The dorsomedial prefrontal cortex represents subjective value across effort-based and risky decision-making.

Decisions that require taking effort costs into account are ubiquitous in real life. The neural common currency theory hypothesizes that a particular neural network integrates different costs (e.g., risk) and rewards into a common scale to facilitate value comparison. Although there has been a surge of interest in the computational and neural basis of effort-related value integration, it is still under debate if effort-based decision-making relies on a domain-general valuation network as implicated in the neural common currency theory. Therefore, we comprehensively compared effort-based and risky decision-making using a combination of computational modeling, univariate and multivariate fMRI analyses, and data from two independent studies. We found that effort-based decision-making can be best described by a power discounting model that accounts for both the discounting rate and effort sensitivity. At the neural level, multivariate decoding analyses indicated that the neural patterns of the dorsomedial prefrontal cortex (dmPFC) represented subjective value across different decision-making tasks including either effort or risk costs, although univariate signals were more diverse. These findings suggest that multivariate dmPFC patterns play a critical role in computing subjective value in a task-independent manner and thus extend the scope of the neural common currency theory.

Gaze-dependent evidence accumulation predicts multi-alternative risky choice behaviour.

Choices are influenced by gaze allocation during deliberation, so that fixating an alternative longer leads to increased probability of choosing it. Gaze-dependent evidence accumulation provides a parsimonious account of choices, response times and gaze-behaviour in many simple decision scenarios. Here, we test whether this framework can also predict more complex context-dependent patterns of choice in a three-alternative risky choice task, where choices and eye movements were subject to attraction and compromise effects. Choices were best described by a gaze-dependent evidence accumulation model, where subjective values of alternatives are discounted while not fixated. Finally, we performed a systematic search over a large model space, allowing us to evaluate the relative contribution of different forms of gaze-dependence and additional mechanisms previously not considered by gaze-dependent accumulation models. Gaze-dependence remained the most important mechanism, but participants with strong attraction effects employed an additional similarity-dependent inhibition mechanism found in other models of multi-alternative multi-attribute choice.

Category-specific memory encoding in the medial temporal lobe and beyond: the role of reward.

The medial temporal lobe (MTL), including the hippocampus (HC), perirhinal cortex (PRC), and parahippocampal cortex (PHC), is central to memory formation. Reward enhances memory through interplay between the HC and substantia nigra/ventral tegmental area (SNVTA). While the SNVTA also innervates the MTL cortex and amygdala (AMY), their role in reward-enhanced memory is unclear. Prior research suggests category specificity in the MTL cortex, with the PRC and PHC processing object and scene memory, respectively. It is unknown, however, whether reward modulates category-specific memory processes. Furthermore, no study has demonstrated clear category specificity in the MTL for encoding processes contributing to subsequent recognition memory. To address these questions, we had 39 healthy volunteers (27 for all memory-based analyses) undergo functional magnetic resonance imaging while performing an incidental encoding task pairing objects or scenes with high or low reward, followed by a next-day recognition test. Behaviorally, high reward preferably enhanced object memory. Neural activity in the PRC and PHC reflected successful encoding of objects and scenes, respectively. Importantly, AMY encoding effects were selective for high-reward objects, with a similar pattern in the PRC. The SNVTA and HC showed no clear evidence of successful encoding. This behavioral and neural asymmetry may be conveyed through an anterior-temporal memory system, including the AMY and PRC, potentially in interplay with the ventromedial prefrontal cortex.

Effects of non-invasive brain stimulation on visual perspective taking: A meta-analytic study.

Visual perspective taking (VPT) is a critical ability required by complex social interaction. Non-invasive brain stimulation (NIBS) has been increasingly used to examine the causal relationship between brain activity and VPT, yet with heterogeneous results. In the current study, we conducted two meta-analyses to examine the effects of NIBS of the right temporoparietal junction (rTPJ) or dorsomedial prefrontal cortex (dmPFC) on VPT, respectively. We performed a comprehensive literature search to identify qualified studies and computed the standardized effect size (ES) for each combination of VPT level (Level-1: visibility judgment; Level-2: mental rotation) and perspective (self and other). Thirteen studies (rTPJ: 12 studies, 23 ESs; dmPFC: 4 studies, 14 ESs) were included in the meta-analyses. Random-effects models were used to generate the overall effects. Subgroup analyses for distinct VPT conditions were also performed. We found that rTPJ stimulation significantly improved participants' visibility judgment from the allocentric perspective, whereas its effects on other VPT conditions are negligible. Stimulation of dmPFC appeared to influence Level-1 performance from the egocentric perspective, although this finding was only based on a small number of studies. Notably, contrary to some theoretical models, we did not find strong evidence that these regions are involved in Level-2 VPT with a higher requirement of mental rotation. These findings not only advance our understanding of the causal roles of the rTPJ and dmPFC in VPT, but also reveal that the efficacy of NIBS on VPT is relatively small. Additionally, researchers should also be cautious about the potential publication bias and selective reporting.

Neural processing of vision and language in kindergarten is associated with prereading skills and predicts future literacy.

The main objective of this longitudinal study was to investigate the neural predictors of reading acquisition. For this purpose, we followed a sample of 54 children from the end of kindergarten to the end of second grade. Preliterate children were tested for visual symbol (checkerboards, houses, faces, written words) and auditory language processing (spoken words) using a passive functional magnetic resonance imaging paradigm. To examine brain-behavior relationships, we also tested cognitive-linguistic prereading skills at kindergarten age and reading performance of 48 of the same children 2 years later. Face-selective response in the bilateral fusiform gyrus was positively associated with rapid automatized naming (RAN). Response to both spoken and written words at preliterate age was negatively associated with RAN in the dorsal temporo-parietal language system. Longitudinally, neural response to faces in the ventral stream predicted future reading fluency. Here, stronger neural activity in inferior and middle temporal gyri at kindergarten age was associated with higher reading performance. Our results suggest that interindividual differences in the neural system of language and reading affect literacy acquisition and thus might serve as a marker for successful reading acquisition in preliterate children.

Emotion Regulation Modulates Dietary Decision-Making via Activity in the Prefrontal-Striatal Valuation System.

The consumption of indulgent, carbohydrate- and fat-rich foods is often used as a strategy to cope with negative affect because they provide immediate self-reward. Such dietary choices, however, can severely affect people's health. One countermeasure could be to improve one's emotion regulation ability. We used functional magnetic resonance imaging to examine the neural activity underlying the downregulation of incidental emotions and its effect on subsequent food choices. We investigated whether emotion regulation leads to healthier food choices and how emotion regulation interacts with the brain's valuation and decision-making circuitry. We found that 1) the downregulation of incidental negative emotions was associated with a subsequent selective increase in decisions for tasty but also for healthy foods, 2) food preferences were predicted by palatability but also by the current emotional state, and 3) emotion regulation modulated decision-related activation in the ventromedial prefrontal cortex and ventral striatum. These results indicate that emotional states are indeed important for food choice and that the process of emotion regulation might boost the subsequent processing of health attributes, possibly via neural reward circuits. In consequence, our findings suggest that increasing emotion regulation ability could effectively modulate food choices by stimulating an incidental upvaluation of health attributes.

Early volumetric changes of hippocampus and medial prefrontal cortex following medial temporal lobe resection.

Previous studies have shown that cognitive demands and physical exercise stimulate adult neurogenesis in the dentate gyrus and hippocampus. Recent observations in healthy humans and patients with mild cognitive impairment moreover suggest that training-induced increases in hippocampal volume may be associated with improved memory performance. The corresponding plasticity processes in hippocampal volume may occur on timescales of months to years. For patients with focal lesions in this region, previous functional imaging studies suggest that increased recruitment of the contralateral hippocampus and extratemporal regions may be an important part of the reorganization of episodic memory. However, it is currently unclear whether focal damage to the medial temporal lobe (MTL) induces gray matter (GM) volume changes in the intact contralateral hippocampus and in connected network regions on a shorter timescale. We therefore investigated whether unilateral resection of the MTL, including the hippocampus, induces measurable volumetric changes in the contralateral hippocampus and in the default mode network (DMN). We recruited 31 patients with unilateral left (N = 19) or right (N = 12) hippocampal sclerosis undergoing MTL resection for treatment of drug-resistant epilepsy. Structural MRI was acquired immediately before and 3 months after surgery. Longitudinal voxel-based morphometry (VBM) analysis revealed a significant increase of right hippocampal volume following resection of the left anterior MTL. Furthermore, this patient group showed GM volume increases in the DMN. These results demonstrate significant structural plasticity of the contralateral hippocampus, even in patients with a long-standing unilateral hippocampal dysfunction and structural reorganization processes extending to distant, but functionally connected brain regions.

Individual differences in envy experienced through perspective-taking involves functional connectivity of the superior frontal gyrus.

Envy is the painful or resentful awareness of another's advantage combined with a desire to possess that same advantage. Recent neuroscientific research has begun to shed light on the brain regions that process the experience of envy, including regions of the prefrontal cortex involved in emotional processing and social cognition. It is still unclear, however, which regions of the brain are functionally connected during the experience of envy. We recorded functional neuroimaging data while inducing simulated envy in participants, experienced through a perspective-taking hypothetical scenario task. In this task, participants took the perspective of a protagonist portrayed in a written description and compared themselves to either i) a self-similar/superior individual, ii) a self-dissimilar/superior individual, or iii) a self-dissimilar/average individual. During each comparison, participants also reported how much envy they experienced while taking the protagonists perspective. We demonstrate an inverse relationship in the connectivity of the left superior frontal gyrus to both the right supramarginal gyrus and the precuneus with respect to self-reported envy ratings across participants. In other words, we show that the greater the functional connectivity that the left superior frontal gyrus shares with the right supramarginal gyrus and precuneus, the less reported envy a participant experiences. Overall, our results are in line with previous research implicating the superior frontal gyrus in the reappraisal of negative emotions and extend these findings by showing this region is also involved in modulating the simulated experience of the social comparative, negative emotion of envy.

Neurobiological mechanisms of social cognition treatment in high-functioning adults with autism spectrum disorder.

BACKGROUND: The promise of precision medicine for autism spectrum disorder (ASD) hinges on developing neuroscience-informed individualized interventions. Taking an important step in this direction, we investigated neuroplasticity in response to an ecologically-valid, computer-based social-cognitive training (SCOTT). METHODS: In an active control group design, 48 adults with ASD were randomly assigned to a 3-month SCOTT or non-social computer training. Participants completed behavioral tasks, a functional and structural magnetic resonance imaging session before and after the training period. RESULTS: The SCOTT group showed social-cognitive improvements on close and distant generalization tasks. The improvements scaled with reductions in functional activity and increases in cortical thickness in prefrontal regions. CONCLUSION: In sum, we provide evidence for the sensitivity of neuroscientific methods to reflect training-induced social-cognitive improvements in adults with ASD. These results encourage the use of neuroimaging data to describe and quantify treatment-related changes more broadly.

Age Differences in the Neural Mechanisms of Intertemporal Choice Under Subjective Decision Conflict.

Older decision-makers may capitalize on their greater experiences in financial decisions and by this offset decline in cognitive abilities. However, this pattern of results should reverse in situations that place high demands on cognitive control functions. In this study, we investigated how decision conflict affects the neural mechanisms of intertemporal decision-making in younger and older adults. To individually adjust the level of decision conflict we determined the indifference point (IDP) in intertemporal decision-making for each participant. During functional magnetic resonance imaging, participants performed choice options close to their IDP (high conflict) or far away from the IDP (low conflict). In younger adults, decision conflict leads to reduced delay discounting and lower discount rates are associated with higher working memory (WM) capacity. In older adults, high decision conflict is associated with enhanced discounting, hypoactivation in the ventral striatum as well diminished ventral striatal representations of differences in subjective values. Taken together, our results show that under enhanced decision conflict, younger adults engage in a more reflective decision mode that reflects individual differences in WM capacity. In contrast, older adults get more present-oriented under high demands on cognitive control and this decision bias is associated with changes in striatal value signaling.

The computational basis of following advice in adolescents.

Advice taking helps one to quickly acquire knowledge and make decisions. This age-comparative study (in children [8- to 10-year-olds], adolescents [13- to 15-year-olds], and adults [18- to 22-year-olds]) investigated developmental differences in how advice, experience, and exploration influence learning. The results showed that adolescents were initially easily swayed to follow peer advice but also switched more rapidly to exploring alternatives like children. Whereas adults stayed with the advice over the task, adolescents put more weight on their own experience compared with adults. A social learning model showed that although social influence most strongly affects adolescents' initial expectations (i.e., their priors), adolescents showed higher exploration and discovered the other good option in the current task. Thus, our model resolved the apparently conflicting findings of adolescents being more and less sensitive to peer influence and provides novel insights into the dynamic interaction between social and individual learning.

Amphetamine modulates brain signal variability and working memory in younger and older adults.

Better-performing younger adults typically express greater brain signal variability relative to older, poorer performers. Mechanisms for age and performance-graded differences in brain dynamics have, however, not yet been uncovered. Given the age-related decline of the dopamine (DA) system in normal cognitive aging, DA neuromodulation is one plausible mechanism. Hence, agents that boost systemic DA [such as d-amphetamine (AMPH)] may help to restore deficient signal variability levels. Furthermore, despite the standard practice of counterbalancing drug session order (AMPH first vs. placebo first), it remains understudied how AMPH may interact with practice effects, possibly influencing whether DA up-regulation is functional. We examined the effects of AMPH on functional-MRI-based blood oxygen level-dependent (BOLD) signal variability (SD(BOLD)) in younger and older adults during a working memory task (letter n-back). Older adults expressed lower brain signal variability at placebo, but met or exceeded young adult SD(BOLD) levels in the presence of AMPH. Drug session order greatly moderated change-change relations between AMPH-driven SD(BOLD) and reaction time means (RT(mean)) and SDs (RT(SD)). Older adults who received AMPH in the first session tended to improve in RT(mean) and RT(SD) when SD(BOLD) was boosted on AMPH, whereas younger and older adults who received AMPH in the second session showed either a performance improvement when SD(BOLD) decreased (for RT(mean)) or no effect at all (for RT(SD)). The present findings support the hypothesis that age differences in brain signal variability reflect aging-induced changes in dopaminergic neuromodulation. The observed interactions among AMPH, age, and session order highlight the state- and practice-dependent neurochemical basis of human brain dynamics.

Memory integration in humans with hippocampal lesions.

Adaptive behavior frequently depends on inference from past experience. Recent studies suggest that the underlying process of integrating related memories may depend on interaction between hippocampus and prefrontal cortex. Here, we investigated how hippocampal damage affects memory integration. Subjects with mediotemporal lesions and healthy controls learned a set of overlapping AB- and BC-associations (object-face- and face-object pairs) and were then tested for memory of these associations ("direct" trials) and of inferential AC-associations ("indirect" trials). The experiment consisted of four encoding/retrieval cycles. In direct trials, performance of patients and controls was similar and stable across cycles. By contrast, in indirect trials, patients and controls showed distinct patterns of behavior. Whereas patients and controls initially showed only minor differences, controls increased performance across subsequent cycles, while patient performance decreased to chance level. Further analysis suggested that this deficit was not merely a consequence of impaired associative memory but rather resulted from an additional hippocampal contribution to memory integration. Our findings further suggest that contextual factors modulate this contribution. Patient deficits in more complex memory-guided behavior may depend on the flexible interaction of hippocampus-dependent and -independent mechanisms of memory integration.

Individual differences in social desirability are associated with white-matter microstructure of the external capsule.

Humans tend to present themselves in a positive light to gain social approval. This behavioral trait, termed social desirability, is important for various types of social success. Surprisingly, investigation into the neural underpinnings of social desirability has been limited and focused only on interindividual differences in dopamine receptor binding. These studies revealed reduced dopamine receptor binding in the striatum of individuals who are high in trait social desirability. Interestingly, high dopamine signaling has been associated with low white-matter integrity, irrespective of social desirability. Based on these findings, we hypothesized that a positive association exists between trait social desirability and the white-matter microstructure of the external capsule, which carries fibers to the striatum from the prefrontal cortex. To test this hypothesis, we collected diffusion tensor imaging data and examined the relationship between fractional anisotropy of the external capsule and participants' social desirability-our analysis revealed a positive association. As a second exploratory step, we examined the association between social desirability and white-matter microstructure throughout the whole brain. Our whole-brain analysis revealed associations within multiple major white-matter tracts, demonstrating that socially desirable behavior relies on connectivity between distributed brain regions.

Changes in Effective Connectivity Between Dorsal and Ventral Prefrontal Regions Moderate Emotion Regulation.

Reappraisal, the cognitive reevaluation of a potentially emotionally arousing event, has been proposed to be based upon top-down appraisal systems within the prefrontal cortex (PFC). It still remains unclear, however, how different prefrontal regions interact to control and regulate emotional responses. We used fMRI and dynamic causal modeling (DCM) to characterize the functional interrelationships among dorsal and ventral PFC regions involved in reappraisal. Specifically, we examined the effective connectivity between the inferior frontal gyrus (IFG), dorsolateral PFC (DLPFC), and other reappraisal-related regions (supplementary motor area, supramarginal gyrus) during the up- and downregulation of emotions in response to highly arousing extreme sports film clips. We found DLPFC to be the central node of the prefrontal emotion regulation network, strongly interconnected with the IFG. The DCM analysis further revealed excitatory changes of connection strength from the DLPFC to the IFG and strong inhibitory changes of connection strength between the IFG and DLPFC during reappraisal. These bidirectional changes in connectivity strength indicate a feedback mechanism by which the IFG may select one out of several possible goal-appropriate reappraisals held active in working memory (represented in the DLPFC) and inhibits the DLPFC once the selection process is completed.

Training-induced changes in subsequent-memory effects: No major differences among children, younger adults, and older adults.

The neural correlates of encoding mode, or the state of forming new memory episodes, have been found to change with age and mnemonic training. However, it is unclear whether neural correlates of encoding success, termed subsequent-memory (SM) effects, also differ by age and mnemonic skill. In a multi-session training study, we investigated whether SM effects are altered by instruction and training in a mnemonic skill, and whether such alterations differ among children, younger adults, and older adults. Before and after strategy training, fMRI data were collected while participants were memorizing word pairs. In all age groups, participants receiving training showed greater performance gains than control group participants. Analysis of task-relevant regions showed training-induced reductions in SM effects in left frontal regions. Reductions in SM effects largely generalized across age and primarily reflected greater training-induced activation increases for omissions than for remembered items, indicating that training resulted in more consistent use of the mnemonic strategy. The present results reveal no major age differences in SM effects in children, younger adults, and older adults.

Neural representation of emotion regulation goals.

The use of top-down cognitive control mechanisms to regulate emotional responses as circumstances change is critical for mental and physical health. Several theoretical models of emotion regulation have been postulated; it remains unclear, however, in which brain regions emotion regulation goals (e.g., the downregulation of fear) are represented. Here, we examined the neural mechanisms of regulating emotion using fMRI and identified brain regions representing reappraisal goals. Using a multimethodological analysis approach, combining standard activation-based and pattern-information analyses, we identified a distributed network of lateral frontal, temporal, and parietal regions implicated in reappraisal and within it, a core system that represents reappraisal goals in an abstract, stimulus-independent fashion. Within this core system, the neural pattern-separability in a subset of regions including the left inferior frontal gyrus, middle temporal gyrus, and inferior parietal lobe was related to the success in emotion regulation. Those brain regions might link the prefrontal control regions with the subcortical affective regions. Given the strong association of this subsystem with inner speech functions and semantic memory, we conclude that those cognitive mechanisms may be used for orchestrating emotion regulation. Hum Brain Mapp 37:600-620, 2016. © 2015 Wiley Periodicals, Inc.

The role of the amygdala in naturalistic mentalising in typical development and in autism spectrum disorder.

BACKGROUND: The substantial discrepancy between mentalising in experimental settings v. real-life social interactions hinders the understanding of the neural basis of real-life social cognition and of social impairments in psychiatric disorders. AIMS: To determine the neural mechanisms underlying naturalistic mentalising in individuals with and without autism spectrum disorder. METHOD: We investigated mentalising with a new video-based functional magnetic resonance imaging task in 20 individuals with autism spectrum disorder and 22 matched healthy controls. RESULTS: Naturalistic mentalising implicated regions of the traditional mentalising network (medial prefrontal cortex, temporoparietal junction), and additionally the insula and amygdala. Moreover, amygdala activity predicted implicit mentalising performance on an independent behavioural task. Compared with controls, the autism spectrum disorder group did not show differences in neural activity within classical mentalising regions. They did, however, show reduced amygdala activity and a reduced correlation between amygdala activity and mentalising accuracy on the behavioural task, compared with controls. CONCLUSIONS: These findings highlight the crucial role of the amygdala in making accurate implicit mental state inferences in typical development and in the social cognitive impairments of individuals with autism spectrum disorder.

Electrophysiological correlates of observational learning in children.

Observational learning is an important mechanism for cognitive and social development. However, the neurophysiological mechanisms underlying observational learning in children are not well understood. In this study, we used a probabilistic reward-based observational learning paradigm to compare behavioral and electrophysiological markers of individual and observational reinforcement learning in 8- to 10-year-old children. Specifically, we manipulated the amount of observable information as well as children's similarity in age to the observed person (same-aged child vs. adult) to examine the effects of similarity in age on the integration of observed information in children. We show that the feedback-related negativity (FRN) during individual reinforcement learning reflects the valence of outcomes of own actions. Furthermore, we found that the feedback-related negativity during observational reinforcement learning (oFRN) showed a similar distinction between outcome valences of observed actions. This suggests that the oFRN can serve as a measure of observational learning in middle childhood. Moreover, during observational learning children profited from the additional social information and imitated the choices of their own peers more than those of adults, indicating that children have a tendency to conform more with similar others (e.g. their own peers) compared to dissimilar others (adults). Taken together, our results show that children can benefit from integrating observable information and that oFRN may serve as a measure of observational learning in children.

Effects of empathic social responses on the emotions of the recipient.

Empathy is highly relevant for social behavior and can be verbally expressed by voicing sympathy and concern (emotional empathy) as well as by paraphrasing or stating that one can mentally reconstruct and understand another person's thoughts and feelings (cognitive empathy). In this study, we investigated the emotional effects and neural correlates of receiving empathic social responses after negative performance feedback and compared the effects of emotionally vs. cognitively empathic comments. 20 participants (10 male) underwent functional magnetic resonance imaging while receiving negative performance feedback for a cognitive task. Performance feedback was followed by verbal comments either expressing cognitive and emotional empathy or demonstrating a lack of empathy. Empathic comments in general led to less negative self-reported feelings and calmer breathing. At the neural level, empathic comments induced activity in regions associated with social cognition and emotion processing, specifically in right postcentral gyrus and left cerebellum (cognitively empathic comments), right precentral gyrus, the opercular part of left inferior frontal gyrus, and left middle temporal gyrus (emotionally empathic comments), as well as the orbital part of the left middle frontal gyrus and left superior parietal gyrus (emotionally empathic vs. unempathic comments). The study shows that cognitively and emotionally empathic comments appear to be processed in partially separable neural systems. Furthermore, confirming and expanding on another study on the same subject, the present results demonstrate that the social display of cognitive empathy exerts almost as positive effects on the recipient's feelings and emotions in states of distress as emotionally empathic response does. This can be relevant for professional settings in which strong negative emotions need to be de-escalated while maintaining professional impartiality, which may allow the display of cognitive but not emotional empathy.