Empathy: Gender effects in brain and behavior

Proactive, that is, unsolicited, prosociality is a key component of our hyper-cooperation, which in turn has enabled the emergence of various uniquely human traits, including complex cognition, morality and cumulative culture and technology. However, the evolutionary foundation of the human prosocial sentiment remains poorly understood, largely because primate data from numerous, often incommensurable testing paradigms do not provide an adequate basis for formal tests of the various functional hypotheses. We therefore present the results of standardized prosociality experiments in 24 groups of 15 primate species, including humans. Extensive allomaternal care is by far the best predictor of interspecific variation in proactive prosociality. Proactive prosocial motivations therefore systematically arise whenever selection favours the evolution of cooperative breeding. Because the human data fit this general primate pattern, the adoption of cooperative breeding by our hominin ancestors also provides the most parsimonious explanation for the origin of human hyper-cooperation.

/pdf/the-evolutionary-origin-of-human-hyper-cooperation-35yfvasjbo.pdf

The evolutionary origin of human hyper-cooperation

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.

/pdf/evolution-of-responses-to-un-fairness-38wurqiy0r.pdf

Evolution of responses to (un)fairness

Take-it or leave-it offers are probably as old as mankind. Our objective here is, first, to provide a, probably subjectively colored, recollection of the initial ultimatum game experiment, its motivation and the immediate responses. Second, we discuss extensions of the standard ultimatum bargaining game in a unified framework, and, third, we offer a survey of the experimental ultimatum bargaining literature containing papers published since the turn of the century. The paper argues that the ultimatum game is a versatile tool for research in bargaining and on social preferences. Finally, we provide examples for open research questions and directions for future studies.

/pdf/more-than-thirty-years-of-ultimatum-bargaining-experiments-2hmhcr5poj.pdf

More than thirty years of ultimatum bargaining experiments: Motives, variations, and a survey of the recent literature

The current review aims to unify existing views of altruism through an examination of the biological bases of a fundamental form of giving: altruistic responding. Altruistic responding is most salient during heroic acts of helping but is also observed any time one perceives another's distress or need, which in turn motivates one to help at a current cost to the self. Such aid is simple, observable across species, and rooted in the instincts and circuits that evolved to maximize inclusive fitness through the care of helpless offspring. By design, the system already biases aid to both kin and nonkin under conditions that are largely adaptive. These inherent benefits are also buttressed in primates and humans by known, later-arriving benefits to helping in group-living animals. Evidence for the proposed homology between altruistic responding and offspring retrieval is presented through 10 key shared factors. Conceptually, both require (a) participation by nonmothers, (b) motor competence and expertise, (c) an adaptive opponency between avoidance and approach, and a facilitating role of (d) neonatal vulnerability, (e) salient distress, and (f) rewarding close contact. Physiologically, they also share neurohormonal support from (g) oxytocin, (h) the domain-general mesolimbocortical system, (i) the cingulate cortex, and (j) the orbitofrontal cortex. The framework intermixes ultimate and proximate levels of analysis and unifies existing views by assuming that even complex human behaviors reflect ancient mammalian neural systems that evolved to solve key problems in adaptive ways, with far-reaching consequences for even our most venerated human traits.

/pdf/the-origins-of-altruism-in-offspring-care-1polv838wc.pdf

The origins of altruism in offspring care.

The study of human and primate altruism faces an evolutionary anomaly: There is ample evidence for altruistic preferences in our own species and growing evidence in monkeys, but one of our closest relatives, the chimpanzee (Pan troglodytes), is viewed as a reluctant altruist, acting only in response to pressure and solicitation. Although chimpanzee prosocial behavior has been reported both in observational captive studies and in the wild, thus far Prosocial Choice Tests have failed to produce evidence. However, methodologies of previous Prosocial Choice Tests may have handicapped the apes unintentionally. Here we present findings of a paradigm in which chimpanzees chose between two differently colored tokens: one “selfish” token resulting in a reward for the actor only (1/0), and the other “prosocial” token rewarding both the actor and a partner (1/1). Seven female chimpanzees, each tested with three different partners, showed a significant bias for the prosocial option. Prosocial choices occurred both in response to solicitation by the partner and spontaneously without solicitation. However, directed requests and pressure by the partner reduced the actor's prosocial tendency. These results draw into question previous conclusions indicating that chimpanzees have a limited sensitivity to the needs of others and behave prosocially only in response to significant prompting.

/pdf/spontaneous-prosocial-choice-by-chimpanzees-1w7ui6em8j.pdf

Spontaneous prosocial choice by chimpanzees

People empathize with fictional displays of behaviour, including those of cartoons and computer animations, even though the stimuli are obviously artificial. However, the extent to which other animals also may respond empathetically to animations has yet to be determined. Animations provide a potentially useful tool for exploring non-human behaviour, cognition and empathy because computer-generated stimuli offer complete control over variables and the ability to program stimuli that could not be captured on video. Establishing computer animations as a viable tool requires that non-human subjects identify with and respond to animations in a way similar to the way they do to images of actual conspecifics. Contagious yawning has been linked to empathy and poses a good test of involuntary identification and motor mimicry. We presented 24 chimpanzees with three-dimensional computer-animated chimpanzees yawning or displaying control mouth movements. The apes yawned significantly more in response to the yawn animations than to the controls, implying identification with the animations. These results support the phenomenon of contagious yawning in chimpanzees and suggest an empathic response to animations. Understanding how chimpanzees connect with animations, to both empathize and imitate, may help us to understand how humans do the same.

/pdf/computer-animations-stimulate-contagious-yawning-in-2tknjwhcpi.pdf

Computer animations stimulate contagious yawning in chimpanzees

In his letter (1), the author argues against prosocial tendencies in chimpanzees because the chimpanzees in our study (2) failed to select the prosocial option all of the time. Instead, he suggests that they show evidence for “mean-spirited” motivations, because any deviation from a perfectly prosocial score must indicate purposeful withholding of benefits for the partner. However, behavioral studies are not conducted in a vacuum, informed by hypothetical analogies. Our study involved real participants in a social environment in which they express myriad conflicting motivations, inhibitions, distractions, emotions, and preexisting social “baggage” brought with them into an experimental setting. Before discussing the potential influence of these factors, let us review some human prosocial choice tests (PCTs) because they yield results that do not substantially differ from our chimpanzee study.

https://www.pnas.org/content/pnas/108/42/E836.full.pdf

Reply to Skoyles: Misplaced assumptions of perfect human prosociality

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Spontaneous prosocial choice by chimpanzees

Computer animations stimulate contagious yawning in chimpanzees

Reply to Skoyles: Misplaced assumptions of perfect human prosociality