The cultural evolution and ecology of institutions.

Human societies are structured by what we refer to as 'institutions', which are socially created and culturally inherited proscriptions on behaviour that define roles and set expectations about social interactions. The study of institutions in several social science fields has provided many important insights that have not been fully appreciated in the evolutionary human sciences. However, such research has often lacked a shared understanding of general processes of change that shape institutional diversity across space and time. We argue that evolutionary theory can provide a useful framework for synthesizing information from different disciplines to address issues such as how and why institutions change over time, how institutional rules co-evolve with other culturally inherited traits, and the role that ecological factors might play in shaping institutional diversity. We argue that we can gain important insights by applying cultural evolutionary thinking to the study of institutions, but that we also need to expand and adapt our approaches to better handle the ways that institutions work, and how they might change over time. In this paper, we illustrate our approach by describing macro-scale empirical comparative analyses that demonstrate how evolutionary theory can be used to generate and test hypotheses about the processes that have shaped some of the major patterns we see in institutional diversity over time and across the world today. We then go on to discuss how we might usefully develop micro-scale models of institutional change by adapting concepts from game theory and agent-based modelling. We end by considering current challenges and areas for future research, and the potential implications for other areas of study and real-world applications. This article is part of the theme issue 'Foundations of cultural evolution'.

An agent-based model of the female rivalry hypothesis for concealed ovulation in humans.

After half a century of debate and few empirical tests, there remains no consensus concerning why ovulation in human females is considered concealed. The predominant male investment hypothesis states that females were better able to obtain material investment from male partners across those females' ovulatory cycles by concealing ovulation. We build on recent work on female competition to propose and investigate an alternative-the female rivalry hypothesis-that concealed ovulation benefited females by allowing them to avoid aggression from other females. Using an agent-based model of mating behaviour and paternal investment in a human ancestral environment, we did not find strong support for the male investment hypothesis, but found support for the female rivalry hypothesis. Our results suggest that concealed ovulation may have benefitted females in navigating their intrasexual social relationships. More generally, this work implies that explicitly considering female-female interactions may inspire additional insights into female behaviour and physiology.

Agent-based modelling reveals strategies to reduce the fitness and metastatic potential of circulating tumour cell clusters.

Metastasis-the ability of cancer cells to disperse throughout the body and establish new tumours at distant locations-is responsible for most cancer-related deaths. Although both single and clusters of circulating tumour cells (CTCs) have been isolated from cancer patients, CTC clusters are generally associated with higher metastatic potential and worse prognosis. From an evolutionary perspective, being part of a cluster can provide cells with several benefits both in terms of survival (e.g. protection) and reproduction (group dispersal). Thus, strategies aimed at inducing cluster dissociation could decrease the metastatic potential of CTCs. However, finding agents or conditions that induce the dissociation of CTC clusters is hampered by the fact that their detection, isolation and propagation remain challenging. Here, we used a mechanistic agent-based model to (a) investigate the response of CTC clusters of various sizes and densities to different challenges-in terms of cell survival and cluster stability, and (b) make predictions as to the combination of factors and parameter values that could decrease the fitness and metastatic potential of CTC clusters. Our model shows that the resilience and stability of CTC clusters are dependent on both their size and density. Also, CTC clusters of distinct sizes and densities respond differently to changes in resource availability, with high-density clusters being least affected. In terms of responses to microenvironmental threats (such as drugs), increasing their intensity is, generally, least effective on high-density clusters. Lastly, we found that combining various levels of resource availability and threat intensity can be more effective at decreasing the survival of CTC clusters than each factor alone. We suggest that the complex effects that cluster density and size showed on both the resilience and stability of the CTC clusters are likely to have significant consequences for their metastatic potential and responses to therapies.

Symmetry-based reciprocity: evolutionary constraints on a proximate mechanism.

Background. While the evolution of reciprocal cooperation has attracted an enormous attention, the proximate mechanisms underlying the ability of animals to cooperate reciprocally are comparatively neglected. Symmetry-based reciprocity is a hypothetical proximate mechanism that has been suggested to be widespread among cognitively unsophisticated animals. Methods. We developed two agent-based models of symmetry-based reciprocity (one relying on an arbitrary tag and the other on interindividual proximity) and tested their ability both to reproduce significant emergent features of cooperation in group living animals and to promote the evolution of cooperation. Results. Populations formed by agents adopting symmetry-based reciprocity showed differentiated "social relationships" and a positive correlation between cooperation given and received: two common aspects of animal cooperation. However, when reproduction and selection across multiple generations were added to the models, agents adopting symmetry-based reciprocity were outcompeted by selfish agents that never cooperated. Discussion. In order to evolve, hypothetical proximate mechanisms must be able to stand competition from alternative strategies. While the results of our simulations require confirmation using analytical methods, we provisionally suggest symmetry-based reciprocity is to be abandoned as a possible proximate mechanism underlying the ability of animals to reciprocate cooperative interactions.

Exchanging grooming, but not tolerance and aggression in common marmosets (Callithrix jacchus).

In this study, we investigated the reciprocal exchanges of grooming, tolerance and reduced aggression in common marmosets (Callithrix jacchus), a cooperatively breeding primate whose groups are typically characterized by uniformly high genetic relatedness and high interdependency between group members. Both partner control and partner choice processes played a role in the reciprocal exchanges of grooming. In contrast, we did not find any evidence of reciprocity between grooming and tolerance over a preferred food source or between grooming and reduced aggression. Thus, reciprocity seems to play a variable role in the exchange of cooperative behaviors in marmosets.

Partner choice promotes cooperation: the two faces of testing with agent-based models.

Reciprocity is one of the most debated among the mechanisms that have been proposed to explain the evolution of cooperation. While a distinction can be made between two general processes that can underlie reciprocation (within-pair temporal relations between cooperative events, and partner choice based on benefits received), theoretical modelling has concentrated on the former, while the latter has been often neglected. We developed a set of agent-based models in which agents adopted a strategy of obligate cooperation and partner choice based on benefits received. Our models tested the ability of partner choice both to reproduce significant emergent features of cooperation in group living animals and to promote the evolution of cooperation. Populations formed by agents adopting a strategy of obligate cooperation and partner choice based on benefits received showed differentiated "social relationships" and a positive correlation between cooperation given and received, two common phenomena in animal cooperation. When selection across multiple generations was added to the model, agents adopting a strategy of partner choice based on benefits received outperformed selfish agents that did not cooperate. Our results suggest partner choice is a significant aspect of cooperation and provides a possible mechanism for its evolution.

Emergent group level navigation: an agent-based evaluation of movement patterns in a folivorous primate.

The foraging activity of many organisms reveal strategic movement patterns, showing efficient use of spatially distributed resources. The underlying mechanisms behind these movement patterns, such as the use of spatial memory, are topics of considerable debate. To augment existing evidence of spatial memory use in primates, we generated movement patterns from simulated primate agents with simple sensory and behavioral capabilities. We developed agents representing various hypotheses of memory use, and compared the movement patterns of simulated groups to those of an observed group of red colobus monkeys (Procolobus rufomitratus), testing for: the effects of memory type (Euclidian or landmark based), amount of memory retention, and the effects of social rules in making foraging choices at the scale of the group (independent or leader led). Our results indicate that red colobus movement patterns fit best with simulated groups that have landmark based memory and a follow the leader foraging strategy. Comparisons between simulated agents revealed that social rules had the greatest impact on a group's step length, whereas the type of memory had the highest impact on a group's path tortuosity and cohesion. Using simulation studies as experimental trials to test theories of spatial memory use allows the development of insight into the behavioral mechanisms behind animal movement, developing case-specific results, as well as general results informing how changes to perception and behavior influence movement patterns.