Evolution of culture-dependent discriminate sociality: a gene-culture coevolutionary model

Animals behave cooperatively towards certain conspecifics while being indifferent or even hostile to others. The distinction is made primarily according to kinship as predicted by the kin selection theory. With regards to humans, however, this is not always the case; in particular, humans sometimes exhibit a discriminate sociality on the basis of culturally transmitted traits, such as personal ornaments, languages, rituals, etc. This paper explores the possibility that the human faculty of cultural transmission and resultant cultural variation among individuals may have facilitated the evolution of discriminate sociality in humans. To this end, a gene-culture coevolutionary model is developed focusing on competition over control of resource as a context in which discriminate sociality may have evolved. Specifically, two types of culture-dependent discriminate sociality are considered: ingroup favouritism, with ingroup and outgroup being distinguished by the presence or absence of a cultural trait; and prestige hierarchies, with the prestige being conferred on the bearer of a cultural trait. The model specifies the conditions under which emergence and evolutionary stability of the two types of discriminate sociality are promoted by the presence of cultural variation among individuals.     

Stasis: a coevolutionary model

Traditional neo-Darwinian explanations for stasis involve limited variation, developmental constraints and stabilizing selection. Of these, stabilizing selection is regarded as the mechanism operating most widely. Arguments based on stabilizing selection, however, implicitly assume a one-way evolutionary relationship between organism and environment. In this paper, I suggest that stasis may arise in a number of different ways as a result of organism/environment coevolution. The chief causes of stasis may be the attainment of coadapted equilibria between organism and environment and periods of quiescence within and between arms races. I also suggest that many cases of stasis in the fossil records may be apparent rather than real due to a reliance on gross morphological trends and that apparently large environmental changes during which stasis persists may not reflect change in the coadapted components of the organisms' environment.     

Metal hyperaccumulation: a model system for coevolutionary studies

Recent years have seen a flurry of research activity concerning the hyperaccumulation of heavy metals by plants. Much of the interest in hyperaccumulation has been fueled by the commercial potential of phytoremediation, the use of plants to clean up contaminated soils (Baker et al ., 1994; Salt et al ., 1995; Chaney et al ., 1997, 2000). These applications have in turn spurred many studies of the genetics and physiology of metal uptake (e.g. Krämer et al ., 1996; Lasat et al ., 1996; Salt & Krämer, 1999; Baker et al ., 2000; see also Krämer, 2000; Lombi et al ., 2000). Although phytoremediation provides an intriguing and potentially profitable backdrop, the ecology and evolution of hyperaccumulation in natural populations are interesting subjects in their own right. Two papers in this issue (Ghaderian et al ., pp. 219–224; Davis & Boyd, pp. 211–217) are exciting contributions to the growing consensus that hyperaccumulation may act as a defense against herbivores and pathogens, and suggest that hyperaccumulation may become a model system for research in this area.     

Evolution of style-length variability in figs and optimization of ovipositor length in their pollinator wasps: A coevolutionary model

Monoecious figs reward their pollinators—agaonid wasps—by allocating a proportion of the flowers for egg laying, and retain the rest for seed production. It has been suggested that these proportions could be regulated by producing short-styled and long-styled flowers such that pollinator wasps could only use the former as their ovipositor does not reach the ovules of the latter. Thus the wasps can lay eggs only in the short-styled flowers and raise their offspring, and the ovules of uninfested, long-styled flowers can develop into seeds. This implied that figs bear dimorphic female flowers, with a bimodal distribution of style length. However, recent studies have shown that style length is distributed normally, with no evidence of bimodality. Therefore the regulation of allocation of flowers to the wasps does not seem to be through the production of two distinct kinds of female flowers. In this article we suggest that two factors govern the proportion of flowers rewarded to the wasps: (i) passive regulation, which is a consequence of the optimization of wasp ovipositor length, and (ii) active regulation, where figs are selected to enhance the variance of style length. We show that these arguments lead to certain predictions about the optimum ovipositor length, the proportion of the flowers available to the wasps, and the coefficient of variation of style length. We also show that data for 18 fig-wasp associations conform well with these predictions. We finally suggest that the regulatory process outlined here can be extended to evolution of style length in dioecious fig species also.     

Evolution of biochemical systems with specific chiralities: a model involving territorial behaviour

In a racemic assembly of stereospecifically reproducing organisms, enantiomeric forms would separate territorially if each organism consisted of two parts, one mobile and the other sessile. Partial destruction could then readily lead to the eventual dominance of organisms of one chirality.     

Modeling of the forest insect population size: a game theory approach

A game theory model of insect population dynamics is proposed. For the case when the population may be in one of two states: when physiological processes are directed to growth and reproduction, and when physiological processes are directed to the development of defense reactions, outbreaks of mass reproduction of insect populations may occur in conditions when population and environment have the "memory", and the state of population and environment depends on their state at the previous time moment. In the framework of the model, the well known effect of insect phase variation during the outbreak of reproduction is explained.     

Mate-guarding in a promiscuous insect: species discrimination influences context-dependent behaviour

Mating strategy is often informed by social context. However, information on social environment may be sensitive to interference by nearby heterospecifics, a process known as reproductive interference (RI). When heterospecific individuals are present in the environment, failures in species discrimination can lead to sub-optimal mating behaviours, such as misplaced courtship, misplaced rivalry behaviours, or heterospecific copulation attempts. All aspects of mating behaviour that are influenced by social context may be prone to RI, including copulatory behaviours associated with mate-guarding in the presence of possible competitors. Here we investigate the effect of three heterospecifics on the mate-guarding behaviour of male Lygaeus equestris seed bugs. We find that, despite previously reported heterospecific mating harassment amongst these species of lygaeid bug, male L. equestris are able to effectively distinguish rival conspecific males from heterospecifics. Thus, heterospecific mating attempts in this group may reflect selection on males to mate opportunistically, rather than a failure of species discrimination.     

Evolution of egg dumping in a subsocial insect

Egg dumping, or abandonment of eggs and young to the care of other conspecifics, frees individuals from costs of maternal care while potentially imposing energetic and ecological costs on egg recipients. It is not clear, however, that egg dumping necessarily represents selfish manipulation of egg recipients, and in some ecological contexts, recipients may benefit from enlarged broods. Thus, egg dumping may either be mutually beneficial for dumpers and recipients or entail costs for dumpers that are compensated by other means, such as improving reproduction of genetically related egg recipients. Here I use field experiments to test the relative importance of manipulation (i.e., "parasitism"), mutualism, and kin selection in the evolution of egg dumping in the tingid lace bug Gargaphia solani. In support of mutualism and kin selection, I found that reproduction of egg recipient G. solani benefits from brood enlargement, most likely because eggs and gregarious nymphs find safety in greater numbers. But contrary to both parasitism and mutualism, egg dumper reproduction was not improved by offspring abandonment. Indeed, dumpers laid smaller clutches than recipients, and dumpers did not convert a survival advantage into greater future reproduction. Genetic analyses of a natural G. solani population revealed, however, that dumpers are related to their egg recipients. Moreover, Hamilton's rule showed that egg-dumping G. solani earn sufficient indirect genetic benefits for kin selection to favor the behavior. Thus, egg dumping in some species may be kin-selected cooperation rather than parasitism or mutualism.     

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.     

Evolution of learned strategy choice in a frequency-dependent game

In frequency-dependent games, strategy choice may be innate or learned. While experimental evidence in the producer-scrounger game suggests that learned strategy choice may be common, a recent theoretical analysis demonstrated that learning by only some individuals prevents learning from evolving in others. Here, however, we model learning explicitly, and demonstrate that learning can easily evolve in the whole population. We used an agent-based evolutionary simulation of the producer-scrounger game to test the success of two general learning rules for strategy choice. We found that learning was eventually acquired by all individuals under a sufficient degree of environmental fluctuation, and when players were phenotypically asymmetric. In the absence of sufficient environmental change or phenotypic asymmetries, the correct target for learning seems to be confounded by game dynamics, and innate strategy choice is likely to be fixed in the population. The results demonstrate that under biologically plausible conditions, learning can easily evolve in the whole population and that phenotypic asymmetry is important for the evolution of learned strategy choice, especially in a stable or mildly changing environment.     

Evolution of host resistance: looking for coevolutionary hotspots at small spatial scales

Natural plant populations are often found to be extremely diverse in their resistance to pathogens. While the potential of pathogens in driving the evolution of resistance in hosts has been widely recognized, empirical evidence linking disease dynamics to host population genetic structure has remained scarce. Here I show that current coevolutionary selection for resistance can be divergent even on a very fine spatial scale. In a natural plant-pathogen metapopulation, disease occurrence patterns were highly aggregated over space and time within host populations. A laboratory inoculation experiment showed higher resistance within areas of the host populations where encounter rates with the pathogen have been high. Higher resistance to sympatric than to allopatric strains of the pathogen suggests that this change has taken place as a response to local selection. These results constitute evidence of adaptive microevolution of resistance resulting from disease epidemics in natural plant-pathogen associations, and highlight the importance of finding the relevant scale at which to address questions of current coevolutionary selection.     

Circadian periodicity of escape behaviour in the mouse: Preliminary chronobiological data

The cosinor methodology previously applied to different studies within the field of ethological rhythms is used in the present report to assess the circadian rhythm of escape behaviour. The peak frequency is located toward the end of the dark span in mice exposed to light between 08.00 and 20.00 h. Such experiments provide a quantitative approach for studying some neurobiological mechanisms regulating adaptive behaviour.     

Evolution of insect olfaction

Neuroethology utilizes a wide range of multidisciplinary approaches to decipher neural correlates of natural behaviors associated with an animal's ecological niche. By placing emphasis on comparative analyses of adaptive and evolutionary trends across species, a neuroethological perspective is uniquely suited to uncovering general organizational and biological principles that shape the function and anatomy of the nervous system. In this review, we focus on the application of neuroethological principles in the study of insect olfaction and discuss how ecological environment and other selective pressures influence the development of insect olfactory neurobiology, not only informing our understanding of olfactory evolution but also providing broader insights into sensory processing.     

A game dynamic model for delayer strategies in vaccinating behaviour for pediatric infectious diseases

Several studies have found that some parents delay the age at which their children receive pediatric vaccines due to perception of higher vaccine risk at the recommended age of vaccination. This has been particularly apparently during the Measles-Mumps-Rubella scare in the United Kingdom. Under a voluntary vaccination policy, vaccine coverage in certain age groups is a potentially complex interplay between vaccinating behaviour, disease dynamics, and age-specific risk factors. Here, we construct an age-structured game dynamic model, where individuals decide whether to vaccinate according to imitation dynamics depending on age-dependent disease prevalence and perceived risk of vaccination. Individuals may be timely vaccinators, delayers, or non-vaccinators. The model exhibits multiple equilibria and a broad range of possible dynamics. For certain parameter regimes, the proportion of timely vaccinators and delayers oscillate in an anti-phase fashion in response to oscillations in infection prevalence. Under an exogenous change to the perceived risk of vaccination as might occur during a vaccine scare, the model can also capture an increase in delayer strategists similar in magnitude to that observed during the Measles-Mumps-Rubella vaccine scare in the United Kingdom. Our model also shows that number of delayers steadily increases with increasing severity of the scare, whereas it saturates to specific value with increases in duration of the scare. Finally, by comparing the model dynamics with and without the option of a delayer strategy, we show that adding a third delayer strategy can have a stabilizing effect on model dynamics. In an era where individual choice—rather than accessibility—is becoming an increasingly important determinant of vaccine uptake, more infectious disease models may need to use game theory or related techniques to determine vaccine uptake.     

Unique coevolutionary dynamics in a predator-prey system

In this paper, we study the predator-prey coevolutionary dynamics when a prey's defense and a predator's offense change in an adaptive manner, either by genetic evolution or phenotypic plasticity, or by behavioral choice. Results are: (1) The coevolutionary dynamics are more likely to be stable if the predator adapts faster than the prey. (2) The prey population size can be nearly constant but the predator population can show very large amplitude fluctuations. (3) Both populations may oscillate in antiphase. All of these are not observed when the handling time is short and the prey's density dependence is weak. (4) The population dynamics and the trait dynamics show resonance: the amplitude of the population fluctuation is the largest when the speed of adaptation is intermediate. These results may explain experimental studies with microorganisms.     

Brood-parasite interactions between great spotted cuckoos and magpies: a model system for studying coevolutionary relationships

Brood parasitism is one of the systems where coevolutionary processes have received the most research. Here, we review experiments that suggest a coevolutionary process between the great spotted cuckoo (Clamator glandarius) and its magpie (Pica pica) host. We focus on different stages of establishment of the relationship, from cuckoos selecting individual hosts and hosts defending their nests from adult cuckoos, to the ability of magpies to detect cuckoo eggs in their nests. Novel coevolutionary insights emerge from our synthesis of the literature, including how the evolution of "Mafia" behaviour in cuckoos does not necessarily inhibit the evolution of host recognition and rejection of cuckoo offspring, and how different populations of black-billed magpies in Europe have evolved specific host traits (e.g. nest and clutch size) as a result of interactions with the great spotted cuckoo. Finally, the results of the synthesis reveal the importance of using a meta-population approach when studying coevolution. This is especially relevant in those cases where gene flow among populations with different degrees of brood parasitism explains patterns of coexistence between defensive and non-defensive host phenotypes. We propose the use of a meta-population approach to distinguish between the "evolutionary equilibrium" hypothesis and the "evolutionary lag" hypothesis.     

Using Drosophila as a model insect

The fruit fly Drosophila melanogaster has become such a popular model organism for studying human disease that it is often described as a little person with wings. This view has been strengthened with the sequencing of the Drosophila genome and the discovery that 60% of human disease genes have homologues in the fruit fly. In this review, I discuss the approach of using Drosophila not only as a model for metazoans in general but as a model insect in particular. Specifically, I discuss recent work on the use of Drosophila to study the transmission of disease by insect vectors and to investigate insecticide function and development.     

Play and the evolution of fairness: a game theory model

Bekoff [J. Consci. Stud. 8 (2001) 81] argued that mammalian social play is a useful behavioral phenotype on which to concentrate in order to learn more about the evolution of fairness. Here, we build a game theoretical model designed to formalize some of the ideas laid out by Bekoff, and to examine whether ‘fair’ strategies can in fact be evolutionarily stable. The models we present examine fairness at two different developmental stages during an individual's ontogeny, and hence we create four strategies—fair at time 1/fair at time 2, not fair at time 1/not fair at time 2, fair at time 1/not fair at time 2, not fair at time 1/fair at time 2. Our results suggest that when considering species where fairness can be expressed during two different developmental stages, acting fairly should be more common than never acting fairly. In addition, when no one strategy was evolutionarily stable, we found that all four strategies we model can coexist at evolutionary equilibrium. Even in the absence of an overwhelming database from which to test our model, the general predictions we make have significant implications for the evolution of fairness.