Increased Contact Rates Research Articles

Genetic co‐structuring in host‐parasite systems: Empirical data from raccoons and raccoon ticks

AbstractMany aspects of parasite biology critically depend on their hosts, and understanding how host‐parasite populations are co‐structured can help improve our understanding of the ecology of parasites, their hosts, and host‐parasite interactions. This study utilized genetic data collected from raccoons (Procyon lotor), and a specialist parasite, the raccoon tick (Ixodes texanus), to test for genetic co‐structuring of host‐parasite populations at both landscape and host scales. At the landscape scale, our analyses revealed a significant correlation between genetic and geographic distance matrices (i.e., isolation by distance) in ticks, but not their hosts. While there are several mechanisms that could lead to a stronger pattern of isolation by distance in tick vs. raccoon datasets, our analyses suggest that at least one reason for the above pattern is the substantial increase in statistical power (due to the ≈8‐fold increase in sample size) afforded by sampling parasites. Host‐scale analyses indicated higher relatedness between ticks sampled from related vs. unrelated raccoons trapped within the same habitat patch, a pattern likely driven by increased contact rates between related hosts. By utilizing fine‐scale genetic data from both parasites and hosts, our analyses help improve our understanding of epidemiology and host ecology.

Dilution versus facilitation: Impact of connectivity on disease risk in metapopulations

Epidemiological studies have suggested that increasing connectivity in metapopulations usually facilitates pathogen transmission. However, these studies focusing on single-host systems usually neglect that increasing connectivity can increase species diversity which might reduce pathogen transmission via the ‘dilution effect’, a hypothesis whose generality is still disputed. On the other hand, studies investigating the generality of the dilution effect were usually conducted without considering habitat structure, which is surprising as species loss is often driven by habitat fragmentation. Using a simple general model to link fragmentation to the dilution effect, we determined the effect of connectivity on disease risk and explored when the dilution effect can be detected. We showed that landscape structure can largely modify the diversity–disease relationship. The net impact of connectivity on disease risk can be either positive or negative, depending on the relative importance of the facilitation effect (through increasing contact rates among patches) versus the dilution effect (via increasing species richness). We also demonstrated that different risk indices (i.e. infection prevalence and abundance of infected hosts) react differently to increasing connectivity and species richness. Our study may contribute to the current debate on the dilution effect, and a better understanding of the impacts of fragmentation on disease risks.

SEROLOGIC EVIDENCE OF CANINE PARVOVIRUS IN DOMESTIC DOGS, WILD CARNIVORES, AND MARSUPIALS IN THE ARGENTINEAN CHACO

The transmission of pathogens between domestic dogs and generalist wildlife species may be modified by environmental degradation, biodiversity losses, host densities, and increased contact rates in remnant forest patches. A serologic survey of canine parvovirus (CPV) in rural domestic dogs and wild mammals was conducted in two neighboring rural areas (disturbed and protected) from Pampa del Indio, northeastern Argentina, between 2008 and 2011. A total of 174 domestic dogs and 26 wild mammals-4 crab-eating foxes (Cerdocyon thous), 3 crab-eating raccoons (Procyon cancrivorus), 17 white-eared opossums (Didelphis albiventris), and 2 gray four-eyed opossums (Philander opossum)-were examined for antibodies to CPV using a hemagglutination inhibition assay. Domestic dogs were numerous and their movements unrestricted. The main function of dogs differed significantly between areas, with more dogs used for herding or hunting around the protected area. The seroprevalence of antibodies to CPV in dogs from both areas was very high (93.9-94.6%) and increased steeply with age. Nearly all carnivores and marsupials showed high exposure to CPV. Although a higher exposure to CPV was expected in wild mammals from disturbed areas as a result of enhanced contact between dogs and wildlife, no significant differences were found between areas. To the authors' knowledge, this study is the first to document exposure to CPV of free-ranging Pr. cancrivorus, D. albiventris, and Ph. opossum, and include a detailed demographic study of the domestic dog populations living in the area. This study highlights that dogs and wildlife have potential opportunities for contact and shows that the edges of the protected area may be as suitable as other fragmented areas for the transmission of CPV. Rural domestic dogs may pose serious threats to the health and conservation of wild carnivores in both disturbed and protected areas, especially in the Gran Chaco, where habitat fragmentation is severely increasing.

Transforming the fight against tuberculosis: targeting catalysts of transmission.

The global tuberculosis control community has committed itself to ambitious 10-year targets. To meet these targets, biomedical advances alone will be insufficient; a more targeted public health tuberculosis strategy is also needed. We highlight the role of "tuberculosis transmission catalysts," defined as variabilities in human behavior, bacillary properties, and host physiology that fuel the propagation of active tuberculosis at the local level. These catalysts can be categorized as factors that increase contact rates, infectiousness, or host susceptibility. Different catalysts predominate in different epidemiological and sociopolitical settings, and public health approaches are likely to succeed only if they are tailored to target the major catalysts driving transmission in the corresponding community. We argue that global tuberculosis policy should move from a country-level focus to a strategy that prioritizes collection of data on key transmission catalysts at the local level followed by deployment of "catalyst-targeted" interventions, supported by strengthened health systems.

Parasites and the conservation of small populations: The case of Baylisascaris procyonis.

Human demands on natural resources result in landscape changes that facilitate the emergence of disease. Most emerging diseases are zoonotic, and some of these pathogens play a role in the decline of vulnerable wildlife species. Baylisascaris procyonis, the common roundworm parasite of raccoons (Procyon lotor), is a well recognized zoonotic infection that has many of the properties associated with a pathogen capable of driving extinction. It is highly non-specific and frequently pathogenic with regard to paratenic hosts, which contact eggs of B. procyonis at raccoon latrines. Eggs accumulate at latrines and remain viable for many years. Transmission of B. procyonis is sensitive to changes in land-use, and fragmented habitats increase contact rates between raccoons, potential paratenic hosts, and the parasite. Raccoons, and subsequently B. procyonis, have been introduced to Europe and Japan, where naïve vertebrates may be exposed to the parasite. Finally, domestic animals and exotic pets can carry patent infections with B. procyonis, thus increasing environmental contamination beyond raccoon latrines, and expanding the area of risk to potential paratenic hosts. This parasite can potentially contribute to extinctions of vulnerable species, as exemplified by the case of the Allegheny woodrat (Neotoma magister), a species that has experienced local declines and extinctions that are linked to B. procyonis. Conservation strategies for vulnerable species should consider the transmission ecology of parasitic pathogens, like B. procyonis.

Comparing mechanisms of host manipulation across host and parasite taxa

Parasites affect host behavior in several ways. They can alter activity, microhabitats or both. For trophically transmitted parasites (the focus of our study), decreased activity might impair the ability of hosts to respond to final-host predators, and increased activity and altered microhabitat choice might increase contact rates between hosts and final-host predators. In an analysis of trophically transmitted parasites, more parasite groups altered activity than altered microhabitat choice. Parasites that infected vertebrates were more likely to impair the host's reaction to predators, whereas parasites that infected invertebrates were more likely to increase the host's contact with predators. The site of infection might affect how parasites manipulate their hosts. For instance, parasites in the central nervous system seem particularly suited to manipulating host behavior. Manipulative parasites commonly occupy the body cavity, muscles and central nervous systems of their hosts. Acanthocephalans in the data set differed from other taxa in that they occurred exclusively in the body cavity of invertebrates. In addition, they were more likely to alter microhabitat choice than activity. Parasites in the body cavity (across parasite types) were more likely to be associated with increased host contact with predators. Parasites can manipulate the host through energetic drain, but most parasites use more sophisticated means. For instance, parasites target four physiological systems that shape behavior in both invertebrates and vertebrates: neural, endocrine, neuromodulatory and immunomodulatory. The interconnections between these systems make it difficult to isolate specific mechanisms of host behavioral manipulation.

Effects of kin-structure on disease dynamics in raccoons (Procyon lotor) inhabiting a fragmented landscape

The spatial aggregation of related individuals can affect disease dynamics by altering contact rates between individuals and/or groups of individuals. However, kin-structure likely affects contact rates in a scale-dependent manner: increasing contact rates within, but decreasing contact rates amongst spatial groups. Thus, the effect of kin-structure on disease exposure risk should depend upon the spatial scale relevant for pathogen transmission. This study was undertaken to test the effects of kin-structure on disease dynamics in raccoons (Procyon lotor) inhabiting a highly fragmented agricultural ecosystem. Raccoons (n=566) were trapped at 31 spatially discrete habitat patches (local populations). Trapped individuals were genotyped at 13 microsatellites and their serum screened for antibodies to canine distemper virus (CDV) and multiple Leptospira spp. serovars. Our analyses revealed that the effects of kin-structure on transmission dynamics are dependent on both spatial scale and the pathogen under consideration. Thus, in the directly transmitted pathogen (CDV), the probability of pathogen co-exposure (the proportion of individual pairs where both individuals were seropositive for a particular pathogen) was positively correlated with genealogical proximity within local populations. Conversely, in the environmentally transmitted leptospires co-exposure was homogeneous within local populations and was positively correlated with gene flow among these populations. Our analyses also revealed that, after controlling for confounding effects of landscape and demographic variables, the effects of kin-structure on disease exposure risk was pathogen-specific. In particular, increasing kin-structure was associated with increased disease exposure risk in the directly transmitted pathogen (CDV), and alternatively with decreased disease exposure risk in environmentally transmitted leptospires. The potential, scale-dependent, effects of kin-structure on disease exposure risk revealed by our analyses could play an important role in elucidating disease dynamics in natural populations and in planning effective disease mitigation strategies.

Evaluating selective culling with vaccination to control wildlife disease: badgers and bovine tuberculosis (bTB)

Emerging infectious diseases often originate in wildlife, but the complex dynamics of wild animal populations mean that disease control is a major scientific and policy challenge. Culling and vaccination can be effective but the ecological characteristics of wild animals may confound the outcomes of simple management programs. In Britain, badgers Meles meles are a recognized reservoir of Mycobacterium bovis, the causative agent of bovine tuberculosis (bTB), and are involved in its transmission to cattle. Experiments have shown that culling badgers can increase bTB incidence in badgers and cattle, by perturbing the social structure of badger populations, and increasing contact rates. Selectively removal of infected badgers and vaccination of the remainder has recently been advocated as a potential solution. Simulation modelling suggests that this intuitively appealing policy could at best deliver only a minor advantage over thoroughly applying either vaccination or culling, but carries a risk of making the disease problem much worse. We suggest this counterintuitive outcome arises because: 1, not all animals can be caught and tested 2, some genuinely infected animals will be test negative 3, selective culling leaves a larger population of susceptible hosts than non-selective culling 4, some susceptible hosts will not respond to vaccination and 5, perturbation increases contact and transmission rates among a relatively high density population of infectious and susceptible animals.

Ocelots on Barro Colorado Island are infected with feline immunodeficiency virus but not other common feline and canine viruses.

Transmission of pathogens from domestic animals to wildlife populations (spill-over) has precipitated local wildlife extinctions in multiple geographic locations. Identifying such events before they cause population declines requires differentiating spillover from endemic disease, a challenge complicated by a lack of baseline data from wildlife populations that are isolated from domestic animals. We tested sera collected from 12 ocelots (Leopardus pardalis) native to Barro Colorado Island, Panama, which is free of domestic animals, for antibodies to feline herpes virus, feline calicivirus, feline corona virus, feline panleukopenia virus, canine distemper virus, and feline immunodeficiency virus (FIV), typically a species-specific infection. Samples also were tested for feline leukemia virus antigens. Positive tests results were only observed for FIV; 50% of the ocelots were positive. We hypothesize that isolation of this population has prevented introduction of pathogens typically attributed to contact with domestic animals. The high density of ocelots on Barro Colorado Island may contribute to a high prevalence of FIV infection, as would be expected with increased contact rates among conspecifics in a geographically restricted population.

Habitat fragmentation as a result of biotic and abiotic factors controls pathogen transmission throughout a host population

1. The hypothesis that habitat fragmentation (biotic or abiotic) alters the transmission of disease within a population is explored using field data from a well-studied amphibian-pathogen system. 2. We used the Ambystoma tigrinum-A. tigrinum virus (ATV) model system to show how habitat fragmentation as a result of emergent vegetation and habitat management affects disease transmission dynamics in ponds across a landscape. 3. We quantified variation in ATV infection over time and across the landscape. ATV infection was significantly higher in ponds modified for livestock use (P = 0.032). Disease incidence decreased with increased amounts of emergent vegetation (P < 0.001). These factors appear to control disease transmission by altering the host contact rate and with it disease transmission. 4. A field experiment to test the effect of emergent vegetation on the distribution of larvae in ponds demonstrated a behavioural change in larvae found in sparsely vegetated ponds. Microhabitat choices resulted in larvae being concentrated at the pond edge resulting in a 'halo effect' in sparsely vegetated ponds, whereas larvae in heavily vegetated ponds were distributed more evenly throughout. Microhabitat choice affects the effective density that larvae experience. This 'halo effect' increases contact rates in the shallows of sparsely vegetated ponds and increases the transmission of a directly transmitted pathogen. 5. Despite recurrent epidemics of a lethal Ranavirus in tiger salamanders on the Kaibab Plateau, Arizona, USA, these populations persist. We discuss the implications of our results in the context of density-dependent transmission and homogeneous mixing, two increases key assumptions of epidemiological theory.

How Host Population Dynamics Translate into Time-Lagged Prevalence: An Investigation of Sin Nombre Virus in Deer Mice

Human cases of hantavirus pulmonary syndrome caused by Sin Nombre virus are the endpoint of complex ecological cascade from weather conditions, population dynamics of deer mice, to prevalence of SNV in deer mice. Using population trajectories from the literature and mathematical modeling, we analyze the time lag between deer mouse population peaks and peaks in SNV antibody prevalence in deer mice. Because the virus is not transmitted vertically, rapid population growth can lead initially to reduced prevalence, but the resulting higher population size may later increase contact rates and generate increased prevalence. Incorporating these factors, the predicted time lag ranges from 0 to 18 months, and takes on larger values when host population size varies with a longer period or higher amplitude, when mean prevalence is low and when transmission is frequency-dependent. Population size variation due to variation in birth rates rather than death rates also increases the lag. Predicting future human outbreaks of hantavirus pulmonary syndrome may require taking these effects into account.

Effects of fragmentation on parasite burden (nematodes) of generalist and specialist small mammal species in secondary forest fragments of the coastal Atlantic Forest, Brazil

AbstractParasites are considered to play an important role in the regulation of wild animal populations. We investigated parasite burden of gastrointestinal nematodes and body condition in specialist and generalist small mammal species in secondary forest fragments in the highly endangered coastal Atlantic Forest. We hypothesized that body condition decreases with increasing parasite load and that parasite burden increases with increasing fragmentation in specialist species but not in generalist species as a consequence of differing responses to fragmentation effects. Investigated species were Akodon montensis, Oligoryzomys nigripes, and Delomys sublineatus (rodents) and the marsupials Marmosops incanus and Gracilinanus microtarsus. Prevalence of parasites was high in all species except for the arboreal G. microtarsus, presumably because of decreased infection probability. No correlation was found between body condition and parasite load in any of the species. Contrary to our expectations, body condition of the specialists D. sublineatus and M. incanus increased in both species with increasing fragmentation. In D. sublineatus, parasite burden increased and body condition decreased in fragments with relatively high density probably due to increased contact rates and facilitation of infection with nematodes. In all generalist species, low or no correlation between parasite burden and fragmentation was detected, suggesting little effect of fragmentation on population health.

A randomized study of electronic mail versus telephone follow-up after emergency department visit

This study was conducted to determine whether electronic mail (e-mail) increases contact rates after patients are discharged from the emergency department (ED). Following discharge, patients were randomized to be contacted by telephone or e-mail. The main outcome was success of contact. Secondary outcome was the median time of response. There were 1561 patients initially screened. Of these, 444 had e-mail and were included in the study. Half were contacted by telephone and the rest via e-mail. Our telephone contact rate was 58% (129/222) after two calls in a 48-h period and our e-mail contact was 41% (90/222). The telephone was nearly two times better than e-mail. The median time of response was 48 h for e-mail and 18 h for telephone. It is concluded that the telephone is a better modality of contact than e-mail for patients discharged from the ED.

Role of bacterial phenotypic traits in selective feeding of the heterotrophic nanoflagellate Spumella sp.

The influence of different bacterial phenotypic traits on the feeding selectivity of a bac- terivorous nanoflagellate was investigated in laboratory experiments. Twelve bacterial isolates from freshwater habitats were characterized in terms of cell size, morphology, capsule formation, surface hydrophobicity and charge, and swimming behavior. Mechanisms of differential flagellate feeding on these isolates were studied in short-term grazing experiments by high-resolution video microscopy using the nanoflagellate Spumella sp. as a model interception-feeding predator. The dif- ferentiation into distinct stages of the feeding process (contact, capture, ingestion) revealed a com- plex selection behavior illustrated by the relatively small proportion of ingested bacteria over cell contacts. We found that bacterial swimming speed increased contact rates but also decreased flagel- late capture efficiency, which had a compensating effect on overall ingestion rates. Bacterial cell size revealed no correlation with flagellate contact rates but an effective inhibition of ingestion rates when exceeding a critical size limit. Correlation analysis presented no bacterial property to account for the residual variability of flagellate ingestion efficiencies. Experiments with specifically coated artificial particles provided evidence for the relative importance of the biochemical surface composi- tion in flagellate food selection compared to physicochemical interaction forces. Only extreme charges beyond -45 mV reduced flagellate ingestion rates. Our results reinforce the idea that bacte- rial cell size strongly affects the feeding success of bacterivorous flagellates, and further implicates size-independent bacterial traits such as swimming speed and biochemical surface composition in this feeding success.

Size-based Prey Selectivity and Dietary Shifts in the Jellyfish, Aurelia aurita

Gut content analyses on field-caught Aurelia aurita showed both quantitative and qualitative change in diet as a function of medusa size. Larger medusae tended towards greater numbers and diversity of prey (up to 1550 individual prey representing as many as 13 different prey groups). We also found that medusa size was a good predictor of prey diversity recovered from the medusa gut. While a shift toward greater prey diversity in larger medusae might be explained by increased contact rates with rare' prey taxa, we also found size-based prey selectivity changes in A. aurita. We used in situ gut content data to describe selectivity by A. aurita for three prey types representing varying degrees of swimming or escape velocity. Fish eggs were used as a non-swimming prey, and small ( 1 mm) copepods were used as slow and fast swimming/escaping prey, respectively. Size-dependent increases in selectivity of small copepods can be explained by increased capture efficiency by larger medusae. Selectivity indices of fish eggs and large copepods did not vary with medusa size. Based on these data, we argue that growth to a large size by scyphomedusae can provide a means to enhance diet which, in turn, may positively influence growth and reproduction.

Can plants use entomopathogens as bodyguards?

For 20 years, ecologists have been gathering evidence in support of the hypothesis that plants can use insect natural enemies such as predators and parasitoids as bodyguards to protect themselves from herbivory, but entomopathogens have escaped this consideration. We extend the bodyguard hypothesis to ask whether plants can use entomopathogens as bodyguards. We first discuss the evolutionary context of such tritrophic interactions and then categorize possible mechanisms as: (1) maintaining a population of bodyguards on the plant surface, (2) increasing contact rates between insect host and pathogen and (3) increasing the susceptibility of the host. We explore these mechanisms further, examining published studies for evidence for the hypothesis. We then discuss potential costs to the plant of promoting pathogens as bodyguards which may include a reduction in the efficiency of other “bodyguard” species, the incidental promotion of plant pathogens and the risk of entomopathogens developing phytopathogenicity. Aside from our intention to stimulate the testing of the bodyguard hypothesis with entompathogens and to provide a conceptual framework for this, we hope to bring evolutionary ecology and insect pathology closer together.

Environmentally driven epizootics

Environmental conditions can be the driving force behind an epizootic. Environmental changes may favor growth of a particular species, which results in increased contact rates and spread of a disease. We examine this particular phenomenon in SI and SIS models and use it to explain the possible disease outbreaks in nature. Either infected individuals recover from the disease ( SIS model) or suffer disease fatalities ( SI model). Epizootic models for a single population are examined where contact rate depends on population size. A reproductive number R is defined that depends on environmental carrying capacity. The single-population models are coupled to form three different two-species models with intra- and interspecies contact rates that depend on the population sizes of both populations. The stability results show that it is possible for the disease to drive one of the populations to extinction, the one with disease fatalities. The surviving species serves as a reservoir for the disease. Single- and two-species epizootic models are examined in a particular case where the contact rates are assumed to be constant. This leads to a new definition for the contact rate. A complete global analysis is possible in these latter cases. The results are compared and contrasted with the models with variable contact rates. The prototype for the models is the spread of disease in wildlife populations, which includes such diseases as plague or Lyme disease.