Chris Cannings: A Life in Games

Chris Cannings was one of the pioneers of evolutionary game theory. His early work was inspired by the formulations of John Maynard Smith, Geoff Parker and Geoff Price; Chris recognized the need for a strong mathematical foundation both to validate stated results and to give a basis for extensions of the models. He was responsible for fundamental results on matrix games, as well as much of the theory of the important war of attrition game, patterns of evolutionarily stable strategies, multiplayer games and games on networks. In this paper we describe his work, key insights and their influence on research by others in this increasingly important field. Chris made substantial contributions to other areas such as population genetics and segregation analysis, but it was to games that he always returned. This review is written by three of his students from different stages of his career

A Hawk-Dove game in kleptoparasitic populations

Kleptoparasitism, the parasitism by theft, is a widespread biological phenomenon. In this paper we extend earlier models to investigate a population of conspecifics involved in foraging and, potentially, kleptoparasitism. We assume that the population is composed of two types of individuals, Hawks and Doves. The types differ according to their strategic choices when faced with an opportunity to steal and to resist a challenge. Hawks use every opportunity to steal and they resist all challenges. Doves never resist and never steal. The fitness of each type of individual depends upon various natural parameters, for example food density, the handling time of a food item, density of the population, as well as the duration of potential fights over the food. We find the Evolutionarily Stable States (ESSs) for all arameter combinations and show that there are three possible ESSs, pure Hawks, pure Doves, and a mixed population of Hawks and Doves. We show that for any set of parameter values there is exactly one ESS. We further investigate the relationship between our findings and the classical Hawk-Dove game as defined in Maynard Smith 1982. We also show how our model extends the classical on

A game-theoretical model of kleptoparasitic behavior in an urban gull (Laridae) population

Kleptoparasitism (food stealing) is a significant behavior for animals that forage in social groups as it permits some individuals to obtain resources while avoiding the costs of searching for their own food. Evolutionary game theory has been used to model kleptoparasitism, with a series of differential equation-based compartmental models providing significant theoretical insights into behavior in kleptoparasitic populations. In this paper, we apply this compartmental modeling approach to kleptoparasitic behavior in a real foraging population of urban gulls (Laridae). Field data was collected on kleptoparasitism and a model developed that incorporated the same kleptoparasitic and defensive strategies available to the study population. Two analyses were conducted: 1) An assessment of whether the density of each behavior in the population was at an equilibrium. 2) An investigation of whether individual foragers were using Evolutionarily Stable Strategies in the correct environmental conditions. The results showed the density of different behaviors in the population could be at an equilibrium at plausible values for handling time and fight duration. Individual foragers used aggressive kleptoparasitic strategies effectively in the correct environmental conditions but some individuals in those same conditions failed to defend food items. This was attributed to the population being composed of 3 species that differed in competitive ability. These competitive differences influenced the strategies that individuals were able to use. Rather than gulls making poor behavioral decisions these results suggest a more complex 3-species model is required to describe the behavior of this population

The stochastic modelling of kleptoparasitism using a Markov process

Kleptoparasitism, the stealing of food items from other animals, is a common behaviour observed across a huge variety of species, and has been subjected to significant modelling effort. Most such modelling has been deterministic, effectively assuming an infinite population, although recently some important stochastic models have been developed. In particular the model of Yates and Broom (Stochastic models of kleptoparasitism. J. Theor. Biol. 248 (2007), 480–489) introduced a stochastic version following the original model of Ruxton and Moody (The ideal free distribution with kleptoparasitism. J. Theor. Biol. 186 (1997), 449–458), and whilst they generated results of interest, they did not solve the model explicitly. In this paper, building on methods used already by van der Meer and Smallegange (A stochastic version of the Beddington-DeAngelis functional response: Modelling interference for a finite number of predators. J. Animal Ecol. 78 (2009) 134–142) we give an exact solution to the distribution of the population over the states for the Yates and Broom model and investigate the effects of some key biological parameters, especially for small populations where stochastic models can be expected to differ most from their deterministic equivalents

Kleptoparasitic melees--modelling food stealing featuring contests with multiple individuals

Kleptoparasitism is the stealing of food by one animal from another. This has been modelled in various ways before, but all previous models have only allowed contests between two individuals. We investigate a model of kleptoparasitism where individuals are allowed to fight in groups of more than two, as often occurs in real populations. We find the equilibrium distribution of the population amongst various behavioural states, conditional upon the strategies played and environmental parameters, and then find evolutionarily stable challenging strategies. We find that there is always at least one ESS, but sometimes there are two or more, and discuss the circumstances when particular ESSs occur, and when there are likely to be multiple ESSs

A game-theoretic model of kleptoparasitic behavior in polymorphic populations

Kleptoparasitism, the stealing of food by one animal from another, is a widespread biological phenomenon. In this paper we build upon earlier models to investigate a population of conspecifics involved in foraging and, potentially, kleptoparasitism. We assume that the population is composed of four types of individuals, according to their strategic choices when faced with an opportunity to steal and to resist an attack. The fitness of each type of individual depends upon various natural parameters, for example food density, the handling time of a food item and the probability of mounting a successful attack against resistance, as well as the choices that they make. We find the evolutionarily stable strategies (ESSs) for all parameter combinations and show that there are six possible ESSs, four pure and two mixtures of two strategies, that can occur. We show that there is always at least one ESS, and sometimes two or three. We further investigate the influence of the different parameters on when each type of solution occurs

An acute bout of cycling does not induce compensatory responses in pre-menopausal women not using hormonal contraceptives

There is a clear need to improve understanding of the effects of physical activity and exercise on appetite control. Therefore, the acute and short-term effects (three days) of a single bout of cycling on energy intake and energy expenditure were examined in women not using hormonal contraceptives. Sixteen active (n = 8) and inactive (n = 8) healthy pre-menopausal women completed a randomised crossover design study with two conditions (exercise and control). The exercise day involved cycling for 1 h (50% of maximum oxygen uptake) and resting for 2 h, whilst the control day comprised 3 h of rest. On each experimental day participants arrived at the laboratory fasted, consumed a standardised breakfast and an ad libitum pasta lunch. Food diaries and combined heart rate-accelerometer monitors were used to assess free-living food intake and energy expenditure, respectively, over the subsequent three days. There were no main effects or condition (exercise vs control) by group (active vs inactive) interaction for absolute energy intake (P > 0.05) at the ad libitum laboratory lunch meal, but there was a condition effect for relative energy intake (P = 0.004, ηp2 = 0.46) that was lower in the exercise condition (1417 ± 926 kJ vs. 2120 ± 923 kJ). Furthermore, post-breakfast satiety was higher in the active than in the inactive group (P = 0.005, ηp2 = 0.44). There were no main effects or interactions (P > 0.05) for mean daily energy intake, but both active and inactive groups consumed less energy from protein (14 ± 3% vs. 16 ± 4%, P = 0.016, ηp2 = 0.37) and more from carbohydrate (53 ± 5% vs. 49 ± 7%, P = 0.031, ηp2 = 0.31) following the exercise condition. This study suggests that an acute bout of cycling does not induce compensatory responses in active and inactive women not using hormonal contraceptives, while the stronger satiety response to the standardised breakfast meal in active individuals adds to the growing literature that physical activity helps improve the sensitivity of short-term appetite control

A game-theoretical model of kleptoparasitic behavior in an urban gull (Laridae) population

Kleptoparasitism (food stealing) is a significant behavior for animals that forage in social groups as it permits some individuals to obtain resources while avoiding the costs of searching for their own food. Evolutionary game theory has been used to model kleptoparasitism, with a series of differential equation-based compartmental models providing significant theoretical insights into behavior in kleptoparasitic populations. In this paper, we apply this compartmental modeling approach to kleptoparasitic behavior in a real foraging population of urban gulls (Laridae). Field data was collected on kleptoparasitism and a model developed that incorporated the same kleptoparasitic and defensive strategies available to the study population. Two analyses were conducted: 1) An assessment of whether the density of each behavior in the population was at an equilibrium. 2) An investigation of whether individual foragers were using Evolutionarily Stable Strategies in the correct environmental conditions. The results showed the density of different behaviors in the population could be at an equilibrium at plausible values for handling time and fight duration. Individual foragers used aggressive kleptoparasitic strategies effectively in the correct environmental conditions but some individuals in those same conditions failed to defend food items. This was attributed to the population being composed of 3 species that differed in competitive ability. These competitive differences influenced the strategies that individuals were able to use. Rather than gulls making poor behavioral decisions these results suggest a more complex 3-species model is required to describe the behavior of this population

Evolutionarily stable sexual allocation by both stressed and unstressed potentially simultaneous hermaphrodites within the same population.

Factors influencing allocation of resources to male and female offspring continue to be of great interest to evolutionary biologists. A simultaneous hermaphrodite is capable of functioning in both male and female mode at the same time, and such a life-history strategy is adopted by most flowering plants and by many sessile aquatic animals. In this paper, we focus on hermaphrodites that nourish post-zygotic stages, e.g. flowering plants and internally fertilising invertebrates, and consider how their sex allocation should respond to an environmental stress that reduces prospects of survival but does not affect all individuals equally, rather acting only on a subset of the population. Whereas dissemination of pollen and sperm can begin at sexual maturation, release of seeds and larvae is delayed by embryonic development. We find that the evolutionarily stable strategy for allocation between male and female functions will be critically dependent on the effect of stress on the trade-off between the costs of male and female reproduction, (i.e. of sperm and embryos). Thus, we identify evaluation of this factor as an important challenge to empiricists interested in the effects of stress on sex allocation. When only a small fraction of the population is stressed, we predict that stressed individuals will allocate their resources entirely to male function and unstressed individuals will increase their allocation to female function. Conversely, when the fraction of stress-affected individuals is high, stressed individuals should respond to this stressor by increasing investment in sperm and unstressed individuals should invest solely in embryos. A further prediction of the model is that we would not expect to find populations in the natural world where both stressed and unstressed individuals are both hermaphrodite

Faculty Productivity and Carnegie Institutional Characteristics within AEJMC Programs

This article reports the results of a content analysis of faculty vitae from eighteen ACEJMC programs drawn using stratified random sampling by Carnegie Classification. The findings indicate that faculty members differ by Carnegie Classification on research productivity, highest earned degrees, professional experience, time assignments (for research, teaching, and service), contact and credit hours, and external grants.Yeshttps://us.sagepub.com/en-us/nam/manuscript-submission-guideline