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PubMed1997

Identification of an Ixodes ricinus salivary gland fraction through its ability to stimulate CD4 T cells present in BALB/c mice lymph nodes draining the tick fixation site.

BALB/c mice infested with larvae or nymphs of Ixodes ricinus develop in their lymph nodes a T cell-specific immune response triggered by salivary gland soluble antigens (SGA). SGA are apparently conserved in the 3 biological stages of I. ricinus ticks and are species specific. SGA derived from partially fed females I. ricinus stimulate lymph node T cells from mice infested with I. ricinus larvae or nymphs. In contrast, lymph node cells from mice infested with Amblyomma hebraeum nymphs do not respond. A chromatographic fraction enriched with a 65 kDa protein (IrSG65) isolated from salivary glands of I. ricinus partially fed females induces in vitro a specific T cell proliferation of lymph node cells from mice infested with I. ricinus nymphs. The depletion of CD4+ T cells drastically reduces the ability of lymphocytes from infested mice to proliferate after IrSG65 stimulation.

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PubMed1999

Transmission of Borrelia afzelii from Apodemus mice and Clethrionomys voles to Ixodes ricinus ticks: differential transmission pattern and overwintering maintenance.

This study deals with the ecology of Lyme borreliosis in Europe. The relationships between Borrelia burgdorferi sensu lato spirochetes, Clethrionomys and Apodemus rodent reservoirs and the Ixodes ricinus tick vector were investigated during 16 consecutive months in an enzootic area in Switzerland. Cultivation of ear skin biopsies was used to isolate spirochetes from C. glareolus, A. sylvaticus, A. flavicollis and Glis glis. Borrelia infection was more frequently observed in Clethrionomys than in Apodemus. Tick xenodiagnosis was used to determine the infectivity of rodents. The infection rate in ticks fed on Clethrionomys was higher than that in ticks fed on Apodemus, but Apodemus yielded more infected ticks than Clethrionomys because of a better tick moulting success. Xenodiagnostic ticks were placed into BSK medium to obtain isolates. Isolates from rodents and rodent-feeding ticks were all identified as B. afzelii. The follow-up of the infectivity status of repeatedly recaptured rodents clearly demonstrated that these hosts remained infective for ticks during winter till the following spring. Comparing C. glareolus and A. sylvaticus, each rodent species showed different host infection, different host infectivity and contributed differently to the moulting success of feeding ticks. These factors influence differentially the pattern of transmission of B. afzelii from Clethrionomys voles and Apodemus mice to I. ricinus ticks.

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PubMed1996

Detection of four species of Borrelia burgdorferi sensu lato in Ixodes ricinus ticks collected from roe deer (Capreolus capreolus) in The Netherlands.

Roe deer (Capreolus capreolus) were investigated for their value as sentinel animals for Lyme borreliosis in the Netherlands. Serum was obtained from 114 roe deer, and 513 Ixodes ricinus, predominantly females (72%), were obtained from 47 animals (41%). The polymerase chain reaction was used to detect DNA of Borrelia burgdorferi sensu lato in a total of 190 ticks, comprising 106 engorged ticks and 84 non-engorged ticks. Borrelia DNA was detected in 24 engorged ticks (23%) and 26 non-engorged ticks (31%). This difference was not significant (P = 0.25). Four species of B. burgdorferi sensu lato were identified in the ticks. B. burgdorferi sensu stricto, Borrelia garinii, Borrelia afzelii and group VS116. B. afzelii was most commonly found and present in 13 mixed infections, and in 28 single infections. Fifteen sera (13%) contained antibodies to Borrelia spp. Ticks are more appropriate sentinel animals for Lyme borreliosis than roe deer, an important host for I. ricinus. Although the viability of borrelia spirochaetes in engorged ticks collected from roe deer was not assessed, a bloodmeal taken from roe deer did not eliminate borrelia spirochaetes from the tick. The relevance of this finding for transovarial transmission of borrelia spirochaetes in ticks is discussed.

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PubMed2002

Detection of four Borrelia burgdorferi genospecies and first report of human granulocytic ehrlichiosis agent in Ixodes ricinus ticks collected in central Italy.

The presence of Borrelia burgdorferi s.l. and of Ehrlichia phagocytophila group was sought by PCR in Ixodes ricinus collected in a protected area of central Italy. Nymphs (n = 1475, gathered in 295 pools of 5 nymphs each) and adult ticks (n = 28) were examined. B. burgdorferi s.l. was detected in 13.8% of the nymph pools; of these, 63.4% were infected by B. valaisiana, 26.8% by B. afzelii, 7.3% by B. garinii, and 2.5% by B. burgdorferi s.s. Only a single adult male tick proved to host B. afzelii. The agent of human granulocytic ehrlichiosis (HGE) was detected in 2.7% of the nymph pools. Two HGE agent-positive nymph pools were also found to be positive for B. garinii and for B. afzelii, respectively. This is the first report from central Italy of the finding of the HGE agent in ticks.

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PubMed1998

Optimal sampling and spatial distribution of Ixodes pacificus, Dermacentor occidentalis and Dermacentor variabilis ticks (Acari: Ixodidae).

A common method for sampling tick populations is flagging, which is a method of dragging a white cloth over a plant substrate for a fixed distance along a transect. Flagging over rough physical surfaces or using long subtransect lengths could lead to the underestimation of tick densities. Using estimates of the drop-off rates of adult Ixodes pacificus in flag sampling, optimal sampling schemes (the length and number of subtransects) were examined using the relationships between the tick drop-off rates (c), the tick density and distribution patterns and the roughness of the sampled plant substrate. It was found that the optimal number of subtransects and Lmax, the longest subtransect length which did not significantly underestimate the tick density from c, were affected by the tick density, substrate roughness and tick distribution pattern. This study also showed that the density and distribution of I. pacificus and Dermacentor occidentalis varied greatly over time in the populations sampled, while the Dermacentor variabilis densities were low and showed no significant changes over time. Both I. pacificus and D. occidentalis had clumped distributions along trails and these clumps were aggregated. However, the clump sizes (or individual clump areas) changed significantly over time because of density fluctuation or the movement of ticks. Finally, a positive association between the number of I. pacificus and D. occidentalis adults was observed from 2 m subtransect flag collections in March 1995; no relationship was found in 1994.

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PubMed1999

Density of deer in relation to the prevalence of Borrelia burgdorferi s.l. in Ixodes ricinus nymphs in Rambouillet forest, France.

The Rambouillet Forest, a Lyme disease-endemic area near Paris, France, was surveyed from September 1994 to October 1995 to determine the risk periods and zones for humans. Firstly, during the period of Ixodes ricinus activity, abundance of nymphs is greater in spring than in autumn. Secondly, we observed significant variation in nymphal abundance between zones according to the density of cervids. The polymerase chain reaction (PCR) was used to detect DNA of Borrelia burgdorferi sensu lato in 461 unfed nymphs. DNA was detected in 38 nymphs (8.2%). By genospecific PCR based on the OspA gene, we detected the three pathogenic spirochetes with occurrences of 10.3, 31.1 and 58.6 for B. burgdorferi s.s., Borrelia garinii and Borrelia afzelii, respectively, indicating that B. afzelii is probably the main Borrelia species in the Rambouillet Forest. Finally, 11.5% of positive nymphs exhibited a double infection. Infection rates of I. ricinus nymphs by B. burgdorferi s.l. were not significantly different throughout the year for a given area, indicating that the risk periods of acquiring Lyme disease are mainly linked to nymph activity and correspond to spring and autumn. Likewise infection rates of nymphs were not significantly different between zones with a high density of deer (more than 100 animals per 100 ha) and zones with lower deer density (less than 20 animals per 100 ha). In addition to the role of deer as an amplifier of tick populations, these data indicate that zones with a high density of cervids should be considered as higher risk areas.

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PubMed1998

Predictive map of Ixodes ricinus high-incidence habitats and a tick-borne encephalitis risk assessment using satellite data.

The main objective of this project was to predict Ixodes ricinus abundant habitats reliably as a means of tick-borne encephalitis (TBE) risk assessment for the prevention of this disease. The vegetation types were used as the indicators of an ecosystem suitable for tick occurrence, for TBE virus circulation and, accordingly, for the existence of natural foci of this infection. Remote sensing methods were used to determine the indicative plant cover. Satellite data covering an experimental area of 70 x 70 km in Central Bohemia, the Czech Republic, was acquired by the Landsat 5 TM scanner. Nine forest classes were recognized in the experimental area by successive supervised and unsupervised classifications and identified in a field-checking botanical survey. An epidemiological TBE map based on human cases contracted in the territory under study was exploited for the evaluation of risk in particular forest classes. Predictive maps are expressed both in digital and in printed forms at a scale of 1:300,000 for an overall risk evaluation and at a scale of 1:25,000 for a detailed local orientation.

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PubMed1998

Interstadial variation in the attachment sites of Ixodes ricinus ticks on sheep.

The spatial aggregation of ticks feeding on vertebrate hosts has been recognized for some time but, for hosts supporting more than one stage of the tick, observations of interstadial variation in the site of attachment have not previously been quantified. This study showed that all three parasitic stages of Ixodes ricinus ticks feeding on sheep attach most commonly to the hair-covered areas of the head and limbs while few ticks attach to the fleeced region of the body. However, significant differences were observed in the site of attachment of the three feeding stages of the tick. Larvae attached to distal limbs and rostral areas of the head and adult females attached to the proximal areas of the limbs and around the neck and ears, while nymphs attached in locations between the larvae and adults. The importance of the spatial aggregation of the ticks and interstadial variation in their distribution on the host, for the transmission of tick-borne pathogens and the epidemiology of the diseases they cause, is discussed.

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PubMed1998

The question of fluctuating asymmetry in the blacklegged tick Ixodes scapularis (Acari: Ixodidae).

An analysis of fluctuating asymmetry was conducted on populations of the blacklegged tick Ixodes scapularis. The eight groups used in this study consisted of larvae and nymphs and males and females from the states of Minnesota, Massachusetts, Maryland, Missouri, North Carolina and Georgia and the F1 progenies of reciprocal crosses between ticks from Massachusetts and Georgia. Measurements included 16 larval, 19 nymphal, ten female and 12 male bilateral characters. Only five differences between the right and left bilateral characters had normal distributions with means of zero and differences in variances between the groups. These five characters included three setal lengths of the larvae, the spiracular plate length of females and the coxa I internal spur widths of males. Bivariate plots of character size ((R + L)/2) and asymmetry (R-L) showed no correlation. In the spiracular plate lengths of females and one of the setal lengths, ticks from Massachusetts had significantly less within-group variance than all the other groups. The only character in which fluctuating asymmetry was observed was the coxa I internal spur width of males, in which ticks from Minnesota, Missouri and North Carolina had significantly greater variance than the remaining groups; fluctuating asymmetry in this character may be explained by sexual selection. The cross progeny did not demonstrate any fluctuating asymmetry, as would be expected if the northern and southern forms of I. scapularis were true species. The virtual lack of fluctuating asymmetry in the characters used in this study further supports the conclusions of other studies which concluded that I. scapularis is a species with clinical variation and a broad geographic distribution.

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PubMed1999

Mosaic pattern of Borrelia infection in a continuous population of the tick Ixodes ricinus (Acari: Ixodidae).

An array of 12 20 x 20 m quadrats in a mixed forest near Poteply, Central Bohemia, Czech Republic, was investigated for the abundance and spatial distribution of host-questing Ixodes ricinus nymphs and their infection with borreliae. Tick densities were estimated by flagging and their borrelia infection status was determined by direct immunofluorescence. While the tick population appeared to be continuous and homogeneous based on quadrat counts, their infection with borreliae evinced a mosaic pattern. The nymphal infection rates ranged between 1.6 and 10.5% (mean = 6.0%) with significant differences between adjacent quadrats. Home ranging of small rodents (Apodemus flavicollis and Clethrionomys glareolus) inhabiting the forest seems to be responsible for the spatial pattern of borrelial infection.

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PubMed1999

Borrelia burgdorferi sensu lato in Ixodes ricinus ticks and rodents in a recreational park in south-western Ireland.

Ixodes ricinus ticks infected with Borrelia burgdorferi sensu lato were numerous on the edges of paths and roads in a recreational park in south-western Ireland. The abundance of ticks at different sites was related to the presence of deer, but a negative relationship was shown between tick abundance and tick infection rates. This is thought to be due to the deposition of large numbers of uninfected ticks by deer, which are apparently not good reservoir hosts of B. burgdorferi s.l. Blood meal analysis only detected deer DNA in uninfected nymphs. Reservoir competent rodents, Apodemus sylvaticus and Clethrionomys glareolus, were abundant at all sites and a high proportion of captured specimens were infested with larval ticks. However, very few rodents were infected with B. burgdorferi s.l. and none of the unfed infected nymphs analysed for the identity of their larval blood meal had fed on rodents. The spirochaetes detected in I. ricinus in the study area may be poorly adapted to rodents or are not transmitted readily because of the absence of nymphal infestation. The majority of spirochaetes in these ticks were apparently acquired from non-rodent hosts, such as birds.

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PubMed2000

Vector competence of Ixodes angustus (Acari: Ixodidae) for Borrelia burgdorferi sensu stricto.

The vector competence of Ixodes angustus for Borrelia burgdorferi sensu stricto (s.s.) was investigated in the laboratory. The larval progeny of female ticks from Washington State were placed on Swiss-Webster mice that had been inoculated intravenously with 10(8) spirochetes each of a Californian isolate of B. burgdorferi. Spirochetes were detected in 6 (12%) of 50 nymphs derived from larvae that had fed on these animals. Ten nymphs from the same cohort of experimentally infected ticks were placed on each of 4 naive deer mice (Peromyscus maniculatus). One of the mice seroconverted to B. burgdorferi and spirochetes were isolated from its ear tissues 4 weeks after exposure to ticks. Further vector competence trials were conducted with I. angustus ticks from California. Larvae were fed on deer mice that had been inoculated intradermally with B. burgdoferi along with larvae of I. spinipalpis as a comparison group. There was no significant difference in the prevalence of infection in nymphs of I. angustus (8.2%) versus those of I. spinipalpis (12.1%). We conclude that I. angustus is a competent experimental vector of B. burgdorferi s.s. and its efficiency for acquiring and transstadially passing such spirochetes is similar to that of I. spinipalpis.

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PubMed1998

Antigenic and genomic analysis of a Borrelia burgdorferi sensu stricto strain isolated from Ixodes ricinus ticks in Alto Adige-South Tyrol, Italy.

A Borrelia burgdorferi sensu lato strain isolated from IXodes ricinus ticks in Alto Adige-South Tyrol (Northern Italy) was analyzed by sodium dodecyl sulfate-polyacrylamide gel electrophoresis (SDS-PAGE) of whole cell proteins, Western immunoblotting analysis (WBA) with polyclonal and monoclonal antibodies, and pulsed-field gel electrophoresis (PFGE). The isolate named BZ6 was identified as belonging to the genospecies B. burgdorferi sensu stricto on the basis of its protein profile and its reactivity with monoclonal and polyclonal antibodies. The PFGE study performed with the two rare-cutting restriction enzymes MluI and SmaI confirmed the SDS-PAGE and WBA characterizations, but showed a genetic diversity between the isolate and two out of the three B. burgdorferi sensu stricto strains used in this study as controls, the American type strain B31 and the locally isolated strain BZ1. No difference in the PFGE patterns between the isolate BZ6 and the Swiss strain IRS was noted. Our findings show the value of PFGE analysis for classifying B. burgdorferi sensu lato isolates and for revealing their genetic diversity, and its usefulness for epidemiological investigations.

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PubMed2000

Occurrence of Borrelia burgdorferi sensu stricto, Borrelia garinii and Borrelia afzelii in the Ixodes ricinus ticks from Eastern Slovakia.

A total of 2816 unfed adults nymphs of Ixodes ricinus ticks were collected from vegetation in Kosice (Eastern Slovakia) from 1994 to 1997. Prevalence of Borrelia burgdorferi s. l. in I. ricinus ticks, detected by dark field microscopy, varied and depended upon the year and the habitat of the collected ticks. The lowest prevalence was observed in 1994 (4.8%). During 1995 it increased to 17.2% and during the next two years decreased to 15.5% and 14.2%. The rate of infection varied from 2.1 to 23.3% within 10 examined habitats of the Kosice area. A different value of relative density of ticks was observed in various habitats. It ranged from 9-212 ticks per collecting hour within one flagged area (600 m2) which is 1.5-35.5 ticks per 100 m2. Eight isolates were obtained from the infected ticks. Electrophoresis and immunoblotting with 6 monoclonal antibodies were used for the identification of Borrelia strains. Three tick isolates were identified as B. burgdorferi s. s. and the other three isolates were found to be B. garinii. One strain reacted as a mixed culture of B. burgdorferi s. s., and B. garinii. The strain originated from the Vihorlat Mountains habitat and was detected by PCR-SSCP as B. burgdorferi s. s. with a small amount of B. afzelii. The obtained results emphasize the epidemiological importance not only of B. garinii and B. afzelii but also of B. burgdorferi s. s. in Central Europe.

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PubMed1999

Genetic diversity of Borrelia burgdorferi sensu lato isolates obtained from Ixodes ricinus ticks collected in Slovakia.

In Europe, Borrelia burgdorferi sensu lato is diverse, including B. burgdorferi s.s., B. garinii, B. afzelii, B. valaisiana and B. lusitaniae. In this study, we focused on the distribution of the different B. burgdorferi species among Ixodes ricinus adult ticks collected in an endemic area within Slovakia. We compared results of prevalence of B. burgdorferi infection in ticks obtained by immunofluorescence (IF) and by isolation. Isolates were characterized by restriction fragment length polymorphism (RFLP) of the rrf-rrl intergenic spacer genes using MseI. Using immunofluorescence we observed that 56/114 (49%) ticks were infected by B. burgdorferi s.l. Males were found to be more often infected (32/57, 56%) than females (24/57, 42%) but the difference was not significant (p = 0.1895). From the same 114 ticks a total of 37 isolates were obtained: 19 from males (33%) and 18 from females (32%). The RFLP identification revealed 25 B. afzelii (68%), 5 B. garinii (14%), 5 B. valaisiana (14%) and 2 B. lusitaniae (5%). The infection in ticks was more often detected by IF than by isolation (p = 0.0153) and isolation success was higher when the infection degree in ticks was high (p = 0.0397). The infection prevalence observed in this area is among the highest observed in Europe.

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PubMed2000

Temporal distribution of the annual nymphal stock of Ixodes ricinus ticks.

The human risk of contracting Lyme disease or other tick borne diseases transmitted by the tick species Ixodes ricinus is broadly linked to the tick nymph density. The study was performed in Rambouillet forest (Yvelines, France), a known focus of Lyme borreliosis, from January 1997 to December 1999. We used a nymph sampling methodology which permitted us to obtain a monthly nymph density index (from 0 to 5). Studying the seasonal nymph and larval activity patterns and estimating the larval developmental duration, we demonstrate the existence of an annual nymphal stock. Secondly, we elucidate how this stock is distributed throughout the year, month by month. Its distribution is principally dependent on two factors: the monthly mean ambient temperature and the proportion of active nymphs which find a host each month. Expected monthly nymph densities derived from a theoretical model describing the temperature-dependent stock distribution gave a good fit to the observed densities, accounting for between 76-86% of the monthly variation in observed nymph densities. Predicting the temporal distribution of nymph activity within a stable Lyme borreliosis focus enables more precise identification of risk periods.

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PubMed2000

Seasonal population dynamics of ixodes ticks and tick-borne encephalitis virus.

Seasonality of the epidemic and epizootic processes of tick-borne encephalitis (TBE) depend on the period of activity of ixodid ticks Ixodes persulcatus Schulze and I. ricinus Linnaeus, which are the main reservoirs and vectors of TBE virus, and also on the process of their activation. The period of activity is the period during which the ticks occur in the active state. Activation is the transition into this state of ticks that moulted from the preceding stage and completed post-moulting development. For I. persulcatus, the first adult ticks generally emerge between April 10 and May 9. Under a variety of natural conditions, activation of adult I. persulcatus after wintering lasts for 45-86 days and this period may be even longer in certain areas of the Far East. The period during which one-half of the entire tick population becomes activated (AT50) comprises no more than 10-20 days. In adult I. ricinus ticks the activation period may last even longer than in I. persulcatus. The data on duration of the period of activity and on activation of larval and nymphal stages of both tick species were considered. Ticks exhausting their nutrient reserves and failing to find a host die quickly. The period during which 50% of the entire tick population die under natural conditions is designated LT50. The main types of I. persulcatus and I. ricinus seasonal activity within their species ranges were reviewed. Data on the relationship between TBE virus reproduction in a natural focus and physiological age, pattern of activation, and seasonal changes in age structure of the tick population were analyzed. Seasonal changes in the prevalence of infection among active unfed adult ticks in a natural population are determined by virus content in individual ticks at the moment of their activation and also by the duration of subsequent virus persistence (the rate of virus loss) in ticks. Apparently, the opportunity and frequency of horizontal TBE virus transmission under natural conditions, change during the season of tick activity.

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