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左侧背外侧前额叶在不平等情境下利他行为中的作用：一项tDCS研究*
The Role of the Left Dorsolateral Prefrontal Cortex in Altruistic Behavior Under Inequity Situations: A tDCS Study
利他是指个体损耗自身资源使他人获益的行为。利他行为塑造了人类进化的历史，广泛渗透于社会和政治生活。利他行为被高度重视，且容易受到不平等情境影响。目前对不平等情境下利他的神经机制知之甚少。为了阐明左侧背外侧前额叶皮层在不平等情境下利他中的作用，对左侧背外侧前额叶皮层施加经颅直流电刺激改变脑区活动，探究其对优势和劣势不平等情境下利他行为影响。具体而言，对左侧背外侧前额叶皮层施加1.5 mA（毫安）阳极刺激，然后让被试完成独裁者博弈任务以测量利他行为。实验结果显示，阳极tDCS刺激左侧背外侧前额叶皮层显著影响被试在优势不平等情境下的利他倾向，降低被试对自身损耗的关注；但是对劣势不平等情境下的利他倾向无显著影响。这一结果表明，左侧背外侧前额叶皮层在优势不平等情境下利他中发挥重要作用，促使个体超越个人利益，提高利他倾向。
Altruism, which is defined as behavior in which individuals deplete their own resources to benefit others, has played a pivotal role in human evolution and is deeply embedded in social and political life. Although highly valued, altruistic behavior is susceptible to influence by inequity situations. Specifically, individuals are more willing to act altruistically when they possess a higher initial endowment than their partners (an advantageous inequity situation) compared to when they have a lower initial endowment (a disadvantageous inequity situation). Neuroimaging studies suggest that advantageous and disadvantageous inequity engage distinct neurocognitive mechanisms. The left dorsolateral prefrontal cortex (lDLPFC), a brain region critical for executive function and impulse control, is hypothesized to play a role in altruistic decision-making. However, whether the lDLPFC differentially modulates altruism in advantageous versus disadvantageous inequity contexts remains unclear.
To address this question, we employed transcranial direct current stimulation (tDCS) to modulate the activity of lDLPFC and examined its effects on altruistic behavior under both advantageous and disadvantageous inequity conditions. Using a 2 × 2 mixed design, we manipulated stimulation condition (between-subjects factor: anodal vs. sham tDCS) and the inequity condition (within-subjects factor: advantageous vs. disadvantageous). A total of 70 healthy participants were recruited in the experiment (35 female, Mage = 20.79 ± 1.84 years). Participants were randomly assigned to receive either anodal tDCS (1.5 mA applied to the lDLPFC for 20 minutes) or sham tDCS (stimulation terminated after 30 seconds). Following the stimulation, participants completed a modified Dictator Game task to assess altruistic propensities. In this task, participants allocated monetary payoffs between themselves and anonymous partners across multiple trials, with no time constraints imposed.
In advantageous inequity trials, participants’ payoffs exceeded those of their partners (e.g., \$50 vs. \$20), while in disadvantageous inequity trials, participants’ payoffs were lower (e.g., \$20 vs. \$50). Altruistic choices involved participants sacrificing a portion of their own payoff (e.g.,\$4) to increase their partner’s payoff (e.g.,\$13). Trials were presented in separate, counterbalanced blocks. Using mixed-effects logistic regression (lme4 package in R), we analyzed the impact of lDLPFC stimulation on altruistic behavior.
The results demonstrated that anodal tDCS over the lDLPFC significantly enhanced altruistic tendencies in advantageous inequity conditions, reducing participants’ focus on personal losses. In advantageous inequity conditions, participants’ altruistic behavior decreased as the cost of altruism increased (β = -1.630, p <.001, Bonferroni-corrected) and increased as the benefit of altruism increased (β =.991, p<.001, Bonferroni-corrected). The interaction between tDCS stimulation and the cost of altruism was significant (β =.564, p <.01, Bonferroni-corrected). Anodal tDCS over the left DLPFC attenuated the impact of altruistic costs on altruism in advantageous inequity conditions, indicating that anodal stimulation of the left DLPFC reduced participants’ focus on self-interest in such contexts. In contrast, no significant effect was observed in disadvantageous inequity conditions. In disadvantageous inequity conditions, participants’ altruistic behavior also decreased as the cost of altruism increased (β = -2.403, p <.001, Bonferroni-corrected) and increased as the benefit of altruism increased (β =.871, p <.001, Bonferroni-corrected). However, the interaction between tDCS stimulation and the cost of altruism was not significant (β =.279, p >.05), nor was the interaction between tDCS stimulation and the benefit of altruism (β = -.218, p >.05). These findings highlight the critical role of the lDLPFC in promoting altruism under advantageous inequity, enabling individuals to transcend self-interest and prioritize the welfare of others.
In summary, this study investigated the impact of tDCS applied to the dorsolateral prefrontal cortex (DLPFC) on altruistic behavior under conditions of inequality. Anodal tDCS stimulation of the left DLPFC was found to reduce participants' sensitivity to altruistic costs in situations of advantageous inequality. However, it did not influence their sensitivity to altruistic costs in disadvantageous inequity condition. These findings suggest that the left DLPFC plays a specific role in promoting altruistic behavior in contexts of advantageous inequality, while its involvement in altruism appears to differ in situation of disadvantageous inequity.
不平等 / 利他 / 左侧背外侧前额叶皮层 / 经颅直流电刺激
inequity / altruism / left dorsolateral prefrontal cortex / transcranial direct current stimulation
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Although neuroscience has made remarkable progress in understanding the involvement of prefrontal cortex (PFC) in human memory, the necessity of dorsolateral PFC (dlPFC) for key competencies of working memory remains largely unexplored. We therefore studied human brain lesion patients to determine whether dlPFC is necessary for working memory function, administering subtests of the Wechsler Memory Scale, the Wechsler Adult Intelligence Scale, and the N-Back Task to three participant groups: dlPFC lesions (n=19), non-dlPFC lesions (n=152), and no brain lesions (n=54). DlPFC damage was associated with deficits in the manipulation of verbal and spatial knowledge, with left dlPFC necessary for manipulating information in working memory and right dlPFC critical for manipulating information in a broader range of reasoning contexts. Our findings elucidate the architecture of working memory, providing key neuropsychological evidence for the necessity of dlPFC in the manipulation of verbal and spatial knowledge.Copyright © 2012 Elsevier Ltd. All rights reserved.
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There is increasing support for the existence of 'social intelligence' [Humphrey (1984) Consciousness Regained], independent of general intelligence. Brothers et al. 1990) J. Cog. Neurosci., 4, 107-118] proposed a network of neural regions that comprise the 'social brain': the orbito-frontal cortex (OFC), superior temporal gyrus (STG) and amygdala. We tested Brothers' theory by examining both normal subjects as well as patients with high-functioning autism or Asperger syndrome (AS), who are well known to have deficits in social intelligence, and perhaps deficits in amygdala function [Bauman & Kemper (1988) J. Neuropath. Exp. Neurol., 47, 369]. We used a test of judging from the expressions of another person's eyes what that other person might be thinking or feeling. Using functional magnetic resonance imaging (fMRI) we confirmed Brothers' prediction that the STG and amygdala show increased activation when using social intelligence. Some areas of the prefrontal cortex also showed activation. In contrast, patients with autism or AS activated the fronto-temporal regions but not the amygdala when making mentalistic inferences from the eyes. These results provide support for the social brain theory of normal function, and the amygdala theory of autism.
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I study the relationships of resources and personality characteristics to charitable giving, postmortem organ donation, and blood donation in a nationwide sample of persons in households in the Netherlands. I find that specific personality characteristics are related to specific types of giving: agreeableness to blood donation, empathic concern to charitable giving, and prosocial value orientation to postmortem organ donation. I find that giving has a consistently stronger relation to human and social capital than to personality. Human capital increases giving; social capital increases giving only when it is approved by others. Effects of prosocial personality characteristics decline at higher levels of these characteristics. Effects of empathic concern, helpfulness, and social value orientations on generosity are mediated by verbal proficiency and church attendance.
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There have been many functional imaging studies of the brain basis of theory of mind (ToM) skills, but the findings are heterogeneous and implicate anatomical regions as far apart as orbitofrontal cortex and the inferior parietal lobe. The functional imaging studies are reviewed to determine whether the diverse findings are due to methodological factors. The studies are considered according to the paradigm employed (e.g., stories vs. cartoons and explicit vs. implicit ToM instructions), the mental state(s) investigated, and the language demands of the tasks. Methodological variability does not seem to account for the variation in findings, although this conclusion may partly reflect the relatively small number of studies. Alternatively, several distinct brain regions may be activated during ToM reasoning, forming an integrated functional "network." The imaging findings suggest that there are several "core" regions in the network-including parts of the prefrontal cortex and superior temporal sulcus-while several more "peripheral" regions may contribute to ToM reasoning in a manner contingent on relatively minor aspects of the ToM task.(c) 2008 Wiley-Liss, Inc.
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The decision to share resources is fundamental for cohesive societies. Humans can be motivated to give for many reasons. Some generosity incurs a definite cost, with no extrinsic reward to the act, but instead provides intrinsic satisfaction (labelled here as 'altruistic' giving). Other giving behaviours are done with the prospect of improving one's own situation via reciprocity, reputation, or public good (labelled here as 'strategic' giving). These contexts differ in the source, certainty, and timing of rewards as well as the inferences made about others' mental states. We executed a combined statistical map and coordinate-based fMRI meta-analysis of decisions to give (36 studies, 1150 participants). Methods included a novel approach for accommodating variable signal dropout between studies in meta-analysis. Results reveal consistent, cross-paradigm neural correlates of each decision type, commonalities, and informative differences. Relative to being selfish, altruistic and strategic giving activate overlapping reward networks. However, strategic decisions showed greater activity in striatal regions than altruistic choices. Altruistic giving, more than strategic, activated subgenual anterior cingulate cortex (sgACC). Ventromedial prefrontal cortex (vmPFC) is consistently involved during generous decisions and processing across a posterior to anterior axis differentiates the altruistic/strategic context. Posterior vmPFC was preferentially recruited during altruistic decisions. Regions of the 'social brain' showed distinct patterns of activity between choice types, reflecting the different use of theory of mind in the two contexts. We provide the consistent neural correlates of decisions to give, and show that many will depend on the source of incentives.Copyright © 2018 Elsevier Inc. All rights reserved.
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Why do we self-sacrifice to help others in distress? Two competing theories have emerged, one suggesting that prosocial behavior is primarily motivated by feelings of empathic other-oriented concern, the other that we help mainly because we are egoistically focused on reducing our own discomfort. Here we explore the relationship between costly altruism and these two sub-processes of empathy, specifically drawing on the caregiving model to test the theory that trait empathic concern (e.g. general tendency to have sympathy for another) and trait personal distress (e.g. predisposition to experiencing aversive arousal states) may differentially drive altruistic behavior. We find that trait empathic concern--and not trait personal distress--motivates costly altruism, and this relationship is supported by activity in the ventral tegmental area, caudate and subgenual anterior cingulate, key regions for promoting social attachment and caregiving. Together, this data helps identify the behavioral and neural mechanisms motivating costly altruism, while demonstrating that individual differences in empathic concern-related brain responses can predict real prosocial choice.Copyright © 2014 The Authors. Published by Elsevier Inc. All rights reserved.
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Disruption of function of left, but not right, lateral prefrontal cortex (LPFC) with low-frequency repetitive transcranial magnetic stimulation (rTMS) increased choices of immediate rewards over larger delayed rewards. rTMS did not change choices involving only delayed rewards or valuation judgments of immediate and delayed rewards, providing causal evidence for a neural lateral-prefrontal cortex-based self-control mechanism in intertemporal choice.
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Unfairness plays an important role in economic decision making. This fMRI study sought to investigate how the loss and the gain contexts could modulate behavioral and brain responses to unfairness by focusing on participants' rejection behaviors during an Ultimatum Game paradigm. Participants were scanned while they were playing the Ultimatum Game as responders in both loss and gain contexts, i.e. receiving ¥50 as gains and paying for ¥50 as losses. At the behavioral level, lower fairness ratings and higher rejection rates were revealed for unfair losses than unfair gains. At the neural level, left dorsolateral prefrontal cortex, bilateral anterior insula, anterior cingulate cortex/anterior middle cingulate cortex and bilateral dorsal striatum were associated with rejection (vs. acceptance) in the loss context, but not in the gain context. Together, our data indicated that participants may experience more unfairness in UG and stronger desire to sanction social norm violations in the loss context than in the gain context, inducing more fairness-related neutral activities when rejecting (vs. accepting) unfair losses than unfair gains. These findings shed light on the significance of context (i.e. loss or gain) in fairness-related social decision-making processes.Copyright © 2013 Elsevier Inc. All rights reserved.
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Fairness is a key concept in social interactions and is influenced by intentionality considerations. In this functional magnetic resonance imaging study, we investigated the neural correlates of fairness by focusing on responder behavior to unfair offers in an Ultimatum Game paradigm with conditions that differed in their intentionality constraints. Brain activity underlying rejection vs acceptance of unfair offers appeared highly dependent on intentionality. Rejection of unfair offers when the proposer had no-alternative as well as acceptance of offers when the proposer had a fair- or hyperfair-alternative was associated with activation in a network of regions including the insula and the dorsal medial prefrontal cortex. These activations were interpreted as neural responses to norm violations because they were mostly involved when behavior was inconsistent with socially accepted behavior patterns. Rejection of unfair offers in the no-alternative condition further resulted in activity in the anterior medial prefrontal cortex and the temporoparietal junction, which was interpreted in terms of higher moral mentalizing demands required in social decision-making when rejection could not be readily justified. Together, results highlight the significance of intentionality considerations in fairness-related social decision-making processes.
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Self-regulation is a core aspect of adaptive human behavior that has been studied, largely in parallel, through the lenses of social and personality psychology as well as cognitive psychology. Here, we argue for more communication between these disciplines and highlight recent research that speaks to their connection. We outline how basic facets of executive functioning (working memory operations, behavioral inhibition, and task-switching) may subserve successful self-regulation. We also argue that temporary reductions in executive functions underlie many of the situational risk factors identified in the social psychological research on self-regulation and review recent evidence that the training of executive functions holds significant potential for improving poor self-regulation in problem populations.Copyright © 2012 Elsevier Ltd. All rights reserved.
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Although altruistic behaviors, e.g., sacrificing one's own interests to alleviate others' suffering, are widely observed in human society, altruism varies greatly across individuals. Such individual differences in altruistic preference have been hypothesized to arise from both individuals' dispositional empathic concern for others' welfare and context-specific cost-benefit integration processes. However, how cost-benefit integration is implemented in the brain and how it is linked to empathy remain unclear. Here, we combine a novel paradigm with the model-based fMRI approach to examine the neurocomputational basis of altruistic behaviors. Thirty-seven adults (16 females) were tested. Modeling analyses suggest that individuals are likely to integrate their own monetary costs with nonlinearly transformed recipients' benefits. Neuroimaging results demonstrate the involvement of an extended common currency system during decision-making by showing that selfish and other-regarding motives were processed in dorsal anterior cingulate cortex and right inferior parietal lobule in a domain-general manner. Importantly, a functional dissociation of adjacent but different sub-regions within anterior insular cortex (aINS) was observed for different sub-processes underlying altruistic behaviors. While dorsal aINS and inferior frontal gyrus were involved in valuation of benefactors' costs, ventral aINS and middle insular cortex (vaINS/mINS), as empathy-related regions, reflected individual variations in valuating recipients' benefits. Multivariate analyses further suggest that both vaINS/mINS and dorsolateral prefrontal cortex reflect individual variations in general altruistic preferences which account for both dispositional empathy and context-specific other-regarding tendency. Together, these findings provide valuable insights into our understanding of psychological and neurobiological basis of altruistic behaviors.Altruistic behaviors play a crucial role in facilitating solidarity and development of human society, but the mechanisms of the cost-benefit integration underlying these behaviors are still unclear. Using model-based neuroimaging approaches, we clarify that people integrate personal costs and non-linearly transformed other's benefits during altruistic decision-making and the implementations of the integration processes are supported by an extended common currency neural network. Importantly, multivariate analyses reveal that both empathy- and cognitive control-related brain regions are involved in modulating individual variations of altruistic preference, which implicate complex psychological and computational processes. Our results provide a neurocomputational account of how people weigh between different attributes to make altruistic decisions and why altruistic preference varies to a great extent across individuals.Copyright © 2021 the authors.
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Humans restrain self-interest with moral and social values. They are the only species known to exhibit reciprocal fairness, which implies the punishment of other individuals' unfair behaviors, even if it hurts the punisher's economic self-interest. Reciprocal fairness has been demonstrated in the Ultimatum Game, where players often reject their bargaining partner's unfair offers. Despite progress in recent years, however, little is known about how the human brain limits the impact of selfish motives and implements fair behavior. Here we show that disruption of the right, but not the left, dorsolateral prefrontal cortex (DLPFC) by low-frequency repetitive transcranial magnetic stimulation substantially reduces subjects' willingness to reject their partners' intentionally unfair offers, which suggests that subjects are less able to resist the economic temptation to accept these offers. Importantly, however, subjects still judge such offers as very unfair, which indicates that the right DLPFC plays a key role in the implementation of fairness-related behaviors.
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A large body of literature is available on wound healing in humans. Nonetheless, a standardizedex vivowound model without disruption of the dermal compartment has not been put forward with compelling justification. Here, we present a novel wound model based on application of negative pressure and its effects for epidermal regeneration and immune cell behaviour. Importantly, the basement membrane remained intact after blister roof removal and keratinocytes were absent in the wounded area. Upon six days of culture, the wound was covered with one to three-cell thick K14+Ki67+keratinocyte layers, indicating that proliferation and migration were involved in wound closure. After eight to twelve days, a multi-layered epidermis was formed expressing epidermal differentiation markers (K10, filaggrin, DSG-1, CDSN). Investigations about immune cell-specific manners revealed more T cells in the blister roof epidermis compared to normal epidermis. We identified several cell populations in blister roof epidermis and suction blister fluid that are absent in normal epidermis which correlated with their decrease in the dermis, indicating a dermal efflux upon negative pressure. Together, our model recapitulates the main features of epithelial wound regeneration, and can be applied for testing wound healing therapies and investigating underlying mechanisms.
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Mounting evidence suggests that multiple mechanisms underlie working memory capacity. Using transcranial direct current stimulation (tDCS), the current study aimed to provide causal evidence for the neural dissociation of two mechanisms underlying visual working memory (WM) capacity, namely, the scope and control of attention. A change detection task with distractors was used, where a number of colored bars (i.e., two red bars, four red bars, or two red plus two blue bars) were presented on both sides (Experiment 1) or the center (Experiment 2) of the screen for 100ms, and participants were instructed to remember the red bars and to ignore the blue bars (in both Experiments), as well as to ignore the stimuli on the un-cued side (Experiment 1 only). In both experiments, participants finished three sessions of the task after 15min of 1.5mA anodal tDCS administered on the right prefrontal cortex (PFC), the right posterior parietal cortex (PPC), and the primary visual cortex (VC), respectively. The VC stimulation served as an active control condition. We found that compared to stimulation on the VC, stimulation on the right PPC specifically increased the visual WM capacity under the no-distractor condition (i.e., 4 red bars), whereas stimulation on the right PFC specifically increased the visual WM capacity under the distractor condition (i.e., 2 red bars plus 2 blue bars). These results suggest that the PPC and PFC are involved in the scope and control of attention, respectively. We further showed that compared to central presentation of the stimuli (Experiment 2), bilateral presentation of the stimuli (on both sides of the fixation in Experiment 1) led to an additional demand for attention control. Our results emphasize the dissociated roles of the frontal and parietal lobes in visual WM capacity, and provide a deeper understanding of the neural mechanisms of WM.Copyright © 2017 Elsevier Inc. All rights reserved.
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Sex differences in social cognitive ability are well established, including measures of Theory of Mind (ToM). The aim of this study was to investigate if sex mediates the effects of high-definition transcranial direct current stimulation (HD-tDCS) administered to a key hub of the social brain (i.e., the dorsomedial prefrontal cortex, dmPFC) on the Reading the Mind in the Eyes Test (RMET). Forty healthy young adults (18-35 years) were randomly allocated to receive either anodal or cathodal HD-tDCS in sham HD-tDCS controlled, double blind designs. In each of the two sessions, subjects completed the RMET. Anodal stimulation to the dmPFC increased accuracy on the RMET in females only. To assure regional specificity we performed a follow-up study stimulating the right temporoparietal junction and found no effect in either sex. The current study is the first to show improved performance on the RMET after tDCS to the dmPFC in females only. The polarity-specific effects and use of focal HD-tDCS provide evidence for sex-dependent differences in dmPFC function in relation to the RMET. Future studies using tDCS to study or improve ToM, need to consider sex.Copyright © 2017 IBRO. Published by Elsevier Ltd. All rights reserved.
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Theories of right temporoparietal junction (rTPJ) function in social cognition include self-other distinction, self-inhibition, or embodied rotation, whereas the dorsomedial prefrontal cortex (dmPFC) is associated with integrating social information. However, no study has provided causal evidence for dissociable roles of the rTPJ and dmPFC in social cognition. A total of 52 healthy young adults were stratified to receive either dmPFC or rTPJ anodal high-definition transcranial direct current stimulation (HD-tDCS) in a sham-controlled, double-blinded, repeated measures design. Self-other processing was assessed across implicit and explicit level 1 (line-of-sight) and level 2 (mental rotation) visual perspective taking (VPT) tasks, and self-other effects on memory. DmPFC stimulation selectively increased the influence of the allocentric perspective during egocentric perspective taking, indexed by an increase in congruency effect across explicit VPT tasks. Moreover, dmPFC stimulation removed the self-reference effect in episodic memory by increasing the recognition of other and decreasing the recognition of self-encoded words. Stimulation of the rTPJ resulted in improved inhibition of the egocentric-perspective during level 2 VPT only, indexed by a reduction of the congruency effect when taking the allocentric perspective. This research supports theories suggesting that the rTPJ facilitates embodied mental rotation of the self into an alternate perspective, whereas the dmPFC integrates social information relevant to self-directed processes.
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Dopamine dysfunction is associated with a wide range of neuropsychiatric disorders commonly treated pharmacologically or invasively. Recent studies provide evidence for a nonpharmacological and noninvasive alternative that allows similar manipulation of the dopaminergic system: transcranial direct current stimulation (tDCS). In rodents, tDCS has been shown to increase neural activity in subcortical parts of the dopaminergic system, and recent studies in humans provide evidence that tDCS over prefrontal regions induces striatal dopamine release and affects reward-related behavior. Based on these findings, we used fMRI in healthy human participants and measured the fractional amplitude of low-frequency fluctuations to assess spontaneous neural activity strength in regions of the mesostriatal dopamine system before and after tDCS over prefrontal regions (= 40, 22 females). In a second study, we examined the effect of a single dose of the dopamine precursor levodopa (l-DOPA) on mesostriatal fractional amplitude of low-frequency fluctuation values in male humans (= 22) and compared the results between both studies. We found that prefrontal tDCS and l-DOPA both enhance neural activity in core regions of the dopaminergic system and show similar subcortical activation patterns. We furthermore assessed the spatial similarity of whole-brain statistical parametric maps, indicating tDCS- and l-DOPA-induced activation, and >100 neuronal receptor gene expression maps based on transcriptional data from the Allen Institute for Brain Science. In line with a specific activation of the dopaminergic system, we found that both interventions predominantly activated regions with high expression levels of the dopamine receptors D2 and D3. Studies in animals and humans provide evidence that transcranial direct current stimulation (tDCS) allows a manipulation of the dopaminergic system. Based on these findings, we used fMRI to assess changes in spontaneous neural activity strength in the human dopaminergic system after prefrontal tDCS compared with the administration of the dopamine precursor and standard anti-Parkinson drug levodopa (l-DOPA). We found that prefrontal tDCS and l-DOPA both enhance neural activity in core regions of the dopaminergic system and show similar subcortical activation patterns. Using whole-brain transcriptional data of >100 neuronal receptor genes, we found that both interventions specifically activated regions with high expression levels of the dopamine receptors D2 and D3.Copyright © 2019 the authors.
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Human altruism shaped our evolutionary history and pervades social and political life. There are, however, enormous individual differences in altruism. Some people are almost completely selfish, while others display strong altruism, and the factors behind this heterogeneity are only poorly understood. We examine the neuroanatomical basis of these differences with voxel-based morphometry and show that gray matter (GM) volume in the right temporoparietal junction (TPJ) is strongly associated with both individuals' altruism and the individual-specific conditions under which this brain region is recruited during altruistic decision making. Thus, individual differences in GM volume in TPJ not only translate into individual differences in the general propensity to behave altruistically, but they also create a link between brain structure and brain function by indicating the conditions under which individuals are likely to recruit this region when they face a conflict between altruistic and selfish acts.Copyright © 2012 Elsevier Inc. All rights reserved.
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Adult functional magnetic resonance imaging (fMRI) literature suggests that a left-right hemispheric dissociation may exist between verbal and spatial working memory (WM), respectively. However, investigation of this type has been obscured by incomparable verbal and spatial WM tasks and/or visual inspection at arbitrary thresholds as means to assess lateralization. Furthermore, it is unclear whether this hemispheric lateralization is present during adolescence, a time in which WM skills are improving, and whether there is a developmental association with laterality of brain functioning. This study used comparable verbal and spatial WM n-back tasks during fMRI and a bootstrap analysis approach to calculate lateralization indices (LIs) across several thresholds to examine the potential of a left-right WM hemispheric dissociation in healthy adolescents. We found significant left hemispheric lateralization for verbal WM, most notably in the frontal and parietal lobes, as well as right hemisphere lateralization for spatial WM, seen in frontal and temporal cortices. Although no significant relationships were observed between LI and age or LI and performance, significant age-related patterns of brain activity were demonstrated during both verbal and spatial WM. Specifically, increased adolescent age was associated with less activity in the default mode brain network during verbal WM. In contrast, increased adolescent age was associated with greater activity in task-positive posterior parietal cortex during spatial working memory. Our findings highlight the importance of utilizing non-biased statistical methods and comparable tasks for determining patterns of functional lateralization. Our findings also suggest that, while a left-right hemispheric dissociation of verbal and spatial WM is apparent by early adolescence, age-related changes in functional activation during WM are also present.Copyright © 2013 Elsevier Inc. All rights reserved.
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| [33] |
Current research on prosocial behavior covers a broad and diverse range of phenomena. We argue that this large research literature can be best organized and understood from a multilevel perspective. We identify three levels of analysis of prosocial behavior: (a) the "meso" level--the study of helper-recipient dyads in the context of a specific situation; (b) the micro level--the study of the origins of prosocial tendencies and the sources of variation in these tendencies; and (c) the macro level--the study of prosocial actions that occur within the context of groups and large organizations. We present research at each level and discuss similarities and differences across levels. Finally, we consider ways in which theory and research at these three levels of analysis might be combined in future intra- and interdisciplinary research on prosocial behavior.
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All known human societies have maintained social order by enforcing compliance with social norms. The biological mechanisms underlying norm compliance are, however, hardly understood. We show that the right lateral prefrontal cortex (rLPFC) is involved in both voluntary and sanction-induced norm compliance. Both types of compliance could be changed by varying the neural excitability of this brain region with transcranial direct current stimulation, but they were affected in opposite ways, suggesting that the stimulated region plays a fundamentally different role in voluntary and sanction-based compliance. Brain stimulation had a particularly strong effect on compliance in the context of socially constituted sanctions, whereas it left beliefs about what the norm prescribes and about subjectively expected sanctions unaffected. Our findings suggest that rLPFC activity is a key biological prerequisite for an evolutionarily and socially important aspect of human behavior.
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Poor social judgment and decision-making abilities have often been attributed to people who have suffered injury to the ventromedial prefrontal cortex (VMPFC). However, few laboratory tests of decision-making have been conducted on these patients. The exception to this is the Iowa Gambling Task which has often, but not always, demonstrated differential performance between patients and controls. Results from patients with prefrontal cortex lesions on a novel test of decision-making are presented. Participants explored and chose from pairs of gambles that differed in their underlying distributions, primarily in the variance of their respective outcomes. In accordance with many findings from the behavioral decision-making literature, both young normal participants and older patient controls demonstrated a marked avoidance of risk and selected largely from secure, low variance gambles. In contrast, patients with ventromedial lesions were divided into two clear sub-groups. One group behaved similarly to normals, showing a risk-averse strategy. The other group displayed a distinctive risk-seeking behavior pattern, choosing predominantly from the high-variance, high-risk decks. This research demonstrates some of the advantages of using methods and theories from traditional decision-making research to study the behavior of patients, as well as the benefits of examining individual participants, and provides new insights into the nature of the decision-making deficit in patients with ventromedial prefrontal cortex lesions.
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Impaired cognitive empathy (ie, understanding the emotional experiences of others) is associated with poor social functioning in schizophrenia. However, it is unclear whether the neural activity underlying cognitive empathy relates to social functioning. This study examined the neural activation supporting cognitive empathy performance and whether empathy-related activation during correctly performed trials was associated with self-reported cognitive empathy and measures of social functioning. Thirty schizophrenia outpatients and 24 controls completed a cognitive empathy paradigm during functional magnetic resonance imaging. Neural activity corresponding to correct judgments about the expected emotional expression in a social interaction was compared in schizophrenia subjects relative to control subjects. Participants also completed a self-report measure of empathy and 2 social functioning measures (social competence and social attainment). Schizophrenia subjects demonstrated significantly lower accuracy in task performance and were characterized by hypoactivation in empathy-related frontal, temporal, and parietal regions as well as hyperactivation in occipital regions compared with control subjects during accurate cognitive empathy trials. A cluster with peak activation in the supplementary motor area (SMA) extending to the anterior midcingulate cortex (aMCC) correlated with social competence and social attainment in schizophrenia subjects but not controls. These results suggest that neural correlates of cognitive empathy may be promising targets for interventions aiming to improve social functioning and that brain activation in the SMA/aMCC region could be used as a biomarker for monitoring treatment response. © The Author 2014. Published by Oxford University Press on behalf of the Maryland Psychiatric Research Center. All rights reserved. For permissions, please email: journals.permissions@oup.com.
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| [37] |
Recent studies explored a network of brain regions involved in economic decision making. The present study focuses on two of those regions, each relevant for specific and distinct functions in economic decision making: the right temporoparietal junction (rTPJ) and the right dorsolateral prefrontal cortex (rDLPFC). In two experiments using transcranial direct current stimulation, we explored two proposed functions of these areas in bargaining situations using the ultimatum game (UG): understanding the others perspective and integration of fairness norms. Participants first took the role of the proposer and then the role of the responder. We showed that stimulation of the rTPJ only affected the proposer condition. Interestingly, inhibition of the rTPJ led to fairer offers, which strengthens the view that the role of the rTPJ in bargaining situations is to differentiate one's own from the other's perspective. Furthermore, we argue that the rDLPFC is most likely involved in suppressing self-interest when a person is confronted with a direct reward but does not play a role in long-term reward anticipation or integrating social fairness norms. We conclude that self-interest inhibition is shown only in responders, and that perspective taking seems to be a necessary specifically for proposers in the UG.© The Author(s) 2019. Published by Oxford University Press.
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| [38] |
Direct current stimulation is a neuromodulatory noninvasive brain stimulation tool, which was first introduced in animal and human experiments in the 1950s, and added to the standard arsenal of methods to alter brain physiology as well as psychological, motor, and behavioral processes and clinical symptoms in neurological and psychiatric diseases about 20 years ago. In contrast to other noninvasive brain stimulation tools, such as transcranial magnetic stimulation, it does not directly induce cerebral activity, but rather alters spontaneous brain activity and excitability by subthreshold modulation of neuronal membranes. Beyond acute effects on brain functions, specific protocols are suited to induce long-lasting alterations of cortical excitability and activity, which share features with long-term potentiation and depression. These neuroplastic processes are important foundations for various cognitive functions such as learning and memory formation and are pathologically altered in numerous neurological and psychiatric diseases. This explains the increasing interest to investigate transcranial direct current stimulation (tDCS) as a therapeutic tool. However, for tDCS to be used effectively, it is crucial to be informed about physiological mechanisms of action. These have been increasingly elucidated during the last years. This review gives an overview of the current knowledge available regarding physiological mechanisms of tDCS, spanning from acute regional effects, over neuroplastic effects to its impact on cerebral networks. Although knowledge about the physiological effects of tDCS is still not complete, this might help to guide applications on a scientifically sound foundation.
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| [39] |
Human social exchange is often characterized by conflicts of interest requiring strategic behavior for their resolution. To investigate the development of the cognitive and neural mechanisms underlying strategic behavior, we studied children's decisions while they played two types of economic exchange games with differing demands of strategic behavior. We show an increase of strategic behavior with age, which could not be explained by age-related changes in social preferences but instead by developmental differences in impulsivity and associated brain functions of the left dorsolateral prefrontal cortex (DLPFC). Furthermore, observed differences in cortical thickness of lDLPFC were predictive of differences in impulsivity and strategic behavior irrespective of age. We conclude that egoistic behavior in younger children is not caused by a lack of understanding right or wrong, but by the inability to implement behavioral control when tempted to act selfishly; a function relying on brain regions maturing only late in ontogeny.Copyright © 2012 Elsevier Inc. All rights reserved.
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| [40] |
Little is known about the positive emotional impact of fairness or the process of resolving conflict between fairness and financial interests. In past research, fairness has covaried with monetary payoff, such that the mental processes underlying preference for fairness and those underlying preference for greater monetary outcome could not be distinguished. We examined self-reported happiness and neural responses to fair and unfair offers while controlling for monetary payoff. Compared with unfair offers of equal monetary value, fair offers led to higher happiness ratings and activation in several reward regions of the brain. Furthermore, the tendency to accept unfair proposals was associated with increased activity in right ventrolateral prefrontal cortex, a region involved in emotion regulation, and with decreased activity in the anterior insula, which has been implicated in negative affect. This work provides evidence that fairness is hedonically valued and that tolerating unfair treatment for material gain involves a pattern of activation resembling suppression of negative affect.
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| [41] |
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| [42] |
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| [43] |
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| [44] |
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| [45] |
People seek the best in every aspect of life. However, little is known about the neurobiological mechanisms supporting this process of maximization. In this study, maximization tendencies were increased by using high-definition transcranial direct current stimulation (HD-tDCS) over the right dorsolateral prefrontal cortex (DLPFC). Participants (n= 64) received 2 mA anodal 4 × 1 HD-tDCS or sham stimulation over the right DLPFC in two sessions and subsequently completed an N-back working memory task and a maximization scale (MS). We observed that maximization tendency scores increased during anodal stimulation. Furthermore, the results indicate that this increase in maximization tendency was driven by motivational changes. On the MS, alternative search subscale scores were significantly increased, demonstrating an increase in motivation to evaluate more alternatives; however, the results did not indicate that the increase in maximization tendencies was due to working memory improvement. These results demonstrated that maximization tendencies can be strengthened through noninvasive interventions and that the right DLPFC has a causal relationship with maximization tendencies.
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| [46] |
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| [47] |
Recent empirical research has shed new light on the perennial question of human altruism. A number of recent studies suggest that from very early in ontogeny young children have a biological predisposition to help others achieve their goals, to share resources with others and to inform others of things helpfully. Humans' nearest primate relatives, such as chimpanzees, engage in some but not all of these behaviors: they help others instrumentally, but they are not so inclined to share resources altruistically and they do not inform others of things helpfully. The evolutionary roots of human altruism thus appear to be much more complex than previously supposed.
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| [48] |
Human's uncertain decision-making involves choices of risk (with known probabilities) and ambiguity (with unknown probabilities). For risky and ambiguous decision-making processes, neural differences are rarely exhibited. To clarify the causal role of the right dorsolateral prefrontal cortex (DLPFC) in uncertain situations, we used transcranial direct current stimulation (tDCS) to demonstrate the involvement of the right DLPFC in decisions involving risk and ambiguity. Participants received either anodal or cathodal tDCS at 1.5 mA or sham stimulation over the right DLPFC and subsequently undertook tasks of risk and ambiguity. The results revealed that a preference for ambiguity could be measurably increased in individuals through anodal stimulation, but no significant differences were observed in the preferences for risky choices among groups. These findings suggest that different neural mechanisms underlie risky and ambiguous decisions because the right DLPFC primarily affects ambiguous behavior.Copyright © 2020 Elsevier B.V. All rights reserved.
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| [49] |
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| [50] |
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| [51] |
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| [52] |
Antisocial behavior and prosocial behavior in the condition of inequality have long been observed in daily life. Understanding the neurological mechanisms and brain regions associated with antisocial and prosocial behavior and the development of new interventions are important for reducing violence and inequality. Fortunately, neurocognitive research and brain imaging research have found a correlation between antisocial or prosocial behavior and the prefrontal cortex. Recent brain stimulation research adopting transcranial direct current stimulation or transcranial magnetic stimulation has shown a causal relationship between brain regions and behaviors, but the findings are mixed. In the present study, we aimed to study whether stimulation of the DLPFC can change participants’ antisocial and prosocial behavior in the condition of inequality. We integrated antisocial and prosocial behavior in a unified paradigm. Based on this paradigm, we discussed costly and cost-free antisocial and prosocial behavior. In addition, we also measured participants’ disadvantageous and advantageous inequality aversion. The current study revealed an asymmetric effect of bilateral stimulation over the DLPFC on costly antisocial behavior, while such an effect of antisocial behavior without cost and prosocial behavior with and without cost were not observed. Moreover, costly antisocial behavior exhibited by men increased after receiving right anodal/left cathodal stimulation and decreased after receiving right cathodal anodal/left anodal stimulation compared with the behavior observed under sham stimulation. However, subjects’ inequality aversion was not influenced by tDCS.
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