psychopathy amygdala.pdf

Opinion

TRENDS in Cognitive Sciences Vol.11 No.9

The amygdala and ventromedial

prefrontal cortex in morality and

psychopathy

R.J.R. Blair

Mood and Anxiety Program, National Institute of Mental Health, Department of Health and Human Services, 15K North Drive,

MSC 2670, Bethesda, MD 20892, USA

Recent work has implicated the amygdala and

ventromedial prefrontal cortex in morality and, when

dysfunctional, psychopathy. This model proposes that

the amygdala, through stimulus-reinforcement learning,

enables the association of actions that harm others with

the aversive reinforcement of the victims’ distress. Con-

sequent information on reinforcement expectancy, fed

forward to the ventromedial prefrontal cortex, can guide

the healthy individual away from moral transgressions.

In psychopathy, dysfunction in these structures means

that care-based moral reasoning is compromised and

the risk that antisocial behavior is used instrumentally to

achieve goals is increased.

particular, moral transgressions are judged to be less

rule-contingent than are conventional transgressions;

individuals are less likely to state that moral, rather than

conventional, transgressions are permissible in the

absence of prohibiting rules [3] .

Early models of moral socialization suggested unitary

accounts of social rule learning, stressing, for example,

either punishment [4] or cultural transmission [5] . How-

ever, such accounts struggle to explain the existence of the

moral–conventional distinction [3] . Moreover, the social

consequences of moral and conventional transgressions

differ; caregivers are more likely to refer transgressors

of moral rules to the consequences of their actions for

the victim and transgressors of conventional rules to the

rules themselves or the sanctions against prohibition [6] .

Early discussions of the development of the moral–

conventional distinction itself suggested that it emerged

as a function of abstract reasoning processes [3] . They

made two clear predictions. First, populations with

impaired abstract reasoning should fail to develop the

moral–conventional distinction. However, no data have

been presented in support of this position. Moreover, at

least one population with pronounced executive function

impairment, children with autism, pass the moral–conven-

tional distinction [7] . Second, populations who show

impairment in the development of the moral–conventional

distinction should show impairment in abstract reasoning

and/or executive dysfunction. Individuals with psychopa-

thy show signiﬁcantly less of a moral–conventional dis-

tinction than do healthy individuals [8] . However, before

considering executive dysfunction in this population, I will

brieﬂy consider the nature of psychopathy.

Introduction

Care-based morality can be considered as those forms of

moral reasoning that concern actions that harm others.

Other forms of moral cognition might exist but these will

not be considered in detail here ( Box 1 ). Why do we care

whether anyone else is hurt? Perhaps equally importantly,

why do some people, individuals with psychopathy, care

less?

Here, I make reference to the functional roles of the

amygdala and ventromedial prefrontal cortex (vmPFC) in

the learning and use of reinforcement expectancies (expec-

tancies of reward or punishment). I argue that (i) the

integrated functioning of these systems enables the basics

of care-based morality; and (ii) dysfunction within these

regions in psychopathy means that reinforcement-based

decision making, including moral decision making, is

impaired.

An early developmental indication of care-based

morality: the moral–conventional distinction

Moral transgressions (e.g. one person hitting another) are

deﬁned by their consequences for the rights and welfare of

others. Social conventional transgressions are deﬁned as

violations of the behavioral uniformities that structure

social interactions within social systems (e.g. dressing in

opposite gender clothing). Healthy individuals distinguish

conventional and moral transgressions in their judgments

from the age of 39 months [1] and across cultures [2] .In

Psychopathy

Psychopathy is a developmental disorder [9] that involves

emotional dysfunction, characterized by reduced guilt,

empathy and attachment to signiﬁcant others, and anti-

social behavior including impulsivity and poor behavioral

control [10] . It is not equivalent to the Diagnostic and

Statistical Manual of Mental Disorders, 4th Edition

(DSM-IV) psychiatric diagnoses of conduct disorder (CD)

and antisocial personality disorder (ASPD), which focus

only on the presence of antisocial behavior rather than any

form of functional impairment that might be causally

related to its emergence.

Corresponding author: Blair, R.J.R. ( blairj@intra.nimh.nih.gov ).

Available online 17 August 2007.

www.sciencedirect.com 1364-6613/$ – see front matter . Published by Elsevier Ltd. doi: 10.1016/j.tics.2007.07.003

388

Opinion

TRENDS in Cognitive Sciences Vol.11 No.9

Box 1. Multiple moralities?

function of abstract reasoning processes [3] , we might

believe that individuals with psychopathy will show

impairment in abstract reasoning and/or executive dys-

function. However, no data suggest that they do. Indeed,

individuals with psychopathy show executive dysfunction

only if the executive function has an affective component

[15] .

Considerable recent work has suggested the importance

of emotional responses for moral development [8,16–19] .A

speciﬁc version of this view suggests that healthy individ-

uals are predisposed to ﬁnd the distress of others aversive

and that we learn to avoid actions associated with this

distress (i.e. acts that harm others) [8] . According to this

view, we distinguish between moral and conventional

transgressions because only moral transgressions are

associated with the distress of others, and this association

remains intact whether there is a rule prohibiting the

action or not. This position predicts that individuals who

show reduced processing of the distress of others should

show a reduced moral–conventional distinction. Individ-

uals with psychopathy show reduced autonomic responses

to the distress of others [20,21] and reduced recognition of

sad and fearful expressions [22,23] . They also, of course,

make signiﬁcantly less of a moral–conventional distinction

than do comparison individuals [8] .

The above suggests that a factor crucial for care-based

moral socialization is appropriate emotional responding to

the distress of others. If this is lacking, as is the case in

psychopathy, there should be interference with socializa-

tion. Individuals with psychopathy are less inﬂuenced by

parental socialization strategies than are healthy individ-

uals [24] . This suggests, in turn, that neural systems

implicated in psychopathy might be importantly involved

in moral development.

It seems likely that there are other forms of ‘moral’ reasoning than

care-based morality, and that these rely on overlapping or perhaps

independent neurocognitive architectures. These might even be

intact in individuals with psychopathy. The two principal possibi-

lities relate to disgust-based moral reasoning and reasoning about

conventional transgressions.

Disgust has been considered a powerful emotive force for

attitudes towards a variety of transgressions, particularly those

concerning sexual activity [51] . It seems likely that we use disgust

expressions in social interaction in a similar fashion to our use of

fearful expressions. Fearful expressions rapidly communicate to

others that the object to which the expression is displayed is

threatening [52] . Fearful expressions activate the amygdala, parti-

cularly if there is a clear object associated with the expression to

enable the initiation of emotional learning [42] . It is likely that

disgust expressions initiate taste-aversion learning. Taste-aversion

learning is mediated by structures such as the insula, which are

activated by disgusted expressions [53] . This suggests that we

might learn disgust reactions to particular transgressions following

the display of disgust expressions directed towards individuals (or

descriptions of individuals) engaged in particular transgressions.

But what about the vmPFC? The vmPFC has a major role in the

representation of reinforcement outcomes. Given that there are

connections between the insula and the vmPFC, it is plausible that

disgust-based valenced outcomes might also be represented in the

vmPFC. However, this remains to be empirically demonstrated.

Although there have been isolated reports of impairment in the

processing of disgust in individuals with psychopathy [54] , this is

not typically seen [22,23] . This suggests the possibility that the

processing of disgust- and potentially disgust-based moral reason-

ing might be intact in individuals with psychopathy. If this is shown,

it will be clear evidence of dissociation between different forms of

emotion-based moral reasoning.

There is already evidence of a dissociation between impaired

care-based moral reasoning and intact social conventional reason-

ing in individuals with psychopathy. Although individuals with

psychopathy show impairment in care-based moral reasoning, as

indexed by their performance on the moral–conventional distinction

task, they seem to be intact in the detection and appropriate rating

of more conventional transgressions and faux pas [55] . Conven-

tional transgressions elicit anger in observers, and both conven-

tional transgressions and induced anger elicit activity in the

ventrolateral prefrontal cortex [56,57] . This region is implicated in

altering current behavior, particularly when a prepotent response is

already engaged [35] . Current data indicate that this region might

not be dysfunctional in individuals with psychopathy.

Dysfunctional neural systems in psychopathy

The neuropsychological literature on psychopathy has

identiﬁed two core neural regions that seem to be dysfunc-

tional in psychopathy: the amygdala and the vmPFC.

Psychopathy is associated with a series of core functional

impairments: deﬁcits in aversive conditioning, the aug-

mentation of the startle reﬂex by visual threat primes

and fearful expression recognition. These impairments

are also seen following lesions of the amygdala [25] .In

addition, psychopathy is associated with problems in

response reversal and in tasks such as the Iowa gambling

task. These impairments are seen following lesions of the

vmPFC [26,27] . However, it is important to recognize that

psychopathy is not a neurological condition. Psychopathy

is not associated with a lesion to a particular region, nor

have all functions mediated by any particular region been

shown to be compromised. Indeed, although the above

functional impairments associated with amygdala dys-

function are seen in psychopathy, other capacities in which

the amygdala seems to be implicated, such as determining

an internal state from information provided by the eyes,

are intact [25] .

The neuroimaging literature on psychopathy has found

that individuals with psychopathy show reduced activation

of both the amygdala and the rostral anterior cingulate

cortex/vmPFC in response to emotional words in the context

A distinction has long been drawn between reactive and

instrumental aggression [11] . Reactive aggression occurs

as an explosive response to threat or frustration and is not

goal directed (e.g. road rage) [11] . Instrumental aggression

is aggression used to achieve a goal (e.g. a mugging) [11] .

Various neurological and psychiatric conditions are at

heightened risk for reactive aggression (e.g. acquired socio-

pathy and childhood bipolar disorder [12,13] ). Individuals

with psychopathy are remarkable, in that they show an

increased risk not only for reactive aggression, but also for

instrumental aggression [14] . There is something about

psychopathy that increases the risk that the individual will

use aggression and other antisocial behaviors, actions that

harm others, to achieve their goals.

Psychopathy and morality

Individuals with psychopathy show signiﬁcantly less of a

moral–conventional distinction than do healthy individ-

uals [8] . If themoral–conventional distinction emerged as a

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Opinion

TRENDS in Cognitive Sciences Vol.11 No.9

389

of emotional memory paradigms [28] and during aversive

conditioning [29] . Work with subclinical populations has

found that individuals with psychopathic traits show

reduced amygdala responses to emotional expressions

[30] and less amygdala and vmPFC differentiation in

responding when making cooperation relative to defection

choices in a prisoner’s dilemma paradigm [31] . The neuroi-

maging literature onpsychopathyhas also sometimes impli-

cated other structures – for example, the superior temporal

cortex and dorsal anterior cingulate cortex (dACC) [32] .

However, these ﬁndings remain unsubstantiated by neu-

ropsychological ﬁndings (and therefore we cannot be sure

that they do not simply reﬂect reduced input from core

regions). For example, impaired Stroop performance would

be a clear predictor of dACC dysfunction, but this is not seen

in psychopathy [25] .

Of course, it is impossible to determine on the basis of

these imaging results whether the reduced orbital frontal

cortex (OFC)/vmPFC activity reﬂects dysfunction in this

region or simply reduced input to this region from the

amygdala. A number of animal studies have stressed the

importance of the interaction between the amygdala and

the OFC/vmPFC [33] . Damage to the amygdala has a

detrimental impact on OFC functioning [33] . However, it

is important to note that animal studies also suggest that

early amygdala dysfunction disrupts the appropriate de-

velopment of the OFC and vmPFC [34] . Moreover, mol-

ecular candidates with respect to the genetic basis for

psychopathy innervate both the amygdala and the OFC

and vmPFC ( Box 2 ). It thus seems likely that psychopathy

is associated with both amygdala and OFC and vmPFC

dysfunction. However, to demonstrate this conclusively, it

will be necessary to demonstrate anomalous activity in the

OFC and vmPFC in a task which does not implicate the

amygdala.

Box 2. Social and genetic causes of psychopathy

Definitive answers with respect to the fundamental etiology of

psychopathy cannot yet be provided. However, some potential basic

causes seem less plausible than others. For example, there have

been suggestions that psychopathy might be due to early physical

or sexual abuse or neglect [58] . However, studies have examined the

impact of such stressors on brain development. Animal studies have

shown that neglect and other stressors increase emotional and

amygdala responsiveness to threatening stimuli [59,60] . Similarly,

in humans, early physical or sexual abuse is a significant risk factor

for the emergence of post-traumatic stress disorder (PTSD), which is

associated with increased emotional and amygdala responsiveness

[61] . Increased emotional and amygdala responsiveness is the

opposite of the pathology seen in psychopathy.

Data suggest a genetic contribution to the disorder [62,63] . For

example, in a recent large study of 3500 twin pairs, callous–

unemotional traits were shown to be strongly heritable (67%

heritability) at both seven and nine years of age [63] . However, an

understanding of psychopathy at the molecular genetic level

remains in its infancy. Suggestive data are provided by recent

investigations of the impact of different genetic polymorphisms on

neural and behavioral responding. For example, several studies

have reported that individuals who are ll homozygotes for the 5-

hydroxytryptamine transporter (5-HTTLPR) gene show a signifi-

cantly reduced amygdala response to emotional expressions

relative to those who have the short-form polymorphism of the

gene [64] . In addition, such individuals show behavioral impairment

on some emotional learning tasks reliant on the amygdala [65] .Itis

possible that there is an array of genes whose polymorphisms

increase or decrease emotional and amygdala responsiveness. The

basic genetic risk for psychopathy might emerge if an individual

possesses a sufficient number of polymorphisms predisposing

towards reduced emotional and amygdala responsiveness.

In short, psychopathy is under considerable genetic influence and

has yet to be reliably related to a social basic cause. However,

although social factors might not be casual, they do influence its

manifestation. For example, socioeconomic status (SES) is asso-

ciated with the emergence of the full syndrome; it is significantly

less likely to appear in individuals of higher SES [66] . Reduced SES

does not predispose towards reduced emotional and amygdala

responsiveness; indeed, as a factor it is unrelated to the emotional

component of psychopathy [67] . However, reduced SES does

supply a motive: a lack of finances. Reduced SES increases the risk

for antisocial behavior in individuals, including those with psycho-

pathy.

The amygdala, vmPFC and morality

The amygdala and vmPFC are implicated not only in the

emergence of psychopathy, but also in moral reasoning.

Thus, the amygdale, and particularly the vmPFC, have

frequently been identiﬁed in neuroimaging studies of

moral reasoning [18,19,35] ( Figure 1 ). For example, Greene

et al. [19] reported increased vmPFC activity in response to

personal as opposed to impersonal moral choices, and the

difference between these two situations related effectively

to the salience of the victim. Similarly, in more recent

work, Luo et al. [35] demonstrated increased amygdala

and vmPFC activity in response to more severe relative to

less severe moral transgressions.

The moral reasoning imaging literature has not

precisely speciﬁed the functional roles of the amygdala

and vmPFC [36] . In one of the more formalized accounts

[37] , it has been argued that the vmPFC is implicated in

representing social and emotional structured event com-

plexes (SEC). These SECs are considered to be long-term

memories of event sequences that guide the perception and

execution of goal-oriented activities, such as going to a

concert or giving a dinner party. However, as the authors

note [37] , the SEC framework does not predict how PFC

regions interact with limbic areas and other cortical

regions.

A cognitive neuroscience approach to care-based

morality and psychopathy

The amygdala

The amygdala is crucial for stimulus-reinforcement

learning [38,39] . This learning enables representations

of conditioned stimuli within temporal cortex to be linked

to emotional responses mediated by the amygdala and

other structures ( Figure 2 ). In short, the amygdala enables

the individual to learn the goodness and badness of objects

and actions. Moreover, because the relationship between

the amygdala and the temporal cortex is reciprocal [40] ,

this learning enables the amygdala to inﬂuence attention

[41] . In humans and other primates, fearfulness serves as a

reinforcer, stimuli associated with expression are avoided,

and this type of stimulus-reinforcement learning relies on

the amygdala too [42,43] .

The argument here is that learning the basics of

care-based morality – learning that some actions harm

others and because of this are to be avoided – relies on this

crucial role of the amygdala in stimulus-reinforcement

learning. Because of this, the amygdala is often seen to

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Opinion

TRENDS in Cognitive Sciences Vol.11 No.9

Figure 1. Neural responses to increasing intensity of moral transgressions and positive actions. Both the amygdala and vmPFC show increased activity in response to

images of high-intensity illegal content (H-il; scenes of interpersonal violence) and high-intensity legal content (H-le; skydiving) relative to images of low intensity illegal

content (L-il; scenes of property damage) and low intensity legal content (L-le; playing the guitar). Reproduced, with permission, from Ref. [35] .

respond in moral reasoning studies [18,44] . Psychopathy

represents an extreme case, where aversive conditioning

[29] and the response to others’ fear [22] is profoundly

impaired. These impairments mean that the individual

with psychopathy is signiﬁcantly more difﬁcult to socialize

[24] . Interestingly, these impairments, at least when they

are indirectly measured by the temperament variable

fearfulness, are also associated with weaker conscience

development in healthy populations [45] .

Figure 2. Core brain regions implicated in the basis of care-based morality and,

when dysfunctional, psychopathy. These regions include the superior temporal

cortex (STC) [(a,b); yellow], the amygdala [(a,b); red] and the vmPFC [(c,b); blue].

(b) Stimulus-reinforcement learning enables the association of representations of

conditioned stimuli within the temporal cortex, including the STC, to be linked to

emotional responses mediated by the amygdala. Of particular importance for care-

based morality is associating actions that cause harm with the aversive

consequences of the victim’s fear and sadness. The connections between

the amygdala and temporal cortex are reciprocal, enabling the amygdala to

influence attention. Reduced emotion-based attention is seen in psychopathy [70] .

The amygdala sends reinforcement expectancy information to the vmPFC. For

care-based morality, this is particularly important when the reinforcement

expectancy information concerns the anticipated distress of another. This

information should guide the healthy information away from the action about to

be committed. However, it is argued that reduced information from the amygdale,

in addition to dysfunction within the vmPFC in psychopathy, means that this

guiding role of the vmPFC functions poorly.

The vmPFC

The amygdala sends outputs not only to the temporal and

visual cortex and regions mediating autonomic responses,

but also forward to the vmPFC. Animal studies have

produced a relatively precise insight into the functional

role of the interaction between the amygdala and the OFC

and vmPFC [33] . In particular, Schoenbaum and Roesch

implicate the amygdala and vmPFC in stimulus–outcome

processing. The basic suggestion is that the amygdala

provides reinforcement expectancy information to the

OFC and vmPFC; the latter then represents this infor-

mation. Human neuroimaging work has conﬁrmed the role

of the OFC and vmPFC in outcome representation [46] .

The individual’s ‘automatic moral attitude’ to a moral

transgression thus involves activation of the amygdala by

the conditioned stimulus that is the individual’s representa-

tion of themoral transgression. The amygdala thenprovides

expected reinforcement (both positively and negatively

valenced) information, which is represented as a valenced

outcome within the vmPFC. Other systems then use

this information to select appropriate responses [47] .This

information is crucial for reinforcement expectancy-based

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Opinion

TRENDS in Cognitive Sciences Vol.11 No.9

391

Box 3. Treatment

individual might consider an act bad and to be avoided, it

does not stipulate the processes necessary to consider that

action immoral [17] . Potentially more crucially, it still

remains unknown how to translate our increased under-

standing of the pathophysiology of psychopathy into viable

treatment options for individuals with this disorder

( Box 3 ).

There are successful psychopharmacological and psychotherapeu-

tic programs that reduce aggression associated with many psychia-

tric conditions [68] . However, currently, psychopathy is regarded as

untreatable [67] .

Two main lines of data provide interesting potential treatment

hypotheses. The first concerns recent work on genetic polymorph-

isms ( Box 2 ). By understanding the molecular genetics of psycho-

pathy, it should be possible to develop theoretically principled

pharmacological treatments. However, this work remains in its

infancy. The second and more immediate line is generated by our

increased understanding of the pathophysiology of psychopathy; in

particular, that it is an emotional disorder associated with reduced

activity in the amygdala and associated structures (see main text).

This is useful information because there are a variety of known

pharmacological agents that modulate amygdala activity. Such

agents target, for example, the serotonergic or noradrenergic

systems. Much clinical research has focused on the identification

of agents that downregulate emotional responding and activity

within the amygdala and associated structures. Such agents have

been used successfully to treat depression and PTSD, disorders

whose primary pathology is increased activity within the amygdala

and associated structures [61] . It is possible that the inverse of these

treatments might be successfully applied to psychopathy. For

example, agents that reduce noradrenergic activity and conse-

quently amygdala activity seem useful in treating PTSD [69] .

Perhaps psychopathy could be treated by agents that increase

noradrenergic activity?

Although pharmacological agents are likely to be necessary for

the successful treatment of psychopathy, they are unlikely to be

sufficient. Common treatments for PTSD are designed to reduce

emotional responding so that cognitive behavior-based extinction

techniques can reduce the emotional significance of the original

trauma. Pharmacological agents might be successful in increasing

the emotional response and activity within the amygdala and

associated structures in individuals with psychopathy. However,

such agents will need to be coupled with cognitive behavior-based

treatments designed to associate actions that hurt others with an

emotional response to the victim’s distress.

Acknowledgements

This work was supported by the Intramural Research Program of the

National Institutes of Health: National Institute of Mental Health.

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decisionmaking, includingmoral reasoning. The disruption

in the amygdala and vmPFC in psychopathy means that

moral and other forms of reinforcement-baseddecisionmak-

ing are disrupted [8,48] . Moreover, a recent neuropsycholo-

gical study demonstrated that vmPFC damage disrupts the

avoidance of actions leading to emotionally aversive con-

sequences (e.g. actively killing another person) shown by

healthy individuals in moral reasoning paradigms [49] .

Interestingly, these systems are recruited in real (virtual)

world moral decision making. Both the amygdala and

vmPFC show signiﬁcantly greater activity during appro-

priate decisions to heal wounded humans or kill attacking

monsters than during less appropriate decisions to heal

attacking monsters and kill humans [50] .

Conclusions

In summary, the model developed here provides a

conceptual framework for understanding the functional

contribution of the amygdala and vmPFC to the basis of

care-based morality and, when dysfunctional, psychopa-

thy. My main argument is that the amygdala enables the

forms of learning necessary to care about the welfare of

others, and, together with the vmPFC, enables this infor-

mation to inform moral decision making. However, many

questions remain unanswered. Although the functional

response described enables an understanding of how the

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