Domestic pigs alter their social strategy in response to social group size

In a series of studies, we investigated the concept that domesticated pigs may adopt cost efficient social strategies, in which individuals become less aggressive due to a higher number of potential competitors, in large social groups. Six blocks, each comprising four pens of 18 pigs (small group, SG) and two pens of 108 pigs (large group, LG) were used. Pigs were 11 weeks old at the initial group formation. Weeks 1, 6 and 12 following SG and LG formation, two randomly selected pigs with SG or LG social experience were incorporated into another SG or LG for a period of 2 h and the aggressive behaviour of pigs was observed. Four test combinations were used (SG to SG (SS), SG to LG (SL), LG to SG (LS) and LG to LG (LL)). There was less aggression in the LL compared to SS, SL and LS combinations (1.6% versus 2.5, 2.3, and 2.5% of time, respectively, P = 0.009). Furthermore, 8 weeks following SG and LG formation, a total of 200 pigs were regrouped for 2 h in groups of four in a neutral test arena to assess the effect of prior social experience (SG vs. LG) on aggressive behaviour. Pigs were regrouped with their own group members (familiar), or with unfamiliar non-group members either from SG or LG. Five test combinations were used (four pigs from the same SG (S), four pigs from the same LG (L), two pigs from a SG and two pigs from a LG (SL), two pigs each from two different LG (LL) and two pigs each from two different SG (SS)). Pigs derived from SG fought aggressively (duration, s) with unfamiliar pigs, compared to those derived from LG (4.9, 6.7, 16.1, 12.1 and 9.5 s for S, L, SS, SL and LL test combinations respectively, P < 0.001). In addition, the latency to initiate first aggressive interaction was shorter (duration, s) in SS than LL treatment combination (23.3, 22.5, 10.8, 11.8 and 19.2 s for S, L, SS, SL and LL test combinations respectively, P = 0.008). Taken together, our results suggest that pigs become less aggressive and may shift to a low-aggressive social strategy in large social groups. This may provide potential benefits for welfare of pigs under commercial production situations.     

Chimpanzees and children avoid mutual defection in a social dilemma

Cooperation often comes with the temptation to defect and benefit at the cost of others. This tension between cooperation and defection is best captured in social dilemmas like the Prisoner's Dilemma. Adult humans have specific strategies to maintain cooperation during Prisoner's Dilemma interactions. Yet, little is known about the ontogenetic and phylogenetic origins of human decision-making strategies in conflict scenarios. To shed light on this question, we compared the strategies used by chimpanzees and 5-year old children to overcome a social dilemma. In our task, waiting for the partner to act first produced the best results for the subject. Alternatively, they could mutually cooperate and divide the rewards. Our findings indicate that the two species differed substantially in their strategies to solve the task. Chimpanzees became more strategic across the study period by waiting longer to act in the social dilemma. Children developed a more efficient strategy of taking turns to reciprocate their rewards. Moreover, children used specific types of communication to coordinate with their partners. These results suggest that while both species behaved strategically to overcome a conflict situation, only children engaged in active cooperation to solve a social dilemma.     

Individual heterogeneity and costly punishment: a volunteer's dilemma

Social control and the enforcement of social norms glue a society together. It has been shown theoretically and empirically that informal punishment of wrongdoers fosters cooperation in human groups. Most of this research has focused on voluntary and uncoordinated punishment carried out by individual group members. However, as punishment is costly, it is an open question as to why humans engage in the punishment of wrongdoers even in one-time-only encounters. While evolved punitive preferences have been advocated as proximate explanations for such behaviour, the strategic nature of the punishment situation has remained underexplored. It has been suggested to conceive of the punishment situation as a volunteer''s dilemma (VOD), where only one individual''s action is necessary and sufficient to punish the wrongdoer. Here, we show experimentally that implementing the punishment situation as a VOD sustains cooperation in an environment where punishers and non-punishers coexist. Moreover, we show that punishment-cost heterogeneity allows individuals to tacitly agree on only the strongest group member carrying out the punishment, thereby increasing the effectiveness and efficiency of social norm enforcement. Our results corroborate that costly peer punishment can be explained without assuming punitive preferences and show that centralized sanctioning institutions can emerge from arbitrary individual differences.     

Habitat patch size and mating system as determinants of social group size in coral-dwelling fishes

It is thought that the size and dispersion of habitat patches can determine the size and composition of animal social groups, however, this has rarely been tested. The relationship between group size, the mating system, and habitat patch size in six species of coral-dwelling gobies was examined. For all species, there was a positive correlation between coral colony size and social group size, however the strength of this relationship varied among species. Paragobiodon xanthosomus exhibited the strongest relationship and a manipulative field experiment confirmed that coral colony size limited group size in this species. For other species including Paragobiodon melanosomus and Eviota bifasciata, either a highly conservative mating system (P. melanosomus), or increased mobility (E. bifasciata) appeared to disrupt the relationship between habitat patch size and group size. There was no consistent relationship between the mating system exhibited and group size among the species investigated. These results demonstrate that habitat patch size, mobility, and mating systems can interact in complex ways to structure group size even among closely related species.     

Dynamic-persistence of cooperation in public good games when group size is dynamic

The evolution of cooperation is possible with a simple model of a population of agents that can move between groups. The agents play public good games within their group. The relative fitness of individuals within the whole population affects their number of offspring. Groups of cooperators evolve but over time are invaded by defectors which eventually results in the group's extinction. However, for small levels of migration and mutation, high levels of cooperation evolve at the population level. Thus, evolution of cooperation based on individual fitness without kin selection, indirect or direct reciprocity is possible. We provide an analysis of the parameters that affect cooperation, and describe the dynamics and distribution of population sizes over time.     

Crystal toxins and the volunteer's dilemma in bacteria

The growth and virulence of the bacteria Bacillus thuringiensis depend on the production of Cry toxins, which are used to perforate the gut of its host. Successful invasion of the host relies on producing a threshold amount of toxin, after which there is no benefit from producing more toxin. Consequently, the production of Cry toxin appears to be a different type of social problem compared with the public goods scenarios that bacteria usually encounter. We show that selection for toxin production is a volunteer's dilemma. We make specific predictions that (a) selection for toxin production depends upon an interplay between the number of bacterial cells that each host ingests and the genetic relatedness between those cells; (b) cheats that do not produce toxin gain an advantage when at low frequencies, and at high bacterial density, allowing them to be maintained in a population alongside toxin‐producing cells. More generally, our results emphasize the diversity of the social games that bacteria play.     

Group size determined by fusion and fission A mathematical modelling with inclusive fitness

We consider a mathematical model for the group size determination by the intra-reactions, self-growth, ostracism and fission within a group, and by the inter-reactions, immigration and fusion between two groups. In some group reactions, a conflict between two groups occurs about the reaction to change the group size. We construct a mathematical model to consider such conflict, taking into account the inclusive fitness of members in each group. In the conflict about the fusion between two groups, our analysis shows that the smaller group wants to fuse, while the larger does not. Also the criterion to resolve the conflict is discussed, and some numerical examples are given, too. It is concluded that, depending on the deviation in the total cost paid for the conflict by counterparts, the group reactions could result in a terminal group size different from that reached only by a sequence of outsider's immigrations into a group.     

Prey size,prey perishability and group foraging in a social spider

Summary Selection might favor group foraging and social feeding when prey are distributed in patches that do not last long enough for a solitary individual to consume more than a small fraction of them (Pulliam and Millikan 1982; Pulliam and Caraco 1984). Here we considered the foraging behavior of a social spider, Anelosimus eximius, in light of this ephemeral resource hypothesis. This species builds large webs in which members cooperate to capture a wide variety of different sizes and types of prey, many of which are very large. The capture success of this species was very high across all prey sizes, presumably due to the fact that they foraged in groups. Group consumption times in natural colonies for all prey larger than five mm were less than the time that dead insects remained on the plastic sheets that we used as artificial webs. Solitary consumption estimates, calculated from the rate at which laboratory individuals extracted insect biomass while feeding, were the same as the residence times of insects on artificial webs in the field for insects between 6 and 15 mm in length and were significantly longer than the persistence of insects on plastic sheets for all larger insects. Large prey, that contribute substantially to colony energy supplies, appeared to be ephemeral resources for these spiders that could not be consumed by a single spider in the time they were available. These factors made the food intake of one spider in a group less sensitive to scavenging by others and could act to reinforce the social system of this species.     

A strategy to increase cooperation in the volunteer's dilemma: reducing vigilance improves alarm calls

One of the most common examples of cooperation in animal societies is giving the alarm in the presence of a predator. A reduction in individual vigilance against predators when group size increases (the "group size effect") is one of the most frequently reported relationships in the study of animal behavior, and is thought to be due to relaxed selection, either because more individuals can detect the predator more easily (the "many eyes" effect) or because the risk of predator attack is diluted on more individuals (the "selfish herd" effect). I show that these hypotheses are not theoretically grounded: because everybody relies on someone else to raise the alarm, the probability that at least one raises the alarm declines with group size; therefore increasing group size does not lead to relaxed selection. Game theory shows, instead, that increasing the risk that the predator is not reported (by reducing vigilance) induces everybody to give the alarm more often. The group size effect, therefore, can be due to strategic behavior to improve the production of a public good. This shows how a selfish behavior can lead to a benefit for the group, and suggests a way to solve social dilemmas in the absence of relatedness and repeated interactions.     

Spatial effects in social dilemmas

Social dilemmas and the evolutionary conundrum of cooperation are traditionally studied through various kinds of game theoretical models such as the prisoner's dilemma, public goods games, snowdrift games or by-product mutualism. All of them exemplify situations which are characterized by different degrees of conflicting interests between the individuals and the community. In groups of interacting individuals, cooperators produce a common good benefitting the entire group at some cost to themselves, whereas defectors attempt to exploit the resource by avoiding the costly contributions. Based on synergistic or discounted accumulation of cooperative benefits a unifying theoretical framework was recently introduced that encompasses all games that have traditionally been studied separately (Hauert, Michor, Nowak, Doebeli, 2005. Synergy and discounting of cooperation in social dilemmas. J. Theor. Biol., in press.). Within this framework we investigate the effects of spatial structure with limited local interactions on the evolutionary fate of cooperators and defectors. The quantitative effects of space turn out to be quite sensitive to the underlying microscopic update mechanisms but, more general, we demonstrate that in prisoner's dilemma type interactions spatial structure benefits cooperation-although the parameter range is quite limited-whereas in snowdrift type interactions spatial structure may be beneficial too, but often turns out to be detrimental to cooperation.     

Evolutionary dynamics of the continuous iterated prisoner's dilemma

The iterated prisoner's dilemma (IPD) has been widely used in the biological and social sciences to model dyadic cooperation. While most of this work has focused on the discrete prisoner's dilemma, in which actors choose between cooperation and defection, there has been some analysis of the continuous IPD, in which actors can choose any level of cooperation from zero to one. Here, we analyse a model of the continuous IPD with a limited strategy set, and show that a generous strategy achieves the maximum possible payoff against its own type. While this strategy is stable in a neighborhood of the equilibrium point, the equilibrium point itself is always vulnerable to invasion by uncooperative strategies, and hence subject to eventual destabilization. The presence of noise or errors has no effect on this result. Instead, generosity is favored because of its role in increasing contributions to the most efficient level, rather than in counteracting the corrosiveness of noise. Computer simulation using a single-locus infinite alleles Gaussian mutation model suggest that outcomes ranging from a stable cooperative polymorphism to complete collapse of cooperation are possible depending on the magnitude of the mutational variance. Also, making the cost of helping a convex function of the amount of help provided makes it more difficult for cooperative strategies to invade a non-cooperative equilibrium, and for the cooperative equilibrium to resist destabilization by non-cooperative strategies. Finally, we demonstrate that a much greater degree of assortment is required to destabilize a non-cooperative equilibrium in the continuous IPD than in the discrete IPD. The continuous model outlined here suggests that incremental amounts of cooperation lead to rapid decay of cooperation and thus even a large degree of assortment will not be sufficient to allow cooperation to increase when cooperators are rare. The extreme degree of assortment required to destabilize the non-cooperative equilibrium, as well as the instability of the cooperative equilibrium, may help explain why cooperation in Prisoner's Dilemmas is so rare in nature.     

Cooperation as a volunteer’s dilemma and the strategy of conflict in public goods games

Conflict and cooperation for the exploitation of public goods are usually modelled as an N‐person prisoner’s dilemma. Many social dilemmas, however, would be described more properly as a volunteer’s dilemma, in which a certain number of individuals are necessary to produce a public good. If volunteering is costly, but so is failure to produce the public good, cheaters can invade and form a stable mixed equilibrium with cooperators. The dilemma is that the benefit for the group decreases with group size because the larger the group is, the less likely it is that someone volunteers. This problem persists even in the presence of a high degree of relatedness between group members. This model provides precise, testable predictions for the stability of cooperation. It also suggests a counterintuitive but practical solution for this kind of social dilemmas: increasing the damage resulting from the failure to produce the public good increases the probability that the public good is actually produced. Adopting a strategy that entails a deliberate risk (brinkmanship), therefore, can lead to a benefit for the society without being detrimental for the individual.     

Social cohesion in groups of sheep: Effect of activity level, sex composition and group size

We investigated the effects of activity, group size and sex composition on the cohesion of merino sheep (Ovis aries) groups. Mixed-sex (50% of each sex) and single-sex groups of 2, 4, 6 and 8 sheep were placed within 491-m² arenas located in natural pastures and were video recorded during 6 daily hours. The behaviour, orientation and location of each sheep were then extracted from the films at 1-s intervals. We analysed the polarisation of individual orientations, mean inter-individual and nearest neighbours’ distances, as well as the frequency of pairs of nearest neighbours according to their sex within mixed-sex groups. Sheep were more aggregated than predicted under the null hypothesis of random spatial distribution for all group compositions and sizes. Sheep were more spread out and less aligned in half-active than in all-active groups, showing that social cohesion was reduced by a lack of activity synchronisation. The highest proximity between individuals was found in resting groups, yet alignment was low. The polarisation peaked in all-active groups. Mean inter-individual distance did not vary and the nearest neighbour distance decreased as group size increased. When sheep were all-active or all-resting, mixed-sex groups were more spread out than single-sex ones, with a greater distance between opposite than between same-sex individuals. Nearest neighbours of the same sex were also more frequent than random. Our results show that social cohesion can be modulated by activity synchrony but also by social affinity.     

Group selection, kin selection, altruism and cooperation: When inclusive fitness is right and when it can be wrong

Group selection theory has a history of controversy. After a period of being in disrepute, models of group selection have regained some ground, but not without a renewed debate over their importance as a theoretical tool. In this paper I offer a simple framework for models of the evolution of altruism and cooperation that allows us to see how and to what extent both a classification with and one without group selection terminology are insightful ways of looking at the same models. Apart from this dualistic view, this paper contains a result that states that inclusive fitness correctly predicts the direction of selection for one class of models, represented by linear public goods games. Equally important is that this result has a flip side: there is a more general, but still very realistic class of models, including models with synergies, for which it is not possible to summarize their predictions on the basis of an evaluation of inclusive fitness.     

Effect of temperature and social environment on worker size in the ant Temnothorax nylanderi

Warm temperatures decrease insect developmental time and body size. Social life could buffer external environmental variations, especially in large social groups, either through behavioral regulation and compensation or through specific nest architecture. Mean worker size and distribution of worker sizes within colonies are important parameters affecting colony productivity as worker size is linked to division of labor in insect societies. In this paper, we investigate the effect of stressful warm temperatures and the role of social environment (colony size and size of nestmate workers) on the mean size and size variation of laboratory-born workers in the small European ant Temnothorax nylanderi. To do so, we reared field-collected colonies under medium or warm temperature treatments after having marked the field-born workers and removed the brood except for 30 first instar larvae. Warm temperature resulted in the production of fewer workers and a higher adult mortality, confirming that this regime was stressful for the ants. T. nylanderi ants followed the temperature size rule observed in insects, with a decreased developmental time and mean size under warm condition. Social environment appeared to play an important role as we observed that (i) larger colonies buffered the effect of temperature better than smaller ones (ii) colonies with larger workers produced larger workers whatever the rearing temperature and (iii) the coefficient of variation of worker size was similar in the field and under medium laboratory temperature. This suggests that worker size variation is not primarily due to seasonal environmental fluctuations in the field. Finally, we observed a higher coefficient of variation of worker size under warm temperature. We propose that this results from a disruption of social regulation, i.e. the control of nestmate workers over developing larvae and adult worker size, under stressful conditions.     

Bet hedging based cooperation can limit kin selection and form a basis for mutualism

Mutualism is a mechanism of cooperation in which partners that differ help each other. As such, mutualism opposes mechanisms of kin selection and tag-based selection (for example the green beard mechanism), which are based on giving exclusive help to partners that are related or carry the same tag. In contrast to kin selection, which is a basis for parochialism and intergroup warfare, mutualism can therefore be regarded as a mechanism that drives peaceful coexistence between different groups and individuals. Here the competition between mutualism and kin (tag) selection is studied. In a model where kin selection and tag-based selection are dominant, mutualism is promoted by introducing environmental fluctuations. These fluctuations cause reduction in reproductive success by the mechanism of variance discount. The best strategy to counter variance discount is to share with agents who experience the most anticorrelated fluctuations, a strategy called bet hedging. In this way, bet hedging stimulates cooperation with the most unrelated partners, which is a basis for mutualism. Analytic results and simulations reveal that, if this effect is large enough, mutualistic strategies can dominate kin selective strategies. In addition, mutants of these mutualistic strategies that experience fluctuations that are more anticorrelated to their partner, can outcompete wild type, which can lead to the evolution of specialization. In this way, the evolutionary success of mutualistic strategies can be explained by bet hedging-based cooperation.     

Evolution of cooperation and altruistic punishment when retaliation is possible

Altruistic punishment is suggested to explain observed high levels of cooperation among non-kin related humans. However, laboratory experiments as well as ethnographic evidence suggest that people might retaliate if being punished, and that this reduces the level of cooperation. Building on existing models on the evolution of cooperation and altruistic punishment, we explore the consequences of the option of retaliation. We find that cooperation and altruistic punishment does not evolve with larger population levels if the option of retaliation is included.     

A unimodal curve for primate group size along a gradient of population density

When plotted along a gradient of population density, the mean group size in populations of several primate species has a unimodal distribution, i.e., mean group size is greater at intermediate population densities than at higher or lower population densities. In this study I present a mathematical model to clarify the cause of this relationship. Population density is assumed to affect group size by enhancing between- or within-group competition and by changing the number of neighboring groups around each group. The mean group size is predicted to decline as population density increases above a critical value, owing to the increasing number of neighboring groups.     

Sex-biased dispersal of adults mediates the evolution of altruism among juveniles

Population viscosity has been proposed as an important mechanism for the evolution of cooperation. The idea is that if individuals do not disperse far during the course of their lives, they will tend to interact with their genealogical relatives, which may give kin-selected benefits for cooperation. However, in the simplest model of population structure, the evolution of cooperation is unaffected by the rate of dispersal, owing to dispersal also mediating competition between social partners. This surprising result has generated much research interest in recent years. Here I show that dispersal does matter if there is a sex difference in dispersal rate, even when the expression of cooperation is not conditional upon the actor's dispersal status or sex. In particular, I show that cooperation among juveniles is relatively favoured when there is a small sex bias in adult dispersal in favour of the sex with the greatest variance in reproductive success, and is relatively disfavoured when this sex bias is large or in the opposite direction. This is because dispersal by individuals of each sex can have different consequences for the genetic structure of the population.     

Genetic relatedness and group size in an aggregation economy

Summary We use Hamilton's Rule to investigate effects of genetic relatedness on the predicted size of social groups. We assume an aggregation economy; individual fitness initially increases with group size, but in sufficiently large groups each member's individual fitness declines with further increments in the size of the group. We model two processes of group formation, designated free entry and group-controlled entry. The first model assumes that solitary individuals decide to join groups or remain alone; group size equilibrates when solitaries no longer choose to join. The second model allows group members to regulate the size of the group, so that the predicted group size results from members' decisions to repel or accept intruding solitaries. Both the Nash equilibrium group size and any change in the equilibrium caused by varying the level of relatedness depend on the particular entry rule assumed. The largest equilibrium group size occurs when solitaries choose between joining or not joining and individuals are unrelated. Increasing genetic relatedness may reduce and can never increase, equilibrium group size when this entry rule applies. The smallest equilibrium group size occurs when group members choose between repelling or accepting intruders and individuals are unrelated. Under this entry rule, increasing genetic relatedness can increase and can never decrease, equilibrium group size. We extend the models' predictions to suggest when individuals should prefer kin vs non-kin as members of the same group.