BackgroundPathogens transmitted by ticks cause human disease on a greater scale than any other vector-borne infections in Europe, and have increased dramatically over the past 2–3 decades. Reliable records of tick-borne encephalitis (TBE) since 1970 show an especially sharp upsurge in cases in Eastern Europe coincident with the end of Soviet rule, including the three Baltic countries, Estonia, Latvia and Lithuania, where national incidence increased from 1992 to 1993 by 64, 175 and 1,065%, respectively. At the county level within each country, however, the timing and degree of increase showed marked heterogeneity. Climate has also changed over this period, prompting an almost universal assumption of causality. For the first time, we analyse climate and TBE epidemiology at sufficiently fine spatial and temporal resolution to question this assumption.Methodology/Principal FindingDetailed analysis of instrumental records of climate has revealed a significant step increase in spring-time daily maximum temperatures in 1989. The seasonal timing and precise level of this warming were indeed such as could promote the transmission of TBE virus between larval and nymphal ticks co-feeding on rodents. These changes in climate, however, are virtually uniform across the Baltic region and cannot therefore explain the marked spatio-temporal heterogeneity in TBE epidemiology.Conclusions/SignificanceInstead, it is proposed that climate is just one of many different types of factors, many arising from the socio-economic transition associated with the end of Soviet rule, that have acted synergistically to increase both the abundance of infected ticks and the exposure of humans to these ticks. Understanding the precise differential contribution of each factor as a cause of the observed epidemiological heterogeneity will help direct control strategies.
Tick-borne encephalitis (TBE), the most serious widespread vector-borne disease of humans in Europe, increased from 2- to 30-fold in many Central and Eastern European countries from 1992 to 1993, coinciding with independence from Soviet rule. Unemployment and low income have been shown in Latvia to be statistically associated with high-risk behaviour involving harvest of wild foods from tick-infested forests, and also with not being vaccinated against TBE. Archival data for 1970--2005 record major changes in the agricultural and industrial sectors, and consequent changes in the abiotic and biotic environment and socio-economic conditions, which could have increased the abundance of infected ticks and the contact of humans with those ticks. For example, abandoned agricultural fields became suitable for rodent transmission hosts; use of pesticides and emissions of atmospheric industrial pollutants plummeted; wildlife hosts for ticks increased; tick populations appear to have responded; unemployment and inequality increased in all countries. These factors, by acting synergistically but differentially between and within each country, can explain the marked spatio-temporal heterogeneities in TBE epidemiology better than can climate change alone, which is too uniform across wide areas. Different degrees of socio-economic upheaval caused by political transition in Estonia, Latvia, Lithuania, Slovenia and the Czech Republic can apparently explain the marked variation in TBE upsurge. Causal linkage between national socio-economic conditions and epidemiology is strongly indicated by striking correlations across eight countries between the degree of upsurge of TBE and both poverty and household expenditure on food (R2 = 0.533 and 0.716, respectively).
The data suggest that the TBE spike was not due to weather-induced variation in tick population dynamics. An alternative explanation, supported by qualitative reports and some data, involves human behavioural responses to weather favourable for outdoor recreational activities, including wild mushroom and berry harvest, differentially influenced by national cultural practices and economic constraints.
Borrelia garinii, a causative agent of Lyme borreliosis in Europe and Asia, is naturally maintained in marine and terrestrial enzootic cycles, which primarily involve birds, including seabirds and migratory passerines. These bird groups associate with, correspondingly, Ixodes uriae and Ixodes ricinus ticks, of which the latter species may bite and transmit the infection to humans. Studies of the overlap between these two natural cycles of B. garinii have been limited, in part due to the absence of representative collections of this spirochete's samples, as well as of the lack of reliable measure of the genetic heterogeneity of its strains. As a prerequisite for understanding the epidemiological correlates of the complex maintenance of B. garinii, the present study sought to assess the diversity and phylogenetic relationships of this species' strains from its natural hosts and patients with Lyme borreliosis from subarctic Eurasia. We used sequence typing of the partial rrs-rrl intergenic spacer (IGS) of archived and prospective samples of B. garinii from I. uriae ticks collected predominantly on Commander Islands in North Pacific, as well as on the islands in northern Sweden and arctic Norway. We also typed B. garinii samples from patients with Lyme borreliosis and I. ricinus ticks infesting migratory birds in southern Sweden, or found questing in selected sites on the islands in the Baltic Sea and Lithuania. Fifty-two (68%) of 77 B. garinii samples representing wide geographical range and associated with I. ricinus and infection of humans contributed 12 (60%) of total 20 identified IGS variants. In contrast, the remaining 25 (32%) samples recovered from I. uriae ticks from a few islands accounted for as many as 10 (50%) IGS types, suggesting greater local diversity of B. garinii maintained by seabirds and their ticks. Two IGS variants of the spirochete in common for both tick species were found in I. ricinus larvae from migratory birds, an indication that B. garinii strains are exchanged between different ecological niches. Notably, B. garinii variants associated with I. uriae ticks were found in each of the six clusters, representing two phylogenetic lineages of this species identified among the studied samples. Our findings suggest that B. garinii in subarctic Eurasia comprises two partially overlapping populations with different levels of genetic heterogeneity, presumably, due to distinctive selective pressures on the spirochete in its marine and terrestrial enzootic cycles.
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