Neural correlates of evaluations of non-binary social feedback: An EEG study.

PURPOSE: In complex and diverse social circumstances, decision making is affected by social feedback. Although previous studies have examined the electrophysiological correlates of social feedback with a binary valence, those related to non-binary feedback, or the magnitude of social feedback, remain unclear. This study investigated the electrophysiological correlates of non-binary social feedback and subsequent action selection processing. METHODS: Participants were asked to complete a Gabor patch direction judgment task in which they were required to make judgments before and after receiving social feedback. They were informed that the feedback stimuli represented the degree to which other participants made the same choice. RESULTS & CONCLUSION: The results revealed that feedback that was highly concordant with the participant's judgments elicited greater P300 activity, which was associated with the fulfillment of expectations regarding social reward. Moreover, moderately concordant feedback induced stronger theta band power, which may indicate monitoring of subjective conflict. Temporal changes in theta power during feedback phase may also relate to adjustments in prediction error. Additionally, when an initial judgment was maintained following social feedback, we observed a stronger increase in beta power, indicating an association with post-social-feedback action processing.

Neural substrates on the judgment of sequential benefits and losses.

People need to adapt to situations where they experience sequential benefits (or losses) to ensure survival. This study investigated the neural substrates involved in judgments of sequential benefits and losses. A total of 29 healthy volunteers participated in this study, in which they were asked to participate in a game of purchasing stocks while a magnetic resonance imaging scan was performed. This game had two main types of trials: (1) participants received four sequential financial benefits (or losses), and (2) participants received an equal amount of benefits (or losses) immediately. The results showed greater activation of the orbitofrontal cortex (OFC) when four benefits were received sequentially than when an equal amount of benefits was received immediately. This indicates that the OFC plays a crucial role in the process of mental integration of sequential benefits and interpretation of their valuations. It also showed greater activation of the dorsal striatum when four sequental losses were received than when an equal amount of losses was received immediately. However, it cannot be concluded that activation of the dorsal striatum reflects the differences between sequential and immediate losses, because previous studies have not confirmed this perspective. Therefore, it is necessary to clarify the function of the striatum in processing these losses.

The Role of the Left Inferior Frontal Gyrus in Introspection during Verbal Communication.

Conversation enables the sharing of our subjective experiences through verbalizing introspected thoughts and feelings. The mentalizing network represents introspection, and successful conversation is characterized by alignment through imitation mediated by the mirror neuron system (MNS). Therefore, we hypothesized that the interaction between the mentalizing network and MNS mediates the conversational exchange of introspection. To test this, we performed hyperscanning functional magnetic resonance imaging during structured real-time conversations between 19 pairs of healthy participants. The participants first evaluated their preference for and familiarity with a presented object and then disclosed it. The control was the object feature identification task. When contrasted with the control, the preference/familiarity evaluation phase activated the dorso-medial prefrontal cortex, anterior cingulate cortex, precuneus, left hippocampus, right cerebellum, and orbital portion of the left inferior frontal gyrus (IFG), which represents introspection. The left IFG was activated when the two participants' statements of introspection were mismatched during the disclosure. Disclosing introspection enhanced the functional connectivity of the left IFG with the bilateral superior temporal gyrus and primary motor cortex, representing the auditory MNS. Thus, the mentalizing system and MNS are hierarchically linked in the left IFG during a conversation, allowing for the sharing of introspection of the self and others.

Dynamic causal model application on hierarchical human motor control estimation in visuomotor tasks.

Introduction: In brain function research, each brain region has been investigated independently, and how different parts of the brain work together has been examined using the correlations among them. However, the dynamics of how different brain regions interact with each other during time-varying tasks, such as voluntary motion tasks, are still not well-understood. Methods: To address this knowledge gap, we conducted functional magnetic resonance imaging (fMRI) using target tracking tasks with and without feedback. We identified the motor cortex, cerebellum, and visual cortex by using a general linear model during the tracking tasks. We then employed a dynamic causal model (DCM) and parametric empirical Bayes to quantitatively elucidate the interactions among the left motor cortex (ML), right cerebellum (CBR) and left visual cortex (VL), and their roles as higher and lower controllers in the hierarchical model. Results: We found that the tracking task with visual feedback strongly affected the modulation of connection strength in ML → CBR and ML↔VL. Moreover, we found that the modulation of VL → ML, ML → ML, and ML → CBR by the tracking task with visual feedback could explain individual differences in tracking performance and muscle activity, and we validated these findings by leave-one-out cross-validation. Discussion: We demonstrated the effectiveness of our approach for understanding the mechanisms underlying human motor control. Our proposed method may have important implications for the development of new technologies in personalized interventions and technologies, as it sheds light on how different brain regions interact and work together during a motor task.

Morphological invariant of the midsagittal deep brain anatomy between humans and African great apes.

OBJECTIVES: Efforts have been made to mathematically reconstruct the brain morphology from human fossil crania to clarify the evolutionary changes in the brain that are associated with the emergence of human cognitive ability. However, because conventional reconstruction methods are based solely on the endocranial shape, deep brain structures cannot be estimated with sufficient accuracy. Our study aims to investigate the possible morphological correspondence between the cranial and deep brain morphologies based on humans and African great apes, with the goal of a more precise reconstruction of fossil brains. MATERIALS AND METHODS: Midsagittal endocranial and deep brain landmarks were obtained from magnetic resonance images of humans and three species of African great apes. The average midsagittal endocranial profile of all four species was calculated after Procrustes registration. The spatial deformation function from each of the endocranial profiles to the average endocranial profile was defined, and the brain landmarks enclosed in the endocranium were transformed using the deformation function to evaluate the interspecific variabilities of the positions of the brain landmarks on the average endocranial profile. RESULTS: The interspecific differences in the shape-normalized positions of the corpus callosum, anterior commissure, thalamus center, and brainstem were approximately within the range of 2% of the human cranial length, indicating that the interspecific variabilities of the positions of these deep brain structures were relatively small among the four species. DISCUSSION: Such an invariant relationship of the deep brain structure and the endocranium that encloses the brain can potentially be utilized to reconstruct the brains of fossil hominins.

Effective Connectivity in the Human Brain for Sour Taste, Retronasal Smell, and Combined Flavour.

The anterior insula and rolandic operculum are key regions for flavour perception in the human brain; however, it is unclear how taste and congruent retronasal smell are perceived as flavours. The multisensory integration required for sour flavour perception has rarely been studied; therefore, we investigated the brain responses to taste and smell in the sour flavour-processing network in 35 young healthy adults. We aimed to characterise the brain response to three stimulations applied in the oral cavity-sour taste, retronasal smell of mango, and combined flavour of both-using functional magnetic resonance imaging. Effective connectivity of the flavour-processing network and modulatory effect from taste and smell were analysed. Flavour stimulation activated middle insula and olfactory tubercle (primary taste and olfactory cortices, respectively); anterior insula and rolandic operculum, which are associated with multisensory integration; and ventrolateral prefrontal cortex, a secondary cortex for flavour perception. Dynamic causal modelling demonstrated that neural taste and smell signals were integrated at anterior insula and rolandic operculum. These findings elucidated how neural signals triggered by sour taste and smell presented in liquid form interact in the brain, which may underpin the neurobiology of food appreciation. Our study thus demonstrated the integration and synergy of taste and smell.

The extrastriate body area is involved in reciprocal imitation of hand gestures, vocalizations, and facial expressions: A univariate and multivariate fMRI study.

The extrastriate body area (EBA) in the lateral occipito-temporal cortex has an important role in reciprocal interaction, as it detects congruence between self and other's hand actions. However, it is unclear whether the EBA can detect congruence regardless of the type of action. In the present study, we examined the neural substrate underlying congruence detection of three types of actions: hand gestures, vocalizations, and facial expressions. A univariate analysis revealed a congruency effect, especially for imitating action, for all three types of actions in the EBA. A multi-voxel pattern analysis classifier in the EBA was able to distinguish between initiating interaction from responding to interaction in all experiments. Correspondingly, the congruency effect in the EBA revealed by univariate analysis was stronger for responding to than for initiating interaction. These findings suggest that the EBA might contribute to detect congruence regardless of the body part used (i.e. face or hand) and the type of action (i.e. gestural or vocal). Moreover, initiating and responding to interaction might be processed differently within the EBA. This study highlights the role of the EBA in comparing between self and other's actions beyond hand actions.Running head: Function of EBA in reciprocal imitation.

Neural substrates of shared visual experiences: a hyperscanning fMRI study.

Sharing experience is a fundamental human social cognition. Since visual experience is a mental state directed toward the world, we hypothesized that sharing visual experience is mediated by joint attention (JA) for sharing directedness and mentalizing for mental state inferences. We conducted a hyperscanning functional magnetic resonance imaging with 44 healthy adult volunteers to test this hypothesis. We employed spoken-language-cued spatial and feature-based JA tasks. The initiator attracts the partner's attention by a verbal command to a spatial location or an object feature to which the responder directs their attention. Pair-specific inter-individual neural synchronization of task-specific activities was found in the right anterior insular cortex (AIC)-inferior frontal gyrus (IFG) complex, the core node of JA and salience network, and the right posterior superior temporal sulcus, which represents the shared categories of the target. The right AIC-IFG also showed inter-individual synchronization of the residual time-series data, along with the right temporoparietal junction and dorsomedial prefrontal cortex-the core components for mentalization and the default mode network (DMN). This background synchronization represents sharing the belief of sharing the situation. Thus, shared visual experiences are represented by coherent coordination between the DMN and salience network linked through the right AIC-IFG.

Aging Impacts the Overall Connectivity Strength of Regions Critical for Information Transfer Among Brain Networks.

Recent studies have demonstrated that connector hubs, regions considered critical for the flow of information across neural systems, are mostly involved in neurodegenerative dementia. Considering that aging can significantly affect the brain's intrinsic connectivity, identifying aging's impact on these regions' overall connection strength is important to differentiate changes associated with healthy aging from neurodegenerative disorders. Using resting state functional magnetic resonance imaging data from a carefully selected cohort of 175 healthy volunteers aging from 21 to 86 years old, we computed an intrinsic connectivity contrast (ICC) metric, which quantifies a region's overall connectivity strength, for whole brain, short-range, and long-range connections and examined age-related changes of this metric over the adult lifespan. We have identified a limited number of hub regions with ICC values that showed significant negative relationship with age. These include the medial precentral/midcingulate gyri and insula with both their short-range and long-range (and thus whole-brain) ICC values negatively associated with age, and the angular, middle frontal, and posterior cingulate gyri with their long-range ICC values mainly involved. Seed-based connectivity analyses further confirmed that these regions are connector hubs with connectivity profile that strongly overlapped with multiple large-scale brain networks. General cognitive performance was not associated with these hubs' ICC values. These findings suggest that even healthy aging could negatively impact the efficiency of regions critical for facilitating information transfer among different functional brain networks. The extent of the regions involved, however, was limited.

Changes in white matter fiber density and morphology across the adult lifespan: A cross-sectional fixel-based analysis.

White matter (WM) fiber bundles change dynamically with age. These changes could be driven by alterations in axonal diameter, axonal density, and myelin content. In this study, we applied a novel fixel-based analysis (FBA) framework to examine these changes throughout the adult lifespan. Using diffusion-weighted images from a cohort of 293 healthy volunteers (89 males/204 females) from ages 21 to 86 years old, we performed FBA to analyze age-related changes in microscopic fiber density (FD) and macroscopic fiber morphology (fiber cross section [FC]). Our results showed significant and widespread age-related alterations in FD and FC across the whole brain. Interestingly, some fiber bundles such as the anterior thalamic radiation, corpus callosum, and superior longitudinal fasciculus only showed significant negative relationship with age in FD values, but not in FC. On the other hand, some segments of the cerebello-thalamo-cortical pathway only showed significant negative relationship with age in FC, but not in FD. Analysis at the tract-level also showed that major fiber tract groups predominantly distributed in the frontal lobe (cingulum, forceps minor) exhibited greater vulnerability to the aging process than the others. Differences in FC and the combined measure of FD and cross section values observed between sexes were mostly driven by differences in brain sizes although male participants tended to exhibit steeper negative linear relationship with age in FD as compared to female participants. Overall, these findings provide further insights into the structural changes the brain's WM undergoes due to the aging process.

Reorganization of brain networks and its association with general cognitive performance over the adult lifespan.

Healthy aging is associated with structural and functional changes in the brain even in individuals who are free of neurodegenerative diseases. Using resting state functional magnetic resonance imaging data from a carefully selected cohort of participants, we examined cross sectional changes in the functional organization of several large-scale brain networks over the adult lifespan and its potential association with general cognitive performance. Converging results from multiple analyses at the voxel, node, and network levels showed widespread reorganization of functional brain networks with increasing age. Specifically, the primary processing (visual and sensorimotor) and visuospatial (dorsal attention) networks showed diminished network integrity, while the so-called core neurocognitive (executive control, salience, and default mode) and basal ganglia networks exhibited relatively preserved between-network connections. The visuospatial and precuneus networks also showed significantly more widespread increased connectivity with other networks. Graph analysis suggested that this reorganization progressed towards a more integrated network topology. General cognitive performance, assessed by Addenbrooke's Cognitive Examination-Revised total score, was positively correlated with between-network connectivity among the core neurocognitive and basal ganglia networks and the integrity of the primary processing and visuospatial networks. Mediation analyses further indicated that the observed association between aging and relative decline in cognitive performance could be mediated by changes in relevant functional connectivity measures. Overall, these findings provided further evidence supporting widespread age-related brain network reorganization and its potential association with general cognitive performance during healthy aging.

Ventromedial prefrontal cortex contributes to performance success by controlling reward-driven arousal representation in amygdala.

When preparing for a challenging task, potential rewards can cause physiological arousal that may impair performance. In this case, it is important to control reward-driven arousal while preparing for task execution. We recently examined neural representations of physiological arousal and potential reward magnitude during preparation, and found that performance failure was explained by relatively increased reward representation in the left caudate nucleus and arousal representation in the right amygdala (Watanabe, et al., 2019). Here we examine how prefrontal cortex influences the amygdala and caudate to control reward-driven arousal. Ventromedial prefrontal cortex (VMPFC) exhibited activity that was negatively correlated with trial-wise physiological arousal change, which identified this region as a potential modulator of amygdala and caudate. Next we tested the VMPFC - amygdala - caudate effective network using dynamic causal modeling (Friston et al., 2003). Post-hoc Bayesian model selection (Friston and Penny, 2011) identified a model that best fit data, in which amygdala activation was suppressively controlled by the VMPFC only in success trials. Furthermore, fixed connectivity strength from VMPFC to amygdala explained individual task performance. These findings highlight the role of effective connectivity from VMPFC to amygdala in order to control arousal during preparation for successful performance.

Short Version Dental Anxiety Inventory Score May Predict the Response in the Insular Cortex to Stimuli Mimicking Dental Treatment.

Background: Dental anxiety is a common reason for avoiding dental visits and is associated with poor dental status. The short version of Dental Anxiety Inventory (SDAxI) is an easy-to-use, multi-faceted questionnaire for assessing the level of trait dental anxiety. However, there was no neurophysiological data indicating if its score associates with the state anxiety when an individual is under real/mock dental environment. We hypothesized that there exists such an association. Materials and Methods: Twenty systemic healthy adults with dental attendance experience and self-claimed free of dental phobia were recruited in this cross-sectional study, with their dental anxiety level assessed by SDAxI. Functional magnetic resonance imaging recorded their brain signals in response to audiovisual footages resembling dental scaler or turbine in action. After the brain imaging, they gave fear ratings to the footages in visual analog scale (VAS). Results: Participants' SDAxI scores positively correlated with their responses in the insular cortex (r2 = 0.388-0.445, P < 0.005). Their SDAxI scores also positively correlated with their fear ratings of the footages (r 2 = 0.415-0.555, P < 0.005). Discussion: Our findings indicated a possible neurobiological relevance of SDAxI, and reinforced its neurobiological validity in assessing dental anxiety level of dental attenders.

Role of the right anterior insular cortex in joint attention-related identification with a partner.

Understanding others as intentional agents is critical in social interactions. We perceive others' intentions through identification, a categorical judgment that others should work like oneself. The most primitive form of understanding others' intentions is joint attention (JA). During JA, an initiator selects a shared object through gaze (initiative joint attention, IJA), and the responder follows the direction of the initiator's gaze (reactive joint attention, RJA). Therefore, both participants share the intention of object selection. However, the neural underpinning of shared intention through JA remains unknown. In this study, we hypothesized that JA is represented by inter-individual neural synchronization of the intention-related activity. Additionally, JA requires eye contact that activates the limbic mirror system; therefore, we hypothesized that this system is involved in shared attention through JA. To test these hypotheses, participants underwent hyperscanning fMRI while performing JA tasks. We found that IJA-related activation of the right anterior insular cortex of participants was positively correlated with RJA-related activation of homologous regions in their partners. This area was activated by volitional selection of the target during IJA. Therefore, identification with others by JA is likely accomplished by the shared intentionality of target selection represented by inter-individual synchronization of the right anterior insular cortex.

Reconstructing the Neanderthal brain using computational anatomy.

The present study attempted to reconstruct 3D brain shape of Neanderthals and early Homo sapiens based on computational neuroanatomy. We found that early Homo sapiens had relatively larger cerebellar hemispheres but a smaller occipital region in the cerebrum than Neanderthals long before the time that Neanderthals disappeared. Further, using behavioural and structural imaging data of living humans, the abilities such as cognitive flexibility, attention, the language processing, episodic and working memory capacity were positively correlated with size-adjusted cerebellar volume. As the cerebellar hemispheres are structured as a large array of uniform neural modules, a larger cerebellum may possess a larger capacity for cognitive information processing. Such a neuroanatomical difference in the cerebellum may have caused important differences in cognitive and social abilities between the two species and might have contributed to the replacement of Neanderthals by early Homo sapiens.

The role of the amygdala in incongruity resolution: the case of humor comprehension.

A dominant theory of humor comprehension suggests that people understand humor by first perceiving some incongruity in an expression and then resolving it. This is called "the incongruity-resolution theory." Experimental studies have investigated the neural basis of humor comprehension, and multiple neural substrates have been proposed; however, the specific substrate for incongruity resolution is still unknown. The reason may be that the resolution phase, despite its importance in humor comprehension, has not been successfully distinguished from the perception phase because both phases occur almost simultaneously. To reveal the substrate, we conducted a functional magnetic resonance study using 51 healthy participants. We used a humor-producing frame of "Given A, I'd say B, because C" so as to focus on the resolution phase independently by suspending humor processing just after the perception phase. This frame allowed us to separate the two phases. Based on our results, incongruity resolution evoked positive emotion and activated the left amygdala, which is known to be related to positive emotion. On the basis of these findings, we argue that the amygdala plays an important role in humor comprehension, considering its functional role in emotional evaluation, particularly the relevance detection for incoming stimuli.

Autistic Traits Affect P300 Response to Unexpected Events, regardless of Mental State Inferences.

Limited use of contextual information has been suggested as a way of understanding cognition in people with autism spectrum disorder (ASD). However, it has also been argued that individuals with ASD may have difficulties inferring others' mental states. Here, we examined how individuals with different levels of autistic traits respond to contextual deviations by measuring event-related potentials that reflect context usage. The Autism Spectrum Quotient (AQ) was used to quantify autistic-like traits in 28 university students, and 19 participants were defined as Low or High AQ groups. To additionally examine inferences about mental state, two belief conditions (with or without false belief) were included. Participants read short stories in which the final sentence included either an expected or an unexpected word and rated the word's degree of deviation from expectation. P300 waveform analysis revealed that unexpected words were associated with larger P300 waveforms for the Low AQ group, but smaller P300 responses in the High AQ group. Additionally, AQ social skill subscores were positively correlated with evaluation times in the Unexpected condition, whether a character's belief was false or not. This suggests that autistic traits can affect responses to unexpected events, possibly because of decreased availability of context information.

Neural evidence for the intrinsic value of action as motivation for behavior.

The intrinsic value of an action refers to the inherent sense that experiencing a behavior is enjoyable even if it has no explicit outcome. Previous research has suggested that a common valuation mechanism within the reward network may be responsible for processing the intrinsic value of achieving both the outcome and external rewards. However, how the intrinsic value of action is neurally represented remains unknown. We hypothesized that the intrinsic value of action is determined by an action-outcome contingency indicating the behavior is controllable and that the outcome of the action can be evaluated by this feedback. Consequently, the reward network should be activated, reflecting the generation of the intrinsic value of action. To test this hypothesis, we conducted a functional magnetic resonance imaging (fMRI) investigation of a stopwatch game in which the action-outcome contingency was manipulated. This experiment involved 36 healthy volunteers and four versions of a stopwatch game that manipulated controllability (the feeling that participants were controlling the stopwatch themselves) and outcome (a signal allowing participants to see the result of their action). A free-choice experiment was administered after the fMRI to explore preference levels for each game. The results showed that the stopwatch game with the action-outcome contingency evoked a greater degree of enjoyment because the participants chose this condition over those that lacked such a contingency. The ventral striatum and midbrain were activated only when action-outcome contingency was present. Thus, the intrinsic value of action was represented by an increase in ventral striatal and midbrain activation.

Age-dependent atypicalities in body- and face-sensitive activation of the EBA and FFA in individuals with ASD.

Individuals with autism spectrum disorder (ASD) have difficuly in recognizing bodies and faces, which are more pronounced in children than adults. If such difficulties originate from dysfunction of the extrastriate body area (EBA) and the fusiform face area (FFA), activation in these regions might be more atypical in children than in adults. We preformed functional magnetic resonance imaging while children and adults with ASD and age-matched typically developed (TD) individuals observed face, body, car, and scene. To examine various aspects, we performed individual region of interest (ROI) analysis, as well as conventional random effect group analysis. At individual ROI analysis, we examined the ratio of participants showing a category-sensitive response, the size of regions, location and activation patterns among the four object categories. Adults with ASD showed no atypicalities in activation of the EBA and FFA, whereas children with ASD showed atypical activation in these regions. Specifically, a smaller percentage of children with ASD showed face-sensitive activation of the FFA than TD children. Moreover, the size of the EBA was smaller in children with ASD than in TD children. Our results revealed atypicalities in both the FFA and EBA in children with ASD but not in adults with ASD.

Being in a Romantic Relationship Is Associated with Reduced Gray Matter Density in Striatum and Increased Subjective Happiness.

Romantic relationship, a widespread feature of human society, is one of the most influential factors in daily life. Although stimuli related to romantic love or being in a romantic relationship commonly result in enhancement of activation or functional connectivity of the reward system, including the striatum, the structure underlying romantic relationship-related regions remain unclear. Because individual experiences can alter gray matter within the adult human brain, we hypothesized that romantic relationship is associated with structural differences in the striatum related to the positive subjective experience of being in a romantic relationship. Because intimate romantic relationships contribute to perceived subjective happiness, this subjective enhancement of happiness might be accompanied by the experience of positive events related to being in a romantic relationship. To test this hypothesis and elucidate the structure involved, we compared subjective happiness, an indirect measure of the existence of positive experiences caused by being in a romantic relationship, of participants with or without romantic partners (N = 68). Furthermore, we also conducted a voxel-based morphometry study of the effects of being in a romantic relationship (N = 113). Being in a romantic relationship was associated with greater subjective happiness and reduced gray matter density within the right dorsal striatum. These results suggest that being in a romantic relationship enhances perceived subjective happiness via positive experiences. Furthermore, the observed reduction in gray matter density in the right dorsal striatum may reflect an increase in saliency of social reward within a romantic relationship. Thus, being in a romantic relationship is associated with positive experiences and a reduction of gray matter density in the right dorsal striatum, representing a modulation of social reward.