Playing the political game: the coevolution of institutions with group size and political inequality.

All societies need to form institutional rules to regulate their social interactions. These specify what actions individuals should take in particular situations, and what sanctions will apply if individuals violate these rules. However, forming these institutional rules involves playing a political game-a process of negotiation between individuals that is costly and time-consuming. Intuitively, this cost should be expected to increase as a group becomes larger, which could then select for a transition to hierarchy to keep the cost of playing the political game down as group size increases. However, previous work has lacked a mechanistic yet general model of political games that could formalize this argument and test the conditions under which it holds. We address this by formalizing the political game using a standard consensus formation model. We show that the increasing cost of forming a consensus over institutional rules selects for a transition from egalitarian to hierarchical organization over a wide range of conditions. Playing a political game to form institutional rules in this way captures and unites a previously disparate set of voluntary theories for hierarchy formation, and can explain why the increasing group size in the Neolithic would lead to strong political inequality. This article is part of the theme issue 'Evolutionary ecology of inequality'.

Evolved Open-Endedness in Cultural Evolution: A New Dimension in Open-Ended Evolution Research.

The goal of Artificial Life research, as articulated by Chris Langton, is "to contribute to theoretical biology by locating life-as-we-know-it within the larger picture of life-as-it-could-be." The study and pursuit of open-ended evolution in artificial evolutionary systems exemplify this goal. However, open-ended evolution research is hampered by two fundamental issues: the struggle to replicate open-endedness in an artificial evolutionary system and our assumption that we only have one system (genetic evolution) from which to draw inspiration. We argue not only that cultural evolution should be seen as another real-world example of an open-ended evolutionary system but that the unique qualities seen in cultural evolution provide us with a new perspective from which we can assess the fundamental properties of, and ask new questions about, open-ended evolutionary systems, especially with regard to evolved open-endedness and transitions from bounded to unbounded evolution. Here we provide an overview of culture as an evolutionary system, highlight the interesting case of human cultural evolution as an open-ended evolutionary system, and contextualize cultural evolution by developing a new framework of (evolved) open-ended evolution. We go on to provide a set of new questions that can be asked once we consider cultural evolution within the framework of open-ended evolution and introduce new insights that we may be able to gain about evolved open-endedness as a result of asking these questions.

An investigation of the role of leadership in consensus decision-making.

Leadership is a widespread phenomena in social organisms and it is recognised to facilitate coordination between individuals. While the role of leadership in group foraging or swarm movement is well understood, it is not clear if leaders would also benefit more complex forms of coordination. In particular, a number of organisms coordinate by consensus decision-making, where individuals explicitly communicate their opinions until they converge toward a common decision. Taking inspiration from physical sciences, we extend a consensus formation model to integrate leaders, which we define by three traits: persuasiveness, talkativeness, and stubbornness. We use numerical simulations to investigate the effect of the number of leaders and their characteristics on the time a group spends to reach consensus, and the bias in the final decision. We show that having a minority of influential individuals (leaders) and a majority of influenceable individuals (followers) reduces the time to reach consensus, but biases the decision towards the preferences of the leaders. This effect emerges solely from the differences in individuals' personality traits, with the most determinant trait being the talkativeness of the individuals. Overall, we provide a comprehensive investigation of the effects of leaders and their traits on consensus decision-making.

Modelling transitions between egalitarian, dynamic leader and absolutist power structures.

Human groups show a variety of leadership dynamics ranging from egalitarian groups with no leader, to groups with changing leaders, to absolutist groups with a single long-term leader. Here, we model transitions between these different phases of leadership dynamics, investigating the role of inequalities in relationships between individuals. Our results demonstrate a novel riches-to-rags class of leadership dynamics where a leader can be replaced by a new individual. We note that the transition between the three different phases of leadership dynamics resembles transitions in leadership dynamics during the Neolithic period of human history. We argue how technological developments, such as food storage and/or weapons which allow one individual to control large quantities of resources, would mean that relationships became more unequal. In general terms, we provide a model of how individual relationships can affect leadership dynamics and structures.

Four levers of reciprocity across human societies: concepts, analysis and predictions.

This paper surveys five human societal types - mobile foragers, horticulturalists, pre-state agriculturalists, state-based agriculturalists and liberal democracies - from the perspective of three core social problems faced by interacting individuals: coordination problems, social dilemmas and contest problems. We characterise the occurrence of these problems in the different societal types and enquire into the main force keeping societies together given the prevalence of these. To address this, we consider the social problems in light of the theory of repeated games, and delineate the role of intertemporal incentives in sustaining cooperative behaviour through the reciprocity principle. We analyse the population, economic and political structural features of the five societal types, and show that intertemporal incentives have been adapted to the changes in scope and scale of the core social problems as societies have grown in size. In all societies, reciprocity mechanisms appear to solve the social problems by enabling lifetime direct benefits to individuals for cooperation. Our analysis leads us to predict that as societies increase in complexity, they need more of the following four features to enable the scalability and adaptability of the reciprocity principle: nested grouping, decentralised enforcement and local information, centralised enforcement and coercive power, and formal rules.

Cooperation in large-scale human societies-What, if anything, makes it unique, and how did it evolve?

To resolve the major controversy about why prosocial behaviors persist in large-scale human societies, we propose that two questions need to be answered. First, how do social interactions in small-scale and large-scale societies differ? By reviewing the exchange and collective-action dilemmas in both small-scale and large-scale societies, we show they are not different. Second, are individual decision-making mechanisms driven by self-interest? We extract from the literature three types of individual decision-making mechanism, which differ in their social influence and sensitivity to self-interest, to conclude that humans interacting with non-relatives are largely driven by self-interest. We then ask: what was the key mechanism that allowed prosocial behaviors to continue as societies grew? We show the key role played by new social interaction mechanisms-change in the rules of exchange and collective-action dilemmas-devised by the interacting individuals, which allow for self-interested individuals to remain prosocial as societies grow.

From disorganized equality to efficient hierarchy: how group size drives the evolution of hierarchy in human societies.

A manifest trend is that larger and more productive human groups shift from distributed to centralized decision-making. Voluntary theories propose that human groups shift to hierarchy to limit scalar stress, i.e. the increase in cost of organization as a group grows. Yet, this hypothesis lacks a mechanistic model to investigate the organizational advantage of hierarchy and its role on its evolution. To fill this gap, we describe social organization by the distribution of individuals' capacity to influence others. We then integrate this formalization into models of social dynamics and evolutionary dynamics. First, our results demonstrate that hierarchy strongly reduces scalar stress, and that this benefit can emerge solely because leaders and followers differ in their capacity to influence others. Second, the model demonstrates that this benefit can be sufficient to drive the evolution of leader and follower behaviours and ultimately, the transition from small egalitarian to large hierarchical groups.

The Institutional Approach for Modeling the Evolution of Human Societies.

Artificial life is concerned with understanding the dynamics of human societies. A defining feature of any society is its institutions. However, defining exactly what an institution is has proven difficult, with authors often talking past each other. This article presents a dynamic model of institutions, which views them as political game forms that generate the rules of a group's economic interactions. Unlike most prior work, the framework presented here allows for the construction of explicit models of the evolution of institutional rules. It takes account of the fact that group members are likely to try to create rules that benefit themselves. Following from this, it allows us to determine the conditions under which self-interested individuals will create institutional rules that support cooperation-for example, that prevent a tragedy of the commons. The article finishes with an example of how a model of the evolution of institutional rewards and punishments for promoting cooperation can be created. It is intended that this framework will allow artificial life researchers to examine how human groups can themselves create conditions for cooperation. This will help provide a better understanding of historical human social evolution, and facilitate the resolution of pressing societal social dilemmas.

When is bigger better? The effects of group size on the evolution of helping behaviours.

Understanding the evolution of sociality in humans and other species requires understanding how selection on social behaviour varies with group size. However, the effects of group size are frequently obscured in the theoretical literature, which often makes assumptions that are at odds with empirical findings. In particular, mechanisms are suggested as supporting large-scale cooperation when they would in fact rapidly become ineffective with increasing group size. Here we review the literature on the evolution of helping behaviours (cooperation and altruism), and frame it using a simple synthetic model that allows us to delineate how the three main components of the selection pressure on helping must vary with increasing group size. The first component is the marginal benefit of helping to group members, which determines both direct fitness benefits to the actor and indirect fitness benefits to recipients. While this is often assumed to be independent of group size, marginal benefits are in practice likely to be maximal at intermediate group sizes for many types of collective action problems, and will eventually become very small in large groups due to the law of decreasing marginal returns. The second component is the response of social partners on the past play of an actor, which underlies conditional behaviour under repeated social interactions. We argue that under realistic conditions on the transmission of information in a population, this response on past play decreases rapidly with increasing group size so that reciprocity alone (whether direct, indirect, or generalised) cannot sustain cooperation in very large groups. The final component is the relatedness between actor and recipient, which, according to the rules of inheritance, again decreases rapidly with increasing group size. These results explain why helping behaviours in very large social groups are limited to cases where the number of reproducing individuals is small, as in social insects, or where there are social institutions that can promote (possibly through sanctioning) large-scale cooperation, as in human societies. Finally, we discuss how individually devised institutions can foster the transition from small-scale to large-scale cooperative groups in human evolution.

Evolutionary Connectionism: Algorithmic Principles Underlying the Evolution of Biological Organisation in Evo-Devo, Evo-Eco and Evolutionary Transitions.

The mechanisms of variation, selection and inheritance, on which evolution by natural selection depends, are not fixed over evolutionary time. Current evolutionary biology is increasingly focussed on understanding how the evolution of developmental organisations modifies the distribution of phenotypic variation, the evolution of ecological relationships modifies the selective environment, and the evolution of reproductive relationships modifies the heritability of the evolutionary unit. The major transitions in evolution, in particular, involve radical changes in developmental, ecological and reproductive organisations that instantiate variation, selection and inheritance at a higher level of biological organisation. However, current evolutionary theory is poorly equipped to describe how these organisations change over evolutionary time and especially how that results in adaptive complexes at successive scales of organisation (the key problem is that evolution is self-referential, i.e. the products of evolution change the parameters of the evolutionary process). Here we first reinterpret the central open questions in these domains from a perspective that emphasises the common underlying themes. We then synthesise the findings from a developing body of work that is building a new theoretical approach to these questions by converting well-understood theory and results from models of cognitive learning. Specifically, connectionist models of memory and learning demonstrate how simple incremental mechanisms, adjusting the relationships between individually-simple components, can produce organisations that exhibit complex system-level behaviours and improve the adaptive capabilities of the system. We use the term "evolutionary connectionism" to recognise that, by functionally equivalent processes, natural selection acting on the relationships within and between evolutionary entities can result in organisations that produce complex system-level behaviours in evolutionary systems and modify the adaptive capabilities of natural selection over time. We review the evidence supporting the functional equivalences between the domains of learning and of evolution, and discuss the potential for this to resolve conceptual problems in our understanding of the evolution of developmental, ecological and reproductive organisations and, in particular, the major evolutionary transitions.

How institutions shaped the last major evolutionary transition to large-scale human societies.

What drove the transition from small-scale human societies centred on kinship and personal exchange, to large-scale societies comprising cooperation and division of labour among untold numbers of unrelated individuals? We propose that the unique human capacity to negotiate institutional rules that coordinate social actions was a key driver of this transition. By creating institutions, humans have been able to move from the default 'Hobbesian' rules of the 'game of life', determined by physical/environmental constraints, into self-created rules of social organization where cooperation can be individually advantageous even in large groups of unrelated individuals. Examples include rules of food sharing in hunter-gatherers, rules for the usage of irrigation systems in agriculturalists, property rights and systems for sharing reputation between mediaeval traders. Successful institutions create rules of interaction that are self-enforcing, providing direct benefits both to individuals that follow them, and to individuals that sanction rule breakers. Forming institutions requires shared intentionality, language and other cognitive abilities largely absent in other primates. We explain how cooperative breeding likely selected for these abilities early in the Homo lineage. This allowed anatomically modern humans to create institutions that transformed the self-reliance of our primate ancestors into the division of labour of large-scale human social organization.

What can ecosystems learn? Expanding evolutionary ecology with learning theory.

BACKGROUND: The structure and organisation of ecological interactions within an ecosystem is modified by the evolution and coevolution of the individual species it contains. Understanding how historical conditions have shaped this architecture is vital for understanding system responses to change at scales from the microbial upwards. However, in the absence of a group selection process, the collective behaviours and ecosystem functions exhibited by the whole community cannot be organised or adapted in a Darwinian sense. A long-standing open question thus persists: Are there alternative organising principles that enable us to understand and predict how the coevolution of the component species creates and maintains complex collective behaviours exhibited by the ecosystem as a whole? RESULTS: Here we answer this question by incorporating principles from connectionist learning, a previously unrelated discipline already using well-developed theories on how emergent behaviours arise in simple networks. Specifically, we show conditions where natural selection on ecological interactions is functionally equivalent to a simple type of connectionist learning, 'unsupervised learning', well-known in neural-network models of cognitive systems to produce many non-trivial collective behaviours. Accordingly, we find that a community can self-organise in a well-defined and non-trivial sense without selection at the community level; its organisation can be conditioned by past experience in the same sense as connectionist learning models habituate to stimuli. This conditioning drives the community to form a distributed ecological memory of multiple past states, causing the community to: a) converge to these states from any random initial composition; b) accurately restore historical compositions from small fragments; c) recover a state composition following disturbance; and d) to correctly classify ambiguous initial compositions according to their similarity to learned compositions. We examine how the formation of alternative stable states alters the community's response to changing environmental forcing, and we identify conditions under which the ecosystem exhibits hysteresis with potential for catastrophic regime shifts. CONCLUSIONS: This work highlights the potential of connectionist theory to expand our understanding of evo-eco dynamics and collective ecological behaviours. Within this framework we find that, despite not being a Darwinian unit, ecological communities can behave like connectionist learning systems, creating internal conditions that habituate to past environmental conditions and actively recalling those conditions.

An evolutionary model explaining the Neolithic transition from egalitarianism to leadership and despotism.

The Neolithic was marked by a transition from small and relatively egalitarian groups to much larger groups with increased stratification. But, the dynamics of this remain poorly understood. It is hard to see how despotism can arise without coercion, yet coercion could not easily have occurred in an egalitarian setting. Using a quantitative model of evolution in a patch-structured population, we demonstrate that the interaction between demographic and ecological factors can overcome this conundrum. We model the coevolution of individual preferences for hierarchy alongside the degree of despotism of leaders, and the dispersal preferences of followers. We show that voluntary leadership without coercion can evolve in small groups, when leaders help to solve coordination problems related to resource production. An example is coordinating construction of an irrigation system. Our model predicts that the transition to larger despotic groups will then occur when: (i) surplus resources lead to demographic expansion of groups, removing the viability of an acephalous niche in the same area and so locking individuals into hierarchy; (ii) high dispersal costs limit followers' ability to escape a despot. Empirical evidence suggests that these conditions were probably met, for the first time, during the subsistence intensification of the Neolithic.

The co-evolution of social institutions, demography, and large-scale human cooperation.

Human cooperation is typically coordinated by institutions, which determine the outcome structure of the social interactions individuals engage in. Explaining the Neolithic transition from small- to large-scale societies involves understanding how these institutions co-evolve with demography. We study this using a demographically explicit model of institution formation in a patch-structured population. Each patch supports both social and asocial niches. Social individuals create an institution, at a cost to themselves, by negotiating how much of the costly public good provided by cooperators is invested into sanctioning defectors. The remainder of their public good is invested in technology that increases carrying capacity, such as irrigation systems. We show that social individuals can invade a population of asocials, and form institutions that support high levels of cooperation. We then demonstrate conditions where the co-evolution of cooperation, institutions, and demographic carrying capacity creates a transition from small- to large-scale social groups.

Punishment can promote defection in group-structured populations.

Pro-social punishment, whereby cooperators punish defectors, is often suggested as a mechanism that maintains cooperation in large human groups. Importantly, models that support this idea have to date only allowed defectors to be the target of punishment. However, recent empirical work has demonstrated the existence of anti-social punishment in public goods games. That is, individuals that defect have been found to also punish cooperators. Some recent theoretical studies have found that such anti-social punishment can prevent the evolution of pro-social punishment and cooperation. However, the evolution of anti-social punishment in group-structured populations has not been formally addressed. Previous work has informally argued that group-structure must favour pro-social punishment. Here we formally investigate how two demographic factors, group size and dispersal frequency, affect selection pressures on pro- and anti-social punishment. Contrary to the suggestions of previous work, we find that anti-social punishment can prevent the evolution of pro-social punishment and cooperation under a range of group structures. Given that anti-social punishment has now been found in all studied extant human cultures, the claims of previous models showing the co-evolution of pro-social punishment and cooperation in group-structured populations should be re-evaluated.

The concurrent evolution of cooperation and the population structures that support it.

The evolution of cooperation often depends upon population structure, yet nearly all models of cooperation implicitly assume that this structure remains static. This is a simplifying assumption, because most organisms possess genetic traits that affect their population structure to some degree. These traits, such as a group size preference, affect the relatedness of interacting individuals and hence the opportunity for kin or group selection. We argue that models that do not explicitly consider their evolution cannot provide a satisfactory account of the origin of cooperation, because they cannot explain how the prerequisite population structures arise. Here, we consider the concurrent evolution of genetic traits that affect population structure, with those that affect social behavior. We show that not only does population structure drive social evolution, as in previous models, but that the opportunity for cooperation can in turn drive the creation of population structures that support it. This occurs through the generation of linkage disequilibrium between socio-behavioral and population-structuring traits, such that direct kin selection on social behavior creates indirect selection pressure on population structure. We illustrate our argument with a model of the concurrent evolution of group size preference and social behavior.