In theory there should be a strong coupling between host and parasite population sizes. Here, we investigated population size and structure in 3 species of acanthocephalans, Corynosoma semerme, C. strumosum and C. magdaleni, in ringed seals (Phoca hispida) from the Bothnian Bay over a period of more than 20 years. During this period, seal numbers first decreased markedly and then increased steadily; at the same time, a paratenic fish host particularly important for C. strumosum has gradually disappeared from the bay due to decreasing salinity. We found no evidence that the mean abundance of any of the 3 acanthocephalan species changed significantly over time, nor was there any relationship between parasite abundance at any point in time and seal numbers in the corresponding year. Based on the proportion of sexually mature female worms per infrapopulation, and on relationships between the sex ratio of worms and infrapopulation size, both C. magdaleni and C. semerme appear to be doing well, independently of the population size of their seal definitive hosts. In contrast, perhaps because of the loss of its main paratenic host, C. strumosum appears more at risk in the Bothnian Bay. Our results show that in complex natural systems, there are not necessarily simple, direct links between definitive host population size or density, and parasite population dynamics.

The number of Teladorsagia circumcincta 4th-stage larvae in naturally infected lambs from a single farm varied among lambs and among different years. Within each year the distribution of 4th-stage larvae among lambs was similar to that expected from a negative binomial distribution. The ratio of 4th-stage larvae to adult T. circumcincta was low in two years with a low mean intensity of infection but high in two years with a higher mean intensity of infection. The negative binomial distribution is defined by the mean and by k, a parameter that measures dispersion; k was low when mean infection intensity was low but higher when mean infection intensity was high. As k is an inverse index of overdispersion this indicated that the distribution of 4th-stage larvae was more overdispersed at low levels of infection. In a combined analysis, the number of adult T. circumcincta and the plasma IgA activity against 4th-stage larvae were both associated with increased numbers of 4th-stage larvae. There was a statistical interaction between the number of adults and IgA activity that moderated their combined effect.

Infection parameters of Posthodiplostomum cuticola, a digenean parasite with a complex life-cycle, were investigated in fish (the second intermediate host) from 6 floodplain water bodies over 2 years. A broad range of factors related to abiotic characteristics of localities, density of the first intermediate (planorbid snails) and definitive (wading birds) hosts and fish community structure were tested for their effects on P. cuticola infection in juvenile and adult fish. Characters of the littoral zone and flood duration were found to be important factors for the presence of the first intermediate and definitive hosts. Visitation time of definitive bird hosts was also related to adult fish host density. Localities with P. cuticola infected fish were visited by a higher number of bird species. Infection of P. cuticola in fish and similarities in infection among fish host assemblages were correlated with fish host density and fish species composition. Parasite infection in both adult and juvenile fishes was associated with the slope of the bank and the bottom type, in particular in juvenile fish assemblages with snail host density. We conclude that habitat characteristics, snail host density and fish community structure contribute significantly to P. cuticola infection in fish hosts.

The skin site at which ticks attach to their hosts to feed is the critical interface between the tick and its host, and tick-borne pathogens. This site is highly modified by the pharmacologically active molecules secreted in tick saliva. For pathogens, it is an ecologically privileged niche that many exploit. Such exploitation is referred to as saliva-activated transmission (SAT) – the indirect promotion of tick-borne pathogen transmission via the actions of bioactive tick saliva molecules on the vertebrate host. Here we review evidence for SAT and consider what are the most likely candidates for SAT factors among the tick pharmacopoeia of anti-haemostatic, anti-inflammatory and immunomodulatory molecules identified to date. SAT factors appear to differ for different pathogens and tick vector species, and possibly even depend on the vertebrate host species. Most likely we are searching for a suite of molecules that act together to overcome the redundancy in host response mechanisms. Whatever they turn out to be, the quest to identify the tick molecules that mediate SAT is an exciting one, and offers new insights to controlling ticks and tick-borne diseases.

Sexual size dimorphism (SSD) in mammals reveals the extent of sexual selection, which may in turn explain why males are often more infected by parasites than females and that parasites may contribute to the association between SSD and male-biased mortality. Here, we investigated the relationship between SSD in small mammals of Central Europe and the differences in sex infection by fleas. A comparative analysis was conducted for 10 species of rodents and insectivores. We found that males harbour higher flea species richness than females and that the abundance of fleas is higher in males than in females. This difference is not related to male-biased density. However, contrary to our hypothesis, we found that an increase in SSD is not related to an increase in male infection by fleas compared with female infection. We discuss our results in term of sex-differences in immunocompetence and/or sex-differences in behaviour.

Due to the low informative value of available morphological characters, cytogenetic and molecular methods, based on rDNA sequencing, were used to characterize adult and larval stages of Phyllodistomum spp. Species studied have 18 chromosomes with comparable absolute and relative lengths. Conventional Giemsa staining and karyometric analysis revealed clear differences in chromosome morphology of larval Phyllodistomum spp. infecting two bivalve host species, Sphaerium corneum and Pisidium amnicum. However, karyotypes of adult P. folium from three-spined sticklebacks and larval stages from S. corneum appear almost identical both with respect to the relative lengths and centromeric indices of the corresponding chromosome pairs. The entire internal transcribed spacer (ITS) region (ITS-1, 5.8S and ITS-2) and the D1-D3 region of 28S gene were sequenced and compared. Again, sufficient differences were observed between larval Phyllodistomum spp., while adult P. folium and larvae from S. corneum showed a high level of similarity. So, both cytogenetic and molecular data support the suggestion that they represent developmental stages of the same species. The results were compared with published data obtained by cytogenetic and molecular studies on the other Phyllodistomum species. Differences revealed in karyotype and rDNA sequences leads to the conclusion that the cercariaeum of P. folium sensu Sinitsin, 1905 could not be regarded as the larva of adult P. folium from three-spined stickleback.

A copro-antigen capture ELISA for the detection of Teladorsagia circumcincta infection in sheep was developed and evaluated. Experiments with faeces from worm-free sheep, that had been spiked with known concentrations of excretory–secretory (E–S) antigen indicated that a positive signal was generated with 180 ng of E–S/ml. A nested design, based on 8 infected and 8 worm-free sheep, was employed to assess the stages during sample preparation contributing to variation in signal from the assay. This showed that 87% of the variance in the optical density readings (ODs) was directly explained by infection status. Variation between individual sheep within infection groups, and between samples at various stages in the assay, collectively accounted for the remaining variance. Initial evaluation of specificity using faeces from animals carrying a range of monospecific infections indicated cross-reactivity with Haemonchus contortus and Nematodirus spathiger. However, by treating the supernatant from faeces for 20 min at 100 °C, the cross-reactive signal was eliminated whilst the specific signal was largely preserved. Heat treatment of faeces from 12 non-infected sheep, 12 sheep with T. circumcincta and 6 with H. contortus resulted in sensitivity being increased from 66·7 to 85·7%, and specificity from 62·5 to 87·5%. OD values showed a significant positive relationship with adult worm burdens, although at low infection intensities there was some overlap between infected and worm-free animals. We discuss the application of CC-ELISAs in facilitating selective chemotherapy of sheep, as a means of avoiding the development of anthelmintic resistance in pastoral regions where sheep are farmed on a large scale.

Since the discovery of the Lyme disease spirochete in North America in 1982 and in Europe in 1983, a plethora of studies on this unique group of spirochetes that comprise Borrelia burgdorferi sensu lato has been accumulated. In an attempt to compare and contrast Lyme borreliosis in Europe and North America we have reviewed the biology of the aetiologic agents, as well as the clinical aspects, diagnosis and treatment of this disease on both continents. Moreover, we have detailed the ecology of the Ixodes ticks that transmit this infection and the reservoir hosts that maintain the spirochete cycle in nature. Finally, we have examined the transmission dynamics of the spirochete on both continents, as well as the available prevention strategies. Although it has been over two decades since the discovery of the Lyme disease spirochete, Lyme borreliosis is an expanding public health problem that has defied our attempts to control it. By comparing the accumulated experience of investigators in North America and Europe, where the disease is most frequently reported, we hope to advance the cause of developing novel approaches to combat Lyme borreliosis.

Published in Parasitology (2004), 128, 503–509.

Chemical cues are so important in the recognition mechanism of social insects that most social parasites (which rely on hosts to rear their brood) have been documented as overcoming the mechanism by which colony residents recognize non-nestmates, by mimicking the odour of the usurped colony. We simulated in the laboratory the process by which the obligate social parasite, Polistes semenowi, invades nests of the host species, Polistes dominulus, in the field and analysed the epicuticular lipid layer before and after host nest usurpation. The experiment documents that P. semenowi social parasites have an epicuticular hydrocarbon pattern which is very similar to that of their host but, after entering host colonies, parasites mimic the odour of the colonies they invade, to the point that they perfectly match the hydrocarbon profile peculiar to the colony they entered. However, both before and after host nest invasion, parasites show a tendency to possess diluted recognition cues with respect to their hosts.

Fibres isolated from the ventral sucker of Fasciola hepatica were identified as muscle on the basis of their contractility, and their actin and myosin staining. They were voltage-clamped at a holding potential of −40 mV and depolarization-activated outward currents were characterized both electrophysiologically and pharmacologically. Activation was well fitted by a Boltzmann equation with a half-maximal potential of +9 mV and a slope factor of −14·3 mV, and the kinetics of activation and deactivation were voltage-sensitive. Tail current analysis showed that the reversal potential was shifted by +46±3 mV when EK was increased by 52 mV, confirming that this was a K+-current with electrophysiological characteristics similar to delayed rectifier and Ca2+-activated K+-currents in other tissues. The peak current at +60 mV was inhibited by 76±6% by tetrapentylammonium chloride (1 mM) and by 84±7% by Ba2+ (3 mM), but was completely resistant to block by tetraethylammonium (30 mM), 3,4-diaminopyridine (100 μM) and 4-aminopyridine (10 mM). Penitrem A, a blocker of high-conductance Ca2+-activated K+-channels reduced the current at +60 mV by 23±5%. When the effects of Ca2+-channel blocking agents were tested, the peak outward current at +60 mV was reduced by 71±7% by verapamil (30 μM) and by 59±4% by nimodipine (30 μM). Superfusion with BAPTA-AM (50 μM), which is hydrolysed intracellularly to release the Ca2+-buffer BAPTA, also decreased the current by 44±16%. We conclude that voltage-and Ca2+-sensitive K+-channels are expressed in this tissue, but that their pharmacology differs considerably from equivalent channels in other phyla.

At least 38 viral species are transmitted by ticks. Virus–tick–vertebrate host relationships are highly specific and less than 10% of all tick species (Argasidae and Ixodidae) are known to play a role as vectors of arboviruses. However, a few tick species transmit several (e.g. Ixodes ricinus, Amblyomma variegatum) or many (I. uriae) tick-borne viruses. Tick-borne viruses are found in six different virus families (Asfarviridae, Reoviridae, Rhabdoviridae, Orthomyxoviridae, Bunyaviridae, Flaviviridae) and at least 9 genera. Some as yet unassigned tick-borne viruses may belong to a seventh family, the Arenaviridae. With only one exception (African swine fever virus, family Asfarviridae) all tick-borne viruses (as well as all other arboviruses) are RNA viruses. Tick-borne viruses are found in all the RNA virus families in which insect-borne members are found, with the exception of the family Togaviridae. Some tick-borne viruses pose a significant threat to the health of humans (Tick-borne encephalitis virus, Crimean-Congo haemorrhagic fever virus) or livestock (African swine fever virus, Nairobi sheep disease virus). Key challenges are to determine the molecular adaptations that allow tick-borne viruses to infect and replicate in both tick and vertebrate cells, and to identify the principal ecological determinants of tick-borne virus survival.

This study describes, for the first time, the characterization of excretory–secretory antigens (ES-Ag) from Echinococcus granulosus protoscoleces, evaluating their usefulness in the immunodiagnosis of human cystic echinococcosis. ES-Ag were obtained from the first 50 h maintenance of protoscoleces in vitro. This preparation contained over 20 major protein components which could be distinguished by 1-dimensional SDS–PAGE with apparent masses between 9 and 300 kDa. The culture of of protoscoleces from liver produced a greater variety of excretory–secretory protein components than those from lung. Determination of enzymatic activities of secreted proteins revealed the presence of phosphatases, lipases and glucosidases, but no proteases. These findings were compared to those obtained from somatic extracts of protoscoleces and hydatid cyst fluid products. Immunochemical characterization was performed by immunoblotting with sera from individuals infected by cystic echinococcosis (n=15), non-hydatidic parasitoses (n=19), various liver diseases (n=24), lung neoplasia (n=16), and healthy donors (n=18). Antigens with apparent masses of 89, 74, 47/50, 32, and 20 kDa showed specificity for immunodiagnosis of human hydatidosis. The 89 and 74 kDa components corresponded to antigens not yet described in E. granulosus, whereas proteins of 41–43 kDa and 91–95 kDa were recognized by the majority of the non-hydatid sera studied.

We compared helminth communities in spiny mice (Acomys dimidiatus) from 4 wadis in the arid montane region of the southern Sinai in Egypt, in a 4-week period in late summer. Total helminth species richness was 14 (8 nematodes, 5 cestodes and 1 acanthocephalan) with 94% of mice carrying at least 1 species and an overall mean species richness of 1·85. The most prevalent parasites were Protospirura muricola (47·8%) and Dentostomella kuntzi (46·3%). One larval cestode, Joyeuxiella rossicum, represents a new host record. The helminth community was dominated by intestinal nematodes (88·7%) of which 58·2% were arthropod-transmitted heteroxenic species. At the component community level, 70% of the worms were recovered from mice in just two wadis (Gharaba and Tlah) and 48·6% of intestinal nematodes were from Wadi Gharaba. Although only 7 species of helminths were recorded from Wadi Gharaba, this site gave the highest Berger-Parker dominance index because of P. muricola. P. muricola was also dominant in Wadi El Arbaein whilst Syphacia minuta was the dominant species in Wadis Gebal and Tlah. At the infracommunity level, mean species richness and Brillouin's index of diversity were highest in Wadi Tlah and lowest in Wadi Gebal, and the former was age dependent. Whilst mice from different wadis differed in the nematodes that were most common, those from Wadi Gharaba carried the highest mean number of worms/mouse. The abundance of P. muricola in particular varied markedly between sites: Wadi Gharaba was distinct as the site showing the highest mean worm burden whereas mice from Wadi Gebal were uninfected. None of the directly transmitted oxyuroid nematodes showed significant variation in abundance between wadis, or host sex or age classes. Overall, the single extrinsic factor in the study, site of capture, was more important than the intrinsic factors in explaining variation in helminth communities in the region. We conclude that in the high mountains of southern Sinai, each wadi is distinct in terms of its rodent parasites, and hence we expect spatially different coevolutionary pressures on their hosts, with resultant variation in life-histories.

Tick fever or cattle fever (babesiosis) is economically the most important arthropod-borne disease of cattle worldwide with vast areas of Australia, Africa, South and Central America and the United States continuously under threat. Tick fever was the first disease for which transmission by an arthropod to a mammal was implicated at the turn of the twentieth century and is the first disease to be eradicated from a continent (North America). This review describes the biology of Babesia spp. in the host and the tick, the scale of the problem to the cattle industry, the various components of control programmes, epidemiology, pathogenesis, immunity, vaccination and future research. The emphasis is on Babesia bovis and Babesia bigemina.

Theileria are economically important, intra-cellular protozoa, transmitted by ixodid ticks, which infect wild and domestic ruminants. In the mammalian host, parasites infect leukocytes and erythrocytes. In the arthropod vector they develop in gut epithelial cells and salivary glands. All four intra-cellular stages of Theileria survive free in the cytoplasm. The schizont stages of certain Theileria species induce a unique, cancer-like, phenotype in infected host leukocytes. Theileria undergoes an obligate sexual cycle, involving fusion of gametes in the tick gut, to produce a transiently diploid zygote. The existence of sexual recombination in T. parva has been confirmed in the laboratory, and is presumed to contribute to the extensive polymorphism observed in field isolates. Key parameters in T. parva population dynamics are the relative importance of asymptomatic carrier cattle and animals undergoing severe disease, in transmission of the parasite to ticks, and the extent of transmission by nymphs as compared to adult ticks. Tick populations differ in vector competence for specific T. parva stocks. Recombinant forms of T. parva and T. annulata sporozoite surface antigens induce protection against parasite challenge in cattle. In future, vaccines might be improved by inclusion of tick peptides in multivalent vaccines.

The tick-borne intracellular pathogen Anaplasma marginale (Rickettsiales: Anaplasmataceae) develops persistent infections in cattle and tick hosts. While erythrocytes appear to be the only site of infection in cattle, A. marginale undergoes a complex developmental cycle in ticks and transmission occurs via salivary glands during feeding. Many geographic isolates occur that vary in genotype, antigenic composition, morphology and infectivity for ticks. In this chapter we review recent research on the host–vector–pathogen interactions of A. marginale. Major surface proteins (MSPs) play a crucial role in the interaction of A. marginale with host cells. The MSP1a protein, which is an adhesin for bovine erythrocytes and tick cells, is differentially regulated and affects infection and transmission of A. marginale by Dermacentor spp. ticks. MSP2 undergoes antigenic variation and selection in cattle and ticks, and contributes to the maintenance of persistent infections. Phylogenetic studies of A. marginale geographic isolates using msp4 and msp1α provide information about the biogeography and evolution of A. marginale: msp1α genotypes evolve under positive selection pressure. Isolates of A. marginale are maintained by independent transmission events and a mechanism of infection exclusion in cattle and ticks allows for only the infection of one isolate per animal. Prospects for development of control strategies by use of pathogen and tick-derived antigens are discussed. The A. marginale/vector/host studies described herein could serve as a model for research on other tick-borne rickettsiae.

The emergence of Lyme borreliosis as a public health burden within the last two decades has stimulated renewed interest in tick-borne infections. This attention towards ticks, coupled with advances in detection technologies, has promoted the recognition of diverse emergent or potentially emerging infections, such as monocytic and granulocytic ehrlichiosis, local variants of spotted fever group rickettsioses, WA-1 babesiosis, or a Lyme disease mimic (Masters' Disease). The distribution of pathogens associated with well-described tick-borne zoonoses such as human babesiosis due to Babesia microti or B. divergens seems wider than previously thought. Bartonellae, previously known to be maintained by fleas, lice or sandflies, have been detected within ticks. Purported ‘new’ agents, mainly identified by sequencing of PCR products and comparison with those sequences present in GenBank, are being increasingly reported from ticks. We briefly review the diversity of these infectious agents, identify aetiological enigmas that remain to be solved, and provide a reminder about ‘old friends’ that should not be forgotten in our pursuit of novelty. We suggest that newly recognised agents or tick/pathogen associations receive careful scrutiny before being declared as potential public health burdens.

Geographic information systems (GIS), their fundamental components and technologies are described. GIS is a computer-based system enabling the storage, integration, query, display and analysis of data using information on data location. Further, remote sensing (RS) methods and their application in landscape characterization are described. Landscape pattern analysis, combined with statistical analysis, allows the determination of landscape predictors of disease risk. This makes RS/GIS a powerful set of tools for disease surveillance, enabling the prediction of potential disease outbreaks and targeting intervention programs. The ‘pre-GIS era’ is briefly described including the early mapping of tick distribution, analyses and the display of biogeographical and medical data. The theory of natural focality of diseases (NFD) is explained and its significance in tick-borne diseases (TBD) research is discussed. Many problems of tick ecology and TBD epidemiology and epizootology have been addressed by means of GIS and examples of these studies are presented and discussed.