The smell of cooperation: rats increase helpful behaviour when receiving odour cues of a conspecific performing a cooperative task

Reciprocity can explain cooperative behaviour among non-kin, where individuals help others depending on their experience in previous interactions. Norway rats (Rattus norvegicus) cooperate reciprocally according to direct and generalized reciprocity. In a sequence of four consecutive experiments, we show that odour cues from a cooperating conspecific are sufficient to induce the altruistic help of rats in a food-exchange task. When rats were enabled to help a non-cooperative partner while receiving olfactory information from a rat helping a conspecific in a different room, they helped their non-cooperative partner as if it was a cooperative one. We further show that the cues inducing altruistic behaviour are released during the act of cooperation and do not depend on the identity of the cue provider. Remarkably, olfactory cues seem to be more important for cooperation decisions than experiencing a cooperative act per se. This suggests that rats may signal their cooperation propensity to social partners, which increases their chances to receive help in return.     

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.     

A friend in need is a friend indeed: Need-based sharing,rather than cooperative assortment,predicts experimental resource transfers among Agta hunter-gatherers

Despite much theorizing, the evolutionary reasons why humans cooperate extensively with unrelated individuals are still largely unknown. While reciprocity explains many instances of non-kin cooperation, much remains to be understood. A recent suite of models based upon ‘cooperative assortativity’ suggest that non-kin cooperation can evolve if individuals preferentially assort with certain cooperative phenotypes, such as helping those who help others. Here, we test these assortative hypotheses among the Agta, a population of Filipino hunter-gatherers, using an experimental resource allocation game in which individuals divide resources between themselves and camp-mates. Individuals preferentially shared with less cooperative individuals, arguing against cooperative assortativity as a mechanism sustaining resource transfers in this population. Rather, sharing was often based on the recipient's level of need, in addition to kin-based transfers and reciprocal sharing. Contrary to several recent theoretical accounts, in this real-world setting we find no evidence for cooperative assortativity influencing patterns of cooperation. These results may reflect the demands of living in a foraging ecology characterized by high resource stochasticity, necessitating need-based sharing as a system of long-term reciprocity to mitigate repeated subsistence shortfalls.     

Density of founder cells affects spatial pattern formation and cooperation in Bacillus subtilis biofilms

In nature, most bacteria live in surface-attached sedentary communities known as biofilms. Biofilms are often studied with respect to bacterial interactions. Many cells inhabiting biofilms are assumed to express ‘cooperative traits'', like the secretion of extracellular polysaccharides (EPS). These traits can enhance biofilm-related properties, such as stress resilience or colony expansion, while being costly to the cells that express them. In well-mixed populations cooperation is difficult to achieve, because non-cooperative individuals can reap the benefits of cooperation without having to pay the costs. The physical process of biofilm growth can, however, result in the spatial segregation of cooperative from non-cooperative individuals. This segregation can prevent non-cooperative cells from exploiting cooperative neighbors. Here we examine the interaction between spatial pattern formation and cooperation in Bacillus subtilis biofilms. We show, experimentally and by mathematical modeling, that the density of cells at the onset of biofilm growth affects pattern formation during biofilm growth. At low initial cell densities, co-cultured strains strongly segregate in space, whereas spatial segregation does not occur at high initial cell densities. As a consequence, EPS-producing cells have a competitive advantage over non-cooperative mutants when biofilms are initiated at a low density of founder cells, whereas EPS-deficient cells have an advantage at high cell densities. These results underline the importance of spatial pattern formation for competition among bacterial strains and the evolution of microbial cooperation.     

感潮河网区环境合作博弈模型及实证

区域决策者往往要选择合适的排污策略,使自己的经济和环境收益最大。当多个决策者进行策略选择时,就形成了区域环境博弈格局。针对感潮河网区排污行为与环境质量具有互相影响的特点,考虑了税收收益、治理成本和环境损失等因素,建立了河网区环境非合作博弈模型和合作博弈模型。非合作博弈的Nash均衡表明,在非合作局面下,区域决策者仅仅考虑最大化自己的收益,选择的排污策略与其他参与者无关。在合作博弈分析中,对大联盟、子联盟博弈进行了分析求解。通过计算所有可能联盟博弈的特征函数,使用常用的Shapley值法进行合作收益公平分配。研究表明,在大联盟合作局面下,区域决策者的策略选择考虑了全局收益的最大化,选择的排污策略与所有参与者的环境损失参数、环境影响参数有关。结合感潮河网区案例对模型进行了验证,研究了3个区域非合作博弈状态和合作博弈状态下的排放量和收益,并使用Shapley值法对合作收益进行分配。对比非合作博弈与合作博弈,合作后3个区域排污量分别比合作前减少了17.98%、15.36%、5.55%。合作收益分别增加了2.17%、3.21%、1.25%。环境质量分别提高了14.24%、13.33%、10.52%。这说明合作局面有利于降低污染排放,分配后的环境合作收益大于非合作收益,河网区环境合作是多赢的。     

Influence of helping and breeding experience on reproductive performance in the Seychelles warbler: a translocation experiment

Reproductive success of the cooperative breeding Seychelleswarbler (Acrocephalus sechellensis) increases with age. Thisage effect is not due to differential survival or increasedreproductive effort, but to accumulated helping and breedingexperience. In their first year of breeding, reproductive performanceof inexperienced warblers with neither helping nor breedingexperience was significandy lower than that of warblers of thesame age with either previous helping or breeding experience.Reproductive performance was the same for primiparae with helpingexperience and for birds with breeding experience. Female primiparaewith helping experience or breeding experience built betternests and spent more time incubating than inexperienced females,which led to increased hatching success. Male primiparae withhelping experience or males with breeding experience guardedthe clutch better than inexperienced males, which led to reducedegg predation. Even-aged warblers with different previous experienceswere transferred to unoccupied islands, where birds startedbreeding immediately in high-quality territories. The experimentshowed that birds with helping experience produced their firstfledgling as fast as experienced breeders, and significandyfaster than inexperienced birds. Breeding performance did notimprove further with experience after the first successful breedingattempt. Only birds with previous breeding experience who pairedwith inexperienced birds, were likely to change mate. The otherpair combinations remained stable. Thus, primiparous birds withhelping experience have greater lifetime reproductive successthan inexperienced primiparae of the same age. This experimentshows that helping behavior has not only been selected for inthe context of promoting an individual's indirect fitness, butalso in the context of gaining helping experience which translatesinto improved reproductive success when a helper becomes a breeder.[Behav Ecol 7: 326-333 (1996)]     

Does Intuition Cause Cooperation?

Recently, researchers claimed that people are intuitively inclined to cooperate with reflection causing them to behave selfishly. Empirical support for this claim came from experiments using a 4-player public goods game with a marginal return of 0.5 showing that people contributed more money to a common project when they had to decide quickly (i.e., a decision based on intuition) than when they were instructed to reflect and decide slowly. This intuitive-cooperation effect is of high scientific and practical importance because it argues against a central assumption of traditional economic and evolutionary models. The first experiment of present study was set up to examine the generality of the intuitive-cooperation effect and to further validate the experimental task producing the effect. In Experiment 1, we investigated Amazon Mechanical Turk (AMT) workers'' contributions to a 4-player public goods game with a marginal return of 0.5 while we manipulated the knowledge about the other players'' contribution to the public goods game (contribution known vs. contribution unknown), the identity of the other players (humans vs. computers randomly generating contributions) and the time constraint (time pressure/intuition vs. forced delay/reflection). However, the results of Experiment 1 failed to reveal an intuitive-cooperation effect. Furthermore, four subsequent direct replications attempts with AMT workers (Experiments 2a, 2b, 2c and Experiment 3, which was conducted with naïve/inexperienced participants) also failed to demonstrate intuitive-cooperation effects. Taken together, the results of the present study could not corroborate the idea that people are intuitively cooperative, hence suggesting that the theoretical relationship between intuition and cooperation should be further scrutinized.     

The structure of a bottlenose dolphin society is coupled to a unique foraging cooperation with artisanal fishermen

Diverse and localized foraging behaviours have been reported in isolated populations of many animal species around the world. In Laguna, southern Brazil, a subset of resident bottlenose dolphins (Tursiops truncatus) uses a foraging tactic involving cooperative interactions with local, beach-casting fishermen. We used individual photo-identification data to assess whether cooperative and non-cooperative dolphins were socially segregated. The social structure of the population was found to be a fission-fusion system with few non-random associations, typical for this species. However, association values were greater among cooperative dolphins than among non-cooperative dolphins or between dolphins from different foraging classes. Furthermore, the dolphin social network was divided into three modules, clustering individuals that shared or lacked the cooperative foraging tactic. Space-use patterns were not sufficient to explain this partitioning, indicating a behavioural factor. The segregation of dolphins using different foraging tactics could result from foraging behaviour driving social structure, while the closer association between dolphins engaged in the cooperation could facilitate the transmission and learning of this behavioural trait from conspecifics. This unique case of a dolphin-human interaction represents a valuable opportunity to explore hypotheses on the role of social learning in wild cetaceans.     

Cooperative and non-cooperative processes of apparent movement of random-dot cinematograms

In this study, we investigated the cooperative and non-cooperative models of stereopsis on apparent movement of the short-range process using spatial frequency filtered random-dot cinematograms. Our results showed that when spatial frequencies were below 4 cycles/degree, maximum displacement (dmax) was decreasing (linearly) with increasing mean frequencies, but at 4 cycles/degree and above dmax stayed constant. For low frequencies, non-cooperative models such as Marr and Poggio's could explain these findings, but not for frequencies above 4 cycles/degree. However, in a previous study we found that the average cooperative neighbourhood for apparent movement of the short-range process is 15 arc min. This fortuitous agreement on 4 cycles/degree could suggest that dmax being constant at frequencies above 4 cycles is related to a cooperative process.     

Cooperation in defence against a predator

The origin and the evolutionary stability of cooperation between unrelated individuals is one of the key problems of evolutionary biology. In this paper, a cooperative defence game against a predator is introduced which is based on Hamilton's selfish herd theory and Eshel's survival game models. Cooperation is altruistic in the sense that the individual, which is not the target of the predator, helps the members of the group attacked by the predator and during defensive action the helper individual may also die in any attack. In order to decrease the long term predation risk, this individual has to carry out a high risk action. Here I show that this kind of cooperative behaviour can evolve in small groups. The reason for the emergence of cooperation is that if the predator does not kill a mate of a cooperative individual, then the survival probability of the cooperative individual will increase in two cases. If the mate is non-cooperative, then—according to the dilution effect, the predator confusion effect and the higher predator vigilance—the survival probability of the cooperative individual increases. The second case is when the mate is cooperative, because a cooperative individual has a further gain, the active help in defence during further predator attacks. Thus, if an individual can increase the survival rate of its mates (no matter whether the mate is cooperative or not), then its own predation risk will decrease.     

Acculturation and end-of-life decision making: comparison of Japanese and Japanese-American focus groups

Variation in decision-making about end-of-life care among ethnic groups creates clinical conflicts. In order to understand changes in preferences for end-of-life care among Japanese who immigrate to the United States, we conducted 18 focus groups with 122 participants: 65 English-speaking Japanese Americans, 29 Japanese-speaking Japanese Americans and 28 Japanese living in Japan. Negative feelings toward living in adverse health states and receiving life-sustaining treatment in such states permeated all three groups. Fear of being meiwaku, a physical, psychological or financial caregiving burden on loved ones, was a prominent concern. They preferred to die pokkuri (popping off) before they become end stage or physically frail. All groups preferred group-oriented decision-making with family. Although advance directives were generally accepted, Japanese participants saw written directives as intrusive whereas Japanese Americans viewed them mainly as tools to reduce conflict created by dying person's wishes and a family's kazoku no jo--responsibility to sustain the dying patient. These findings suggest that in the United States Japanese cultural values concerning end-of-life care and decision-making process are largely preserved.     

Time Pressure Increases Cooperation in Competitively Framed Social Dilemmas

What makes people willing to pay costs to benefit others? Does such cooperation require effortful self-control, or do automatic, intuitive processes favor cooperation? Time pressure has been shown to increase cooperative behavior in Public Goods Games, implying a predisposition towards cooperation. Consistent with the hypothesis that this predisposition results from the fact that cooperation is typically advantageous outside the lab, it has further been shown that the time pressure effect is undermined by prior experience playing lab games (where selfishness is the more advantageous strategy). Furthermore, a recent study found that time pressure increases cooperation even in a game framed as a competition, suggesting that the time pressure effect is not the result of social norm compliance. Here, we successfully replicate these findings, again observing a positive effect of time pressure on cooperation in a competitively framed game, but not when using the standard cooperative framing. These results suggest that participants'' intuitions favor cooperation rather than norm compliance, and also that simply changing the framing of the Public Goods Game is enough to make it appear novel to participants and thus to restore the time pressure effect.     

Asymmetric interaction will facilitate the evolution of cooperation

Explaining the evolution of cooperation remains one of the greatest problems for both biology and social science. The classical theories of cooperation suggest that cooperation equilibrium or evolutionary stable strategy between partners can be maintained through genetic similarity or reciprocity relatedness. These classical theories are based on an assumption that partners interact symmetrically with equal payoffs in a game of cooperation interaction. However, the payoff between partners is usually not equal and therefore they often interact asymmetrically in real cooperative systems. With the Hawk-Dove model, we find that the probability of cooperation between cooperative partners will depend closely on the payoff ratio. The higher the payoff ratio between recipients and cooperative actors, the greater will be the probability of cooperation interaction between involved partners. The greatest probability of conflict between cooperative partners will occur when the payoff between partners is equal. The results show that this asymmetric relationship is one of the key dynamics of the evolution of cooperation, and that pure cooperation strategy (i.e., Nash equilibrium) does not exist in asymmetrical cooperation systems, which well explains the direct conflict observed in almost all of the well documented cooperation systems. The model developed here shows that the cost-to-benefit ratio of cooperation is also negatively correlated with the probability of cooperation interaction. A smaller cost-to-benefit ratio of cooperation might be created by the limited dispersal ability or exit cost of the partners involved, and it will make the punishment of the non-cooperative individuals by the recipient more credible, and therefore make it more possible to maintain stable cooperation interaction.     

Social Transmission of Avoidance Behavior under Situational Change in Learned and Unlearned Rats

Background

Rats receive information from other conspecifics by observation or other types of social interaction. Such social interaction may contribute to the effective adaptation to changes of environment such as situational switching. Learning to avoid dangerous places or objects rapidly occurs with even a single conditioning session, and the conditioned memory tends to be sustained over long periods. The avoidance is important for adaptation, but the details of the conditions under which the social transmission of avoidance is formed are unknown. We demonstrate that the previous experience of avoidance learning is important for the formation of behaviors for social transmission of avoidance and that the experienced rats adapt to a change of situation determined by the presence or absence of aversive stimuli. We systematically investigated social influence on avoidance behavior using a passive avoidance test in a light/dark two-compartment apparatus.

Methodology/Principal Findings

Rats were divided into two groups, one receiving foot shocks and another with no aversive experience in a dark compartment. Experienced and inexperienced rats were further divided into subjects and partners. In Experiment 1, each subject experienced (1) interaction with an experienced partner, (2) interaction with an inexperienced partner, or (3) no interaction. In Experiment 2, each subject experienced interaction with a partner that received a shock. The entering latency to a light compartment was measured. The avoidance behavior of experienced rats was inhibited by interaction with inexperienced or experienced partners in a safely-changed situation. The avoidance of experienced rats was reinstated in a dangerously-changed situation by interaction with shocked rats. In contrast, the inexperienced rats were not affected by any social circumstances.

Conclusions/Significance

These results suggest that transmitted information among rats can be updated under a situational change and that the previous experience is crucial for social enhancement and inhibition of avoidance behavior in rats.     

Iterated prisoner's dilemma in an asocial world dominated by loners, not by defectors

Cooperation among genetically unrelated individuals can arise when pairs of individuals interact repeatedly in the Prisoner’s Dilemma. However, the conditions allowing the evolution of reciprocal cooperation become extremely restrictive as the size of the cooperative group increases, because defectors can exploit cooperators more efficiently in larger groups. Here we consider three strategies: Tit for Tat, defector, and loner. Loner beats defector in a non-cooperative world. However, a cooperative strategy Tit for Tat (TFT₀) that stops cooperation after the first iteration when there is at least one defector in the group, can invade a world of loners, even in sizable groups, if both the TFT₀ and the defector strategies arise at the same frequency by mutation.     

Decision-Making for Risky Gains and Losses among College Students with Internet Gaming Disorder

Individuals with Internet gaming disorder (IGD) tend to exhibit disadvantageous risky decision-making not only in their real life but also in laboratory tasks. Decision-making is a complex multifaceted function and different cognitive processes are involved in decision-making for gains and losses. However, the relationship between impaired decision-making and gain versus loss processing in the context of IGD is poorly understood. The main aim of the present study was to separately evaluate decision-making for risky gains and losses among college students with IGD using the Cups task. Additionally, we further examined the effects of outcome magnitude and probability level on decision-making related to risky gains and losses respectively. Sixty college students with IGD and 42 matched healthy controls (HCs) participated. Results indicated that IGD subjects exhibited generally greater risk taking tendencies than HCs. In comparison to HCs, IGD subjects made more disadvantageous risky choices in the loss domain (but not in the gain domain). Follow-up analyses indicated that the impairment was associated to insensitivity to changes in outcome magnitude and probability level for risky losses among IGD subjects. In addition, higher Internet addiction severity scores were associated with percentage of disadvantageous risky options in the loss domain. These findings emphasize the effect of insensitivity to losses on disadvantageous decisions under risk in the context of IGD, which has implications for future intervention studies.     

Facial expressions as honest signals of cooperative intent in a one-shot anonymous Prisoner's Dilemma game

Previous research has posited that facial expressions of emotion may serve as honest signals of cooperation. Although findings from several empirical studies support this position, prior studies have not used comprehensive and dynamic measures of facial expression as potential predictors of behaviorally defined cooperation. The authors investigated (a) specific positive and negative facial actions displayed among strangers immediately following verbal promises of commitment within an unrestricted acquaintance period and (b) anonymous, behaviorally defined decisions of cooperation or defection in a one-shot, two-person Prisoner's Dilemma game occurring directly following the acquaintance period. The Facial Action Coding System [Ekman P. & Friesen W.V. (1978). Facial Action Coding System. Palo Alto, CA: Consulting Psychology Press] was used to measure affect-related facial actions. It was found that facial actions related to enjoyment were predictive of cooperative decisions within dyads; additionally, facial actions related to contempt were predictive of noncooperative decisions within dyads. Furthermore, and consistent with previous works, participants were able to accurately predict their partner's decisions following the acquaintance period. These results suggest that facial actions may function as honest signals of cooperative intent. These findings also provide a possible explanation for the association between subjective affective experience and facial expression that advances understanding of cooperative behavior.     

Impact of Dental Atmosphere and Behaviour of the Dentist on Children’s Cooperation

During dental treatment children are usually under psychological pressure. With the Sarnat Behaviour Score five different types of patients can be distinguished. There is no method that measures the impact of dental atmosphere and dentist's behaviour on the young patients' readiness to cooperate. The objective of the present study was the implementation and evaluation of a questionnaire on this subject. Eighty-eight patients participated in this study. In the first part of the new questionnaire personal information was collected. The second part consists of 43 items and investigates the relationship between dentist and patient and reflects the atmosphere of the environment. Statistical analysis was performed using the Chi-square test. There were statistically significant differences between the cooperative and non-cooperative group, as regards the perceived honesty of the dentist, the ability to explain and wish to help. Uncooperative children are significantly more often afraid of the dental environment. Sympathy alone has only a minor effect on children's cooperation. Children should be treated with empathy. Especially younger patients appreciate detailed explanations by the dentist. Children's non-cooperative behaviour results often from their aroused interest in the unknown environment, which causes an unpleasant perception of the whole setting.     

Evolutionary causes and consequences of consistent individual variation in cooperative behaviour

Behaviour is typically regarded as among the most flexible of animal phenotypic traits. In particular, expression of cooperative behaviour is often assumed to be conditional upon the behaviours of others. This flexibility is a key component of many hypothesized mechanisms favouring the evolution of cooperative behaviour. However, evidence shows that cooperative behaviours are often less flexible than expected and that, in many species, individuals show consistent differences in the amount and type of cooperative and non-cooperative behaviours displayed. This phenomenon is known as ‘animal personality’ or a ‘behavioural syndrome’. Animal personality is evolutionarily relevant, as it typically shows heritable variation and can entail fitness consequences, and hence, is subject to evolutionary change. Here, we review the empirical evidence for individual variation in cooperative behaviour across taxa, we examine the evolutionary processes that have been invoked to explain the existence of individual variation in cooperative behaviour and we discuss the consequences of consistent individual differences on the evolutionary stability of cooperation. We highlight that consistent individual variation in cooperativeness can both stabilize or disrupt cooperation in populations. We conclude that recognizing the existence of consistent individual differences in cooperativeness is essential for an understanding of the evolution and prevalence of cooperation.     

Sex roles, parental experience and reproductive success of eastern kingbirds, Tyrannus tyrannus

We quantified parental behaviour of eastern kingbirds during the incubation and nestling periods to determine parental roles, and to examine the impact of previous breeding experience (defined as having bred on the territory in the past) on behaviour and reproductive success. Females performed all incubation, while males spent more than 60% of their time in vigilant or nest guarding behaviour during incubation. Parental roles were not defined as sharply during the nestling period. Females spent more time vigilant, but males provisioned young at only 54% of the rate of females. Vigilance and nest watching were still primarily male duties. Male and female behaviour did not vary with the pair's combination of experience (e.g. experienced-experienced versus inexperienced-inexperienced in previous-current breeding season, respectively) during either phase of reproduction, but experienced males were more vigilant during incubation and fed young relatively more than inexperienced males. Experienced females were also more efficient foragers. Although behaviour did not differ among the four combinations of pair experience, inexperienced pairs none the less lost the most young to starvation and predation. Consequently, inexperienced pairs fledged one less nestling per nesting attempt than did pairs with at least one experienced breeder. Our results suggest that having at least one experienced breeder substantially improved a pair's reproductive success. We propose that female site fidelity is a safeguard to avoid the lower breeding success a female would incur if she were to move to a new territory and breed with an inexperienced male. Copyright 1999 The Association for the Study of Animal Behaviour.