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Logo of nihpaAbout Author manuscriptsSubmit a manuscriptHHS Public Access; Author Manuscript; Accepted for publication in peer reviewed journal;
CNS Spectr. Author manuscript; available in PMC 2011 August 29.
Published in final edited form as:
CNS Spectr. 2009 November; 14(11): 608–620.
PMCID: PMC3163302

The Neurobiology of Moral Behavior: Review and Neuropsychiatric Implications

Dr. Mario F. Mendez, MD, PhD, Professor


Morality may be innate to the human brain. This review examines the neurobiological evidence from research involving functional magnetic resonance imaging of normal subjects, developmental sociopathy, acquired sociopathy from brain lesions, and frontotemporal dementia. These studies indicate a “neuromoral” network for responding to moral dilemmas centered in the ventromedial prefrontal cortex and its connections, particularly on the right. The neurobiological evidence indicates the existence of automatic “prosocial” mechanisms for identification with others that are part of the moral brain. Patients with disorders involving this moral network have attenuated emotional reactions to the possibility of harming others and may perform sociopathic acts. The existence of this neuromoral system has major clinical implications for the management of patients with dysmoral behavior from brain disorders and for forensic neuropsychiatry.


For years, scientists and philosophers have proposed a sixth human sense for morality. Recently, there is increasing evidence that there is, in fact, an intrinsic morality network. The presence of a moral sense is consistent with a focus of human evolution on mechanisms of individual behavior that maximize survival in social groups. Evolution has promoted social cooperation through emotions against harming others, a need for fairness and the enforcement of moral rules, empathy and “Theory of Mind” (ToM), as well as other behaviors that feed into the concept of morality. ToM is the ability to appreciate the thoughts, feelings, and beliefs of others.

If there is a “moral sense,” then there should be specific brain mechanisms for morality as well as brain disordered patients with impaired morality. Convergent evidence that this is the case comes from studies of functional magnetic resonance imaging (fMRI) in normals, neurological investigations of sociopaths, and the examination of patients with focal brain lesions or with frontotemporal dementia (FTD). This neurobiological evidence points to an automatic, emotionally- mediated moral network that is centered in the ventromedial prefrontal cortex (VMPFC), particularly in the right hemisphere. Although this literature is still young, disparate, and heavily reliant on fMRI correlations, the convergence of evidence supports the presence of a neuromoral brain network. This report reviews this burgeoning literature and discusses the theoretical implications for brain-behavior relationships, and its clinical and legal implications. Although much of the presented evidence is still debated, a picture of moral neuroscience is beginning to emerge.


Morality is a code of values and customs that guide social conduct. An example of a moral value is the avoidance of harm to others (“no-harm” rule). Philosophers often divide morality into “descriptive” or “normative” types. Descriptive morality is a code of conduct held by a particular society or group as authoritative in all matters of right and wrong. It focuses on areas beyond no-harm, such as purity, accepting authority, and emphasizing loyalty to the group.1 Normative morality, on the other hand, is a universal code of moral actions and prohibitions held by all rational people, irregardless of their society or group’s descriptive morality.1,2 It focuses predominantly on no-harm and fairness but also includes the other aspects of morality. Philosophers since pre-Socratic times have long pondered the existence of a universal normative morality in addition to the descriptive codes proposed by each society, religion, or legal system.

Neurobiology is concerned with normative morality, which can lead to different codes of moral behavior when it interacts with socio-cultural learning.24 The interface of evolutionary psychology with social neuroscience points to universal “neuromoral” emotions and drives that strengthen social cohesion and cooperation. 5,6 Studies with apes and other social animals describe moral emotions such as empathy, gratitude, a sense of fairness, feelings of reciprocity, righteousness, consolation, and group loyalty.79 In humans, moral emotions, such as guilt, shame, embarrassment, gratitude, compassion, pride, fear of negative evaluation by others, and outrage at unfair treatment, are strong motivators to act in a socially favorable way.10,11 These emotions or sentiments allow humans to quickly grasp the moral implications of social interactions and then act to enhance their personal reputations and the likelihood of future social cooperation.12 Furthermore, moral emotions are manifestations of evolutionary-based neuromoral drives including no-harm, fairness or equity, community, authority, and purity.1,13 The two most prominent of these may be no-harm, as evidenced by the discomfort felt on directly hurting others,14 and fairness, as evidenced by the need to punish “free-riders” or those who cheat and break the rules.15


Investigators have used fMRI in normals to define the neuroanatomy of moral behavior.4,14,16 These studies usually involve tasks or dilemmas of moral judgment or reasoning.14,1619 The main neuromoral areas involved are the VMPFC and adjacent orbitofrontal, plus ventrolateral, cortex (OFC/VL), amygdalae, and the dorsolateral prefrontal cortex (DLPFC) (Figure 1).4,5,14,16,18 The VMPFC (defined here as Brodmann’s areas [BA]10–12, 25, 32 plus the frontopolar region of BA10) attaches moral and emotional value to social events, anticipates their future outcomes, and participates in ToM, empathy, attribution of intention, and related tasks.2023 The OFC/VL region (defined here as BA47, parts of BA10–12 and 25, plus VL BA44), mediates socially aversive responses, changes responses based on feedback, and inhibits impulsive, automatic, or amygdalar responses.2426 The amygdalae, located in the anteromedial temporal lobes, mediate the response to threat and aversive social and moral learning.6,27,28 The DLPFC can override this neuromoral network through the application of reasoned analysis to moral situations. 18 Finally, some fMRI morality and related tasks activate additional regions such as the anterior insula,29 posterior superior temporal sulcus (pSTS),3032 anterior cingulate gyrus,33 the inferior parietal lobules and temporoparietal junctions,32,3436 ventral striatum and mesolimbic reward system,37 precuneus,35 and posterior cingulate.30,32 Other regions, such as subcortical limbic and anterior temporal lobes, can lead to impaired moral behavior as well.5,38

Anatomic areas in morality network

The VMPFC, particularly on the right, is core to this neuromoral system. In fMRI studies, this brain region becomes activated with tasks requiring explicit moral judgments, passive viewing of morally salient photos (Figure 2), and the elicitation of charity, fairness, guilt, and other moral emotions and sentiments.16,22,30,39 Greene and colleagues14 have found VMPFC activation on presenting “personal” moral dilemmas involving the possibility that the participant’s direct action could cause someone serious harm (Table 1), compared to “impersonal” moral dilemmas where the possibility of causing someone serious harm does not involve the participant’s direct action on another. These findings indicate that the VMPFC mediates automatic moral and “prosocial” reactions, such as discomfort at the prospect of being a direct agent of a personal moral violation or of harm to someone else.4042 In fact, fMRI studies indicate that the VMPFC participates in prosocial, affiliative, or social attachment emotions in general, including guilt, embarrassment, and compassion. 22,43,44 In contrast, Greene and colleagues14 have found DLPFC activation on presentation of the impersonal moral dilemmas, suggesting a later, dispassionate, reasoned, or cost-benefit assessment for moral judgments in the absence of a sufficient VMPFC “moral reaction.”8,18,30,4550

Ventromedial prefrontal cortex in moral judgment and emotion16
Example of Reasoned and Personal Moral Dilemmas14

In addition to emotions to do no harm, moral emotions serve to enforce moral rules by attributing negative intentions and seeking to punish “cheaters” who violate them.15,44,51 This “altruistic punishment” is a manifestation of the moral drive for fairness and equity. There is increased VMPFC activation from a sense of fairness (Figure 3), which contributes to the drive to punish violators or non-cooperators, even if costly for the punisher.44.51,52 This is further exemplified in the Ultimatum Game, where one player must divide a sum of money with a second player, but if the second player rejects the division as unfair, neither player receives anything. The second player’s rejection of unfair offers, and the foregoing of any money whatsoever, usually reflects both their sense of fairness and desire for altruistic punishment.53 Altruistic punishment is strongly dependent on determining that others, particularly those with a negative reputation, are deliberately not playing by the rules.44,54 The VMPFC is involved through its role in attribution or in inferring the intention of others’ behavior. 55 The OFC/VL (BA47) region and neighboring anterior insula and amygdala, especially on the right,52,56 subsequently effect altruistic punishment through sentiments linked to social aversion/exclusion, such as anger, indignation, disgust, and contempt.22,43,44,5761

fMRI study of fairness52

ToM and empathy are two processes very closely related to morality. ToM involves the VMPFC, which facilitates the appreciation that others have thoughts, feelings, and beliefs.33,62,63 The “cognitive” aspects of empathy, such as taking someone else’s perspective and vicariously identifying with it, involve the VMPFC (BA10,11) in a phylogenetically new system that only occurs in great apes and advanced mammals.42,6468 The most emotional aspects of empathy, such as emotional contagion, include the OFC/VL (BA44) in a phylogenetically old system.64,65,6971 Variables that strongly affect “cognitive” empathy, and impact on VMPFC (BA10) activity, include the self as the agent of an action and the perceived similarity between the self and others.58,72 This suggests that the VMPFC deals with complex “self-other conjoining,” or a resonating of the protagonist’s mental and emotional states with that of someone else. Other areas modulate self-other conjoining, including OFC/VL mirror neurons, when observed intentions and emotions of others are internally mapped or imitated;65,7382 the anterior cingulate cortex, when self-concepts are threatened by the outperformance of others (envy);33 and the ventral striatum, when pleasure results from another’s misfortune (“schadenfreude”).33


Neuroscience has come a long way since the claims of an innate “criminal mind” from Cesare Lombroso and others.83 Neuroimaging and other techniques have revealed a great deal about the neurobiology of morality through the study of sociopathy, or chronic antisocial behavior. Sociopaths lack moral emotions, empathy, conscience, or remorse and guilt for their acts. Although they have difficulty distinguishing between moral (victim-based) transgressions and conventional (social disorder-based), they have normal moral knowledge and reasoning.60,61 Sociopaths have instrumental (cold-blooded and goal-directed) aggression with decreased sympathetic arousal. On psychophysiological measures, they show minimal alterations in heart rate, skin conductance, or respirations when they are subjected to fear or stressful or unpleasant pictures, and they have reduced autonomic responses to the distress of others, as well as reduced recognition of sad and fearful expressions.8486

Those who have committed violent offenses have a high incidence of neurological changes. In one study, nearly two-thirds of murderers had neurological diagnoses, including brain injuries, mental retardation, cerebral palsy, epilepsy, dementia, and others.87 Neurological examinations often show marked frontal or temporal deficits or changes on neuroimaging or electroencephalography.88 Some of these deficits could be due to alcohol or substance abuse or other confounding variables, and future studies will need to control for these variables. Deficits in frontal functions such as inability to change their responses (response reversal learning) or to inhibit risk-taking behavior after negative feedback occur among institutionalized and violence-prone patients.60,8994 Functional neuroimaging studies can show frontotemporal hypometabolism, hypoperfusion, or changes in spectroscopy in murderers pleading not guilty by reason of insanity and in violent psychiatric inpatients.86,88,89,9598 Voxel-based morphometry reveals a correlation of frontopolar and OFC/VL grey matter reduction with increased psychopathic traits or scores.98,99 A reduction in prefrontal gray matter volume associated with reduced autonomic arousal occurs among violent offenders.100 Moreover, the smaller the volume of prefrontal cortex, the greater the tendency towards sociopathic behavior is among known sociopaths.101 Paradoxically, boys with callous unemotional conduct problems have increased gray matter concentrations in medial frontal regions, suggesting a delay in cortical maturation.102

In addition to the predominant frontal lobe abnormalities, sociopaths have reduced function of the amygdalae.60,103 These structures are involved in aversive or fear conditioning, instrumental learning (reward), and the retrieval of socially relevant knowledge, such as facial trust-worthiness, approachability, or fear.27 Sociopaths are impaired in these and in anticipatory stimulus- reinforcement learning,104 information which is needed by the VMPFC for the development of normal moral socialization.60,61,105 Animal studies further show that the early amygdalar dysfunction disrupts the appropriate development of the VMPFC and the OFC/VL.60,105 In developmental sociopathy, early amygdalar dysfunction may result in VMPFC and OFC/VL dysfunction, through the impaired association of actions that harm others with the aversive reinforcement of the victims’ distress. Finally, there is evidence for subtle changes in a whole network of areas in developmental psychopathy.98


Ever since the descriptions of Phineas Gage, perhaps neurology’s most famous patient, clinicians have viewed patients with VMPFC lesions as characterized by alterations in social and moral behavior (Figure 4). Focal lesions affecting VMPFC and adjacent OF/VLC include strokes, trauma, tumors, infections, and a ruptured anterior commissure aneurysm.83,106 Right frontal lesions may be especially associated with deviant social behavior and left frontal lesions associated with violent or angry outbursts.83,107,108 Consistent with other data, investigations show that focal lesions in the VMPFC and OFC/VL impair moral judgment, and early lesions of these areas impair the development of moral knowledge and judgment. 109,110 When VMPFC lesions are acquired before 16 months of age, they may lead to severe antisocial behavior, insensitivity to future consequences of decisions, and repeated failure to respond to behavioral interventions.109

Phineas Gage’s skull

The lesions implicated in “acquired sociopathy” involve the VMPFC and the OFC/VL. Patients with focal VMPFC lesions, especially on the right, have attenuated feelings of emotional discomfort for sociomoral violations, severe deficits in empathy, and reduced responsiveness to victims.9,22,68,89,110120 VMPFC lesions, especially on the right, disturb “fortune-of-others” and related emotions such as compassion, shame, guilt, envy, inappropriate pride, and gloating; emotions that correlate with perspective-taking and ToM.109,116,117,121124 Although first-order ToM may be intact, they may be particularly unable to read the feelings and emotions (“affective ToM”) of others as evident on irony and faux pas tasks.118,125 Like developmental sociopaths, patients with VMPFC lesions had autonomic hyporesponsivity, especially in response to social stimuli,66,100,116,120 but compared to developmental sociopaths, their hyporesponsivity is more general and not selective for fearful or sad expressions.126,127 They also differ in that patients with VMPFC lesions do not crave constant stimulation, and they are not deceitful, manipulative, or instrumentally aggressive.59 Finally, OFC/VL lesions impair the use of immediate feedback from social and emotional cues, and the control of emotional and impulsive responses.24,57,59,60,128

Several recent research studies have used moral dilemmas to investigate moral decision-making among patients with VMPFC lesions. Ciaramelli and colleagues114 compared 7 patients with VMPFC lesions with 12 normal controls on personal, impersonal, and non-moral dilemmas. Compared to normal controls, the patients were significantly more willing to judge personal moral violations as acceptable behaviors, and they did so quickly and with little hesitation. In a similar study, Koenigs and colleagues117 carefully examined six patients with focal bilateral damage to the VMPFC (Figure 5). Their patients maximized the good for the many (utilitarianism) on moral dilemmas, had impaired autonomic activity in responses to emotionally charged pictures, and had diminished empathy, embarrassment, and guilt. In both of these studies, compared to controls, the patients with VMPFC damage tend to make utilitarian choices in conflicting moral dilemmas.114,117 In contrast, patients with VMPFC lesions may continue to reject unfair offers in the Ultimatum Game,129 a finding that suggests that their intact OFC/VL region, with relatively preserved socially aversive sentiments,22 continues to apply altruistic punishment in situations where fairness and intentionality are overt or predefined.12,22 The sparing of the VL areas related to aversion may add to the emotional negative aspects of their behavior and render them vehement self-centered moralists.12,114

Ventromedial prefrontal cortex in moral judgment and emotion117

Despite their deficits in per sonal moral responses, patients with adult-onset VMPFC lesions retain moral knowledge and reasoning. Although early-onset VMPFC lesions can impair the acquisition of moral knowledge,109,110 patients with adult-onset VMPFC lesions have preserved moral reasoning and retain the knowledge of moral rules and conventions.129132 On moral tasks, they can verbalize the differences between right and wrong responses, but they may not act on that knowledge and can score well on self-report moral behavioral inventories (Table 2). In sum, patients with VMPFC lesions do not act according to their retained moral knowledge because they have a deficit in prosocial or affiliative sentiments, and do not use their moral reasoning abilities to anticipate the future consequences, outcomes, or feelings of their actions.1012,22,112,113,133137 One reason for this partially retained moral knowledege is that it is located outside the VMPFC, particularly in the right anterior temporal lobe.43

Moral Behavior Inventory150


Brain disorders including slow, insidious neurodegenerative dementias, are another window to the organization of morality in the brain. There are many brain disorders that can disturb sociomoral behavior, such as Huntington’s disease, traumatic brain injury, frontal tumors and other conditions (Table 3). Perhaps the most characteristic is FTD. This disease can serve as a model to illustrate alterations in moral behavior from brain diseases.

Neurological Diseases with Potential Disturbances of Moral Behavior

In contrast to the memory and cognitive deficits of Alzheimer’s disease and other dementias, the core features of FTD are transgression of social norms including sociopathic behavior, a loss of empathy or appreciation of the feelings of others, and a loss of insight for their behavior and its consequences.138 In FTD, asymmetric right-sided frontal involvement is especially associated with socially undesirable behaviors, loss of empathy, and a distorted appreciation of others. 139143 FTD patients manifest violations of social and moral rules or norms early in their disease.138 Most commonly there is a loss of social tact and propriety, unacceptable physical contact, and improper verbal or nonverbal communication.144 Sociopathic behavior occurs in more than half of patients with FTD (Table 4).145 Investigators have described FTD patients with stealing and shoplifting, 145147 inappropriate sexual behavior,138,148 physical aggression and acts of violence,146,148 frequent traffic violations and hit-and-run accidents,148 pedophilia,149 and other transgressions.145 These sociopathic acts are associated with right frontal, presumably VMPFC, involvement on imaging and on neuropathology.143,144

Sociopathic Acts Among 16 Patients with Frontotemporal Dementia145

Early FTD affects the VMPFC more than the DLPFC, and studies show corresponding impairments of emotionally-based moral judgments. In one study, FTD patients were impaired in their ability to immediately respond to personal moral dilemmas, compared to Alzheimer’s disease patients and normal controls.150 Using relatively intact DLPFC processes, the FTD patients solved the moral dilemmas in a logical, cold, and calculating fashion. These patients have problems with empathy, both empathic concern and perspective taking.136 Investigations of personality among FTD patients have also shown decreased empathy and decreased agreeableness with right OFC/VL involvement and inter-personal coldness or decreased emotional empathy with anterior temporal involvement.151154 On voxel-based morphometry studies, empathy correlates with a right medial frontotemporal network (VMPFC and anterior temporal areas).151,155 FTD patients are particularly impaired in gauging the seriousness of moral transgressions.136 Other studies document defects in ToM in patients with FTD,125,136,156158 as well as deficits in social concepts from right anterior temporal lobe involvement.43

In FTD, defective moral emotions along with decreased self-other conjoining could account for defective moral judgment as well as most of their observed sociopathic acts. There is a selective impairment in decision-making in personal moral judgment in FTD in the face of relatively preserved moral knowledge and moral reasoning ability.150 Their impersonal responses to personal moral violations are consistent with the early focus of neuropathology in the VMPFC.144 These changes, coupled with insufficient control of impulsivity from adjacent OFC/VL involvement, explain the tendency to impulsive moral violations in full knowledge of the potential consequences. Furthermore, involvement of the rigth anterior temporal in FTD can lead to social knowledge deficits and contribute to the moral behavioral changes in this disorder.43,151


The evidence from fMRI studies in normals, sociopathy, brain lesions, and FTD suggest a neuromoral network of prosocial emotions and drives that promote social cohesiveness and cooperation.1,6,159 Most moral judgments are rapid, involuntary, and intuitive; whereas, deliberate rational reasoning is often post hoc rationalization for judgments which have already occurred.8,13,45 Normative morality appears to be rooted in an intrinsic neuromoral network.

The neuromoral network comprises the VMPFC, especially on the right, the OFC/VL, the amygdalae, and related structures.16 The VMPFC, with its rich interconnections with limbic structures, mediate these strong automatic reactions to moral violations.4,14,16,18,45,81 Brain lesions or diseases that involve the right VMPFC reduce moral emotions and responses to dilemmas that involve either harm to others or a sense of fairness or equity. The OFC/VL region manages socially aversive emotions and may inhibit immediate and amygdalar responses, suppresses impulsive behavior, and responds to feedback learning.24,59 The amygdalae mediate the response to immediate threat, aversive social learning, and help incorporate social or moral prohibitions.27 Some investigators also propose that this morality network can be over-ridden by DLPFC-mediated reasoning processes, resulting in utilitarianism, ie, the greatest good for the greatest many.22

If the brain has a “moral grammar,” it is less in the Chomsky sense than in an interaction of moral drives with the process of self-other conjoining. 160,161 Biological moral drives, such as no-harm and fairness, are the forces that result in moral emotions. In the VMPFC, this is coupled with automatic, complex self-other conjoining. The result is the creation of joint attention and “intersubjective space,”162 ie, the activation of one’s representations of the state and situation of others.8,76 Unless actively inhibited, activation of these shared representations probably occurs automatically through mirror neurons and results in ToM, empathy, moral emotions, and moral behavior.163167

These new findings have implications for clinical neuropsychiatry. Patients may present with alterations in moral behavior due to brain disorders. They can manifest as shoplifting, hit-and-run accidents, lack of empathic aid, or the spectrum from subtle changes in personality all the way to serious crimes like pedophilia.149 When patients present with dysmoral behavior for the first time, as a change from a prior pervasive pattern of behavior, clinicians need to consider a possible, causative brain disorder (Table 3). Family and friends need education as to the significance of the patient’s behavior, and the question of whether their dysmoral behavior is their “fault” may need frank discussion. Lastly, medications can be useful in controlling related behaviors such as impulsivity, but do not selectively suppress dysmoral behavior. Selective serotonin reuptake inhibitors, beta-blockers, and mood stabilizing antiepileptic agents (such as valproate, carbamazepine, and lamotrigine) could be of help in this regard.

The neurobiology of morality raises additional questions of legal culpability. Patients with VMPFC lesions or FTD with disturbed volition have committed crimes and been arrested.145,146,148 The US federal insanity defense hardens the original M’Naughton rule, requiring the defendant to prove, by “clear and convincing evidence,” that “at the time of the commission of the acts constituting the offense, the defendant, as a result of a severe mental disease or defect, was unable to appreciate the nature and quality or the wrongfulness of his acts” (18 U.S.C. § 17).168 Without the restraint of intuitive moral emotions and self-other conjoining, however, patients may not be able to deter an impulse to act in an unacceptable manner, even as they know right and wrong and understand the nature of their acts. Furthermore, Anglo-American jurisprudence distinguishes between reason-based law and a natural law based on what a “reasonable person” would do in like circumstance.169 Paradoxically, under the law a “reasonable person” is someone who actually responds to intact moral emotions; therefore, the proof that these patients lack the faculties of a “reasonable person” is the sociopathic behaviors themselves. These considerations demand a reappraisal of the how we view culpability and criminal violations among brain-injured patients.83

Of necessity, this review summarizes a large number of studies that are disparate in their scope and variable in their approach. Most published studies focus on social behavior, rather than specifically on moral behavior. The case for a neuromoral network is primarily based on cross-sectional studies correlating fMRI findings with either task performance or with clinical characteristics supplemented by developmental and acquired sociopathy, and findings from patients with FTD. As a result, the conclusions from this review require future hypothesis-driven studies with specific causal inferences.


Current research is beginning to outline a neuromoral network with a hub in the VMPFC. This research has implications for understanding the organization of our moral sense in the brain and has implications for clinical and forensic neuropsychiatry. The findings reviewed here are preliminary, but this story promises to rapidly unfold as more research is done on the neurobiological basis of morality in normals and in brain-injured patients.


  • Humans have an innate moral sense based in a neuromoral network centered in the ventromedial prefrontal cortex and its connections.
  • The neuromoral network works through moral emotions and moral drives, such as the avoidance of harm to others and the need for fairness and punishment of violators; it includes self-other conjoining processes, such as Theory of Mind and empathy, which also involve the ventromedial prefrontal cortex.
  • Disorders of this region, such as focal lesions or frontotemporal dementia, disturb personal, intrinsic moral emotions and decision-making.
  • Clinicians must recognize and manage “acquired sociopathy” and other dysmoral behaviors associated with disorders of the neuromoral network.
  • Patients with these disorders pose a special problem for forensic neuropsychiatry.


Funding/Support: This work was supported by grant #R01AG034499-02.


Faculty Disclosures: Dr. Mendez reports no affiliations with or financial interest in any organiztion that may pose a conflict of interest.


1. Haidt J. The new synthesis in moral psychology. Science. 2007;316:998–1002. [PubMed]
2. Wilson JQ. The Moral Sense. New York, NY: Simon & Schuster; 1993.
3. Hauser MD. Moral Minds: How Nature Designed our Universal Sense of right and Wrong. New York, NY: Ecco/Harper Collins; 2006.
4. Moll J, de Oliveira-Souza R, Eslinger PJ. Morals and the human brain: a working model. Neuroreport. 2003;14:299–305. [PubMed]
5. Moll J, Zahn R, de Oliveira-Souza R, Krueger F, Grafman J. Opinion: the neural basis of human moral cognition. Nat Rev Neurosci. 2005;6:799–809. [PubMed]
6. Berthoz S, Grezes J, Armony JL, Passingham RE, Dolan RJ. Affective response to one’s own moral violations. Neuroimage. 2006;31:945–950. [PubMed]
7. De Waal FB. How animals do business. Sci Am. 2005;292:54–61. [PubMed]
8. Haidt J. The emotional dog and its rational tail: A social intuitionist approach to moral judgment. Psychol Rev. 2001;108:814–834. [PubMed]
9. Hauser MD, Cushman FA, Young LL. A dissociation between moral judgments and justifications. Mind Language. 2006;22:1–21.
10. Fiske AP. Moral emotions provide the self-control needed to sustain social relationships. Self Identity. 2002;1:169–175.
11. Tangney JP, Stuewig J, Mashek DJ. Mirror neuron system: basic findings and clinical applications. Ann Neurol. 2007;62:213–218. [PubMed]
12. Moll J, Schulkin J. Social attachment and aversion in human moral cognition. Neurosci Biobehav Rev. 2009;33:456–465. [PubMed]
13. Pinker S. The New York Times Magazine. Jan 13, 2008. The moral instinct.
14. Greene JD, Sommerville RB, Nystrom LE, Darley JM, Cohen JD. An fMRI investigation of emotional engagement in moral judgment. Science. 2001;293:2105–2108. [PubMed]
15. Cosmides L, Tooby J, Fiddick L, Bryant GA. Detecting cheaters. Trends Cogn Sci. 2005;9:505–506. [PubMed]
16. Moll J, Oliveira-Souza R, Eslinger PJ, et al. The neural correlates of moral sensitivity: a functional magnetic resonance imaging investigation of basic and moral emotions. J Neurosci. 2002;22:2730–2736. [PubMed]
17. Schaich Borg J, Hynes C, Van Horn J, Grafton S, Sinnott-Armstrong W. Consequences, action, and intention as factors in moral judgments: an FMRI investigation. J Cogn Neurosci. 2006;18:803–817. [PubMed]
18. Greene JD, Nystrom LE, Engell AD, Darley JM, Cohen JD. The neural bases of cognitive conflict and control in moral judgment. Neuron. 2004;44:389–400. [PubMed]
19. Heekeren HR, Wartenburger I, Schmidt H, Schwintowski HP, Villringer A. An fMRI study of simple ethical decision-making. Neuroreport. 2003;14:1215–1219. [PubMed]
20. D’Argembeau A, Xue G, Lu ZL, Van der Linden M, Bechara A. Neural correlates of envisioning emotional events in the near and far future. Neuroimage. 2008;40:398–407. [PMC free article] [PubMed]
21. Damasio AR. Descartes’ Error: Emotion, Reason, and the Human Brain. New York, NY: Putnam; 1994.
22. Moll J, de Oliveira-Souza R. Moral judgments, emotions and the utilitarian brain. Trends Cogn Sci. 2007;11:319–321. [PubMed]
23. Moll J, De Oliveira-Souza R, Zahn R. The neural basis of moral cognition: sentiments, concepts, and values. Ann NY Acad Sci. 2008;1124:161–180. [PubMed]
24. Baxter MG, Parker A, Lindner CC, Izquierdo AD, Murray EA. Control of response selection by reinforcer value requires interaction of amygdala and orbital prefrontal cortex. J Neurosci. 2000;20:4311–4319. [PubMed]
25. Roelofs K, Minelli A, Mars RB, van Peer J, Toni I. On the neural control of social emotional behavior. Soc Cogn Affect Neurosci. 2009;4:50–58. [PMC free article] [PubMed]
26. Rolls ET, Hornak J, Wade D, McGrath J. Emotion-related learning in patients with social and emotional changes associated with frontal lobe damage. J Neurol Neurosurg Psychiatry. 1994;57:1518–1524. [PMC free article] [PubMed]
27. Adolphs R, Tranel D, Damasio AR. The human amygdala in social judgment. Nature. 1998;393:470–474. [PubMed]
28. Luo Q, Nakic M, Wheatley T, Richell R, Martin A, Blair RJ. The neural basis of implicit moral attitude–an IAT study using event-related fMRI. Neuroimage. 2006;30:1449–1457. [PubMed]
29. Hsu M, Anen C, Quartz SR. The right and the good: distributive justice and neural encoding of equity and efficiency. Science. 2008;320:1092–1095. [PubMed]
30. Harenski CL, Hamann S. Neural correlates of regulating negative emotions related to moral violations. Neuroimage. 2006;30:313–324. [PubMed]
31. Singer T, Kiebel SJ, Winston JS, Dolan RJ, Frith CD. Brain responses to the acquired moral status of faces. Neuron. 2004;41:653–662. [PubMed]
32. Robertson D, Snarey J, Ousley O, et al. The neural processing of moral sensitivity to issues of justice and care. Neuropsychologia. 2007;45:755–766. [PubMed]
33. Takahashi H, Lato M, Matsuura M, Mobbs D, Suhara T, Okubo Y. When your gain is my pain and your pain is my gain: Neural correlates of envy and schadenfreude. Science. 2009;323:937–939. [PubMed]
34. Frith CD, Frith U. The neural basis of mentalizing. Neuron. 2006;50:531–544. [PubMed]
35. Young L, Saxe R. The neural basis of belief encoding and integration of moral judgment. Neuroimage. 2008;40:1912–1920. [PubMed]
36. Young L, Cushman F, Hauser M, Saxe R. The neural basis of the interaction between theory of mind and moral judgment. Proc Nat Acad Sci. 2007;104:8235–8240. [PubMed]
37. Schaefer A, Collette F, Philippot P, et al. Neural correlates of “hot” and “cold” emotional processing: a multilevel approach to the functional anatomy of emotion. Neuroimage. 2003;18:938–949. [PubMed]
38. Zahn R, Moll J, Krueger F, Huey ED, Garrido G, Grafman J. Social concepts are represented in the superior anterior temporal cortex. Proc Nat Acad Sci. 2007;104:6430–6435. [PubMed]
39. Decety J, Jackson PL. The functional architecture of human empathy. Behav Cogn Neurosci Rev. 2004;3:71–100. [PubMed]
40. Heekeren HR, Wartenburger I, Schmidt H, Prehn K, Schwintowski HP, Villringer A. Influence of bodily harm on neural correlates of semantic and moral decision-making. Neuroimage. 2005;24:887–897. [PubMed]
41. Waldemann MR, Dieterich JH. Throwing a bomb on a person versus throwing a person on a bomb: intervention myopia in moral intuitions. Psychol Sci. 2007;18:247–253. [PubMed]
42. Young L, Saxe R. An fMRI investigation of Spontaneous Mental State Inference for Moral judgment. J Cogn Neurosci. 2009;21:1396–1405. [PubMed]
43. Zahn R, Moll J, Iyengar V, et al. Social conceptual impairments in frontotemporal lobar degeneration with right anterior temporal hypometabolism. Brain. 2009;132:604–616. [PMC free article] [PubMed]
44. Takahashi H, Kato M, Matsuura M, et al. Neural correlates of human virtue judgment. Cerebral Cortex. 2008;18:1886–1891. [PubMed]
45. Greene J, Haidt J. How (and where) does moral judgment work? Trends Cogn Sci. 2002;6:517–523. [PubMed]
46. Greene J. From neural ‘is’ to moral ‘ought’: what are the moral implications of neuroscientific moral psychology? Nat Rev Neurosci. 2003;4:846–849. [PubMed]
47. Greene JD. Why are VMPFC patients more utilitarian? A dual-process theory of moral judgment explains. Trends Cogn Sci. 2007;11:322–323. [PubMed]
48. Nichols S, Mallon R. Moral dilemmas and moral rules. Cognition. 2006;100:530–542. [PubMed]
49. Prehn K, Wartenberger I, Mériau K, et al. Individual differences in moral judgment competence influence neural correlates of socio-normative judgments. Soc Cogn Affect Neurosci. 2008;2:33–46. [PMC free article] [PubMed]
50. Miller G. Neurobiology. The roots of morality. Science. 2008;320:734–737. [PubMed]
51. De Quervain DJ-F, Fischbacher U, Treyer V, et al. The neural basis of altruistic punishment. Science. 2004;305:1254–1258. [PubMed]
52. Tabibnia G, Satpute AB, Lieberman MD. The sunny side of fairness: preference for fairness activates reward circuitry (and disregarding unfairness activates self-control circuitry) Psychol Sci. 2008;19:339–347. [PubMed]
53. Talmi D, Frith C. Neurobiology–feeling right about doing right. Nature. 2007;446:865–866. [PubMed]
54. Kliemann D, Young L, Scholz J, Saxe R. The influence of prior record on moral judgment. Neuropsychologia. 2008;46:2949–2957. [PubMed]
55. Amodio DM, Frith CD. Meeting of minds: the medial frontal cortex and social cognition. Nat Rev Neurosci. 2006;7:268–277. [PubMed]
56. Knoch D, Nitsche MA, Fischbacher U, Eisenegger C, Pascual-Leone A, Fehr E. Studying the neurobiology of social interaction with transcranial direct current stimulation-- the example of punishing unfairness. Cereb Cortex. 2008;18:1987–1990. [PubMed]
57. Bechara A, Damasio H, Damasio AR. Emotion, decision making and the orbitofrontal cortex. Cereb Cortex. 2000;10:295–307. [PubMed]
58. Zahn R, Moll J, Paiva M, et al. The neural basis of human social values: evidence from functional MRI. Cerebr Cortex. 2009;19:276–283. [PMC free article] [PubMed]
59. Blair RJR. The roles of orbital frontal cortex in the modulation of antisocial behavior. Brain Cogn. 2004;55:198–208. [PubMed]
60. Blair J, Mitchell D, Blair K. The Psychopath. Emotion and the Brain. Oxford, England: Blackwell Publishing; 2005.
61. Blair RJ. Applying a cognitive neuroscience perspective to the disorder of psychopathy. Dev Psychopathol. 2005;17:865–891. [PubMed]
62. Bird CM, Castelli F, Malik O, Frith U, Husain M. The impact of extensive medial frontal lobe damage on ‘Theory of Mind’ and cognition. Brain. 2004;127:914–928. [PubMed]
63. Berthoz S, Armony JL, Blair RJ, Dolan RJ. An fMRI study of intentional and unintentional (embarrassing) violations of social norms. Brain. 2002;125:1696–1708. [PubMed]
64. De Waal FB. Putting the altruism back into altruism: the evolution of empathy. Annu Rev Psychol. 2007;59:1–22. [PubMed]
65. Shamay-Tsoory SG, Aharon-Peretz J, Perry D. Two systems for empathy: a double dissociation between emotional and cognitive empathy in inferior frontal gyrus versus ventromedial prefrontal lesions. Brain. 2009;132:617–627. [PubMed]
66. Eslinger PJ. Neurological and neuropsychological bases of empathy. Eur Neurol. 1998;39:193–199. [PubMed]
67. Gallese V. Before and below ‘theory of mind’: embodied simulation and the neural correlates of social cognition. Philos Trans R Soc Lond B Biol Sci. 2007;362:659–669. [PMC free article] [PubMed]
68. Shamay-Tsoory SG, Tomer R, Berger BD, Aharon-Peretz J. Characterization of empathy deficits following prefrontal brain damage: the role of the right ventromedial prefrontal cortex. J Cogn Neurosci. 2003;15:324–337. [PubMed]
69. Singer T. The neuronal basis and ontogeny of empathy and mind reading: review of literature and implications for future research. Neurosci Biobehav Rev. 2006;30:855–863. [PubMed]
70. Carr L, Iacoboni M, Dubeau M-C, Mazziotta JC, Lenzi JL. Neural mechanisms of empathy in humans: a relay from neural systems for imitation to limbic areas. Proc Natl Acad Sci USA. 2003;100:5497–5502. [PubMed]
71. Nummenmaa L, Hirvonen J, Parkkola R, Hietanen JK. Is emotional contagion special? An fMRI study on neural systems for affective and cognitive empathy. Neuroimage. 2008;43:571–580. [PubMed]
72. Mitchell JP, Banaji MR, Macrae CN. The link between social cognition and self-referential thought in the medial prefrontal cortex. J Cogn Neurosci. 2005;17:1306–1315. [PubMed]
73. Iacoboni M, Mazziotta JC. Mirror neuron system: basic findings and clinical applications. Ann Neurol. 2007;62:213–218. [PubMed]
74. Rizzolatti G, Fabbri-Destro M. The mirror system and its role in social cognition. Curr Opin Neurobiol. 2008;18:179–184. [PubMed]
75. Oberman LM, Pineda JA, Ramachandran VS. The human mirror neuron system: A link between action observation and social skills. Soc Cogn Affect Neurosci. 2007;2:62–66. [PMC free article] [PubMed]
76. Preston SD, de Waal FB. Empathy: Its ultimate and proximate bases. Behav Brain Sci. 2002;25:1–20. [PubMed]
77. Schulte-Ruther M, Markowitsch HJ, Fink GR, Piefke M. Mirror neuron and theory of mind mechanisms involved in face-to-face interactions: a functional magnetic resonance imaging approach to empathy. J Cogn Neurosci. 2007;19:1354–1372. [PubMed]
78. Kaplan JT, Iacoboni M. Getting a grip on other minds: mirror neurons, intention understanding, and cognitive empathy. Soc Neurosci. 2006;1:175–183. [PubMed]
79. Kédia G, Berthoz S, Wessa M, Hilton D, Martinot JL. An agent harms a victim: a functional magnetic resonance imaging study on specific moral emotions. Cogn Neurosci. 2008;20:1788–1798. [PubMed]
80. Lamm C, Batson CD, Decety J. The neural substrate of human empathy: effects of perspective-taking and cognitive appraisal. J Cogn Neurosci. 2007;19:42–58. [PubMed]
81. Lieberman MD. Social cognitive neuroscience: a review of core processes. Ann Rev Psychol. 2007;58:259–289. [PubMed]
82. Schilbach L, Wohlschlaeger AM, Kraemer NC, et al. Being with virtual others: neurological correlates of social interaction. Neuropsychologia. 2006;44:718–730. [PubMed]
83. Markowitsch HJ. Neuroscience and crime. Neurocase. 2008;14:1–6. [PubMed]
84. Intrator J, Hare R, Stritzke P, et al. A brain imaging (single photon emission computerized tomography) study of semantic and affective processing in psychopaths. Biol Psychiatry. 1997;42:96–103. [PubMed]
85. Levenston GK, Patrick CJ, Bradley MM, Lang PJ. The psychopath as observer: emotion and attention in picture processing. J Abnorm Psychol. 2000;109:373–385. [PubMed]
86. Raine A, Buchsbaum M, LaCasse L. Brain abnormalities in murderers indicated by positron emission tomography. Biol Psychiatry. 1997;42:495–508. [PubMed]
87. Blake PY, Pincus JH, Buckner C. Neurologic abnormalities in murderers. Neurology. 1995;45:1641–1647. [PubMed]
88. Wong MT, Lumsden J, Fenton GW, Fenwick PB. Electroencephalography, computed tomography and violence ratings of male patients in a maximum-security mental hospital. Acta Psychiatr Scand. 1994;90:97–101. [PubMed]
89. Critchley HD, Simmons A, Daly EM, et al. Prefrontal and medial temporal correlates of repetitive violence to self and others. Biol Psychiatry. 2000;47:928–934. [PubMed]
90. Ishikawa SS, Raine A, Lencz T, Bihrle S, Lacasse L. Autonomic stress reactivity and executive functions in successful and unsuccessful criminal psychopaths from the community. J Abnorm Psychol. 2001;110:423–432. [PubMed]
91. Krakowski M, Czobor P, Carpenter MD, et al. Community violence and inpatient assaults: neurobiological deficits. J Neuropsychiatry Clin Neurosci. 1997;9:549–555. [PubMed]
92. LaPierre D, Braun CMJ, Hodgins S. Ventral frontal deficits in psychopathy: neuropsychological test findings. Neuropsychologia. 1995;33:139–151. [PubMed]
93. Moffitt TE. Adolescence-limited and life-course-persistent antisocial behavior: a developmental taxonomy. Psychol Rev. 1993;100:674–701. [PubMed]
94. Raine A, Meloy JR, Bihrle S, Stoddard J, LaCasse L, Buchsbaum MS. Reduced prefrontal and increased subcortical brain functioning assessed using positron emission tomography in predatory and affective murderers. Behav Sci Law. 1998;16:319–332. [PubMed]
95. Seidenwurm D, Pounds TR, Globus A, Valk PE. Abnormal temporal lobe metabolism in violent subjects: correlation of imaging and neuropsychiatric findings. Am J Neuroradiol. 1997;18:625–631. [PubMed]
96. Soderstrom H, Hultin L, Tullberg M, Wikkelso C, Ekholm S, Forsman A. Reduced frontotemporal perfusion in psychopathic personality. Psychiatry Res. 2002;114:81–94. [PubMed]
97. Hoptman MJ. Neuroimaging studies of violence and antisocial behavior. J Psychiatr Pract. 2003;9:265–278. [PubMed]
98. De Oliveira-Souza R, Hare RD, Bramati IE, et al. Psychopathy as a disorder of the moral brain: fronto-temporo-limbic grey matter reductions demonstrated by voxel-based morphometry. Neuroimage. 2008;40:1202–1213. [PubMed]
99. Tiihonen J, Rossi R, Laakso MP, et al. Brain anatomy of persistent violent offenders: more rather than less. Psychiatry Res. 2008;163:201–212. [PubMed]
100. Raine A, Lencz T, Bihrle S, LaCasse L, Colletti P. Reduced prefrontal gray matter volume and reduced autonomic activity in antisocial personality disorder. Arch Gen Psychiatry. 2000;57:119–127. [PubMed]
101. Sapolsky RM. The frontal cortex and the criminal justice system. Philos Trans R Soc Lond B Biol Sci. 2004;359:1787–1796. [PMC free article] [PubMed]
102. De Brito SA, Mechelli A, Wilke M, et al. Size matters: Increased grey matter in boys with conduct problems and callous-unemotional traits. Brain. 2009;132:843–852. [PubMed]
103. Veit R, Flor H, Erb M, et al. Brain circuits involved in emotional learning in antisocial behavior and social phobia in humans. Neurosci Lett. 2002;328:233–236. [PubMed]
104. Finger EC, Marsh AA, Mitchell DG, et al. Abnormal ventromedial prefrontal cortex function in children with psychopathic traits during reversal learning. Arch Gen Psychiatry. 2008;65:586–594. [PMC free article] [PubMed]
105. Blair RJ. The amygdala and ventromedial prefrontal cortex in morality and psychopathy. Trends Cogn Sci. 2007;11:387–392. [PubMed]
106. Cohen L, Angladette L, Benoit N, Pierrot-Deseilligny C. A man who borrowed cars. Lancet. 1999;353:34. [PubMed]
107. Paradiso S, Robinson RG, Arndt S. Self-reported aggressive behavior in patients with stroke. J Nerv Ment Dis. 1996;184:746–753. [PubMed]
108. Pillmann F, Rohde A, Ullrich S, Draba S, Sannemüller U, Marneros A. Violence, criminal behavior, and the EEG: significance of left hemispheric focal abnormalities. J Neuropsychiatry Clin Neurosci. 1999;11:454–457. [PubMed]
109. Anderson SW, Bechara A, Damasio H, Tranel D, Damasio AR. Impairment of social and moral behavior related to early damage in human prefrontal cortex. Nat Neurosci. 1999;2:1031–1037. [PubMed]
110. Eslinger PJ, Flaherty-Craig CV, Benton AL. Developmental outcomes after early prefrontal cortex damage. Brain Cogn. 2004;55:84–103. [PubMed]
111. Bechara A, Tranel D, Damasio H. Characterization of the decision-making deficit of patients with ventromedial prefrontal cortex lesions. Brain. 2000;123:2189–2202. [PubMed]
112. Bechara A, Damasio AR, Damasio H, Anderson SW. Insensitivity to future consequences following damage to human prefrontal cortex. Cognition. 1994;50:7–15. [PubMed]
113. Blair RJR, Cipolotti L. Impaired social response reversal. A case of ‘acquired sociopathy’ Brain. 2000;123:1122–1141. [PubMed]
114. Ciaramelli E, Muccioli M, Ladavas E, di Pellegrino G. Selective deficit in personal moral judgment following damage to ventromedial prefrontal cortex. Soc Cogn Affect Neurosci. 2007;2:84–92. [PMC free article] [PubMed]
115. Cushman FA, Young LL, Hauser MD. The role of conscious reasoning and intuition in moral judgments: Testing three principles of permissible harm. Psychol Sci. 2006;17:1082–1089. [PubMed]
116. Damasio AR, Tranel D, Damasio H. Individuals with sociopathic behavior caused by frontal damage fail to respond autonomically to social stimuli. Behav Brain Res. 1990;41:81–94. [PubMed]
117. Koenigs M, Young L, Adolphs R, Tranel D, Cushman F, Hauser M, Damasio A. Damage to the prefrontal cortex increases utilitarian moral judgments. Nature. 2007;446:908–911. [PMC free article] [PubMed]
118. Shamay-Tsoory SG, Tomer R, Berger BD, Goldsher D, Aharon-Peretz J. Impaired “affective theory of mind” is associated with right ventromedial prefrontal damage. Cogn Behav Neurol. 2005;18:55–67. [PubMed]
119. Tranel D, Bechara A, Denburg NL. Asymmetric functional roles of right and left ventromedial prefrontal cortices in social conduct, decision-making, and emotional processing. Cortex. 2002;38:589–612. [PubMed]
120. Tranel D. “Acquired sociopathy”: the development of sociopathic behavior following focal brain damage. Prog Exp Pers Psychopathol Res. 1994:285–311. [PubMed]
121. Beer JS, Heerey EA, Keltner D, Scabini D, Knight RT. The regulatory function of self-conscious emotion: Insights from patients with orbitofrontal damage. J Pers Soc Psychol. 2003;85:594–604. [PubMed]
122. Beer JS, John OP, Scabini D, Knight RT. Orbitofrontal cortex and social behavior: integrating self-monitoring and emotion-cognition interactions. J Cong Neurosci. 2006;18:871–879. [PubMed]
123. Eslinger PJ, Grattan LM, Damasio AF. Developmental consequences of childhood frontal lobe damage. Arch Neurol. 1992;49:764–769. [PubMed]
124. Shamay-Tsoory SG, Tibi-Elhanany Y, Aharon-Peretz J. The green-eyed monster and malicious joy: the neuroanatomical bases of envy and gloating (schadenfreude) Brain. 2007;130:1663–1678. [PubMed]
125. Lough S, Hodges JR. Measuring and modifying abnormal social cognition in frontal variant frontotemporal dementia. J Psychosom Res. 2002;53:639–646. [PubMed]
126. Scarpa A, Raine A. Psychophysiology of anger and violent behavior. Psychiatr Clin North Am. 1997;20:375–394. [PubMed]
127. Zlotnick C. Antisocial personality disorder, affect dysregulation and childhood abuse among incarcerated women. J Personal Disord. 1999;13:90–95. [PubMed]
128. Brower MC, Price BH. Neuropsychiatry of frontal lobe dysfunction in violent and criminal behaviour: a critical review. J Neurol Neurosurg Psychiatry. 2001;71:720–726. [PMC free article] [PubMed]
129. Koenigs M, Tranel D. Irrational economic decision-making after ventromedial prefrontal damage: evidence from the ultimatum game. J Neurosci. 2007;21:951–956. [PMC free article] [PubMed]
130. Anderson SW, Barrash J, Bechara A, Tranel D. Impairments of emotion and realword complex behavior following childhood- or adult-onset damage to ventromedial prefrontal cortex. J Int Neuropsychol Soc. 2006;12:224–235. [PubMed]
131. Burgess PW, Alderman N, Forbes C, et al. The case for the development and use of “ecologically valid” measures of executive functions in experimental and clinical neuropsychology. J Int Neuropsychol Soc. 2006;12:194–209. [PubMed]
132. Saver JL, Damasio AR. Preserved access and processing of social knowledge in a patient with acquired sociopathy due to ventromedial frontal damage. Neuropsychologia. 1991;29:1241–1249. [PubMed]
133. Barrash J, Tranel D, Anderson SW. Acquired personality disturbances associated with bilateral damage to the ventromedial prefrontal region. Develop Neuropsychol. 2000;18:355–381. [PubMed]
134. Bechara A, Van Der Linden M. Decision-making and impulse control after frontal lobe injuries. Curr Opin Neurol. 2005;18:734–739. [PubMed]
135. Camille N, Coricelli G, Sallet J, Pradat-Diehl P, Duhamel JR, Sirigu A. The involvement of the orbitofrontal cortex in the experience of regret. Science. 2004;304:1167–1170. [PubMed]
136. Lough S, Kipps CM, Treise C, Watson P, Blair JR, Hodges JR. Social reasoning, emotion and empathy in frontotemporal dementia. Neuropsychologia. 2006;44:950–958. [PubMed]
137. Mah LW, Arnold MC, Grafman J. Deficits in social knowledge following damage to ventromedial prefrontal cortex. J Neuropsychiatry Clin Neurosci. 2005;17:66–74. [PubMed]
138. Neary D, Snowden J, Gustafson L, et al. Frontotemporal lobar degeneration: a consensus on clinical diagnostic criteria. Neurology. 1998;51:1546–1552. [PubMed]
139. Miller BL, Chang L, Mena I, Boone K, Lesser IM. Progressive right frontotemporal degeneration: clinical, neuropsychological and SPECT characteristics. Dementia. 1993;4:204–213. [PubMed]
140. Edwards-Lee T, Miller BL, Benson DF, et al. The temporal variant of frontotemporal dementia. Brain. 1997;120:1027–1040. [PubMed]
141. Mychack P, Kramer JH, Boone KB, Miller BL. The influence of right frontotemporal dysfunction on social behavior in frontotemporal dementia. Neurology. 2001;56(Suppl 4):S11–S15. [PubMed]
142. Perry RJ, Rosen HR, Kramer JH, Beer JS, Levenson RL, Miller BL. Hemispheric dominance for emotions, empathy and social behaviour: Evidence from right and left handers with frontotemporal dementia. Neurocase. 2001;7:145–160. [PubMed]
143. Mendez MF, Lim GTH. Alterations in the sense of ‘humanness’ in right hemisphere predominant FTD patients. Cogn Behav Neurol. 2004;17:133–138. [PubMed]
144. Mendez MF, Lauterbach E, Sampson S. ANPA Committee on Research. An Evidence- Based Review of the Psychopathology of Frontotemporal Dementia: A Report of the ANPA Committee on Research. J Neuropsychiatry Clin Neurosci. 2008;20:130–149. [PubMed]
145. Mendez MF, Chen AK, Shapira JS, Miller BL. Acquired sociopathy and frontotemporal dementia. Dement Geriatr Cogn Disord. 2005;20:99–104. [PubMed]
146. Gustafson L. Clinical picture of frontal lobe degeneration of non-Alzheimer type. Dementia. 1993;4:143–148. [PubMed]
147. Lynch T, Sano M, Marder KS, et al. Clinical characteristics of a family with chromosome 17-linked disinhibition-dementia-parkinsonism-amyotrophy complex. Neurology. 1994;44:1878–1884. [PubMed]
148. Miller BL, Darby A, Benson DF, Cummings JL, Miller MH. Aggressive, socially disruptive and antisocial behaviour associated with fronto-temporal dementia. Br J Psychiatry. 1997;170:150–154. [PubMed]
149. Mendez MF, Chow T, Ringman J, Twitchell G, Hinkin CH. Pedophilia and disturbances of the temporal lobes. J Neuropsychiatry Clin Neurosci. 2000;12:71–76. [PubMed]
150. Mendez MF, Anderson E, Shapira JS. An investigation of moral judgment in frontotemporal dementia. Cogn Behav Neurol. 2005;18:193–197. [PubMed]
151. Rankin KP, Kramer JH, Miller BL. Patterns of cognitive and emotional empathy in frontotemporal lobar degeneration. Cogn Behav Neurol. 2005;18:28–36. [PubMed]
152. Rankin KP, Kramer JH, Mychack P, Miller BL. Double dissociation of social functioning in frontotemporal dementia. Neurology. 2003;60:266–271. [PMC free article] [PubMed]
153. Rankin KP, Rosen HJ, Kramer JH, et al. Right and left medial orbitofrontal volumes show an opposite relationship to agreeableness in FTD. Dement Geriatr Cogn Disord. 2004;17:328–332. [PMC free article] [PubMed]
154. Gorno-Tempini ML, Rankin KP, Woolley JD, Rosen HJ, Phengrasamy L, Miller BL. Cognitive and behavioral profile in a case of right anterior temporal lobe neurodegeneration. Cortex. 2004;40:631–644. [PubMed]
155. Rankin KP, Gorno-Tempini ML, Allison SC, et al. Structural anatomy of empathy in neurodegenerative disease. Brain. 2006;129:2945–2956. [PMC free article] [PubMed]
156. Snowden JS, Gibbons ZC, Blackshaw A, et al. Social cognition in frontotemporal dementia and Huntington’s disease. Neuropsychologia. 2003;41:688–701. [PubMed]
157. Gregory C, Lough S, Stone V, et al. Theory of mind in patients with frontal variant frontotemporal dementia and Alzheimer’s disease: theoretical and practical implications. Brain. 2002;125:752–764. [PubMed]
158. Lough S, Gregory C, Hodges JR. Dissociation of social cognition and executive function in frontal variant frontotemporal dementia. Neurocase. 2001;7:123–130. [PubMed]
159. Hauser MD. The liver and the moral organ. Soc Cogn Affect Neurosci. 2006;1:214–220. [PMC free article] [PubMed]
160. Mikhail J. Universal moral grammar: theory, evidence and the future. Trends Cogn Sci. 2007;11:143–152. [PubMed]
161. Pijnenburg YA. The roots of social inappropriateness in frontotemporal dementia. J Neurol Neurosurg Psychiatry. 2007;78:441. [PMC free article] [PubMed]
162. Moll J, de Oliveira-Souza R, Garrido GJ, et al. The self as a moral agent: linking the neural bases of social agency and moral sensitivity. Soc Neurosci. 2007;2:336–352. [PubMed]
163. Shallice T. ‘Theory of mind’ and the prefrontal cortex. Brain. 2001;124:247–248. [PubMed]
164. Stone VE, Baron-Cohen S, Calder A, Keane J, Young A. Acquired theory of mind impairments in individuals with bilateral amygdalar lesions. Neuropsychologia. 2003;41:209–220. [PubMed]
165. Stuss DT, Gallup GG, Jr, Alexander MP. The frontal lobes are necessary for theory of mind. Brain. 2001;124:279–286. [PubMed]
166. Tankersley D, Stowe CJ, Huettel SA. Altruism is associated with an increased neural response to agency. Nat Neurosci. 2007;10:150–151. [PubMed]
167. Calarge C, Andreasen NC, O’Leary DS. Visualizing how one brain understands another: a PET study of theory of mind. Am J Psychiatry. 2003;160:1954–1964. [PubMed]
168. Borum R, Fulero SM. Empirical research on the insanity defense and attempted reforms: evidence toward informed policy. Law Hum Behav. 1999;23:375–393. [PubMed]
169. Goodenough OR, Prehn K. A neuroscientific approach to normative judgment in law and justice. Philos Trans R Soc Lond B Biol Sci. 2004;359:1709–1726. [PMC free article] [PubMed]