility/resistance, with some fish able to control S. salaris infections; (2) trout stocks are innately resistant to G. salaris; (3) individual trout show a spectrum in susceptibility/resistance to G. derjavini, ranging from innate resistance through slightly susceptible to highly susceptible but with acquired resistance controlling infection; (4) although G. derjavini infections grow poorly on salmon, this host stock is susceptible to the parasite, but can limit infection by a host reaction; (5) susceptibility/resistance traits to gyrodactylids are genetically controlled and resistance can be transferred as a dominant trait through interspecific crosses between different salmonids; (6) interspecific hybrids between susceptible and resistant salmonids have a pattern of susceptibility to gyrodactylids intermediate to that of the parents; (7) resistance to gyrodactylids may be controlled by relatively few genes in salmonids; (8) epidemiologically, hybrids may act as a reservoir for gyrodactylids, may support a wider diversity of species than either parent and may disseminate gyrodactylids of both host species.

The importance of host cues to three species of steinernematid nematodes (Rhabdita: Steinernematidae) with different foraging strategies was compared. We presented host materials to nematodes in series to test responses to combinations of host cues. If a fixed hierarchy of cues is followed during foraging, parasites should respond most strongly to cues offered in natural order. Steinernema carpocapsae, an ambush forager, aggregated at the source of volatile host cues only after attachment to host cuticle. They also parasitized hosts more efficiently after contact with cuticle. Steinernema glaseri, a cruise forager, was unaffected by exposure to combinations of host cues. Steinernema feltiae, a nematode with characteristics of both ambushing and cruising, was affected by cue hierarchies when either contact or volatile cues were presented first. Host-associated materials encountered out of the context may not qualify as host cues for the ambush forager, S. carpocapsae. Perhaps the order in which cues are encountered is more predictable for ambushers than for cruisers. Therefore an ambusher's response to host materials has a more fixed contextual framework.

Melittobia is a chalcid, ectoparasitic upon a number of species of Hymenoptera and upon the pupae of certain flies. The insect was bred in the laboratory and the life-history is described in detail.

A number of experiments were made with regard to possible hosts and as to feeding and reproduction and certain points are of some interest.

For instance, the male apparently does not feed but the female uses her ovipositor with which she punctures the eggs, larvae or pupae of various insects, afterwards sucking the blood which oozes from the puncture. Such eggs, larvae or pupae, in spite of repeated punctures, may continue to develop.

Before beginning oviposition upon the surface of a suitable host, the female punctures the host one or more times with her ovipositor and apparently injects an anaesthetizing fluid, since insect larvae and pupae, upon which eggs have been laid, very rarely develop farther and yet remain fresh for a considerable time extending to many months.

With regard to reproduction, inbreeding appears to be normal, mating usually taking place between two individuals of the same brood and later between the female and one of her own offspring. Virgin females, unable to find a male, lay a few eggs, which always produce males, the first of which to emerge mates with the female.

The female is very prolific and may lay as many as twelve hundred eggs in two or three batches, mating taking place before each batch is laid. The last eggs of a batch are almost invariably male eggs, indicating that the female has exhausted her supply of spermatozoa.

A virgin female, whose eggs are removed so that she cannot rear a mate, may lay as many as ninety eggs during her life-time, her length of life being more than doubled, and such a female will survive through the winter, even withstanding frost, whereas fertilized females apparently all die in October or November at the latest.

All the eggs of a virgin female are apparently capable of developing and all such eggs produce males.

The recent development of genetic methods allows the delineation of species boundaries, especially in organisms where morphological characters are not reliable to differentiate species. However, few empirical studies have used these tools to delineate species among parasitic metazoans. Here we investigate the species boundaries of Clinostomum, a cosmopolitan trematode genus with complex life cycle. We sequenced a mitochondrial [cytochrome c oxidase subunit I (COI)] gene for multiple individuals (adults and metacercariae) from Middle-America. Bayesian phylogenetic analysis of the COI uncovered five reciprocally monophyletic clades. COI sequences were then explored using the Automatic Barcode Gap Discovery to identify putative species; this species delimitation method recognized six species. A subsample was sequenced for a nuclear gene (ITS1, 5·8S, ITS2), and a concatenated phylogenetic analysis was performed through Bayesian inference. The species delimitation of Middle-American Clinostomum was finally validated using a multispecies coalescent analysis (species tree). In total, five putative species are recognized among our samples. Mapping the second intermediate hosts (fish) onto the species tree suggests that metacercariae of these five species exhibit some level of host specificity towards their fish intermediate host (at the family level), irrespective of geographical distribution.

Analyses of small subunit ribosomal RNA gene sequences of representatives of major taxa of Monopisthocotylea were performed to identify the sister group of Gyrodactylus. Nuclear ribosomal DNA sequencesNucleotide sequence data reported in this paper are available in the GenBank, DDBJ and EMBL databases under the accession numbers AJ566375-79, AJ566768, AJ567670-74, AJ581657. from the complete internal transcribed spacer (ITS) region were used to infer phylogeny of 37 Gyrodactylus species and Gyrodactyloides bychowskii, Macrogyrodactylus polypteri and Gyrdicotylus gallieni, using maximum likelihood, parsimony and Bayesian inference. The genus Gyrodactylus appeared to be a monophyletic group in all analyses, based on the present data set. Within the genus, there were 3 major groups recognized by high bootstrap values and posterior probabilities. None of the 6 subgenera appeared to be monophyletic, and the most basal subgenus G. (Gyrodactylus) was paraphyletic. Characteristics of the excretory system of Gyrodactylus do not seem to be conservative enough to reveal subgenera within Gyrodactylus and we suggest abandoning existing subgenera as indicators of phylogeny. The grouping of species based on the morphology of the ventral bar and marginal hooks seems to have sufficient power to infer relationships between the Gyrodactylus species.

The small intestine of rats was divided into twenty sections in a reproducible manner in order to study the distribution of Nippostrongylus brasiliensis by stretching it under a tension of 5g in adrenaline saline.

A small but significant difference between the distribution of parasites in male and female rats was observed.

As larvae had virtually ceased to reach the intestine by the fifth day all changes in distribution after that day were due to movements of the established adult population.

Up to the twelfth day of a primary infestation the majority of the worms were found between the third and tenth sections, the population mode being in the fifth or sixth section. After the thirteenth day the number of worms in this region fell sharply.

The female worms had not ceased from egg-laying by the time most of the worms were being rejected.

The posterior half of the small intestine, i.e. the eleventh to twentieth sections, was not heavily parasitized, many of the worms seen being in passage to the anus.

The first section was not parasitized until the seventh day, but thereafter remained parasitized until long after worms had disappeared from the more posterior sections.

The relative number of male worms present increased as the infestation aged.

Throughout the experiment the relative number of male worms present at the anterior end of the smaller intestine was higher than that at the posterior.

Fourth-stage larvae were found chiefly in the sections that were later most heavily parasitized by adult worms.