Artificial feeding of ixodid ticks

Ixodid ticks are economically important as they cause direct damage to livestock and are vectors of several pathogens that cause diseases in humans and animals. Some of the important tick-borne pathogens of livestock are Theileria parva, T. annulata, Babesia bigemina, B. bovis, Anaplasma marginale and Cowdria ruminantium. These pathogens are responsible for causing enormous losses in livestock. Identification of factors that influence transmission and development of these pathogens in ticks will greatly facilitate development of rational strategies for control of tick-borne diseases. This research has been hampered by the lack of suitable artificial feeding methods. In this paper, Sam Waladde, Aian Young and Subhash Morzaria review recent developments in the artificial feeding of ixodid ticks and evaluate how this method can potentially be exploited. They use an example the transmission of an important livestock pathogen, T. parva, by Rhipicephalus appendiculatus.     

Host resistance in cattle tick control

Cattle ticks are an important constraint on the livestock industry, particularly in tropical and subtropical areas, mainly because of the diseases they transmit and the costs of control. Conventional control is by means of acaricides; although there are still serious drawbacks, these can be minimized by a strategic approach. In this review Julio de Castro and Robin Newson look at alternatives. One is to make use of the host's innate abilities. In the simplest situation, where no control measures are applied, the animals develop their own protective immunity against ticks and tick-borne diseases (TBDs). Alternatively, breeds of cattle with naturally high resistance to ticks can be used, enhanced by selection. Methods of immunizing cattle against ticks are also under development. Ideally, ticks should be managed at an economically acceptable level by a combination of techniques, supported where necessary by vaccination against TBDs.     

Is endemic stability of tick-borne disease in cattle a useful concept?

Endemic stability is a widely used term in the epidemiology of ticks and tick-borne diseases. It is generally accepted to refer to a state of a host-tick-pathogen interaction in which there is a high level of challenge of calves by infected ticks, absence of clinical disease in calves despite infection, and a high level of immunity in adult cattle with consequent low incidence of clinical disease. Although endemic stability is a valid epidemiological concept, the modelling studies that underpinned subsequent studies on the epidemiology of tick-borne diseases were specific to a single host-tick-pathogen system, and values derived from these models should not be applied in other regions or host-tick-pathogen systems.     

Control of ticks of ruminants, with special emphasis on livestock farming systems in India: present and future possibilities for integrated control—a review

India is predominantly an agricultural country with about 70% of her population dependent on income from agriculture. Although India accounts for a significant share of world’s livestock resources, livestock production is greatly affected by ticks and tick-borne diseases (TTBDs). Therefore, India represents a particularly interesting scenario for the study of TTBDs. Herein, we review the problems and opportunities for the integrated control of ticks of ruminants with special emphasis on livestock farming systems in India. Developments discussed in the review in the area of tick vaccines and other tick control measures should have an impact on the future of Indian livestock production.     

Identification of novel tick salivary gland proteins for vaccine development

Methods currently used to control Ixodes scapularis ticks rely principally on acaricidal applications which suffer from a number of limitations. Recently, host vaccination against ticks has been shown to be a promising alternative tick control method. In tick salivary glands, numerous genes are induced during the feeding process. Many of these newly expressed proteins are secreted in tick saliva and may play a role in modulating host immune responses and pathogen transmission. We have performed suppression subtraction hybridization to identify unique I. scapularis salary gland proteins specifically expressed during engorgement. We have cloned and sequenced ten unique salivary gland-associated cDNAs that are up-regulated during feeding. The protein products of these genes represent potential vaccine candidates for use in the control of ticks and to prevent transmission of tick-borne diseases.     

Tick-borne infections of animals and humans: a common ground

A wide variety of pathogens is transmitted from ticks to vertebrates including viruses, bacteria, protozoa and helminths, of which most have a life cycle that requires passage through the vertebrate host. Tick-borne infections of humans, farm and companion animals are essentially associated with wildlife animal reservoirs. While some flying insect-borne diseases of humans such as malaria, filariasis and Kala Azar caused by Leishmania donovani target people as their main host, major tick-borne infections of humans, although potentially causing disease in large numbers of individuals, are typically an infringement of a circulation between wildlife animal reservoirs and tick vectors. While new tick-borne infectious agents are frequently recognised, emerging agents of human tick-borne infections were probably circulating among wildlife animal and tick populations long before being recognised as clinical causes of human disease as has been shown for Borrelia burgdorferi sensu lato. Co-infection with more than one tick-borne infection is common and can enhance pathogenic processes and augment disease severity as found in B. burgdorferi and Anaplasma phagocytophilum co-infection. The role of wild animal reservoirs in co-infection of human hosts appears to be central, further linking human and animal tick-borne infections. Although transmission of most tick-borne infections is through the tick saliva, additional routes of transmission, shown mostly in animals, include infection by oral uptake of infected ticks, by carnivorism, animal bites and transplacentally. Additionally, artificial infection via blood transfusion is a growing threat in both human and veterinary medicine. Due to the close association between human and animal tick-borne infections, control programs for these diseases require integration of data from veterinary and human reporting systems, surveillance in wildlife and tick populations, and combined teams of experts from several scientific disciplines such as entomology, epidemiology, medicine, public health and veterinary medicine.     

Tick genomics: the Ixodes genome project and beyond

Ticks and mites (subphylum Chelicerata; subclass Acari) include important pests of animals and plants worldwide. The Ixodes scapularis (black-legged tick) genome sequencing project marks the beginning of the genomics era for the field of acarology. This project is the first to sequence the genome of a blood-feeding tick vector of human disease and a member of the subphylum Chelicerata. Genome projects for other species of Acari are forthcoming and their genome sequences will likely feature significantly in the future of tick research. Parasitologists interested in advancing the field of tick genomics research will be faced with specific challenges. The development of genetic tools and resources, and the size and repetitive nature of tick genomes are important considerations. Innovative approaches may be required to sequence, assemble, annotate and analyse tick genomes. Overcoming these challenges will enable scientists to investigate the genes and genome organisation of this important group of arthropods and may ultimately lead to new solutions for control of ticks and tick-borne diseases.     

Integrated control of vector-borne diseases of livestock--pyrethroids: panacea or poison?

Tick- and tsetse-borne diseases cost Africa approximately US$4-5 billion per year in livestock production-associated losses. The use of pyrethroid-treated cattle to control ticks and tsetse promises to be an increasingly important tool to counter this loss. However, uncontrolled use of this technology might lead to environmental damage, acaricide resistance in tick populations and a possible exacerbation of tick-borne diseases. Recent research to identify, quantify and to develop strategies to avoid these effects are highlighted.     

Present status of tick-borne diseases in Sudan

Ticks and tick-borne diseases are widespread in the Sudan, cause substantial economic losses and constitute major obstacles to the development of animal wealth. Most important among these diseases are tropical theileriosis, malignant ovine theileriosis, cowdriosis, babesiosis, anaplasmosis and avian spirochaetosis. However, knowledge about ticks and tick-borne diseases is still fragmentary and far from complete. The large number of tick species, the multplicity of transmitted agents and the diverse ecoclimatic zones of the Sudan provide a unique opportunity to host diverse research activities that could benefit other regions in Africa.     

How ticks keep ticking in the adversity of host immune reactions

Journal of Mathematical Biology - Ixodid ticks are acknowledged as one of the most important hematophagous arthropods because of their ability in transmitting a variety of tick-borne diseases....     

Prevalence of tick-borne pathogens in ixodid ticks (Acari: Ixodidae) collected from European wild boar (<Emphasis Type="Italic">Sus scrofa</Emphasis>) and Iberian red deer (<Emphasis Type="Italic">Cervus elaphus hispanicus</Emphasis>) in central Spain

Emerging tick-borne diseases of humans and animals have occurred frequently during the past 30 years. These disease outbreaks appear to result from changes in the distribution of tick and vertebrate hosts, and the introduction of humans and domestic animals into tick–pathogen–wildlife cycles. Use of molecular technologies now available for identification of pathogens in ticks can provide valuable information that allows for risk analysis of emerging tick-borne diseases. In this study, the prevalence of selected pathogens in ticks collected in six locations in central Spain from the major wild ungulate species, European wild boar (Sus scrofa) and Iberian red deer (Cervus elaphus hispanicus), was determined by PCR. Tick species collected included Ixodes ricinus, Dermacentor marginatus, Rhipicephalus bursa and Hyalomma m. marginatum. Pathogens identified in ticks included piroplasmids, Anaplasma spp., Ehrlichia spp. and Rickettsia spp. Piroplasmids were identified in all tick species except I. ricinus. Ehrlichia spp. were detected in all tick species and collection locations, while Rickettsia spp., which proved to be R. slovaca and a recently identified Rickettsia sp. DnS28, were identified only in D. marginatus. A. marginale and A. phagocytophilum were detected in D. marginatus, R. bursa and Hy. m. marginatum. Concurrent infections of these pathogens were frequently observed in ticks. Notably, A. phagocytophilum, which is infective for a broad host range that includes humans and domestic and wild animals, was identified in ticks from all collection locations. The variety of ticks and tick-borne pathogens demonstrated in this study suggests a risk in central Spain for the emergence of tick-borne diseases in humans and domestic animals.     

Effect of Prior Exposure to Noninfected Ticks on Susceptibility of Mice to Lyme Disease Spirochetes

To determine whether prior exposure to Nearctic Ixodes vector ticks protects native reservoir mice from tick-borne infection by Lyme disease spirochetes, we compared their infectivities for white-footed mice and laboratory mice that had been repeatedly infested by noninfected deer ticks. Nymphal ticks readily engorged on tick-exposed laboratory mice, but their feeding success on white-footed mice progressively declined. Tick-borne spirochetes readily infected previously tick-infested mice. Thus, prior infestation by Nearctic ticks does not protect sympatric reservoir mice or Palearctic laboratory mice from infection by sympatric tick-borne spirochetes.     

Co-feeding ticks: Epidemiological significance for tick-borne pathogen transmission

Until recently, the transmission of tick-borne pathogens via vertebrates was thought to depend on the development of a systemic infection in the vertebrate hosts. Pathogen transmission has now been shown to occur between infected and uninfected ticks co-feeding in time or space in the absence of a systemic infection, originally for viruses, but now also for bacteria. The epidemiological consequences of this new non-systemic transmission pathway necessitate a major reassessment of the components and dynamics of tick-borne pathogen enzootic cycles. Here Sarah Randolph, Lise Gern and Pat Nuttall show that a much wider range of natural hosts than was previously recognized may contribute significantly to the transmission of tick-borne diseases, and compare quantitatively the relative contributions made by the systemic and non-systemic transmission pathways.     

Projecting the potential distribution of ticks in China under climate and land use change

Ticks are known as vectors of several pathogens causing various human and animal diseases including Lyme borreliosis, tick-borne encephalitis, and Crimean-Congo hemorrhagic fever. While China is known to have more than 100 tick species well distributed over the country, our knowledge on the likely distribution of ticks in the future remains very limited, which hinders the prevention and control of the risk of tick-borne diseases. In this study, we selected four representative tick species which have different regional distribution foci in mainland China. i.e., Dermacentor marginatus, Dermacentor silvarum, Haemaphysalis longicornis and Ixodes granulatus. We used the MaxEnt model to identify the key environmental factors of tick occurrence and map their potential distributions in 2050 under four combined climate and socioeconomic scenarios (i.e., SSP1-RCP2.6, SSP2-RCP4.5, SSP3-RCP7.0 and SSP5-RCP8.5). We found that the extent of the urban fabric, cropland and forest, temperature annual range and precipitation of the driest month were the main determinants of the potential distributions of the four tick species. Under the combined scenarios, with climate warming, the potential distributions of ticks shifted to further north in China. Due to a decrease in the extent of forest, the distribution probability of ticks declined in central and southern China. In contrast with previous findings on an estimated amplification of tick distribution probability under the extreme emission scenario (RCP8.5), our studies projected an overall reduction in the distribution probability under RCP8.5, owing to an expected effect of land use. Our results could provide new data to help identify the emerging risk areas, with amplifying suitability for tick occurrence, for the prevention and control of tick-borne zoonoses in mainland China. Future directions are suggested towards improved quantity and quality of the tick occurrence database, comprehensiveness of factors and integration of different modelling approaches, and capability to model pathogen spillover at the human-tick interface.     

Protective immune mechanisms to ticks and tick-borne diseases of ruminants.

A workshop of the European Union (EU) Concerted Action Project on the Integrated Control of Ticks and Tick-borne diseases (CA-ICTTD) recently assessed protective immune mechanisms to ticks and tick-borne diseases. The consensus achieved there is summarized here by Patricia Preston and Frans Jongejan. The current understanding of this field is expanded upon by the accompanying articles, and Poster, in this special issue of Parasitology Today.     

The effects of the brown ear-tick, Rhipicephalus appendiculatus, on milk production in dairy cattle

Abstract. The effect is reported of artificially controlled levels of infestation with adults of Rhipicephalus appendiculatus on the milk yield of twenty commercial Bos taurus dairy cattle on a high plane of nutrition and eighteen crossbed B.taurus × Sanga cattle on a lower plane of nutrition in the highveld of Zimbabwe. The results showed no significant effect on milk yield of infestations averaging twenty engorging ticks per animal per day, despite severe ear damage in some animals. They indicate that milk production of dairy cattle under commercial management is not sensitive to infestation with R.appendiculatus. The results have important implications for management of ticks in Africa, but need to be interpreted within the context of the control of tick-borne diseases.     

Tick-borne pathogens in Dermacentor reticulatus collected from dogs in eastern Poland

In recent years, the distribution of Dermacentor reticulatus ticks has expanded into new territories in many European countries, including Poland, with increased population densities in areas of their regular occurrence. The spread of D. reticulatus enhances the risk of exposure of domestic animals and their owners to tick-borne diseases. The objective of this study was to assess the prevalence of infection of D. reticulatus ticks feeding on dogs with the pathogens Borrelia burgdorferi sensu lato and Anaplasma phagocytophilum. The study material comprised 152 D. reticulatus ticks collected from dogs in the northeastern part of Lublin Province (eastern Poland). A ready-made AmpliSens® TBEV, B.burgdorferi sl, A.phagocytophilum, E.chaffeensis/E.muris-FRT PCR kit was used for qualitative detection and differentiation of tick-borne infections. The assessment of the degree of infection of the analyzed ticks with the two pathogens revealed that 9.2% (14/152) of the examined ticks were infected with one of the pathogens. No co-infections with the pathogens were detected in any of the ticks. The highest specific percentage of infections (8.6%, 13/152) was associated with A. phagocytophilum. The presence of B. burgdorferi s.l. was detected in only one of the examined ticks (0.7%). The spread of D. reticulatus to new territories and the increase in population density in areas of their regular occurrence implies the need for further studies of the prevalence of pathogens with medical and veterinary importance in order to assess the risk of tick-borne diseases.

     

Experimental Transmission of Karshi (Mammalian Tick-Borne Flavivirus Group) Virus by Ornithodoros Ticks >2,900 Days after Initial Virus Exposure Supports the Role of Soft Ticks as a Long-Term Maintenance Mechanism for Certain Flaviviruses

Background

Members of the mammalian tick-borne flavivirus group, including tick-borne encephalitis virus, are responsible for at least 10,000 clinical cases of tick-borne encephalitis each year. To attempt to explain the long-term maintenance of members of this group, we followed Ornithodoros parkeri, O. sonrai, and O. tartakovskyi for >2,900 days after they had been exposed to Karshi virus, a member of the mammalian tick-borne flavivirus group.

Methodology/Principal Findings

Ticks were exposed to Karshi virus either by allowing them to feed on viremic suckling mice or by intracoelomic inoculation. The ticks were then allowed to feed individually on suckling mice after various periods of extrinsic incubation to determine their ability to transmit virus by bite and to determine how long the ticks would remain infectious. The ticks remained efficient vectors of Karshi virus, even when tested >2,900 d after their initial exposure to virus, including those ticks exposed to Karshi virus either orally or by inoculation.

Conclusions/Significance

Ornithodoros spp. ticks were able to transmit Karshi virus for >2,900 days (nearly 8 years) after a single exposure to a viremic mouse. Therefore, these ticks may serve as a long-term maintenance mechanism for Karshi virus and potentially other members of the mammalian tick-borne flavivirus group.     

A Stochastic Tick-Borne Disease Model: Exploring the Probability of Pathogen Persistence

We formulate and analyse a stochastic epidemic model for the transmission dynamics of a tick-borne disease in a single population using a continuous-time Markov chain approach. The stochastic model is based on an existing deterministic metapopulation tick-borne disease model. We compare the disease dynamics of the deterministic and stochastic models in order to determine the effect of randomness in tick-borne disease dynamics. The probability of disease extinction and that of a major outbreak are computed and approximated using the multitype Galton–Watson branching process and numerical simulations, respectively. Analytical and numerical results show some significant differences in model predictions between the stochastic and deterministic models. In particular, we find that a disease outbreak is more likely if the disease is introduced by infected deer as opposed to infected ticks. These insights demonstrate the importance of host movement in the expansion of tick-borne diseases into new geographic areas.     

Ecological correlates of a tick-borne disease, Anaplasma phagocytophilum, in moose in southern Norway

As the distribution and abundance of ticks increase, so do the risks of tick-borne diseases. Anaplasma phagocytophilum, transmitted by Ixodes spp. ticks, is a widespread tick-borne infection causing tick-borne fever (TBF) in domestic ruminants and human granulocytic anaplasmosis. However, the role of wildlife in its epidemiology is poorly understood. Evidence of infection has been detected in wild cervids, but the pathogenicity and ecological consequences are unknown. We conducted a serological study of moose (Alces alces) in two populations in southern Norway, one where TBF was endemic (Telemark) and the other where sheep ticks (Ixodes ricinus) were essentially absent (Hedmark). Seroprevalence to A. phagocytophilum antibodies was 79 and 0 %, respectively. In Telemark, seroprevalence was significantly higher among females that calved successfully (85 %) than among others (50 %). Body mass and winter mass change were unrelated to serostatus. Relative abundance of questing ticks in Telemark was highest in deciduous forest and lowest in mature coniferous forest and higher at easterly aspects and altitudes below 350 m. Habitat factors associated with high tick abundance were risk factors for seropositivity among moose. Our findings were consistent with anaplasmosis causing a persistent subclinical infection in moose without population-level effects. Further work is needed to establish the importance of moose as a reservoir for the disease in sympatric domestic livestock.