Innocent intentions: A correlation between forgiveness for accidental harm and neural activity

Thinking in Patterns: Using Multi-Voxel pattern Analyses to Find Neural Correlates of Moral Judgment in Neurotypical and ASD Populations

In today’s society, people are very concerned about morality. News and events about morality are full of our daily life. People often make moral judgment and evaluation on the behavior of others through the network, media and self-media. In the process of moral judgment in which the general public participates, the behavior of high-class or low-class actors is the most concerned. Then, will people be influenced by the social class characteristics of the actors, leading to more lenient or stricter moral judgment? In order to discover and confirm the existence of social class effect in moral judgment, this article will discuss the influence of social class characteristics on the judges’ moral judgment from two aspects of moral behavior and immoral behavior through two studies respectively. Study 1 found that there was social class effect in moral behavior judgment. Through the overall moral behavior judgment materials, the social class characteristics of the actors would affect the severity of moral judgment on the moral behavior. Compared with the low-class actors, the participants had looser moral judgment on the moral behavior of the high-class actors. The class effect in the judgment of moral behavior is similar to the Matthew effect —“for unto every one that hath shall be given, and he shall have abundance: But from him that hath not shall be taken away even that which he hath”. Any individual who has achieved success and progress in a certain aspect (such as money, reputation, status, etc.) will have an advantage accumulation and more opportunities to achieve greater success and progress. Study 2 found the social class effect of immoral behavior judgment, the participants was more severe in moral judgment of immoral behavior of the high-class actors than the low-class actors; the participants thought the immoral behavior of the high-class actors were more immoral than that of the low-class actors. The social class effect in the judgment of immoral behavior is similar to tall poppy syndrome—“when any individual makes a great success in society, when he makes a mistake, he will be criticized spontaneously and collectively by the community”. To sum up, the social class characteristics of the actor would affect the judge’s moral judgment of his behavior. In terms of moral behavior, the judges were more lenient in their moral judgment towards the high-class actors than those of the low-class actors. As for the immoral behavior, judges were more severe in their moral judgment of the high-class actors than those of the low-class actors. On the basis of these results, we put forward the social class effect model of moral judgment. The judge’s moral judgment standard is influenced by the moral valence ( X -axis) and the actor’s social class ( Y -axis). The moral judgment standard is the product of the actor’s social class and the behavior’s morality in each quadrant. When the product is positive, the moral judgment standard becomes loose; when the product is negative, the moral judgment standard becomes strict. This model provides a more clear perspective for the research results. At the same time, this model can also be applied to similar research which the influence of social class characteristics and behavior moral valence on moral judgment standards.

Judgments about whether an action is morally right or wrong typically depend on our capacity to infer the actor’s beliefs and the outcomes of the action. Prior neuroimaging studies have found that mental state (e.g., beliefs, intentions) attribution for moral judgment involves a complex neural network that includes the temporoparietal junction (TPJ). However, neuroimaging studies cannot demonstrate a direct causal relationship between the activity of this brain region and mental state attribution for moral judgment. In the current study, we used transcranial direct current stimulation (tDCS) to transiently alter neural activity in the TPJ. The participants were randomly assigned to one of three stimulation treatments (right anodal/left cathodal tDCS, left anodal/right cathodal tDCS, or sham stimulation). Each participant was required to complete two similar tasks of moral judgment before receiving tDCS and after receiving tDCS. We studied whether tDCS to the TPJ altered mental state attribution for moral judgment. The results indicated that restraining the activity of the right temporoparietal junction (RTPJ) or the left the temporoparietal junction (LTPJ) decreased the role of beliefs in moral judgments and led to an increase in the dependance of the participants’ moral judgments on the action’s consequences. We also found that the participants exhibited reduced reaction times both in the cases of intentional harms and attempted harms after receiving right cathodal/left anodal tDCS to the TPJ. These findings inform and extend the current neural models of moral judgment and moral development in typically developing people and in individuals with neurodevelopmental disorders such as autism.

Moral judgment often requires making difficult tradeoffs (e.g., is it appropriate to torture to save the lives of innocents at risk?). Previous research suggests that both emotional appraisals and more deliberative utilitarian appraisals influence such judgments and that these appraisals often conflict. However, it is unclear how these different types of appraisals are represented in the brain, or how they are integrated into an overall moral judgment. We addressed these questions using an fMRI paradigm in which human subjects provide separate emotional and utilitarian appraisals for different potential actions, and then make difficult moral judgments constructed from combinations of these actions. We found that anterior cingulate, insula, and superior temporal gyrus correlated with emotional appraisals, whereas temporoparietal junction and dorsomedial prefrontal cortex correlated with utilitarian appraisals. Overall moral value judgments were represented in an anterior portion of the ventromedial prefrontal cortex. Critically, the pattern of responses and functional interactions between these three sets of regions are consistent with a model in which emotional and utilitarian appraisals are computed independently and in parallel, and passed to the ventromedial prefrontal cortex where they are integrated into an overall moral value judgment. Significance statement: Popular accounts of moral judgment often describe it as a battle for control between two systems, one intuitive and emotional, the other rational and utilitarian, engaged in winner-take-all inhibitory competition. Using a novel fMRI paradigm, we identified distinct neural signatures of emotional and utilitarian appraisals and used them to test different models of how they compete for the control of moral behavior. Importantly, we find little support for competitive inhibition accounts. Instead, moral judgments resembled the architecture of simple economic choices: distinct regions represented emotional and utilitarian appraisals independently and passed this information to the ventromedial prefrontal cortex for integration into an overall moral value signal.

Morality judgment usually refers to the evaluation of moral behavior`s ability to affect others` interests and welfare, while moral aesthetic judgment often implies the appraisal of moral behavior's capability to provide aesthetic pleasure. Both are based on the behavioral understanding. To our knowledge, no study has directly compared the brain activity of these two types of judgments. The present study recorded and analyzed brain activity involved in the morality and moral aesthetic judgments to reveal whether these two types of judgments differ in their neural underpinnings. Results reveled that morality judgment activated the frontal, parietal and occipital cortex previously reported for motor representations of behavior. Evaluation of goodness and badness showed similar patterns of activation in these brain regions. In contrast, moral aesthetic judgment elicited specific activations in the frontal, parietal and temporal cortex proved to be involved in the behavioral intentions and emotions. Evaluation of beauty and ugliness showed similar patterns of activation in these brain regions. Our findings indicate that morality judgment and moral aesthetic judgment recruit different cortical networks that might decode others' behaviors at different levels. These results contribute to further understanding of the essence of the relationship between morality judgment and aesthetic judgment.

Although individual differences in the application of moral principles, such as utilitarianism, have been documented, so too have powerful context effects-effects that raise doubts about the durability of people's moral principles. In this article, we examine the robustness of individual differences in moral judgment by examining them across time and across different decision contexts. In Study 1, consistency in utilitarian judgment of 122 adult participants was examined over two different survey sessions. In Studies 2A and 2B, large samples (Ns = 130 and 327, respectively) of adult participants made a series of 32 moral judgments across eight different contexts that are known to affect utilitarian endorsement. Contrary to some contemporary theorizing, our results reveal a strong degree of consistency in moral judgment. Across time and experimental manipulations of context, individuals maintained their relative standing on utilitarianism, and aggregated moral decisions reached levels of near-perfect consistency. Results support the view that on at least one dimension (utilitarianism), people's moral judgments are robustly consistent, with context effects tailoring the application of principles to the particulars of any given moral judgment.

Morality has been a hot topic in the psychological literature, and moral judgment, appraising process that enables individuals to differentiate good from evil and right from wrong according to moral principles, is a critical component of human morality. This review aims to introduce the application of multinomial models in moral judgment research. Several theories have been proposed to explain how moral judgment is made. Dual-process theory of moral judgment proposed that moral judgment is shaped by two moral principles (i.e., utilitarianism and deontology). The principle of utilitarianism emphasizes that the morality of an action is determined by its consequence whereas the principle of deontology states that the morality of an action depends on its consistency with moral norms. To measure people’s moral inclinations in moral judgment, researchers developed moral dilemma paradigm that pit the utilitarian principle against the deontological principle. Based on this moral dilemma paradigm, moral psychologists have revealed gender difference (i.e., men are more utilitarian) and individual differences (e.g., psychopathy individuals are more utilitarian) in moral judgment. However, the traditional moral dilemma paradigms, which did not distinguish the consequences and moral norms in moral dilemmas, can’t precisely measure an individual’s inclinations to utilitarian and deontological nor preclude potential confounders (e.g., people’s preference to act or not in moral dilemmas). To overcome these shortcomings of the traditional moral dilemma paradigms, researchers have recently developed a multinomial model (i.e., the CNI model) that allows researchers to quantify sensitivity to consequences ( C ), sensitivity to moral norms ( N ), and a general preference for inaction versus action irrespective of consequences and norms ( I ) in responses to moral dilemmas. This review aims to discuss the disadvantages of traditional moral dilemma paradigm and the advantages of the CNI model, and to summarize how CNI model-based findings contribute to our understanding of moral judgment. Specifically, recent work based on the CNI model of moral judgment has elaborated on how cognitive load and incidental happiness affect specific psychological processes of moral judgment, which challenges the dual-process theory of moral judgment. Moreover, the cognitive mechanisms underlying the individual differences (i.e., gender and psychopathy) in moral judgment have been clarified. We also point out future directions for moral judgment research. First, due to a small number of observation data in the model (i.e., data from only 24 moral dilemmas), the CNI model may not fit the data well at an individual-level and thus not be suitable for correlational designs. Future studies need to develop a larger set of moral dilemmas to increase the observations for the model and allow model fitting at an individual level. Second, it has been recognized that Chinese people showed stronger deontological inclination than British people, thus it would be interesting to uncover the underlying neurocognitive mechanism of culture differences in moral judgment. Third, several neuromodulators, such as oxytocin and serotonin, have been shown to affect moral judgment, future research is thus needed to clarify the influence of neuromodulators on specific cognitive processes underlying moral preferences. In conclusion, this review highlights the advantages of multinomial modeling and summarizes how the CNI model-based findings deepen our understanding of moral judgment, and the individual differences (i.e., gender and psychopathy) in moral judgment, which yields insights into human morality.

THE STUDY OF NORMAL AND ABNORMAL MORAL COGNITION AND BEHAVIOR Studies on normal volunteers and patients presenting with acquired or developmental antisocial behavioral disorders have begun to reveal with unprecedented clarity the neural substrates of moral cognition and behavior.1 This knowledge stems from 2 sources. The oldest is represented by observations on previously normal individuals who develop persistent antisocial behaviors as a result of acquired brain damage, a condition currently known as acquired sociopathy.2 Clinicoanatomic correlations in these cases have contributed to the development of models depicting the brain regions whose integrity is critical for moral behavior.3,4 Although most observations to date were done on adults, acquired sociopathy in children and adolescents has also shed light on the cerebral structures that are necessary for moral development.5 The other source of data is represented by the study of the morphological and functional abnormalities in the brains of individuals with antisocial behaviors and poor social adjustment beginning in childhood or adolescence, but without acquired brain damage.6 The personality and behavioral characteristics of these individuals were depicted by Cleckley,7 who called them psychopaths. Psychopaths began to be studied objectively thanks to the development of psychometrically sound instruments, which are known as the Psychopathy Checklists,8 and the development of neuroimaging gears that allow the in vivo scanning of the human brain with a high degree of spatial resolution.9 A PRACTICAL TAXONOMY OF ANTISOCIAL BEHAVIORAL DISORDERS The possibility afforded by magnetic resonance imaging (MRI) to study normal volunteers and patients represents a major advance in the field of moral cognition and behavior. The validation of MRI studies will increase the diagnostic accuracy in doubtful cases and its use in court and insurance decisions. However, the formulation of predictions on the nature and neural correlates of deviant social behaviors must necessarily depart from psychometrically sound diagnostic criteria. Table 1 presents the main conditions in which enduring and recurrent antisocial behaviors constitute the sole or predominant reason for referral, diagnosis, and counseling.TABLE 1: Taxonomy of Antisocial Behavioral DisordersNEURAL STRUCTURES INVOLVED IN SOCIAL BEHAVIORAL DISORDERS Charting of the brain structures whose damage gives rise to persistent sociopathy is an essential early step in the ascertainment of the brain circuits that underpin moral cognition and behavior. This analysis must be contrasted with those brain regions whose lesion does not predictably give rise to acquired sociopathy. From this double dissociation perspective,10 damage to the frontal and temporal lobes has been most consistently implicated in the production of acquired sociopathy, Cluster B personality disorders, and psychopathy. However, neither lobe is equipotential in this regard because acquired sociopathy results preferentially from lesions of certain sectors of the frontal lobes, the same being true of the temporal lobes and a set of related structures lying in a continuum along the rostral basal forebrain that includes the preoptic area/ventromedial hypothalamus and the subgenual/septal regions (Fig. 1). The relative contribution of cerebellar,12 thalamic, and pontine13 damage in the determination of sociopathy is still obscure and will not be further discussed in this article.FIGURE 1: Brain regions implicated in human moral cognition (adapted from Moll and Schulkin11 and Moll et al3). Cortical regions include the frontopolar cortex (FPC), medial and lateral ventral PFC, right anterior dorsolateral PFC, anterior temporal lobes, superior temporal sulcus (posterior STS) region. Subcortical structures include the extended amygdalae, the hypothalamus, the basal forebrain (especially the septal region), the basal ganglia, and the midbrain regions. We have postulated that integration across these corticolimbic structures gives rise to event-feature-emotion complexes, possibly by temporal binding mechanisms.Acquired Sociopathy and Corticosubcortical Circuits The most thoroughly investigated cases of acquired sociopathy are those with long-standing frontal lobe damage. (A brief anatomical account of the ventromedial and orbital frontal cortices). Despite its obvious relevance for a correct interpretation of clinical and experimental findings, there is as yet no consensus on what is exactly meant, in anatomical terms, by ventromedial prefrontal cortex [vmPFC] and to what extent it encompasses at least part of the orbitofrontal cortex [OFC; refer to, for example, the different partitions of the PFC by Bechara14 and Davidson et al15]. Part of the uncertainty may derive from differences in the criteria used by different authors; for example, cytoarchitectural16 versus clinicoanatomic.17 For the purposes of this article, the vmPFC encompasses the subgenual area and the lower medial frontal lobe up to the medial orbital sulcus, on the orbital surface.18 The orbital surface, which is covered by the OFC, is medially grooved by the olfactory sulcus, which delimits the gyrus rectus medially from the OFC proper, laterally. Two longitudinally oriented sulci [medial and lateral] delimit the medial orbital gyrus [between the olfactory and medial orbital sulci] and the lateral orbital gyrus [sitting laterally to the lateral orbital sulcus], which is an extension of the orbital division of the frontal operculum. The orbital sulci are joined together by the transverse orbital sulcus forming an H or K pattern.19 The gyri lying rostral and caudal to the transverse sulcus are, respectively, the anterior and posterior orbital gyri). In individual cases, the vmPFC lesion may variably extend to the frontopolar (rostromedial/rostrolateral), dorsolateral,1,20 and orbitofrontal cortices.21 One of our patients, previously adjusted and productive, developed a severe antisocial personality disorder after he fell from a train and sustained a circumscribed injury of the left dorsolateral PFC (Fig. 2). Acquired sociopathy has also been documented in patients with lesions of the OFC, with relative sparing of the vmPFC.22-24 As a group, these patients show a fairly uniform syndrome, which is characterized by poor social adjustment and an erratic pattern of interpersonal and occupational behavior. The stability of former alliances and commitments breaks down and is replaced by a flamboyant pattern of behavior marked by disregard toward relatives and peers, cheerfulness, impulsivity, and severe failure to keep occupation and level of productivity. A previously gifted student loses his talent and spends his days on simple manual tasks that must be constantly organized and checked by others.25 A prosperous controller in a home-building firm, married, with 2 children, and active in church affairs, is unable to stay on the job and gets involved in short-term partnerships, loses wife and children, and eventually moves in with his parents, unable to hold a productive occupation.2 These changes in personality are not necessarily followed by a decrease of premorbid intelligence or gross impairments in social knowledge, language, memory, perception, and praxic skills.26FIGURE 2: Acquired sociopathy in a case of traumatic left frontal lobe damage involving mainly the rostrolateral and dorsolateral PFC. The coronal cuts show that the connections of the lateral OFC may have been compromised by the extensive damage of the white matter underlying the anterior prefrontal lobe.The anterior temporal lobes, especially their polar division, are also often implicated in the social regulation of conduct, both in experimental animals27 and in human patients.28-30 Their importance in this respect is revealed by cases of acquired sociopathy in which most symptoms can be explained solely by damage to the temporal poles.31,32 Damage to the superior anterior sector of the temporal lobe, in particular, leads to impairments of social conceptual knowledge.33 In practice, most cases of acquired sociopathy display variable combinations of asymmetric vmPFC/OFC and anterior temporal lesions, which may extend ventrally into the anterior insula and the rostral basal forebrain and dorsolaterally into the anterior third of the first and second temporal gyri.34,35 In other cases, the lesions spare the cortex and are predominantly found in rostral prosencephalic structures, especially the amygdala,36 the ventromedial hypothalamus,37,38 and septum.39 This is true of surgical cases2 and of the major categories of diseases that may affect these structures, namely, vascular,40 traumatic,25 neoplastic,41 demyelinating,42 and degenerative.43 The cases with basal forebrain damage, however, are so complex in symptom heterogeneity and diversity that seldom do the personality changes appear in isolation.44 The anatomical picture that emerges from clinicoanatomic observations is consistent with the view that lesions that cause acquired sociopathy usually fall within the limits of the limbic circuits described by Livingston and Escobar.45 The basolateral limbic circuit is represented by the basolateral nuclei of the amygdala and the temporopolar and orbitofrontal cortices, which are interconnected by the uncinate fasciculus.46 This circuit receives highly patterned information through sequential pathways that ultimately project onto the isocortex of the frontal and temporal lobes allowing ongoing ideational events to be experienced as emotions and retained as memories.47 The ventral amygdalofugal pathway, a massive, loosely arranged system of fibers that connects the basolateral limbic circuit with the basal forebrain and the hypothalamus, provides information on the physiological state of the body.48 The basolateral structures are further connected with the septal nuclei via the diagonal band of Broca, thus establishing a bridge with the medial limbic circuit.49 The latter is anatomically represented by the gyrus fornicatus (the cortical component of the circuit of Papez), its subcortical connections, and the corticomedial nuclei of the amygdala, which project to the septal nuclei through the stria terminalis and its bed nucleus.50 (The gyrus fornicatus comprises the"… almost annular cortical region nearest the edge of the cerebral cortex [consisting] of the subcallosal gyrus, the cingulate gyrus, the retrosplenial area, the hippocampal gyrus, and the cuneus." The whole of this almost annular cortical locus is Broca's great limbic lobe.... It is called the gyrus fornicatus because of its arciform shape [fornix is Latin for arch].... Early anatomists saw that the vertical part of a massive, circumscribed bundle in the medial wall of the cerebral hemisphere had the look of a column supporting the gyrus fornicatus from under the corpus callosum. They called it the columna fornicis: the pillar supporting the arch).51 Depending on the site of injury within the medial limbic circuit, variable degrees of memory impairment,52 akinesia,53 and abulia54-56 result. Acquired sociopathy is mainly associated with lesions of the ventromedial frontal division of the medial circuit. It is likely that to be expressed as antisocial behaviors, which are mostly engendered by basal forebrain structures,35,38,57-59 parts of the medial limbic circuit must be intact, particularly those subserving motivation and drive.60 Developmental Psychopathy as a Disorder of the Moral Brain Recorded throughout history, psychopathy was the first personality disorder recognized in psychiatry.61 In the first half of the 19th century, it was characterized as a medical, in contrast to legal, anomaly.61 The departure was given by the observation of a type of individual who behaved in evil ways without being "mad" (ie, without acting out on delusions or hallucinations) and with an intact intellect (ie, who was able to tell right from wrong). This paradox is implied in the term moral insanity62 and remains one of the most fascinating challenges in the study of antisocial behavior.63 The severity of psychopathy is related to the additive effect of discrete personality traits, which are tapped by current assessment systems, such as the PCL8 and the DSM-IV.64 Individuals with psychopathy present with a personality disorder that begins in childhood or adolescence.65 Clinical investigations indicate that the dysmoral conduct of psychopaths results from functional impairments in neural circuits that subserve moral behavior.43 In contrast with acquired sociopathy, the search for the neuroanatomical substrates of psychopathy has met with little success until recently.66 Studies using in vivo volumetry by means of manual tracing of prespecified brain structures67 and studies using novel techniques, such as voxel-based morphometry (VBM), have shown that the personalities of psychopaths have discrete anatomic counterparts.68 One of the first studies on psychopaths using VBM found (a) a remarkable pattern of anatomic changes in a set of relatively discrete regions of the forebrain in noncriminal adult psychopaths living in the community and (b) a volumetric decrease of these regions against a background of normal total gray and white matter and cerebrospinal fluid volumes (Fig. 3). Remarkably, the gray matter decreases in the FPC, the OFC, and the caudal STS showed an inverse correlation with callousness (Fig. 4).68 Subsequent studies confirmed and extended these findings.69 Furthermore, Kruesi et al70 found a volumetric reduction of the temporal lobe in a sample of youths with a diagnosis of conduct disorder and without associated substance abuse, suggesting that the morphometric changes observed in adulthood may already be present in the first years of life. Another VBM study found decreased gray matter in the right anterior temporal lobe in a sample of 10 criminal psychopaths recruited from a forensic psychiatric facility.71 These results, albeit encouraging, are still preliminary because the sample sizes are too small and heterogeneous in what concerns age, sex, social background, and comorbidity. Notwithstanding these limitations, they lend support to the proposal that the anatomical correlates of psychopathy map onto neural circuits underlying moral cognition, emotion, and behavior.67,68,72FIGURE 3: Brain region showing decreased gray matter in psychopathy. Results from a VBM study demonstrating areas of gray matter reductions in community-dwelling psychopaths (assessed with the PCL, screening version [PCL-sv]), compared with normal control subjects (matched for age, sex, and education). The FPC and the STS showed the most prominent gray matter decreases. Adapted from de Oliveira-Souza et al.68FIGURE 4: Brain region showing correlations between gray matter decreases and psychopathy. The FPC (not shown in this picture), the medial OFC (mOFC), and the STS showed parametric relationships between the level of gray matter reductions and scores on the factor 1 (emotional callousness) of the PCL-sv. Results from a VBM study, adapted from de Oliveira-Souza et al.6The most striking finding of the functional neuroimaging literature has been the demonstration that the brain regions that are engaged by moral tasks in normal volunteers are roughly the same as those involved in developmental and acquired sociopathy. Such studies have further indicated that different regions involved in moral cognition in experimental settings play specific roles in the ultimate production of moral experience and behavior in more ecologic settings. For example, the septal-subgenual area is essential for the mediation of affiliative and altruistic behaviors73-75 (Fig. 5), and in economic behaviors that involve trusting others76 (Fig. 6), whereas the posterior STS integrates the social cues that allow the intuitive formulation of hypotheses about the intentions and feelings of others.77 These prosocial actions are closely related to the concomitant experience of moral emotions or sentiments78,79 and well-being80 that reinforce the enactment of prosocial decisions and actions. Affiliative/altruistic and theory of mind processes are major elements for the attribution of moral qualities to social events.72FIGURE 5: Brain regions involved in donation and opposition to charitable organizations. Brain regions showing increased activation in a functional MRI (fMRI) study of charitable donations to societal causes (eg, abortion, children's rights, nuclear energy, war, and euthanasia). A, Both pure monetary rewards (an experimental control condition) and decisions to donate (with or without personal financial costs) activated the mesolimbic reward system, including the ventral tegmental area (VTA) and the ventral and dorsal striatum (STR). B, The subgenual-septal area (SG), however, was selectively activated by decisions to donate, as compared with pure monetary rewards (both by costly and noncostly decisions [conjunction analysis]). The lateral OFC (latOFC) was activated by decisions to oppose charities. This activation extended to the anterior insula and to the inferior dorsolateral PFC and was present both for costly and noncostly decisions (conjunction analysis). The FPC and ventral medial PFC (mPFC), in green, were activated when volunteers made costly decisions, that is, when they voluntarily chose to sacrifice own monetary resources either to donate to a charity or to oppose to it (conjunction analysis). Areas activated by donations are coded in red, and areas activated by opposition are coded in blue. Adapted from Moll and Schulkin.11Figure 5 can be viewed online in color at www.topicsinmri.com.FIGURE 6: Brain responses for trust maintenance. In the group of participants who trusted their partners in the Trust Game even when the monetary incentive to defect was high, decisions to trust contrasted with decisions to reciprocate revealed a higher activation in the septal area. Pairs of participants who showed the highest trust-reciprocate history (frequency) in their decisions also showed the highest activation (parameter estimates) in the septal area.FUNCTIONAL IMAGING STUDIES ON MORAL COGNITION AND BEHAVIOR Functional MRI studies have substantiated the notion that the human brain is tuned to detect morality in trivial stimuli even in conditions of passive exposure without previous warning as to the moral nature of stimuli.79 In such conditions, the main activations fall in the vmPFC and the STS, as well as in sectors of the thalamus, the amygdala, and the midbrain (Fig. 7). When individuals are also asked to explicitly judge the moral content of the stimulus, for example, as either right or wrong, the frontopolar cortices and the right anterior temporal lobe become major activation foci.81-84 These studies also explored the influence of emotional engagement in moral judgments, providing direct evidence that moral sensitivity and judgment are decisively influenced by emotional experience and how these interactive "hot" and "cool" moral systems are organized in the brain. In fact, more recent fMRI studies are starting to explore the functional anatomy of specific moral sentiments regarding shared and distinct neural components (Fig. 8).78,85FIGURE 7: Brain regions activated by unpleasant moral and nonmoral pictures. Areas activated by both moral unpleasant and basic (nonmoral) unpleasant conditions relative to the neutral condition (assessed with conjunction analysis) are shown (upper row). Activations were observed in the amygdala and upper midbrain. Areas showing increased activation in the unpleasant moral condition as compared with the basic unpleasant condition included the anterior mOFC/FPC and the STS. Adapted from Moll et al.79FIGURE 8: Functional MRI results for specific moral sentiments. Activation of the FPC sector of the anterior PFC, and STS/temporoparietal junction (STS/TPJ) was observed when comparing guilt, compassion, and embarrassment to neutral condition (upper row). Analysis restricted to the empathic moral sentiments (guilt and compassion) revealed activation in limbic regions, including the ventral striatum/septal area, and in the FPC (lower row). Adapted from Moll et al.78Fewer studies have investigated the brains of psychopaths with fMRI. So far, the outstanding finding is that, in agreement with the fact that individuals with a diagnosis of psychopathy show major abnormalities in the emotional sphere, in comparison with a normal control group, psychopaths show distinct patterns of cerebral activation in response to emotionally charged pictures.86 In a prisoner's dilemma game, individuals with high scores on psychopathy defected more often and were less obliging after establishing mutual cooperation with a partner.87 In this dyadic social interactive setting, these individuals showed weaker OFC activation when they chose to cooperate and weaker activation within the dorsolateral PFC and anterior cingulate when they chose to defect. Some of these regions, especially the subgenual cortex, were highly active in normal individuals engaged in anonymous altruistic donations75 (Fig. 9), suggesting that psychopaths lack a basic attachment disposition toward other people, which may be one of their characteristics that we feel as "callousness."FIGURE 9: Brain responses related to the level of engagement in real-life altruistic behavior. Relationship between self-reported engagement in real-life voluntary activities and anterior mOFC activity to costly donations. Adapted from Moll et al.74CAVEATS AND CONCLUSIONS Neuroimaging studies on antisocial behavior must be interpreted with caution. This is due, in part, to the evolving neuroimaging methods and the conceptual issues they ensure and to the subtleties involved in the selection of patients and controls. Concerning the selection of subjects for research, acquired sociopathy and psychopathy must be differentiated from phenomenologically related conditions, such as bipolar disorder88 and attention deficit hyperactivity disorder.89 This distinction may not be as obvious as it may appear at first sight. For example, one of the paradigmatic cases with acquired sociopathy due to frontal lobe damage may, in fact, have had secondary mania.90 Comorbidity is another major confounder in such studies. Because antisocial behavioral disorders are often associated with alcohol and illicit drug abuse,91 the pattern of substance use must be carefully controlled before the neuroimaging findings may confidently be to the personality From a forensic is beginning to influence the means by which evidence is and interpreted in of personality in cases of brain injury may be against the pattern of the anatomical damage in individual Although the study of the neural of morality is still in its it will play an in the clinical assessment of patients with social behavioral

Moral judgments are increasingly being understood as showing context dependent variability. A growing literature has identified a range of specific contextual factors (e.g., emotions, intentions) that can influence moral judgments in predictable ways. Integrating these diverse influences into a unified approach to understanding moral judgments remains a challenge. Recent work by Railton (2017) attempted to address this with a causal-evaluative modelling approach to moral judgment. In support of this model Railton presents evidence from novel variations of classic trolley type dilemmas. We present results from a pre-registered pilot study that highlight a significant confound and demonstrate that it likely influenced Railton's results. Building on this, our registered report presents a replication-extension of Railton's study, using larger more diverse samples, and more rigorous methods and materials, specifically controlling for potential confounds. We found that participants' judgments in sacrificial dilemmas are influenced by both direct personal force, and by whether harm occurs as a means or as a side-effect of action. We also show the relationship between a range of individual difference variables and responses to sacrificial moral dilemmas. Our results provide novel insights into the factors that influence people's moral judgments, and contribute to ongoing theoretical debates in moral psychology.

In contemporary moral psychology, an often-heard claim is that knowing how we make moral judgments can help us make better moral judgments. Discussions about moral development and improvement are often framed in terms of the question of which mental processes have a better chance of leading to good moral judgments. However, few studies elaborate on the question of what makes a moral judgment a good moral judgment. This article examines what is needed to answer questions of moral improvement and development. It distinguishes 3 types of claims that are at stake: descriptive claims, metaethical claims, and normative claims. To find out what makes certain moral judgments better than others, one needs to have insight in the psychological processes and capacities underlying moral judgment formation. However, one also needs to address the question of what makes a moral judgment justified, and this in turn requires a view on the nature of moral goodness and on the question of what makes a judgment moral at all. The author discusses possible ways in which philosophical theories in the areas of metaethics and normative ethics can contribute to the answering of such questions. Also, she provides concrete suggestions for doing interdisciplinary research that is able to address those questions in moral psychology that have both normative and descriptive aspects.

Causal evidence of the roles of the prefrontal and occipital cortices in modulating the impact of color on moral judgement

Is the basis of criminality an act that causes harm, or an act undertaken with the belief that one will cause harm? The present study takes a cognitive neuroscience approach to investigating how information about an agent's beliefs and an action's consequences contribute to moral judgment. We build on prior developmental evidence showing that these factors contribute differentially to the young child's moral judgments coupled with neurobiological evidence suggesting a role for the right temporoparietal junction (RTPJ) in belief attribution. Participants read vignettes in a 2 x 2 design: protagonists produced either a negative or neutral outcome based on the belief that they were causing the negative outcome ("negative" belief) or the neutral outcome ("neutral" belief). The RTPJ showed significant activation above baseline for all four conditions but was modulated by an interaction between belief and outcome. Specifically, the RTPJ response was highest for cases of attempted harm, where protagonists were condemned for actions that they believed would cause harm to others, even though the harm did not occur. The results not only suggest a general role for belief attribution during moral judgment, but also add detail to our understanding of the interaction between these processes at both the neural and behavioral levels.

Article Figures and data Abstract Editor's evaluation Introduction Results Discussion Materials and methods Appendix 1 Data availability References Decision letter Author response Article and author information Metrics Abstract Altruism is critical for cooperation and productivity in human societies but is known to vary strongly across contexts and individuals. The origin of these differences is largely unknown, but may in principle reflect variations in different neurocognitive processes that temporally unfold during altruistic decision making (ranging from initial perceptual processing via value computations to final integrative choice mechanisms). Here, we elucidate the neural origins of individual and contextual differences in altruism by examining altruistic choices in different inequality contexts with computational modeling and electroencephalography (EEG). Our results show that across all contexts and individuals, wealth distribution choices recruit a similar late decision process evident in model-predicted evidence accumulation signals over parietal regions. Contextual and individual differences in behavior related instead to initial processing of stimulus-locked inequality-related value information in centroparietal and centrofrontal sensors, as well as to gamma-band synchronization of these value-related signals with parietal response-locked evidence-accumulation signals. Our findings suggest separable biological bases for individual and contextual differences in altruism that relate to differences in the initial processing of choice-relevant information. Editor's evaluation In this important paper, the authors use a sophisticated combination of computational modeling and EEG to show that variation in generosity produced by changes in context (i.e., disadvantageous vs. advantageous inequality) and variation due to individual differences in concern for others both seem to occur early, during the perceptual or valuation stage of a choice, rather than later on during choice comparison. However, these two sources of variation also appear to operate through distinct mechanisms during this stage of processing, which spurs further questions about the drivers of human prosocial behavior. This paper will be of considerable interest to researchers studying the psychological and neural basis of variation in prosocial behavior. https://doi.org/10.7554/eLife.80667.sa0 Decision letter Reviews on Sciety eLife's review process Introduction Altruism – incurring own costs to benefit others – is fundamental for cooperation and productivity in human societies (de Waal, 2008; Piliavin and Charng, 1990). It not only plays crucial roles in shaping socio-political ideology and welfare (e.g. via tax policies and charity; Bechtel et al., 2018; Offer and Pinker, 2017) but is also essential for collective management of challenging situations, such as political, financial, and public health crises. While altruism is thought to be a stable behavioral tendency shaped by the evolutionary advantages of the ability to cooperate, it is unclear why this tendency varies so strongly across individuals, contexts, and cultures (Bester and Güth, 1998; Hamilton, 1964a; Hamilton, 1964b; Lebow, 2018; Piliavin and Charng, 1990). Is altruism governed by a set of unitary neuro-cognitive mechanisms that are engaged to varying degrees in different situations or different people (Tricomi et al., 2010)? Or are there fundamentally different types of altruistic actions that are guided by different neuro-cognitive processes triggered by different contexts (Hein et al., 2016)? From a neurobiological perspective, both these possibilities appear plausible. On the one hand, all altruistic actions necessitate the ability to override self-interest, a parsimonious brain mechanism (Bester and Güth, 1998) that is thought to be facilitated more or less by different contexts and that could be expressed to different degrees in different people (Morishima et al., 2012; Trivers, 1971). On the other hand, empirical observations suggest that altruism varies with a range of factors such as others' previous actions (e.g. empathy-based vs. reciprocity-based altruism) or their perceived similarity (e.g. social distance; Hein et al., 2016; Vekaria et al., 2017). It is thus often argued that in different contexts or different individuals, superficially similar altruistic actions can be guided by distinct motives (such as personal moral norms, responsibility, or empathy), which may be controlled by fundamentally different types of neurocognitive mechanisms (Hein et al., 2016; Piliavin and Charng, 1990; Zaki and Mitchell, 2011). One specific context factor that is often discussed in this context is the inequality in resources held by the actor and the recipient of a possible distribution: People are more willing to share if they possess more than the recipient (advantageous inequality, ADV) than if they possess less (disadvantageous inequality; DIS) (Charness and Rabin, 2002; Fehr and Schmidt, 1999; Gao et al., 2018; Güroğlu et al., 2014; Morishima et al., 2012; Tricomi et al., 2010). Although this consistent effect has been formalized with the same utility model across contexts, this model needs to comprise two distinct latent parameters quantifying altruism in the two contexts (i.e. decision weights on others' payoffs that are specific for ADV and DIS), and these are often uncorrelated and differ strongly from each other (Gao et al., 2018; Morishima et al., 2012). These observations, together with distinct psychological accounts for the distribution behaviors in different contexts (i.e. 'guilt' in the advantageous and 'envy' in the disadvantageous inequality context), imply that altruistic choices in the two contexts may be driven by fundamentally different psychological processes (Fehr and Schmidt, 1999; Gao et al., 2018). Moreover, modeling studies often reveal that these altruism parameters vary strongly between different people for the same choice set (Fehr and Schmidt, 1999), and neuroimaging studies have shown that while distributional behavior in both contexts correlates with activity in brain regions commonly associated with motivation (e.g. the putamen and orbitofrontal cortex), either context also leads to activity in a set of distinct areas (the dorsolateral and dorsomedial prefrontal cortex in advantageous and the amygdala and anterior cingulate cortex in disadvantageous inequality; Gao et al., 2018; Yu et al., 2014). Finally, neuroanatomical research shows that only for advantageous inequality, individual variations in altruistic preferences relate to gray matter volume in the temporoparietal junction (TPJ; Morishima et al., 2012). While these behavioral modeling and neural findings suggest clear contextual and individual differences in altruism, it is still unclear what specific neurocognitive mechanisms these differences could arise from. Previous research on individual and contextual differences in altruism has largely used unitary computational models focusing exclusively on valuation (rather than attempting to separate distinct aspects of the choice process), and has used functional magnetic resonance imaging (fMRI) to identify spatial patterns of neural activity that correlate with valuation processes during wealth distribution behaviors in different contexts (Charness and Rabin, 2002; Fehr and Schmidt, 1999; Gao et al., 2018; Güroğlu et al., 2014; Morishima et al., 2012; Tricomi et al., 2010). For example, recent studies combined computational modeling with fMRI techniques to show that the value of altruistic choice can be modeled as the weighted sum of self- and other-interest, and that different attributes are integrated into an overall value signal correlating with BOLD activity in the ventromedial prefrontal cortex (vmPFC) (Crockett et al., 2017; Crockett et al., 2013; Hutcherson et al., 2015; Hare et al., 2010). However, since these studies neither formally examined the difference in altruistic choices between advantageous and disadvantageous inequality contexts, nor focused on separating different aspects of the decision mechanisms of altruistic choice, they can hardly address the question of whether and how different mechanisms are involved in different types of altruistic actions in different contexts (Crockett et al., 2013; Crockett et al., 2008; Gao et al., 2018). To systematically investigate this issue, it would be beneficial to harness the fact that altruistic decisions – like all choices – are guided by processes unfolding at different temporal stages (Seo and Lee, 2012; Shin et al., 2021; Tump et al., 2020). These processes include (1) initial perception of the objective information related to wealth distribution (e.g. payoff numbers) (Nieder, 2016; Pinel et al., 2004), (2) biased representations of the subjectively decision-relevant information attributes, such as attention-guided weighing of self- vs other-payoffs (Chen and Krajbich, 2018; Teoh et al., 2020), (3) integration of all these attributes and subjective preferences into decision values (Collins and Frank, 2018; Harris et al., 2018; Hutcherson et al., 2015), and (4) final decision processes that transform the decision values into motor responses (O'Connell et al., 2012; Polanía et al., 2014). Taking into account this temporal unfolding of the neurocognitive processes further refines the questions about the origins of differences in altruistic behavior: Do altruistic choices involve different sets of computations throughout all the temporally different processing stages (i.e., initial perceptual processing, valuation, final integrative choice mechanisms) in these different contexts and by different individuals (as suggested by Gao et al., 2018; Tricomi et al., 2010)? Or do individuals mainly perceive and attend to the choice-relevant information differently, before passing on this information to valuation and integrative decision mechanisms devoted to all types of altruistic choices (as suggested by Yu et al., 2014)? Answering these questions by means of modelling and neural recording techniques that allow a detailed focus on different temporal stages of altruistic choice processes could help us understand the biological origins of altruism, reveal why people differ strongly in altruistic behavior, and develop more efficient strategies to facilitate altruism. In the current study, we take such an approach. We combined a modified dictator game that independently varies payoffs to a player versus another person, and thereby also the inequality between both players, with electroencephalography (EEG) and sequential sampling modeling (SSM). This allowed us to identify electrophysiological markers of the initial perceptual processing and biased representation of the decision-relevant information (i.e. stimulus-locked event-related potentials [ERPs] related to the payoffs and the inequality context) as well as of the processes integrating this information into a decision variable used to guide choice (i.e. response-locked evidence accumulation [EA] signals; Balsdon et al., 2021; Hutcherson et al., 2015; Krajbich et al., 2015; Nassar et al., 2019). Thus, our approach differs from that of fMRI studies identifying brain areas involved in the valuation of own and others' payoffs (Fehr and Schmidt, 1999; Morishima et al., 2012; Sáez et al., 2015), since the temporal resolution of fMRI measures makes it difficult to separate response-locked decision-making processes from stimulus-locked perceptual processes and to examine the independent dynamics of these processes during distribution decisions. Our approach is also motivated by studies of nonsocial decisions showing that SSMs may provide a useful framework for investigating the temporal dynamics of the processes that integrate different choice attributes into the decision outcome (Harris et al., 2018; Maier et al., 2020). Many studies have shown that SSMs can identify these processes not just computationally, but also at the neural level, for both the perceptual (Brunton et al., 2013; Kelly and O'Connell, 2013; Ossmy et al., 2013) and value-based decision making (Glaze et al., 2015; Hutcherson et al., 2015; Pisauro et al., 2017; Polanía et al., 2014). The SSM framework provides a formal way to predict the temporal dynamics of processes that integrate evidence for one choice option over another for the temporal period leading up until choice, and to separate these from initial perceptual processes time-locked to stimulus presentation. Neural signals corresponding to these predicted evidence-accumulation signals have been identified with EEG for perceptual decision making across different sensory modalities or stimulus features (Kelly and O'Connell, 2013; O'Connell et al., 2012; Wyart et al., 2012) as well as for value-based decision making (Pisauro et al., 2017; Polanía et al., 2014). These studies have identified evidence accumulation processes either as the model-free build-up rate of the centroparietal positivity (CPP) (Kelly and O'Connell, 2013; Loughnane et al., 2018; Loughnane et al., 2016; O'Connell et al., 2012) or in SSM-prediction-based neural signals measured over parietal and/or frontal regions (Pisauro et al., 2017; Polanía et al., 2014). Both types of neural signals are commonly interpreted as reflecting integration of the choice-relevant evidence to reach a decision, rather than basic motor planning which is usually identified by a fundamentally different neural signal, the contralateral action readiness potential (Kornhuber and Deecke, 2016; Schurger et al., 2021). The cortical origins of these signals may in principle correspond to locations identified by fMRI studies of corresponding SSM-predicted evidence accumulation traces, but note that these studies were not able to study the temporal dynamics of such signals and to unambiguously separate them into stimulus-locked perceptual versus response-locked decision processes (Gluth et al., 2012; Hare et al., 2011; Hutcherson et al., 2015; Rodriguez et al., 2015). Studies using this approach to investigate different types of decisions have identified different cortical areas that implement evidence-accumulation signals in different choice contexts (e.g. parietal regions specifically for perceptual decision making vs. both frontal and parietal regions for value-based decision making Polanía et al., 2014). This shows that different types of decisions may, even if they are reported via the same manual actions, draw on evidence accumulation computations that are instatiated in distinct brain regions. Moreover, altruistic decisions driven by different motives, or made by individuals with different social preferences, have also been found to involve activity in different neural networks (Hein et al., 2016). Therefore, it is necessary to differentiate whether the contextual and individual differences in altruistic decisions reflect recruitment of different brain areas/signals and/or of different computations that are performed within these brain areas. If different final decision mechanisms (i.e. computational and/or neural mechanisms) were to be involved in the two types of altruistic choices, or in different individuals, we should observe response-locked evidence-accumulation signals in different brain areas (e.g. frontal vs. parietal regions), or even different types of computations, in the two types of inequality contexts and/or different individuals. Conversely, if the same final decision mechanism is employed for both types of choice contexts, we should observe similar evidence-accumulation neural signals in similar brain areas, but systematic variations across contexts and/or individuals in those signals (e.g. responses in different brain areas and/or with different temporal characteristics) related to early perceptual/attentional processing of choice-relevant information, such as the available payoff magnitudes (Harris et al., 2018). Here, we apply this approach and use SSMs fitted to individuals' wealth distribution behaviors to predict the underlying neural evidence accumulation dynamics. We then employ these predicted EA signals in our EEG analyses to examine whether a similar neural choice system accumulates the choice-relevant evidence in both inequality contexts, or whether distinct neural systems implement this decision process for the different contexts. Then, we examine whether the different features of each choice problem that ultimately need to be integrated into the choice-relevant evidence – that is, the specific payoffs available to oneself and the other person – are initially processed in a different manner for different contexts and in different individuals. This allows us to directly approach the question of whether contextual and individual differences in altruism arise from differences in the decision mechanisms that integrate and compare choice-relevant information at the final stage of the choice process, or rather from differences in the initial processing and biased representation of the choice-relevant information that is ultimately integrated into the final decision mechanism. Results We recorded 128-channel EEG data from healthy participants playing a modified Dictator Game (DG). On each trial of this task, participants played as proposers and chose between two possible allocations of monetary tokens between themselves and an unknown partner. We systematically varied the allocation options from trial to trial so that in half of the trials, participants received less than their partners for both choice options (disadvantageous context [DIS]) and in the other half they got more than their partners for both options (advantageous context [ADV]). These two types of trials were randomly intermixed and were only defined by the size of the payoffs presented on the screen. On each trial, we presented the two options sequentially, to allow clear identification of time points at which the information associated with each option was processed (Figure 1A, see Materials and methods for details). This sequential presentation allowed us to establish the inequality context with the presentation of the first option, without having to explicitly instruct participants about thetwo contexts. We then studied individuals' sensitivity to self-payoff and other-payoff by focusing on how the choice of the second option depended on the change in these variables from the first to the second option. Importantly, as shown in the payoff schedule of all trials (Figure 1—figure supplement 1), we matched self-/other-payoff differences and the resulting absolute levels of inequality across both contexts and also across the second and the first options (Figure 1—figure supplement 1 middle and right panels). This allowed us to compare choices and response times, model-defined neural choice processes time-locked to the response, and neural processing of different stimulus information (self- and other-payoff) between the two contexts. Figure 1 with 2 supplements see all Download asset Open asset Experimental design and behavioral results. We employed a modified dictator game to measure individuals' wealth distribution behaviors. (A) Example of display in a single trial. In the task, participants played as proposers to allocate a certain amount of monetary tokens between themselves and anonymous partners. At the beginning of each trial, participants were presented with one reference option in blue and were asked to keep their eyes on the central cross for at least 1 s to start the trial, as indicated by the change in font color from blue to green. When the second option was presented, participants had to choose between the two options within 3 s. The selected option was highlighted in blue before the inter-trial interval. Font color assignment to phases (i.e. blue and green to response) was counterbalanced across participants. (B) Payoff information and context affect choice systematically. The generalized linear mixed-effects model shows the effects of multiple predictors on the probability to choose the second option; (C) Payoff information and context affect response times systematically. The linear mixed-effects model shows the effects of multiple predictors on response times (RTs). ΔS, Self-payoff Change; ΔO, Other-payoff Change; CON, Context; C, Constant; •••, p < 0.001; ••, p < 0.01; •, p < 0.05. Error bars indicate 95% confidence interval (CI) of the estimates, N=38. Based on the model fits and their predicted response-locked evidence accumulation EEG traces, we first tested whether similar or different neural processes (i.e. brain regions or physiological markers) underlie the ultimate choice process in the two inequality contexts, in similarity to how this has been studied for other types of decisions (Polanía et al., 2014). Then, we clarified whether neural processing of the stimulus information – which subsequently feeds into the decision processes – differs across contexts and individuals. For this analysis, we examined stimulus-locked event-related potentials (ERPs), in a way that has also been used to differentiate neural processing of decision-relevant features in non-social value-based decision making (e.g. perceptions of health and taste of food items) (Harris et al., 2018). Finally, we explored how individual differences in altruism are related to large-scale information communications between regions associated with these two sets of processes (i.e. response-locked decision processes and stimulus-locked perceptual processes), by examining inter-regional synchronization in the gamma-band frequency (30–90 Hz). This last analysis was motivated by the consideration that evidence accumulation processes need to integrate evidence input from different neural sources (e.g. perceptual processes) (Polanía et al., 2014), and by the proposal that coherent phase-coupling in the gamma band between different groups of neurons may serve as a fundamental process of neural communication for information transmission (Bosman et al., 2014; Fries, 2009; Fries, 2005; Vinck et al., 2013), as already shown for non-social value-based decisions (Polanía et al., 2014; Siegel et al., 2008). Behavior: Altruism depends differentially on self- and other-payoffs across contexts Before performing model-based analyses, we ran model-free linear mixed-effects regressions to establish that the choice-relevant information (i.e. self-payoff, other-payoff, and inequality context [ADV and DIS]) indeed systematically affects individual wealth distribution choices. These analyses confirmed that both self-payoff and other-payoff were important factors underlying individuals' choices. Specifically, participants chose the second option more often when either they or the receiver profited more from this choice (main effect Self-payoff Change (ΔS): beta = 3.77, 95% CI [3.65–3.89], p < 0.001; main effect Other-payoff Change (ΔO): beta = 0.56, 95% CI [0.51–0.61], p < 0.001, ΔS(ΔO): participants' own (partners') payoff change between the second and the first option) (Supplementary file 1, Figure 1B). However, participants were less influenced by changes in their own payoff when they had more money than the other (ADV, interaction Self-payoff Change and beta = 95% CI to p < or when the receiver got payoffs from this choice other-payoff, interaction Self-payoff Change and Other-payoff Change (ΔO): beta = 95% CI p = This effect was when the participants had more money than the receiver interaction Self-payoff Change Other-payoff Change and beta = 95% CI p < 0.001; file 1, Figure 1B). For of these see Appendix 1 and Figure 1—figure supplement for by model-based analyses see Appendix that we also models without interaction effects and/or main but model analyses the model (Supplementary file linear mixed-effects model suggested that the presentation (i.e. first or of options would not affect individuals' choices Appendix 1 and file other-payoff, and context also how participants their decisions. were for absolute values of self-payoff change (main effect Self-payoff Change beta = 95% CI to p < and other-payoff change (main effect of Other-payoff Change beta = 95% CI to p = (Figure both these effects were different for the two inequality contexts, with response times more strongly in the disadvantageous inequality context between Self-payoff Change and beta = 95% CI p < 0.001; interaction between Other-payoff Change and beta = 95% CI p = file Figure These effects are consistent with the central of the SSM framework that evidence will up evidence accumulation and resulting choice, thereby already that an decision process may integrate self- and other-payoff to guide individual decisions of these see Figure 1—figure supplement EEG similar parietal evidence accumulation across contexts To address the question of whether distribution choices are by similar or different neural decision processes across both inequality contexts, we fitted a sequential sampling model to participants' behavioral data and used it to predict neural evidence accumulation signals for the two contexts. Our analyses EA signals over similar parietal regions for both contexts and EA signals that would indicate the use of fundamentally different final choice mechanisms in the different contexts. Specifically, we first fitted the SSM by trials as or choices, on whether the selected the option with more or less distribution of monetary tokens between both For each trial, the model used the subjective value difference between the more option and the more option using the utility see Materials and as input to predict evidence accumulation signals until the when the decision was For we used the choice model which a process et al., (1) (2) with for = for disadvantageous inequality = ADV for advantageous inequality context), s for participants = and for trials = participants' payoff of the option in for s and trial the payoff in the option in for s and trial This model allowed us to parameters that correspond to different aspects of and the choice the decision on others decision and rate to as well as parameters which are less to be to the or neural mechanisms underlying valuation or decision and time Materials and methods for a detailed model these we could examine the effects of context on both basic altruistic (i.e. and the final decision process that the subjective values on from perception and valuation processes (i.e. and Although the payoff of each option was for each trial, participants still had to evidence by and the difference in payoffs between so the decision time may have the evidence accumulation when the decision process the The thus how the decision process the or of evidence see Materials and methods for the of models and the of the model we used for our To the evidence accumulation process, we EA by the fitted model for the context and the payoffs on each trial. that these were of the EA processes underlying choice, since the fitted model could both choices and across the two contexts. For both types of choices and contexts and DIS), the of the data by the model was than (Figure and the was than (Figure right Materials and The model also response that choices are during advantageous inequality overall in ADV vs. 95% CI = = p < and for

Concepts can be related in many ways. They can belong to the same taxonomic category (e.g., "doctor" and "teacher," both in the category of people) or be associated with the same event context (e.g., "doctor" and "stethoscope," both associated with medical scenarios). How are these two major types of semantic relations coded in the brain? We constructed stimuli from three taxonomic categories (people, manmade objects, and locations) and three thematic categories (school, medicine, and sports) and investigated the neural representations of these two dimensions using representational similarity analyses in human participants (10 men and nine women). In specific regions of interest, the left anterior temporal lobe (ATL) and the left temporoparietal junction (TPJ), we found that, whereas both areas had significant effects of taxonomic information, the taxonomic relations had stronger effects in the ATL than in the TPJ ("doctor" and "teacher" closer in ATL neural activity), with the reverse being true for thematic relations ("doctor" and "stethoscope" closer in TPJ neural activity). A whole-brain searchlight analysis revealed that widely distributed regions, mainly in the left hemisphere, represented the taxonomic dimension. Interestingly, the significant effects of the thematic relations were only observed after the taxonomic differences were controlled for in the left TPJ, the right superior lateral occipital cortex, and other frontal, temporal, and parietal regions. In summary, taxonomic grouping is a primary organizational dimension across distributed brain regions, with thematic grouping further embedded within such taxonomic structures.SIGNIFICANCE STATEMENT How are concepts organized in the brain? It is well established that concepts belonging to the same taxonomic categories (e.g., "doctor" and "teacher") share neural representations in specific brain regions. How concepts are associated in other manners (e.g., "doctor" and "stethoscope," which are thematically related) remains poorly understood. We used representational similarity analyses to unravel the neural representations of these different types of semantic relations by testing the same set of words that could be differently grouped by taxonomic categories or by thematic categories. We found that widely distributed brain areas primarily represented taxonomic categories, with the thematic categories further embedded within the taxonomic structure.

The profound nature of moral judgment has been discussed and debated for centuries. When facing the trade-off between pursuing moral rights and seeking better consequences, most people make different moral choices between two kinds of dilemmas. Such differences were explained by the dual-process theory involving an automatic emotional response and a controlled application of utilitarian decision-rules. In neurocognitive studies, the bilateral dorsolateral prefrontal cortex (DLPFC) has been demonstrated to play an important role in cognitive “rational” control processes in moral dilemmas. However, the profile of results across studies is not entirely consistent. Although one transcranial magnetic stimulation (TMS) study revealed that disrupting the right DLPFC led to less utilitarian responses, other TMS studies indicated that inhibition of the right DLPFC led to more utilitarian choices. Moreover, the right temporoparietal junction (TPJ) is essential for its function of integrating belief and intention in moral judgment, which is related to the emotional process according to the dual-process theory. Relatively few studies have reported the causal relationship between TPJ and participants' moral responses, especially in moral dilemmas. In the present study, we aimed to demonstrate a direct link between the neural and behavioral results by application of transcranial direct current stimulation (tDCS) in the bilateral DLPFC or TPJ of our participants. We observed that activating the right DLPFC as well as inhibiting the left DLPFC led to less utilitarian judgments, especially in moral-personal conditions, indicating that the right DLPFC plays an essential role, not only through its function of moral reasoning but also through its information integrating process in moral judgments. It was also revealed that altering the excitability of the bilateral TPJ using tDCS negligibly altered the moral response in non-moral, moral-impersonal and moral-personal dilemmas, indicating that bilateral TPJ may have little influence over moral judgments in moral dilemmas.