1. Ecoimmunologists aim to understand the costs, benefits, and net fitness consequences of different strategies for immune defense. 2. Measuring the fitness consequences of immune responses is ...difficult, partly because of complex relationships between host fitness and the within-host density of parasites and immunological cells or molecules. In particular, neither the strongest immune responses nor the lowest parasite densities necessarily maximize host fitness. 3. Here, we propose that ecoimmunologists should routinely endeavour to measure three intertwined parameters: host fitness, parasite density, and relevant immune responses. We further propose that analyses of relationships among these traits would benefit from the statistical machinery used for analyses of phenotypic plasticity and/or methods that are robust to the bi-directional causation inherent in host-parasite relationships. For example, analyses of how host fitness depends upon parasite density, which is an evolutionary ecological definition of tolerance, would benefit from these more robust methods. 4. Together, these steps promote rigorous quantification of the fitness consequences of immune responses. Such quantification is essential if ecoimmunologists are to decipher causes of immune polymorphism in nature and predict trajectories of natural selection on immune defense.
Parasite transmission in a natural multihost–multiparasite community Auld, Stuart K. J. R.; Searle, Catherine L.; Duffy, Meghan A.
Philosophical transactions of the Royal Society of London. Series B. Biological sciences,
05/2017, Letnik:
372, Številka:
1719
Journal Article
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Understanding the transmission and dynamics of infectious diseases in natural communities requires understanding the extent to which the ecology, evolution and epidemiology of those diseases are ...shaped by alternative hosts. We performed laboratory experiments to test how parasite spillover affected traits associated with transmission in two co-occurring parasites: the bacterium Pasteuria ramosa and the fungus Metschnikowia bicuspidata. Both parasites were capable of transmission from the reservoir host (Daphnia dentifera) to the spillover host (Ceriodaphnia dubia), but this occurred at a much higher rate for the fungus than the bacterium. We quantified transmission potential by combining information on parasite transmission and growth rate, and used this to compare parasite fitness in the two host species. For both parasites, transmission potential was lower in the spillover host. For the bacterium, virulence was higher in the spillover host. Transmission back to the original host was high for both parasites, with spillover influencing transmission rate of the fungus but not the bacterium. Thus, while inferior, the spillover host is not a dead-end for either parasite. Overall, our results demonstrate that the presence of multiple hosts in a community can have important consequences for disease transmission, and host and parasite fitness.
This article is part of the themed issue ‘Opening the black box: re-examining the ecology and evolution of parasite transmission’.
The Red Queen hypothesis can explain the maintenance of host and parasite diversity. However, the Red Queen requires genetic specificity for infection risk (i.e., that infection depends on the exact ...combination of host and parasite genotypes) and strongly virulent effects of infection on host fitness. A European crustacean (Daphnia magna)--bacterium (Pasteuria ramosa) system typifies such specificity and high virulence. We studied the North American host Daphnia dentifera and its natural parasite Pasteuria ramosa, and also found strong genetic specificity for infection success and high virulence. These results suggest that Pasteuria could promote Red Queen dynamics with D. dentifera populations as well. However, the Red Queen might be undermined in this system by selection from a more common yeast parasite (Metschnikowia bicuspidata). Resistance to the yeast did not correlate with resistance to Pasteuria among host genotypes, suggesting that selection by Metschnikowia should proceed relatively independently of selection by Pasteuria.
Celotno besedilo
Dostopno za:
DOBA, IZUM, KILJ, NUK, PILJ, PNG, SAZU, SIK, UILJ, UKNU, UL, UM, UPUK
Populations experiencing varying levels of ionizing radiation provide an excellent opportunity to study the fundamental drivers of evolution. Radiation can cause mutations and thus supply genetic ...variation; it can also selectively remove individuals that are unable to cope with the physiological stresses associated with radiation exposure, or non‐selectively cull swathes of the population, reducing genetic variation. Since the nuclear power plant explosion in 1986, the Chernobyl area has experienced a spatially heterogeneous exposure to varying levels of ionizing radiation. We sampled Daphnia pulex (a freshwater crustacean) from lakes across the Chernobyl area, genotyped them at ten microsatellite loci and also calculated the current radiation dose rates. We then investigated whether the pattern of genetic diversity was positively associated with radiation dose rates, consistent with radiation‐mediated supply of de novo mutations, or negatively associated with radiation dose rates, as would be expected with strong radiation‐mediated selection. We found that measures of genetic diversity, including expected heterozygosity and mean allelic richness (an unbiased indicator of diversity), were significantly higher in lakes that experienced the highest radiation dose rates. This suggests that mutation outweighs selection as the key evolutionary force in populations exposed to high radiation dose rates. We also found significant but weak population structure, indicative of low genetic drift and clear evidence for isolation‐by‐distance between populations. This further suggests that gene flow between nearby populations is eroding population structure and that mutational input in high radiation lakes could, ultimately, supply genetic variation to lower radiation sites.
We examined population genetic diversity across Chernobyl Daphnia populations. Measures of genetic diversity, including expected heterozygosity and mean allelic richness, were higher in lakes that experienced the highest radiation dose rates. This suggests that mutation outweighs selection as the key evolutionary force in populations exposed to high radiation dose rates.
All animals and plants respond to changes in the environment during their life cycle. This flexibility is known as phenotypic plasticity and allows organisms to cope with variable environments. A ...common source of environmental variation is predation risk, which describes the likelihood of being attacked and killed by a predator. Some species can respond to the level of predation risk by producing morphological defences against predation. A classic example is the production of so‐called ‘neckteeth’ in the water flea, Daphnia pulex, which defend against predation from Chaoborus midge larvae. Previous studies of this defence have focussed on changes in pedestal size and the number of spikes along a gradient of predation risk. Although these studies have provided a model for continuous phenotypic plasticity, they do not capture the whole‐organism shape response to predation risk. In contrast, studies in fish and amphibians focus on shape as a complex, multi‐faceted trait made up of different variables. In this study, we analyse how multiple aspects of shape change in D. pulex along a gradient of predation risk from Chaoborus flavicans. These changes are dominated by the neckteeth defence, but there are also changes in the size and shape of the head and the body. We detected change in specific modules of the body plan and a level of integration among modules. These results are indicative of a complex, multi‐faceted response to predation and provide insight into how predation risk drives variation in shape and size at the level of the whole organism.
Recent advances in statistical methods for measuring shape were combined with a classic example of continuous phenotypic plasticity to measure the whole‐organism shape response to predation in Daphnia pulex. We found complex, multi‐faceted shape changes, with a mixture of locally and globally co‐ordinated features.
Climate change is causing warmer and more variable temperatures as well as physical flux in natural populations, which will affect the ecology and evolution of infectious disease epidemics. Using ...replicate seminatural populations of a coevolving freshwater invertebrate‐parasite system (host: Daphnia magna, parasite: Pasteuria ramosa), we quantified the effects of ambient temperature and population mixing (physical flux within populations) on epidemic size and population health. Each population was seeded with an identical suite of host genotypes and dose of parasite transmission spores. Biologically reasonable increases in environmental temperature caused larger epidemics, and population mixing reduced overall epidemic size. Mixing also had a detrimental effect on host populations independent of disease. Epidemics drove parasite‐mediated selection, leading to a loss of host genetic diversity, and mixed populations experienced greater evolution due to genetic drift over the season. These findings further our understanding of how diversity loss will reduce the host populations’ capacity to respond to changes in selection, therefore stymying adaptation to further environmental change.
Using replicate semi‐natural populations of a coevolving invertebrate‐parasite system, we demonstrated that biologically reasonable increases in environmental temperature led to larger epidemics, and population mixing reduced overall epidemic size. Mixing also had a detrimental effect on host populations independent of disease. Epidemics drove parasite‐mediated selection, leading to a loss of host genetic diversity, and mixed populations experienced greater evolution due to genetic drift over the season.
Epidemics commonly exert parasite-mediated selection and cause declines in host population genetic diversity. This can lead to evolution of resistance in the long term and smaller subsequent ...epidemics. Alternatively, the loss of genetic diversity can increase host vulnerability to future disease spread and larger future epidemics. Matters are made more complex by the fact that a great many host organisms produce diapausing life stages in response to environmental change (often as a result of sexual reproduction; e.g. plant seeds and invertebrate resting eggs). These diapausing stages can disrupt the relationship between past epidemics, host genetic diversity and future epidemics because they allow host dispersal through time. Specifically, temporally dispersing hosts avoid infection and thus selection from contemporary parasites, and also archive genetic variation for the future. We studied 80 epidemics in 20 semi-natural populations of the temporally dispersing crustacean
and its sterilizing bacterial parasite
, and half of these populations experienced a simulated environmental disturbance treatment. We found that early initiation of diapause relative to the timing of the epidemic led to greater host genetic diversity and reduced epidemic size in the subsequent year, but this was unaffected by environmental disturbance.
Sex as a strategy against rapidly evolving parasites Auld, Stuart K. J. R.; Tinkler, Shona K.; Tinsley, Matthew C.
Proceedings of the Royal Society. B, Biological sciences,
12/2016, Letnik:
283, Številka:
1845
Journal Article
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Why is sex ubiquitous when asexual reproduction is much less costly? Sex disrupts coadapted gene complexes; it also causes costs associated with mate finding and the production of males who do not ...themselves bear offspring. Theory predicts parasites select for host sex, because genetically variable offspring can escape infection from parasites adapted to infect the previous generations. We examine this using a facultative sexual crustacean, Daphnia magna, and its sterilizing bacterial parasite, Pasteuria ramosa. We obtained sexually and asexually produced offspring from wild-caught hosts and exposed them to contemporary parasites or parasites isolated from the same population one year later. We found rapid parasite adaptation to replicate within asexual but not sexual offspring. Moreover, sexually produced offspring were twice as resistant to infection as asexuals when exposed to parasites that had coevolved alongside their parents (i.e. the year two parasite). This fulfils the requirement that the benefits of sex must be both large and rapid for sex to be favoured by selection.
Parasite prevalence shows tremendous spatiotemporal variation. Theory indicates that this variation might stem from life-history characteristics of parasites and key ecological factors. Here, we ...illustrate how the interaction of an important predator and the schedule of transmission potential of two parasites can explain parasite abundance. A field survey showed that a noncastrating fungus (Metschnikowia bicuspidata) commonly infected a dominant zooplankton host (Daphnia dentifera), while a castrating bacterial parasite (Pasteuria ramosa) was rare. This result seemed surprising given that the bacterium produces many more infectious propagules (spores) than the fungus upon host death. The fungus’s dominance can be explained by the schedule of within-host growth of parasites (i.e., how transmission potential changes over the course of infection) and the release of spores from “sloppy” predators (Chaoborusspp., who consumeDaphniaprey whole and then later regurgitate the carapace and parasite spores). In essence, sloppy predators create a niche that the faster-schedule fungus currently occupies. However, a selection experiment showed that the slower-schedule bacterium can evolve into this faster-schedule, predator-mediated niche (but pays a cost in maximal spore yield to do so). Hence, our study shows how parasite life history can interact with predation to strongly influence the ecology, epidemiology, and evolution of infectious disease.
How hosts respond to and cope with infectious agents can change the environment and in so doing, alter selective pressures and evolutionary trajectories. To date, such eco-evolutionary feedbacks are ...best known from simplified mathematical models and laboratory experiments with a limited number of genetically homogenous model systems. However, the extent to which ecology and (co)-evolution interact to shape disease over space and time in natural communities remains poorly understood. Studies in this theme issue break new ground by integrating empirical and field studies stemming from a diverse array of taxa and ecosystems to understand how ecological and evolutionary feedbacks shape host-parasite interactions. Contributing papers synthesize emerging research and diverse perspectives on pathogen life history, virulence, resistance, leveraging novel methodological advances and integrating infection processes across multiple scales of biological organization. Two key take-aways emerge from this theme issue. First, more research tackling the eco-evolutionary dynamics of infectious disease in truly multi-species contexts is needed. Second, while we are far from understanding eco-evolutionary dynamics in the ‘natural theater’ where hosts encounter their parasites, exciting advances and foundational studies provide a well-defined road map.
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