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Here we develop a tug-of-war game theory model of mixed-strategy facultative parasitism. In this framework, individuals decide how to strategically invest in parental care effort, parasitism, and resistance to being parasitized, choosing their investments to respond optimally to their opponent's behavior and vice versa. We have implemented the model in the well-studied case of mixed-strategy conspecific brood parasitism, which occurs when a female raises her own clutch of eggs and flexibly lays some eggs in a conspecific's nest. However, the nest of a parasitic female in this case may also be parasitized. We model this as a tug-of-war, allowing parasitic and resistance efforts to strategically coevolve with each other. We then derive expressions for parasitism outcomes commonly observed in the field. The model also captures the trade-offs between parental care and parasitism when both are possible. We make a number of novel predictions about the rate of successful versus attempted parasitism as well as how parental care effort, resistance to parasitism, and fitness vary as functions of group size and individual differences between players in energy budget and production efficiency. Although we focus primarily on conspecific brood parasitism, the model is general enough to be extended to other systems in which facultative parasitism is possible, including the parasitism of food, fertilizations, nest sites, or other resources from conspecifics. [ABSTRACT FROM AUTHOR]

The determinants of diet breadth are of interest to nutritionists, ecologists, and evolutionary biologists. A recent synthesis addressing this issue found conflicting evidence for the relationship between diet breadth and mean individual fitness. Specifically, it found that while, on average, a mixed diet does increase mean fitness, in some instances, a single food provides equal (or higher) fitness than a mixed diet. Critical to ecological and evolutionary considerations of diet, however, is not only mean fitness but also variance in fitness. We combine contemporary meta-analytic methods with models of nutritional geometry to evaluate how diet affects between-individual variance in fitness within generalist consumers froma range of trophic levels. As predicted by nutritional geometry, we found that between-individual variance in fitness-related traits is higher on single-food than mixed diets. The effect was strong for longevity traits (57% higher) and reproductive traits (37%) and present but weaker for size-related traits (10%). Further, the effect became stronger as the number of available foods increased. The availability ofmultiple foods likely allows individuals with differing nutritional optima to customize intake, each maximizing their own fitness. Importantly, these findings may suggest that selection on traits correlated with nutritional requirements is weak in heterogeneous nutritional environments. [ABSTRACT FROM AUTHOR]

The near-neutral model of B chromosome evolution predicts that the invasion of a new population should last some tens of generations, but the details on how it proceeds in real populations are mostly unknown. Trying to fill this gap, we analyze here a natural population of the grasshopper Eyprepocnemis plorans at three time points during the last 35 years. Our results show that B chromosome frequency increased significantly during this period and that a cline observed in 1992 had disappeared in 2012 once B chromosome frequency reached an upper limit at all sites sampled. This indicates that, during B chromosome invasion, transient clines for B chromosome frequency are formed at the invasion front on a microgeographic scale. Computer simulation experiments showed that the pattern of change observed for genotypic frequencies is consistent with the existence of B chromosome drive through females and selection against individuals with a high number of B chromosomes. [ABSTRACT FROM AUTHOR]

The integration of physiological mechanisms into life-history theory is an emerging frontier in our understanding of the constraints and drivers of life-history evolution. Dynamic patterns of antagonism between developmental and immunological pathways in juvenile insects illustrate the importance of mechanisms for determining life-history strategy optima in the face of trade-offs. For example, developmental interference occurs when developmental processes transiently take priority over resources or pathway architecture, preventing allocation to immunity or other traits. We designed a within-host model of infected larval development to explore the impact of developmental dynamics on optimal resource mobilization and allocation strategies as well as on larval resistance and tolerance phenotypes. The model incorporates mechanism-inspired functional forms of developmental interference with immunity against parasites that attack specific larval stages. We find that developmental interference generally increases optimal investment in constitutive immunity and decreases optimal resource mobilization rates, but the results are sensitive to the developmental stage at first infection. Moreover, developmental interference reduces resistance but generally increases tolerance of infection. We demonstrate the potential impact of these dynamics on empirical estimates of host susceptibility and discuss the general implications of incorporating realistic physiological mechanisms and developmental dynamics for life-history theory in insects and other organisms. [ABSTRACT FROM AUTHOR]

Intragenomic conflict may arise when social partners are more related through one parent than the other—for example, owing to individuals or gametes of one sex dispersing further prior to fertilization. In particular, genes originating from the former parent are favored to promote selflessness, and those originating from the latter parent are favored to promote selfishness. While the impact of patterns of dispersal on the evolution of intragenomic conflict has received recent attention, the consequences of intragenomic conflict for the evolution of dispersal remain to be explored. We suggest that if the evolution of dispersal is driven at least in part by kin selection, differential relatedness of social partners via their mothers versus their fathers may lead to an intragenomic conflict, with maternal-origin genes and paternal-origin genes favoring different rates of dispersal. As an illustration, we extend a classic model of the evolution of dispersal to explore how intragenomic conflict may arise between an individual's maternal-origin and paternal-origin genes over whether that individual should disperse in order to ease kin competition. Our analysis reveals extensive potential for intragenomic conflict over dispersal and predicts that genes underpinning dispersal phenotypesmay exhibit parent-of-origin-specific expression, which may facilitate their discovery. [ABSTRACT FROM AUTHOR]

Host-parasite infection genetics can be more complex than envisioned by classic models such as the gene-for-gene or matching-allele models. By means of a mathematical model, I investigate the coevolutionary dynamics arising from a large set of generalized models of infection genetics in which hosts are either fully resistant or fully susceptible to a parasite, depending on the genotype of both individuals. With a single diploid interaction locus in the hosts, many of the infection genetic models produce stable or neutrally stable genotype polymorphisms. However, only a few models, which are all different versions of the matching-allele model, lead to sustained cycles of genotype frequency fluctuations in both interacting species ("Red Queen" dynamics). By contrast, with two diploid interaction loci in the hosts, many infection genetics models that cannot be classified as one of the standard infection genetics models produce Red Queen dynamics. Sexual versus asexual reproduction and, in the former case, the rate of recombination between the interaction loci have a large impact on whether Red Queen dynamics arise from a given infection genetics model. This may have interesting but as yet unexplored implications with respect to the Red Queen hypothesis for the evolution of sex. [ABSTRACT FROM AUTHOR]

Worker honeybees, Apis mellifera, police each other's reproduction by killing worker-laid eggs. Previous experiments demonstrated that worker policing is effective, killing most (~98%) worker-laid eggs. However, many queen-laid eggs were also killed (~50%) suggesting that effective policing may have high costs. In these previous experiments, eggs were transferred using forceps into test cells, mostly into unrelated discriminator colonies. We measured both the survival of unmanipulated queen-laid eggs and the proportion of removal errors that were rectified by the queen laying a new egg. Across 2 days of the 3-day egg stage, only 9.6% of the queen-laid eggs in drone cells and 4.1% in worker cells were removed in error. When queen-laid eggs were removed from cells, 85% from drone cells and 61% from worker cells were replaced within 3 days. Worker policing in the honeybee has a high benefit to policing workers because workers are more related to the queen's sons (brothers, r p 0.25) than sister workers' sons (0.15). This study shows that worker policing also has a low cost in terms of the killing of queenlaid eggs, as only a small proportion of queen-laid eggs are killed, most of which are rapidly replaced. [ABSTRACT FROM AUTHOR]

The evolution of traits involved in ecological interactions such as predator-prey, host-parasite, and plant-pollinator interactions, are likely to be shaped by the phylogenetic history of both parties. We develop generalized linear mixed-effects models (GLMM) that estimate the effect of both parties' phylogenetic history on trait evolution, both in isolation but also in terms of how the two histories interact. Using data on the incidence and abundance of 206 flea species on 121 mammal species, we illustrate our method and compare it to previously used methods for detecting host-parasite coevolution. At large spatial scales we find that the phylogenetic interaction effect was substantial, indicating that related parasite species were more likely to be found on related host species. At smaller spatial scales, and when sampling effort was not controlled for, phylogenetic effects on the number and types of parasite species harbored by hosts were found to dominate. We go on to show that in situations where these additional phylogenetic effects exist, then previous methods have very high Type I error rates when testing for the phylogenetic interaction. Our GLMM method represents a robust and reliable approach to quantify the phylogenetic effects of traits determined by, or defined by, ecological interactions and has the advantage that it can easily be extended and interpreted in a broader context than existing permutation tests. [ABSTRACT FROM AUTHOR]

Adaptive speciation with gene flow via the evolution of assortative mating has classically been studied in one of two different scenarios. First, speciation can occur if frequency-dependent competition in sympatry induces disruptive selection, leading to indirect selection for mating with similar phenotypes. Second, if a subpopulation is locally adapted to a specific environment, then there is indirect selection against hybridizing with maladapted immigrants. While both of these mechanisms have been modeled many times, the literature lacks models that allow direct comparisons between them. Here we incorporate both frequency-dependent competition and local adaptation into a single model and investigate whether and how they interact in driving speciation. We report two main results. First, we show that individually, the two mechanisms operate under separate conditions, hardly influencing each other when one of them alone is sufficient to drive speciation. Second, we also find that the two mechanisms can operate together, leading to a third speciation mode in which speciation is initiated by selection against maladapted migrants but completed by within-deme competition in a distinct second phase. While this third mode bears some similarity to classical reinforcement, it is considerably faster, and both newly formed species go on to coexist in sympatry. [ABSTRACT FROM AUTHOR]

In many species, nongenetic phenotypic variation helps mitigate risk associated with an uncertain environment. In some cases, developmental cues can be used to match phenotype to environment-- a strategy known as predictive plasticity. When environmental conditions are entirely unpredictable, generating random phenotypic diversity may improve the long-term success of a lineage-- a strategy known as diversified bet hedging. When partially reliable information is available, a well-adapted developmental strategy may strike a balance between the two strategies. We use information theory to analyze a model of development in an uncertain environment, where cue reliability is affected by variation both within and between generations. We show that within-generation variation in cues decreases the reliability of cues without affecting their fitness value. This transpires because the optimal balance of predictive plasticity and diversified bet hedging is unchanged. However, within-generation variation in cues does change the developmental mechanisms used to create that balance: developmental sensitivity to such cues not only helps match phenotype to environment but also creates phenotypic diversity that may be useful for hedging bets against environmental change. Understanding the adaptive role of developmental sensitivity thus depends on a proper assessment of both the predictive power and the structure of variation in environmental cues. [ABSTRACT FROM AUTHOR]

Disentangling individual selection from kin selection is one of the greatest challenges of evolutionary biology. Even solitary organisms that do not interact directly with conspecifics may interact indirectly with them through competition for resources. As a result, traits that appear to affect individual fitness alone can also modify the fitness of relatives nearby and thus may evolve partially through these cryptic indirect fitness effects. Here we develop a method to quantitatively separate direct and indirect fitness consequences when some microbes become dormant, while neighbors of the same genotype remain active. Dormant microbes typically survive stresses that kill metabolically active cells, but dormancy also has a social side effect, sparing resources that may be used by nondormant individuals for growth. In structured populations, spared resources may be preferentially consumed by nondormant clonemates, providing an indirect benefit. Without population structure, however, exploitation by a never-dormant competitor imposes an indirect fitness cost on dormant cells. Cryptic indirect fitness effects may play a significant role in the evolution of many ostensibly asocial traits. [ABSTRACT FROM AUTHOR]

Transmission from host to host is a crucial step in the life cycle of pathogens, particularly of viruses, ensuring spread and maintenance in host populations. The immobile nature of plants and the strong pectin and cellulose barrier surrounding cells have constrained most plant virus species to use vectors (mainly insects) for exit, transfer and entry from one host to another., A growing body of evidence is showing that plant viruses can influence vector physiology and behaviour to increase their chances of transmission, either directly or through modification of the host plant. In contrast, little is known on the possible reciprocal interaction, where the vector way of life would significantly impact on the viral behaviour and/or phenotype within the infected plants, on its population genetics and its evolution., The complex possible reaches of these three-way interactions on the ecology of each partner have not been exhaustively explored., After briefly summarizing the current knowledge on how viruses can induce changes in insect vector behaviour, physiology and population dynamics, this review focuses on presenting unforeseen aspects related to (i) the impacts that the feeding habits of different insect vectors can have on the evolution of plant viruses and (ii) the possibility that vector-related stresses induce major switches in the 'behaviour' of viruses in planta, affecting primarily the efficiency of transmission by insect vectors. [ABSTRACT FROM AUTHOR]

The transmission-virulence trade-off hypothesis is one of the few adaptive explanations of virulence evolution, and assumes that there is an overall positive correlation between parasite transmission and virulence. The shape of the transmission-virulence relationship predicts whether virulence should evolve toward either a maximum or to an intermediate optimum. A positive correlation between each of these traits and within-host growth is often suggested to underlie the relationship between virulence and transmission. There are few experimental tests of this hypothesis; this study reports on the first empirical test on a plant pathogen. We infected Brassica rapa plants with nine natural isolates of Cauliflower mosaic virus and then estimated three traits: transmission, virulence, and within-host viral accumulation. As predicted by the trade-off hypothesis, we observed a positive correlation between transmission and virulence, suggestive of the existence of an intermediate optimum. We discovered the unexpected existence of two groups of within-host accumulation, differing by at least an order of magnitude. When accumulation groups were not accounted for, within-host accumulation was correlated neither to virulence nor transmission, although our results suggest that within each group these correlations exist. [ABSTRACT FROM AUTHOR]

Background: The question whether Plasmodium falciparum infection affects the fitness of mosquito vectors remains open. A hurdle for resolving this question is the lack of appropriate control, non-infected mosquitoes that can be compared to the infected ones. It was shown recently that heating P. falciparum gametocyte-infected blood before feeding by malaria vectors inhibits the infection. Therefore, the same source of gametocyte-infected blood could be divided in two parts, one heated, serving as the control, the other unheated, allowing the comparison of infected and uninfected mosquitoes which fed on exactly the same blood otherwise. However, before using this method for characterizing the cost of infection to mosquitoes, it is necessary to establish whether feeding on previously heated blood affects the survival and fecundity of mosquito females. Methods: Anopheles gambiae M molecular form females were exposed to heated versus non-heated, parasite-free human blood to mimic blood meal on non-infectious versus infectious gametocyte-containing blood. Life history traits of mosquito females fed on blood that was heat-treated or not were then compared. Results: The results reveal that heat treatment of the blood did not affect the survival and fecundity of mosquito females. Consistently, blood heat treatment did not affect the quantity of blood ingested. Conclusions: The study indicates that heat inactivation of gametocyte-infected blood will only inhibit mosquito infection and that this method is suitable for quantifying the fitness cost incurred by mosquitoes upon infection by P. falciparum. [ABSTRACT FROM AUTHOR]

The authors comment on the study by J. Smith and colleagues, which challenges transmission-virulence trade-off hypothesis in virulence evolution. They note that despite experimental results have good quality, they opposes the study as such hypothesis is one of the adaptive accounts of virulence evolution and is important to describe virulence levels of a parasite. They also develop a mathematical model to address their contrary to the author's concepts on transmission and superinfection.

Transitions from sexual to asexual reproduction are often coupled with elevations in ploidy. As a consequence, the importance of ploidy per se for the maintenance and spread of asexual populations is unclear. To examine the effects of ploidy and asexual reproduction as independent determinants of the success of asexual lineages, we sampled diploid sexual, diploid asexual, and triploid asexual Eucypris virens ostracods across a European wide range. Applying nuclear and mitochondrial markers, we found that E. virens consists of genetically highly differentiated diploid sexual populations, to the extent that these sexual clades could be considered as cryptic species. All sexual populations were found in southern Europe and North Africa and we found that both diploid asexual and triploid asexual lineages have originated multiple times from several sexual lineages. Therefore, the asexual lineages show a wide variety of genetic backgrounds and very strong population genetic structure across the wide geographic range. Finally, we found that triploid, but not diploid, asexual clones dominate habitats in northern Europe. The limited distribution of diploid asexual lineages, despite their shared ancestry with triploid asexual lineages, strongly suggests that the wider geographic distribution of triploids is due to elevated ploidy rather than to asexuality. [ABSTRACT FROM AUTHOR]

The authors discuss aspects of the study "Large Bottleneck Size in Cauliflower Mosaic Virus Populations during Host Plant Colonization," by B. Monsion and colleagues in the 2008 issue. Topics include the estimated bottleneck size of cauliflower mosaic virus (CaMV) during host plant colonization, the two methods use in the study which were on the evolution of neutral marker frequencies, and the Poisson-distributed random variable.