Foundations of Human Social Behavior


International Symposium "Foundations of Human Social Behavior"

June 18-21, 2008

The University Research Priority Program “Foundations of Human Social Behavior” is organizing a small, invited symposium that will take place at the University of Zurich from June 18-21, 2008.

The international symposium will bring together key scientists from different social science and biomedical disciplines to address the actionable knowledge needed to answer crucial questions associated with adaptive and maladaptive human social behavior.

The symposium will be comprised of sessions on: Prosocial Behavior, Antisocial Behavior, Understanding Others, and Social Behavior in Cross-Cultural and Cross-Species Perspective.

Prosocial Behavior

Oxytocin, vasopressin and social monogamy: Implications for mental illness
C. Sue Carter, University of Illinois at Chicago, Chicago, IL

Oxytocin is an ancient neuropeptide hormone, previously best known for its role in birth and breastfeeding. Recent research reveals that oxytocin and the related molecule vasopressin regulate the novel traits of social monogamy, including social bonding and parental care. In addition, these same neuropeptides, as well as associated social relationships, have important roles in the regulation of neural and autonomic systems associated with stress and anxiety. We will discuss the causes and consequences of social bonds and related social behaviors, using evidence from humans and other socially monogamous mammals, such as prairie voles. For example, in prairie voles social isolation or exposure to a social stimulus, such as an infant, can causes dynamic changes in oxytocin, vasopressin, and hormones of the hypothalamic-pituitary-adrenal axis, including CRF and adrenal steroids, and may even regulate neurogenesis. Early experiences, including handling and exposure to neuropeptides, can have long lasting effects on the expression of these systems. However, in many cases these effects differ in males and females. We also will describe possible implications of these same neuropeptides and their actions for mental health and mental illnesses, including disorders such as depression, autism and schizophrenia.

Neural bases of individual differences in social and emotional behavior
Richard J. Davidson, University of Wisconsin-Madison, Madison, WI

Variation in reactivity to social and emotional stimuli is the norm rather than the exception in human behavior. This variation is crucial in understanding fundamental modes of adaptation including resilience and psychopathology. By considering the contributions of underlying neural systems to this variation, better measures of individual differences and their neural bases can be obtained. This talk will feature several examples from our laboratory of individual differences in both normal and abnormal social and emotional behavior and their neural bases probed with functional magnetic resonance imaging. Individual differences in prefrontal and amygdala circuitry plays an important role in modulating individual differences in behavior. In one series of studies, individual differences in emotion regulation will be highlighted using a paradigm in which participants are instructed to voluntarily regulate their emotions using cognitive reappraisal strategies. These studies highlight the role of ventromedial and frontopolar regions of prefrontal cortex and their connectivity to the amygdala. Good emotion regulators show greater activation of these prefrontal regions during the down-regulation of negative affect, and lower levels of activation in the amygdala, compared with poor emotion regulators. This pattern is abnormal in patients with depression. In another series of studies, eye tracking is used as a measure of sensitivity to social cues. Variations in eye tracking to facial emotion is related to individual differences in amygdala reactivity. In individuals with autism these patterns are abnormal and are related to structural variation in the amygdala. Together these findings underscore the critical importance of individual differences in any comprehensive understanding of social and emotional behavior.

Cooperative breeding and the origin of prosociality
Carel van Schaik, University of Zurich

Humans are strikingly more cooperative than other primates, and our exceptional cooperation is based on a psychological predisposition known as spontaneous prosociality, other-regarding preferences, or concern for the welfare of others. Its evolutionary roots, however, remain poorly understood. Recent evidence suggests a discontinuity between humans and our closest relatives. Chimpanzees are highly cooperative, yet three independent attempts failed to elicit prosociality in a food donation experiment, suggesting hominins evolved a new psychology of cooperation. However, an experiment found spontaneous food donation among common marmosets, our closest primate relatives to share extensive allomaternal care with us humans. Since most other primates show far less cooperation than chimpanzees, this suggests that we should look to the origin of extensive allomaternal care, i.e. cooperative breeding, to explain the origin of spontaneous prosociality.
We therefore examine and extend Sarah Hrdy’s cooperative breeding hypothesis. Cooperative breeding enabled our hominin ancestors to break through the brain size ceiling found among primates. Cooperative breeding led to improved social cognition in marmosets, relative to their sister group, partly because prosociality leads to an interest in imparting skills and knowledge to others. In hominins, therefore, cooperative breeding probably promoted joint intentionality, intentional teaching, and even the evolution of language. In sum, we suggest that the adoption of cooperative breeding is ultimately responsible for the social and physical cognitive abilities and the skill-intensive foraging niche that are so different from the other great apes.

Antisocial Behavior

The amygdala and ventromedial prefrontal cortex: Functional contributions and impairment in psychopathy
James Blair, National Institute of Mental Health, Bethesda, MD

In this talk, four issues will be addressed. First, I will consider the functional contributions of the amygdala and ventromedial prefrontal cortex that are relevant to understanding the impairment seen in psychopathy. In short, I will consider the amygdala’s role in stimulus-reinforcement association formation (including its role in responding to the reinforcement provided by some facial expressions) and the role of ventromedial frontal cortex in the representation of reinforcement information. Second, I will consider the bases of psychopathy and conclude that there is a substantial genetic contribution to the development of this disorder though social influences clearly influence how this genetic contribution is expressed. Third, I will provide data showing that the functional contributions of the amygdala and ventromedial frontal cortex are profoundly compromised in individuals with psychopathy. I will describe how these impairments disrupt the individual’s socialization, leading to an increased risk for goal-directed, instrumental antisocial behavior. Fourth, I will provide data showing that the role of ventromedial prefrontal cortex in the representation of prediction errors and the readjustment of stimulus-reinforcement associations following contingency change is also profoundly disrupted in psychopathy. I will describe how this impairment is likely to lead to increased frustration and consequently the increased risk for reactive aggression seen in this population.

The impact of neural, psychological, and social factors on the responsible assignment of moral responsibility for action
Shaun Gallagher, University of Central Florida, Orlando, FL

Generally speaking we want to hold individuals responsible for their actions. Recent research in neuroscience and cognitive neuropsychology, however, confirms long-held philosophical suspicions that individuals may not be in full control of their actions – that neural, psychological, and social factors causally impact action decisions. This view raises questions about how responsibility should be assigned. After reviewing some of the recent science, I argue that although science can tell us a lot about causal responsibility it cannot resolve questions about moral responsibility. For example, it may very well be the case that psychopaths and sociopaths have abnormal brains that prevent them from being causally responsible; it remains an open question, however, to be decided in each case, whether we should hold the individual morally responsible for their actions. For all practical purposes in the moral and social realms, the correct question is not “Is the individual responsible for her actions?” but “Should we hold the individual responsible for her actions?” Part of this decision should include reference to causal responsibility, but other factors enter into judgments of moral responsibility. On this side of the issue there are also interesting empirical studies that demonstrate a variety of problems with regard to how people actually do make judgments of moral responsibility. How we make such judgments may be affected by some of the same neural, psychological, and social factors that causally impact our action decisions.

Some key principles of juvenile disruptive behavior and delinquency
Rolf Loeber, University of Pittsburgh, Pittsburgh, PA

Several principles of development have emerged from the scientific literature, which are relevant to disruptive and delinquent children: The Early-onset principle: an early-onset of delinquency or persistent disruptive behaviors, compared to later-onset, is predictive of later chronic, serious, and violent offending. The Age-crime curve: is a universal phenomenon showing that the prevalence of offenders is low in late childhood and early adolescence, peaks in middle to late adolescence and decreases subsequently. The Developmental Pathway Principle refers to findings that the development of serious forms of delinquency almost always is preceded by successive forms of less serious disruptive behavior and minor to moderate serious levels of delinquency. Dose-response principle: there is a dose-response relationship between risk factors and outcomes, meaning that the higher the number of risk factors that a child is exposed to the more likely it is that that child will become seriously delinquent. There also is an inverse dose-response relationship between promotive factors and outcomes, meaning that the higher the number of promotive factors that a child is exposed to, the lower the likelihood that that child will become seriously delinquent. The Buffering principle: the higher the number of promotive compared to risk factors juveniles are exposed to the more likely it is that the promotive factors will buffer or negate the impact of risk factors. Thus, non-delinquency is more likely when promotive factors outstrip risk factors. The same applies to desistance from delinquency: desistance becomes likely the higher the number of promotive compared to risk factors. The Developmental prominence principle: there is emerging evidence that promotive factors tend to predominate in childhood, whereas the number of risk factors that children are exposed to tends to increase between childhood and adolescence. These six principles are of importance in assessing children, and particularly their risk of future serious delinquency, but the principles are also highly relevant for the formulation and evaluation of interventions.

Antisocial behavior: Dysfunctional brain mechanisms and moral decision-making
Adrian Raine, University of Pennsylvania, Philadelphia, PA

The past decade has witnessed an exponential increase in brain imaging research on antisocial individuals which has documented significant structural and functional brain impairments in these individuals. A meta-analysis of 43 imaging studies of the prefrontal cortex shows reduced prefrontal structure and function in antisocial individuals (d = -.64), with significant effect sizes for both structural (d = -.45) and functional (d = -.74) studies, as well as for non-violent antisocial (d = -.57) as well as violent antisocial (d = -.57) groups. Evidence for a neurodevelopmental hypothesis of antisocial behavior is presented based on structural brain imaging findings, together with data showing localization of structural impairments to specific amygdala nuclei in psychopaths. Data are presented suggesting that the gender difference in antisocial behavior is part-mediated by gender differences in orbitofrontal gray. A model of the broader cognitive, affective, and motor systems and brain regions that are dysfunctional in antisocial individuals is also presented. The neural circuitry underlying moral decision-making is outlined, a circuitry argued to be compromised in antisocial populations. Research findings raise significant neuroethical issues that impact society and the criminal justice system, issues which focus on responsibility, freedom of will, punishment, and treatment. Biological manipulations which improve brain structure and function hold the promise of reducing violence and risk for homicide, and one benign method of manipulating neurobiological functioning to reduce risk for aggression through dietary intervention is outlined.

Understanding Others

Psychological antecedents of empathy-induced altruism
C. Daniel Batson, University of Kansas, Lawrence, KS

Do we humans ever, in any degree, care about others for their sakes and not simply for our own? The empathy-altruism hypothesis offers an affirmative answer to this question. It claims that empathic concern (an other-oriented emotional response elicited by and congruent with the perceived welfare of another in need) produces altruistic motivation (a motivational state with the ultimate goal of relieving the need). Over the past 35 years, social psychologists have tested the empathy-altruism hypothesis using laboratory experiments and have found quite strong support. To find that empathy-induced altruistic motivation is within the human repertoire raises the question of what produces empathic concern. Proposed proximal antecedents include: self-other merging, similarity, sharing the other’s feelings, two forms of perspective-taking (imagine-other and imagine-self), and valuing the other’s welfare. Research to date suggests that self-other differentiation, not merging, is necessary to experience empathic concern. Research also suggests that neither similarity nor sharing the other’s feelings is necessary, but that imagining the thoughts and feelings of another in need (imagine-other perspective) is a consistent and powerful antecedent. Imagining oneself in the other’s shoes (imagine-self perspective) may serve as a stepping stone to imagining how the other feels. But it may also lead to a self-centered focus on one’s own feelings, inhibiting empathic concern. In everyday life, intrinsic valuing of another’s welfare may be the key proximal antecedent. The most plausible distal (genetic) antecedent of empathy-induced altruism is not reciprocal altruism, kin selection, or group selection. It is parental nurturance.

A genetic approach to social behavior in humans: Regulatory mechanisms and new phenotypes
Andreas Meyer-Lindenberg, Central Institute of Mental Health (ZI), Mannheim

The regulation of social behavior is a fundamental brain function essential for survival and, as such, under strong genetic control. The investigation of how genetic variation shapes neural circuits relevant for social function in humans is a new line of inquiry. We used imaging genetics in healthy human samples as well as in participants with Williams syndrome to identify neural circuits for human social behavior under genetic control. We also describe work investigating promising target phenotypes in the social domain that show a high degree of conservation across the mammalian lineage and thus have a high prior probability of genetic contribution. We have characterized a regulatory circuit linking amygdala with several regions in prefrontal cortex that implies mechanisms for risk for anxiety and depression as a function of variation in the serotonin transporter gene as well as for genetically abnormal social behavior in Williams syndrome. New data show a strong modulatory effect of prosocial neuropeptides, as well as of genetic variation in neuropeptide receptor genes linked to autism (OXTR, AVPR1A) on this circuit, extending to the human a large body of preclinical work on neuropeptide regulation of complex behavior. A pronounced impact on this same circuit emerges from recent results from studying variation in MAO-A, a gene linked to risk for violence mediated by early abuse. We present data on the neural processing of social hierarchies showing that similar regulatory mechanisms (especially through ventromedial prefrontal cortex) apply for this phenotype. Taken together, these results begin to define neural mechanisms both for genetic influences on complex behavior and their mediation through gene-environment interactions, and may indicate new targets for treating psychiatric disorders where social dysfunction is prominent, such as social phobia, schizophrenia, and autism.

Using self to understand others: fMRI studies of mentalizing
Jason P. Mitchell, Harvard University, Cambridge, MA

Human social interaction requires the recognition that other people are governed by the same type of mental states – beliefs, desires, intentions – that guide one's own behavior. One useful strategy for inferring others' mental states (i.e., mentalizing) may be to use knowledge of one's own thoughts, feelings, and desires as a proxy for those of other people. These self-referential accounts of social cognition are supported by recent research suggesting that a single brain region – ventromedial prefrontal cortex (vmPFC) – is engaged both by tasks that require self-reference and those that require inferences about the minds of others. In this talk, I will review our group's recent work suggesting the boundary conditions of such self-referential mentalizing, such as the selective deployment of such processes for those that are presumed to be similar to self. Conversely, I will review additional work suggesting that one can increase the use of vmPFC-mediated mentalizing processes by engaging in explicit perspective-taking manipulations. Together, these results suggest that for particular mentalizing contexts, the vmPFC subserves both introspecting about one's own mind and considering the mind of others.

Theory or simulation: Is there a third option?
Dan Zahavi, University of Copenhagen

In recent years, much of the discussion of the nature of social cognition has taken place within the framework of the so-called theory of mind debate. Although it was originally assumed that it was the possession and use of a theory that provided the individual with the capacity to attribute mental states, the contemporary debate is split on the issue, and is generally considered to be a dispute between two views. On one side, we find the theory-theory of mind and on the other the simulation theory of mind. The theory-theory of mind and the simulation theory of mind are frequently depicted as quite opposed accounts of the basic nature of social cognition. However, both accounts share certain presuppositions that underlie and shape the very debate. Indeed, despite their many differences, the theory-theory of mind and the simulation theory of mind both share the view that the minds of others are hidden, and they consider one of the main challenges facing a theory of social cognition to be the question of how and why we start ascribing such hidden mental entities or processes to certain publicly observable bodies. Both accounts deny that it is possible to experience other minded creatures. It is precisely because of the absence of an experiential access to the minds of others that we need to rely on and employ either theoretical inferences or internal simulations. In my talk, I will argue that the forced choice between theory-theory and simulation-theory is a false choice, and defend what I consider to be an interesting and significant – though overlooked – alternative; one that to a larger extent emphasizes the embodied and embedded nature of the mind.

Social Behavior in Cross-Cultural and Cross-Species Perspective

Cross-cultural differences norm enforcement
Simon Gächter, University of Nottingham

There is substantial evidence that altruistic punishment of defectors is a powerful mechanism for sustaining social cooperation. In this talk I will present evidence that punishment might not only be directed at defectors, but also at cooperators. One important reason for this 'anti-social punishment' is that punished defectors might retaliate by punishing cooperators (who typically punish the defectors). We document the widespread existence of anti-social punishment in public goods experiments which we conducted in sixteen comparable subject pools around the world. We found a very large cross-societal variation in anti-social punishment. In some subject pools anti-social punishment is as severe as altruistic punishment of defectors. In some subject pools, in particular those were previous research on altruistic punishment has been conducted, anti-social punishment is negligible (and has therefore largely been ignored in previous research). As a consequence of (anti-social) punishment, cooperation is stabilized at vastly different levels. In some subject pools anti-social punishment strongly undermines the cooperation-enhancing effect of punishment opportunities as observed in previous research. Our findings provide novel support for culture-gene co-evolutionary theories, which predict that multiple equilibria can arise even in identical environments. To our knowledge, our study reports the largest cross-cultural variation observed in experimental cooperation games with punishment. Moreover, we are also able to show for the first time that punishment behavior is related to societal-level social norms of cooperation. In particular, we show that weak norms of civic cooperation and the weakness of the Rule of Law in a society are significant predictors of anti-social punishment. We conclude that private punishment opportunities are socially beneficial only if they are complemented by strong social norms of cooperation which constrain anti-social punishment.

The coevolution of human sociality and the prosocial institutions: Behavioral experiments and ethnography from 15 small-scale societies
Joseph Henrich, University of British Columbia, Vancouver, BC

Speculation on the nature of human sociality has a long history among social theorists. Only recently, however, have social scientists begun to systematically deploy experimental tools to chart the depth and topography of human social behavior and psychology. Because of the predictive clarity of Economics' formalized theory, the impact of this still rising tide of empirical findings has been felt most acutely there, where foundational assumptions once taken as axiomatic are being rapidly reconsidered and revised. Unfortunately, the robust empirical findings upon which these new theories are built come almost exclusively from university student populations in the U.S. and Europe – with occasional "cross-cultural" forays to universities in Asia – and are devoid of detailed individual-level data about peoples' lives outside the laboratory. To address both of these empirical deficiencies, my co-researchers and I have combined ethnographic and behavioral experimental techniques in two synchronized studies of 15 small-scale societies (23 different societies across the two projects) that includes hunter-gatherers, pastoralists, swidden horticulturalists and small-scale sedentary farmers from Africa, South America, Oceania, New Guinea and Northern Asia. Using Dictator, Ultimatum, and Third Party Punishment Games (as well as some Public Goods and Trust Games), we find four important patterns in human preferences. First, experimental measures across all societies reveal consistent prosocial deviations from the predictions based on pure self interest. Second, while all societies studied violate the predictions of the canonical model, different societies violate them to substantially differing degrees, and sometimes in qualitatively different ways. Third, despite the group-level variation, we find consistent evidence across our sample that the threat of punishment crowds out intrinsic prosocial motivations. Finally, both our measures of fairness and punishment are predicted independently by measures of market integration, population size, and association with a world religion (vs. a traditional-local religion). Taken together, these findings greatly weaken the species-level generality of most theoretical reformulations in Economics and elsewhere, and favor a new approach to human sociality that endogenizes the cultural coevolution of human social motivations and institutions.

How do emotions relate to morality? A review of competing models
Jesse J. Prinz, University of North Carolina at Chapel Hill, Chapel Hill, NC

There is a growing body of evidence that emotions occur when people make moral judgments. This fact alone, however, is consistent with a variety of processing models. Emotions might be the effects of moral judgment, or the causes of moral judgments, or components. Emotions might be involved in all moral judgments, or just some (a dual process theory). I argue that emotions are components of moral judgments and against dual processing theory. I also consider competing “philosophical models,” i.e., metaethical theories of the role that emotions play in morality. I present empirical evidence of a “response-dependent” view as opposed to an “error theory” or “emotivism.”

Cooperation and helping in children and chimpanzees
Michael Tomasello, Max Planck Institute for Evolutionary Anthropology, Leipzig

Recent studies in our laboratory have focused on the behavior of great apes and human children in situations involving: (1) helping, (2) cooperation, and (3) punishment and norms. Apes engage in some helping, when food is not involved; children help much more widely and are internally motivated to do so. Apes can cooperate when sharing the spoils at the end has been made less contentious and when they are already more tolerant of one another; children cooperate in more complex ways either for its own sake or with a view to sharing the spoils at the end. Apes retaliate against those who have harmed them, but they do not enforce norms from a third-party perspective; children enforce norms from a third-party perspective not only for moral behaviors but also for simple conventions ("This is the way it's done"). A tentative evolutionary account based on these findings will be proposed.

Poster Sessions

Young researchers (doctoral or postdoctoral students with less than five years’ postgraduate experience) from relevant fields were invited to apply for an invitation to attend the symposium and present their research in a poster session. The Organizing Committee reviewed their applications and has sent invitations to selected applicants. (Please understand that, as the submission deadline has passed, it is no longer possible to apply for an invitation.)

Travel Awards

Ten of the young researchers invited to participate in the symposium will receive a Travel Award of 500 Swiss francs.

Organizing Committee

Ingolf U. Dalferth
Ernst Fehr
Urs Fischbacher
Markus Heinrichs
Michael Kosfeld
Tania Singer

Please address questions concerning the symposium to Tamara Herz (