Brosnan & De Waal Science 2014 with Summary
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RESEARCH ◥ have a more explicitly social focus (11). Disad- REVIEW vantageous IA is also the most common type in animals (see below). However, responses to over- compensation are also expected, as they, too, SOCIAL EVOLUTION provide a long-term benefit (12). We have called disadvantageous IA “first-order inequity aversion” to indicate that it is the primary reaction, using Evolution of responses to (un)fairness “second-order inequity aversion” for the less com- mon and less pronounced advantageous IA Sarah F. Brosnan1* and Frans B. M. de Waal2 (Fig. 1) (13). The connection between IA and fairness is not The human sense of fairness is an evolutionary puzzle. To study this, we can look to straightforward. The hallmark of the human other species, in which this can be translated empirically into responses to reward sense of fairness is the idea of impartiality; that distribution. Passive and active protest against receiving less than a partner for the same is, human fairness or justice is based on the idea task is widespread in species that cooperate outside kinship and mating bonds. There is of appropriate outcomes applied to everyone less evidence that nonhuman species seek to equalize outcomes to their own detriment, within the community, not just a few individuals, yet the latter has been documented in our closest relatives, the apes. This reaction and, in particular, not just oneself. Thus, out- probably reflects an attempt to forestall partner dissatisfaction with obtained outcomes comes are judged against a standard, or an ideal. and its negative impact on future cooperation. We hypothesize that it is the evolution of There is variation in this ideal across cultures or this response that allowed the development of a complete sense of fairness in humans, situations, but there is consistency within a given which aims not at equality for its own sake but for the sake of continued cooperation. context. This complete sense of fairness likely requires abstraction at the community level as ooperation could not have evolved without been explored in a number of species and found well as language (to establish a consistent set of mechanisms to ensure the sharing of pay- to be most pronounced in animals that cooperate ideals), both of which capacities may be restricted offs. For an individual to cooperate with an outside of the bonds of mating and kinship. to our species (14). Community concern is not unrelated partner to achieve goals that it We propose that sensitivity to (in)equity offers wholly absent in other primates, however, and cannot achieve alone or to exchange favors several evolutionary benefits. First, animals need neither is impartiality, such as when policing Cover time requires an ability to compare payoffs to recognize when they receive less than a part- males break up fights (15). with investments. Given the ample evidence for ner, because this tells them that the benefits of Inasmuch as social ideals escape measure- mutualistic cooperation and reciprocal altruism cooperation may be in danger. By protesting ment, a sense of fairness is impossible to prove or (1, 2)inhumansaswellasotherspecies(here- against this situation, they show a response known disprove in animals. Reactions to inequity, on after, animals), we therefore expect well-developed as inequity aversion (IA).Evidenceindicatesthat the other hand, are open to empirical investiga- capacities for payoff evaluation in species that this behavior is widespread in cooperative spe- tion by creating situations in which one individ- flexibly cooperate with individually known part- cies under many circumstances. As the reliance ual receives more or less than another. These ners. We also expect negative reactions to exces- on cooperation increases, individuals also benefit reactions typically manifest as a rejection of a sive payoff imbalances, because such imbalances from sensitivity to receiving more than another, received reward or an unwillingness to partici- undermine cooperation among nonrelatives, which which risks undermining cooperative partner- pate in the interaction (Fig. 2A). In most experi- requires proportionalitybetweeneffortandgain ships. This behavior is likely taxonomically re- ments, subjects must complete a simple task to so that gains among parties jointly contributing stricted, because it requires prediction of the receive a reward (Table 1). To control for the so- to a given enterprise are shared. partner's reaction to getting less and its effect on cial aspect of the interaction, these experiments Along with the human sense of fairness and the relationship. It also requires restraint to re- often combine with ones on contrast effects that justice, responses to inequity have enjoyed a long frain from an immediately advantageous out- measure how subjects respond to a lesser reward history of scholarship in philosophy, law, eco- come. The pressure for increased cooperation after having just received a better reward (con- nomics, and psychology. Yet the evolution of combined with advanced cognitive abilities and trast) or another lesser one (control). It has been these responses and possible parallels in other emotional control allowed humans to evolve a found that the mere visibility of better rewards is species have only recently come into focus. Even complete sense of fairness. Here, we review the not the issue, because primates reliably perform though “contrast effects,” which describe how literature on IA in humans and other animals tasks for lesser rewards regardless of whether or animals respond to unanticipated individual re- within an evolutionary framework of cooperation, not better ones are immediately in front of them ward outcomes, have been known for nearly a social reciprocity, and conflict resolution. Our main (6, 16). Experiments on IA have shown that there century (3), the first study to measure reactions conclusion is that the sense of fairness did not is substantial variation among species in this to interindividual outcome contrasts was pub- evolve for the sake of fairness per se but in order response, even within the primates; some species lished only in 2003 (4). In this study, brown capu- to reap the benefits of continued cooperation. respond more strongly to contrast effects (17), chin monkeys (Cebus apella)becameagitated others more strongly to disadvantageous inequity and refused to perform a task for which a com- Responses to inequity (4, 16); some respond to both (18), and some seem panion received superior rewards [see (5)fora IA has been defined as a negative reaction to indifferent to either condition (19, 20). video]. The monkeys’ protest was not due to the unequal outcomes (7). It is further subdivided There are also important individual differences mere sight of unavailable superior rewards, be- into “disadvantageous IA,” or reactions to in- in response that hint at the situations in which cause they showed it only if these rewards actual- equity to the detriment of the actor, and “advan- inequity responses provide an advantage. For in- ly went to their partner. If superior rewards were tageous IA,” also known as overcompensation, or stance, merely feeding unequal foods fails to merely visible, they were mostly ignored (4, 6). reactions to inequity that benefits the actor (7). generate the same reaction; hence, an effortful Since this early study, inequity responses have Responses to the first kind of IA offer a clear task is essential (6, 16, 20)(Table1),eventhough advantage if they help increase one’sownshare. the nature of the task may be irrelevant (20). A 1Departments of Psychology and Philosophy, Neuroscience Not surprisingly, human studies indicate that dis- second methodological issue emerges when we Institute and Language Research Center, Georgia State advantageous IA emerges earlier (8)andismore consider all reported studies regardless of spe- 2 University, Atlanta, GA, USA. Living Links, Yerkes National pronounced than advantageous IA (9). Young cies. Animals tested with an effortful task respond Primate Research Center and Psychology Department, Emory University, Atlanta, GA, USA. children may even pay a cost to maintain their to inequity almost exclusively when seated close- *Corresponding author. E-mail: [email protected] advantage (10), although advantageous IA may ly side-by-side, compared with tests in which SCIENCE sciencemag.org 17 OCTOBER 2014 • VOL 346 ISSUE 6207 1251776-1 RESEARCH | REVIEW they sat far apart or across from each other, in to unfair outcomes. Additional work to deter- up a less-preferred, but still beneficial, reward) which few IA responses were observed (Table 1). mine the influence of these and other factors while increasing inequity (the partner still re- This suggests that physical proximity may be in- on animal IA responses will provide additional ceives the preferred reward versus the other re- tegral to IA outcomes, possibly because of the nuance in our understanding of the evolution ceiving nothing). If the goal of IA is to minimize relationship between proximity and cooperation of IA (Table 1). current inequity (7), these animals show the and the way proximity facilitates information First-order IA has been documented in controlled wrong response. gathering about the partner (21). Finally, individ- experiments in capuchin monkeys [(4, 6, 27–29), New lines of evidence, however, have led to a ual differences have been found in some species, but see also (30)], macaques (18, 31), chimpanzees reassessment of this evaluation. First, humans, notably chimpanzees, who show substantial varia- [(16, 22), but see also (32, 23)], dogs (33–35), and too, respond in this way. The workhorse of in- tion even within the same experiment (16, 22, 23). crows (36), and it has been implied in rodents equity studies has been the ultimatum game (UG), Responses also seem influenced by dominance (37). These animals refuse lesser rewards if a in which one individual, the proposer, must de- rank, sex, and relationship quality. This is the partner receives better ones and/or stop perform- cide how to divide a set sum of money. The sec- case in humans as well, where factors such as ing after multiple exposures to such outcomes.