Social evolution in the shadow of asymmetrical relatedness.

The persistence of altruism and spite remains an enduring problem of social evolution. It is well known that selection for these actions depends on the structure of the population-that is, on actors' genetic relationships to recipients and to the 'neighbourhood' upon which the effects of their actions redound. Less appreciated, however, is that population structure can cause genetic asymmetries between partners whereby the relatedness (defined relative to the neighbourhood) of an individual i to a partner j will differ from the relatedness of j to i.

Measures of relative fitness of social behaviors in finite structured population models.

How should we measure the relative selective advantage of different behavioral strategies? The various approaches to this question have fallen into one of the following categories: the fixation probability of a mutant allele in a wild type population, some measures of gene frequency and gene frequency change, and a formulation of the inclusive fitness effect. Countless theoretical studies have examined the relationship between these approaches, and it has generally been thought that, under standard simplifying assumptions, they yield equivalent results. Most of this theoretical work, however, has assumed homogeneity of the population interaction structure--that is, that all individuals are equivalent.

Local mate competition and transmission bottlenecks: a new model for understanding malaria parasite and other sex ratios.

The local mate competition model from sex ratio theory predicts female-biased sex ratios in populations that are highly subdivided during mating, and is thought to accord well with the population structure of malaria parasites. However, the selective advantage of female-biased sex ratios comes from the resulting increase in total reproductive output, an advantage the transmission biology of malaria parasite likely reduces. We develop a mathematical model to determine how bottlenecks in transmission that cause diminishing fitness returns from female production affect sex ratio evolution.

Hamilton's inclusive fitness in finite-structured populations.

Hamilton's formulation of inclusive fitness has been with us for 50 years. During the first 20 of those years attention was largely focused on the evolutionary trajectories of different behaviours, but over the past 20 years interest has been growing in the effect of population structure on the evolution of behaviour and that is our focus here. We discuss the evolutionary journey of the inclusive-fitness effect over this epoch, nurtured as it was in an essentially homogeneous environment (that of 'transitive' structures) having to adapt in different ways to meet the expectations of heterogeneous structures.

Enhanced kin recognition through population estimation.

Kin recognition systems enable organisms to predict genetic relatedness. In so doing, they help to maximize the fitness consequences of social actions. Recognition based on phenotypic similarity-a process known as phenotype matching-is thought to depend upon information about one's own phenotype and the phenotypes of one's partners.

Relatedness with different interaction configurations.

In an inclusive fitness model of social behaviour, a key concept is that of the relatedness between two interactants. This is typically calculated with reference to a "focal" actor taken to be representative of all actors, but when there are different interaction configurations, relatedness must be constructed as an average over all such configurations. We provide an example of such a calculation in an island model with local reproduction but global mortality, leading to variable island size and hence variable numbers of individual interactions.

Joint evolution of multiple social traits: a kin selection analysis.

General models of the evolution of cooperation, altruism and other social behaviours have focused almost entirely on single traits, whereas it is clear that social traits commonly interact. We develop a general kin-selection framework for the evolution of social behaviours in multiple dimensions. We show that whenever there are interactions among social traits new behaviours can emerge that are not predicted by one-dimensional analyses.

Evolution of cooperation by phenotypic similarity.

The emergence of cooperation in populations of selfish individuals is a fascinating topic that has inspired much work in theoretical biology. Here, we study the evolution of cooperation in a model where individuals are characterized by phenotypic properties that are visible to others. The population is well mixed in the sense that everyone is equally likely to interact with everyone else, but the behavioral strategies can depend on distance in phenotype space.

From inclusive fitness to fixation probability in homogeneous structured populations.

The methods of inclusive fitness provide a powerful analysis of the action of selection on social behaviour. The key component of this analysis is the concept of relatedness R. In infinite populations, a standard method of calculating relatedness coefficients is through coefficients of consanguinity using the notion of genetic identity by descent.

Evolution of cooperation in a finite homogeneous graph.

Recent theoretical studies of selection in finite structured populations have worked with one of two measures of selective advantage of an allele: fixation probability and inclusive fitness. Each approach has its own analytical strengths, but given certain assumptions they provide equivalent results. In most instances the structure of the population can be specified by a network of nodes connected by edges (that is, a graph), and much of the work here has focused on a continuous-time model of evolution, first described by ref.

Spiteful soldiers and sex ratio conflict in polyembryonic parasitoid wasps.

The existence of spiteful behaviors remains controversial. Spiteful behaviors are those that are harmful to both the actor and the recipient, and they represent one of the four fundamental types of social behavior (alongside selfishness, altruism, and mutual benefit). It has generally been assumed that the conditions required for spite to evolve are too restrictive, and so spite is unlikely to be important.

The evolutionary consequences of plasticity in host-pathogen interactions.

Interactions between individuals such as hosts and pathogens are often characterized by substantial phenotypic plasticity. Pathogens sometimes alter their exploitation strategies in response to defensive strategies adopted by their host and vice versa. Nevertheless, most game-theoretic models developed to explain the evolution of pathogen and host characteristics assume that no such plasticity occurs.

Sex allocation and dispersal in a heterogeneous two-patch environment.

We investigate the evolution of sex allocation and dispersal in a two-habitat environment using a game theoretic analysis. One habitat is of better quality than the other and increased habitat quality influences the competitive ability of offspring in a sex-specific manner. Unlike previous work, we allow incomplete mixing of the population during mating.

Modelling information exchange in worker-queen conflict over sex allocation.

We investigate the conflict between queen and worker over sex allocation, specifically the allocation of the queen's eggs between workers and reproductives and the allocation of the reproductive eggs between male and female. In contrast to previous models, we allow workers to observe and use information about the strategy of the queen. We consider three conflict models: simultaneous (no information exchange), sequential (a one-way information exchange) and negotiated (an iterated two-way information exchange).

A kin-selection approach to the resolution of sex-ratio conflict between mates.

We investigate an instance of conflict between mates over the sex ratio of their brood. We construct a kin-selection model for the evolution of the sex ratio assuming local resource competition (LRC) among females. We explore two basic scenarios: (a) the case where parents make simultaneous sex-ratio decisions (the simultaneous allocation model); and (b) the case where parental sex-ratio decisions occur one after the other (the sequential allocation model).

Fitness and evolutionary stability in game theoretic models of finite populations.

We investigate two methods of measuring fitness in evolutionary games played among members of a finite population. Classical notions of stability account for the action of selection only, and use immediate reproductive gains as a measure of fitness. This classical interpretation of fitness is what we call reproductive fitness (RF), and is found in the early studies of evolutionary stability in finite populations.

Stability in negotiation games and the emergence of cooperation.

Consider a two-player game in which each player contributes a costly resource to the common good of the pair. For such contests, the Nash equilibrium contribution, x*, is one for which neither player can increase its pay-off by unilaterally altering its contribution from x*. We study an elaboration of this game, which allows the players to exchange x-offers back and forth in a negotiation phase until they converge to a final pair of contributions, x1 and x2.

SEX-RATIO BIAS WITH ASYMMETRIC EXCHANGE OF POLLEN BETWEEN DEMES.

In a patch of hermaphroditic plants, with a low level of pollen migration between patches, a prevailing wind creates a gradient, within the patch, in the strength of local competition among pollen for reproductive success. This leads to a sex ratio gradient, with a male (pollen) bias in downwind individuals, which can be quite strong even for large patches. The effect can be understood as follows: downwind individuals have relatively low reproductive value and respond by putting more resources into the gamete (pollen) with the best long-range (extra-patch) reproductive success.