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PubMed1989

Structural instability of models of sexual selection.

Models for sexual selection by the coevolution of female preference and a male trait have as a generic feature the existence of a neutrally stable line of equilibria up and down which the system can drift at random. However, this feature is structurally unstable since it is destroyed by introducing either mutation or weak direct selection on female preference into the model, to be replaced by a single equilibrium (or a finite set of disconnected equilibria). This process is investigated in detail under a simple but general population genetic model. It is concluded that the level of female preference is determined by mutation, selection, and genetic drift acting directly on it, and drags the male trait along with it along the line of equilibria. More attention should be paid to selective forces acting directly on female preference.

Biological Evolution↗
PubMed1996

Sexually antagonistic male adaptation triggered by experimental arrest of female evolution.

Each sex is part of the environment of the other sex. This may lead to perpetual coevolution between the sexes, when adaptation by one sex reduces fitness of the other. Indirect evidence comes from experiments with Drosophila melanogaster indicating that seminal fluid reduces the competitive ability of sperm from other males, thereby increasing male fitness. It also reduces a female's propensity to remate and increase her egg-laying rate. In contrast to these benefits to males, seminal fluid has substantial toxic side effects in females, with increasing quantity leading to decreasing female survival. Here I show that when female D. melanogaster are experimentally prevented from coevolving with males, males rapidly adapt to the static female phenotype. This male adaptation leads to a reduction in female survivorship, which is mediated by an increased rate of remating and increased toxicity of seminal fluid.

Adaptation, Physiological↗
PubMed2000

On meme--gene coevolution.

In this article we examine the effects of the emergence of a new replicator, memes, on the evolution of a pre-existing replicator, genes. Using a version of the NKCS model we examine the effects of increasing the rate of meme evolution in relation to the rate of gene evolution, for various degrees of interdependence between the two replicators. That is, the effects of memes' (suggested) more rapid rate of evolution in comparison to that of genes is investigated using a tunable model of coevolution. It is found that, for almost any degree of interdependence between the two replicators, as the rate of meme evolution increases, a phase transition-like dynamic occurs under which memes have a significantly detrimental effect on the evolution of genes, quickly resulting in the cessation of effective gene evolution. Conversely, the memes experience a sharp increase in benefit from increasing their rate of evolution. We then examine the effects of enabling genes to reduce the percentage of gene-detrimental evolutionary steps taken by memes. Here a critical region emerges as the comparative rate of meme evolution increases, such that if genes cannot effectively select memes a high percentage of the time, they suffer from meme evolution as if they had almost no selective capability.

Animals↗
PubMed2001

Coevolution in host-parasite systems: behavioural strategies of slave-making ants and their hosts.

Recently, avian brood parasites and their hosts have emerged as model systems for the study of host-parasite coevolution. However, empirical studies of the highly analogous social parasites, which use the workers of another eusocial species to raise their own young, have never explicitly examined the dynamics of these systems from a coevolutionary perspective. Here, we demonstrate interpopulational variation in behavioural interactions between a socially parasitic slave-maker ant and its host that is consistent with the expectations of host-parasite coevolution. Parasite pressure, as inferred by the size, abundance and raiding frequency of Protomognathus americanus colonies, was highest in a New York population of the host Leptothorax longispinosus and lowest in a West Virginia population. As host-parasite coevolutionary theory would predict, we found that the slave-makers and the hosts from New York were more effective at raiding and defending against raiders, respectively, than were conspecifics from the West Virginia population. Some of these variations in efficacy were brought about by apparently simple shifts in behaviour. These results demonstrate that defence mechanisms against social parasites can evolve, and they give the first indications of the existence of a coevolutionary arms race between a social parasite and its host.

Animals↗
PubMed2000

Cooperative coevolution: an architecture for evolving coadapted subcomponents.

To successfully apply evolutionary algorithms to the solution of increasingly complex problems, we must develop effective techniques for evolving solutions in the form of interacting coadapted subcomponents. One of the major difficulties is finding computational extensions to our current evolutionary paradigms that will enable such subcomponents to "emerge" rather than being hand designed. In this paper, we describe an architecture for evolving such subcomponents as a collection of cooperating species. Given a simple string-matching task, we show that evolutionary pressure to increase the overall fitness of the ecosystem can provide the needed stimulus for the emergence of an appropriate number of interdependent subcomponents that cover multiple niches, evolve to an appropriate level of generality, and adapt as the number and roles of their fellow subcomponents change over time. We then explore these issues within the context of a more complicated domain through a case study involving the evolution of artificial neural networks.

Adaptation, Physiological↗
PubMed1995

Artificial symbiogenesis.

Symbiosis is the phenomenon in which organisms of different species live together in close association, resulting in a raised level of fitness for one or more of the organisms. Symbiogenesis is the name given to the process by which symbiotic partners combine and unify-forming endosymbioses and then potentially transferring genetic material-giving rise to new morphologies and physiologies evolutionarily more advanced than their constitutents. In this article we begin by using the NKC model of coevolution to examine endosymbiosis and its effect on the evolutionary performance of the partners involved. We are then able to suggest the conditions under which endosymbioses are more likely to occur and why; we find they emerge between organisms within a window of their respective "chaotic gas regimes" and hence that the association represents a more stable state for the partners. The conditions under which gene transfer is more likely to represent an advantage for such endosymbionts are then examined within the same model. We find that, providing a suitable pathway exists, such a process can lead to a more efficient genetic configuration for the symbionts within a window that overlaps that in which endosymbioses occur. Finally, the results are used as grounds for implementing symbiogenesis within artificial evolutionary multiagent systems.

Algorithms↗
PubMed1995

Slow coevolution of a viral pathogen and its diploid host.

We study a population exposed to a lethal infectious disease. Host response is carried at one locus with two alleles while the pathogen occurs in two variants. Based on an SI-type epidemic model we derive explicit equations for the dynamics of each genotype. By assuming small variations in both host and disease, we obtain a separation in time scales between epidemic and evolutionary processes. This allows us to describe explicitly the changes in host and disease gene frequencies. The resulting model has a rich behaviour including multiple stable states and oscillations. However, in the oscillatory situation the model is degenerate excluding the possibility of limit cycles. We show that the degeneracy can only be removed by frequency dependent selection in the pathogen, for example by including direct interaction of virus in a free-living stage. The qualitative conclusions extend to an SIR-type epidemic model, where recovery with immunity from the disease is possible.

Biological Evolution↗
PubMed2002

The ecology of fish parasites with particular reference to helminth parasites and their salmonid fish hosts in Welsh rivers: a review of some of the central questions.

Ecological studies carried out in Welsh rivers on the feeding behaviour of salmonid fish, their helminth parasites and intermediate hosts in the early 1950s and in 1998 have been used as a basis to review the literature dealing with the following questions. First, how are the helminth populations dispersed in space-time? Second, to what extent are the distributional patterns and the life history strategies of the parasites influenced by physicochemical factors? Third, to what extent are populations of helmith parasites in salmonid fish influenced by host characteristics including the genome, sex, age, size, social position and Feeding behaviour? Fourth, are the populations of parasites regulated in a density-dependent manner? Fifth, do the parasites influence the survival and wellbeing of their salmonid hosts and the evolution of sex? Sixth, to what extent is the parasite community influenced by environmental changes including those of an anthropogenic nature and can the parasites be used as bioindicators of pollution? As with most parasites the helminth species found were highly overdispersed thus making it necessary to undertake a log10 (1 + x) conversion for statistical analyses. Statistical analyses confirm that the genome, age and sex of salmonid fish hosts, the station and seasonal change in radiation levels were significant factors in predicting the number of parasites. The evidence given supports the hypothesis that the feeding behaviour and habitat selection by the host fish, their position in the social hierarchy and the overdispersed nature of the transmission sites are the key factors in causing differences in the parasitic fauna related to host species, age, size and sex. Differences in the helminth parasite community related to station can be explained on the basis of differences in water types, sediments and chemistry. Although the evidence presented is in accord with the consensus view that temperature is correlated with seasonal changes in the abundance of many species of helminth parasites, it is argued that it may not be the direct causative mechanism. It is postulated that the life history strategy that results in a decline in abundance of the more vulnerable adult parasites in the gut of the salmonid hosts during the summer has arisen as a result of evolutionary pressures. At this time, the gut environment is particularly inhospitable because of the temperature-related enhancement of the host's immune mechanism and the increased gut turnover rate. In contrast, the larval stages in the immunologically and metabolically more benign intermediate host would be under less intensive selective pressures. It is postulated therefore that evolutionary pressures have caused the parasites to leave the definitive host and concentrate their reproductive efforts in the intermediate hosts during the warmer months. Evidence is given in support of the hypothesis that the parasite populations are regulated in a density-dependent manner and that the regulatory mechanisms may involve the host's immune mechanisms and intraspecies competition and interspecies competition of an exploitative or interference nature. Quantitative studies using 'K' factor analysis and biochemical research to elucidate the nature of the interference mechanisms are required to test this hypothesis. The absence of age-related resistance indicates an old and stable relationship in which the immunosuppressive and immunoavoidance mechanisms of the parasites and hosts, respectively, are in balance. This indicates that the introduction of novel parasites or new genetic strains of host fish could result in harmful epidemics. Despite causing tissue damage, there was no evidence of parasite-induced mortality among the salmonids in the Teifi. This finding is in accord with the generally accepted view that most freshwaters are not troubled by parasite problems. although parasites are present in abundance. In fact, parasite abundance in the salmonid fish in the Teifi was positively correlated with the condition factor and the adipose index. Two testable hypotheses were advanced to explain these observations. First, the more dominant well-conditioned fish in the hierarchy are more likely to acquire parasites because they ingest more food items and spend more time in sheltered habitats with depositing sediments where transmission mainly occurs. Second, the parasites may release factors that stimulate the host's immune and endocrinological systems to produce factors that enhance somatic growth and inhibit reproduction of the host. This benign relationship is considered to be indicative of long-term coevolution. The sex of the fish had a significant influence on the abundance of the parasites in total and also on particular species with the bias in all cases being in favour of the female fish. This review shows that sex bias in parasitism is generally not strong and that male bias in parasitism is not a general rule. Taken as a whole, the results fail to support most of the predictions based on the Hamilton-Zuk and the immunocompetence hypotheses. Possible hypotheses to explain why parasitism tends to be higher in female than in male trout include testosterone immunosuppression, corticosteroid-based immune suppression and differences between the size and behaviour of the sexes. However, the latter two hypotheses have more credence, although testosterone levels are higher in female than male trout. Between the early 1950s and 1998 there has been a marked decline in the prevalence, abundance and diversity of the helminth parasite communities in salmonid fish as well as their intermediate hosts. Possible reasons for these declines include heavy metal pollution, increased acidity and habitat degradation linked to changes in land use. It is concluded that although helminth parasites can provide supplementary information on pollution. the use of biotic indices based on the Biological monitoring working party (BMWP) or River invertebrate prediction and classification system (RIVPACS) methods are preferable. However, as these methods were designed to measure the impact of organic pollution they lack the sensitivity for measuring metal pollution. It is advocated therefore that new biomonitoring methods should be developed to measure the impact of heavy metal pollution using biotic indices based on the sampling of the susceptible invertebrate communities inhabiting depositing sediments in the transmission sites of helminth parasites.

Animals↗
PubMed2002

Ecological bistability and evolutionary reversals under asymmetrical competition.

How does the process of life-history evolution interplay with population dynamics? Almost all models that have addressed this question assume that any combination of phenotypic traits uniquely determine the ecological population state. Here we show that if multiple ecological equilibria can exist, the evolution of a trait that relates to competitive performance can undergo adaptive reversals that drive cyclic alternation between population equilibria. The occurrence of evolutionary reversals requires neither environmentally driven changes in selective forces nor the coevolution of interactions with other species. The mechanism inducing evolutionary reversals is twofold. First, there exist phenotypes near which mutants can invade and yet fail to become fixed; although these mutants are eventually eliminated, their transitory growth causes the resident population to switch to an alternative ecological equilibrium. Second, asymmetrical competition causes the direction of selection to revert between high and low density. When ecological conditions for evolutionary reversals are not satisfied, the population evolves toward a steady state of either low or high abundance, depending on the degree of competitive asymmetry and environmental parameters. A sharp evolutionary transition between evolutionary stasis and evolutionary reversals and cycling can occur in response to a smooth change in ecological parameters, and this may have implications for our understanding of size-abundance patterns.

Animals↗
PubMed2004

Host-parasite coevolution and optimal mutation rates for semiconservative quasispecies.

In this paper, we extend a model of host-parasite coevolution to incorporate the semiconservative nature of DNA replication for both the host and the parasite. We find that the optimal mutation rate for the semiconservative and conservative hosts converge for realistic genome lengths, thus maintaining the admirable agreement between theory and experiment found previously for the conservative model and justifying the conservative approximation in some cases. We demonstrate that, while the optimal mutation rate for a conservative and semiconservative parasite interacting with a given immune system is similar to that of a conservative parasite, the properties away from this optimum differ significantly. We suspect that this difference, coupled with the requirement that a parasite optimize survival in a range of viable hosts, may help explain why semiconservative viruses are known to have significantly lower mutation rates than their conservative counterparts.

Adaptation, Physiological↗
PubMed2006

Bacteriophage migration via nematode vectors: host-parasite-consumer interactions in laboratory microcosms.

Pathogens vectored by nematodes pose serious agricultural, economic, and health threats; however, little is known of the ecological and evolutionary aspects of pathogen transmission by nematodes. Here we describe a novel model system with two trophic levels, bacteriophages and nematodes, each of which competes for bacteria. We demonstrate for the first time that nematodes are capable of transmitting phages between spatially distinct patches of bacteria. This model system has considerable advantages, including the ease of maintenance and manipulation at the laboratory bench, the ability to observe many generations in short periods, and the capacity to freeze evolved strains for later comparison to their ancestors. More generally, experimental studies of complex multispecies interactions, host-pathogen coevolution, disease dynamics, and the evolution of virulence may benefit from this model system because current models (e.g., chickens, mosquitoes, and malaria parasites) are costly to maintain, are difficult to manipulate, and require considerable space. Our initial explorations centered on independently assessing the impacts of nematode, bacterium, and phage population densities on virus migration between host patches. Our results indicated that virus transmission increases with worm density and host bacterial abundance; however, transmission decreases with initial phage abundance, perhaps because viruses eliminate available hosts before migration can occur. We discuss the microbial growth dynamics that underlie these results, suggest mechanistic explanations for nematode transmission of phages, and propose intriguing possibilities for future research.

Animals↗
PubMed1995

Computer models: killing mosquitoes with information.

This paper looks at the relationship between man and mosquitoes from the perspective of coevolution. From this perspective, the primacy of information processing in vector control programs becomes acutely evident. A composite mosquito control program is developed and illustrated to show the benefits derived from incremental increases in information. The use of computer modeling is seen as the next logical step to be taken by vector control personnel to add the next increment of efficiency and effectiveness. This step could well lead to significant reductions or perhaps the elimination of the need for pesticide use. The author encourages the use of computer modeling in teams as the means to learn across disciplines. The feasibility of this approach has been greatly enhanced by the availability of off-the-shelf modeling programs. The author appeals to university and vector control professionals to support students and staff in learning computer modeling techniques.

Animals↗
PubMed2006

Persistent hepatitis C virus infection in vitro: coevolution of virus and host.

The virological and cellular consequences of persistent hepatitis C virus (HCV) infection have been elusive due to the absence of the requisite experimental systems. Here, we report the establishment and the characteristics of persistent in vitro infection of human hepatoma-derived cells by a recently described HCV genotype 2a infectious molecular clone. Persistent in vitro infection was characterized by the selection of viral variants that displayed accelerated expansion kinetics, higher peak titers, and increased buoyant densities. Sequencing analysis revealed the selection of a single adaptive mutation in the HCV E2 envelope protein that was largely responsible for the variant phenotype. In parallel, as the virus became more aggressive, cells that were resistant to infection emerged, displaying escape mechanisms operative at the level of viral entry, HCV RNA replication, or both. Collectively, these results reveal the existence of coevolutionary events during persistent HCV infection that favor survival of both virus and host.

Adaptation, Biological↗
PubMed2006

Self-organized similarity, the evolutionary emergence of groups of similar species.

Ecologists have long been puzzled by the fact that there are so many similar species in nature. Here we show that self-organized clusters of look-a-likes may emerge spontaneously from coevolution of competitors. The explanation is that there are two alternative ways to survive together: being sufficiently different or being sufficiently similar. Using a model based on classical competition theory, we demonstrate a tendency for evolutionary emergence of regularly spaced lumps of similar species along a niche axis. Indeed, such lumpy patterns are commonly observed in size distributions of organisms ranging from algae, zooplankton, and beetles to birds and mammals, and could not be well explained by earlier theory. Our results suggest that these patterns may represent self-constructed niches emerging from competitive interactions. A corollary of our findings is that, whereas in species-poor communities sympatric speciation and invasion of open niches is possible, species-saturated communities may be characterized by convergent evolution and invasion by look-a-likes.

Animals↗
PubMed1998

Evolution of transmission bias in cultural inheritance.

Evolution of transmission bias in cultural inheritance is investigated using simple models of cultural selection. Conventional models of cultural transmission describe cultural changes by incorporating transmission bias and non-vertical pathways into the ordinary population genetic framework. The methodology has been successful in understanding cultural changes in terms of natural selection, but it is difficult to see from the theoretical framework how biased transmission in favor of maladaptive traits might have evolved. To show that ordinary cultural processes lead at times to the evolution of a preference that favors a deleterious cultural variant, this study presents an alternative model of cultural transmission, where cultural elements are transmitted in a manner more like infections in epidemiological transmission. An ordinary equilibrium analysis indicates that, under certain conditions, runaway dynamics emerges and the coevolution of a maladaptive cultural variant and an associated preference in favor of the maladaptive variant is observed. If the preference of an individual does not change during its ontogeny (e.g., if it is transmitted genetically), however, then cultural selection alone does not produce such runaway dynamics, and only those preferences that favor adaptive variants should eventually evolve. Since cultural processes may at times result in a reduction in the fitness of individuals, simplistic adaptive interpretations of culture are unconvincing without detailed specification of the cultural processes involved. Moreover, cultural runaway of this kind may help to explain the existence of traits that are apparently maladaptive at the individual level but may be advantageous for the group. Inferences are also made regarding the observed differences between human and non-human social information transfer.

Adaptation, Physiological↗
PubMed2006

Visualizing coevolution with CIAO plots.

In a previous article, we introduced a number of visualization techniques that we had developed for monitoring the dynamics of artificial competitive coevolutionary systems. One of these techniques involves evaluating the performance of an individual from the current population in a series of trials against opponents from all previous generations, and visualizing the results as a 2D grid of shaded cells or pixels: qualitative patterns in the shading can indicate different classes of coevolutionary dynamics. As this technique involves pitting a current individual against ancestral opponents, we referred to the visualizations as CIAO plots. Since then, a number of other authors studying the dynamics of competitive coevolutionary systems have used CIAO plots or close derivatives to help illuminate the dynamics of their systems, and it has become something of a de facto standard visualization technique. In this very brief article we summarize the rationale for CIAO plots, explain the method of constructing a CIAO plot, and review important recent results that identify significant limitations of this technique.

Artificial Intelligence↗
PubMed2005

Measure dynamics on a one-dimensional continuous trait space: theoretical foundations for adaptive dynamics.

The measure dynamics approach to modelling single-species coevolution with a one-dimensional trait space is developed and compared to more traditional methods of adaptive dynamics and the Maximum Principle. It is assumed that individual fitness results from pairwise interactions together with a background fitness that depends only on total population size. When fitness functions are quadratic in the real variables parameterizing the one-dimensional traits of interacting individuals, the following results are derived. It is shown that among monomorphisms (i.e. measures supported on a single trait value), the continuously stable strategy (CSS) characterize those that are Lyapunov stable and attract all initial measures supported in an interval containing this trait value. In the cases where adaptive dynamics predicts evolutionary branching, convergence to a dimorphism is established. Extensions of these results to general fitness functions and/or multi-dimensional trait space are discussed.

Adaptation, Physiological↗
PubMed2003

Shared control of epidemiological traits in a coevolutionary model of host-parasite interactions.

Most models concerning the evolution of a parasite's virulence and its host's resistance assume that each component of the relationship (transmission, virulence, recovery, etc.) is controlled by either the host or the parasite but not by both. We present a model that describes the coevolution of host and parasite, assuming that the rate of transmission or the virulence depends on both genotypes. The evolution of these traits is constrained by trade-offs that account for costs of defense and attack strategies, in line with previous studies on the separate evolution of the host and the parasite. Considering shared control by the host and the parasite in determining the traits of the relationship leads to several novel predictions. First, the host should evolve maximal investment in defense against parasites with an intermediate replication rate. Second, the evolution of the parasite strongly depends on the way the host's defense is described. Third, the coevolutionary process may lead to decreasing the parasite's virulence as a response to a rise in the host's background mortality, contrary to classical predictions.

Adaptation, Physiological↗