c o r t e x x x x ( 2 0 1 4 ) 1 e9

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Emotional detachment in psychopathy:

Involvement of dorsal default-mode connections

Arjun Sethi a,b,*, Sarah Gregory b, Flavio Dell'Acqua a,

Eva Periche Thomas a, Andy Simmons c, Declan G.M. Murphy b,
Sheilagh Hodgins d, Nigel J. Blackwood b,1 and Michael C. Craig a,b,1
a
NatBrainLab, London, UK
b
Department of Forensic and Neurodevelopmental Sciences, Institute of Psychiatry, London, UK
c
NIHR Biomedical Research Centre for Mental Health, Institute of Psychiatry, King's College London, London, UK
d
Department de Psychiatrie, Universite de Montreal, Canada

article info abstract

Article history: Criminal psychopathy is defined by emotional detachment [Psychopathy Checklist e

Received 29 November 2013 Revised (PCL-R) factor 1], and antisocial behaviour (PCL-R factor 2). Previous work has
Reviewed 31 March 2014 associated antisocial behaviour in psychopathy with abnormalities in a ventral temporo-
Revised 3 July 2014 amygdala-orbitofrontal network. However, little is known of the neural correlates of
Accepted 28 July 2014 emotional detachment. Imaging studies have indicated that the ‘default-mode network’
Published online xxx (DMN), and in particular its dorsomedial (medial prefrontal e posterior cingulate)
component, contributes to affective and social processing in healthy individuals.
Keywords: Furthermore, recent work suggests that this network may be implicated in psychopathy.
Psychopathy However, no research has examined the relationship between psychopathy, emotional
Emotional detachment detachment, and the white matter underpinning the DMN. We therefore used diffusion
Default-mode network tensor imaging (DTI) tractography in 13 offenders with psychopathy and 13 non-offenders
Cingulum to investigate the relationship between emotional detachment and the microstructure of
Diffusion MRI white matter connections within the DMN. These included the dorsal cingulum (containing
the medial prefrontal e posterior cingulate connections of the DMN), and the ventral
cingulum (containing the posterior cingulate e medial temporal connections of the DMN).
We found that fractional anisotropy (FA) was reduced in the left dorsal cingulum in the
psychopathy group (p ¼ .024). Moreover, within this group, emotional detachment was
negatively correlated with FA in this tract portion bilaterally (left: r ¼ .61, p ¼ .026; right:
r ¼ .62, p ¼ .023). These results suggest the importance of the dorsal DMN in the emotional
detachment observed in individuals with psychopathy. We propose a ‘dual-network’
model of white matter abnormalities in the disorder, which incorporates these with pre-
vious findings.
© 2014 Elsevier Ltd. All rights reserved.

* Corresponding author. NatBrainLab, Department of Forensic and Neurodevelopmental Sciences, Institute of Psychiatry, 16 De Cres-
pigny Park, London, SE5 8AF, UK.
E-mail address: arjun.sethi@kcl.ac.uk (A. Sethi).
1
Joint last authors.
http://dx.doi.org/10.1016/j.cortex.2014.07.018
0010-9452/© 2014 Elsevier Ltd. All rights reserved.

Please cite this article in press as: Sethi, A., et al., Emotional detachment in psychopathy: Involvement of dorsal default-mode
connections, Cortex (2014), http://dx.doi.org/10.1016/j.cortex.2014.07.018
2 c o r t e x x x x ( 2 0 1 4 ) 1 e9
1. Introduction
Criminal psychopathy is characterised by antisocial behav-
iour and a constellation of affective and interpersonal traits
including callousness, shallow affect, and manipulativeness
(R. D. Hare, 1991). These behavioural characteristics have a
significant negative impact on society. For instance, in-
dividuals with psychopathy constitute approximately 15e20%
of the prison population, and commit a disproportionate
number of violent and recidivistic offences (Hart & Hare, 1997;
Hemphill, Hare, & Wong, 1998) that cost the USA over $460
billion a year (Kiehl & Hoffman, 2011). It has been proposed
that psychopathy is a disorder of the ‘paralimbic’ system
(Kiehl, 2006), including structures such as the amygdala and
ventromedial prefrontal cortex (Blair, 2008).
Investigation into the neural basis of psychopathy has
been facilitated by the development of in vivo brain imaging
and reliable, well-validated, instruments that permit quanti-
fication of specific traits and behavioural tendencies. Factor
analysis of these traits, using the Psychopathy Checklist e
Revised (PCL-R), suggests that they can be divided into di-
mensions of ‘emotional detachment’ (factor 1) and ‘antisocial
behaviour’ (factor 2) (Hare, 1991, 2003; Hare et al., 1990; Harpur,
Hakstian, & Hare, 1988; Harpur, Hare, & Hakstian, 1989). Factor
2 antisocial scores have been reported to be negatively
correlated with the microstructure of the right uncinate
fasciculus (Craig et al., 2009), a ventral limbic tract connecting
the anterior temporal cortex and amygdala with orbitofrontal
regions (Catani & Thiebault De Schotten, 2012). The relation-
ship between psychopathy/antisocial behaviour and the un-
cinate has since been confirmed (Motzkin, Newman, Kiehl, &
Koenigs, 2011; Sundram et al., 2012). Moreover, the associa-
tion between specific regions within this network and psy-
chopathy and antisocial behaviour is supported by an
increasing number of neuropsychological (Blair, Colledge,
Murray, & Mitchell, 2001; Budhani & Blair, 2005; Budhani,
Richell, & Blair, 2006; Levenston, Patrick, Bradley, & Lang,
2000), lesion (Barrash, Tranel, & Anderson, 2000; Blair &
Cipolotti, 2000; Damasio, Grabowski, Frank, Galaburda, &
Damasio, 1994; Harlow, 1993, 1999; Kluver & Bucy, 1997;
Saver & Damasio, 1991), stimulation (King, 1961) and in vivo
brain imaging studies (Boccardi et al., 2011; Kiehl et al., 2001;
Raine, Buchsbaum, & LaCasse, 1997; Raine, Lencz, Bihrle,
LaCasse, & Colletti, 2000; Veit et al., 2002).
These prior studies have been important first steps in un-
derstanding psychopathy. However, antisocial behaviour is
not specific to psychopathic individuals (Harpur, et al., 1989;
Skeem & Cooke, 2010), and it is emotional detachment (fac-
tor 1) that differentiates psychopathic personality from the
broader diagnosis of Antisocial Personality Disorder. More-
over, emotional detachment in adults with psychopathy is
presumed to reflect a heritable developmental trajectory from
callous-unemotional traits in childhood (Barry et al., 2000;
Forsman, Lichtenstein, Andershed, & Larsson, 2008; Frick,
Kimonis, Dandreaux, & Farell, 2003; Frick & Viding, 2009;
Viding, Blair, Moffitt, & Plomin, 2005; Wootton, Frick,
Shelton, & Silverthorn, 1997). Investigating the neural corre-
lates of factor 1 traits is therefore likely to be of central
importance to understanding the neurodevelopment of
Please cite this article in press as: Sethi, A., et al., Emotional detachment in psychopathy: Involvement of dorsal default-mode
connections, Cortex (2014), http://dx.doi.org/10.1016/j.cortex.2014.07.018
psychopathic personality. Mounting evidence suggests that
the ‘default-mode’ network (DMN) is linked to psychopathy,
and may be related to emotional detachment in the disorder.
The DMN is a subdivision of the limbic system that has
largely been associated with introspective and self-referent
processing (Gusnard, Akbudak, Shulman, & Raichle, 2001;
Johnson et al., 2006; Kelley et al., 2002). This network con-
sists of a set of regions that are active and functionally
intercorrelated under resting-state conditions (Raichle et al.,
2001). These include the posterior cingulate cortex (PCC), the
medial prefrontal cortex (mPFC), the medial temporal lobe
(MTL), and the angular gyrus (Fox et al., 2005; Fransson, 2005;
Raichle et al., 2001; Shulman et al., 1997). These regions are of
specific interest due to their overlap with areas involved in
affective processing (Kiehl et al., 2001; Maddock, Garrett, &
Buonocore, 2003). This network has also been strongly impli-
cated in social processing (Buckner, Andrews-Hanna, &
Schacter, 2008; Vollm et al., 2006) and moral judgement
(Greene, Sommerville, Nystrom, Darley, & Cohen, 2001;
Harrison et al., 2008). The DMN is therefore well placed to
play an important role in the profound emotional detachment
in psychopathy.
The relevance of the DMN to psychopathy is further sup-
ported by recent functional MRI studies that have reported
abnormal activation and connectivity within this network
among men with psychopathy (Glenn, Raine, & Schug, 2009;
Motzkin, et al., 2011; Pujol et al., 2011). Similarly, structural
imaging studies of both adults with psychopathy (Boccardi
et al., 2011; Ermer, Cope, Nyalakanti, Calhoun, & Kiehl, 2012;
Gregory et al., 2012; de Oliveira-Souza et al., 2008; Yang,
Raine, Colletti, Toga, & Narr, 2009) and boys with conduct
disorder and callous-unemotional traits (De Brito et al., 2009;
Rijsdijsk et al., 2010) have reported abnormal grey matter
volume in DMN regions. Importantly, a preliminary study has
reported that the degree of functional connectivity within a
network containing DMN regions was related to emotional
detachment in individuals with psychopathy (Juarez, Kiehl, &
Calhoun, 2012). These studies collectively point towards the
importance of the DMN in psychopathy, and perhaps
emotional detachment in particular. However, it is unknown
whether (i) previously observed functional differences in
psychopathy are associated with abnormalities in the white
matter anatomy of this network; (ii) whether any such dif-
ferences in this network are related specifically to emotional
detachment.
Direct white matter connections between the medial DMN
regions (Greicius, Supekar, Menon, & Dougherty, 2009) lie
within the cingulum, a long association tract which can be
subdivided into functionally and anatomically distinct por-
tions. For the purposes of this study, we identify two distin-
guishable portions: (i) the dorsal cingulum, connecting the
PCC to the mPFC, which is related to social and emotional
aspects of cognition; and (ii) the ventral cingulum, connecting
PCC to the MTL, which is involved in memory and spatial
orientation (Catani & Thiebault De Schotten, 2012).
In the current study we used DTI tractography to analyse
the dorsal and ventral cingulum in offenders with psychopa-
thy and age- and IQ-matched non-offenders. Based on prior
work, we hypothesised that surrogate indices of
 c o r t e x x x x ( 2 0 1 4 ) 1 e9
microstructure in these tract portions, and the dorsal segment
in particular, would be reduced in the psychopathy group and
that this would be correlated with factor 1 scores.
2. Methods and materials
2.1. Ethics and consent
This study was approved by the Camberwell St Giles NHS
research ethics committee (formerly the Joint South London
and Maudsley and the Institute of Psychiatry Research Ethics
Committee/South East London REC 4) (reference 06/Q0706/87).
Participants were fully informed of the study requirements
and risks and given the opportunity to ask any questions
before giving informed consent. This consent included
authorising members of the research group to access partici-
pants' official criminal records.
2.2. Participants
All participants were right-handed men who spoke English as
their first language and had a reading age of above 10 years
old. Participants had no history of significant head injury (i.e.,
leading to loss of consciousness for an hour or longer), no
neurological problems, or any significant visual or hearing
impairment. Patients had no history of claustrophobia or
other contraindications to MRI scanning as assessed by self-
report. Participants were also screened for psychiatric disor-
ders using the Structured Clinical Interview for the DSM-IV
(Spitzer, Williams, Gibbon, & First, 1992), and were excluded
if they had any lifetime history of Axis 1 major mental ill-
nesses, or any substance use disorders in the previous month.
Participants were also administered the Weschler Adult In-
telligence Scale (Weschler, 1997).
All offenders with psychopathy were recruited via the
National Probation Service, and had committed and been
incarcerated for at least one serious violent crime including
murder, attempted murder, rape, and serious assault.
Forensic psychologists assessed psychiatric, medical and
criminal history by preliminary screening with self-report
measures, probation files and probation officer reports.
Criminal records were retrieved from the Police National
Database.
The PCL-R (Hare, 1991) was conducted by forensic psychi-
atrists and psychologists. Interviews were videotaped, and
25% of these were randomly selected and rerated by a second
trained psychiatrist with a reasonable intraclass correlation
coefficient (ICC) for total PCL-R scores (.81). A PCL-R score of
25 was used to define offenders as having psychopathy. The
cut-off of 25 has been used in accordance with previous work
(Craig et al., 2009; Gregory et al., 2012), and based on obser-
vations of cross-cultural differences in psychopathy (Cooke &
Michie, 1999). Healthy non-offenders were recruited via
bulletin boards in local unemployment offices and commu-
nity websites. These individuals did not meet the PCL-R 25
criteria for psychopathy, and had never been convicted of a
criminal offence.
Participants were encouraged to abstain from all substance
misuse for a period of two weeks prior to the scan. Participants
Please cite this article in press as: Sethi, A., et al., Emotional detachment in psychopathy: Involvement of dorsal default-mode
connections, Cortex (2014), http://dx.doi.org/10.1016/j.cortex.2014.07.018
3
were required to have their saliva tested for alcohol use, and a
urine test to detect illicit drug use before the scanning session.
This revealed some offenders tested positive for substances,
despite being requested to refrain from substance use prior to
the scan. A forensic psychologist, and a member of the
research team (S.G.), assessed the participants to ascertain
their suitability to enter the scanning environment and
adhere to the safety protocol. Participants were reimbursed
for their time at the National Minimum Wage rate.
2.3. Scanning
Scanning was performed using a GE 1.5T Sigma Excite MRI
Scanner (actively shielded magnetic field gradients: max amp.
40 mT m1) using a multi-slice doubly refocused spin echo EPI
sequence acquisition based on (Jones et al., 2002) using body
coil RF transmission and 8 channel head coil NMR reception.
60 contiguous near axial slices were acquired with
2.5  2.5  2.5 mm voxels. Acquisition had an echo time of
101.3 msec, with an effective repetition time of 12e20 RR in-
tervals (using cardiac gating). A maximum diffusion weighting
of 1300 sec mm1 was used, and 64 diffusion-weighted brain
volumes with diffusion gradients uniformly distributed in
space were collected at each slice location, as well as 7 vol-
umes without diffusion weighting applied.
2.4. Analysis
Data was corrected for eddy currents and motion distortion,
and the diffusion tensor was estimated. Whole brain trac-
tography (step size: .5 mm; FA threshold: .2; angle threshold:
30 ) was then performed using Euler integration (Basser,
Pajevic, Pierpaoli, Duda, & Aldroubi, 2000). Data was pro-
cessed using ExploreDTI. The dorsal and ventral cingulum
were identified and dissected on the axial plane (Fig. 1) and
tract-specific measures were extracted using TrackVis. The
cingulum was initially defined by one region of interest (ROI)
on the sagittal plane and dissected bilaterally, with an ROI at
the midline to exclude callosal fibres. The division between
the dorsal and ventral portions of the cingulum was defined
anatomically as the point above the splenium of the corpus
callosum on the midline slice. An exclusion ROI was used at
this slice, to exclude fibres of the ventral cingulum that
continued into the region defined as the dorsal cingulum.
Dissections were performed blind to diagnosis by a member of
the research team (AS), and then repeated by a second blinded
member independently (EPT).
Tract measures of fractional anisotropy (FA; an indirect
measure of fibre myelination and axonal integrity and orga-
nisation) and radial diffusivity (an ostensibly more specific
marker of fibre myelination) were examined for between
group differences using one-way ANOVA, and correlations
with PCL-R measures were assessed using Pearson's product-
moment (two-tailed) correlation in SPSS. To control for vari-
ables of no interest, we performed an ANCOVA analysis to
assess difference between the psychopathy and control group.
Dissection reliability was assessed using ICC analyses (two
way mixed absolute agreement model model) for FA mea-
surements in our tracts of interest.
4 c o r t e x x x x ( 2 0 1 4 ) 1 e9

Fig. 1 e (a) Fractional anisotropy colour maps with overlaid ROI for the dorsal (green) and ventral (blue) cingulum. (b) DTI
tractography reconstruction of the dorsal (green) and ventral (blue) cingulum.

Finally, to assess reliability of these dissections we

3. Results
We compared 13 offenders with psychopathy (PCL-R ¼ 28 ± 2),
aged 40 ± 10 years, with full scale IQ 90 ± 12, to 13 non-
offenders (PCL-R ¼ 4 ± 3), aged 34 ± 9 years, with full scale
IQ 98 ± 13. The two groups did not significantly differ in age or
IQ.
One-way ANOVAs revealed that the psychopathy group
had significantly lower FA in the left dorsal cingulum [F (1,
24) ¼ 5.83, p ¼ .024] and ventral cingulum bilaterally [left: F (1,
24) ¼ 7.60, p ¼ .011; right: F (1, 24) ¼ 4.81, p ¼ .038] (Fig. 2). The
observed FA differences in the left dorsal cingulum FA
remained significant even after controlling for alcohol,
cocaine and cannabis dependency (as determined by the SCID;
Table 1) [F (1, 15) ¼ 5.78, p ¼ .030]. FA differences in the ventral
cingulum were no longer significant at p ¼ .05 [left: F (1,
15) ¼ 3.00, p ¼ .104; right: F (1, 15) ¼ 1.46, p ¼ .245]. Radial
diffusivity did not differ significantly between the groups in
either tract portion.
We also report a negative correlation between PCL-R factor
1 scores and FA in the left [r (11) ¼ .61, p ¼ .026] and right [r
(11) ¼ .62, p ¼ .023] dorsal cingulum within the psychopathy
group. Importantly, there was no relationship between factor
2 scores and dorsal cingulum FA, or between either factor and
ventral cingulum FA (Table 2).
employed ICC analyses (two way mixed absolute agreement
model), which showed high agreement between raters (AS &
EPT) in all tracts (Left dorsal cingulum FA: ICC ¼ .99, p < .001;
Left ventral cingulum FA: ICC ¼ .96, p < .001; Right dorsal
cingulum FA: ICC ¼ .99, p < .001; Right ventral cingulum FA:
ICC ¼ .93, p < .001).
4. Discussion
The results from this study extend our understanding of the
role of the paralimbic system in psychopathy. We suggest a
model of the disorder where the main diagnostic features are
at least partly dissociable at the network level. We previously
reported that antisocial behaviour (factor 2) in psychopathy is
associated with abnormalities in the microstructure of a
ventral ‘temporo-amygala-orbitofrontal’ network (connected
by the uncinate fasciculus) (Craig et al., 2009). The current
study extends that work, and suggests that emotional
detachment (factor 1) in psychopathy is associated with ab-
normalities in a different neural circuit e the dorsal ‘default-
mode’ network (Fig. 3). Such a relationship is also supported
by previous functional imaging studies in individuals with
psychopathy (Glenn et al., 2009; Juarez et al., 2012).

Table 1 e Prevalence of substance use disorders within psychopathy and non-offender groups.

Drug % Dependency Fischer's exact test p-value

Offenders with psychopathy Non-offenders
Alcohol 40% 0% 5.00 .087
Cannabis 30% 10% 1.25 .582
Cocaine 30% 0% 3.53 .211
Stimulants 0% 0% e e
Sedatives 0% 0% e e
Opioids 10% 0% 1.05 1.000
Hallucinogens 0% 0% e e
Other 0% 0% e e

N.B. Data incomplete for three participants from each group.

Please cite this article in press as: Sethi, A., et al., Emotional detachment in psychopathy: Involvement of dorsal default-mode
connections, Cortex (2014), http://dx.doi.org/10.1016/j.cortex.2014.07.018
 c o r t e x x x x ( 2 0 1 4 ) 1 e9 5

Fig. 2 e Differences in Fractional Anisotropy (FA) between individuals with psychopathy and non-offenders *p < .05.

Our findings suggest that this relationship is specific to the
dorsal component of the DMN. This is consistent with func-
tions associated with this subdivision of the DMN. The mPFC
is involved in introspection and social function, including
emotional reflection (Gusnard et al., 2001), and evaluating self-
and other-emotional states (Ochsner et al., 2004). The PCC has
been observed to be responsive to emotional stimuli (Maddock
et al., 2003), and is also activated by evaluation of emotional
states (Ochsner et al., 2004). Collectively, these regions are
active during moral judgement (Greene et al., 2001; Harrison
et al., 2008), where they are understood to reflect ‘emotional
engagement’ with moral dilemmas (Greene & Haidt, 2002;
Greene et al., 2001). Similarly, both have also been observed
to be active during socio-affective processing [i.e., empathy
(Vollm et al., 2006)]. Abnormalities in such functions could
plausibly mediate emotional detachment in psychopaths.
Indeed, this is not only consistent with the nature of the traits
described by emotional detachment (e.g., lack of guilt, shallow
affect), but also with specific socio-affective (Blair, 2005;
Shamay-Tsoory, Harari, Aharon-Peretz, & Levkovitz, 2010)
and moral-affective (Koenigs, Kruepke, Zeier, & Newman,
2012) behavioural abnormalities observed in individuals with
psychopathy.
The selective relationship between emotional detachment
and the dorsal DMN also partially distinguishes the DMN
abnormalities observed in psychopathy from those observed
in other psychiatric populations. DMN differences have, for
example, been observed in depression (Sheline et al., 2009),
attention deficit hyperactivity disorder (ADHD) (Castellanos &
Proal, 2012; Sonuga-Barke & Castellanos, 2007), dementias (Bai
et al., 2012; Filippi et al., 2013; Greicius, Srivastava, Reiss, &
Menon, 2004), schizophrenia (Koch et al., 2013), and autism
(Assaf et al., 2010). However, the contribution of the DMN has
been conceptualised differently in each of these disorders. For
example, in ADHD, it has been suggested that they reflect a
difficulty ‘switching’ between default-mode and fronto-
parietal control networks (Castellanos & Proal, 2012; Sonuga-
Barke & Castellanos, 2007). In Alzheimer's disease, dysfunc-
tion has been reported to be associated with the more ventral
PCC-MTL component of this network (Greicius et al., 2004),
thus disconnecting the medial temporal memory system from
other regions.
In psychopathy, abnormalities in a dorsal subdivision of
the DMN associated with introspection and social, moral and
affective processing appear to contribute to emotional
detachment. This is consistent with autistic spectrum condi-
tions which exhibit a similar relationship to the DMN, with
social deficits related to dorsal mPFC and PCC subnetworks
(Assaf et al., 2010). Further work is however required to

Table 2 e Psychopathy checklist revised correlations with

fractional anisotropy in the dorsal and ventral cingulum.

Measure Pearon's r DF p
Factor 1
Left
Dorsal cingulum .61 11 .026*
Ventral cingulum .08 11 .794
Right
Dorsal cingulum .62 11 .023*
Ventral cingulum .01 11 .979
Factor 2
Left
Dorsal cingulum .01 11 .988
Ventral cingulum .30 11 .322
Right
Dorsal cingulum .14 11 .646
Ventral cingulum .43 11 .146 Fig. 3 e The default mode network, connecting the mPFC
and PCC. The temporo-amygdala-orbitofrontal network
*p < .05.
connecting the orbitofrontal cortex (OFC) and amygdala.

Please cite this article in press as: Sethi, A., et al., Emotional detachment in psychopathy: Involvement of dorsal default-mode
connections, Cortex (2014), http://dx.doi.org/10.1016/j.cortex.2014.07.018
6 c o r t e x x x x ( 2 0 1 4 ) 1 e9
determine the exact contribution of overlapping functional
anatomical networks to these disorders and others, and how
differences in distinct sub networks and networkenetwork
interactions lead to the array of psychiatric manifestations
associated with the DMN. Such work is essential for refining
future neuroanatomical models of these disorders.
The proposed ‘dual network’ model of psychopathy sup-
ports a shift in the prevailing conceptualisation of psychopa-
thy, which has focused on differences within, and between,
the amygdala and orbitofrontal cortex (Blair, 2008). One of the
main limitations of this earlier model was that similar dif-
ferences had been reported in other, non-psychopathic, anti-
social populations (Hoptman et al., 2010; Sarkar et al., 2013;
Sundram et al., 2012). The current model can accommodate
these findings, and provide a putative explanation of the two
core features of criminal psychopathy. Although it is probable
that other regions/networks are relevant to psychopathy
(Juarez et al., 2012), findings from this study and others
strongly suggest the importance of the identified dorsal
(Ermer et al., 2012; Glenn et al., 2009; Gregory et al., 2012;
Juarez et al., 2012; Motzkin et al., 2011; de Oliveira-Souza
et al., 2008; Pujol et al., 2011; Yang et al., 2009) and ventral
(Blair, 2008; Craig et al., 2009; Motzkin et al., 2011) networks. In
summary, this dual network model suggests a differential
contribution of ventral and dorsal paralimbic sub-networks to
the discrete features of psychopathy.
The biological mechanism(s) underpinning these network
abnormalities are not yet clear, though they may reflect a
primary deficit in white matter maturation. FA increases with
age during the first two decades of life, and in some tracts
(including the cingulum and uncinate fasciculus) this process
is protracted over a longer period (Lebel, Walker, Leemans,
Phillips, & Beaulieu, 2008). Therefore, reduced FA in psychop-
athy could be related to an aberrant maturational trajectory of
these limbic pathways. This is consistent with findings that
boys with conduct disorder and callous-unemotional traits
exhibit a divergent pattern of white matter development from
typically developing controls (De Brito et al., 2009). However,
white matter differences could also be secondary to develop-
mental changes within ventral and/or dorsal brain regions.
Further work is needed to investigate these hypotheses, and to
characterise the aetiology of any such changes.
Twin studies provide a unique method for studying
development, and may provide clues as to the aetiology of
neurobiological differences in psychopathy. For example,
using this approach, boys with callous-unemotional traits
have been reported to exhibit heritable grey matter anomalies
within the DMN (Rijsdijsk et al., 2010). This suggests that at
least some differences within the DMN appear to be heritable
in psychopathy. However, it is also possible that the reported
differences in white matter microstructure could be influ-
enced by environmental factors. In particular, drug de-
pendency is known to exert a significant effect on FA (Ashtari
et al., 2009; Harris et al., 2008; Lim, Choi, Pomara, Wolkin, &
Rotrosen, 2002; Pfefferbaum, Rosenbloom, Rohlfing, &
Sullivan, 2009; Pfefferbaum & Sullivan, 2005). This potential
confound does not appear to have influenced our results,
however, as the observed reductions in FA remained signifi-
cant after controlling for group differences in life-time di-
agnoses of alcohol, cocaine, and cannabis dependence
Please cite this article in press as: Sethi, A., et al., Emotional detachment in psychopathy: Involvement of dorsal default-mode
connections, Cortex (2014), http://dx.doi.org/10.1016/j.cortex.2014.07.018
between groups. This suggests that the reported association
between emotional detachment and FA in the dorsal
cingulum cannot simply be explained by these environmental
factors. Nevertheless, future studies would benefit from a
control group matched for substance misuse and include
twin, and longitudinal, designs to tease apart putative envi-
ronmental and genetic influences more clearly.
Further studies are also required to overcome other limi-
tations of the current study. This includes the relatively small
sample size, and the need to replicate our findings in a larger
sample. The current study may have also benefited from
assessing the connectivity profile of the angular gyrus, which
contributes to the DMN. Reconstruction of the angular gyrus'
connections with other DMN regions is limited by the inability
of the tensor model to resolve crossing fibres (Greicius et al.,
2009). However, this could potentially be addressed in future
studies, as high angular resolution diffusion imaging (HARDI)
techniques (Tuch et al., 2002), such as spherical deconvolution
(Dell'Acqua et al., 2010; Tournier, Calamante, Gadian, &
Connelly, 2004), continue to develop. Lastly, the proposed
model should be interpreted within the context of its predic-
tive limitations. For example, it is probable that there are other
regions/networks which also contribute to factor 1 and 2 in
psychopaths. In addition, other neurobiological differences
may contribute to functional abnormalities that are not
encapsulated by the PCL-R. For example, we also observed
differences in the ventral cingulum that did not relate to
either PCL-R factor. This is consistent with our understanding
that this tract is more associated with memory and spatial
orientation, than emotional and social processing (Catani &
Thiebault De Schotten, 2012).
5. Conclusions
The current study, combined with previous work, suggests
that it may be possible to fractionate the behavioural pheno-
type of psychopathy [that is, emotional detachment (PCL-R
factor 1) and antisocial behaviour (factor 2)] on the network
level. Whilst differences in a ventral ‘temporo-amygdala-
orbitofrontal’ network are related to antisocial behaviour in
psychopathy, we report that emotional detachment is related
to abnormalities in a dorsal ‘default-mode’ network. Further
work is required to determine the cause of these differences,
and if they predict outcome. Due to the apparent heritability
and early-emergence of affective and interpersonal traits, the
DMN may represent a more specific target for future studies
aimed at understanding the neurodevelopmental trajectory of
psychopathy and potential treatments.
Funding
This research was funded by research grants from: The
Department of Health (the National Forensic Mental Health
R&D programme; MRD 12/102); The Ministry of Justice (a DSPD
programme grant); The Psychiatry Research Trust; The NIHR
Biomedical Research Centre, South London and Maudsley
NHS Foundation Trust, and Institute of Psychiatry (King's
College London).
 c o r t e x x x x ( 2 0 1 4 ) 1 e9
Conflict of interest
None.
Human and animal rights
The authors assert that all procedures contributing to this
work comply with the ethical standards of the relevant na-
tional and institutional committees on human experimenta-
tion and with the Helsinki Declaration of 1975, as revised in
2008. Informed consent was obtained for all human subjects.
Acknowledgements
We gratefully acknowledge the work of Ms. Sam Prior, Ms.
Clare Goodwin, Mr. William Wainwright, Mr. Ruben Azevedo,
Mr. Francis Vergunst, Ms. Lucy Butler, Ms. Leila Niknejad, Dr.
Anna Plodowski, Dr. Philip Baker, Dr. Timothy Rogers, Dr.
Preethi Chabbra, Dr. Stephen Attard, Dr. Seema Sukhwal, Dr.
Nathan Kolla, Dr. Paul Wallang, and Dr. Clare Conway in
participant recruitment and assessment.
references
Ashtari, M., Cervellione, K., Cottone, J., Ardekani, B. A., Sevy, S., &
Kumra, S. (2009). Diffusion abnormalities in adolescents and
young adults with a history of heavy cannabis use. Journal of
Psychiatric Research, 43(3), 189e204. http://dx.doi.org/10.1016/
j.jpsychires.2008.12.002.
Assaf, M., Jagannathan, K., Calhoun, V. D., Miller, L.,
Stevens, M. C., Sahl, R., et al. (2010). Abnormal functional
connectivity of default mode sub-networks in autism
spectrum disorder patients. NeuroImage, 53(1), 247e256. http://
dx.doi.org/10.1016/j.neuroimage.2010.05.067.
Bai, F., Shi, Y., Yuan, Y., Wang, Y., Yue, C., Teng, Y., et al. (2012).
Altered self-referential network in resting-state amnestic type
mild cognitive impairment. Cortex, 48(5), 604e613. http://
dx.doi.org/10.1016/j.cortex.2011.02.011.
Barrash, J., Tranel, D., & Anderson, S. W. (2000). Acquired
personality disturbances associated with bilateral damage to
the ventromedial prefrontal region. Developmental
Neuropsychology, 18(3), 355e381. http://dx.doi.org/10.1207/
S1532694205Barrash.
Barry, C. T., Frick, P. J., DeShazo, T. M., McCoy, M. G., Ellis, M., &
Loney, B. R. (2000). The importance of callous-unemotional
traits for extending the concept of psychopathy to children.
Journal of Abnormal Psychology, 109(2), 335e340.
Basser, P. J., Pajevic, S., Pierpaoli, C., Duda, J., & Aldroubi, A. (2000).
In vivo fiber tractography using DT-MRI data. Journal of
Abnormal Psychology, 44(4), 625e632.
Blair, R. J. (2005). Responding to the emotions of others:
dissociating forms of empathy through the study of typical
and psychiatric populations. Consciousness and Cognition, 14(4),
698e718. http://dx.doi.org/10.1016/j.concog.2005.06.004.
Blair, R. J. (2008). The amygdala and ventromedial prefrontal
cortex: functional contributions and dysfunction in
psychopathy. Philosophical Transactions of the Royal Society of
London, Series B: Biological Sciences, 363(1503), 2557e2565. http://
dx.doi.org/10.1098/rstb.2008.0027.
Please cite this article in press as: Sethi, A., et al., Emotional detachment in psychopathy: Involvement of dorsal default-mode
connections, Cortex (2014), http://dx.doi.org/10.1016/j.cortex.2014.07.018
7
Blair, R. J., & Cipolotti, L. (2000). Impaired social response reversal.
A case of ‘acquired sociopathy’. Brain, 123(Pt 6), 1122e1141.
Blair, R. J., Colledge, E., Murray, L., & Mitchell, D. G. (2001). A
selective impairment in the processing of sad and fearful
expressions in children with psychopathic tendencies. Journal
of Abnormal Child Psychology, 29(6), 491e498.
Boccardi, M., Frisoni, G. B., Hare, R. D., Cavedo, E., Najt, P.,
Pievani, M., et al. (2011). Cortex and amygdala morphology in
psychopathy. Psychiatry Research, 193(2), 85e92. http://
dx.doi.org/10.1016/j.pscychresns.2010.12.013.
Buckner, R. L., Andrews-Hanna, J. R., & Schacter, D. L. (2008). The
brain's default network: anatomy, function, and relevance to
disease. Annuals of the New York Academy of Sciences, 1124,
1e38. http://dx.doi.org/10.1196/annals.1440.011.
Budhani, S., & Blair, R. J. (2005). Response reversal and children
with psychopathic tendencies: success is a function of
salience of contingency change. Journal of Child Psychology and
Psychiatry, 46(9), 972e981. http://dx.doi.org/10.1111/j.1469-
7610.2004.00398.x.
Budhani, S., Richell, R. A., & Blair, R. J. (2006). Impaired reversal
but intact acquisition: probabilistic response reversal deficits
in adult individuals with psychopathy. Journal of Abnormal
Psychology, 115(3), 552e558. http://dx.doi.org/10.1037/0021-
843x.115.3.552.
Castellanos, F. X., & Proal, E. (2012). Large-scale brain systems in
ADHD: beyond the prefrontal-striatal model. Trends in
Cognitive Sciences, 16(1), 17e26. http://dx.doi.org/10.1016/
j.tics.2011.11.007.
Catani, M., & Thiebault de Schotten, M. (2012). Atlas of human brain
connections. Oxford: Oxford University Press.
Cooke, D. J., & Michie, C. (1999). Psychopathy across cultures:
North America and Scotland compared. Journal of Abnormal
Psychology, 108(1), 58e68.
Craig, M. C., Catani, M., Deeley, Q., Latham, R., Daly, E.,
Kanaan, R., et al. (2009). Altered connections on the road to
psychopathy. Molecular Psychiatry, 14(10), 946e953. http://
dx.doi.org/10.1038/mp.2009.40, 907.
Damasio, H., Grabowski, T., Frank, R., Galaburda, A. M., &
Damasio, A. R. (1994). The return of Phineas Gage: clues about
the brain from the skull of a famous patient. Science, 264(5162),
1102e1105.
De Brito, S. A., Mechelli, A., Wilke, M., Laurens, K. R., Jones, A. P.,
Barker, G. J., et al. (2009). Size matters: increased grey matter
in boys with conduct problems and callous-unemotional
traits. Brain, 132(Pt 4), 843e852. http://dx.doi.org/10.1093/
brain/awp011.
Dell'Acqua, F., Scifo, P., Rizzo, G., Catani, M., Simmons, A.,
Scotti, G., et al. (2010). A modified damped Richardson-Lucy
algorithm to reduce isotropic background effects in spherical
deconvolution. NeuroImage, 49(2), 1446e1458. http://dx.doi.org/
10.1016/j.neuroimage.2009.09.033.
Ermer, E., Cope, L. M., Nyalakanti, P. K., Calhoun, V. D., &
Kiehl, K. A. (2012). Aberrant paralimbic gray matter in criminal
psychopathy. Journal of Abnormal Psychology, 121(3), 649e658.
http://dx.doi.org/10.1037/a0026371.
Filippi, M., Agosta, F., Scola, E., Canu, E., Magnani, G., Marcone, A.,
et al. (2013). Functional network connectivity in the behavioral
variant of frontotemporal dementia. Cortex, 49(9), 2389e2401.
http://dx.doi.org/10.1016/j.cortex.2012.09.017.
Forsman, M., Lichtenstein, P., Andershed, H., & Larsson, H. (2008).
Genetic effects explain the stability of psychopathic
personality from mid- to late adolescence. Journal of Abnormal
Psychology, 117(3), 606e617. http://dx.doi.org/10.1037/0021-
843X.117.3.606.
Fox, M. D., Snyder, A. Z., Vincent, J. L., Corbetta, M., Van
Essen, D. C., & Raichle, M. E. (2005). The human brain is
intrinsically organized into dynamic, anticorrelated functional
networks. Proceedings of the National Academy of the Sciences of
8 c o r t e x x x x ( 2 0 1 4 ) 1 e9
the United States of America, 102(27), 9673e9678. http://
dx.doi.org/10.1073/pnas.0504136102.
Fransson, P. (2005). Spontaneous low-frequency BOLD signal
fluctuations: an fMRI investigation of the resting-state default
mode of brain function hypothesis. Human Brain Mapping,
26(1), 15e29. http://dx.doi.org/10.1002/hbm.20113.
Frick, P. J., Kimonis, E. R., Dandreaux, D. M., & Farell, J. M. (2003).
The 4 year stability of psychopathic traits in non-referred
youth. Behavioural Sciences & the Law, 21(6), 713e736. http://
dx.doi.org/10.1002/bsl.568.
Frick, P. J., & Viding, E. (2009). Antisocial behavior from a
developmental psychopathology perspective. Developmental
Psychopathology, 21(4), 1111e1131. http://dx.doi.org/10.1017/
s0954579409990071.
Glenn, A. L., Raine, A., & Schug, R. A. (2009). The neural correlates
of moral decision-making in psychopathy. Molecular
Psychiatry, 14(1), 5e6. http://dx.doi.org/10.1038/mp.2008.104.
Greene, J., & Haidt, J. (2002). How (and where) does moral
judgment work? Trends in Cognitive Science, 6(12), 517e523.
Greene, J. D., Sommerville, R. B., Nystrom, L. E., Darley, J. M., &
Cohen, J. D. (2001). An fMRI investigation of emotional
engagement in moral judgment. Science, 293(5537), 2105e2108.
http://dx.doi.org/10.1126/science.1062872.
Gregory, S., Ffytche, D., Simmons, A., Kumari, V., Howard, M.,
Hodgins, S., et al. (2012). The antisocial brain: psychopathy
matters: a structural MRI investigation of antisocial male
violent offenders. Archives of General Psychiatry. http://
dx.doi.org/10.1001/archgenpsychiatry.2012.222.
Greicius, M. D., Srivastava, G., Reiss, A. L., & Menon, V. (2004).
Default-mode network activity distinguishes Alzheimer's
disease from healthy aging: evidence from functional MRI.
Proceedings of the National Academy of the Sciences of the United
States of America, 101(13), 4637e4642. http://dx.doi.org/10.1073/
pnas.0308627101.
Greicius, M. D., Supekar, K., Menon, V., & Dougherty, R. F. (2009).
Resting-state functional connectivity reflects structural
connectivity in the default mode network. Cerebral Cortex,
19(1), 72e78. http://dx.doi.org/10.1093/cercor/bhn059.
Gusnard, D. A., Akbudak, E., Shulman, G. L., & Raichle, M. E.
(2001). Medial prefrontal cortex and self-referential mental
activity: relation to a default mode of brain function.
Proceedings of the National Academy of the Sciences of the United
States of America, 98(7), 4259e4264. http://dx.doi.org/10.1073/
pnas.071043098.
Hare, R. D. (1991). The hare psychopathy checklist e Revised. Toronto,
Ontario, Canada: Multi-Health Systems.
Hare, R. D. (2003). The hare psychopathy checklist e Revised (PCLeR)
manual (2nd ed.). Toronto, Ontario, Canada: Multi-Health
Systems.
Hare, R. D., Harpur, T. J., Hakstian, A. R., Forth, A. E., Hart, S. D., &
Newman, J. P. (1990). The revised psychopathy checklist:
reliability and factor structure. Psychological Assessment, 2(3),
338e341. http://dx.doi.org/10.1037/1040-3590.2.3.338.
Harlow, J. M. (1993). Recovery from the passage of an iron bar
through the head. 1868. History of Psychiatry, 4(14), 274e281.
http://dx.doi.org/10.1177/0957154x9300401407.
Harlow, J. M. (1999). Passage of an iron rod through the head. 1848.
The Journal of Neuropsychiatry and Clinical Neurosciences, 11(2),
281e283.
Harpur, T. J., Hakstian, A. R., & Hare, R. D. (1988). Factor structure
of the psychopathy checklist. Journal of Consulting and Clinical
Psychology, 56(5), 741e747.
Harpur, T. J., Hare, R. D., & Hakstian, A. R. (1989). Two-factor
conceptualization of psychopathy: construct validity and
assessment implications. Psychological Assessment, 1(1), 6e17.
http://dx.doi.org/10.1037/1040-3590.1.1.6.
Harris, G. J., Jaffin, S. K., Hodge, S. M., Kennedy, D., Caviness, V. S.,
Marinkovic, K., et al. (2008). Frontal white matter and
Please cite this article in press as: Sethi, A., et al., Emotional detachment in psychopathy: Involvement of dorsal default-mode
connections, Cortex (2014), http://dx.doi.org/10.1016/j.cortex.2014.07.018
cingulum diffusion tensor imaging deficits in alcoholism.
Alcoholism: Clinical and Experimental Research, 32(6), 1001e1013.
http://dx.doi.org/10.1111/j.1530-0277.2008.00661.x.
Harrison, B. J., Pujol, J., Lo  pez-Sola
 , M., Herna
 ndez-Ribas, R.,
Deus, J., Ortiz, H., et al. (2008). Consistency and functional
specialization in the default mode brain network. Proceedings
of the National Academy of the Sciences of the United States of
America, 105(28), 9781e9786. http://dx.doi.org/10.1073/
pnas.0711791105.
Hart, S. D., & Hare, R. D. (1997). Psychopathy: assessment and
association with criminal conduct. In D. M. Stoff, J. Breiling, &
J. D. Maser (Eds.), Handbook of antisocial behaviour. New York: John
Wiley & Sons, Inc.
Hemphill, J. F., Hare, R. D., & Wong, S. (1998). Psychopathy and
recidivism: a review. Legal and Criminological Psychology, 3(1),
139e170. http://dx.doi.org/10.1111/j.2044-8333.1998.tb00355.x.
Hoptman, M. J., D'Angelo, D., Catalano, D., Mauro, C. J.,
Shehzad, Z. E., Kelly, A. M., et al. (2010). Amygdalofrontal
functional disconnectivity and aggression in schizophrenia.
Schizophrenia Bulletin, 36(5), 1020e1028. http://dx.doi.org/
10.1093/schbul/sbp012.
Johnson, M. K., Raye, C. L., Mitchell, K. J., Touryan, S. R.,
Greene, E. J., & Nolen-Hoeksema, S. (2006). Dissociating medial
frontal and posterior cingulate activity during self-reflection.
Social Cognitive & Affective Neuroscience, 1(1), 56e64. http://
dx.doi.org/10.1093/scan/nsl004.
Jones, D. K., Williams, S. C., Gasston, D., Horsfield, M. A.,
Simmons, A., & Howard, R. (2002). Isotropic resolution diffusion
tensor imaging with whole brain acquisition in a clinically
acceptable time. Human Brain Mapping, 15(4), 216e230.
Juarez, M., Kiehl, K. A., & Calhoun, V. D. (2012). Intrinsic limbic
and paralimbic networks are associated with criminal
psychopathy. Human Brain Mapping. http://dx.doi.org/10.1002/
hbm.22037.
Kelley, W. M., Macrae, C. N., Wyland, C. L., Caglar, S., Inati, S., &
Heatherton, T. F. (2002). Finding the self? An event-related
fMRI study. Journal of Cognitive Neuroscience, 14(5), 785e794.
http://dx.doi.org/10.1162/08989290260138672.
Kiehl, K. A. (2006). A cognitive neuroscience perspective on
psychopathy: evidence for paralimbic system dysfunction.
Psychiatry Research, 142(2e3), 107e128. http://dx.doi.org/
10.1016/j.psychres.2005.09.013.
Kiehl, K. A., & Hoffman, M. B. (2011). The criminal psychopath:
history, neuroscience, treatment, and economics. Jurimetrics,
51(4), 355e397.
Kiehl, K. A., Smith, A. M., Hare, R. D., Mendrek, A., Forster, B. B.,
Brink, J., et al. (2001). Limbic abnormalities in affective processing
by criminal psychopaths as revealed by functional magnetic
resonance imaging. Biological Psychiatry, 50(9), 677e684.
King, H. E. (1961). Psychological effects of excitation of the limbic
system. In D. E. Sheer (Ed.), Electrical stimulation of the brain.
Houston, Texas,: Hoggs Foundation, University of Texas Press.
Kluver, H., & Bucy, P. C. (1997). Preliminary analysis of functions
of the temporal lobes in monkeys. 1939. The Journal of
Neuropsychiatry and Clinical Neurosciences, 9(4), 606e620.
Koch, K., Schultz, C. C., Wagner, G., Schachtzabel, C.,
Reichenbach, J. R., Sauer, H., et al. (2013). Disrupted white
matter connectivity is associated with reduced cortical
thickness in the cingulate cortex in schizophrenia. Cortex,
49(3), 722e729. http://dx.doi.org/10.1016/j.cortex.2012.02.001.
Koenigs, M., Kruepke, M., Zeier, J., & Newman, J. P. (2012).
Utilitarian moral judgment in psychopathy. Social Cognitive &
Affective Neuroscience, 7(6), 708e714. http://dx.doi.org/10.1093/
scan/nsr048.
Lebel, C., Walker, L., Leemans, A., Phillips, L., & Beaulieu, C. (2008).
Microstructural maturation of the human brain from
childhood to adulthood. NeuroImage, 40(3), 1044e1055. http://
dx.doi.org/10.1016/j.neuroimage.2007.12.053.
 c o r t e x x x x ( 2 0 1 4 ) 1 e9
Levenston, G. K., Patrick, C. J., Bradley, M. M., & Lang, P. J. (2000).
The psychopath as observer: emotion and attention in picture
processing. Journal of Abnormal Psychology, 109(3), 373e385.
Lim, K. O., Choi, S. J., Pomara, N., Wolkin, A., & Rotrosen, J. P.
(2002). Reduced frontal white matter integrity in cocaine
dependence: a controlled diffusion tensor imaging study.
Biological Psychiatry, 51(11), 890e895.
Maddock, R. J., Garrett, A. S., & Buonocore, M. H. (2003). Posterior
cingulate cortex activation by emotional words: fMRI evidence
from a valence decision task. Human Brain Mapping, 18(1),
30e41. http://dx.doi.org/10.1002/hbm.10075.
Motzkin, J. C., Newman, J. P., Kiehl, K. A., & Koenigs, M. (2011).
Reduced prefrontal connectivity in psychopathy. Journal of
Neuroscience, 31(48), 17348e17357. http://dx.doi.org/10.1523/
JNEUROSCI.4215-11.2011.
de Oliveira-Souza, R., Hare, R. D., Bramati, I. E., Garrido, G. J.,
Azevedo Ignacio, F., Tovar-Moll, F., et al. (2008). Psychopathy
as a disorder of the moral brain: fronto-temporo-limbic grey
matter reductions demonstrated by voxel-based
morphometry. NeuroImage, 40(3), 1202e1213. http://dx.doi.org/
10.1016/j.neuroimage.2007.12.054.
Ochsner, K. N., Knierim, K., Ludlow, D. H., Hanelin, J.,
Ramachandran, T., Glover, G., et al. (2004). Reflecting upon
feelings: an fMRI study of neural systems supporting the
attribution of emotion to self and other. Journal of Cognitive
Neuroscience, 16(10), 1746e1772. http://dx.doi.org/10.1162/
0898929042947829.
Pfefferbaum, A., Rosenbloom, M., Rohlfing, T., & Sullivan, E. V.
(2009). Degradation of association and projection white matter
systems in alcoholism detected with quantitative fiber
tracking. Biological Psychiatry, 65(8), 680e690. http://dx.doi.org/
10.1016/j.biopsych.2008.10.039.
Pfefferbaum, A., & Sullivan, E. V. (2005). Disruption of brain white
matter microstructure by excessive intracellular and
extracellular fluid in alcoholism: evidence from diffusion
tensor imaging. Neuropsychopharmacology, 30(2), 423e432.
http://dx.doi.org/10.1038/sj.npp.1300623.
Pujol, J., Batalla, I., Contreras-Rodriguez, O., Harrison, B. J.,
Pera, V., Hernandez-Ribas, R., et al. (2011). Breakdown in the
brain network subserving moral judgment in criminal
psychopathy. Social Cognitive & Affective Neuroscience. http://
dx.doi.org/10.1093/scan/nsr075.
Raichle, M. E., MacLeod, A. M., Snyder, A. Z., Powers, W. J.,
Gusnard, D. A., & Shulman, G. L. (2001). A default mode of
brain function. Proceedings of the National Academy of the
Sciences of the United States of America, 98(2), 676e682. http://
dx.doi.org/10.1073/pnas.98.2.676.
Raine, A., Buchsbaum, M., & LaCasse, L. (1997). Brain
abnormalities in murderers indicated by positron emission
tomography. Biological Psychiatry, 42(6), 495e508. http://
dx.doi.org/10.1016/s0006-3223(96)00362-9.
Raine, A., Lencz, T., Bihrle, S., LaCasse, L., & Colletti, P. (2000).
Reduced prefrontal gray matter volume and reduced
autonomic activity in antisocial personality disorder. Archives
of General Psychiatry, 57(2), 119e127. discussion 128e119.
Rijsdijsk, F. V., Viding, E., De Brito, S., Forgiarini, M., Mechelli, A.,
Jones, A. P., et al. (2010). Heritable variations in gray matter
concentration as a potential endophenotype for psychopathic
traits. Archives of General Psychiatry, 67(4), 406e413. http://
dx.doi.org/10.1001/archgenpsychiatry.2010.20.
Sarkar, S., Craig, M. C., Catani, M., Dell'acqua, F., Fahy, T.,
Deeley, Q., et al. (2013). Frontotemporal white-matter
microstructural abnormalities in adolescents with conduct
disorder: a diffusion tensor imaging study. Psychological
Medicine, 43(2), 401e411. http://dx.doi.org/10.1017/
S003329171200116X.
Saver, J. L., & Damasio, A. R. (1991). Preserved access and
processing of social knowledge in a patient with acquired
Please cite this article in press as: Sethi, A., et al., Emotional detachment in psychopathy: Involvement of dorsal default-mode
connections, Cortex (2014), http://dx.doi.org/10.1016/j.cortex.2014.07.018
9
sociopathy due to ventromedial frontal damage.
Neuropsychologia, 29(12), 1241e1249.
Shamay-Tsoory, S. G., Harari, H., Aharon-Peretz, J., &
Levkovitz, Y. (2010). The role of the orbitofrontal cortex in
affective theory of mind deficits in criminal offenders with
psychopathic tendencies. Cortex, 46(5), 668e677. http://
dx.doi.org/10.1016/j.cortex.2009.04.008.
Sheline, Y. I., Barch, D. M., Price, J. L., Rundle, M. M.,
Vaishnavi, S. N., Snyder, A. Z., et al. (2009). The default mode
network and self-referential processes in depression.
Proceedings of the National Academy of the Sciences of the United
States of America, 106(6), 1942e1947. http://dx.doi.org/10.1073/
pnas.0812686106.
Shulman, G. L., Fiez, J. A., Corbetta, M., Buckner, R. L.,
Miezin, F. M., Raichle, M. E., et al. (1997). Common blood flow
changes across visual tasks: II. Decreases in cerebral cortex.
Journal of Cognitive Neuroscience, 9(5), 648e663. http://
dx.doi.org/10.1162/jocn.1997.9.5.648.
Skeem, J. L., & Cooke, D. J. (2010). Is criminal behavior a central
component of psychopathy? Conceptual directions for
resolving the debate. Psychological Assessment, 22(2), 433e445.
http://dx.doi.org/10.1037/a0008512.
Sonuga-Barke, E. J., & Castellanos, F. X. (2007). Spontaneous
attentional fluctuations in impaired states and pathological
conditions: a neurobiological hypothesis. Neuroscience &
Biobehavioural Reviews, 31(7), 977e986. http://dx.doi.org/
10.1016/j.neubiorev.2007.02.005.
Spitzer, R. L., Williams, J. B., Gibbon, M., & First, M. B. (1992). The
structured clinical interview for DSM-III-R (SCID). I: history,
rationale, and description. Archives of General Psychiatry, 49(8),
624e629.
Sundram, F., Deeley, Q., Sarkar, S., Daly, E., Latham, R., Craig, M.,
et al. (2012). White matter microstructural abnormalities in the
frontal lobe of adults with antisocial personality disorder. Cortex,
48(2), 216e229. http://dx.doi.org/10.1016/j.cortex.2011.06.005.
Tournier, J. D., Calamante, F., Gadian, D. G., & Connelly, A. (2004).
Direct estimation of the fiber orientation density function
from diffusion-weighted MRI data using spherical
deconvolution. NeuroImage, 23(3), 1176e1185. http://dx.doi.org/
10.1016/j.neuroimage.2004.07.037.
Tuch, D. S., Reese, T. G., Wiegell, M. R., Makris, N., Belliveau, J. W.,
& Wedeen, V. J. (2002). High angular resolution diffusion
imaging reveals intravoxel white matter fiber heterogeneity.
Journal of Abnormal Psychology, 48(4), 577e582. http://
dx.doi.org/10.1002/mrm.10268.
Veit, R., Flor, H., Erb, M., Hermann, C., Lotze, M., Grodd, W., et al.
(2002). Brain circuits involved in emotional learning in
antisocial behavior and social phobia in humans. Neuroscience
Letters, 328(3), 233e236.
Viding, E., Blair, R. J., Moffitt, T. E., & Plomin, R. (2005). Evidence for
substantial genetic risk for psychopathy in 7-year-olds. Journal
of Child Psychology and Psychiatry, 46(6), 592e597. http://
dx.doi.org/10.1111/j.1469-7610.2004.00393.x.
Vollm, B. A., Taylor, A. N., Richardson, P., Corcoran, R., Stirling, J.,
McKie, S., et al. (2006). Neuronal correlates of theory of mind
and empathy: a functional magnetic resonance imaging study
in a nonverbal task. NeuroImage, 29(1), 90e98. http://
dx.doi.org/10.1016/j.neuroimage.2005.07.022.
Weschler, D. (1997). WAIS-III: Administration and scoring manual.
San Antonio, Texas: Harcourt Brace and Co.
Wootton, J. M., Frick, P. J., Shelton, K. K., & Silverthorn, P. (1997).
Ineffective parenting and childhood conduct problems: the
moderating role of callous-unemotional traits. Journal of
Consulting and Clinical Psychology, 65(2), 301e308.
Yang, Y., Raine, A., Colletti, P., Toga, A. W., & Narr, K. L. (2009).
Abnormal temporal and prefrontal cortical gray matter
thinning in psychopaths. Molecular Psychiatry, 14(6), 561e562.
http://dx.doi.org/10.1038/mp.2009.12, 555.