Longitudinal evidence that infants develop their imitation abilities by being imitated.

Humans are widely considered the most socially sophisticated species on the planet. Their remarkable abilities in navigating the social world have given rise to complex societies and the advancement of cultural intelligence.1,2,3,4,5 But what characterizes us as ultra-social beings? Theoretical advances in social sciences over the last century purport imitation as a central mechanism for the emergence of humans' unique social-cognitive abilities.6,7,8 Uncovering the ontogeny of imitation is therefore paramount for understanding human cultural evolution. Yet how humans become able to imitate is unclear and intensely debated. Recently, multidisciplinary findings have challenged long-standing assumptions that imitation is inborn.9,10,11,12 So what are the underlying processes supporting the development of imitation? One fascinating possibility is that infants become able to imitate by being imitated.13,14,15 Cognitive theories have suggested that by perceiving others imitating one's own behavior, visual and motor representations of that behavior are coactivated and associated, leading to the emergence of imitation abilities.14,15 Here, we show that being imitated by sensitive caregivers in infancy constitutes a psychological process giving rise to infants' imitation abilities. Results demonstrated (1) that maternal imitation at 14 months positively predicted infants' imitation abilities at 18 months and (2) that maternal imitation at 14 months mediated the positive effect of maternal sensitivity at 6 months on infants' imitation abilities at 18 months. This offers substantial evidence for the role of social interactions in the emergence of imitation as a key factor for human cultural learning.

Compliance or empathy-What links maternal sensitivity and toddlers' emotional helping?

Emotional helping-that is, helping based on others' emotional distress-has been suggested to be a central prosocial response to others in need. Developmental theorizing proposed that emotional helping has social origins. Whereas research indeed demonstrated a link between maternal sensitivity and children's emotional helping, developmental theories stress different mediating processes. Emotion-sharing theories claim empathic concern to be the crucial link for helping, whereas internalization theories base children's helping on children's compliance. To investigate these hypotheses, the current study explored empathy and compliance as two possible mediators for the relation between maternal sensitivity and children's emotional helping at 18 months of age. Overall, maternal sensitivity was positively related to children's empathy, children's compliance, and children's emotional helping. Interestingly, children's empathy-but not children's compliance-mediated the link between maternal sensitivity and children's emotional helping. These findings deepen our understanding of the psychological processes subserving emotional helping during infancy and support theories that stress the socioemotional origins of children's prosocial behavior.

How does the moral self-concept relate to prosocial behaviour? Investigating the role of emotions and consistency preference.

The moral self-concept has been proposed as a central predictor of prosocial behaviour. In two experiments (one preregistered), we explored the nature of the relation between the moral self-concept (explicit and implicit) and prosocial behaviour. Specifically, we investigated the role of emotions associated with prosocial behaviour (consequential or anticipated) and preference for consistency. The results revealed a relation between the explicit moral self-concept and sharing behaviour. The explicit moral self-concept was linked to anticipated and consequential emotions regarding not-sharing. Importantly, anticipated and consequential emotions about not-sharing mediated the relation between self-concept and behaviour. Yet, the relation was independent of preference for consistency. The implicit moral self-concept was neither related to prosocial behaviour nor to emotions associated with behaviour. Overall, our study demonstrates the interplay between cognitive and emotional processes in explaining prosocial behaviour. More specific, it underlines the link between the moral self-concept and prosocial behaviour and highlights the role of emotions about the omission of prosocial behaviour.

Neural processing of moral content reflects moral identity in 10-year-old children.

Middle childhood seems to be crucial for the emergence of a moral identity, that is, an evaluative stance of how important it is for someone's sense of self to be moral. This study investigates the effects of moral identity on the neural processing of moral content in 10-year-old children. Participants were presented with scenes portraying prosocial and antisocial behavior, while electroencephalographic responses were collected. Analyses of event-related potentials (ERPs) showed that, for children with a strong moral identity, antisocial scenes elicited a greater early posterior negativity (EPN) as compared to prosocial scenes. Thus, for children with a strong moral identity, antisocial scenes capture more attentional resources than prosocial ones in early processing stages. In contrast to previous findings with adults, the implicit moral self-concept was not related to any ERP differences. Overall, the results show that, even in its developmental emergence, moral identity relates to the neurocognitive processing of third-party moral content. Together, the study supports the social-cognitive model of the development of moral identity, according to which moral identity is based on a chronical activation of moral schemas that guide a person's perception of the social world.

Detached empathic experience of others' pain in remitted states of depression - An fMRI study.

BACKGROUND: Major depressive disorder is strongly associated with impairments and difficulties in social interactions. Deficits in empathy, a vital skill for social interactions, have been identified as a risk factor for relapse. However, research on empathy in remitted states of depression is scarce. We chose a social neuroscience approach to investigate potentially altered neural processes involved in sub-components of empathy in remitted states of depression. We expected aberrations in cognitive components of empathy, based on previous reports regarding their role as risk factors for relapse. METHODS: Employing functional magnetic resonance imaging and a pain empathy task (video clips of painful medical treatments), we compared behavioral and neural empathic responses of unmedicated remitted depressive patients (N = 32) to those of untreated acutely depressed patients (N = 29) and healthy controls (N = 35). Self-report ratings of pain evaluation and affect-sharing were obtained. RESULTS: Compared to controls and acutely depressed patients, remitted depressive patients reported higher pain evaluation and showed increased activity in the right temporo-parietal junction. This region, which is central to self-other distinction and which has been linked to adopting a detached perspective, also exhibited reduced connectivity to the anterior insula. Furthermore, we observed reduced activity in regions involved in emotion processing (amygdala) and perception of affective facial expressions (fusiform face area, posterior superior temporal sulcus). CONCLUSIONS: Remitted states of depression are associated with a detached empathic style in response to others' pain, characterized by increased self-other distinction, lowered affective processing, and reduced connectivity between empathy-related brain regions. Although this may prevent emotional harm in specific situations, it may reduce opportunities for positive experiences in social interactions in the long run.

Does agency matter? Neural processing of robotic movements in 4- and 8-year olds.

Despite the increase in interactions between children and robots, our understanding of children's neural processing of robotic movements is limited. The current study theorized that motor resonance hinges on the agency of an actor: its ability to perform actions volitionally. As one of the first studies with a cross-sectional sample of preschoolers and older children and with a specific focus on robotic action (rather than abstract non-human action), the current study investigated whether the perceived agency of a robot moderated children's motor resonance for robotic movements, and whether this changed with age. Motor resonance was measured using electroencephalography (EEG) by assessing mu power while 4 and 8-year-olds observed actions performed by agentic versus non-agentic robots and humans. Results show that older children resonated more strongly with non-agentic than agentic robotic or human movement, while no such differences were found for preschoolers. This outcome is discussed in terms of a predictive coding account of motor resonance. Importantly, these findings contribute to the existing set of studies on this topic by showing that, while keeping all kinematic information constant, there is a clear developmental difference in how children process robotic movement depending on the level of agency of a robot.

Neural processing of equitable and inequitable distributions in 5-year-old children.

This study aimed at investigating the neurocognitive correlates of the perception and evaluation of equitable and inequitable distributions in five-year-old children. Children observed one character distributing toys or candies between two recipients. One of the recipients already possessed many resources, and the other possessed just a few. We used event-related potentials to compare brain activity elicited by equitable (the poor receives more) and inequitable (the rich receives more) distribution. On a behavioral level, children evaluated inequitable distribution as worse than equitable and considered the distributor as mean and worthy of punishment when she distributed inequitably as compared to equitably. On the neural level, we expected to find a MFN effect between 250 and 350 ms after picture onset. Instead, we found a frontal positivity (P2), which was greater for inequitable vs. equitable distributions, indicating greater saliency and attentional capture. This was followed by marginally significant greater positivity for equitable distributions between 600 and 1000 ms after picture onset (LPP), which indicates greater allocation of processing resources. Furthermore, a greater LPP was associated with more extreme evaluations for both conditions. This suggests that the more resources children invest in processing the distribution, the more they endorse equity and condemn inequity.

Antidepressant treatment, not depression, leads to reductions in behavioral and neural responses to pain empathy.

Major depressive disorder (MDD) has been hypothesized to lead to impairments in empathy. Previous cross-sectional studies did not disentangle effects of MDD itself and antidepressant treatment. In this first longitudinal neuroimaging study on empathy in depression, 29 patients with MDD participated in two functional magnetic resonance imaging (fMRI) sessions before and after 3 months of antidepressant therapy. We compared their responses to an empathy for pain task to a group of healthy controls (N = 35). All participants provided self-report ratings targeting cognitive (perspective taking) and affective (unpleasant affect) aspects of empathy. To control for general effects on processing of negative affective states, participants additionally underwent an electrical pain task. Before treatment, we found no differences in empathic responses between controls and patients with MDD. After treatment, patients showed significant decreases in both affective empathy and activity of three a priori selected brain regions associated with empathy for pain. Decreases in affective empathy were moreover correlated with symptom improvement. Moreover, functional connectivity during the empathy task between areas associated with affective (anterior insula) and cognitive (precuneus) empathy decreased between sessions in the MDD group. Neither cognitive empathy nor responses to painful electrical shocks were changed after treatment. These findings contradict previous cross-sectional reports of empathy deficits in acute MDD. Rather, they suggest that antidepressant treatment reduces the aversive responses triggered by exposure to the suffering of others. Importantly, this cannot be explained by a general blunting of negative affect, as treatment did not change self-experienced pain.

Moral identity relates to the neural processing of third-party moral behavior.

Moral identity, or moral self, is the degree to which being moral is important to a person's self-concept. It is hypothesized to be the `missing link' between moral judgment and moral action. However, its cognitive and psychophysiological mechanisms are still subject to debate. In this study, we used event-related potentials to examine whether the moral self-concept is related to how people process prosocial and antisocial actions. To this end, participants' implicit and explicit moral self-concept were assessed. We examined whether individual differences in moral identity relate to differences in early, automatic processes [i.e. Early Posterior Negativity (EPN), N2] or late, cognitively controlled processes (i.e. late positive potential) while observing prosocial and antisocial situations. Results show that a higher implicit moral self was related to a lower EPN amplitude for prosocial scenarios. In addition, an enhanced explicit moral self was related to a lower N2 amplitude for prosocial scenarios. The findings demonstrate that the moral self affects the neural processing of morally relevant stimuli during third-party evaluations. They support theoretical considerations that the moral self already affects (early) processing of moral information.

Daytime REM sleep affects emotional experience but not decision choices in moral dilemmas.

Moral decision-making depends on the interaction between automatic emotional responses and rational cognitive control. A natural emotional regulator state seems to be sleep, in particular rapid eye movement (REM) sleep. We tested the impact of daytime sleep, either with or without REM, on moral decision. Sixty participants were presented with 12 sacrificial (6 Footbridge- and 6 Trolley-type) and 8 everyday-type moral dilemmas at 9 AM and at 5 PM. In sacrificial dilemmas, participants had to decide whether or not to kill one person to save more people (utilitarian choice), and to judge how morally acceptable the proposed choice was. In everyday-type dilemmas, participants had to decide whether to endorse moral violations involving dishonest behavior. At 12 PM, 40 participants took a 120-min nap (17 with REM and 23 with NREM only) while 20 participants remained awake. Mixed-model analysis revealed that participants judged the utilitarian choice as less morally acceptable in the afternoon, irrespective of sleep. We also observed a negative association between theta activity during REM and increased self-rated unpleasantness during moral decisions. Nevertheless, moral decision did not change across the day and between groups. These results suggest that although both time and REM sleep may affect the evaluation of a moral situation, these factors did not ultimately impact the individual moral choices.

It's immoral, but I'd do it! Psychopathy traits affect decision-making in sacrificial dilemmas and in everyday moral situations.

This research investigated whether emotional hyporeactivity affects moral judgements and choices of action in sacrificial moral dilemmas and in everyday moral conflict situations in which harm to other's welfare is differentially involved. Twenty-six participants with high trait psychopathy (HP) and 25 with low trait psychopathy (LP) were selected based on the primary psychopathy scale of the Levenson Self-Report Psychopathy Scale. HP participants were more likely to sacrifice one person to save others in sacrificial dilemmas and to pursue a personal advantage in everyday moral situations entailing harm to another's good. While deciding in these situations, HP participants experienced lower unpleasantness as compared to LP participants. Conversely, no group differences emerged in choice of action and unpleasantness ratings for everyday moral situations that did not entail harm to others. Importantly, moral judgements did not differ in the two groups. These results suggest that high psychopathy trait affects choices of action in sacrificial dilemmas because of reduced emotional reactivity to harmful acts. The dissociation between choice of action and moral judgement suggests that the former is more closely related to emotional experience. Also, emotion seems to play a critical role in discriminating harmful from harmless acts and in driving decisions accordingly.

Will I Regret It? Anticipated Negative Emotions Modulate Choices in Moral Dilemmas.

We tested if post-decisional emotions of regret, guilt, shame, anger, and disgust can account for individuals' choices in moral dilemmas depicting the choice of letting some people die (non-utilitarian option) or sacrificing one person to save them (utilitarian option). We collected participants' choices and post-decisional emotional ratings for each option using Footbridge-type dilemmas, in which the sacrifice of one person is the means to save more people, and Trolley-type dilemmas, in which the sacrifice is only a side effect. Moreover, we computed the EEG Readiness Potential to test if the neural activity related to the last phase of decision-making was related to the emotional conflict. Participants reported generally stronger emotions for the utilitarian as compared to the non-utilitarian options, with the exception of anger and regret, which in Trolley-type dilemmas were stronger for the non-utilitarian option. Moreover, participants tended to choose the option that minimized the intensity of negative emotions, irrespective of dilemma type. No significant relationship between emotions and the amplitude of the Readiness Potential emerged. It is possible that anticipated post-decisional emotions play a role in earlier stages of decision-making.

Gaze cuing of attention in snake phobic women: the influence of facial expression.

Only a few studies investigated whether animal phobics exhibit attentional biases in contexts where no phobic stimuli are present. Among these, recent studies provided evidence for a bias toward facial expressions of fear and disgust in animal phobics. Such findings may be due to the fact that these expressions could signal the presence of a phobic object in the surroundings. To test this hypothesis and further investigate attentional biases for emotional faces in animal phobics, we conducted an experiment using a gaze-cuing paradigm in which participants' attention was driven by the task-irrelevant gaze of a centrally presented face. We employed dynamic negative facial expressions of disgust, fear and anger and found an enhanced gaze-cuing effect in snake phobics as compared to controls, irrespective of facial expression. These results provide evidence of a general hypervigilance in animal phobics in the absence of phobic stimuli, and indicate that research on specific phobias should not be limited to symptom provocation paradigms.

Evaluation of the legal consequences of action affects neural activity and emotional experience during the resolution of moral dilemmas.

In any modern society killing is regarded as a severe violation of the legal codes that is subjected to penal judgment. Therefore, it is likely that people take legal consequences into account when deciding about the hypothetical killing of one person in classic moral dilemmas, with legal concerns contributing to decision-making. In particular, by differing for the degree of intentionality and emotional salience, Footbridge- and Trolley-type dilemmas might promote differential assignment of blame and punishment while implicating the same severity of harm. The present study was aimed at comparing the neural activity, subjective emotional reactions, and behavioral choices in two groups of participants who either took (Legal group) or did not take (No Legal group) legal consequences into account when deciding on Footbridge-type and Trolley-type moral dilemmas. Stimulus- and response-locked ERPs were measured to investigate the neural activity underlying two separate phases of the decision process. No difference in behavioral choices was found between groups. However, the No Legal group reported greater overall emotional impact, associated with lower preparation for action, suggesting greater conflict between alternative motor responses representing the different decision choices. In contrast, the Legal group showed an overall dampened affective experience during decision-making associated with greater overall action readiness and intention to act, reflecting lower conflict in responding. On these bases, we suggest that in moral dilemmas legal consequences of actions provide a sort of reference point on which people can rely to support a decision, independent of dilemma type.