Evolution of responses to (un)fairness

Background The human sense of fairness presents an evolutionary puzzle because it appears to run counter to the short-term interests of at least some parties. We not only react negatively to getting less than a partner but sometimes also to getting more, which seems illogical. Following an ideal of impartiality, we seek appropriate outcomes for everyone within the community, not just a few individuals, and, in particular, not just ourselves. Why do we react this way? Are we the only species to do so? Here, we consider the evidence with regard to nonhuman primates and other animals to illuminate the evolution of the sense of fairness. Because social ideals escape measurement, we focus on behavioral responses to equal versus unequal reward division. Moreover, a true sense of fairness includes taking account of receiving both less than a partner and more than a partner. Therefore, we consider the evidence for both of these in other species and how this informs our understanding of the evolution of fairness in humans. A diagram of the relationship between first-order IA and second-order IA. Individual A received high-level rewards, and individual B received low-level rewards. Individuals who recognize when they receive less than another may react against this situation—for instance, by finding a new cooperative partner. As reliance on cooperation increases, individuals also benefit from recognizing when they receive more, as this allows them to forestall first-order IA reactions in their partners and thereby maintain a successful cooperative relationship. This is the foundation of the full-blown human sense of fairness. A diagram of the relationship between first-order IA and second-order IA. Individual A received high-level rewards, and individual B received low-level rewards. Individuals who recognize when they receive less than another may react against this situation—for instance, by finding a new cooperative partner. As reliance on cooperation increases, individuals also benefit from recognizing when they receive more, as this allows them to forestall first-order IA reactions in their partners and thereby maintain a successful cooperative relationship. This is the foundation of the full-blown human sense of fairness. Advances There is widespread evidence for sensitivity to receiving less than a partner, or “first-order inequity aversion” (IA), in species that cooperate outside mating bonds and kinship. In these studies, animals are paired with a social partner who receives a preferred reward for completing a task. Subjects may respond by refusing to participate or refusing to accept the food reward, and such reactions are compared with those following control tests in which both subjects receive the same reward for the same effort. Increased responses when the partner receives a preferred reward are indicative of a sensitivity to inequity. Thus far, passive and active protest against unfavorable outcomes has been documented in monkeys, apes, dogs, and birds. It is thought that these species compare their outcomes with those of others so as to judge the merit of their partnerships. They may turn away from partners that appropriate more than their fair share of the yields of cooperation. A complete sense of fairness also includes second-order IA, however, which seeks to equalize outcomes even at a short-term cost to the self. This requires individuals to give up an immediate benefit to stabilize a long-term valuable cooperative relationship. Second-order IA reactions have thus far been found only in humans and apes. We hypothesize that second-order IA requires anticipation of first-order IA in the partner and its negative impact on the relationship. To forestall these consequences, and ensure continued cooperation, outcomes are equalized between partners. Outlook Thus, humans and other species seem to share basic reactions to inequity, which serves to sustain cooperation. We postulate that the basic emotional reactions and calculations underlying our sense of fairness are rooted in our primate background and offer a model that places these reactions in the context of cooperative relationships. Future research should more explicitly investigate the key variables underlying IA, such as the degree of dependence on cooperation, anticipation of the way resource division affects relationships, and the freedom to choose among and change partners. A cross-species investigation with a standardized paradigm, including both first- and second-order IA, may further illuminate the factors involved and help verify or falsify the model proposed. The evolutionary benefits of behaving fairly Humans have a deep and innate sense of fairness. Humans, however, are not the only species to react to apparent inequities. Brosnan and de Waal propose that inequity aversion can be broken down into two levels. At the most basic level, individuals react to immediate unequal distribution of a reward for equal effort expended, whereas at the second, they show the ability to accept a current unequal distribution with the expectation that over time distribution will equalize. This second level facilitates cooperation over time and requires the cognitive abilities both to assess current distribution and envision future opportunities for equalization. As cognitive abilities advanced across the primate lineage, this more complex accounting of equal distribution and cooperation may have developed into the complete sense of fairness we see in humans today. Science, this issue 10.1126/science.1251776 The human sense of fairness is an evolutionary puzzle. To study this, we can look to other species, in which this can be translated empirically into responses to reward distribution. Passive and active protest against receiving less than a partner for the same task is widespread in species that cooperate outside kinship and mating bonds. There is less evidence that nonhuman species seek to equalize outcomes to their own detriment, yet the latter has been documented in our closest relatives, the apes. This reaction probably reflects an attempt to forestall partner dissatisfaction with obtained outcomes and its negative impact on future cooperation. We hypothesize that it is the evolution of this response that allowed the development of a complete sense of fairness in humans, which aims not at equality for its own sake but for the sake of continued cooperation.