Group Cooperation without Group Selection: Modest Punishment Can Recruit Much Cooperation

Humans everywhere cooperate in groups to achieve benefits not attainable by individuals. Individual effort is often not automatically tied to a proportionate share of group benefits. This decoupling allows for free-riding, a strategy that (absent countermeasures) outcompetes cooperation. Empirically and formally, punishment potentially solves the evolutionary puzzle of group cooperation. Nevertheless, standard analyses appear to show that punishment alone is insufficient, because second-order free riders (those who cooperate but do not punish) can be shown to outcompete punishers. Consequently, many have concluded that other processes, such as cultural or genetic group selection, are required. Here, we present a series of agent-based simulations that show that group cooperation sustained by punishment easily evolves by individual selection when you introduce into standard models more biologically plausible assumptions about the social ecology and psychology of ancestral humans. We relax three unrealistic assumptions of past models. First, past models assume all punishers must punish every act of free riding in their group. We instead allow punishment to be probabilistic, meaning punishers can evolve to only punish some free riders some of the time. This drastically lowers the cost of punishment as group size increases. Second, most models unrealistically do not allow punishment to recruit labor; punishment merely reduces the punished agent’s fitness. We instead realistically allow punished free riders to cooperate in the future to avoid punishment. Third, past models usually restrict agents to interact in a single group their entire lives. We instead introduce realistic social ecologies in which agents participate in multiple, partially overlapping groups. Because of this, punitive tendencies are more expressed and therefore more exposed to natural selection. These three moves toward greater model realism reveal that punishment and cooperation easily evolve by direct selection—even in sizeable groups.     

The cost of cowardice: punitive sentiments towards free riders in Turkana raids

Models indicate that large-scale cooperation can be sustained by indirect reciprocity or direct punishment, but the relative importance of these mechanisms is unresolved. Unlike direct punishment, indirect sanctions can be meted out without cost to the sanctioner, but direct punishment is advantageous when the scale of cooperation exceeds the network size of individuals. It is of great interest to assess the importance of these mechanisms in small-scale acephalous groups in which people have lived for most of our evolutionary history. Here we evaluate sentiments towards free riders in combat among the Turkana, an acephalous nomadic pastoral society in East Africa who periodically mobilize for cattle-raids against neighboring ethnic groups. Using vignette studies, we probed participants’ motivation to sanction fictitious warriors who were cowards or deserters in a raid and compared it respectively to their reactions to an unskilled warrior or a warrior who turns back due to illness. Our results indicate that the Turkana are motivated to impose both indirect and direct sanctions on cowards consistent with indirect reciprocity and punishment models of cooperation. Our findings imply that both these mechanisms have shaped human cooperative psychology, and sheds light on how prestate societies solve the collective action problem in warfare.     

Reputational benefits for altruistic punishment

Many studies show that people act cooperatively and are willing to punish free riders (i.e., people who are less cooperative than others). However, nonpunishers benefit when free riders are punished, making punishment a group-beneficial act. Presented here are four studies investigating whether punishers gain social benefits from punishing. Undergraduate participants played public goods games (PGGs) (cooperative group games involving money) in which there were free riders, and in which they were given the opportunity to impose monetary penalties on free riders. Participants rated punishers as being more trustworthy, group focused, and worthy of respect than nonpunishers. In dyadic trust games following PGGs, punishers did not receive monetary benefits from punishing free riders in a single-round PGG, but did benefit monetarily from punishing free riders in iterated PGGs. Punishment that was not directed at free riders brought no monetary benefits, suggesting that people distinguish between justified and unjustified punishment and only respond to punishment with enhanced trust when the punishment is justified.     

Do descriptive social norms drive peer punishment? Conditional punishment strategies and their impact on cooperation

Peer punishment is widely considered a key mechanism supporting cooperation in human groups. Although much research shows that human behavior is shaped by the prevailing social norms, little is known about how punishment decisions are impacted by the social context. We present a set of large-scale incentivized experiments in which participants (999 American participants recruited via Amazon Mechanical Turk) could punish their partner conditional on either the level of cooperation or the level of punishment displayed by others who previously interacted in the same setting. While many participants punish independently of levels of cooperation or punishment, a substantial portion punishes free riding more severely when cooperation is more common (‘norm enforcement’), or when free riding is more severely punished by others (‘conformist punishment’). With a dynamic model we demonstrate that conditional punishment strategies can substantially promote cooperation. In particular, conformist punishment helps cooperation to gain a foothold in a population, and norm enforcement helps to maintain cooperation at high levels. Our results provide solid empirical evidence of conditional punishment strategies and illustrate their possible implications for the dynamics of human cooperation.     

Love or fear: Can punishment promote cooperation?

Cooperation is a paradox: Why should one perform a costly behavior only to increase the fitness of another? Human societies, in which individuals cooperate with genetically unrelated individuals on a considerably larger scale than most mammals do, are especially puzzling in this regard. Recently, the threat of punishment has been given substantial attention as one of the mechanisms that could help sustain human cooperation in such situations. Nevertheless, using punishment to explain cooperation only leads to further questions: Why spend precious resources to penalize free‐riders, especially if others can avoid this investment and cheaters can punish you back? Here, it is argued that current evidence supports punishment as an efficient means for the maintenance of cooperation, and that the gravity of proposed limitations of punishment for maintaining cooperation may have been overestimated in previous studies due to the features of experimental design. Most notably, the importance of factors as characteristic of human societies as reputation and language has been greatly neglected. Ironically, it was largely the combination of the two that enabled humans to shape costly punishment into numerous low‐cost and less detrimental strategies that clearly can promote human cooperation.     

A study of physical demands in riding

Thirteen experienced riders and three elite riders underwent bicycle ergometer tests at submaximal and maximal workloads. Oxygen uptake, pulmonary ventilation and heart rate were also studied during riding at a walk, a trot and a canter. The mean maximal oxygen uptake of the experienced riders in the ergometer test (2.71 . min-1) was superior to the average maximal oxygen uptake of other groups of the same age and sex. The average oxygen uptake of the experienced riders in trot sitting was 1.701 . min-1, trot rising 1.681 . min-1 and in canter 1.801 . min-1. The experienced riders used at least 60% of their maximal aerobic power in trot and canter, which is an exercise intensity that may induce some training effect. Two elite riders consistently had lower oxygen uptakes in riding than the other riders. The heart rate -- oxygen uptake relationships in riding and in the ergometer tests were similar, except during trot sitting when the heart rate tended to be higher, indicating a larger share of static muscle contraction in this gait. Static muscle strength was measured in nine riders and seven non-riders. Six muscle groups were investigated, but no significant difference in muscle strength could be demonstrated between riders and controls.     

The evolution of cooperation by negotiation in a noisy world

Cooperative interactions among individuals are ubiquitous despite the possibility of exploitation by selfish free riders. One mechanism that may promote cooperation is ‘negotiation’: individuals altering their behaviour in response to the behaviour of others. Negotiating individuals decide their actions through a recursive process of reciprocal observation, thereby reducing the possibility of free riding. Evolutionary games with response rules have shown that infinitely many forms of the rule can be evolutionarily stable simultaneously, unless there is variation in individual quality. This potentially restricts the conditions under which negotiation could maintain cooperation. Organisms interact with one another in a noisy world in which cooperative effort and the assessment of effort may be subject to error. Here, we show that such noise can make the number of evolutionarily stable rules finite, even without quality variation, and so noise could help maintain cooperative behaviour. We show that the curvature of the benefit function is the key factor determining whether individuals invest more or less as their partner's investment increases, investing less when the benefit to investment has diminishing returns. If the benefits of low investment are very small then behavioural flexibility tends to promote cooperation, because negotiation enables cooperators to reach large benefits. Under some conditions, this leads to a repeating cycle in which cooperative behaviour rises and falls over time, which may explain between‐population differences in cooperative behaviour. In other conditions, negotiation leads to extremely high levels of cooperative behaviour, suggesting that behavioural flexibility could facilitate the evolution of eusociality in the absence of high relatedness.     

How domesticating fire facilitated the evolution of human cooperation

Controlled fire use by early humans could have facilitated the evolution of human cooperation. Individuals with regular access to the benefits of domestic fire would have been at an advantage over those with limited or no access. However, a campfire would have been relatively costly for an individual to maintain and open to free riders. By cooperating, individuals could have reduced maintenance costs, minimized free riding and lessened the risk of being without fire. Cooperators were more likely to survive and reproduce than uncooperative individuals because the former would have been better able to maximize a fire’s returns and enjoy regular access to its benefits. This is how the emergence of controlled fire use in Pleistocene human populations could have facilitated the evolution of cooperation.     

Assessing the rider's seat and horse's behavior: difficulties and perspectives

A correct seat and position are the basis for a good performance in horseback riding. This study aimed to measure deviations from the correct seat, test a seat improvement program (dismounted exercises), and investigate whether horse behavior was affected by the rider's seat. Five experienced trainers defined 16 seat deviations and scored the occurrence in 20 riders in a dressage test. Half the riders then carried out an individual training program; after 9 weeks, riders were again scored. The study took no video or heart-rate recordings of horses and riders. Panel members did not agree on the deviations in the rider's seat; the study detected no differences-with the exception of improvement of backward-tilted pelvis-between the groups. Horse behavior, classified as "evasive," increased; horse heart rate decreased in the experimental group. Heart rates of riders in both groups decreased. Seven of 9 riders in the experimental group had the impression that the exercises improved their riding performance. There is a clear need to develop a robust system that allows trainers to objectively evaluate the rider's seat.     

The evolution of punishment through reputation

Punishment of non-cooperators has been observed to promote cooperation. Such punishment is an evolutionary puzzle because it is costly to the punisher while beneficial to others, for example, through increased social cohesion. Recent studies have concluded that punishing strategies usually pay less than some non-punishing strategies. These findings suggest that punishment could not have directly evolved to promote cooperation. However, while it is well established that reputation plays a key role in human cooperation, the simple threat from a reputation of being a punisher may not have been sufficiently explored yet in order to explain the evolution of costly punishment. Here, we first show analytically that punishment can lead to long-term benefits if it influences one''s reputation and thereby makes the punisher more likely to receive help in future interactions. Then, in computer simulations, we incorporate up to 40 more complex strategies that use different kinds of reputations (e.g. from generous actions), or strategies that not only include punitive behaviours directed towards defectors but also towards cooperators for example. Our findings demonstrate that punishment can directly evolve through a simple reputation system. We conclude that reputation is crucial for the evolution of punishment by making a punisher more likely to receive help in future interactions, and that experiments investigating the beneficial effects of punishment in humans should include reputation as an explicit feature.     

Evolutionary emergence and maintenance of horizontally transmitted mutualism that do not rely on the supply of standing variation in symbiont quality

Mutualism based on reciprocal exchange of costly services must avoid exploitation by ‘free‐rides’. Accordingly, hosts discriminate against free‐riding symbionts in many mutualistic relationships. However, as the selective advantage of discriminators comes from the presence of variability in symbiont quality that they eliminate, discrimination and thus mutualism have been considered to be maintained with exogenous supply of free‐riders. In this study, we tried to resolve the ‘paradoxical’ co‐evolution of discrimination by hosts and cooperation by symbionts, by comparing two different types of discrimination: ‘one‐shot’ discrimination, where a host does not reacquire new symbionts after evicting free‐riders, and ‘resampling’ discrimination, where a host does from the environment. Our study shows that this apparently minor difference in discrimination types leads to qualitatively different evolutionary outcomes. First, although it has been usually considered that the benefit of discriminators is derived from the variability of symbiont quality, the benefit of a certain type of discriminators (e.g. one‐shot discrimination) is proportional to the frequency of free‐riders, which is in stark contrast to the case of resampling discrimination. As a result, one‐shot discriminators can invade the free‐rider/nondiscriminator population, even if standing variation for symbiont quality is absent. Second, our one‐shot discriminators can also be maintained without exogenous supply of free‐riders and hence is free from the paradox of discrimination. Therefore, our result indicates that the paradox is not a common feature of evolution of discrimination but is a problem of specific types of discrimination.     

Energy expenditure during bicycling

This study was designed to measure the O2 uptake (VO2) of cyclists while they rode outdoors at speeds from 32 to 40 km/h. Regression analyses of data from 92 trials using the same wheels, tires, and tire pressure with the cyclists riding in their preferred gear and in an aerodynamic position indicated the best equation (r = 0.84) to estimate VO2 in liters per minute VO2 = -4.50 + 0.17 rider speed + 0.052 wind speed + 0.022 rider weight where rider and wind speed are expressed in kilometers per hour and rider weight in kilograms. Following another rider closely, i.e., drafting, at 32 km/h reduced VO2 by 18 +/- 11%; the benefit of drafting a single rider at 37 and 40 km/h was greater (27 +/- 8%) than that at 32 km/h. Drafting one, two, or four riders in a line at 40 km/h resulted in the same reduction in VO2 (27 +/- 7%). Riding at 40 km/h at the back of a group of eight riders reduced VO2 by significantly more (39 +/- 6%) than drafting one, two, or four riders in a line; drafting a vehicle at 40 km/h resulted in the greatest decrease in VO2 (62 +/- 6%). VO2 was also 7 +/- 4% lower when the cyclists were riding an aerodynamic bicycle. An aerodynamic set of wheels with a reduced number of spokes and one set of disk wheels were the only wheels to reduce VO2 significantly while the cyclists were riding a conventional racing bicycle at 40 km/h.(ABSTRACT TRUNCATED AT 250 WORDS)     

Park or Ride? Evolution of Infant Carrying in Primates

Few mammalian orders carry their infants clinging to the mother's fur. I investigated the evolution of carrying behavior in primates and the life-history and ecological correlates of infant care patterns. Primates are ideal for the study as there is variation in infant care patterns. Primate infants are left hidden in nests or parked in trees, both of which strategies I term parking, and are carried orally or ride clinging to the mother's fur: riding. Infant carrying has evolved several times in the Primates and, once evolved, it has been conserved. Significant energetic costs of riding are indicated as riding species maintain smaller home ranges than those of non-riders of the same body size. With body size and phylogenetic influences taken into account, riders appear to incur a reproductive cost by weaning and breeding later than parkers. Although riders do not have lower birth rates than those of parkers, their later age at first reproduction leads to their having a lower reproductive rate, measured by the intrinsic rate of population increase. Precociality of infants is not correlated with either riding or nesting behavior. Although non-nesting species have larger litter sizes, their infants are not significantly smaller, nor are their neonatal brains relatively smaller. Although riding may have some energetic and reproductive costs, its repeated evolution in the Primates suggests that it also has some benefits, the most likely being a reduced mortality risk for carried infants.     

Explaining the evolution of parochial punishment in humans

Humans usually favour members of their own group, ethnicity or culture (parochial cooperation), and punish out-group wrongdoers more harshly (parochial punishment). The evolution of parochial cooperation is mainly explained by intergroup conflict, as restricting cooperation to in-groups can provide a relative advantage during conflict. However, explanations for the evolution of parochial punishment are still lacking. It is unclear whether conflict can also explain parochial punishment, because conflict is expected to lead to full hostility towards out-groups, irrespective of their behaviour. Here, we use an agent-based simulation to explore which conditions favour the evolution of parochial third-party punishment. We show that when groups interact and then engage in conflict with each other, third-party punishment is not parochial but spiteful, and directed towards all out-groups. A parochial bias in punishment decisions evolves (i) without conflict, when groups compete against nature and enforcing cooperation requires many punitive actions, and (ii) with conflict, when groups come into conflict with a group other than one they previously interacted with. Our findings suggest that intergroup conflict does not always lead to parochial punishment, and that stable collaborative relations between groups is a key factor promoting third-party parochial punishment. Our findings also provide novel predictions on how punishment and intergroup conflict influence in-group bias in human societies.     

Does horse temperament influence horse-rider cooperation?

Cooperation between rider and horse is of major importance in equitation. A balanced team of horse and rider improves (sport) performances and welfare aspects by decreasing stress, frustration, risks of injuries, and accidents. Important features affecting the cooperation are the physical skills, knowledge, and personality of the rider on one hand and the temperament, experience, and physical abilities of the horse on the other. A study with 16 riders and 16 warm-blood riding horses tested the effect of personality of riders and temperament of horses on cooperation between riders and horses. More emotionally reactive horses showed more evasive behavior during riding. Riders preferred to ride those horses who were assessed by the riders as being attentive to the rider's aid. The frequency of evasive behaviors during riding--as assessed by riders, in contrast to the assessments made by an external judge--influenced the cooperation between rider and horse. On average, a rider's personality did not affect the cooperation between rider and horse; however, it is suggested that a rider's personality does affect the cooperation with more emotionally reactive horses.     

Punishing free riders: direct and indirect promotion of cooperation

Human cooperation in a large group of genetically unrelated people is an evolutionary puzzle. Despite its costly nature, cooperative behavior is commonly found in all human societies—a fact that has interested researchers from a wide range of disciplines, including biology, economics, and psychology, to name a few. Many behavioral experiments have demonstrated that cooperation within a group can be sustained when free riders are punished. We argue that punishment has both a direct effect and an indirect effect on promoting cooperation. The direct effect of punishment alters the consequences of cooperation and defection in such a way as to make a rational person prefer cooperation. The indirect effect of punishment promotes cooperation among conditional cooperators by providing the condition necessary for their cooperation (i.e., the expectation that other members will also cooperate). Here we present data from two one-shot n-person prisoner's dilemma games, demonstrating that the indirect effect of punishment complements the direct effect to increase cooperation in the game. Furthermore, we show that direct and indirect effects are robust across two forms of punishment technology: either when punishment is voluntarily provided by game players themselves or when it is exogenously provided by the experimenter.     

The Impact of Therapeutic Riding for People with Disabilities on Variables Related to Mental Health

Substantial anecdotal evidence and some research indicate that therapeutic riding for people with disabilities has a positive impact on mental health. There is similar evidence showing that equine programs that include a psychotherapeutic component also have a positive impact. However, much of the existing research has many methodological problems, the most common being the absence of a comparison or control group. Therefore the present study investigated the mental health benefits of therapeutic riding for people with disabilities in comparison to therapeutic skiing for people with disabilities. The age range of participants was 16 to 70 years. They had a wide range of physical disabilities. In a quasi-experimental pre-test post-test design, all participants completed The Beck Depression Inventory, The Beck Anxiety Inventory, and The Rosenberg Self-Esteem Scale. They also completed an adjective checklist, consisting of 26 items that focused on mood, emotion, and sense of self. Open-ended interviews were conducted with a subset of riders. In addition, the riders completed the Pet Attitude Scale (PAS-M). Scores on the PAS-M were not related to the three psychometric measures. Both groups improved on the three psychometric measures during the course of their respective programs. Although the improvement was statistically significant, it was not clinically significant. Contrary to the hypothesis, there was no difference in improvement between riders and skiers on these measures. However, on the adjective checklist riders felt more positive regarding 21 of the 26 items, and of those 21, eight were statistically significant. The interview data suggest that the riding program improves motivation, relationship building, and self-esteem. Although these results must be interpreted with caution, due to the exploratory nature of the checklist and the interviews, they suggest that equine activities have a positive impact for people with disabilities and may have a greater impact than other activities for some aspects of mental health. However, the positive impact may be related to general wellbeing rather than psychopathology per se.     

Rare but severe concerted punishment that favors cooperation

As one of the mechanisms that are supposed to explain the evolution of cooperation among unrelated individuals, costly punishment, in which altruistic individuals privately bear the cost to punish defection, suffers from such drawbacks as decreasing individuals’ welfare, inducing second-order free riding, the difficulty of catching defection, and the possibility of triggering retaliation. To improve this promising mechanism, here we propose an extended Public Goods game with rare but severe concerted punishment, in which once a defector is caught punishment is triggered and the cost of punishment is equally shared among the remainder of the group. Analytical results show that, when the probability for concerted punishment is above a threshold, cooperating is, while defecting is not, an evolutionarily stable strategy in finite populations, and that this way of punishment can considerably decrease the total cost of inhibiting defection, especially in large populations.     

Punishment of Minor Female Genital Ritual Procedures: Is the Perfect the Enemy of the Good?

Female genital alteration (FGA) is any cutting, removal or destruction of any part of the external female genitalia. Various FGA practices are common throughout the world. While most frequent in Africa and Asia, transglobal migration has brought ritual FGA to Western nations. All forms of FGA are generally considered undesirable for medical and ethical reasons when performed on minors. One ritual FGA procedure is the vulvar nick (VN). This is a small laceration to the vulva that does not cause morphological changes. Besides being performed as a primary ritual procedure it has been proposed as a substitute for more extensive forms of FGA. Measures advocated or taken to reduce the burden of FGA can be punitive or non‐punitive. Even if it is unethical to perform VN, we argue that it also is unethical to attempt to suppress it through punishment. First, punishment of VN is likely to cause more harm than good overall, even to those ostensibly being protected. Second, punishment is likely to exceed legitimate retributive ends. We do not argue in favor of performing VN. Rather, we argue that non‐punitive strategies such as education and harm reduction should be employed.     

Self-serving punishment of a common enemy creates a public good in reef fishes

A key challenge for evolutionary biologists is to determine conditions under which individuals benefit from a contribution to public goods [1, 2]. For humans, it has been observed that punishment of free riders may promote contributions [3,?4], but the conditions that lead to stable cooperation based on punishment remain hotly debated [5-8]. Here we present empirical evidence that public goods may emerge as a by-product of self-serving punishment in interactions between coral reef fishes and parasitic saber-tooth blennies that stealthily attack their fish victims from behind to take a bite [9]. We first show that chasing the blenny functions as punishment [10], because it decreases the probability of future attacks. We then provide evidence that in female scalefin anthias, a shoaling species, punishment creates a public good because it increases the probability that the parasite switches to another species for the next attack. A final experiment suggests that punishment is nevertheless self-serving because blennies appear to be able to discriminate between look-alike punishers and nonpunishers. Thus, individuals that do contribute to the public good may risk being identified by the parasite as easy targets for future attacks.