Fatal attraction: vegetation responses to nutrient inputs attract herbivores to infectious anthrax carcass sites

Parasites can shape the foraging behaviour of their hosts through cues indicating risk of infection. When cues for risk co-occur with desired traits such as forage quality, individuals face a trade-off between nutrient acquisition and parasite exposure. We evaluated how this trade-off may influence disease transmission in a 3-year experimental study of anthrax in a guild of mammalian herbivores in Etosha National Park, Namibia. At plains zebra (Equus quagga) carcass sites we assessed (i) carcass nutrient effects on soils and grasses, (ii) concentrations of Bacillus anthracis (BA) on grasses and in soils, and (iii) herbivore grazing behaviour, compared with control sites, using motion-sensing camera traps. We found that carcass-mediated nutrient pulses improved soil and vegetation, and that BA is found on grasses up to 2 years after death. Host foraging responses to carcass sites shifted from avoidance to attraction, and ultimately to no preference, with the strength and duration of these behavioural responses varying among herbivore species. Our results demonstrate that animal carcasses alter the environment and attract grazing hosts to parasite aggregations. This attraction may enhance transmission rates, suggesting that hosts are limited in their ability to trade off nutrient intake with parasite avoidance when relying on indirect cues.     

Predation and disease: understanding the effects of predators at several trophic levels on pathogen transmission

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The cost of phage resistance in a plant pathogenic bacterium is context‐dependent

Parasites are ubiquitous features of living systems and many parasites severely reduce the fecundity or longevity of their hosts. This parasite‐imposed selection on host populations should strongly favor the evolution of host resistance, but hosts typically face a trade‐off between investment in reproductive fitness and investment in defense against parasites. The magnitude of such a trade‐off is likely to be context‐dependent, and accordingly costs that are key in shaping evolution in nature may not be easily observable in an artificial environment. We set out to assess the costs of phage resistance for a plant pathogenic bacterium in its natural plant host versus in a nutrient‐rich, artificial medium. We demonstrate that mutants of Pseudomonas syringae that have evolved resistance via a single mutational step pay a substantial cost for this resistance when grown on their tomato plant hosts, but do not realize any measurable growth rate costs in nutrient‐rich media. This work demonstrates that resistance to phage can significantly alter bacterial growth within plant hosts, and therefore that phage‐mediated selection in nature is likely to be an important component of bacterial pathogenicity.     

Evolution of Drosophila resistance against different pathogens and infection routes entails no detectable maintenance costs

Pathogens exert a strong selective pressure on hosts, entailing host adaptation to infection. This adaptation often affects negatively other fitness‐related traits. Such trade‐offs may underlie the maintenance of genetic diversity for pathogen resistance. Trade‐offs can be tested with experimental evolution of host populations adapting to parasites, using two approaches: (1) measuring changes in immunocompetence in relaxed‐selection lines and (2) comparing life‐history traits of evolved and control lines in pathogen‐free environments. Here, we used both approaches to examine trade‐offs in Drosophila melanogaster populations evolving for over 30 generations under infection with Drosophila C Virus or the bacterium Pseudomonas entomophila, the latter through different routes. We find that resistance is maintained after up to 30 generations of relaxed selection. Moreover, no differences in several classical life‐history traits between control and evolved populations were found in pathogen‐free environments, even under stresses such as desiccation, nutrient limitation, and high densities. Hence, we did not detect any maintenance costs associated with resistance to pathogens. We hypothesize that extremely high selection pressures commonly used lead to the disproportionate expression of costs relative to their actual occurrence in natural systems. Still, the maintenance of genetic variation for pathogen resistance calls for an explanation.     

Toxoplasma gondii infection enhances testicular steroidogenesis in rats

The protozoan parasite Toxoplasma gondii enhances the sexual attractiveness of infected male rats and attenuates the innate fear of cat odour in infected individuals. These behavioural changes plausibly lead to greater transmission of parasites through sexual and trophic routes, respectively. Testosterone, a testicular steroid, is known to reduce fear and enhance sexual attractiveness in males. Here, we show that Toxoplasma gondii infection enhances expression of genes involved in facilitating synthesis of testosterone, resulting in greater testicular testosterone production in male rats. In several species, testosterone mediates trade‐offs between sexually selected traits and life history decisions. Augmentation of testosterone synthesis by Toxoplasma gondii suggests that parasites may manipulate these trade‐offs in rats.     

Herbivore species identity rather than diversity of the non‐host community determines foraging behaviour of the parasitoid wasp Cotesia glomerata

Extensive research has been conducted to reveal how species diversity affects ecosystem functions and services. Yet, consequences of diversity loss for ecosystems as a whole as well as for single community members are still difficult to predict. Arthropod communities typically are species‐rich, and their species interactions, such as those between herbivores and their predators or parasitoids, may be particularly sensitive to changes in community composition. Parasitoids forage for herbivorous hosts by using herbivore‐induced plant volatiles (indirect cues) and cues produced by their host (direct cues). However, in addition to hosts, non‐suitable herbivores are present in a parasitoid's environment which may complicate the foraging process for the parasitoid. Therefore, ecosystem changes in the diversity of herbivores may affect the foraging efficiency of parasitoids. The effect of herbivore diversity may be mediated by either species numbers per se, by specific species traits, or by both. To investigate how diversity and identity of non‐host herbivores influence the behaviour of parasitoids, we created environments with different levels of non‐host diversity. On individual plants in these environments, we complemented host herbivores with 1–4 non‐host herbivore species. We subsequently studied the behaviour of the gregarious endoparasitoid Cotesia glomerata L. (Hymenoptera: Braconidae) while foraging for its gregarious host Pieris brassicae L. (Lepidoptera: Pieridae). Neither non‐host species diversity nor non‐host identity influenced the preference of the parasitoid for herbivore‐infested plants. However, after landing on the plant, non‐host species identity did affect parasitoid behaviour, whereas non‐host diversity did not. One of the non‐host species, Trichoplusia ni Hübner (Lepidoptera: Noctuidae), reduced the time the parasitoid spent on the plant as well as the number of hosts it parasitized. We conclude that non‐host herbivore species identity has a larger influence on C. glomerata foraging behaviour than non‐host species diversity. Our study shows the importance of species identity over species diversity in a multitrophic interaction of plants, herbivores, and parasitoids.     

Variation in costs of parasite resistance among natural host populations

Organisms that can resist parasitic infection often have lower fitness in the absence of parasites. These costs of resistance can mediate host evolution during parasite epidemics. For example, large epidemics will select for increased host resistance. In contrast, small epidemics (or no disease) can select for increased host susceptibility when costly resistance allows more susceptible hosts to outcompete their resistant counterparts. Despite their importance for evolution in host populations, costs of resistance (which are also known as resistance trade‐offs) have mainly been examined in laboratory‐based host–parasite systems. Very few examples come from field‐collected hosts. Furthermore, little is known about how resistance trade‐offs vary across natural populations. We addressed these gaps using the freshwater crustacean Daphnia dentifera and its natural yeast parasite, Metschnikowia bicuspidata. We found a cost of resistance in two of the five populations we studied – those with the most genetic variation in resistance and the smallest epidemics in the previous year. However, yeast epidemics in the current year did not alter slopes of these trade‐offs before and after epidemics. In contrast, the no‐cost populations showed little variation in resistance, possibly because large yeast epidemics eroded that variation in the previous year. Consequently, our results demonstrate variation in costs of resistance in wild host populations. This variation has important implications for host evolution during epidemics in nature.     

Evolution of host resistance and trade‐offs between virulence and transmission potential in an obligately killing parasite

Standard epidemiological theory predicts that parasites, which continuously release propagules during infection, face a trade‐off between virulence and transmission. However, little is known how host resistance and parasite virulence change during coevolution with obligate killers. To address this question we have set up a coevolution experiment evolving Nosema whitei on eight distinct lines of Tribolium castaneum. After 11 generations we conducted a time‐shift experiment infecting both the coevolved and the replicate control host lines with the original parasite source, and coevolved parasites from generation 8 and 11. We found higher survival in the coevolved host lines than in the matching control lines. In the parasite populations, virulence measured as host mortality decreased during coevolution, while sporeload stayed constant. Both patterns are compatible with adaptive evolution by selection for resistance in the host and by trade‐offs between virulence and transmission potential in the parasite.     

Spread of an introduced parasite across the Hawaiian archipelago independent of its introduced host

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Identity and combinations of arbuscular mycorrhizal fungal isolates influence plant resistance and insect preference

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Turbidity amplifies the non‐lethal effects of predation and affects the foraging success of characid fish shoals

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QTL mapping of freezing tolerance: links to fitness and adaptive trade‐offs

Local adaptation, defined as higher fitness of local vs. nonlocal genotypes, is commonly identified in reciprocal transplant experiments. Reciprocally adapted populations display fitness trade‐offs across environments, but little is known about the traits and genes underlying fitness trade‐offs in reciprocally adapted populations. We investigated the genetic basis and adaptive significance of freezing tolerance using locally adapted populations of Arabidopsis thaliana from Italy and Sweden. Previous reciprocal transplant studies of these populations indicated that subfreezing temperature is a major selective agent in Sweden. We used quantitative trait locus (QTL) mapping to identify the contribution of freezing tolerance to previously demonstrated local adaptation and genetic trade‐offs. First, we compared the genomic locations of freezing tolerance QTL to those for previously published QTL for survival in Sweden, and overall fitness in the field. Then, we estimated the contributions to survival and fitness across both field sites of genotypes at locally adaptive freezing tolerance QTL. In growth chamber studies, we found seven QTL for freezing tolerance, and the Swedish genotype increased freezing tolerance for five of these QTL. Three of these colocalized with locally adaptive survival QTL in Sweden and with trade‐off QTL for overall fitness. Two freezing tolerance QTL contribute to genetic trade‐offs across environments for both survival and overall fitness. A major regulator of freezing tolerance, CBF2, is implicated as a candidate gene for one of the trade‐off freezing tolerance QTL. Our study provides some of the first evidence of a trait and gene that mediate a fitness trade‐off in nature.     

Alternative parasite development in transmission strategies: how time flies!

Among parasitic platyhelminths with complex life cycles, it has been well documented that transmission opportunities are the main forces shaping the diversity of life‐history traits and parasite developmental strategies. While deviations in the development pathway usually involve shortening of life cycles, their extension may also occur following perception of remaining time by parasites. Polystoma gallieni, the monogenean parasite of Hyla meridionalis, is able to trigger two alternative developmental strategies depending on the physiological stage of the tadpoles upon which larvae attach. The distribution and reproductive outputs of both resulting phenotypes were surveyed to address questions about the dynamics of transmission in natural environments. Because modifications in the completion of life cycles can have drawbacks which may perturb the dynamic equilibrium of the resulting host–parasite systems, experimental infestations were also performed to assess parasite–parasite interactions. Our results suggest that the bladder adult phenotype, which involves transmission between frogs and tadpoles, is supplied secondarily by the branchial phenotype which involves transmission between tadpoles and metamorphs. They also support the occurrence of finely tuned trade‐offs between hosts and parasites and highlight positive trends behind the extension of direct life cycles, in which host‐derived signals account for the remaining time to achieve parasitic transmission.     

Do‐or‐die life cycles and diverse post‐infection resistance mechanisms limit the evolution of parasite host ranges

In light of the dynamic nature of parasite host ranges and documented potential for rapid host shifts, the observed high host specificity of most parasites remains an ecological paradox. Different variants of host‐use trade‐offs have become a mainstay of theoretical explanations of the prevalence of host specialism, but empirical evidence for such trade‐offs is rare. We propose an alternative theory based on basic features of the parasite life cycle: host selection and subsequent intrahost replication. We introduce a new concept of effective burst size that accounts for the fact that successful host selection does not guarantee intrahost replication. Our theory makes a general prediction that a parasite will expand its host range if its effective burst size is positive. An in silico model of bacteria‐phage coevolution verifies our predictions and demonstrates that the tendency for relatively narrow host ranges in parasites can be explained even in the absence of trade‐offs.     

Grazing in heterogeneous environments: infra- and supra-parasite distributions determine herbivore grazing decisions

We tested the hypothesis that the infra-gastrointestinal parasite population of herbivores affects their grazing behaviour in relation to the supra-parasite population of parasites in the environment. Our first objective was to create a naturally heterogeneous sward structure of gaps and tussocks using a continuous grazing scheme. We then demonstrate that a nutrition vs. parasitism grazing trade-off occurs within that sward structure and that infra-gastrointestinal parasite populations affect the grazing decisions of herbivores faced with the trade-off. A pool of 50 naturally parasitised female Soay sheep and their lambs were used to create a heterogeneous tall, faeces-contaminated tussock/short, non-contaminated gap sward structure in a 1-ha experimental plot. Tussocks offered approximately 1.5 times greater forage intake but contained 5.5 times the number of strongyle parasites compared to the gaps. Following a 10-week period in which the heterogeneous sward structure was created, two 5-day periods of observations of sward structure selection (i.e. gap vs. tussock) were carried out. Twenty female Soay lambs were divided into two groups of ten (balanced for live-weight) immediately prior to the start of the observation period. One of the groups of lambs was treated with an anthelmintic drench before the start of the second observation period creating two levels of parasitism (high and low). On each observation day 5-min focal observations were carried out on each animal at least twice a day, during which time the number of bites taken from gaps and tussocks were recorded along with the number of steps. During the first period of observations, all animals rejected the relatively tall, faeces-contaminated tussocks for grazing to a similar extent and had similar bite and step rates. During the second period of observations all animals showed reduced rejection of the tussocks relative to the first week, however, animals with a reduced parasite population showed a greater reduction in rejection as compare to the highly parasitised animals. We conclude that the infra- and supra-distributions of parasites within herbivore hosts and the environment greatly impact on herbivore grazing behaviour and foraging decisions and thus the structure and heterogeneity of grazed ecosystems.     

Variation in resource acquisition and use among host clones creates key epidemiological trade‐offs

Parasites can certainly harm host fitness. Given such virulence, hosts should evolve strategies to resist or tolerate infection. But what governs those strategies and the costs that they incur? This study illustrates how a fecundity‐susceptibility trade‐off among clonally reared genotypes of a zooplankton (Daphnia dentifera) infected by a fungal parasite (Metschnikowia) arises due to variation in resource acquisition and use by hosts. To make these connections, we used lab experiments and theoretical models that link feeding with susceptibility, energetics, and fecundity of hosts. These feeding‐based mechanisms also produced a fecundity‐survivorship trade‐off. Meanwhile, a parasite spore yield–fecundity trade‐off arose from variation in juvenile growth rate among host clones (another index of resource use), a result that was readily anticipated and explained by the models. Thus, several key epidemiological trade‐offs stem from variation in resource acquisition and use among clones. This connection should catalyze the creation of new theory that integrates resource‐ and gene‐based responses of hosts to disease.     

Within-host population dynamics and the evolution of microparasites in a heterogeneous host population

Abstract Why do parasites harm their hosts? The general understanding is that if the transmission rate and virulence of a parasite are linked, then the parasite must harm its host to maximize its transmission. The exact nature of such trade‐offs remains largely unclear, but for vertebrate hosts it probably involves interactions between a microparasite and the host immune system. Previous results have suggested that in a homogeneous host population in the absence of super‐ or coinfection, within‐host dynamics lead to selection of the parasite with an intermediate growth rate that is just being controlled by the immune system before it kills the host (Antia et al. 1994). In this paper, we examine how this result changes when heterogeneity is introduced to the host population. We incorporate the simplest form of heterogeneity–random heterogeneity in the parameters describing the size of the initial parasite inoculum, the immune response of the host, and the lethal density at which the parasite kills the host. We find that the general conclusion of the previous model holds: parasites evolve some intermediate growth rate. However, in contrast with the generally accepted view, we find that virulence (measured by the case mortality or the rate of parasite‐induced host mortality) increases with heterogeneity. Finally, we link the within‐host and between‐host dynamics of parasites. We show how the parameters for epidemiological spread of the disease can be estimated from the within‐host dynamics, and in doing so examine the way in which trade‐offs between these epidemiological parameters arise as a consequence of the interaction of the parasite and the immune response of the host.     

Escherichia coli populations adapt to complex,unpredictable fluctuations by minimizing trade‐offs across environments

In nature, organisms are simultaneously exposed to multiple stresses (i.e. complex environments) that often fluctuate unpredictably. Although both these factors have been studied in isolation, the interaction of the two remains poorly explored. To address this issue, we selected laboratory populations of Escherichia coli under complex (i.e. stressful combinations of pH, H₂O₂ and NaCl) unpredictably fluctuating environments for ~900 generations. We compared the growth rates and the corresponding trade‐off patterns of these populations to those that were selected under constant values of the component stresses (i.e. pH, H₂O₂ and NaCl) for the same duration. The fluctuation‐selected populations had greater mean growth rate and lower variation for growth rate over all the selection environments experienced. However, whereas the populations selected under constant stresses experienced trade‐offs in the environments other than those in which they were selected, the fluctuation‐selected populations could bypass the across‐environment trade‐offs almost entirely. Interestingly, trade‐offs were found between growth rates and carrying capacities. The results suggest that complexity and fluctuations can strongly affect the underlying trade‐off structure in evolving populations.     

Environmental context alters ecological trade‐offs controlling ant coexistence in a spatially heterogeneous region

1. Ecological trade‐offs in ant (Hymenoptera: Formicidae) assemblages and their implications for coexistence boast a rich history in entomology. Yet investigations of trade‐offs have largely been limited to homogeneous environments. We examined how environmental context modifies trade‐off expression in an ant assemblage spanning a heterogeneous region in central Florida, U.S.A. 2. We examined how trade‐off expression is altered among two contrasting habitat types: open shrub and forest. We tested for the presence of the dominance‐discovery trade‐off and two dominance‐thermal tolerance trade‐offs by estimating behavioral dominance, discovery ability, and thermal tolerance (foraging thermal limit, lethal temperature, and maximal abundance temperature) for a wide range of interacting ant species. 3. We found significantly linear dominance hierarchies in both shrub and forest habitats, showing dominant species out‐compete subordinates for food resources. In thermally stressful shrub habitats, subordinates exhibit higher thermal tolerances, take greater thermal risks, and reach maximum forager abundances at higher temperatures than do dominant species. This suggests temperature mediated trade‐offs control coexistence in shrub habitat. In thermally moderate forest habitat, we found limited evidence for trade‐offs between competitive dominance and resource discovery or between dominance and thermal traits, implying other processes control coexistence. These results demonstrate that trade‐offs controlling ant coexistence may be contingent on environmental context.     

The combined effects of hypoxia and fish kairomones on several physiological and life history traits of Daphnia

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