A Quantitative Prioritisation of Human and Domestic Animal Pathogens in Europe

Disease or pathogen risk prioritisations aid understanding of infectious agent impact within surveillance or mitigation and biosecurity work, but take significant development. Previous work has shown the H-(Hirsch-)index as an alternative proxy. We present a weighted risk analysis describing infectious pathogen impact for human health (human pathogens) and well-being (domestic animal pathogens) using an objective, evidence-based, repeatable approach; the H-index. This study established the highest H-index European pathogens. Commonalities amongst pathogens not included in previous surveillance or risk analyses were examined. Differences between host types (humans/animals/zoonotic) in pathogen H-indices were explored as a One Health impact indicator. Finally, the acceptability of the H-index proxy for animal pathogen impact was examined by comparison with other measures. 57 pathogens appeared solely in the top 100 highest H-indices (1) human or (2) animal pathogens list, and 43 occurred in both. Of human pathogens, 66 were zoonotic and 67 were emerging, compared to 67 and 57 for animals. There were statistically significant differences between H-indices for host types (humans, animal, zoonotic), and there was limited evidence that H-indices are a reasonable proxy for animal pathogen impact. This work addresses measures outlined by the European Commission to strengthen climate change resilience and biosecurity for infectious diseases. The results include a quantitative evaluation of infectious pathogen impact, and suggest greater impacts of human-only compared to zoonotic pathogens or scientific under-representation of zoonoses. The outputs separate high and low impact pathogens, and should be combined with other risk assessment methods relying on expert opinion or qualitative data for priority setting, or could be used to prioritise diseases for which formal risk assessments are not possible because of data gaps.     

A stakeholder-informed approach to the identification of criteria for the prioritization of zoonoses in Canada

Background

Zoonotic diseases account for over 60% of all communicable diseases causing illness in humans and 75% of recently emerging infectious diseases. As limited resources are available for the control and prevention of zoonotic diseases, it is necessary to prioritize diseases in order to direct resources into those with the greatest needs. The selection of criteria for prioritization has traditionally been on the basis of expert opinion; however, details of the methods used to identify criteria from expert opinion often are not published and a full range of criteria may not be captured by expert opinion.

Methodology/Principal Findings

This study used six focus groups to identify criteria for the prioritization of zoonotic diseases in Canada. Focus groups included people from the public, animal health professionals and human health professionals. A total of 59 criteria were identified for prioritizing zoonotic diseases. Human-related criteria accounted for the highest proportion of criteria identified (55%), followed by animal-related criteria (26%) then pathogen/disease-related criteria (19%).Similarities and differences were observed in the identification and scoring of criteria for disease prioritization between groups; the public groups were strongly influenced by the individual-level of disease burden, the responsibility of the scientific community in disease prioritization and the experiences of recent events while the professional groups were influenced by the societal- and population-level of disease burden and political and public pressure.

Conclusions/Significance

This was the first study to describe a mixed semi-quantitative and qualitative approach to deriving criteria for disease prioritization. This was also the first study to involve the opinion of the general public regarding disease prioritization. The number of criteria identified highlights the difficulty in prioritizing zoonotic diseases. The method presented in this paper has formulated a comprehensive list of criteria that can be used to inform future disease prioritization studies.     

Modeling climate impact on an emerging disease,the Phytophthora alni‐induced alder decline

Alder decline caused by Phytophthora alni is one of the most important emerging diseases in natural ecosystems in Europe, where it has threatened riparian ecosystems for the past 20 years. Environmental factors, such as mean site temperature and soil characteristics, play an important role in the occurrence of the disease. The objective of the present work was to model and forecast the effect of environment on the severity of alder Phytophthora outbreaks, and to determine whether recent climate change might explain the disease emergence. Two alder sites networks in NE and SW France were surveyed to assess the crown health of trees; the oomycete soil inoculum was also monitored in the NE network. The main factors explaining the temporal annual variation in alder crown decline or crown recovery were the mean previous winter and previous summer temperatures. Both low winter temperatures and high summer temperatures were unfavorable to the disease. Cold winters promoted tree recovery because of poor survival of the pathogen, while hot summer temperature limited the incidence of tree decline. An SIS model explaining the dynamics of the P. alni‐induced alder decline was developed using the data of the NE site network and validated using the SW site network. This model was then used to simulate the frequency of declining alder over time with historical climate data. The last 40 years' weather conditions have been generally favorable to the establishment of the disease, indicating that others factors may be implicated in its emergence. The model, however, showed that the climate of SW France was much more favorable for the disease than that of the Northeast, because it seldom limited the overwintering of the pathogen. Depending on the European area, climate change could either enhance or decrease the severity of the alder decline.     

Altitudinal patterns of tick and host abundance: a potential role for climate change in regulating tick-borne diseases?

The impact of climate change on vector-borne infectious diseases is currently controversial. In Europe the primary arthropod vectors of zoonotic diseases are ticks, which transmit Borrelia burgdorferi sensu lato (the agent of Lyme disease), tick-borne encephalitis virus and louping ill virus between humans, livestock and wildlife. Ixodes ricinus ticks and reported tick-borne disease cases are currently increasing in the UK. Theories for this include climate change and increasing host abundance. This study aimed to test how I. ricinus tick abundance might be influenced by climate change in Scotland by using altitudinal gradients as a proxy, while also taking into account the effects of hosts, vegetation and weather effects. It was predicted that tick abundance would be higher at lower altitudes (i.e. warmer climates) and increase with host abundance. Surveys were conducted on nine hills in Scotland, all of open moorland habitat. Tick abundance was positively associated with deer abundance, but even after taking this into account, there was a strong negative association of ticks with altitude. This was probably a real climate effect, with temperature (and humidity, i.e. saturation deficit) most likely playing an important role. It could be inferred that ticks may become more abundant at higher altitudes in response to climate warming. This has potential implications for pathogen prevalence such as louping ill virus if tick numbers increase at elevations where competent transmission hosts (red grouse Lagopus lagopus scoticus and mountain hares Lepus timidus) occur in higher numbers.     

Host nutritional status: the neglected virulence factor

The emergence of new infectious diseases and old diseases with new pathogenic properties is a burgeoning worldwide problem. Severe acute respiratory syndrome (SARS) and acquired immune deficiency syndrome (AIDS) are just two of the most widely reported recent emerging infectious diseases. What are the factors that contribute to the rapid evolution of viral species? Various hypotheses have been proposed, all involving opportunities for virus spread (for example, agricultural practices, climate changes, rainforest clearing or air travel). However, the nutritional status of the host, until recently, has not been considered a contributing factor to the emergence of infectious disease. In this review, we show that host nutritional status can influence not only the host response to the pathogen, but can also influence the genetic make-up of the viral genome. This latter finding markedly changes our concept of host-pathogen interactions and creates a new paradigm for the study of such phenomena.     

Within-colony variation in inducibility of coral disease resistance

The abiotic environment influences a variety of ecological processes, including the emergence, transmission, and distribution of disease. In the oceans, increased temperatures associated with climate change are hypothesized to decrease host resistance and/or increase pathogen growth, virulence, or infectivity. Colonial organisms, such as corals, could face a unique challenge with respect to temperature and disease stress: heterogeneous within-colony distribution of constitutive and temperature-induced resistance to infection. This could facilitate disease if warming temperatures promote pathogen growth while decreasing resistance of some areas of the coral colony. Here, an experiment was used to test the hypothesis that temperature-induced disease resistance is heterogeneous within colonies of the sea fan coral, Gorgonia ventalina. Resistance, measured as activity of antifungal metabolites, increased (approx. 30%) with temperature only in young edge tissue, not in older center tissue, consistent with patterns of infection in older, larger sea fan colonies on Caribbean reefs.     

Downy mildew (Plasmopara viticola) epidemics on grapevine under climate change

As climate is a key agro‐ecosystem driving force, climate change could have a severe impact on agriculture. Many assessments have been carried out to date on the possible effects of climate change (temperature, precipitation and carbon dioxide concentration changes) on plant physiology. At present however, likely effects on plant pathogens have not been investigated deeply. The aim of this work was to simulate future scenarios of downy mildew (Plasmopara viticola) epidemics on grape under climate change, by combining a disease model to output from two general circulation models (GCMs). Model runs corresponding to the SRES‐A2 emissions scenario, characterized by high projections of both population and greenhouse gas emissions from present to 2100, were chosen in order to investigate impacts of worst‐case scenarios, among those currently available from IPCC. Three future decades were simulated (2030, 2050, 2080), using as baseline historical series of meteorological data collected from 1955 to 2001 in Acqui Terme, an important grape‐growing area in the north‐west of Italy. Both GCMs predicted increase of temperature and decrease of precipitation in this region. The simulations obtained by combining the disease model to the two GCM outputs predicted an increase of the disease pressure in each decade: more severe epidemics were a direct consequence of more favourable temperature conditions during the months of May and June. These negative effects of increasing temperatures more than counterbalanced the effects of precipitation reductions, which alone would have diminished disease pressure. Results suggested that, as adaptation response to future climate change, more attention would have to be paid in the management of early downy mildew infections; two more fungicide sprays were necessary under the most negative climate scenario, compared with present management regimes. At the same time, increased knowledge on the effects of climate change on host–pathogen interactions will be necessary to improve current predictions.     

Interannual patterns of avian diseases in wild New Zealand avifauna near conservation areas

The past few years have seen a noticeable increase in the emergence of infectious diseases in wildlife, especially vector-borne diseases, presenting a challenge for the conservation of endangered species. One such vector-borne disease, avian malaria (Plasmodium spp.) is on the rise in New Zealand avifauna, threatening bird populations that are among the most extinction-prone in the world. Furthermore, recent reports have outlined an increase in deaths of native iconic bird species specifically due to this disease. In order to help manage breakouts of this pathogen at a local scale, we need a better understanding of potential drivers of the emergence of avian malaria in wild New Zealand avifauna. Here, we set to test the role of climatic drivers in synchronizing contacts between avian hosts and vectors, assess the temporal stability of transmission dynamics between years, and determine the role of introduced species in causing spill-over of this pathogen towards native species. Our study focused on three sites that were sampled regularly during two consecutive years in the austral summer, each site being adjacent to a breeding colony of Yellow-eyed penguins (Megadyptes antipodes). Our results reveal an overall temporal stability of avian malaria incidence patterns, with a decrease in infection throughout the austral summer for both sampled years. Moreover, we highlight a phylogenetic signal among sampled bird species, with introduced species being more heavily infected by avian malaria than their native counterparts. In contrast, we found no effect of the two climatic drivers investigated, temperature and rainfall, on mosquito abundance. Our results suggest a strong effect of alien species acting as reservoirs for diseases spilling-over towards immunologically naïve species, and provide conservation managers with a critical timeframe to control avian malaria breakouts.     

Climate change accelerates local disease extinction rates in a long‐term wild host–pathogen association

Pathogens are a significant component of all plant communities. In recent years, the potential for existing and emerging pathogens of agricultural crops to cause increased yield losses as a consequence of changing climatic patterns has raised considerable concern. In contrast, the response of naturally occurring, endemic pathogens to a warming climate has received little attention. Here, we report on the impact of a signature variable of global climate change – increasing temperature – on the long‐term epidemiology of a natural host–pathogen association involving the rust pathogen Triphragmium ulmariae and its host plant Filipendula ulmaria. In a host–pathogen metapopulation involving approximately 230 host populations growing on an archipelago of islands in the Gulf of Bothnia we assessed changes in host population size and pathogen epidemiological measures over a 25‐year period. We show how the incidence of disease and its severity declines over that period and most importantly demonstrate a positive association between a long‐term trend of increasing extinction rates in individual pathogen populations of the metapopulation and increasing temperature. Our results are highly suggestive that changing climatic patterns, particularly mean monthly growing season (April‐November) temperature, are markedly influencing the epidemiology of plant disease in this host–pathogen association. Given the important role plant pathogens have in shaping the structure of communities, changes in the epidemiology of pathogens have potentially far‐reaching impacts on ecological and evolutionary processes. For these reasons, it is essential to increase understanding of pathogen epidemiology, its response to warming, and to invoke these responses in forecasts for the future.     

Implications of host genetic variation on the risk and prevalence of infectious diseases transmitted through the environment

Previous studies have shown that host genetic heterogeneity in the response to infectious challenge can affect the emergence risk and the severity of diseases transmitted through direct contact between individuals. However, there is substantial uncertainty about the degree and direction of influence owing to different definitions of genetic variation, most of which are not in line with the current understanding of the genetic architecture of disease traits. Also, the relevance of previous results for diseases transmitted through environmental sources is unclear. In this article a compartmental genetic-epidemiological model was developed to quantify the impact of host genetic diversity on epidemiological characteristics of diseases transmitted through a contaminated environment. The model was parameterized for footrot in sheep. Genetic variation was defined through continuous distributions with varying shape and degree of dispersion for different disease traits. The model predicts a strong impact of genetic heterogeneity on the disease risk and its progression and severity, as well as on observable host phenotypes, when dispersion in key epidemiological parameters is high. The impact of host variation depends on the disease trait for which variation occurs and on environmental conditions affecting pathogen survival. In particular, compared to homogeneous populations with the same average susceptibility, disease risk and severity are substantially higher in populations containing a large proportion of highly susceptible individuals, and the differences are strongest when environmental contamination is low. The implications of our results for the recording and analysis of disease data and for predicting response to selection are discussed.     

The effect of climate change on the occurrence and prevalence of livestock diseases in Great Britain: a review

There is strong evidence to suggest that climate change has, and will continue to affect the occurrence, distribution and prevalence of livestock diseases in Great Britain (GB). This paper reviews how climate change could affect livestock diseases in GB. Factors influenced by climate change and that could affect livestock diseases include the molecular biology of the pathogen itself; vectors (if any); farming practice and land use; zoological and environmental factors; and the establishment of new microenvironments and microclimates. The interaction of these factors is an important consideration in forecasting how livestock diseases may be affected. Risk assessments should focus on looking for combinations of factors that may be directly affected by climate change, or that may be indirectly affected through changes in human activity, such as land use (e.g. deforestation), transport and movement of animals, intensity of livestock farming and habitat change. A risk assessment framework is proposed, based on modules that accommodate these factors. This framework could be used to screen for the emergence of unexpected disease events.     

Climate,vegetation, introduced hosts and trade shape a global wildlife pandemic

Global factors, such as climate change, international trade and introductions of exotic species are often elicited as contributors to the unprecedented rate of disease emergence, but few studies have partitioned these factors for global pandemics. Although contemporary correlative species distribution models (SDMs) can be useful for predicting the spatial patterns of emerging diseases, they focus mainly on the fundamental niche (FN) predictors (i.e. abiotic climate and habitat factors), neglecting dispersal and propagule pressure predictors (PP, number of non-native individuals released into a region). Using a validated, predictive and global SDM, we show that both FN and PP accounted for significant, unique variation to the distribution of the chytrid fungus Batrachochytrium dendrobatidis (Bd), a pathogen implicated in the declines and extinctions of over 200 amphibian species worldwide. Bd was associated positively with vegetation, total trade and introduced amphibian hosts, nonlinearly with annual temperature range and non-significantly with amphibian leg trade or amphibian species richness. These findings provide a rare example where both FN and PP factors are predictive of a global pandemic. Our model should help guide management of this deadly pathogen and the development of other globally predictive models for species invasions and pathogen emergence influenced by FN and PP factors.     

Extreme climatic events and host–pathogen interactions: The impact of the 1976 drought in the UK

Intense, long droughts have increased in occurrence since the 1970s and have been linked with global climate change. Extreme climate alters the risk of pathogen infections and diseases in both animals and plants, although little is known about the impact of any single event on host–pathogen dynamics in a wide range of species. Evaluating past climatic events can provide valuable information on complex interactions that occur between hosts, pathogens, and the environment, thereby paving the way for predictive models and ultimately early and efficient response to disease threats. The present study reviews the substantial impact of the 1976 UK drought on climate-driven host–pathogen associations. This 16-month drought had a devastating effect on flora and fauna and is considered a benchmark for dry conditions in this country. Changes to the occurrence of infections in farmed and wild animals and plants are presented in terrestrial, freshwater, and marine ecosystems and the implications for pathogen transmission under extreme climate conditions are assessed.     

Elicitation of Expert Prior Opinion: Application to the MYPAN Trial in Childhood Polyarteritis Nodosa

Objectives

Definitive sample sizes for clinical trials in rare diseases are usually infeasible. Bayesian methodology can be used to maximise what is learnt from clinical trials in these circumstances. We elicited expert prior opinion for a future Bayesian randomised controlled trial for a rare inflammatory paediatric disease, polyarteritis nodosa (MYPAN, Mycophenolate mofetil for polyarteritis nodosa).

Methods

A Bayesian prior elicitation meeting was convened. Opinion was sought on the probability that a patient in the MYPAN trial treated with cyclophosphamide would achieve disease remission within 6-months, and on the relative efficacies of mycophenolate mofetil and cyclophosphamide. Expert opinion was combined with previously unseen data from a recently completed randomised controlled trial in ANCA associated vasculitis.

Results

A pan-European group of fifteen experts participated in the elicitation meeting. Consensus expert prior opinion was that the most likely rates of disease remission within 6 months on cyclophosphamide or mycophenolate mofetil were 74% and 71%, respectively. This prior opinion will now be taken forward and will be modified to formulate a Bayesian posterior opinion once the MYPAN trial data from 40 patients randomised 1:1 to either CYC or MMF become available.

Conclusions

We suggest that the methodological template we propose could be applied to trial design for other rare diseases.     

Root pathogen diversity and composition varies with climate in undisturbed grasslands,but less so in anthropogenically disturbed grasslands

Soil-borne pathogens structure plant communities, shaping their diversity, and through these effects may mediate plant responses to climate change and disturbance. Little is known, however, about the environmental determinants of plant pathogen communities. Therefore, we explored the impact of climate gradients and anthropogenic disturbance on root-associated pathogens in grasslands. We examined the community structure of two pathogenic groups—fungal pathogens and oomycetes—in undisturbed and anthropogenically disturbed grasslands across a natural precipitation and temperature gradient in the Midwestern USA. In undisturbed grasslands, precipitation and temperature gradients were important predictors of pathogen community richness and composition. Oomycete richness increased with precipitation, while fungal pathogen richness depended on an interaction of precipitation and temperature, with precipitation increasing richness most with higher temperatures. Disturbance altered plant pathogen composition and precipitation and temperature had a reduced effect on pathogen richness and composition in disturbed grasslands. Because pathogens can mediate plant community diversity and structure, the sensitivity of pathogens to disturbance and climate suggests that degradation of the pathogen community may mediate loss, or limit restoration of, native plant diversity in disturbed grasslands, and may modify plant community response to climate change.Subject terms: Fungal ecology, Soil microbiology, Grassland ecology     

Emergence and prevalence of human vector-borne diseases in sink vector populations

Vector-borne diseases represent a major public health concern in most tropical and subtropical areas, and an emerging threat for more developed countries. Our understanding of the ecology, evolution and control of these diseases relies predominantly on theory and data on pathogen transmission in large self-sustaining 'source' populations of vectors representative of highly endemic areas. However, there are numerous places where environmental conditions are less favourable to vector populations, but where immigration allows them to persist. We built an epidemiological model to investigate the dynamics of six major human vector borne-diseases in such non self-sustaining 'sink' vector populations. The model was parameterized through a review of the literature, and we performed extensive sensitivity analysis to look at the emergence and prevalence of the pathogen that could be encountered in these populations. Despite the low vector abundance in typical sink populations, all six human diseases were able to spread in 15-55% of cases after accidental introduction. The rate of spread was much more strongly influenced by vector longevity, immigration and feeding rates, than by transmission and virulence of the pathogen. Prevalence in humans remained lower than 5% for dengue, leishmaniasis and Japanese encephalitis, but substantially higher for diseases with longer duration of infection; malaria and the American and African trypanosomiasis. Vector-related parameters were again the key factors, although their influence was lower than on pathogen emergence. Our results emphasize the need for ecology and evolution to be thought in the context of metapopulations made of a mosaic of sink and source habitats, and to design vector control program not only targeting areas of high vector density, but working at a larger spatial scale.     

Interacting Symbionts and Immunity in the Amphibian Skin Mucosome Predict Disease Risk and Probiotic Effectiveness

Pathogenesis is strongly dependent on microbial context, but development of probiotic therapies has neglected the impact of ecological interactions. Dynamics among microbial communities, host immune responses, and environmental conditions may alter the effect of probiotics in human and veterinary medicine, agriculture and aquaculture, and the proposed treatment of emerging wildlife and zoonotic diseases such as those occurring on amphibians or vectored by mosquitoes. Here we use a holistic measure of amphibian mucosal defenses to test the effects of probiotic treatments and to assess disease risk under different ecological contexts. We developed a non-invasive assay for antifungal function of the skin mucosal ecosystem (mucosome function) integrating host immune factors and the microbial community as an alternative to pathogen exposure experiments. From approximately 8500 amphibians sampled across Europe, we compared field infection prevalence with mucosome function against the emerging fungal pathogen Batrachochytrium dendrobatidis. Four species were tested with laboratory exposure experiments, and a highly susceptible species, Alytes obstetricans, was treated with a variety of temperature and microbial conditions to test the effects of probiotic therapies and environmental conditions on mucosome function. We found that antifungal function of the amphibian skin mucosome predicts the prevalence of infection with the fungal pathogen in natural populations, and is linked to survival in laboratory exposure experiments. When altered by probiotic therapy, the mucosome increased antifungal capacity, while previous exposure to the pathogen was suppressive. In culture, antifungal properties of probiotics depended strongly on immunological and environmental context including temperature, competition, and pathogen presence. Functional changes in microbiota with shifts in temperature provide an alternative mechanistic explanation for patterns of disease susceptibility related to climate beyond direct impact on host or pathogen. This nonlethal management tool can be used to optimize and quickly assess the relative benefits of probiotic therapies under different climatic, microbial, or host conditions.     

Temperature mediates vector transmission efficiency: inoculum supply and plant infection dynamics

Climate, particularly environmental temperature, frequently plays an important role in disease epidemiology. This study investigated the role of environmental temperature on transmission of the generalist plant pathogen Xylella fastidiosa by its leafhopper vectors. In this system temperature is known to influence both vector performance and feeding rate, yet the implications for pathogen transmission have not been documented. Experiments were conducted over a range of temperatures to document effects on transmission efficiency of the California native Graphocephala atropunctata (blue–green sharpshooter) and the invasive Homalodisca vitripennis (glassy-winged sharpshooter). Inoculation efficiency of H. vitripennis was positively related to temperature. Graphocephala atropunctata mortality and transmission responded non-linearly to temperature, with the highest rates of both at the highest temperature. The experiment also evaluated whether differences in inoculum supply contributed to plant infection level using quantitative PCR. Although total X. fastidiosa population within G. atropunctata was not related to plant infection, the number of infectious vectors was a strong predictor of plant infection level–suggesting that the number of inoculation events is important in the development of systemic infection of X. fastidiosa in grapevines. These results, along with existing evidence from the literature, point to wide-ranging impacts of climate on the epidemiology of X. fastidiosa diseases.     

Maize potential yields and yield gaps in the changing climate of northeast China

Northeast China (NEC) is not only one of the major agricultural production areas in China, but it is also the most susceptible to climate variability. This led us to investigate the impact of climate change on maize potential yield and yield gaps in this region, where maize accounts for about 30% of the nation's production. The APSIM‐Maize model was calibrated and validated for maize phenology and yields. The validated model was then used to estimate potential yields, rain‐fed potential yields, and yield gaps for assessing the climate impacts on maize productivity in NEC. During maize growing seasons from 1981 to 2010, the analysis indicates a warming trend all across NEC, whereas the trends in solar radiation and total precipitation tended to decrease. When the same hybrid was specified in APSIM for all years, a simulated increase of maximum temperature resulted in a negative impact on both potential yield and rain‐fed potential yield. A simulated increase in minimum temperature produced no significant changes in potential or rain‐fed potential yield. However, the increase of minimum temperature was shown to result in a positive impact on the on‐farm yield, consistent with our finding that farmers adopted longer season hybrids for which the increase in minimum temperature provided better conditions for germination, emergence, and grain filling during night time. The gap between potential and rain‐fed potential yields was shown to be larger at locations with lower seasonal precipitation (<500 mm). Our results indicate that regions with the largest yield gaps between rain‐fed potential and on‐farm yields were located in the southeast of NEC. Within NEC, on‐farm maize yields were, on average, only 51% of the potential yields, indicating a large exploitable yield gap, which provides an opportunity to significantly increase production by effective irrigation, fertilization, herbicide, and planting density in NEC.     

Modelling the effect of temperature variation on the seasonal dynamics of Ixodes ricinus tick populations

Seasonal variation in temperature is known to drive annual patterns of tick activity and can influence the dynamics of tick-borne diseases. An age-structured model of the dynamics of Ixodes ricinus populations was developed to explore how changes in average temperature and different levels of temperature variability affect seasonal patterns of tick activity and the transmission of tick-borne diseases. The model produced seasonal patterns of tick emergence that are consistent with those observed throughout Great Britain. Varying average temperature across a continuous spectrum produced a systematic pattern in the times of peak emergence of questing ticks which depends on cumulative temperature over the year. Examination of the effects of between-year stochastic temperature variation on this pattern indicated that peak emergence times are more strongly affected by temperature stochasticity at certain levels of average temperature. Finally the model was extended to give a simple representation of the dynamics of a tick-borne disease. A threshold level of annual cumulative temperature was identified at which disease persistence is sensitive to stochastic temperature variation. In conclusion, the effect of changing patterns of temperature variation on the dynamics of I. ricinus ticks and the diseases they transmit may depend on the cumulative temperature over the year and will therefore vary across different locations. The results also indicate that diapause mechanisms have an important influence on seasonal patterns of tick activity and require further study.