The mind minds minds: The effect of intentional stance on the neural encoding of joint attention Abstract Recent neuroimaging studies have observed that the neural processing of social cues from a virtual reality character appears to be affected by "intentional stance" (i.e., attributing mental states, agency, and "humanness"). However, this effect could also be explained by individual differences or perceptual effects resulting from the design of these studies. The current study used a new design that measured centro-parietal P250, P350, and N170 event-related potentials (ERPs) in 20 healthy adults while they initiated gaze-related joint attention with a virtual character (“Alan”) in two conditions. In one condition, they were told that Alan was controlled by a human; in the other, they were told that he was controlled by a computer. When participants believed Alan was human, his congruent gaze shifts, which resulted in joint attention, generated significantly larger P250 ERPs than his incongruent gaze shifts. In contrast, his incongruent gaze shifts triggered significantly larger increases in P350 ERPs than his congruent gaze shifts. These findings support previous studies suggesting that intentional stance affects the neural processing of social cues from a virtual character. The outcomes also suggest the use of the P250 and P350 ERPs as objective indices of social engagement during the design of socially approachable robots and virtual agents.

Links between adolescent bullying and neural activation to viewing social exclusion Abstract Neuroscientists who have studied bullying have primarily focused on the psychopathology of diagnosable offenders or the resulting symptomatology of victimization. Less attention has been given to theories that suggest that bullying may be an interpersonal strategy. In an exploratory study, we recruited a sample of adolescents (N = 24) who engaged in high rates of delinquent behavior and collected self-report ratings of bullying behaviors. During an fMRI scan, adolescents observed instances of social exclusion and social inclusion. The adolescents’ self-reported bullying was associated with greater ventral striatum, amygdala, medial prefrontal cortex, and insula activation when viewing social exclusion > social inclusion. Activation in these regions is commonly associated with reward-learning, salience monitoring, and motivational processes, suggesting that bullies show altered processing of interpersonal cues and social dynamic experiences in their environment. Our findings highlight the need for developmental neuroscientists to further explore the role of social motivation in processing socio-affective information, with a particular focus on goal-directed antisocial behavior.

Overnight sleep benefits both neutral and negative direct associative and relational memory Abstract Strong evidence suggests that sleep plays a role in memory consolidation, which involves both stabilizing memory into long-term storage as well as integrating new information into existing stores. The current study investigated consolidation, across a day of wakefulness or night of sleep, of emotional and neutral directly learned visual paired associates (A-B/B-C pairs) as well as formation of memory for relational pairs formed via overlapping learned components (A-C pairs). Participants learned 40 negative and 40 neutral face-object pairs followed by a baseline test in session 1 either in the morning or evening. They then spent a 12-hour retention period during which participants either went about their normal day or spent the night in the sleep lab. During session 2, participants completed a surprise test to assess their memory for relational pairs (A-C) as well as memory for direct associates (A-B/B-C). As hypothesized, the results demonstrated that a 12-hour retention period predominantly spent asleep, compared to awake, benefited memory for both relational and direct associative memory. However, contrary to the hypothesis that emotional salience would promote preferential consolidation, sleep appeared to benefit both negative and neutral information similarly for direct associative and relational memories, suggesting that sleep may interact with other factors affecting encoding (e.g., depth of encoding) to benefit direct and relational associative memory. As one of the few studies examining the role of nocturnal sleep and emotion on both direct and relational associative memory, our findings suggest key insights into how overnight sleep consolidates these different forms of memory.

Neural tracking of subjective value under riskand ambiguity in adolescence Abstract Although many neuroimaging studies on adolescent risk taking have focused on brain activation during outcome valuation, less attention has been paid to the neural correlates of choice valuation. Subjective choice valuation may be particularly influenced by whether a choice presents risk (known probabilities) or ambiguity (unknown probabilities), which has rarely been studied in developmental samples. Therefore, we examined the neural tracking of subjective value during choice under risk and ambiguity in a large sample of adolescents (N = 188, 12–22 years). Specifically, we investigated which brain regions tracked subjective value coding under risk and ambiguity. A model-based approach to estimate individuals’ risk and ambiguity attitudes showed prominent variation in individuals’ aversions to risk and ambiguity. Furthermore, participants subjectively experienced the ambiguous options as being riskier than the risky options. Subjective value tracking under risk was coded by activation in ventral striatum and superior parietal cortex. Subjective value tracking under ambiguity was coded by dorsolateral prefrontal cortex (PFC) and superior temporal gyrus activation. Finally, overlapping activation in the dorsomedial PFC was observed for subjective value under both conditions. Overall, this is the first study to chart brain activation patterns for subjective choice valuation under risk and ambiguity in an adolescent sample, which shows that the building blocks for risk and ambiguity processing are already present in early adolescence. Finally, we highlight the potential of combining behavioral modeling with fMRI for investigating choice valuation in adolescence, which may ultimately aid in understanding who takes risks and why.

High sensation seeking is associated with behavioral and neural insensitivity to increased negative outcomes during decision-making under uncertainty Abstract High-level sensation seeking (HSS) has been linked to a range of risky and unhealthy behavior; however, the neural mechanisms underlying such linkage remain unclear. In the present study, we used event-related potential (ERP) with a Balloon Analogue Risk Task (BART) to investigate how sensation seeking modulates brain responses to sequential decision-making with variable reward/loss outcome magnitudes. Behavior data showed that decision-making behavior was significantly affected by the large compared with the small magnitude of monetary outcome in the BART for individuals with low-level sensation seeking (LSS), but not for individuals with HSS. Similarly, HSS individuals displayed no changes in feedback-related negativity (FRN) in response to negative outcomes from decision-making with large or small reward/loss magnitudes, whereas LSS individuals showed greater FRN in response to decision-making with large loss magnitude than those with small loss magnitude. In addition, FRN amplitude differences correlated with decision-making behavior changes from small to large outcome magnitude for LSS, while such correlation was not observed for HSS. These findings suggest that a high-level of sensation seeking is associated with behavioral and neural insensitivity to increased negative outcome during decision-making under uncertainty, which may lead to greater risky behavior in these individuals when facing potential loss.

Cerebral responses to self-initiated action during social interactions Abstract Social interaction involves self-initiated actions that engage subjective awareness of one's own volition. Individuals with social communication needs or social anxiety find it particularly difficult to initiate social interactions. However, extant studies have not specifically addressed how perceived exclusion may influence self-initiated actions during social interaction. As a first step to address this question, we scanned 24 healthy adults participating in a Cyberball game with two fictive players. By contrasting events of observing, receiving, and initiating ball toss during a scenario of fair game (FG) and of exclusion (EX), we examined the neural correlates of self-initiated action during social interactions. Behaviorally, participants were faster in catching but slower in tossing the ball in EX compared with FG, suggesting a burden during self-initiated actions during social exclusion. Tossing versus receiving (or observing) engaged higher activity during EX than FG in the precuneus and angular gyrus, regions that have been widely implicated in theory of mind processing and social emotions. Across subjects these cortical activities correlated positively with the difference between EX and FG in the percentage of trials where participants tossed the ball back to the same player (r = 0.69, p < 0.001). Together, the results suggested that, in healthy adults, social exclusion encumbered and engaged higher posterior cortical activations during self-initiated actions. The findings may facilitate future research of neural markers of social behavioral disorders.

The importance of agency in human reward processing Abstract Converging evidence suggests that reinforcement learning (RL) signals exist within the human brain and that they play a role in the modification of behaviour. According to RL theory, prediction errors are used to update values associated with actions and/or predictive cues, thus facilitate decision-making. For example, the reward positivity—a feedback-sensitive component of the event-related brain potential (ERP)—is thought to index an RL prediction error. An unresolved question, however, is whether or not action is required to elicit a reward positivity. Reinforcement learning theory would predict that the reward positivity should diminish or disappear in the absence of action, but evidence for this claim is conflicting. To investigate the impact of cue, choice, and action on the amplitude of the reward positivity, we altered a two-armed bandit task by systematically removing these factors. The reward positivity was greatly reduced or absent in the altered versions of the task. This result highlights the key role of agency in producing learning signals, such as the reward positivity.

The neural representation of mental beliefs held by two agents Abstract Neuroimaging research has demonstrated that mentalizing about false beliefs held by other people recruits the temporo-parietal junction (TPJ). However, earlier work was limited to a single agent that held a false belief. We investigated the effect of two agents that held similar or mixed false and/or true beliefs. Participants saw animated stories with two smurfs holding true or false beliefs (Story phase). At the end of each trial, they were requested to take the perspective of the self or one of the smurfs (Question phase). We predicted that an increasing number of smurfs holding a false belief would increase activation in the TPJ when participants have to report the belief of the smurf, because the incongruent belief should have a stronger influence if it is held by two compared with one agent. This prediction was confirmed as activation in the TPJ during the Story and Question phase increased when more smurfs held a false belief. Taking the perspective of the self led to stronger activation of the TPJ in the two conditions that involved a true belief and weakest activation in the condition of two false beliefs. These data suggest that activation in TPJ depends on the perspective participants take, and that the number of agents holding a false belief influences activation in the TPJ only when taking the agent’s perspective.

Age-related differences in neural activation and functional connectivity during the processing of vocal prosody in adolescence Abstract The ability to recognize others’ emotions based on vocal emotional prosody follows a protracted developmental trajectory during adolescence. However, little is known about the neural mechanisms supporting this maturation. The current study investigated age-related differences in neural activation during a vocal emotion recognition (ER) task. Listeners aged 8 to 19 years old completed the vocal ER task while undergoing functional magnetic resonance imaging. The task of categorizing vocal emotional prosody elicited activation primarily in temporal and frontal areas. Age was associated with a) greater activation in regions in the superior, middle, and inferior frontal gyri, b) greater functional connectivity between the left precentral and inferior frontal gyri and regions in the bilateral insula and temporo-parietal junction, and c) greater fractional anisotropy in the superior longitudinal fasciculus, which connects frontal areas to posterior temporo-parietal regions. Many of these age-related differences in brain activation and connectivity were associated with better performance on the ER task. Increased activation in, and connectivity between, areas typically involved in language processing and social cognition may facilitate the development of vocal ER skills in adolescence.

Neural correlates of aversive anticipation: An activation likelihood estimate meta-analysis across multiple sensory modalities Abstract Anticipation is a universal preparatory response essential to the survival of an organism. Although meta-analytic synthesis of the literature exists for the anticipation of reward, a neuroimaging-based meta-analysis of the neural mechanisms of aversive anticipation is lacking. To address this gap in the literature, we ran an activation likelihood estimate (ALE) meta-analysis of 63 fMRI studies of aversive anticipation across multiple sensory modalities. Results of the ALE meta-analysis provide evidence for a core circuit involved in aversive anticipation, including the anterior insula, anterior cingulate cortex, mid-cingulate cortex, amygdala, thalamus, and caudate nucleus among other regions. Direct comparison of aversive anticipation studies using tactile versus visual stimuli identified additional regions involved in sensory specific aversive anticipation across these sensory modalities. Results from complementary multi-study voxel-wise and NeuroSynth analyses generally provide converging evidence for a core circuit involved in aversive anticipation. The multi-study voxel-wise analyses also implicate a more widespread preparatory response across sensory, motor, and cognitive control regions during more prolonged periods of aversive anticipation. The potential roles of these structures in anticipatory processing as well as avenues for future research are discussed.