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Many theoretical models have been formulated to better understand the coevolutionary patterns that emerge from antagonistic interactions. These models usually assume that the attacks by the exploiters are random, so the effect of victim selection by exploiters on coevolutionary patterns remains unexplored. Here we analytically studied the payoff for predators and prey under coevolution assuming that every individual predator can attack only a small number of prey any given time, considering two scenarios: (i) predation occurs at random; (ii) predators select prey according to phenotype matching. We also develop an individual based model to verify the robustness of our analytical prediction. We show that both scenarios result in well known similar coevolutionary patterns if population sizes are sufficiently high: symmetrical coevolutionary branching and symmetrical coevolutionary cycling (Red Queen dynamics). However, for small population sizes, prey selection can cause unexpected coevolutionary patterns. One is the breaking of symmetry of the coevolutionary pattern, where the phenotypes evolve towards one of two evolutionarily stable patterns. As population size increases, the phenotypes oscillate between these two values in a novel form of Red Queen dynamics, the episodic reversal between the two stable patterns. Thus, prey selection causes prey phenotypes to evolve towards more extreme values, which reduces the fitness of both predators and prey, increasing the likelihood of extinction., (Copyright © 2020 Elsevier Ltd. All rights reserved.)

Obligate symbiosis evolved from free-living individuals most likely via the intermediate stage of facultative symbiosis. However, why should facultative symbionts, who can live independently but also benefit from their partners if these are available, give up this best of both worlds? Using the adaptive dynamics approach, we analyse a simple model, focusing on one partner of the symbiosis, to gain more insight into the selective forces that make individuals forgo the ability to reproduce in the free-living state. Our results suggest that, similar to the parasitism-mutualism continuum, the free-living way of life and obligate symbiosis are two extremes of a continuum of the ability to reproduce independently of a partner. More importantly, facultative symbiosis should be the rule as for many parameter combinations completely giving up independent reproduction or adopting a pure free-living strategy is not so easy. We also show that if host encounter comes at a cost, individuals that put more effort into increasing the chances to meet with their partners are more likely to give up the ability to reproduce independently. Finally, our model does not specify the ecological interactions between hosts and symbionts but we discuss briefly how the ecological nature of an interaction can influence the transition from facultative to obligate symbiosis., Competing Interests: The authors have declared that no competing interests exist.

Spatial structure is known to affect the evolution of social behaviour, but little is known on how this evolution depends on simultaneous competition for resources. In simple models, competition for resources tends to counteract altruism, but ecologically more realistic models suggest that competition for resources might actually reinforce altruism. Here we set up a probabilistic cellular automaton (PCA) model and analyse it using the Correlation Dynamics (CD) approach, to study how competition for resources affects the evolution of altruism. If the resource diffuses across space, spatiallyseparateclustersofrelativesmaystillcompeteforresources, thuscreatingalargercompetitive kernelthantheimmediateneighbourhood. Thisincreasestheseparationofclustersofrelativesandthus reinforces the selection of altruistic behaviour.

Information is a crucial currency for living organisms as it allows them to adjust their behaviour to environmental fluctuations. Thus, natural selection should have favoured the capacity of collecting information from different sources, including social interactions whereby individuals could quickly gain reliable information. However, such conditions may also favour the gathering of potentially detrimental information, such as false or misinterpreted accounts of environmental and social phenomena such as rumours, which may spread via informational cascades. We applied ecological and evolutionary principles to investigate how the propagation of social information at a populational level affects the propensity to assimilate it, here defined as the gullibilty. Our results show that the evolution of an individual's susceptibility to assimilate information strongly depends on eco-evolutionary feedbacks, in particular when both useful and detrimental information circulate. We discuss our results regarding the different information transmission mechanisms involved with particular attention to specific cases of social learning., (© 2018 The Author(s).)

In kin selection theory in the evolution of social behaviours, the relatedness between interacting individuals is influenced by the spatial structure of the population. It is generally considered that in the 'viscous' population competition among individuals tends to suppress the evolution of altruism. We consider that more complex interactions produce specific spatial patterns in the presence of competitive interaction, which could alter the process of kin selection in a given space. Here, we theoretically studied the joint evolution of altruism and resource exploitation in a spatially structured population. The simulations indicated that joint evolution can result in self-organisation of regularly arranged cluster structures, which creates a new 'level of selection', and significantly promotes the evolution of altruism through a promotion of kin selection. The analysis also suggested synergetic effects of the joint evolution of two traits, including an evolutionary suicide of the population., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Like many other bacteria, Pseudomonas aeruginosa sequesters iron from the environment through the secretion, and subsequent uptake, of iron-binding molecules. As these molecules can be taken up by other bacteria in the population than those who secreted them, this is a form of cooperation through a public good. Traditionally, this problem has been studied by comparing the relative fitnesses of siderophore-producing and non-producing strains, but this gives no information about the fate of strains that do produce intermediate amounts of siderophores. Here, we investigate theoretically how the amount invested in this form of cooperation evolves. We use a mechanistic description of the laboratory protocols used in experimental evolution studies to describe the competition and cooperation of the bacteria. From this dynamical model we derive the fitness following the adaptive dynamics method. The results show how selection is driven by local siderophore production and local competition. Because siderophore production reduces the growth rate, local competition decreases with the degree of relatedness (which is a dynamical variable in our model). Our model is not restricted to the analysis of small phenotypic differences and allows for theoretical exploration of the effects of large phenotypic differences between cooperators and cheats. We predict that an intermediate ESS level of cooperation (molecule production) should exist. The adaptive dynamics approach allows us to assess evolutionary stability, which is often not possible in other kin-selection models. We found that selection can lead to an intermediate strategy which in our model is always evolutionarily stable, yet can allow invasion of strategies that are much more cooperative. Our model describes the evolution of a public good in the context of the ecology of the microorganism, which allows us to relate the extent of production of the public good to the details of the interactions., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

For a communication system to be stable, senders should convey honest information. Providing dishonest information, however, can be advantageous to senders, which imposes a constraint on the evolution of communication systems. Beyond single populations and bitrophic systems, one may ask whether stable communication systems can evolve in multitrophic systems. Consider cross-species signalling where herbivore-induced plant volatiles (HIPVs) attract predators to reduce the damage from arthropod herbivores. Such plant signals may be honest and help predators to identify profitable prey/plant types via HIPV composition and to assess prey density via the amount of HIPVs. There could be selection for dishonest signals that attract predators for protection from possible future herbivory. Recently, we described a case in which plants release a fixed, high amount of HIPVs independent of herbivore load, adopting what we labelled a 'cry-wolf' strategy. To understand when such signals evolve, we model coevolutionary interactions between plants, herbivores and predators, and show that both 'honest' and 'cry-wolf' types can emerge, depending on the assumed plant-herbivore encounter rates and herbivore population density. It is suggested that the 'cry-wolf' strategy may have evolved to reduce the risk of heavy damage in the future. Our model suggests that eco-evolutionary feedback loops involving a third species may have important consequences for the stability of this outcome., (© 2015 The Author(s).)

The social amoeba Dictyostelium discoideum is widely studied for its multicellular development program as a response to starvation. Aggregates of up to 10 (6) cells form fruiting bodies containing (i) dormant spores (~80%) that can persist for months in the absence of nutrients, and (ii) dead stalk cells (~20%) that promote the dispersion of the spores towards nutrient-rich areas. It is often overlooked that not all cells aggregate upon starvation. Using a new quantitative approach based on time-lapse fluorescence microscopy and a low ratio of reporting cells, we have quantified this fraction of non-aggregating cells. In realistic starvation conditions, up to 15% of cells do not aggregate, which makes this third cell fate a significant component of the population-level response of social amoebae to starvation. Non-aggregating cells have an advantage over cells in aggregates since they resume growth earlier upon arrival of new nutrients, but have a shorter lifespan under prolonged starvation. We find that phenotypic heterogeneities linked to cell nutritional state bias the representation of cells in the aggregating vs. non-aggregating fractions, and thus affect population partitioning. Next, we report that the fraction of non-aggregating cells depends on genetic factors that regulate the timing of starvation, signal sensing efficiency and aggregation efficiency. In addition, interactions between clones in mixtures of non-isogenic cells affect the partitioning of each clone into both fractions. We further build a numerical model to test the evolutionary significance of the non-aggregating cell fraction. The partitioning of cells into aggregating and non-aggregating fractions is optimal in fluctuating environments with an unpredictable duration of starvation periods. Our study highlights the unicellular component of the response of social amoebae to starvation, and thus extends its evolutionary and ecological framework.

The social amoeba Dictyostelium discoideum is widely studied for its multicellular development program as a response to starvation and constitutes a model of choice in microbial cooperation studies. Aggregates of up to 10 (6) cells form fruiting bodies containing two cell types: (i) dormant spores (~80%) that can persist for months in the absence of nutrients, and (ii) dead stalk cells (~20%) that promote the dispersion of the spores towards nutrient-rich areas. It is often overlooked that not all cells aggregate upon starvation. Using a new quantitative approach based on time-lapse fluorescence microscopy and a low ratio of reporting cells, we have quantified this fraction of non-aggregating cells. In realistic starvation conditions, up to 15% of cells do not aggregate, which makes this third cell fate a significant component of the population-level response of social amoebae to starvation. Non-aggregating cells have an advantage over cells in aggregates since they resume growth earlier upon arrival of new nutrients, but have a shorter lifespan under prolonged starvation. We find that phenotypic heterogeneities linked to cell nutritional state bias the representation of cells in the aggregating vs. non-aggregating fractions, and thus regulate population partitioning. Next, we report that the fraction of non-aggregating cells depends on genetic factors that regulate the timing of starvation, signal sensing efficiency and aggregation efficiency. In addition, interactions between clones in mixtures of non-isogenic cells affect the partitioning of each clone into both fractions. We further test the evolutionary significance of the non-aggregating cell fraction. The partitioning of cells into aggregating and non-aggregating fractions is optimal in fluctuating environments with an unpredictable duration of starvation periods. D. discoideum thus constitutes a model system lying at the intersection of microbial cooperation and bet hedging, defining a new frontier in microbiology and evolution studies.

Evolution can be characterized as a process that shapes and maintains information across generations. It is also widely acknowledged that information may play a pivotal role in many other ecological processes. Most of the ecologically relevant information (and some important evolutionary information too) is of a very subjective and analogue kind: individuals use cues that may carry information useful only to them but not to others. This is a problem because most information theory has been developed for objective and discrete information. Can information theory be extended to this theory to incorporate multiple forms of information, each with its own (physical) carriers and dynamics? Here, I will not review all the possible roles that information can play, but rather what conditions an appropriate theory should satisfy. The most promising starting point is provided by entropy measures of conditional probabilities (using the so-called Kullback-Leibler divergence), allowing an assessment of how acquiring information can lead to an increase in fitness. It is irrelevant (to a certain extent) where the information comes from-genes, experience or culture-but it is important to realize that information is not merely subjective but its value should be evaluated in fitness terms, and it is here that evolutionary theory has an enormous potential. A number of important stumbling points remain, however; namely, the identification of whose fitness it concerns and what role the spatio-temporal dynamics plays (which is tightly linked to the nature of the physical carriers of the information and the processes that impact on it).

Vaccines that elicit protective cytotoxic T lymphocytes (CTL) may improve on or augment those designed primarily to elicit antibody responses. However, we have little basis for estimating the numbers of CTL required for sterilising immunity at an infection site. To address this we begin with a theoretical estimate obtained from measurements of CTL surveillance rates and the growth rate of a virus. We show how this estimate needs to be modified to account for (i) the dynamics of CTL-infected cell conjugates, and (ii) features of the virus lifecycle in infected cells. We show that provided the inoculum size of the virus is low, the dynamics of CTL-infected cell conjugates can be ignored, but knowledge of virus life-histories is required for estimating critical thresholds of CTL densities. We show that accounting for virus replication strategies increases estimates of the minimum density of CTL required for immunity over those obtained with the canonical model of virus dynamics, and demonstrate that this modeling framework allows us to predict and compare the ability of CTL to control viruses with different life history strategies. As an example we predict that lytic viruses are more difficult to control than budding viruses when net reproduction rates and infected cell lifetimes are controlled for. Further, we use data from acute SIV infection in rhesus macaques to calculate a lower bound on the density of CTL that a vaccine must generate to control infection at the entry site. We propose that critical CTL densities can be better estimated either using quantitative models incorporating virus life histories or with in vivo assays using virus-infected cells rather than peptide-pulsed targets.

To many pathogenic bacteria, human hosts are an evolutionary dead end. This begs the question what evolutionary forces have shaped their virulence traits. Why are these bacteria so virulent? The coincidental evolution hypothesis suggests that such virulence factors result from adaptation to other ecological niches. In particular, virulence traits in bacteria might result from selective pressure exerted by protozoan predator. Thus, grazing resistance may be an evolutionarily exaptation for bacterial pathogenicity. This hypothesis was tested by subjecting a well characterized collection of 31 Escherichia coli strains (human commensal or extra-intestinal pathogenic) to grazing by the social haploid amoeba Dictyostelium discoideum. We then assessed how resistance to grazing correlates with some bacterial traits, such as the presence of virulence genes. Whatever the relative population size (bacteria/amoeba) for a non-pathogenic bacteria strain, D. discoideum was able to phagocytise, digest and grow. In contrast, a pathogenic bacterium strain killed D. discoideum above a certain bacteria/amoeba population size. A plating assay was then carried out using the E. coli collection faced to the grazing of D. discoideum. E. coli strains carrying virulence genes such as iroN, irp2, fyuA involved in iron uptake, belonging to the B2 phylogenetic group and being virulent in a mouse model of septicaemia were resistant to the grazing from D. discoideum. Experimental proof of the key role of the irp gene in the grazing resistance was evidenced with a mutant strain lacking this gene. Such determinant of virulence may well be originally selected and (or) further maintained for their role in natural habitat: resistance to digestion by free-living protozoa, rather than for virulence per se.

We study the evolution of a spatially structured population with two age classes using spatial moment equations. In the model, adults can either help juveniles by increasing their survival, or adopt a cannibalistic behaviour and consume juveniles. While cannibalism is the sole evolutionary outcome when the population is well-mixed, both cannibalism and parental care can be evolutionarily stable if the population is viscous. Our analysis allows us to make two main technical points. First, we present a method to define invasion fitness in class-structured viscous populations, which allows us to apply adaptive dynamics methodology. Second, we show that ordinary pair approximation introduces an important quantitative bias in the evolutionary model, even on random networks. We propose a correction to the ordinary pair approximation that yields quantitative accuracy, and discuss how the bias associated with this approach is precisely what allows us to identify subtle aspects associated with the evolutionary dynamics of spatially structured populations.

There is ample theoretical and experimental evidence that virulence evolution depends on the immune response of the host. In this article, we review a number of recent studies that attempt to explicitly incorporate the dynamics of the immune system (instead of merely representing it by a single black box parameter) in models for the evolution of parasite virulence. A striking observation is that the type of infection (acute or chronic) is invariably considered to be a constraint that model assumptions have to satisfy rather than as a potential outcome of the interaction of the parasite with its host's immune system. We argue that avoiding making assumptions about the type of infection will lead to a better understanding of infectious diseases, even though a number of fundamental and technical problems remain. Dynamical modeling of the immune system opens a wide range of perspectives: for understanding how the immune system eradicates a parasite (which it does for most pathogens but not for all, HIV being a notorious example of a virus that is not completely eliminated), for studying multiple infections through concomitant immunity, for understanding the emergence and evolution of the immune system in animals, and for evolutionary epidemiology in general (e.g., predicting evolutionary consequences of new therapies and public health policies). We conclude by discussing new approaches based on embedded (or nested) models and identify future perspectives for the modeling of infectious diseases.

Understanding the effect of multiple infections is essential for the prediction (and eventual control) of virulence evolution. Some theoretical studies have considered the possibility that several strains coexist in the same host (coinfection), but few have taken their within-host dynamics explicitly into account. Here, we develop a nested approach based on a simple model for the interaction of parasite strains with their host's immune system. We study virulence evolution by linking the within-host dynamics to an epidemiological framework that incorporates multiple infections. Our model suggests that antigenically similar parasite strains cannot coexist in the long term inside a host. We also find that the optimal level of virulence increases with the efficiency of multiple infections. Finally, we notice that coinfections create heterogeneity in the host population (with susceptible hosts and infected hosts), which can lead to evolutionary branching in the parasite population and the emergence of a hypervirulent parasite strategy. We interpret this result as a parasite specialization to the infectious state of the hosts. Our study has experimental and theoretical implications in a virulence management perspective.

Though it is commonly supposed that there is a trade-off between virulence and transmission, there is little data and little insight into what it should look like. Here, we consider the specific case of vector-borne parasites (inspired by human malaria) and analyse an embedded model to understand how specific life-cycle aspects may affect this trade-off. First, we find that, for such parasites, the transmission function may have an S-shape. Second, we find that the trade-off obtained for vector-borne parasites is less sensitive to parameter variations than the trade-off obtained for directly transmitted parasites. Third, we find that other parasite traits, such as the conversion from replicative to infective stages, could have important epidemiological implications. Finally, we compare the effect of treatments targeting either the asexual or the sexual parasite life-stage.

Offspring quality and quantity are subject to a trade-off. Depending on species, ant colonies can produce either numerous winged queens that start new colonies alone (independent colony foundation [ICF]) or one propagule consisting of reproductives helped by workers (fission). Some species do both. In the Rhytidoponera impressa group, colonies with winged queens use ICF, whereas colonies with mated workers use fission. We assessed the effect of latitude and environment on colonial reproduction by collecting 79 colonies along the Australian east coast. A population dynamical model predicted that from tropical north to temperate south, seasonal fluctuations and harsher conditions should lead to a decrease in ICF success (-48%), making fission a suitable alternative. Yet, our empirical data showed that ICF persists at a higher rate than expected, presumably because it allows aerial dispersal, unlike fission. Investigation of colony demography and measurement of lean and fat weights in virgin winged queens and workers showed that, from north to south, colonies increase quality of queens relative to workers (+50%) at the cost of quantity (-86%). This modification limits the decrease in ICF success (-34%). A tremendous range of offspring phenotypes associated with quality-quantity trade-offs make ants competitive in diverse habitats.

In harsh environments, sessile organisms can make their habitat more hospitable by buffering environmental stress or increasing resource availability. Although the ecological significance of such local facilitation is widely established, the evolutionary aspects have been seldom investigated. Yet addressing the evolutionary aspects of local facilitation is important because theoretical studies show that systems with such positive interactions can exhibit alternative stable states and that such systems may suddenly become extinct when they evolve (evolutionary suicide). Arid ecosystems currently experience strong changes in climate and human pressures, but little is known about the effects of these changes on the selective pressures exerted on the vegetation. Here, we focus on the evolution of local facilitation in arid ecosystems, using a lattice-structured model explicitly considering local interactions among plants. We found that the evolution of local facilitation depends on the seed dispersal strategy. In systems characterized by short-distance seed dispersal, adaptation to a more stressful environment leads to high local facilitation, allowing the population to escape extinction. In contrast, systems characterized by long-distance seed dispersal become extinct under increased stress even when allowed to adapt. In this case, adaptation in response to climate change and human pressures could give the final push to the desertification of arid ecosystems.

Spatial self-structuring has been a focus of recent interest among evolutionary ecologists. We review recent developments in the study of the interplay between spatial self-structuring and evolution. We first discuss the relative merits of the various theoretical approaches to spatial modelling in ecology. Second, we synthesize the main theoretical studies of the evolution of cooperation in spatially structured populations. We show that population viscosity is generally beneficial to cooperation, because cooperators can reap additional benefits from being clustered. A similar mechanism can explain the evolution of honest communication and of reduced virulence in host-parasite interactions. We also discuss some recent innovative empirical results that test these theories. Third, we show the relevance of these results to the general field of evolutionary ecology. An important conclusion is that kin selection is the main process that drives evolution of cooperation in viscous populations. Many results of kin selection theory can be recovered as emergent properties of spatial ecological dynamics. We discuss the implications of these results for the study of multilevel selection and evolutionary transitions. We conclude by sketching some perspectives for future research, with a particular emphasis on the topics of evolutionary branching, criticality, spatial fluctuations and experimental tests of theoretical predictions. Space is the place - Sun Ra.

We investigate the evolution of parental care and cannibalism in a spatially structured population where adults can either help or kill juveniles in their neighborhood. We show that spatial structure can reverse the selective pressures on adult behavior, leading to the evolution of parental care, whereas the nonspatial model predicts that cannibalism is the sole evolutionary outcome. Our analysis emphasizes that evolution of such spatially structured populations is best understood at the level of the cluster of invading mutants, and we define invasion fitness as the growth rate of that cluster. We derive an analytical expression for the selective pressures on the trait and show that relatedness and Hamilton's rule are recovered as emergent properties of the spatial ecological dynamics. When adults can also help other adults, the benefits to each class of recipients are weighted by the class reproductive value, a result consistent with that of other models of kin selection. Finally, we advocate a different approach to moment equations and argue that even though the development of moment closure approximations is a necessary line of research, much-needed ecological and evolutionary insight can be gained by studying the unclosed moment equations.

Though predation, productivity (nutrient richness), spatial heterogeneity, and disturbance regimes are known to influence species diversity, interactions between these factors remain largely unknown. Predation has been shown to interact with productivity and with spatial heterogeneity, but few experimental studies have focused on how predation and disturbance interact to influence prey diversity. We used theory and experiments to investigate how these factors influence diversification of Pseudomonas fluorescens by manipulating both predation (presence or absence of Bdellovibrio bacteriovorus) and disturbance (frequency and intensity of disturbance). Our results show that in a homogeneous environment, predation is essential to promote prey species diversity. However, in most but not all treatments, elevated diversity was transitory, implying that the effect of predation on diversity was strongly influenced by disturbance. Both our experimental and theoretical results suggest that disturbance interacts with predation by modifying the interplay of resource and apparent competition among prey.

Arid ecosystems are liable to undergo sudden discontinuous transitions from a vegetated to a desert state as a result of human pressure and climate change. A predictive framework about the conditions under which such transitions occur is lacking. Here, we derive and analyze a general model describing the spatial dynamics of vegetation in arid ecosystems considering local facilitation as an essential process. We investigate the conditions under which continuous or discontinuous transitions from a vegetated to a desert state are likely to occur. We focus on arid ecosystems but our approach is sufficiently general to be applied to other ecosystems with severe environmental conditions. The model exhibits bistability and vegetation patchiness. High local facilitation decreases the risk of discontinuous transitions. Moreover, for arid ecosystems where local facilitation is a driving process, vegetation patchiness indicates proximity to a transition point, but does not allow distinguishing between continuous and discontinuous transitions.

We investigate the evolution of manipulation of host dispersal behaviour by parasites using spatially explicit individual-based simulations. We find that when dispersal is local, parasites always gain from increasing their hosts' dispersal rate, although the evolutionary outcome is determined by the costs-to-benefits ratio. However, when dispersal can be non-local, we show that parasites investing in an intermediate dispersal distance of their hosts are favoured even when the manipulation is not costly, due to the intrinsic spatial dynamics of the host-parasite interaction. Our analysis highlights the crucial importance of ecological spatial dynamics in evolutionary processes and reveals the theoretical possibility that parasites could manipulate their hosts' dispersal.

The occurrence of polygyny and polyandry in social insects has long puzzled evolutionary biologists. If cooperation requires genetic relatedness, how do we explain the occurrence and maintenance of mechanisms that reduce the degree of relatedness among colony members? A much-discussed hypothesis states that genetically diverse colonies are more resistant to parasitism than homogenous colonies because genetic diversity reduces the spread of a disease within a colony. However, as we will argue in this note, a necessary condition for the parasite hypothesis is that genetically heterogeneous colonies have a larger suite of parasites that are capable of infecting them. This implicit relationship is important because it implies that even if the cost per infection is reduced, this may not be sufficient to offset the increased rate of acquiring infections. The advantages of genetic heterogeneity as a defense against parasites thus may not be as big as commonly thought.

The evolution of altruism, a behaviour that benefits others at one's own fitness expense, poses a darwinian paradox. The paradox is resolved if many interactions are with related individuals so that the benefits of altruism are reaped by copies of the altruistic gene in other individuals, a mechanism called kin selection. However, recognition of altruists could provide an alternative route towards the evolution of altruism. Arguably the simplest recognition system is a conspicuous, heritable tag, such as a green beard. Despite the fact that such genes have been reported, the 'green beard effect' has often been dismissed because it is unlikely that a single gene can code for altruism and a recognizable tag. Here we model the green beard effect and find that if recognition and altruism are always inherited together, the dynamics are highly unstable, leading to the loss of altruism. In contrast, if the effect is caused by loosely coupled separate genes, altruism is facilitated through beard chromodynamics in which many beard colours co-occur. This allows altruism to persist even in weakly structured populations and implies that the green beard effect, in the form of a fluid association of altruistic traits with a recognition tag, can be much more prevalent than hitherto assumed.

Most models of virulence evolution assume that a parasite cannot raise its transmission rate without causing more harm to its host. However, the existence of such trade-off relationships has recently been challenged. Here, we study how a trade-off can emerge from a model that explicitly incorporates within-host dynamics. We find that the existence and the convexity of the trade-off are robust, which implies a definite level of evolutionarily stable virulence (ESV) for the parasite. However, we also show that the dependence of the ESV on parameter values may be very strong. One possible consequence of this sensitivity is that relationships between transmission and virulence observed across populations need not conform to the patterns expected on the basis of a common (fixed) trade-off. We discuss possible experiments and implications of our results for the development of virulence management strategies.

Temporal and spatial variations of the environment are important factors favoring the evolution of dispersal. With few exceptions, these variations have been considered to be exclusively fluctuations of habitat quality. However, since the presence of conspecifics forms part of an individual's environment, demographic stochasticity may be a component of this variability as well, in particular when local populations are small. To study this effect, we analyzed the evolution of juvenile dispersal in a metapopulation model in which habitat quality is constant in space and time but occupancy fluctuates because of demographic stochasticity. Our analysis extends previous studies in that it includes competition of resources and competition for space. Also, juvenile dispersal is not given by a fixed probability but is made conditional on the presence of free territories in a patch, whereas individuals born in full patches will always disperse. Using a combination of analytical and numerical approaches, we show that demographic stochasticity in itself may provide enough variability to favor dispersal even from patches that are not fully occupied. However, there is no simple relationship between the evolution of dispersal and various indicators of demographic stochasticity. Selected dispersal depends on all aspects of the life-history profile, including kin selection.

We investigate how the evolution of communication strategies affects signal credibility when there is common interest as well as a conflict between communicating individuals. Taking alarm calls as an example, we show that if the temptation to cheat is low, a single signal is used in the population. If the temptation increases cheaters will erode the credibility of a signal, and an honest mutant using a different signal ('a private code') will be very successful until this, in turn, is cracked by cheaters. In such a system, signal use fluctuates in time and space and hence the meaning of a given signal is not constant. When the temptation to cheat is too large, no honest communication can maintain itself in a Tower of Babel of many signals. We discuss our analysis in the light of the Green Beard mechanism for the evolution of altruism.

Recent studies indicate that the time required for virus-infected cells to become vulnerable for the activity of CTL is of significance for the capacity of CTL to control ongoing viral reproduction. To investigate whether this applies to the effectiveness of HIV-1-specific CTL, we measured virus production in cultures containing CD4(+) T cells inoculated with HIV at low multiplicity of infection, and CTL directed against an early protein, Rev, or a late protein, RT. The Rev-specific CTL prevented at least 2 log(10) more HIV-1 production, in 10 days, than similar numbers of RT-specific CTL. To study how CTL effectiveness depends on variations in the potency of effector functions and kinetics of HIV protein expression, we developed a mathematical model describing CTL-target cell interactions during successive infection cycles. The results show that substantially higher CTL-mediated target cell elimination rates are required to achieve control as there is less time for CTL to act before infected cells release progeny virions. Furthermore, in vitro experiments with HIV recombinant viruses showed that the RT-specific CTL were at least as effective as the Rev-specific CTL, but only if the RT epitope was expressed as part of the early protein Nef. Together these results indicate that CTL control ongoing HIV reproduction more effectively if they are able to recognize infected cells earlier during individual viral replication cycles. This provides rationale for immunization strategies that aim at inducing, boosting or skewing CTL responses to early regulatory proteins in AIDS vaccine development.

We analyze the evolutionary consequences of host resistance (the ability to decrease the probability of being infected by parasites) for the evolution of parasite virulence (the deleterious effect of a parasite on its host). When only single infections occur, host resistance does not affect the evolution of parasite virulence. However, when superinfections occur, resistance tends to decrease the evolutionarily stable (ES) level of parasite virulence. We first study a simple model in which the host does not coevolve with the parasite (i.e., the frequency of resistant hosts is independent of the parasite). We show that a higher proportion of resistant host decreases the ES level of parasite virulence. Higher levels of the efficiency of host resistance, however, do not always decrease the ES parasite virulence. The implications of these results for virulence management (evolutionary consequences of public health policies) are discussed. Second, we analyze the case where host resistance is allowed to coevolve with parasite virulence using the classical gene-for-gene (GFG) model of host-parasite interaction. It is shown that GFG coevolution leads to lower parasite virulence (in comparison with a fully susceptible host population). The model clarifies and relates the different components of the cost of parasitism: infectivity (ability to infect the host) and virulence (deleterious effect) in an evolutionary perspective.

Aggregated spatial distributions of prey and predators promote stability of the otherwise unstable Nicholson-Bailey model. Nevertheless, when both predators and prey choose patches in an ideal free way, sufficiently aggregated distributions will only arise if patch quality (e.g., reflected in prey fecundity) is very heterogeneous. This requirement profoundly limits the possibilities for simultaneous population dynamical and evolutionary stability. However, stability is not necessary for coexistence. Under cyclic or chaotic dynamics, the rate at which species change their distributions becomes important. Here we consider two endpoints of a continuum of rates. The first is "rigid" selection of patch types based on the expected long-term distribution of conditions. The second is "flexible" selection based on current conditions. We carried out systematic surveys of the population-level consequences of coevolutionarily stable patch-selection strategies for different combinations of rigid and flexible strategies of predator and prey. First, if both prey and predators have rigid strategies, the evolutionary end result is either a stable population dynamical equilibrium or diverging cycles eventually leading to extinction. Second, for the case with rigid prey and flexible predators, the persistence boundary in parameter space is shifted from the boundaries obtained for rigid predators. The mechanism underlying persistence is different in that the flexible strategies of the predators destabilize the equilibrium, while the evolutionary response of the rigid prey leads to reduced cycles. Third, if both prey and predators are flexible, simulations lead either to chaotic fluctuations or to extinction (but not to stable equilibria, nor to limit cycles), and the conditions for coexistence are much wider than those under rigid patch selection. These simulations suggest that information constraints on adaptive patch choice have a major impact on predator-prey persistence under nonequilibrium conditions. We discuss how these predictions can be tested by field observations on expansions and contractions in dietary range (or habitat range) in relation to population fluctuations.