Host Population Dynamics Research Articles

Impacts of infection avoidance for populations affected by sexually transmitted infections

Sexually transmitted infections are ubiquitous in nature and affect many populations. The key process for their transmission is mating, usually preceded by mate choice. Susceptible individuals may avoid mating with infected individuals to prevent infection provided it is recognizable. We show that accounting for infection avoidance significantly alters host population dynamics. We observe bistability between the disease-free and endemic or disease-induced extinction equilibria, significant abrupt reduction in the host population size and disease-induced host extinction. From the population persistence perspective, the best strategy is either not to avoid mating with the infected individuals, to prevent disease-induced host extinction, or to completely avoid mating with the infected individuals, to prevent pathogen invasion. Increasing sterilization efficiency of the infection leads to lower population sizes and reduced effect of mating avoidance. We also find that the disease-free state is more often attained by populations with strong polyandry, whereas a high-density endemic state is more often observed for populations with strong polygyny, suggesting that polygamy rather than monogamy may be promoted in denser host populations.

Spatiotemporal and gender-specific parasitism in two species of gobiid fish.

Parasitism is considered a major selective force in natural host populations. Infections can decrease host condition and vigour, and potentially influence, for example, host population dynamics and behavior such as mate choice. We studied parasite infections of two common marine fish species, the sand goby (Pomatoschistus minutus) and the common goby (Pomatoschistus microps), in the brackish water Northern Baltic Sea. We were particularly interested in the occurrence of parasite taxa located in central sensory organs, such as eyes, potentially affecting fish behavior and mate choice. We found that both fish species harbored parasite communities dominated by taxa transmitted to fish through aquatic invertebrates. Infections also showed significant spatiotemporal variation. Trematodes in the eyes were very few in some locations, but infection levels were higher among females than males, suggesting differences in exposure or resistance between the sexes. To test between these hypotheses, we experimentally exposed male and female sand gobies to infection with the eye fluke Diplostomum pseudospathaceum. These trials showed that the fish became readily infected and females had higher parasite numbers, supporting higher susceptibility of females. Eye fluke infections also caused high cataract intensities among the fish in the wild. Our results demonstrate the potential of these parasites to influence host condition and visual abilities, which may have significant implications for survival and mate choice in goby populations.

Association of spring-summer hydrology and meteorology with human West Nile virus infection in West Texas, USA, 2002\u20132016

BackgroundThe emergence of West Nile virus (WNV) in the Western Hemisphere has motivated research into the processes contributing to the incidence and persistence of the disease in the region. Meteorology and hydrology are fundamental determinants of vector-borne disease transmission dynamics of a region. The availability of water influences the population dynamics of vector and host, while temperature impacts vector growth rates, feeding habits, and disease transmission potential. Characterization of the temporal pattern of environmental factors influencing WNV risk is crucial to broaden our understanding of local transmission dynamics and to inform efforts of control and surveillance.MethodsWe used hydrologic, meteorological and WNV data from west Texas (2002–2016) to analyze the relationship between environmental conditions and annual human WNV infection. A Bayesian model averaging framework was used to evaluate the association of monthly environmental conditions with WNV infection.ResultsFindings indicate that wet conditions in the spring combined with dry and cool conditions in the summer are associated with increased annual WNV cases. Bayesian multi-model inference reveals monthly means of soil moisture, specific humidity and temperature to be the most important variables among predictors tested. Environmental conditions in March, June, July and August were the leading predictors in the best-fitting models.ConclusionsThe results significantly link soil moisture and temperature in the spring and summer to WNV transmission risk. Wet spring in association with dry and cool summer was the temporal pattern best-describing WNV, regardless of year. Our findings also highlight that soil moisture may be a stronger predictor of annual WNV transmission than rainfall.

Resource-driven changes to host population stability alter the evolution of virulence and transmission

What drives the evolution of parasite life-history traits? Recent studies suggest that linking within- and between-host processes can provide key insight into both disease dynamics and parasite evolution. Still, it remains difficult to understand how to pinpoint the critical factors connecting these cross-scale feedbacks, particularly under non-equilibrium conditions; many natural host populations inherently fluctuate and parasites themselves can strongly alter the stability of host populations. Here, we develop a general model framework that mechanistically links resources to parasite evolution across a gradient of stable and unstable conditions. First, we dynamically link resources and between-host processes (host density, stability, transmission) to virulence evolution, using a 'non-nested' model. Then, we consider a 'nested' model where population-level processes (transmission and virulence) depend on resource-driven changes to individual-level (within-host) processes (energetics, immune function, parasite production). Contrary to 'non-nested' model predictions, the 'nested' model reveals complex effects of host population dynamics on parasite evolution, including regions of evolutionary bistability; evolution can push parasites towards strongly or weakly stabilizing strategies. This bistability results from dynamic feedbacks between resource-driven changes to host density, host immune function and parasite production. Together, these results highlight how cross-scale feedbacks can provide key insights into the structuring role of parasites and parasite evolution.This article is part of the theme issue 'Anthropogenic resource subsidies and host-parasite dynamics in wildlife'.

Prediction and attenuation of seasonal spillover of parasites between wild and domestic ungulates in an arid mixed-use system.

Transmission of parasites between host species affects host population dynamics, interspecific competition, and ecosystem structure and function. In areas where wild and domestic herbivores share grazing land, management of parasites in livestock may affect or be affected by sympatric wildlife due to cross‐species transmission.We develop a novel method for simulating transmission potential based on both biotic and abiotic factors in a semi‐arid system in Botswana. Optimal timing of antiparasitic treatment in livestock is then compared under a variety of alternative host scenarios, including seasonally migrating wild hosts.In this region, rainfall is the primary driver of seasonality of transmission, but wildlife migration leads to spatial differences in the effectiveness of treatment in domestic animals. Additionally, competent migratory wildlife hosts move parasites across the landscape.Simulated transmission potential matches observed patterns of clinical disease in livestock in the study area. Increased wildlife contact is correlated with a decrease in disease, suggesting that non‐competent wild hosts may attenuate transmission by removing infective parasite larvae from livestock pasture.Optimising the timing of treatment according to within‐year rainfall patterns was considerably more effective than treating at a standard time of year. By targeting treatment in this way, efficient control can be achieved, mitigating parasite spillover from wildlife where it does occur. Synthesis and applications. This model of parasite transmission potential enables evidence‐based management of parasite spillover between wild and domestic species in a spatio‐temporally dynamic system. It can be applied in other mixed‐use systems to mitigate parasite transmission under altered climate scenarios or changes in host ranges.

Phage-host population dynamics promotes prophage acquisition in bacteria with innate immunity.

Temperate bacteriophages integrate in bacterial genomes as prophages and represent an important source of genetic variation for bacterial evolution, frequently transmitting fitness-augmenting genes such as toxins responsible for virulence of major pathogens. However, only a fraction of bacteriophage infections are lysogenic and lead to prophage acquisition, whereas the majority are lytic and kill the infected bacteria. Unless able to discriminate lytic from lysogenic infections, mechanisms of immunity to bacteriophages are expected to act as a double-edged sword and increase the odds of survival at the cost of depriving bacteria of potentially beneficial prophages. We show that although restriction-modification systems as mechanisms of innate immunity prevent both lytic and lysogenic infections indiscriminately in individual bacteria, they increase the number of prophage-acquiring individuals at the population level. We find that this counterintuitive result is a consequence of phage-host population dynamics, in which restriction-modification systems delay infection onset until bacteria reach densities at which the probability of lysogeny increases. These results underscore the importance of population-level dynamics as a key factor modulating costs and benefits of immunity to temperate bacteriophages.

Macroparasite dynamics of migratory host populations

Spatial variability in host density is a key factor affecting disease dynamics of wildlife, and yet there are few spatially explicit models of host–macroparasite dynamics. This limits our understanding of parasitism in migratory hosts, whose densities change considerably in both space and time. In this paper, we develop a model for host–macroparasite dynamics that considers the directional movement of host populations and their associated parasites. We include spatiotemporal changes in the mean and variance in parasite burden per host, as well as parasite-mediated host mortality and parasite-mediated migratory ability. Reduced migratory ability with increasing parasitism results in heavily infested hosts halting their migration, and higher parasite burdens in stationary hosts than in moving hosts. Simulations reveal the potential for positive feedbacks between parasite-reduced migratory ability and increasing parasite burdens at infection hotspots, such as stopover sites, that may lead to parasite-induced migratory stalling. This framework could help understand how global change might influence wildlife disease via changes to migratory patterns and parasite demographic rates.

Coinfection Timing Drives Host Population Dynamics through Changes in Virulence.

Infections of one host by multiple parasites are common, and several studies have found that the order of parasite invasion can affect both within-host competition and disease severity. However, it is unclear to what extent coinfection timing might be important to consider when modeling parasite impacts on host populations. Using a model system of two viruses infecting barley, we found that simultaneous infections of the two viruses were significantly more damaging to hosts than sequential coinfections. While priority effects were evident in within-host concentrations of sequential coinfections, priority did not influence any parameters (such as virulence or transmission rate) that affect host population dynamics. We built a susceptible-infected model to examine whether the observed difference in coinfection virulence could impact host population dynamics under a range of scenarios. We found that coinfection timing can have an important but context-dependent effect on projected host population dynamics. Studies that examine only simultaneous coinfections could inflate disease impact predictions.

Iron Chloride Flocculation of Bacteriophages from Seawater.

Viruses influence ecosystem dynamics by modulating microbial host population dynamics, evolutionary trajectories and metabolic outputs. While they are ecologically important across diverse ecosystems, viruses are challenging to study due to minimal biomass often obtained when sampling natural communities. Here we describe a technique using chemical flocculation, filtration and resuspension to recover bacteriophages from seawater and other natural waters. The method uses iron to precipitate viruses which are recovered by filtration onto large-pore size membranes and then resuspended using a buffer containing magnesium and a reductant (ascorbic acid or oxalic acid) at slightly acid pH (6-6.5). The recovery of bacteriophages using iron flocculation is efficient (>90%), inexpensive and reliable, resulting in preparations that are amenable to downstream analysis by next generation DNA sequencing, proteomics and, in some cases, can be used to study virus-host interactions.

Alterations in microbial community composition with increasing <i>f</i>CO<sub>2</sub>: a mesocosm study in the eastern Baltic Sea

Abstract. Ocean acidification resulting from the uptake of anthropogenic carbon dioxide (CO2) by the ocean is considered a major threat to marine ecosystems. Here we examined the effects of ocean acidification on microbial community dynamics in the eastern Baltic Sea during the summer of 2012 when inorganic nitrogen and phosphorus were strongly depleted. Large-volume in situ mesocosms were employed to mimic present, future and far future CO2 scenarios. All six groups of phytoplankton enumerated by flow cytometry ( < 20 µm cell diameter) showed distinct trends in net growth and abundance with CO2 enrichment. The picoeukaryotic phytoplankton groups Pico-I and Pico-II displayed enhanced abundances, whilst Pico-III, Synechococcus and the nanoeukaryotic phytoplankton groups were negatively affected by elevated fugacity of CO2 (fCO2). Specifically, the numerically dominant eukaryote, Pico-I, demonstrated increases in gross growth rate with increasing fCO2 sufficient to double its abundance. The dynamics of the prokaryote community closely followed trends in total algal biomass despite differential effects of fCO2 on algal groups. Similarly, viral abundances corresponded to prokaryotic host population dynamics. Viral lysis and grazing were both important in controlling microbial abundances. Overall our results point to a shift, with increasing fCO2, towards a more regenerative system with production dominated by small picoeukaryotic phytoplankton.

Egyptian rousette bats maintain long-term protective immunity against Marburg virus infection despite diminished antibody levels

Although bats are natural reservoir hosts for numerous zoonotic viruses, little is known about the long-term dynamics of the host immune response following infection and how these viruses are maintained in nature. The Egyptian rousette bat (ERB) is a known reservoir host for Marburg virus (MARV). Following infection of ERBs with MARV, virus-specific IgG antibodies are induced but rapidly wane and by 3 months post-infection the bats are seronegative. To determine whether reinfection of ERBs plays a role in MARV maintenance, we challenge groups of ERBs that were “naturally” or experimentally infected with MARV 17–24 months prior. No bats in either group exhibit evidence of MARV replication or shedding and all bats develop virus-specific secondary immune responses. This study demonstrates that infection of ERBs with MARV induces long-term protective immunity against reinfection and indicates that other factors, such as host population dynamics, drive MARV maintenance in nature.

Epiphytic Curtobacterium flaccumfaciens strains isolated from symptomless solanaceous vegetables are pathogenic on leguminous but not on solanaceous plants

In Iran, during 2013–16, 16 Gram‐positive corynebacteria‐like strains were recovered from the epiphytic parts of solanaceous vegetables including eggplant, pepper and tomato. The strains were recovered accidentally as a result of monitoring for other bacterial pathogens in solanaceous fields. The strains were phenotypically different from Clavibacter michiganensis strains. Although none of the strains were pathogenic on their host of isolation or on any other solanaceous plants, 12 out of 16 strains were pathogenic on common bean, cowpea, mung bean and soybean. Colonization by strains was observed on maize, zucchini, faba bean, honeydew melon, rapeseed, sugar beet and sunflower plants under greenhouse conditions. In PCR tests, the primer pair CffFOR2/CffREV4, specific for Curtobacterium flaccumfaciens pv. flaccumfaciens, enabled the amplification of the appropriately sized fragment in 12 out of 16 strains, and all 12 strains were pathogenic on dry beans. Phylogenetic analysis, using the gyrB and recA genes, showed all 16 bacterial strains clustered within several pathovars of C. flaccumfaciens. A nonpathogenic yellow‐pigmented strain (Xeu15) was clustered with the type strains of C. flaccumfaciens pv. betae and C. flaccumfaciens pv. oortii. Bacteriocin profiling assays revealed no significant differences among the pathogenic and nonpathogenic strains. Host range and population dynamics of four representative strains on 17 plant species showed population build‐up of the strains only on common bean, cowpea, wheat and red nightshade plants. The results provide important insights into the possible role of nonhost plants as reservoirs of plant pathogenic bacteria, which has important implications in plant disease epidemiology and management.

Marine viruses discovered via metagenomics shed light on viral strategies throughout the oceans

Marine viruses are key drivers of host diversity, population dynamics and biogeochemical cycling and contribute to the daily flux of billions of tons of organic matter. Despite recent advancements in metagenomics, much of their biodiversity remains uncharacterized. Here we report a data set of 27,346 marine virome contigs that includes 44 complete genomes. These outnumber all currently known phage genomes in marine habitats and include members of previously uncharacterized lineages. We designed a new method for host prediction based on co-occurrence associations that reveals these viruses infect dominant members of the marine microbiome such as Prochlorococcus and Pelagibacter. A negative association between host abundance and the virus-to-host ratio supports the recently proposed Piggyback-the-Winner model of reduced phage lysis at higher host densities. An analysis of the abundance patterns of viruses throughout the oceans revealed how marine viral communities adapt to various seasonal, temperature and photic regimes according to targeted hosts and the diversity of auxiliary metabolic genes.

The basic reproduction number of vector-borne plant virus epidemics

The basic reproduction number R0 is a key parameter in plant disease epidemiology, which largely determines whether or not an epidemic will occur in a plant population. The next generation matrix approach to deriving and calculating the basic reproduction number of a plant virus epidemic is described. The approach is illustrated through a series of examples of increasing complexity, ranging from the simplest case of one vector transmitting one virus to a single host, to the case of multiple vectors, to combined horizontal (vector) and vertical (seed) transmission, and where vector control using insecticides is practised. The importance of parameters representing host and vector population dynamics and their interaction in the absence of disease is stressed, and the constraints these place on the calculation of the basic reproduction number. Finally, mention is made of further elaborations to the approach that could prove useful in plant virus epidemiology.

ASSESSMENT OF EPIZOOTIC ACTIVITY AND EPIDEMIC RISK IN HEMORRHAGIC FEVER WITH RENAL SYNDROME FOREST FOCI OF PRIMORSKY KRAI

In forest foci of hemorrhagic fever with renal syndrome (HFRS) of Primorsky Krai Amur and Hokkaido hantaviruses are circulated in the natural host population, i.e. Apodemus рeninsulae and Myodes rufocanus. The indicators of epizootic activity and their threshold values in rodent population dynamics were determined in order to forecast HFRS outbreaks. The material for the research was obtained throughout 2001–2013. The number of trap-nights was 33890. 2737 of A. peninsulae and 1643 of M. rufocanus were investigated. ELISA, RT-PCR, IFA were used for the detection of hantavirus antigen, RNA, antibodies/avidity. The parameters of epizootic activity per 100 TN (N – population, n i – infected rodents, n ai – individuals with acute infection) were proposed. The results showed that the active foci of Amur viral infection are located on the slopes of the Sikhote-Alin ridge. The epizootic activity in A. peninsulae populations is characterized by the expressed cyclicity: phase I is rise, phase II is high activity, phase III is recession/low activity. Phases I and II are valid up to a year, whereas phase III is valid within a few years, determining the whole epizootic cycle. In phase I the number of rodents with an acute infection increased from summer to late autumn. In phase II the number of rodents with acute infection was considerable in spring/summer and decreased in autumn more than 3 times. In all phases of the cycle the threshold values were considerably higher for A. peninsulae, than for M. rufocanus, thus showing the former’s epidemiological role in forest ecosystems. HFRS infection rate in phases I/II of the epizootic cycle in A. peninsulae populations had two peaks: during late autumn - winter (≥ 37% of cases) and spring - first half of summer (≥ 55% of cases). The obtained threshold values of Amur hantavirus active circulation in population dynamics of the natural host allows to predict the periods of the increased risk of infection in HFRS forest foci for humans.

Infections of Wolbachia may destabilize mosquito population dynamics

Recent efforts in controlling mosquito-borne diseases focus on biocontrol strategies that incapacitate pathogens inside mosquitoes by altering the mosquito's microbiome. A case in point is the introduction of Wolbachia into natural mosquito populations in order to eliminate Dengue virus. However, whether this strategy can successfully control vector-borne diseases is debated; particularly, how artificial infection affects population dynamics of hosts remains unclear. Here, we show that natural Wolbachia infections are associated with unstable mosquito population dynamics by contrasting Wolbachia-infected versus uninfected cage populations of the Asian tiger mosquito (Aedes albopictus). By analyzing weekly data of adult mosquito abundances, we found that the variability of the infected populations is significantly higher than that of the uninfected. The elevated population variability is explained by increased instability in dynamics, as quantified by system nonlinearity (i.e., state-dependence). In addition, predictability of infected populations is substantially lower. A mathematical model analysis suggests that Wolbachia may alter mosquito population dynamics by modifying larval competition of hosts. These results encourage examination for effects of artificial Wolbachia establishment on mosquito populations, because an enhancement of population variability with reduced predictability could pose challenges in management. Our findings have implications for application of microbiome alterations in biocontrol programs.

Plant genotype and induced defenses affect the productivity of an insect-killing obligate viral pathogen.

Plant-mediated variations in the outcomes of host-pathogen interactions can strongly affect epizootics and the population dynamics of numerous species, including devastating agricultural pests such as the fall armyworm. Most studies of plant-mediated effects on insect pathogens focus on host mortality, but few have measured pathogen yield, which can affect whether or not an epizootic outbreak occurs. Insects challenged with baculoviruses on different plant species and parts can vary in levels of mortality and yield of infectious stages (occlusion bodies; OBs). We previously demonstrated that soybean genotypes and induced anti-herbivore defenses influence baculovirus infectivity. Here, we used a soybean genotype that strongly reduced baculovirus infectivity when virus was ingested on induced plants (Braxton) and another that did not reduce infectivity (Gasoy), to determine how soybean genotype and induced defenses influence OB yield and speed of kill. These are key fitness measures because baculoviruses are obligate-killing pathogens. We challenged fall armyworm, Spodoptera frugiperda, with the baculovirus S. frugiperda multi-nucleocapsid nucleopolyhedrovirus (SfMNPV) during short or long-term exposure to plant treatments (i.e., induced or non-induced genotypes). Caterpillars were either fed plant treatments only during virus ingestion (short-term exposure to foliage) or from the point of virus ingestion until death (long-term exposure). We found trade-offs of increasing OB yield with slower speed of kill and decreasing virus dose. OB yield increased more with longer time to death and decreased more with increasing virus dose after short-term feeding on Braxton compared with Gasoy. OB yield increased significantly more with time to death in larvae that fed until death on non-induced foliage than induced foliage. Moreover, fewer OBs per unit of host tissue were produced when larvae were fed induced foliage than non-induced foliage. These findings highlight the potential importance of plant effects, even at the individual plant level, on entomopathogen fitness, which may impact epizootic transmission events and host population dynamics.

Variations in infection levels and parasite-induced mortality among sympatric cryptic lineages of native amphipods and a congeneric invasive species: Are native hosts always losing?

Shared parasites can strongly influence the outcome of competition between congeneric, sympatric hosts, and thus host population dynamics. Parasite-mediated competition is commonly hypothesized as an important factor in biological invasion success; invasive species often experience lower infection levels and/or parasite-induced mortality than native congeneric hosts. However, variation in infection levels among sympatric hosts can be due to contrasting abilities to avoid infection or different parasite-induced mortality rates following infection. Low parasite infection levels in a specific host can be due to either factor but have drastically different implications in interaction outcomes between sympatric hosts.We assessed acanthocephalan infection levels (prevalence and abundance) among cryptic molecular taxonomic units (MOTU) of the native G. pulex/G. fossarum species complex from multiple populations where they occur in sympatry. We concomitantly estimated the same parameters in the invasive Gammarus roeseli commonly found in sympatry with G. pulex/G. fossarum MOTUs. We then tested for potential differences in parasite-induced mortality among these alternative hosts. As expected, the invasive G. roeseli showed relatively low infection level and was not subject to parasite-induced mortality. We also found that both acanthocephalan infection levels and parasite-induced mortality varied greatly among cryptic MOTUs of the native amphipods. Contrary to expectations, some native MOTUs displayed levels of resistance to their local parasites similar to those observed in the invasive G. roeseli. Overall, cryptic diversity in native amphipods coupled with high levels of variability in infection levels and parasite-induced mortality documented here may strongly influence inter-MOTU interactions and native population dynamics as well as invasion success and population dynamics of the congeneric invasive G. roeseli.

Drivers of salamander extirpation mediated by Batrachochytrium salamandrivorans.

The recent arrival of Batrachochytrium salamandrivorans in Europe was followed by rapid expansion of its geographical distribution and host range, confirming the unprecedented threat that this chytrid fungus poses to western Palaearctic amphibians. Mitigating this hazard requires a thorough understanding of the pathogen's disease ecology that is driving the extinction process. Here, we monitored infection, disease and host population dynamics in a Belgian fire salamander (Salamandra salamandra) population for two years immediately after the first signs of infection. We show that arrival of this chytrid is associated with rapid population collapse without any sign of recovery, largely due to lack of increased resistance in the surviving salamanders and a demographic shift that prevents compensation for mortality. The pathogen adopts a dual transmission strategy, with environmentally resistant non-motile spores in addition to the motile spores identified in its sister species B. dendrobatidis. The fungus retains its virulence not only in water and soil, but also in anurans and less susceptible urodelan species that function as infection reservoirs. The combined characteristics of the disease ecology suggest that further expansion of this fungus will behave as a 'perfect storm' that is able to rapidly extirpate highly susceptible salamander populations across Europe.

Temporal dynamics of the tick Ixodes ricinus in northern Europe: epidemiological implications

BackgroundTick-borne pathogens pose an increasing threat to human and veterinary health across the northern hemisphere. While the seasonal activity of ticks is largely determined by climatic conditions, host-population dynamics are also likely to affect tick abundance. Consequently, abundance fluctuations of rodents in northern Europe are expected to be translated into tick dynamics, and can hence potentially affect the circulation of tick-borne pathogens. We quantified and explained the temporal dynamics of the tick Ixodes ricinus in the northernmost part of its European geographical range, by estimating (i) abundance in vegetation and (ii) infestation load in the most common rodent species in the study area, the bank vole Myodes glareolus.ResultsIxodes ricinus nymphs and adult females, the life stages responsible for the most of tick bites in humans, peaked in May-June and August-September. Larvae and nymphs were simultaneously active in June and abundance of questing larvae and nymphs in the vegetation showed a positive association with bank vole abundance. Moreover, infesting larvae and nymphs were aggregated on bank voles, and the infestation of bank voles with I. ricinus larvae and nymphs was positively associated with bank vole abundance.ConclusionOur results indicate early summer and early autumn as periods of increased risk for humans to encounter I. ricinus ticks in boreal urban forests and suggest a 2 years life-cycle for I. ricinus with two cohorts of ticks during the same year. Moreover, we identified a simultaneous activity of larvae and nymphs which allows co-feeding on the rodent host, which in turn supports the transmission of several important zoonotic tick-borne pathogens. Finally, we showed that a high density of the rodent host may enhance the risk that ticks and, potentially, tick-borne pathogens pose to human health.