A category of stimuli of great importance for primates, humans in particular, is that formed by actions done by other individuals. If we want to survive, we must understand the actions of others. Furthermore, without action understanding, social organization is impossible. In the case of humans, there is another faculty that depends on the observation of others' actions: imitation learning. Unlike most species, we are able to learn by imitation, and this faculty is at the basis of human culture. In this review we present data on a neurophysiological mechanism--the mirror-neuron mechanism--that appears to play a fundamental role in both action understanding and imitation. We describe first the functional properties of mirror neurons in monkeys. We review next the characteristics of the mirror-neuron system in humans. We stress, in particular, those properties specific to the human mirror-neuron system that might explain the human capacity to learn by imitation. We conclude by discussing the relationship between the mirror-neuron system and language.

The mirror-neuron system.

Behavioral scientists routinely publish broad claims about human psychology and behavior in the world's top journals based on samples drawn entirely from Western, Educated, Industrialized, Rich, and Democratic (WEIRD) societies. Researchers - often implicitly - assume that either there is little variation across human populations, or that these "standard subjects" are as representative of the species as any other population. Are these assumptions justified? Here, our review of the comparative database from across the behavioral sciences suggests both that there is substantial variability in experimental results across populations and that WEIRD subjects are particularly unusual compared with the rest of the species - frequent outliers. The domains reviewed include visual perception, fairness, cooperation, spatial reasoning, categorization and inferential induction, moral reasoning, reasoning styles, self-concepts and related motivations, and the heritability of IQ. The findings suggest that members of WEIRD societies, including young children, are among the least representative populations one could find for generalizing about humans. Many of these findings involve domains that are associated with fundamental aspects of psychology, motivation, and behavior - hence, there are no obvious a priori grounds for claiming that a particular behavioral phenomenon is universal based on sampling from a single subpopulation. Overall, these empirical patterns suggests that we need to be less cavalier in addressing questions of human nature on the basis of data drawn from this particularly thin, and rather unusual, slice of humanity. We close by proposing ways to structurally re-organize the behavioral sciences to best tackle these challenges.

/pdf/the-weirdest-people-in-the-world-480wr4m3kb.pdf

The WEIRDest People in the World

Research on moral judgment has been dominated by rationalist models, in which moral judgment is thought to be caused by moral reasoning. The author gives 4 reasons for considering the hypothesis that moral reasoning does not cause moral judgment; rather, moral reasoning is usually a post hoc construction, generated after a judgment has been reached. The social intuitionist model is presented as an alternative to rationalist models. The model is a social model in that it deemphasizes the private reasoning done by individuals and emphasizes instead the importance of social and cultural influences. The model is an intuitionist model in that it states that moral judgment is generally the result of quick, automatic evaluations (intuitions). The model is more consistent than rationalist models with recent findings in social, cultural, evolutionary, and biological psychology, as well as in anthropology and primatology.

https://www.motherjones.com/files/emotional_dog_and_rational_tail.pdf

The emotional dog and its rational tail: a social intuitionist approach to moral judgment.

We demonstrate that a simple model, constructed on the premise that people are motivated by both their pecuniary payoff and their relative payoff standing, organizes a large and seemingly disparate set of laboratory observations as one consistent pattern The model is incomplete information but nevertheless posed entirely in terms of directly observable variables The model explains observations from games where equity is thought to be a factor, such as ultimatum and dictator, games where reciprocity is thought to play a role, such as the prisoner's dilemma and gift exchange, and games where competitive behavior is observed, such as Bertrand markets (JEL C78, C90, D63, D64, H41)

/pdf/erc-a-theory-of-equity-reciprocity-and-competition-9lhkqwqjzg.pdf

ERC: A Theory of Equity, Reciprocity, and Competition

The anterior insular cortex (AIC) is implicated in a wide range of conditions and behaviours, from bowel distension and orgasm, to cigarette craving and maternal love, to decision making and sudden insight. Its function in the re-representation of interoception offers one possible basis for its involvement in all subjective feelings. New findings suggest a fundamental role for the AIC (and the von Economo neurons it contains) in awareness, and thus it needs to be considered as a potential neural correlate of consciousness.

How do you feel--now? The anterior insula and human awareness.

There is disagreement in the literature about the exact nature of the phenomenon of empathy. There are emotional, cogni- tive, and conditioning views, applying in varying degrees across species. An adequate description of the ultimate and proximate mecha- nism can integrate these views. Proximately, the perception of an object's state activates the subject's corresponding representations, which in turn activate somatic and autonomic responses. This mechanism supports basic behaviors (e.g., alarm, social facilitation, vicar- iousness of emotions, mother-infant responsiveness, and the modeling of competitors and predators) that are crucial for the reproduc- tive success of animals living in groups. The Perception-Action Model (PAM), together with an understanding of how representations change with experience, can explain the major empirical effects in the literature (similarity, familiarity, past experience, explicit teach- ing, and salience). It can also predict a variety of empathy disorders. The interaction between the PAM and prefrontal functioning can also explain different levels of empathy across species and age groups. This view can advance our evolutionary understanding of empa- thy beyond inclusive fitness and reciprocal altruism and can explain different levels of empathy across individuals, species, stages of de- velopment, and situations.

/pdf/empathy-its-ultimate-and-proximate-bases-3gnr9okjki.pdf

Empathy: Its ultimate and proximate bases.

Evolutionary theory postulates that altruistic behavior evolved for the return-benefits it bears the performer. For return-benefits to play a motivational role, however, they need to be experienced by the organism. Motivational analyses should restrict themselves, therefore, to the altruistic impulse and its knowable consequences. Empathy is an ideal candidate mechanism to underlie so-called directed altruism, i.e., altruism in response to anothers's pain, need, or distress. Evidence is accumulating that this mechanism is phylogenetically ancient, probably as old as mammals and birds. Perception of the emotional state of another automatically activates shared representations causing a matching emotional state in the observer. With increasing cognition, state-matching evolved into more complex forms, including concern for the other and perspective-taking. Empathy-induced altruism derives its strength from the emotional stake it offers the self in the other's welfare. The dynamics of the empathy mechanism a...

/pdf/putting-the-altruism-back-into-altruism-the-evolution-of-3jxkydidt6.pdf

Putting the Altruism Back into Altruism: The Evolution of Empathy

Prologue Darwinian Dilemmas Survival of the Unfittest Biologicizing Morality Calvinist Sociobiology A Broader View The Invisible Grasping Organ Ethology and Ethics Photo Essay: Closeness Sympathy Warm Blood in Cold Waters Special Treatment of the Handicapped Responses to Injury and Death Having Broad Nails The Social Mirror Lying and Aping Apes Simian Sympathy A World without Compassion Photo Essay: Cognition and Empathy Rank and Order A Sense of Social Regularity The Monkey's Behind Guilt and Shame Unruly Youngsters The Blushing Primate Two Genders, Two Moralities? Umbilical versus Confrontational Bonds Primus inter Pares Quid pro Quo The Less-than-Golden Rule Mobile Meals At the Circle's Center A Concept of Giving Testing for Reciprocity From Revenge to Justice Photo Essay: Help from a Friend Getting Along The Social Cage The Relational Model Peacemaking Rope Walking Baboon Testimony Draining the Behavioral Sink Community Concern Photo Essay: War and Peace Conclusion What Does It Take to Be Moral? Floating Pyramids A Hole in the Head Notes Bibliography Acknowledgments Index

Good Natured: The Origins of Right and Wrong in Humans and Other Animals

During the evolution of cooperation it may have become critical for individuals to compare their own efforts and pay-offs with those of others. Negative reactions may occur when expectations are violated. One theory proposes that aversion to inequity can explain human cooperation within the bounds of the rational choice model1, and may in fact be more inclusive than previous explanations2,3,4,5,6,7,8. Although there exists substantial cultural variation in its particulars, this ‘sense of fairness’ is probably a human universal9,10 that has been shown to prevail in a wide variety of circumstances11,12,13. However, we are not the only cooperative animals14, hence inequity aversion may not be uniquely human. Many highly cooperative nonhuman species seem guided by a set of expectations about the outcome of cooperation and the division of resources15,16. Here we demonstrate that a nonhuman primate, the brown capuchin monkey (Cebus apella), responds negatively to unequal reward distribution in exchanges with a human experimenter. Monkeys refused to participate if they witnessed a conspecific obtain a more attractive reward for equal effort, an effect amplified if the partner received such a reward without any effort at all. These reactions support an early evolutionary origin of inequity aversion.

https://courses.washington.edu/anmind/Brosnan%20&%20de%20Waal%20-%20monkeys%20reject%20unequal%20pay%20-%20Nature%202003.pdf

Monkeys reject unequal pay

Does biology condemn the human species to violence and war? Previous studies of animal behavior incline us to answer yes, but the message of this book is considerably more optimistic. Without denying our heritage of aggressive behavior, Frans de Waal describes powerful checks and balances in the makeup of our closest animal relatives, and in so doing he shows that to humans making peace is as natural as making war. In this meticulously researched and absorbing account, we learn in detail how different types of simians cope with aggression, and how they make peace after fights. Chimpanzees, for instance, reconcile with a hug and a kiss, whereas rhesus monkeys groom the fur of former adversaries. By objectively examining the dynamics of primate social interactions, de Waal makes a convincing case that confrontation should not be viewed as a barrier to sociality but rather as an unavoidable element upon which social relationships can be built and strengthened through reconciliation. The author examines five different species--chimpanzees, rhesus monkeys, stump-tailed monkeys, bonobos, and humans--and relates anecdotes, culled from exhaustive observations, that convey the intricacies and refinements of simian behavior. Each species utilizes its own unique peacemaking strategies. The bonobo, for example, is little known to science, and even less to the general public, but this rare ape maintains peace by means of sexual behavior divorced from reproductive functions; sex occurs in all possible combinations and positions whenever social tensions need to be resolved. "Make love, not war" could be the bonobo slogan. De Waal's demonstration of reconciliation in both monkeys andapes strongly supports his thesis that forgiveness and peacemaking are widespread among nonhuman primates--an aspect of primate societies that should stimulate much needed work on human conflict resolution.

Frans B. M. de Waal

Papers

Empathy: Its ultimate and proximate bases.

Putting the Altruism Back into Altruism: The Evolution of Empathy

Good Natured: The Origins of Right and Wrong in Humans and Other Animals

Monkeys reject unequal pay

Peacemaking among primates